UNIVERSITY OF
ILLINOIS LIBRARY
AT URBANA CHAMPAIGN
BIOLOGY
3 1 1984
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V
FLORA OF GUATEMALA
PAUL C. STANDLEY
AND
LOUIS 0. WILLIAMS
THE
LOGANIAGEAE
BY
DOROTHY N. GIBSON
FIELDIANA: BOTANY
VOLUME 24, PART VIII, NUMBER 4
Published by
FIELD MUSEUM OF NATURAL HISTORY
JUNE 30, 1969
MAY 17 1972
The t*f*fV of the
MAY 15 v
University of HHnois
•t Urburw-Champaipn
FLORA OF GUATEMALA
PART VIII
FLORA OF GUATEMALA
PAUL C. STANDLEY
The Late Curator of the Herbarium
AND
LOUIS 0. WILLIAMS
Chief Curator, Botany
THE
LOGANIACEAE
BY
DOROTHY N. GIBSON
Supervisor of the Herbarium
FIELDIANA: BOTANY
VOLUME 24, PART VIII, NUMBER 4
Published by
FIELD MUSEUM OF NATURAL HISTORY
JUNE 30, 1969
Library of Congress Catalog Card Number: 1+8-3076
PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS
CONTENTS
Families Included in Part VIII, Number 4
PAGE PAGE
Contortae 263 Gentianaceae . . 302
Oleaceae 264 Apocynaceae 334
Loganiaceae 276 Asclepiadaceae 407
LIST OF ILLUSTRATIONS
TEXT FIGURES
PAGE
72. Forestiera rhamnifolia 265
73. Fraxinus uhdei 268
74. Ligustrum lucidum 272
75. Linociera oblanceolata 274
76. Buddleia megalocephala 282
77. Cynoctonum petiolatum 285
78. Gelsemium sempervirens 287
79. Plocosperma buxifoliiim 289
80. Polypremum procumbens 290
81. Spigelia pygmaea 295
82. Strychnos peckii 299
83. Centarium rosans 306
84. Chelonanthus alatus 309
85. Coutoubea spicata 311
86. Centiana lewisiae 314
87. Halenia crassiuscula 317
88. Lisianthus nigrescens var. ctispidatus 325
89. Nymphoides humboldtianum 327
90. Schultesia guianensis 329
91. Voyria thalesioides 333
92. Aspidosperma megalocarpon 339
93. Cameraria latifolia 342
94. Couma macrocarpa 344
95. Echites tuxtlensis 347
96. Fernaldia pandurata 349
97. Forsteronia myriantha 351
98. Forsteronia viridescens 353
99. Haplophyton cinereum 355
100. Lacmellea standleyi 357
101. Malouetia guatemalensis 360
102. Mandevilla donnell-smithii 363
103. Mesechites trifida 367
104. Odontadcnia caudigera 369
105. Plumeria rubra 372
106. Prestonia concolor 377
107. Rauvolfia tetraphylla 382
108. Rhabdadenia biflora 384
109. Stemmadenia obovata 388
110. Tabernaemontana amygdalifolia 391
111. Thevetia ahouai 395
vii
PAGE
112. Thevetia ovata 397
113. Tintinnabularia mortonii 399
114. Tondnzia macrantha 403
115. Urechites andrieuxii 404
116. Vallesia mexicana 406
117. Asclepias woodsoniana 414
118. Blepharodon mucronatum 416
119. Cynanchum schlechtendahlii 423
120. Fischeria martiana 427
121. Gonolobus barbatus 431
122. Gonolobus donnellsmithianus 433
123. Gonolobus uniflorus 439
124. Marsdenia macrophylla 445
125. Matelea diffusa 456
126. Matelea quirosii 462
127. Oxypetalum cordifolium 467
128. Sarcostemma clausum 469
viii
Flora of Guatemala - Part VIII, Number 4
CONTORTAE1
The plants are herbs, shrubs or trees often of great size, the leaves are usually
opposite and entire although pinnately compound leaves are to be found, especially
in Oleaceae; they are estipulate or stipulate. The corolla is sympetalous or the
petals nearly free, or sometimes none, with lobes convolutive, the flowers bisexual
or unisexual (then petals may be missing), the stamens usually 2, 4 or 5, the ovary
is bilocular and superior.
The families here called "the Contortae" were treated by Engler
and Prantl as a group of six families of which five are to be found in
Guatemala — Oleaceae, Loganiaceae, Gentianaceae, Apocynaceae and
Asclepiadaceae. The sixth family, Salvadoraceae, was removed and
eventually the Desfontaineaceae added to the order. The recent
(12th) edition of the Syllabus der Pflanzenfamilien divides the order
into two as Oleales and Gentianales, discarding the name "Con-
tortae." The families remain the same with the notable addition of
the Rubiaceae to the Gentianales. The Rubiaceae follows later in
the system which we are using.
Hutchinson in the second edition of his Families of Flowering
Plants places the families mentioned in the paragraph above into
three orders: the Loganiales, the Apocynales and the Gentianales.
The first of these, the Loganiales, is divided into seven families (of
which five in Guatemala) but essentially the Loganiaceae and Olea-
ceae of our flora; the second, the Apocynales, consists of the Apocyna-
ceae and Asclepiadaceae; the third, the Gentianales, divided into two
families is the Gentianaceae of this flora.
Many ornamental plants belong in this order, perhaps the best
known in the temperate regions is the lilac. Olives are perhaps the
most important of the economic food plants of the group; there are
many that produce alkaloides used in medicine or for other purposes,
1 Much of the field work; work in the herbarium and library; and assistants for
the principal investigator (and junior author) have been made possible by continu-
ing grants from National Science Foundation.
263
264 FIELDIANA: BOTANY, VOLUME 24
strychnine is one of the better known ones. Timbers are to be found
in the group. Some grow into large trees in the wet tropics.
OLEACEAE
Reference: Knoblauch, Oleaceae in Engler & Prantl, Pflanzenf.
IV, 2: 1-16. 1895.
Trees, shrubs or herbs, often scandent. Leaves usually opposite, rarely alter-
nate or verticillate, simple or pinnately compound; stipules absent; flowers actino-
morphic, bisexual or rarely unisexual, borne in racemose, paniculate or thyrisiform
terminal or lateral inflorescences; calyx usually 4-lobed (sometimes more or none);
corolla sympetalous or the petals free or nearly so, with usually 4 lobes imbricated
or induplicate-valvate in bud; stamens 2, rarely 4, epipetalous, bilocular, the loculi
opening lengthwise and the cells usually back to back, often muticious; disk none;
ovary superior, bilocular, style simple with a thickened or bifid stigma; ovules usu-
ally 2 in each cell, rarely 1 or 4-8; fruit capsular and loculicidally bivalvate, or
indehiscent, often of samaras, or baccate or drupaceous, embryo straight.
A small family of some 30 genera and 500 species in temperate
and tropical regions of the world. The most important species of the
family is the olive, Olea europea L., from which the fruit is eaten when
pickled. Olive oil, one of the best food and cooking oils, is expressed
from the ripe fruits.
Olives were said to have been planted in Guatemala as early as
1600 but the cultivation apparently was not successful. So far as we
know there are now no olive trees in the country. Several fine orna-
mentals belong in the family, lilacs perhaps the best known in tem-
perate regions.
Fruit a samara; leaves pinnate; trees Fraxinus.
Fruit not a samara; leaves simple — or pinnate in one cultivated species of Jas-
minum; trees, shrubs or woody vines.
Fruits didymous; cultivated shrubs or vines; leaves simple or pinnate.
Jasminum.
Fruits entire, not didymous; trees or shrubs, never vines; leaves simple and
entire.
Calyx none or minute; corolla none or of one or two petals Forestiera.
Calyx well developed; corolla sympetalous or of four separate or nearly sep-
arate petals.
Corolla of separate or nearly separate linear petals; native trees. .Linociera.
Corolla sympetalous; native or cultivated trees or shrubs.
Corolla lobes imbricate; inflorescence axillary; native trees. .Osmanthus.
Corolla lobes induplicate-valvate; inflorescences terminal; cultivated street
trees Ligustrum.
FORESTIERA Poiret
Dioecious trees or shrubs. Leaves opposite, often on short branches, usually
deciduous, entire or denticulate; inflorescence a fascicle of one to few flowers, rarely
FIG. 72. Forestiera rhamnifolia. A, habit, X 1A; B, fruit, X 3. Osmanthus
mexicana. C, habit, X 1A', D, flower, X 5; E, pistil and calyx, X 5.
265
266 FIELDIANA: BOTANY, VOLUME 24
a few-flowered raceme, borne on old wood or in the axils of leaves, bracteate,
flowers small, commonly imperfect; calyx tube short, the limb 4-6-lobate; corolla
none or sometimes of 1-2 small, free petals; stamens 2 or 4; ovary 2-celled, usually
with 2 seeds in each cell; stigmas thickened, sometimes bilobate; fruit drupaceous,
usually 1-seeded.
The genus is an uncommon one in Central America, three species
are known, each from one or few collections. There are several spe-
cies in Mexico.
Forestiera rhamnifolia Griseb. Cat. PL Cub. 169. 1866.
Along the edge of mangrove swamps; British Honduras (Corn-
house Creek, H. H. Bartlett 11283). Reported from Socorro Islands,
west coast of Mexico; Tamaulipas; West Indies.
Small dioecious trees, in Cuba as much as 12 m. high, the branchlets minutely
puberulent or glabrous, the older ones with numerous large pale elevated lenticels;
leaves membranaceous, short-petiolate, broadly elliptic to lance-oblong, 5-8 cm.
long, 2-5 cm. broad, usually abruptly contracted at the apex, with an obtuse tip,
rarely attenuate, acute at the base or abruptly contracted, entire or remotely and
obscurely crenulate, glabrous above, glabrous or pubescent beneath and with
numerous small scattered pores; flowers unisexual, very small, in small few-flowered
racemes mostly from defoliated nodes, the racemes glabrous, little if at all longer
than the petioles; staminate calyx almost obsolete; filaments rather thick, much
longer than the subglobose anthers; fruit ellipsoid, purple, glaucous, about 8 mm.
long, obtuse at each end, often tipped by the slender short persistent style.
FRAXINUS L. Ash
Reference: A. Lingelsheim, Oleaceae-Oleoideae-Fraxineae, Pflan-
zenreich IV. 243, I: 1-65. 1920.
Small or large, deciduous trees; leaves opposite, pinnate, with few or numerous
leaflets, or simple, these usually dentate, usually appearing before the flowers;
flowers small, perfect or unisexual; calyx small, 4-parted or 4-lobate or wanting;
corolla of 2-6, usually 4, distinct petals, sometimes none; stamens generally 2; ovary
2-celled, with 2 stigmas, ovules 2 in each cell; fruit a 1-seeded nutlet or samara,
commonly with an elongate wing at the apex; seed 1, oblong, with endosperm.
Species about 65, in the northern hemisphere, in America extend-
ing southward to Honduras. The wood of ash (Spanish "fresno") is
dark brown, often hard, heavy, tough and strong, rather coarse in
texture, of excellent working qualities, not durable. In the United
States it is much used for tool handles, agricultural implements, bent
work, oars, kitchen furniture and interior trim. In Central America
the trees are not common enough to be of economic importance. Al-
though we have collected material of this genus assiduously in Guate-
mala, the amount at hand is disappointingly small and unsatisfactory.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 267
Only rarely have we found the trees with flowers or fruit. If a large
amount of material in good condition could be collected, it would be
possible to present a much more satisfactory account of the few
Central American species.
Lateral leaflets sessile F. vellerea.
Lateral leaflets petiolulate.
Fruits 4.5-5 cm. long F. cavekiana.
Fruits 1.5-3 cm. long.
Fruit 2.5-3 cm. long F. uhdei.
Fruits about 1.5 cm. long F. sp.
Fraxinus cavekiana Standl. & Steyerm. Field Mus. Bot. 23:
74. 1944.
Usually along stream banks, sometimes in moist mixed forest,
1,500-2,000 m.; endemic; Guatemala; Quiche" (type from Nebaj,
Skutchl660).
A small or large tree, sometimes 23 m. high, the branchlets thick, at first puber-
ulent, soon glabrate, blackish-brown, bearing sparse large pale lenticels; leaves
large, 5-9-foliolate, the leaflets all petiolulate, the petiolules often much elongate;
leaflets ovate to oblong-lanceolate, mostly 8-14 cm. long and 3-7 cm. broad, acu-
minate to narrowly long-acuminate, usually acute at the base or abruptly con-
tracted, membranaceous, green above and somewhat lustrous, glabrous, the costa
and nerves impressed, somewhat paler beneath, glabrous or sparsely pubescent,
often barbate along the costa, the nerves very prominent, the veins prominulous,
laxly reticulate; fruiting panicles large, about 22 cm. long, lax, the short pedicels
filiform; fruit 4.5-5 cm. long, the seed-bearing portion 1.5 cm. long, subterete, the
wing obtuse, 6-7 mm. broad, decurrent to the middle of seed-bearing portion.
Only two collections with fruit are available, and the determina-
tion of some of the others referred here is therefore uncertain. The
species name is derived from Cavek, the name of the royal house of
Quiche", from its foundation until its overthrow by the Spaniards.
Fraxinus uhdei (Wenzig) Lingelsheim, Bot. Jahrb. 40: 221. 1907.
Fraxinus americana var. uhdei Wenzig, Bot. Jahrb. 4: 182. 1883. Frax-
inus chiapensis Lundell, Contrib. Univ. Mich. Herb. 7: 45. 1942.
Madre de agua.
Usually along stream banks, 800-2,000 m.; Guatemala; Huehue-
tenango. Western and southern Mexico.
A tree of 12-15 m., the young branchlets puberulent, soon glabrous; leaves
large, mostly 5-7-foliolate; leaflets mostly long-petiolulate, firm-membranaceous,
lanceolate, oblong-lanceolate or elliptic, mostly 8-15 cm. long, long acuminate,
obtuse or rounded at the base and often abruptly contracted, the terminal leaflet
usually acute at the base, obscurely serrulate, green and glabrous above, the vena-
FIG. 73. Fraxinus uhdei. A, a leaf, X 1A', B, fruiting inflorescence, X
C, fruit, X 2.y2.
268
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 269
tion prominulous and closely reticulate, little paler beneath, almost glabrous or
often densely velutinous-pubescent; panicles lax or dense, mostly much shorter
than the leaves; calyx minute, acutely 4-dentate; fruit 2.5-3 cm. long, the wing
about 5 mm. broad, obtuse or rounded at the apex, decurrent to about the middle
of the seed-bearing portion.
The available Mexican material of this species is not ample enough
for satisfactory study. In some of the Guatemalan specimens the
leaflets are glabrous, in others, not otherwise distinguishable, they are
densely pubescent beneath.
Fraxinus vellerea Standl. & Styerm. Field Mus. Bot. 23: 74.
1944 (type, Standley 76162 from Jutiapa); F. bicolor Standl. & Stey-
erm. I.e. 73 (type, Standley 80408 from Guatemala).
Moist or dry rocky hillsides, 800-1,400 meters; Chiquimula; Juti-
apa; Guatemala. Mexico (Chiapas).
Small trees (or shrubs) te 10 m. or perhaps more. Leaves from densely pubes-
cent below to quite glabrous; leaflets 5-7, the lateral ones sessile, broadly ovate to
oblong-ovate, 4-8 cm. long and 2-4 cm. broad, acute or acuminate, from almost
entire to serrate along the whole margin, lower surface lighter in color than the
upper surface; fruit (from Deam 6067, not type) 2.5-3 cm. long and 0.4-0.5 cm.
broad, acute or obtuse, the seed-bearing portion about 1 cm. long.
Guatemalan material, mostly sterile, determined as F. vellerea,
F. bicolor and F. purpusii Brandegee has been placed together. This
brings together material that is vegetatively similar but from almost
glabrous to densely puberulent. The group is in need of monographic
study.
Fraxinus sp.
Guatemala: Finca Nacional La Aurora.
Leaf 40 cm. long; leaflets 9, lanceolate to ovate-lanceolate, acuminate, serrate
above the middle, glabrous or pilosulose along the veins below, petiolulate, the
petiolule about 15 mm. long, leaflets 10-15 cm. long; inflorescence paniculate,
about 20 cm. long; fruits about 15 mm. long and 3 mm. broad, narrowly elliptic,
acute, the seed up to 5 mm. long.
A single specimen from a tree on the national farm, where pos-
sibly introduced.
JASMINUM L. Jasmine
Erect or scandent shrubs; leaves usually opposite, simple, 3-foliolate, or odd-
pinnate; flowers arising from the ends of the branches or from scale buds, solitary
or usually cymose, often showy, usually fragrant, white or yellow or red outside;
calyx 4-9-dentate or lobate; corolla salverform, the tube cylindric, the limb spread-
270 FIELDIANA: BOTANY, VOLUME 24
ing, with 4-5 or more numerous lobes, these obtuse, imbricate; stamens 2, included,
the filaments short; anthers dorsifixed near the base, the connective commonly
mucronate; ovary 2-celled, the style filiform, the stigma oblong or capitate, gener-
ally bifid in age; ovules 2 in each cell or rarely 3-4, affixed laterally near the base;
fruit baccate, didymous, one of the carpels sometimes abortive, the pericarp carnose
or membranaceous; seeds usually solitary, erect; endosperm none.
About 200 species, in the Old World tropics or in subtemperate
regions. Several of them are cultivated for their showy or fragrant
flowers. The generic name is said to be of Arabic derivation.
Leaves pinnate J. grandiflorum.
Leaves simple.
Leaves glabrous or nearly so, barbate in the axils beneath, rounded at the apex;
flowers single or double J. sambac.
Leaves softly pubescent beneath, acute or acuminate; flowers not double.
J. multiflorum.
Jasminum grandiflorum L. Sp. PL ed. 2: 9. 1762. Jazmin.
Planted commonly for ornament, at middle and low elevations.
Native of India but widely cultivated in other warm regions.
A sarmentose or scandent shrub, sometimes 10 m. long, usually smaller, the
slender branchlets glabrous or nearly so; leaves pinnate, the leaflets usually 5 or 7,
elliptic-ovate to oblong-ovate, 1-5 cm. long, acuminate to obtuse, entire, sessile,
the terminal leaflet larger and short-stalked, glabrous; flowers white, fragrant,
slender-pedicellate, in 2-10-flowered terminal cymes; calyx teeth linear; corolla
limb 3-4 cm. broad, the 4-5 lobes oblong or obovate, about equaling the slender
tube.
This is perhaps best treated as a form of J. officinale L., J. offici-
nale var. grandiflorum (L.) Kobuski. In El Salvador it is called
"jazmin de parra." In Guatemala this species is particularly com-
mon in the region of Coban, where the flowers are sold in the market.
Jasminum multiflorum (Burm. f.) Andr. Bot. Repos. 8: L 4.96.
1807. Nyctanthes multiflora Burm. f. Fl. Ind. 5. L 3, f. 1. 1768.
N. pubescens Retz. Obs. Bot. 5: 9. 1789. J. pubescens Willd. Sp. PI.
1 : 37. 1797. Jasmin; jazmin de novia.
Native of India; planted commonly in the lowlands of Guatemala,
chiefly on the Pacific plains and foothills and in the North Coast,
probably also sometimes at higher elevations; grown for ornament
in most tropical regions.
Usually a shrub of 1.5-2 m., erect, the stems and leaves densely pubescent;
leaves short-petiolate, ovate or broadly ovate, 4-7 cm. long, acute or acuminate,
usually subcordate at the base; flowers clustered at the ends of the branches, almost
sessile, white; calyx lobes numerous, filiform, 1 cm. long, densely pubescent; corolla
3 cm. long, the lobes about 10, acuminate, as long as the slender tube.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 271
Jasminum sambac (L.) Ait. Hort. Kew. 1: 8. 1789. Nyctanthes
sambac L. Sp. PI. 6. 1753. Gemela; diamela; jazmin de Amelia (Pet£n).
Native of tropical Asia, now grown for ornament in most tropical
regions; planted commonly in Guatemalan gardens at low and middle
elevations; one of the common ornamental shrubs.
An erect shrub, 1-2 m. high, the stout branches sparsely or densely pubescent;
leaves almost sessile, mostly verticillate, broadly oval to oval-obovate, generally
5-7 cm. long, rounded at the apex, rounded or subcordate at the base, glabrous or
nearly so but barbate beneath in the axils of the nerves, the nerves coarse and con-
spicuous beneath, lustrous; flowers white, fragrant, densely clustered at the ends
of the branches, sessile or short-pedicellate; calyx lobes numerous, less than 5 mm.
long, pubescent; corollas simple or very "double," with numerous crowded lobes,
these often longer than the conspicuously costate tube.
Both the single and double form of this species are found in Gua-
temala. The doubled form is the more common and the corolla lobes
are so numerous and so crowded that the flowers resemble small
double roses.
LIGUSTRUM L.
Deciduous or evergreen shrubs or trees; leaves opposite, petiolate, entire;
flowers perfect, small, white, in terminal panicles; calyx campanulate, 4-dentate;
corolla salverform, the tube usually short, the 4 lobes spreading, induplicate-val-
vate; stamens 2, included or exserted; style cylindric, not exceeding the stamens;
ovary 2-celled, the cells 2-ovulate; fruit drupaceous, 1-4-seeded, black or bluish
black.
Species about 50, mostly in eastern Asia. Several are cultivated
for ornament in temperate and tropical America.
Leaves coriaceous, usually acuminate and 7-13 cm. long; tree L. lucidum.
Leaves membranaceous, obtuse or rounded at the apex, commonly 3-4 cm. long;
shrub L. vulgare.
Ligustrum lucidum Ait. Hort. Kew. ed. 2. 1: 19. 1810. Trueno.
Native of eastern Asia; planted commonly in Guatemala from
sea level up to 2,500 m. or perhaps even higher.
A large shrub or tree, sometimes 12 m. high or more, with a rather short and
thick trunk, the crown broad and rounded, very dense, the lower branches often
somewhat pendent, glabrous or nearly so; leaves on stout petioles, coriaceous,
broadly ovate to ovate-oblong, acuminate, rounded at the base, the margins and
costa often reddish, the lateral nerves indistinct, 4-5 pairs; flowers creamy white,
scarcely 4 mm. long, forming dense panicles 6-15 cm. long, short-pedicellate; tube
of the corolla longer than the calyx; stamens exserted; fruit bluish black, oval or
subglobose, 6-8 mm. long.
This is one of the commonest street trees of Guatemalan cities,
and is admirably suited to that purpose. It withstands neglect and
272
FIELDIANA: BOTANY, VOLUME 24
FIG. 74. Ligustrum lucidum, a cultivated Asian plant. A, habit, X K; B, co-
rolla, X 5; C, calyx and style, X 5; D, fruits, X 2.
ill treatment, also the long dry season, and when even fairly well
grown is a handsome tree, with dense, deep green foliage. Although
it grows well at lower elevations, it is seen at its best in such places
as Guatemala, Chimaltenango, Quezaltenango and San Marcos, where
there are many long avenues. The finest trees of the country are
probably those in the northwestern part of the city of Quezaltenango
in the suburb known as Garibaldi.
Ligustrum vulgare L. Sp. PI. 7. 1753. Jupiter bianco ; jazmin
de Persia; privet.
Native of Europe and northern Africa; cultivated as a hedge plant
in many temperate and subtropical regions; planted occasionally for
ornament or hedges in Guatemala, especially about Coban, but not
common.
Usually a densely branched shrub of 1-2 m. but often larger, the branchlets and
panicles minutely puberulent; leaves short-petiolate, oblong-lanceolate to ovate,
mostly obtuse, glabrous; flowers white or greenish white, in rather dense, narrow
panicles 3-6 cm. long; anthers short-exserted ; fruit subglobose or ovoid, 6-8 mm.
long, black, lustrous.
In the United States this is the shrub much grown as a hedge
plant, at least in cities. If allowed to grow untrimmed, it becomes
a rather straggling bush, but if trimmed closely, it makes a dense
and handsome hedge, shedding its leaves in winter.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 273
LINOCIERA Swartz
Large shrubs or trees; leaves opposite, entire, more or less coriaceous; inflores-
cence thyrsoid (ours), umbellate, capitulate, cymulose, racemose-paniculate or
racemose-fasciculate, lateral and terminal; calyx small, 4-fid or 4-dentate; petals 4,
free or nearly so, linear or oblong, induplicate-valvate; stamens normally 2, affixed
to the bases of the sepals, the filaments short; anthers ovate to linear, the con-
nective apiculate or naked; ovary 2-celled, the style short, the stigma oblong,
clavate, or subglobose, entire or emarginate; ovules 2 in each cell, laterally affixed
near the apex, pendulous; fruit drupaceous, ovoid to oblong or subglobose, the
endocarp usually hard and osseous; seeds usually solitary, pendulous, the testa
thin or thick; endosperm carnose, subcartilaginous, or none; cotyledons flat or
thick-carnose, the radicle short, superior.
Species 50 or more, in the tropics of both hemispheres. One other
Central American species is known, in Costa Rica and Panama.
Petals rather broadly linear; petioles mostly 1.5-3 cm. long L. domingensis.
Petals filiform; petioles mostly 5 mm. long or shorter L. oblanceolata.
Linociera domingensis (Lam.) Knobl. Bot. Centralbl. 61: 87.
1895. Chionanthus domingensis Lam. Tabl. Encycl. 1: 30. 1791.
Moist or wet, mixed forest, 1,500 m. or less; Izabal; Quezalte-
nango. British Honduras; West Indies.
A tree of 9-12 m., glabrous except in the inflorescence, the trunk sometimes
70 cm. in diameter, the bark smooth, gray; leaves elliptic-lanceolate, chartaceous
or subcoriaceous, on long slender petioles, 16 cm. long and 6 cm. broad or smaller,
acuminate, gradually attenuate into the petiole, usually with barbate pits in the
axils below; panicles small or large, terminal and axillary, glabrous or sparsely
pubescent, sometimes equaling the leaves, many-flowered, the pedicels 2 mm. long
or shorter; calyx glabrous, scarcely 2 mm. long; petals white or pink, flat, 15-
20 mm. long, 1.5 mm. broad, rounded or very obtuse at the apex; drupes oval or
ellipsoid, 1.5-2 cm. long.
Linociera oblanceolata Robinson, Proc. Am. Acad. 49: 504.
1913. Cachicha macho.
Upland forest, 700 m. or less; Peten. Mexico (Tabasco); British
Honduras (type from upper Moho River, M. E. Peck 719}.
A large shrub or tree, sometimes 17 meters high with a trunk 25 cm. in diam-
eter, glabrous almost throughout; leaves subcoriaceous, on petioles mostly 3-4 mm.
long, oblanceolate to obovate-elliptic, 10-17 cm. long, 3-7 cm. broad, narrowly
long-acuminate or merely abruptly short-acute, usually attenuate to the base and
there abruptly and narrowly obtuse, not decurrent, usually barbate beneath in the
axils of the nerves; panicles lax, arising in the axils of the upper leaves, 6-10 cm.
long, many-flowered, the branches almost filiform, puberulent, the pedicels 3-5 mm.
long; calyx puberulent, the lobes ovate, acuminate, 1.2 mm. long; petals white or
greenish, 7-10 mm. long, filiform, involute; connective of the anther not produced
at the apex; fruit broadly ellipsoid, 1.5-2 cm. long, bluish black.
274
FIELDIANA: BOTANY, VOLUME 24
FIG. 75. Linociera oblanceolata. A, branch showing leaves and inflorescence,
X 3^; B, flower, X 8; C, stamens and pistil, X 22.
In general appearance this is much like L. caribaea (Jacq.) Knobl.,
with which it was compared by Robinson, but it seems to differ con-
stantly in its muticous rather than appendaged anther connective
and in having shorter petiolate, less coriaceous leaves.
OSMANTHUS Loureiro
Trees or shrubs. Leaves opposite, entire (ours) or dentate; inflorescence in
congested racemes, thyrses or fascicles, terminal or lateral (ours); calyx short,
4-dentate or divided; corolla 4-lobate, campanulate; the lobes imbricate; stamens 2,
rarely 4; anthers subextrorse; ovary 2-celled, the style short, stigma entire; ovules 2
in each cell; fruit an ovoid or globose drupe, the exocarp fleshy, the endocarp hard
and bone-like.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 275
A small genus of not more than ten species in both hemispheres,
two others in America in temperate regions; one of these, Osmanthus
americana (L.) B. & H., extending to northern Mexico, is similar
to ours.
Osmanthus mexicana Lundell, Phytologia 1 : 308. 1939.
Native of Chiapas, Mexico (type, Matuda 2023} not far from
Guatemala and to be expected.
Small trees 6-7 m. tall. Leaves narrowly lanceolate or oblanceolate, acuminate*
decurrent on the petioles, coriaceous, glabrous, 4-9 cm. long and 1-2.5 cm. broad>
petioles 1-2 cm. long; inflorescences short, axillary, few-flowered thyrse; sepals 4>
about 1 mm. long, the lobes ciliolate, triangular, obtuse; corolla subcampanulate>
about 4 mm. long, the lobes ovate, obtuse, glabrous or obscurely ciliolate; stamens
2, attached at about the middle of the corolla; stigma capitate; fruit ellipsoid,
about 13 mm. long and 8 mm. in diameter.
Dr. Lundell thought the flowers to be unisexual but those on our
specimen seem to be perfect.
The common lilac, Syringa vulgaris L., has been tested in Guate-
mala but, like some other shrubs of temperate regions, will not grow
well here, or elsewhere in the tropics. Senorita Diaz of Coban told
the senior author that many years ago she asked a friend to obtain
seed for her, wishing a small packet for trial purposes. He sent her
from France five pounds of lilac seed, with a rather fantastic bill.
The seeds germinated well and many young plants were raised and
distributed widely in the Coban region. They never reached a height
of more than a meter, then would die down to the roots. During the
winter they shed their leaves, and the gardeners, thinking the plants
were dead, would dig them out of the ground, so that within a few
years all were gone, and none of them ever flowered.
LOGANIACEAE. Logania Family
DOROTHY N. GIBSON
References: Endlicher, Gen. PL 574. 1840. A. DC. Prodr. 9: 1-37.
1845; 10: 432. 1846. Bentham & Hooker, Gen. PI. 2: 786. 1876.
Solereder in Engler & Prantl, Pflanzenf. IV (2) : 19-30. 1892. Klett,
Umlang und Inhalt der Familie der Loganiaceen. Bot. Arch. 5:
312-338. 1924. Raymond J. Moore, Cytotaxonomic Studies in the
Loganiaceae, I-III. Journ. Bot. 34: 527-538. 1947; 35: 404-410.
1948; 36: 511-516. 1949.
Herbs, shrubs, vines and lianas or trees; leaves opposite or rarely verticillate,
simple, entire or dentate, connected by a transverse line or stipular sheath ; inflores-
cences terminal or axillary, normally 2-3 times dichotomous and cymose, some-
times paniculate; flowers regular or nearly so, bisexual, usually bracteate; calyx
4-5 parted, usually short, the segments imbricate; corolla gamopetalous, variously
colored, funnelform, salverform, rarely campanulate or rotate, the lobes valvate,
imbricate, or contorted; stamens as many as the corolla lobes and alternate with
them, inserted in the throat or tube of the corolla, the filaments usually short;
anthers introrse, dorsifixed, the 2 cells distinct and parallel, dehiscing longitudi-
nally; ovary superior (rarely half -inferior), usually 2-celled (rarely 1, 3 or 5); style
simple or bifid; ovules usually many (rarely 1), amphitropous or anatropous, pla-
centation axile; fruit usually capsular and septicidally bivalvate, sometimes bac-
cate or drupaceous and indehiscent; seeds variable in form, sometimes winged;
endosperm carnose or cartilaginous, usually copious; embryo small, usually straight,
rarely incurved.
The Loganiaceae are well represented in the tropical and warm
temperate regions of both hemispheres. Seven of the 32 genera occur
in Guatemala; an eighth, Potalia, is known from Costa Rica.
An interesting discussion of the obviously close relationship be-
tween this family and the Rubiaceae is P. Jovet's "Aux confins de
Rubiace"es et des Loganiace"es" in Not. Syst. 10: 39-53. 1941.
Herbs or subshrubs.
Leaves broad; flowers 5-merous; corolla lobes valvate.
Corolla conspicuous, tubular-campanulate to funnelform; style 1; capsule
circumcissile above the persistent cupular base Spigelia.
Corolla inconspicuous, urceolate; styles 2, connate at summit with a common
stigma but divergent after anthesis; capsule 2-horned, dehiscent along
inner margin of each Cynodonum.
276
GIBSON: FLORA OF GUATEMALA 277
Leaves linear; flowers 4-merous; corolla lobes imbricate Polypremum.
Shrubs, vines, lianas or trees.
Stigmas entire, or nearly so.
Shrubs or trees, never vining; leaves pinnate-nerved with stellate indumen-
tum; fruits capsular, less than 1 cm. in diameter Buddleia.
Vines or lianas developing from erect shrubs; leaves 3-5 plinerved, indumen-
tum (if present) never stellate; fruits baccate, globose, at least 4 cm. in
diameter Strychnos.
Stigmas 4 (style bifid, each branch bilobate).
Leaves lanceolate to ovate, acuminate; corolla yellow; capsule oblong-ovate,
2.5 cm. long or less, beaked; seeds not linear, flat, apically winged.
Gelsemium.
Leaves elliptic-oblong or oval-oblong, obtuse; corolla purplish-magenta; cap-
sule linear, 5-9 cm. long, not beaked; seeds linear, bearing tuft of long
hairs at apex Plocosperma.
BUDDLEIA L.
References: Linnaeus, Sp. PL 112. 1753; Linnaeus, Gen. PI. ed. 5.
51. 1754. Eliane M. Norman, The genus Buddleia in North America,
Gentes Herb. 10(1): 1-116. [1966] 1967.
Shrubs or trees to 30 m. tall, the bark usually furrowed, the young branches
usually tomentose; leaves opposite, decussate, petiolate or sessile, with stipules
often reduced to a line, sometimes foliaceous, the blades membranaceous to cori-
aceous, stellate-pubescent, often glandular, rarely glabrous, lanceolate, elliptic or
ovate, serrate, serrulate, crenate, dentate, or entire; inflorescences paniculate, sub-
tended by small leaves, bracts or bracteoles; flowers in dichasial capitate clusters,
usually fragrant; calyx 4-parted, the lobes usually shorter than the tube; corolla
campanulate, funnelform, or salverform, usually stellate-tomentose outside, the
inner surface more or less pubescent with long, unicellular, pitted hairs, the lobes
usually smaller than the tube; stamens 4, sessile or with short filaments inserted
below the throat of the tube, directly below the sinuses; style short, the clavate to
to clavellate stigma obscurely to clearly bilobate; ovary superior, usually ovoid
but may be short-cylindrical or subglobose, bilocular, with numerous ovules multi-
seriate on the placentae; mature capsule small (3-6 mm. long in our species), par-
tially puberulent to tomentose, dehiscing septicidally and loculicidally (rarely in-
dehiscent); seeds numerous, oblong or ovoid, yellow or brown, the testa often
extended into wings; endosperm carnose, the embryo usually small and straight.
America, Africa, and Asia.
Corolla lobes imbricate, with scattered hairs on inner surface and in throat of tube.
Flowers borne in capitate clusters 1.2-2 cm. in diameter B. megalocephala.
Flowers borne in capitate clusters less than 1.2 cm. in diameter.
Flowers 10-15 in each cluster; corollas usually less than 3 mm. long.
B. skutchii.
Flowers 2-8 in each cluster; corollas 3-5 mm. long.
Stigma clearly divided, the lobes incurved B. euryphylla.
Stigma undivided, clavellate.
Leaves lanceolate or ovate, average width 4-13 cm.; inflorescence large,
averaging 20 cm. long, 18 cm. across; capsule glabrescent. .B. cordata.
278 FIELDIANA: BOTANY, VOLUME 24
Leaves oblong-lanceolate, average width 0.5-3 cm.; inflorescence small,
averaging 8 cm. long, 7 cm. across; capsule tomentulose. . .B. nitida.
Corolla lobes valvate, with crescent-shaped line of hairs on inner surface.
Upper surface of leaves glabrate to stellate-pubescent, with loose stellate tomen-
tum on lower surface; leaf bases attenuate, acute or obtuse. .B. americana.
Upper surface of leaves usually stellate-tomentose, with very thick, floccose
tomentum on lower surface; leaf bases truncate, cuneate, subcordate, auric-
ulate or amplexicaul B. crotonoides.
Buddleia americana L. Sp. PL 112. 1753 (as Buddleja) . B. oc-
cidentalisL. Sp. PL ed. 2. 162. 1762. B. spicata R. & P. Prodr. 1: 53,
t. 81. 1798. B. callicarpoides HBK. Nova Gen. & Sp. 2: ed. qu. 350,
ed. fol. 282. 1818. B. dentata HBK. I.e., ed. qu. 352, ed. fol. 283.
B. floribunda HBK. I.e. B. verbascifolia HBK. I.e., ed. qu. 351, ed.
fol. 283. B. cana Willd. ex J. A. & J. H. Schultes, Mant. 3: 94. 1827,
in syn. B. rufescens Willd. I.e. 97. B. americana albiflora Gomez,
Anal. Hist. Nat. Madrid 19: 259. 1890. B. americana var. Roth-
schulii Loes. Bot. Jahrb. 23: 118, 129. 1896. Arnica (Huehuetenan-
go); Sactzam (Alta Verapaz); Salvia (Jutiapa and Sacatepequez) ;
Salva santa (Guatemala and Izabal).
Usually in dry or damp thickets, sometimes in waste ground or
in oak forests, 80-2,100 m.; Alta Verapaz; Baja Verapaz; Chimalte-
nango; Escuintla; Guatemala; Huehuetenango; Izabal; Jalapa; Juti-
apa; Pete"n; Quiche"; Sacatepequez; San Marcos; Santa Rosa; Zacapa.
Southern Mexico, Central and South America to Bolivia; West Indies.
Shrubs or small trees 2-5 m. tall (rarely 10 m.), the young branches tomentose;
leaves subsessile or with petioles to 2 cm. long, blades membranaceous, serrate or
entire, upper surface glabrate or stellate-pubescent, lower surface with loose stel-
late-tomentum underlain by glandular trichomes, narrowly lanceolate, elliptic,
lance-ovate, or ovate, averaging 10-15 cm. in length (4-26 cm.), usually 5-8 cm.
broad (2-13 cm.), acuminate, base often decurrent but may be attenuate, acute, or
obtuse; inflorescences 8-22 cm. long, the flowering clusters lowest on the branches
usually short-pedunculate, the remaining ones sessile, fragrant; calyx 1.5-2 mm.
long, tubular with lanceolate, acuminate lobes, stellate-tomentose outside; corolla
4-5 mm. long, funnelform, the lobes about equalling the tube, yellow inside, whitish
outside, stellate-tomentose outside, the inner surface of lobes with a crescent-
shaped line or tuft of pitted hairs; stamens inserted at sinuses or just below; ovary
ovoid, 1-1.5 mm. long, tomentose on upper half, style short, the clavate stigma
obscurely bilobed; mature capsule short-cylindrical to ovoid, 3.5-5 mm. long, septi-
cidally dehiscent for half its length, loculicidally usually only at apex; seeds numer-
ous, oblong, 0.8-1 mm. long, the testa reticulate, extending into short wings.
This is the most abundant and variable species in Central Amer-
ica and is often weedy.
Buddleia cordata HBK. Nova Gen. & Sp. 2: ed. qu. 348, ed.
fol. 280, t. 185. 1818 (as Buddleja). B. acuminata HBK. I.e., ed. qu.
GIBSON: FLORA OF GUATEMALA 279
349, ed. fol. 281, t. 187, not B. acuminata Poiret. B. Humboldtiana
J. A. & J. H. Schultes, Mant. 3: 93. 1827. B. decurrens Schlecht. &
Cham. Linnaea 5: 105. 1830. B. floccosa Kunth, Ind. Sem. Hort.
Berol. 1844 and in Linnaea 18: 500. 1844. B. macrophylla Kunth,
I.e. B. ovalifolia Kunth, I.e. B. propinqua Kunth, Ind. Sem. Hort.
Berol. 1844 and in Linnaea 18: 501. 1844. B. spectabilis Kunth &
Bouche", Ind. Sem. Hort. Berol. 1845: 11. 1845, and in Ann. Sci. Nat.
ser. 3, Bot. 5: 358. 1846. B. cordata var. teposan Loes. Verh. Bot.
ver. Brand. 53: 73. 1911. B. floccosa var. crassifolia Loes. I.e. 72.
1944. B. astralis Standl. & Steyerm. Field Mus. Bot. 23: 72. 1944.
B. cordata subsp. cordata Norman, Gent. Herb. 10: 67. 1967.
The only Guatemalan collection seen is from a brushy field, south-
ern slopes of Volcan de Tajumulco, San Marcos, 1,400-1,700 m.,
Steyermark 37276, and is the type of B. astralis Standl. & Steyerm.
In Mexico, it grows on rocky ledges and barrancas in oak and pine
forests, 1,500-3,000 m.
Shrubs or trees, 2-20 m. tall (a tree of 12 m. in Guatemala), the branches
densely tomentose; leaf blades usually 4-23 cm. long with petioles usually 1-4 cm.
long (ours is 19-27 cm. long, 9-11 cm. wide, with petioles 5-7 cm. long), ovate to
lanceolate, acuminate, acute at base, entire in ours but may be serrulate, green
above and almost glabrous, covered beneath with closely appressed tomentum and
with lax, floccose candelabra hairs, prominently veined; inflorescences terminal,
paniculate, 6-30 cm. long (ca. 25 cm. in ours), the heads few-flowered and lax;
calyx tubular, puberulent to stellate tomentose, ca. 2 mm. long, the lobes broadly
triangular, obtuse; corolla yellow, campanulate, twice as long as the calyx, stellate-
tomentulose outside, with pitted hairs on inner surface of lobes sometimes extend-
ing down into upper part of tube; stamens subsessile or with short filaments, in-
serted near throat; ovary ovoid, 1-1.8 mm. long, tomentulose on upper part, the
style to 1.4 mm. long, the clavellate stigma obscurely bilobate; mature capsule
short-cylindrical, 3.5-5 mm. long, septicidally dehiscent for half its length, locu-
licidally only at apex, glabrescent (fide Norman); seeds numerous, 1.2-2 mm. long,
the reticulate testa extended into prominent wings.
Buddleia crotonoides A. Gray, Proc. Am. Acad. Sci. 5: 165.
1861. B. tuxtlica Loes. Verh. Bot. Ver. Brand. 53: 73. 1911. B. pur-
pusii Standley, Journ. Wash. Acad. Sci. 16: 15. 1926. B. stenoptera
Standl. & Steyerm. Field Mus. Bot. 23: 70. 1944. B. amplexicaulis
Standl. & Steyerm. I.e. 71. B. crotonoides subsp. amplexicaulis Nor-
man, Gentes Herb. 10: 87-89. 1966. Salvia (Huehuetenango and
Quezaltenango).
Usually on rocky slopes in pine-oak forests, sometimes in wet or
dry thickets, 1,200-2,500 m.; Baja Verapaz; Chimaltenango; Guate-
mala; Huehuetenango; Jalapa; Quezaltenango; Quiche"; Sacatape"-
280 FIELDIANA: BOTANY, VOLUME 24
quez; San Marcos; Totonicapan. Mexico (where it sometimes grows
near sea level); Honduras; Costa Rica.
Shrubs or small trees 2-5 m. tall, the young branches densely covered with lax
tomentum; leaves short-petiolate or sessile or amplexicaul, the blades lanceolate,
elliptic-ovate or broadly ovate, closely dentate or serrate, 4.5-20 cm. long, 1-8 cm.
broad, usually stellate-tomentose above (rarely puberulent), densely covered be-
neath with floccose tomentum underlain by glandular trichomes, acute or acumi-
nate at apex, base subcordate, truncate, cuneate, or broadly spatulate to auriculate
and clasping the stem; inflorescences 6-18 cm. long, the flower clusters sessile,
0.5-1.2 cm. in diameter, usually interrupted but occasionally continuous along the
branches, the first pair of heads on all except the lowermost lateral branches borne
close to the main axis; calyx tubular, stellate-tomentose outside, 2-3.5 mm. long,
the lobes lanceolate or subobtuse; corolla greenish-white to pale yellow, 3-5 mm.
long, funnelform, stellate-tomentose outside, the lobes acute or obtuse, with pitted
hairs in tufts, often in a lunate line inside the lobes; stamens inserted near sinuses;
ovary ovoid or short-cylindrical, 1-1.5 mm. long, usually densely tomentose (some-
times woolly, rarely glabrate) on upper part, style short, with clavellate stigma;
mature capsule ovoid, 2.5-3.5 mm. long, opening septicidally for half its length,
loculicidally at apex; seeds numerous, to 0.7 mm. long, the reticulate testa extended
into short wings.
The collections with obviously amplexicaul and/or auriculate
leaves cited by Norman as B. crotonoides subsp. amplexicaulis were
carefully studied and compared with two recent collections not seen
by her (Breedlove 8570 from Huehuetenango and Raven & Breedlove
200^1 from Chiapas) before it was decided to place these plants with
variant leaf form in synonymy. The Chiapas plant, on which most
of the leaves are auriculate and amplexicaul, possesses one pair of
leaves with petioles 0.5 cm. long. The Guatemalan plant, on which
many leaves are merely sessile with attenuate or cuneate bases, pos-
sesses three pairs of leaves with broadly spatulate bases, two of which
have very small auricles and are definitely clasping. Therefore, as
only one character differs and it appears to be somewhat unstable,
neither subspecific nor varietal rank seems justified.
Two additional variant specimens were observed, both from Ja-
lapa: Steyermark 32083, with leaf blades long-attenuate and 9-15
lateral veins (other specimens of B. crotonoides average only 5-9 lat-
eral veins), and Standley 7677^., with only 5-6 flowers in each cymule
(other specimens average 12-20 flowers).
Buddleia euryphylla Standl. & Steyerm. Field Mus. Bot. 23:
223. 1947.
Forested ravines or hillsides, 2,000-3,000 m.; known only from
the type locality, Sierra de las Minas, on Montana Piamonte, El
Progreso.
GIBSON: FLORA OF GUATEMALA 281
Trees 7-15 m. tall, branches stout, obtusely tetragonous, densely stellate-
tomentose, the tomentum brownish, lax; leaves membranaceous, large, the blades
entire, broadly ovate to ovate-elliptic, 15-26 cm. long, 8-15 cm. broad, on stout
petioles 3-7 cm. long, subacute to short-acuminate, base obtuse or acute, upper
surface glabrous or nearly so at maturity, lower surface densely tomentose with
both appressed and floccose stellate hairs, prominently veined; inflorescences panic-
ulate, ca. 30 cm. long and as broad, much branched, the flowers short-pedunculate
in lax cymules, fragrant; calyx 2-3 mm. long, densely stellate-tomentose, the lobes
broadly triangular, obtuse; corolla golden yellow, 3-4 mm. long, the lobes rounded,
densely stellate-tomentose outside with pitted hairs within on lower part of lobes,
extending into upper tube; stamens inserted ca. 0.5 mm. below sinuses; ovary
ovoid, 1.5 mm. long, tomentulose, style to 1.3 mm. long with stigma clearly divided,
the 2 lobes arching toward each other; mature capsule not seen, immature seeds
numerous, winged.
Buddleia megalocephala Donn.-Sm. Bot. Gaz. 23: 10. 1897.
B. megalocephala f. albilantha Moldenke, Phytologia 2: 363. 1947.
B. hypsophila I. M. Johnston, Journ. Arn. Arb. 19: 127. 1938. Salvia,
Patushe (Quezaltenango) .
High mountain forests, often in association with Pinus, Abies,
Cupressus or Juniperus, sometimes forming small, dense stands, 2,400-
4,050 m.; Chimaltenango; Huehuetenango; Quezaltenango; San Mar-
cos; Solola; Totonicapan. Mexico.
Trees to 12 m. high, usually with a thick trunk, sparsely branched, the stout
branches covered with dense tomentum; leaves subcoriaceous, lanceolate or elon-
gate-lanceolate, entire, acute to acuminate, obtuse at base, petioles 1-2 cm. long,
the blades 7-20 cm. long, 2-5 cm. wide, lustrous, green and glabrate above, lower
surface covered with dense, whitish to brown stellate tomentum; inflorescences
terminal, 6-30 cm. long, the heads very dense, globose, 1.5-2 cm. in diameter at
maturity, long-pedunculate, usually in short, simple racemes, or the racemes
branched at the base; calyx tubular, densely tomentose, 3-4.5 mm. long, the lobes
acuminate or narrowly triangular, ca. 2 mm. long; corolla 6-8 mm. long, deep
orange inside, paler outside, funnelform, stellate tomentose outside, with scattered,
pitted hairs inside on upper part of tube, fragrant; stamens inserted ca. 1 mm.
below sinuses; ovary ovoid, 1.5-2 mm. long, tomentulose, style 2-3 mm. long,
stigma clavate; mature capsule ovoid, glabrous or stellate-puberulent, 4.5-6 mm.
long, primarily septicidally dehiscent; seeds numerous, to 2 mm. long, the reticu-
late testa extended into wings.
I believe that Johnston's B. hypsophila is a hybrid between B.
megalocephala and B. nitida. It differs from those specimens cited
by Norman as possible hybrids (Standley 84407 and Breedlove 8489)
only in having more leaves with an acute apex. Breedlove 8489 has
several leaves with acute apices rather than acuminate. The type
of B. hypsophila, Skutch 843, has the same short, broadly triangular
282
FIELDIANA: BOTANY, VOLUME 24
FIG. 76. Buddleia megalocephala. A, branch with inflorescence, X 1A', B, fruit-
ing "head," X 1%.
calyx lobes as those of the supposed hybrids, and a few specimens
which are obviously B. megalocephala exhibit thinner, appressed in-
dument and smaller leaves than typical.
Buddleia nitida Bentham in DC. Prodr. 10: 437. 1846. B. alpina
Oersted, Vid. Medd. Nat. For. Kjoebenh. 5: 25-26. 1853. Sacumis
(Huehuetenango) .
GIBSON: FLORA OF GUATEMALA 283
On open or brushy, limestone, sand, or lava slopes and para-
mos, frequently in oak, oak-pine, or Cupressus forests, sometimes in
Baccharis forests, 2,000-4,000 m.; Chimaltenango; El Progreso; Gua-
temala; Huehuetenango; Jalapa; Quezaltenango; Quiche"; Sacatepe*-
quez; San Marcos; Solola; Totonicapan. Southern Mexico to Panama.
Shrubs or trees usually 4-10 m. high (rarely to 15 m.), usually densely branched,
the young branches covered with dense, closely appressed, white or brownish to-
rn en turn, becoming glabrescent; leaves subcoriaceous, deep green and glabrate
above (sometimes with scattered stellate hairs), covered beneath with dense,
closely appressed, usually silvery-white but may be brownish, stellate tomentum,
oblong-lanceolate or elongate-lanceolate, the blades usually 5-7 cm. long, 1.5-
2 cm. broad (rarely to 10 cm. long, 3 cm. broad, or in the dwarfed form1 only ca.
1 cm. long, 0.4 cm. broad), acute to long acuminate, base acute or obtuse, entire
(rarely serrulate), petioles 0.5-3 cm. long; inflorescences terminal, paniculate, usu-
ally 7-8 cm. long, 5-7 cm. broad (rarely 3-14 cm. long, 3-10 cm. broad), the flowers
in short pedunculate cymules, fragrant; calyx tubular-campanulate, 1.5-2.4 mm.
long, stellate-tomentose outside, the lobes short, broadly triangular; corolla yellow
to orange, 3.7-5 mm. long, campanulate-funnelform, stellate-tomentose on outside
of lobes and upper part of tube, with pitted hairs in throat and on lower part of
lobes; stamens inserted just below the sinuses; ovary subglobose, 1-1.5 mm. long,
tomentulose on upper part, style 1-1.5 mm. long, the clavellate stigma obscurely
bilobate; mature capsule oblong or ellipsoid, 4-5 mm. long, tomentulose, septi-
cidally dehiscent for most of its length, loculicidally only at apex; seeds numerous,
to 1.7 mm. long, the reticulate testa extended into prominent wings.
Buddleia parviflora HBK of Mexico is much like B. nitida but
the flowers of B. parviflora are distinctly smaller. As noted by Nor-
man, there is evidence that B. nitida crosses with B. skutchii Morton
and B. megalocephala Donn.-Sm.
Buddleia skutchii Morton, Phytologia 1: 148. 1935. B. matu-
dae Standl. Field Mus. Bot. 22: 96. 1940. Saclajac (Huehuetenango),
Salvia, Salvia blanca, Flor de Santa Maria (Quezaltenango).
Usually in pine-oak or Cupressus forests, sometimes in open fields,
hedgerows, or on brushy slopes, 1,600-3,800 m.; Chimaltenango;
Guatemala; Huehuetenango; Quezaltenango; Quiche"; San Marcos;
Totonicapan. Chiapas, Mexico.
Trees, 6-16 m. tall, the young branches densely covered with whitish stellate
tomentum; leaves subcoriaceous, usually entire (rarely serrulate), lanceolate or
ovate-lanceolate, petioles 2-3.5 cm. long, the blades 6-20 cm. long, 2-7 cm. broad,
1 There is now no doubt that certain dwarfed and usually sterile specimens
from Guatemala and Costa Rica with leaves ca. 1 cm. long, are indeed B. nitida.
Fertile specimens (J. R. Johnston 1 725, Standley 85252, and Sharp 254.06) check in
all other characters and a few branches bear larger leaves, to 3.8 cm. The dwarfing
is thought to be due to browsing.
284 FIELDIANA: BOTANY, VOLUME 24
usually acuminate, sometimes acute, base obtuse or cuneate, green and glabrate
above, covered beneath with whitish to brownish tomentum, the outer layer of
floccose candelabra hairs; inflorescences terminal, paniculate, ca. 8-15 cm. long,
8-20 cm. broad, the flowers 10-15 in each small, dense, short-pedunculate cymule,
fragrant; calyx tubular-campanulate, 1.5-2.5 mm. long, tomentulose, the lobes
broadly triangular; corolla yellow to orange, 2-3 mm. long, campanulate, the lobes
spreading, stellate tomentose outside, with pitted hairs sparsely scattered inside
on lobes and in tube; stamens inserted near the sinuses; ovary ovoid, 0.5-1 mm.
long, tomentulose at least on upper part, style 0.5-0.7 mm. long, with clavellate
stigma; mature capsule ovoid, 3-4 mm. long, tomentulose or glabrescent, septi-
cidally dehiscent for half its length, loculicidally at apex; seeds numerous, to 1.8
mm. long, the reticulate testa extended into prominent wings.
CYNOCTONUM J. F. Gmel.
Reference: J. F. Gmelin, Syst. Nat. 443. 1791.
Erect annual or perennial herbs; leaves opposite, entire, minutely ciliate, lan-
ceolate to ovate, membranaceous, the petioles connected by a narrow membrane
or by small stipules; inflorescences terminal or axillary in pedunculate, dichotomous
cymes, the flowers secund along the branches, almost sessile, usually bracteate;
calyx 5-parted, the lobes lanceolate; corolla urceolate, exceeding the calyx, the
tube ventricose, contracted at the throat, the 5 lobes short, valvate; stamens 5,
included, inserted on the corolla tube, filaments short, anthers ovate, cordate at
base, the cells parallel; styles 2, short, connate below the small capitate pilose
stigma, divergent after an thesis; ovary superior, bilocular, broad at the apex;
ovules numerous, peltately affixed; capsule broad, compressed contrary to the sep-
tum, truncate or bilobate at apex, the carpels divergent above, erect or incurved,
dehiscent along inner margin; seeds subglobose or compressed, tuberculate-rugose;
endosperm carnose, embryo linear.
A genus of chiefly tropical regions, it ranges from the southern
United States through Mexico, the West Indies, Central and South
America, and in the East Indies, India, northern Australia and Mada-
gascar. Two of the three species occur in Guatemala.
The genus very closely resembles Ophiorrhiza of the Rubiaceae,
from which it differs only by the superior ovary, and there is certainly
justification for confusion of the two in the past.
Lobes of capsule widely divergent; calyx lobes 1-nerved; corolla lobes short, a
quarter to a third the length of corolla; throat of corolla with a ring of minute
hairs C. mitreola.
Lobes of capsule curved toward each other; calyx lobes 3-nerved, corolla lobes
elongate, one-half the length of corolla; sinuses with minute fringe of hairs.
C. petiolatum.
Cynoctonum mitreola (L.) Britton, Mem. Torr. Bot. Club 5:
258. 1894. Ophiorriza Mitreola L. Sp. PI. 150. 1753. C. mitreola
vars. intermedia and orthocarpa Hochr. Bull. N. Y. Bot. Gard. 6(21):
284. 1910. C. oldenlandioides (A. DC.) Robinson, Proc. Am. Acad.
GIBSON: FLORA OF GUATEMALA
285
FIG. 77. Cynoctonum petiolatum. A, habit, fruiting plant, J^ natural size; B,
capsule, after Bentham in Hooker, Icon. 9: t. 828. 1852; C, corolla dissected to
show stamens and pistil, after Bentham, I.e.
45: 396. 1910. C. pedicellatum (Benth.) Robinson, I.e. Mitreola olden-
landioides A. DC. Prodr. 9: 9. 1845; Hooker, Icon. 9: t. 827. 1852.
M. pedicellata Bentham, Journ. Linn. Soc. 1: 91. 1857. Altanecia
(Pet&i, fide Lundell).
In damp thickets and along streams and lakes, 150-1,700 m., Chi-
quimula; Huehuetenango; Pete"n; Santa Rosa. Southeastern United
286 FIELDIANA: BOTANY, VOLUME 24
States, Mexico to Panama; East and West Indies; northwestern
South America; northern Australia; Madagascar; tropical Africa.
A slender, erect annual, 15-75 cm. tall, sparsely branched, the stems usually
pale, glabrous; leaves almost sessile or on slender petioles 1-2 cm. long, appearing
entire but actually very minutely ciliate, lanceolate to ovate, 1.5-8 cm. long, acute
or acuminate, rounded at the bases and decurrent, forming narrow wings on the
petioles, almost glabrous, paler beneath; cymes long-pedunculate, terminal and
arising in the upper leaf axils, few-many flowered; calyx lobes 1-nerved; corolla
white, 1-2 mm. long, ca. 1.5 mm. broad, sessile or nearly so along one side of the
slender, elongate branches, lobes short, \i - y$ corolla length, the tube of the
corolla with a ring of minute hairs at the throat; ovary glabrous or at least the
upper half minutely puberulous or tuberculate; capsule 2-3 mm. long, the lobes
mitre-shaped, separate, widely divergent or upright and spreading outward.
Cynoctonum petiolatum J. F. Gmel. Syst. Nat. 2: 443. 1791.
C. mitreola var. campylocarpa Hochr. Bull. N. Y. Bot. Gard. 6(21):
284. 1910. C. paniculatum (A. DC.) Robinson, Proc. Am. Acad. Sci.
45: 396. 1910. Mitreola petiolata T. & G. Fl. N. Amer. 2: 45. 1841.
M. paniculata A. DC. Prodr. 9: 9. 1845; Hooker, Icon. 9: t. 828. 1852.
M. paniculata var. glabra Hoehne, Comm. Linh. Telegr. Estrat.
Matto Grosso, Annexe 5, Bot. pt. 6: 68. 1915.
In damp thickets, along stream banks, 220-1,000 m., Santa Rosa.
Mexico; Honduras; West Indies; South America (Brazil, Colombia,
Ecuador and Peru) ; India.
Differs from C. mitreola by the 3-nerved calyx lobes, slightly
smaller corolla, 1-1.5 mm. long, ca. 0.5 mm. broad, elongate corolla
lobes about equalling the tube, a minute fringe of hairs appearing
only in the sinuses, not ringing the throat as in C. mitreola, and by the
strongly incurved capsule lobes which are almost connivent, so that
the capsule often appears nearly globular.
The marked differences in the two species are well illustrated by
Hooker, in Icones Plantarum 9: t. 827 and 828, 1852.
GELSEMIUM Juss.
Reference: A. L. de Jussieu, Gen. PL 150. 1789.
Perennial woody, evergreen vines, the stems and branches glabrous; leaves
opposite, entire, membranaceous or chartaceous, glabrous except for some minute
puberulence on under surface near leaf bases, lanceolate to ovate, the petioles con-
nected by a transverse line; flowers dimorphic, often fragrant, large and showy, in
terminal or axillary, 1-5-flowered cymes subtended by several small bracts; calyx
5-parted, the imbricate segments appearing dry, with submembranaceous margins;
corolla funnelform, dilated at the throat, the 5 lobes ovate to oblong, imbricate;
stamens 5, included, inserted on corolla tube, the anthers linear-oblong, bilobate at
FIG. 78. Gelsemium sempervirens. A, habit, ]/% natural size; B, pistil, with calyx
and bracts, X 4; C, corolla dissected to show stamens, X 3; D, capsule, X 2.
287
288 FIELDIANA: BOTANY, VOLUME 24
base, the cells parallel, the filaments slender (the short filaments accompanying
the long style and the long ones accompanying the short style); style filiform or
subulate-filiform, bifid, each branch bilobate; ovary superior, seated on a disc, ob-
long, bilocular, with numerous ovules, 3-4 seriate upon a linear placenta; mature
capsule ovoid to oblong, obcompressed, beaked, the 2 carpels dehiscent at the apex,
the seeds usually numerous, compressed, orbicular, tuberculate-rugose, sometimes
winged; endosperm carnose, embryo straight or slightly curved.
One species, G. elegans Benth., is known only from eastern Asia;
of the remaining two, G. rankinii Small is apparently confined to the
coastal plain of the southeastern United States, and only G. semper-
virens (L.) Persoon ranges through the southern United States to
Mexico and Guatemala.
Gelsemium sempervirens (L.) Persoon, Syn. PI. 1: 267. 1805.
Bignonia sempervirens L. Sp. PI. 623. 1753. Gelseminum sempervirens
Catesby, Nat. Hist. Carolina, Edwards rev. 1: 53. 1754. Gelsemium
nitidum Michx. Fl. Bor. Am. 1: 120. 1803. G. sempervirens (Catesby)
Jaume-Saint-Hilaire, Exposition des Fam. Nat. 1: 338. 1805. G.
sempervirens Ait. f. Hort. Kew. 2(2): 64. 1811.
In damp thickets or forest, 1,200-2,600 m.; Alta Verapaz; Baja
Verapaz; Quiche"; Zacapa. Southern United States; southern Mexico.
Perennial, woody vine, glabrous throughout; leaves short-petiolate, subcoria-
ceous or chartaceous, the venation inconspicuous, dark green and lustrous above,
lanceolate or lance-oblong, 3-8 cm. long, 1-4 cm. broad, acuminate or attenuate,
rounded or obtuse at the base, entire; sepals ovate, 3-4 mm. long, often very mi-
nutely ciliate; corolla bright yellow, 2-3.5 cm. long, the lobes less than half as long
as the tube; stamens included, adnate near base of tube; ovary oblong; mature cap-
sule oblong-ovate, 14-23 mm. long, 8-11 mm. wide, greenish-brown, prominently
veined, with a beak 1.3-3 mm. long; seeds ca. 7 in each locule, flat, tuberculate-
rugose, variously notched, winged apically.
PLOCOSPERMA Benth.
References: G. Bentham in Benth. & Hook. Gen. PI. 2: 789. 1876.
Solereder in Engler & Prantl, Pflanzenf . IV(2) : 19-30. 1892. Leeu-
wenberg, Acta Bot. Neerl. 16(2): 56-61. 1967.
Stiff, erect, glabrous shrubs or small trees; leaves opposite, coriaceous, lustrous,
apices usually obtuse to retuse, occasionally acute; flowers axillary, in clusters of
2-4, the pedicels short, filiform; calyx small, 5-6 parted (rarely 4), the segments
lanceolate; corolla funnelform-campanulate, the tube short, the throat broad, the
broad lobes imbricate; stamens 5, the filaments slender to filiform, affixed to the
tube of the corolla; anthers ovate, cordate at the base, the cells parallel; ovary
1-celled, contracted and stipitate at the base, the style filiform, twice bifid, the ulti-
mate branches stigmatose; ovules 4, affixed to the walls of the cell in decussate or
superposed pairs; capsule elongate, subterete, multicostate, 2-valvate from the
FIG. 79. Plocosperma buxifolium. A, fruiting branch, J^ natural size; B, flower,
X 2; C, corolla opened to show stamens and pistil, after Bentham in Hooker,
Icon. 12, t. 1195. 1876.
289
290 FIELDIANA: BOTANY, VOLUME 24
apex; perfect seed 1, linear, subterete, bearing at the apex a dense tuft of long hairs,
the testa subcoriaceous; endosperm thin, carnose; embryo linear, straight.
One species, southern Mexico and Guatemala.
Plocosperma buxifolium Bentham in Hooker, Icon. 12: 82,
t. 1195. 1876. P. microphyllum Baill. ex Solereder, Bull. Mens. Soc.
Linn. Paris 780. 1899. P. anomalum Blake, Contr. U. S. Nat. Herb.
24: 17. 1922. Frutitta (Zacapa).
Dry, brushy slopes and hillsides, 200-650 m.; El Progreso; Jalapa;
Santa Rosa; Zacapa. Mexico.
A shrub or small tree of 2-5 m., the young branches puberulent, becoming
glabrous; leaves sessile or on petioles 1-2 mm. long, the blades coriaceous, elliptic-
oblong, oblong-ovate or lanceolate-ovate, 1.5-5 cm. long, 1-2.5 cm. wide, usually
narrowly rounded and subemarginate at the apex (very rarely acute), rounded and
slightly unequal at the base, somewhat lustrous above, dull beneath, sparsely pilos-
ulous, puberulent, or glabrate; pedicels 2-13 mm. long; calyx 1-3 mm. long, 5-6
parted (rarely 4), the sepals lanceolate or lance-oblong, subacute, ciliate; corolla
blue-violet to purplish-magenta, the tube 5-9 mm. long, the limb 13-25 mm. in
diameter, with 5-6 rounded lobes; stamens included, the filaments inserted 3-5 mm.
above the base of the corolla; capsule linear, 5-9 cm. long, 2-4 mm. thick, glabrous,
finely costate; seeds subterete or compressed, 1.2-2.3 cm. long, the tuft of hairs
7-15 mm. long.
The shrub seems to be rare, and only a few collections of it have
been obtained.
POLYPREMUM L.
Reference: Linnaeus, Sp. PL 111. 1753.
Herbaceous annuals, glabrous, low, diffusely branched from base, the branches
dichotomous; leaves opposite, linear, connected by their dilated bases; flowers very
small, solitary and subsessile in the forks of branches and in leaf axils, or 2-4 in
terminal, cymose clusters; calyx 4-parted (rarely 5-parted), the segments linear-
subulate, erect, green, somewhat rigid; corolla shorter than the sepals, tubular-
campanulate, villous within throat, the lobes usually 4, sometimes 5, obtuse, im-
bricate; stamens 4 (rarely 5), the short filaments inserted at a point just above the
middle of the corolla tube, the anthers ovate, the cells parallel; ovary superior,
bilocular, the style short, with the capitate stigma entire or obscurely bilobate;
ovules numerous on the oblong placentae which are affixed at the base to the sep-
tum; capsule ovoid or obovoid, somewhat bilobate, subcompressed contrary to the
septum, loculicidally bivalvate; seeds numerous, small, subglobose, smooth, the
endosperm carnose, the embryo straight.
Polypremum procumbens L. Sp. PL 111. 1753.
Usually in dry, sandy soil, open fields or on open slopes, some-
times on gravel along streams, occasionally in oak or pine-oak forests,
GIBSON: FLORA OF GUATEMALA
291
FIG. 80. Polypremum procumbens. A, habit of plant, natural size; B, flower,
X 5; C, corolla opened to show stamen insertion; D, capsule, X 10.
sea level to 2,500 m.; Chimaltenango; Escuintla; Guatemala; Hue-
huetenango; Izabal; Jalapa; Quezaltenango ; San Marcos; Solola.
Eastern and southern United States; Mexico; West Indies; Central
America; northern South America.
A low annual, usually 8-20 cm. high, stems diffusely branched, often dense,
erect or becoming prostrate, the branches rather stiff, 4-angulate, leaves narrowly
linear, 4-20 mm. long, acute, sessile, minutely serrulate on the margins, often with
fascicles of smaller leaves in the axils; flowers sessile, axillary or 2-4 in terminal,
cymose clusters, subtended by bracts similar to the leaves; calyx segments 2-3 mm.
long, with strong midrib and scarious margins; corolla white, to 2 mm. long; cap-
292 FIELDIANA: BOTANY, VOLUME 24
sule crustaceous, obovoid, ca. 2 mm. in diameter, somewhat bilobate; seeds numer-
ous, subglobose, minute.
A common plant throughout Central America, and perhaps one
of the least conspicuous.
SPIGELIA L.
References: Linnaeus, Sp. PL 149. 1753; Progel in Martius, Fl.
Bras. 6(1): 253. 1868; Solereder, Pflanzenf. 4, Abt. 2: 32. 1892; L. B.
Smith, Wrightia 2: 90. 1960.
Annual or perennial herbs or subshrubs, glabrous or pubescent, with stems
terete or tetragonous; leaves simple, entire, decussate or in whorls at stem apices,
connected at base of petioles by a transverse line or by interpetiolar stipules; in-
florescences usually terminal, unilaterally spicate with several to many flowers, or
1-2-flowered in forks of branches, the flowers essentially sessile; calyx persistent,
5-parted, the segments narrow, with 2 or more linear glands inside at the base of
each; corolla tubular-campanulate to funnelform, the short lobes valvate in bud;
stamens 5, equal, the filaments inserted on the corolla tube, anthers oblong or lan-
ceolate, the 2 cells parallel, dehiscing longitudinally; ovary superior, bilocular, the
style filiform, the upper stigmatose portion soon caducous, the lower segment per-
sistent on mature capsule; ovules several to many on the peltate placentae; capsule
bilobate, more or less compressed contrary to the septum, loculicidal and septicidal,
circumcissile above the persistent, cupular base; seeds usually few, turbinate or
obliquely ellipsoid to ovoid, tuberculate, verrucose or reticulate; embryo short and
straight, endosperm copious.
Probably about 50 species, all American, six of which occur in
Guatemala. A seventh species, S. splendens Wendl. ex Hook., is
known from southern Mexico and Costa Rica and is therefore in-
cluded.
Capsules muricate or muriculate.
Plants 20-40 cm. tall; upper surface of leaf blades minutely scaberulous; cap-
sules 4-6 mm. broad; persistent style segment considerably exceeding lobes
of capsule S. anthelmia.
Plants 6-14 cm. tall; upper surface of leaf blades smooth; capsules 2 mm. broad;
style articulate at apex of capsule S. pygmaea.
Capsules smooth, never muricate.
Leaves linear to linear-lanceolate, usually 1-2 cm. long; corollas 2-3 mm. long.
S. polystachya.
Leaves ovate or lanceolate or obovate, never linear.
Corollas rose-red, showy, 25-30 mm. long; stamens attached to tube by short
filaments directly below the sinuses S. splendens.
Corollas white or pallid, 20 mm. long or less; stamens attached to tube by
short filaments inserted about midway of tube.
Average length of leaf blades 3-8 cm.
Leaves thin; persistent style segment longer than sepals, exceeding apex
of capsule S. humboldtiana.
Leaves thick and succulent; persistent style segment equalling or shorter
than sepals, not exceeding apex of capsule S. carnosa.
Average length of leaf blades 1-2 cm S. coulteriana.
GIBSON: FLORA OF GUATEMALA 293
Spigelia anthelmia L. Sp. PL 149. 1753; Sims in Curtis Bot.
Mag. 50: t. 2359. 1823; A. DC., Prodr. 9: 7. 1845; Leeuwenberg, Acta
Bot. Neerl. 10: 460. 1961. S. quadrifolia Stokes, Bot. Mat. Med. 1:
307. 1812. S. nervosa Steud. Flora 26: 764. 1843. S. anthelmia var.
obliquinervia A. DC. Prodr. 9: 7. 1845. S. fruticulosa Lam. Illust. 1:
478. 1851. S. stipularis Prog, in Martius, Fl. Bras. 6(1): 262. 1868.
S. anthelmia var. nervosa (Steud.) Prog. I.e. S. domingensis Gando-
ger, Bull. Soc. Bot. Fr. 70: 921. 1923.
In damp thickets, 500 m. or less; Chiquimula; Santa Rosa; Za-
capa. Southern Florida, Mexico and Central America; West Indies;
Colombia, Venezuela, British and French Guiana, Peru and Brazil;
naturalized in tropical Africa and Indonesia.
Annuals, erect, simple or sparsely branched, usually 20-40 cm. high, the stems
glabrous, stipules inconspicuous; leaves entire, the blades thin, papyraceous when
dry, minutely ciliolate, lanceolate or ovate-lanceolate, attenuate, the lower ones
opposite, short-petiolate, the upper ones usually in a whorl of 4 (2 decussate pairs),
sessile or nearly so, 4-15 cm. long, one pair considerably larger than the other, the
blades deep green and minutely scaberulous above, pale and glabrous beneath; in-
florescences terminal, usually in slender secund spikes of 10-20 bracteolate flowers
(occasionally only 1 or 2 flowers); calyx segments linear-lanceolate, 2-3 mm. long;
corolla white or pale pink (rarely light yellow), 5-10 mm. long, tubular-campanu-
late, the lobes short, ovate, acute; stamens included, the glabrous filaments inserted
about midway of tube; ovary nearly globose, style equalling the corolla or slightly
exserted, considerably exceeding calyx; capsule 3-5 mm. long, 4-6 mm. broad,
bilobate, finely muricate, with persistent portion of style to 2 mm. long, exceeding
lobes of capsule; seeds 12-15 per capsule.
Spigelia carnosa Standl. & Steyerm. Field Mus. Bot. 23: 72.
1944.
Dense, wet, mixed forest, 1,300-2,000 m.; endemic. Alta Vera-
paz, Huehuetenango.
Perennials, erect or decumbent, the lower portion often prostrate and rooting,
glabrous throughout; stems terete, stipules persistent, triangular, 1.5-3 mm. long;
leaves opposite or the uppermost quaternate, short-petiolate, the blades thick and
fleshy when fresh, almost coriaceous when dry, deep green above, much paler be-
neath with nerves often obscure, elliptic to elliptic-oblong, ovate or obovate, 4-
10 cm. long, short-acuminate to obtuse, acute at base; inflorescences spicate, 4-
13 cm. long, usually many-flowered, the flowers sessile; calyx segments ca. 5 mm.
long, linear-lanceolate, subrecurved; corolla 1.5 cm. long, white, pinkish, or pale
lavender outside, white within; stamens included, the short filaments inserted
about midway of tube; ovary nearly globose; style in flowers from which the corolla
has just fallen equalling or shorter than the sepals, equalling the apex of the mature
capsule; capsule 4-5 mm. broad, smooth, glabrous; seeds 10-12 per capsule, yellow
or brown, ca. 2.5 mm. long, minutely reticulate.
294 FIELDIANA: BOTANY, VOLUME 24
Spigelia coulteriana Bentham, Journ. Linn. Soc. 1: 90. 1857.
Huehuetenango, Sierra de los Cuchumatanes, 1,400-2,100 m.
Mexico.
Plants perennial, small, 5-8 cm. high, the erect stems tetragonous, arising from
a prostrate base; leaves opposite, decussate, the uppermost quaternate, the blades
obovate to ovate, 1.5-2.8 cm. long, 1.5-2.5 cm. broad, acute or subobtuse; spicate
inflorescences terminal or 1-3 from leaf axils, simple or branching, 3-10-flowered,
the flowers sessile or subsessile; calyx segments linear, 2-3 mm. long, ciliolate,
longer than the persistent style segment; corolla 1.5-2 cm. long, white with laven-
der outside; stamens included, the short filaments inserted about midway of the
tube; ovary subglobose, persistent style segment ca. 1.5 mm. long; capsule bilobate,
smooth, 2-3 mm. long, 3-4 mm. broad.
Spigelia humboldtiana Cham. & Schlecht. Linnaea 1: 200.
1826. S. scabra Cham. & Schlecht. I.e. 202. Lombricera (Quezalte-
nango) ; Lombriz quen (Alta Verapaz) .
Wet forests or thickets, along streams and on gravel bars, rarely
in open ground, sea level to 2,000 m.; Alta Verapaz; Chimaltenango;
Chiquimula; Guatemala; Huehuetenango; Izabal; Jalapa; Pete"n;
Quezaltenango; Sacatepe"quez; San Marcos; Santa Rosa; Zacapa.
Southern Mexico to Panama; South America.
Perennial from long, horizontal rootstocks, erect or decumbent, 12-40 cm. high
(usually 20 cm. or more), usually glabrous except for scattered puberulence on
stems, petioles, and lower surface of leaves, the stems tetragonous or subterete,
simple or branched; leaves short-petiolate or sessile, opposite, the uppermost qua-
ternate, the blades thin, lance-oblong to ovate, minutely ciliolate, 2.5-14 cm. long
(usually 3-7 cm.), 1.5-4 cm. wide, acuminate to subobtuse, rounded and abruptly
decurrent at the base or attenuate or obtuse, paler beneath; stipules small, triangu-
lar or ovate- triangular; inflorescences terminal, solitary or binate, usually few-
flowered, the flowers sessile or subsessile; calyx segments linear-lanceolate, 1.5-
3.5 mm. long; corolla 8-10 mm. long, usually white or cream, sometimes pale pink
or tinged with purple, red or green; stamens included, the short filaments inserted
about midway of tube; ovary nearly globose; style remnant conspicuously longer
than the calyx; mature capsule 4-6 mm. wide, bilobate, glabrous; the persistent
style segment extending beyond the lobes of the capsule; seeds grayish.
Although an inconspicuous plant, this is well known in Central
America because the rootstocks are often used in decoction to expel
intestinal parasites in man. It has been stated that excessive doses
of this and other species may result in death.
Spigelia polystachya Klotzsch ex Prog, in Martius, Fl. Bras.
6(1): 265. 1868.
Wet plains, borders of swamps, mud flats, 50-950 m.; Jutiapa,
Peten. Honduras; El Salvador northern South America.
GIBSON: FLORA OF GUATEMALA
295
FIG. 81. Spigelia pygmaea. A, habit, X 1; B, flower, X 15; C, corolla dis-
sected to show stamens, X 15; D, pistil, X 15; E, capsule showing muriculate apex,
X 10. Drawing by Davida Simon.
Plants annual, small, erect, usually 6-10 cm. high but may attain 20 cm., gla-
brous, the stems branching, tetragonous; leaves opposite, the blades linear, linear-
oblong, or linear-lanceolate, usually 1-2 cm. long, 0.1-0.2 cm. broad, but may
attain 3.5 cm. X 0.5 cm., subacute, attenuate to the sessile base; spicate inflores-
ces solitary in the leaf axils, 5 cm. long or shorter, usually many-flowered, the
296 FIELDIANA: BOTANY, VOLUME 24
flowers sessile; calyx ca. one-third as long as the corolla, ca. one-half as long as the
mature capsule, the segments ovate-lanceolate, subobtuse, glabrous; corolla sub-
campanulate, 2-3 mm. long, villous within; stamens included, the very short fila-
ments inserted about midway of the tube; ovary subglobose, style articulate at or
very near apex of capsule; mature capsule bilobate, 1.5-2 mm. long, 2-3 mm. broad,
smooth, glabrous, seeds ridged.
Spigelia pygmaea D. Gibson, Fieldiana: Botany 32: 5, t. 1968.
In zapotal, on La Gloria Road, Dos Lagunas, ca. 7 km. west of
village, Pete'n. Endemic.
Plants annual or possibly perennial, the lower portion sometimes creeping and
rooting, small, erect, 6-14 cm. tall (average height 6-8 cm.), the stems short, 1-
3 cm. long, tetragonous, the stipules broadly triangular; leaves decussate or in a
whorl of 4 (2 decussate pairs), one pair much larger than the other, the blades gla-
brous, entire, tapering to the short, narrowly winged petioles, thin, obtuse, those
of one pair ovate-rhombate, the second and larger pair oblong-ovate or oblong-
elliptic, obtuse to acute, the smaller pair 1-2 cm. long, 0.8-1.5 cm. broad, the larger
pair 2-5 cm. long, 1.5-2.5 cm. broad, deep green above, paler beneath, the nerves
prominent, few; inflorescences multispicate, the flowering spikes erect, some arising
from near the base, others, especially on older plants, from leaf axils, sometimes
branching, 1.5-9 cm. tall (averaging 4-6 cm.), minutely granular-verrucose, many-
flowered, the flowers sessile or subsessile, bracteolate; sepals lanceolate, ciliolate,
minutely granular, especially along the midnerve, ca. 1 mm. long; corolla white,
ca. 1.5 mm. long, subcampanulate, smooth within, the lobes broadly lanceolate,
acute; stamens included, the very short filaments inserted just below the middle
of the tube; ovary subglobose, the style minute, the stigma articulating directly
above the apex of the capsule; mature capsule bilobate, muriculate, 1-1.5 mm.
long, ca. 2 mm. broad; seeds ridged.
Although about the same height as S. polystachya, and with the
same minute style segment, S. pygmaea may be readily separated by
its broad leaves, multispicate inflorescences, and muriculate capsules.
Spigelia splendens Wendl. ex Hooker, Bot. Mag. t. 5268. 1861.
S. platyphylla Prog, in Martius, Fl. Bras. 6(1): 256. 1868.
In shaded, moist places, wooded or shrubby slopes, 1,500 m. or
less; Mexico and Costa Rica.
Perennials, erect, herbaceous, the simple stems 45 cm. high or less, subterete,
usually more or less villous; leaves thin, sessile or short-petiolate, ovate or broadly
obovate-rounded, 7-20 cm. long, 5-12 cm. wide, acute or cuspidate, cuneate at
base or broadly and abruptly contracted, sparsely villous or nearly glabrous, mi-
nutely ciliate, conspicuously nerved, 4 leaves in a whorl near the apex of the stem,
occasionally an additional pair of smaller leaves shortly below the top whorl, and
the lowermost pair usually reduced to scales; spicate inflorescences 1-10, flowers
short-pedicellate, the upper ones sessile; calyx segments linear-lanceolate, 4-5 mm.
long; corolla tubular, 2.5-3 cm. long, glabrous or minutely puberulent, rose-red;
GIBSON: FLORA OF GUATEMALA 297
anthers conspicuous but not exceeding the corolla, the short filaments attached to
tube shortly below the sinuses; ovary globose, the persistent portion of the style
becoming 1-1.5 cm. long; capsule 5-6 mm. broad, glabrous or minutely puberulent,
smooth; seeds reticulate-rugose.
Although no specimens from Guatemala were seen, it is to be
expected there.
STRYCHNOS L.
References: Linnaeus, Sp. PI. 189. 1753 and Gen. PI. ed. 5. 86.
1754. B. A. Krukoff and J. Monachino, The American species of
Strychnos, Brittonia 4: 248-322. 1942. B. A. Krukoff, Supplemen-
tary notes on the American species of Strychnos, VII, Mem. N. Y.
Bot. Gard. 12(2): 1-94. 1965.
Woody plants, erect shrubs in the juvenile stage and in open situations but
becoming lianas in moist woods, usually with spines or tendrils or both; leaves
opposite, simple, entire, the petiole bases connected by stipular lines, the blades
longest toward the ends of the branchlets, those near the bases of branchlets some-
times reduced to cataphylls, membranaceous to coriaceous, 3-5 plinerved (usually
2 principal nerves, one on each side of the midrib, in ours) ; inflorescences terminal
or axillary, short-thyrsoid or corymbose-paniculate, composed of 1-3-flowered
cymes, calyx usually 5-parted (sometimes 4); corolla salverform or rotate, the 5-4
lobes valvate; stamens 5 or 4, the short filaments affixed in the throat of the corolla,
anthers ovate (in ours), dorsifixed; ovary bilocular (rarely unilocular), the style
filiform (in ours), stigma more or less capitate and obscurely bilobate at the apex;
ovules axile, numerous; fruit baccate, globose or nearly so, often very large, inde-
hiscent, the exocarp usually rather thick and hard, the endocarp sometimes devel-
oping a wool that adheres to the testa of the seed; seeds usually many, variable in
shape, sometimes irregularly 3-sided, discoid, spheroid, or elliptic-oblong.
Of the some 200 reported species generally distributed in tropical
regions, 64 are American, with five occurring in Guatemala. The
genus Strychnos is well known as the source of various alkaloids hav-
ing paralyzing and poisonous properties. The best known species is
the Indian S. Nux-vomica L., source of strychnine and Nux-vomica.
Unfortunately, the bulk of the collections from Guatemala is
sterile material. I have therefore accepted Krukoff's annotations
and my treatment of the genus follows his interpretation of the species.
Corolla equalling or shorter than the calyx; plants often armed with stout spines;
leaf blades usually lanceolate and usually less than 8 cm. long . . .S. nigricans.
Corolla much longer than the calyx; plants always unarmed.
Inflorescences axillary; mature leaf blades usually elliptic-oblong and large, to
30 cm. long, usually twice as long as broad S. peckii.
Inflorescences terminal.
Filaments not distinct; styles pilose; leaf blades long-lanceolate or elliptic-
obovate, to 17 cm. long, usually 3 or more times longer than broad.
S. chloranlha.
298 FIELDIANA: BOTANY, VOLUME 24
Filaments distinct (much longer than anthers); styles glabrous; leaf blades
variable but most often ovate to broadly lanceolate.
Corolla glabrous outside S. panamensis.
Corolla pubescent outside S. labascana.
Strychnos chlorantha Prog, in Martius, Fl. Bras. 6(1): 273.
1868.
Wet, mixed forest, on limestone, about 900 m.; Alta Verapaz.
Costa Rica.
Unarmed woody vines; leaves short-petiolate, the blades coriaceous, lustrous,
glabrous, elliptic-obovate to lanceolate, 5-17.5 cm. long, 2.5-5 cm. broad, acumi-
nate to long-acuminate, attenuate or acute at the base; inflorescences in terminal
cymes, corymbose, sparsely puberulent, the pedicels 4 mm. long or less; calyx lobes
broadly ovate, 1 mm. long, rounded or obtuse, ciliate; corolla tube 1.5 cm. long,
glabrous, not papillose, the lobes 4 mm. long; anthers subsessile, partly exserted,
2.7 mm. long or less; styles pilose; fruits globose, ca. 6 cm. in diameter, with shell
nearly 1 cm. thick.
Strychnos nigricans Prog, in Martius, Fl. Bras. 6(1) : 280. 1868.
S. brachistantha Standl. Field Mus. Bot. 12: 412. 1936. Ichbolay
(Alta Verapaz) .
Wet forests, wooded bluffs, sea level to 800 m.; Alta Verapaz;
Huehuetenango ; Izabal; Pet£n. Southern Mexico; Honduras; Nica-
ragua; Panama; Venezuela; southern Brazil.
Woody vines, the branches minutely puberulent, often armed with stout, re-
curved spines 5-20 mm. long; leaves chartaceous or membranous, short-petiolate,
the blades lanceolate, ovate-lanceolate or ovate-elliptic, 3-12 cm. long (usually
less than 8 cm. long), 1-3 cm. broad, acuminate to long acuminate, rounded or
acute at the base, with sparse puberulence often on midribs and petioles; inflores-
cences small, terminal, many-flowered, glabrous or puberulent, pedicels equalling
or shorter than the calyx; calyx lobes ovate-lanceolate, ca. 1 mm. long, acute or
acuminate, glabrous; corolla equalling or shorter than the calyx, the lobes obscurely
papillose outside, barbate within; anthers glabrous or pilose; ovary and style gla-
brous; fruits globose, 5 cm. or more in diameter, shell 5-6 mm. thick, seeds numer-
ous, ca. 17 mm. long, pale brown to orange.
Strychnos panamensis Seem. Bot. Voy. Herald 166. 1854. S.
hachensis Karst. Fl. Columb. 2: 75. 1863. S. longissima Loes. Repert.
Sp. Nov. 9: 357. 1911. S. tepicensis Standl. Contr. U. S. Nat. Herb.
23: 1142. 1924. Aguacate de mico (Santa Rosa).
Damp thickets or forests of lowlands, sometimes in rocky thickets
along stream banks, rarely in dry thickets (in shrubby form), 100-
1,600 m.; Escuintla; Quezaltenango; Retalhuleu; Sacatepe" quez ; San
FIG. 82. Strychnos peckii. A, flowering branch, X 1A; B, flower, X
C, corolla dissected to show interior, stamens and pistil, X 4; D, fruit, X 1A-
299
300 FIELDIANA: BOTANY, VOLUME 24
Marcos; Santa Rosa. Western Mexico to Panama. Northeastern
Venezuela and northern Colombia.
Unarmed woody vines, the stems sometimes attaining a diameter of 10 cm.
or more, the branches and tendrils often puberulent to hirsute; leaves on short,
often pubescent petioles, the petioles 1-5 mm. long, the blades quite variable, ovate
to elliptic or lanceolate, 4-13 cm. long, 1.5-6 cm. wide, acute or acuminate, rounded,
subcordate or acute at the base, membranaceous to chartaceous, sparsely pubescent
beneath along base and midrib or quite glabrous; inflorescences in terminal cymes,
corymbose, the branches hirsutulous, the pedicels 3 mm. long or less, usually hir-
sutulous; calyx lobes lanceolate, acuminate, 2-3 mm. long, ciliate; corolla white or
yellowish, 1.5-2 cm. long, papillose and glabrous outside, the tube pilose within
but the lobes densely papillose within and glabrous; stamens exserted, the filaments
more than twice as long as the anthers; style glabrous; fruit globose, 4-8 cm. in
diameter, many-seeded.
This is probably the most frequently collected species in Central
America. Sterile collections may be confused with S. nigricans,
which when young may also be unarmed; however, the leaf blades
of S. panamensis are usually broader and more ovate than those of
S. nigricans.
Strychnos peckii Robinson, Proc. Am. Acad. Sci. 49: 504. 1913.
Wet forest or thickets, at or a little above sea level; Izabal; Pete"n.
Honduras; Costa Rica. South America.
Unarmed woody vines, attaining a diameter of 15-20 cm., the branchlets puber-
ulent; leaves on puberulent petioles 5-18 mm. long, the blades elliptic to elliptic-
oblong or lance-oblong, 7-30 cm. long, 3.5-15 cm. wide, acuminate, rounded to
obtuse at the bass, coriaceous, minutely puberulent beneath when young, becoming
almost glabrous with age; inflorescences axillary, cymose, many-flowered, fulvous-
puberulent, the pedicels 2 mm. long or less; calyx lobes broadly ovate- triangular,
ca. 1 mm. long, subacute, densely puberulent, ciliate; corolla pale yellowish-green,
the tube 6-9 mm. long, densely papillose outside and pubescent, lanate within;
anthers subsessile, included except at the tip, 1.5 mm. long or less; style glabrous;
fruit globose, 6 cm. or more in diameter, buff; seeds many.
Strychnos tabascana Sprague & Sandw. Kew Bull. 128. 1927.
S. panamensis var. hirtiflora Standl. Field Mus. Bot. 11: 138. 1932.
S. hirtiflora Lundell, Bull. Torr. Bot. Club 64: 556. 1937.
Wet forest or thickets, sea level to 900 m.; Alta Verapaz, Izabal,
Pete"n. Mexico; Honduras; Costa Rica.
Unarmed woody vines, the branchlets and tendrils often puberulent or hirsut-
ulous; leaves on pubescent petioles 1-3 mm. long, the blades ovate to elliptic-
lanceolate, 5-11 cm. long, 2.5-5 cm. wide, acuminate, cuneate to obcordate at the
base, often puberulent beneath when young, but glabrous or nearly so in age, mem-
branaceous; inflorescences in terminal cymes, corymbose, the branches hirsutulous,
GIBSON: FLORA OF GUATEMALA 301
the pedicels 1-3 mm. long, hirsutulous; calyx lobes lanceolate, 2-3 mm. long,
papillose, ciliate; corolla white or cream, 11-17 mm. long, the lobes and upper
third of tube pilose to subsetulose outside, white-lanate within; stamens exserted,
the filaments about twice as long as the anthers; style glabrous; fruit globose,
4-8 cm. in diameter, many seeded.
GENTIANACEAE. Gentian Family
PAUL C. STANDLEY
AND
Louis 0. WILLIAMS
Annual or perennial herbs, glabrous throughout, with usually bitter sap ; leaves
opposite (except in Nymphoides), often connate at the base or connected by a
transverse line, without stipules,1 rarely reduced to scales, entire, the whole plant
sometimes destitute of chlorophyll, but the plants usually with normal green leaves;
inflorescence various in form but most often several times dichotomous; flowers
large or small, various in color, regular or nearly so, perfect, rarely polygamous;
calyx inferior, the tube campanulate or often very short or none, the teeth or lobes
4-5, imbricate or open in bud; corolla gamopetalous, funnelform, salverform, cam-
panulate, or rotate, the limb 4-5-lobate, the lobes usually dextrorsely contorted;
stamens as many as the corolla lobes and alternate with them, inserted in the throat
or tube of the corolla, the filaments filiform or dilated at the base; anthers dorsifixed
above the base, erect, versatile, or reflexed, bilobate at the base, the cells distinct,
parallel, dehiscent by longitudinal slits; disk none, or annular or 5-glandular, incon-
spicuous; ovary superior, usually sessile, generally 1-celled, with 2 parietal placen-
tae; style simple, the stigma terminal, capitate or bilamellate or shallowly bifid at
the apex; ovules usually numerous, 1-many-seriate on the placentae, anatropous
or amphitropous; fruit capsular, membranaceous or indurate, usually bivalvate;
seeds globose, angulate, or rarely compressed, sometimes narrowly winged, the
testa membranaceous or crustaceous, often foveolate, reticulate, or tuberulate;
endosperm usually copious; embryo small, subterete or conic, the radicle usually
superior.
Genera about 80, best represented in temperate regions of both
hemispheres, but species rather numerous also in the tropics, at low
or high elevations. Two other genera, Symbolanthus and Enicostema,
are represented in southern Central America (Costa Rica) and
Panama.
Plants without chlorophyll; leaves reduced to small scales.
Capsule dehiscent by longitudinal slits between the apex and base, but not
through the base and apex; plants very slender and delicate, the leaf scales
inconspicuous, few, closely appressed to the stem; seeds with elongated
wings Leiphaimos.
1 In some Lisianthus the connate leaf bases appear very much like the inter-
petiolar stipules to be found in Rubiaceae.
302
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 303
Capsule dehiscent through its whole length; stems stout, the leaf scales numer-
ous and conspicuous, not appressed but loose and somewhat spreading;
seeds wingless Voyria.
Plants with normal green leaves.
Leaves alternate; corolla lobes induplicate-valvate; aquatic plants with mostly
orbicular or reniform leaves Nymphoides.
Leaves all of chiefly opposite; corolla lobes contorted or rarely imbricate; plants
not aquatic, the leaves not orbicular or reniform.
Stigma capitate.
Plants small and rather delicate; flowers about 5 mm. long; leaves linear.
Curtia.
Plants large and coarse; flowers much larger; leaves lanceolate to broadly
ovate.
Plants tricotomously branched, rarely simple; corolla not gibbous on one
side Lisianthus.
Plants usually simple or dichotomously branched; corolla gibbous ven-
trally Chelonanthus.
Stigma bilamellate.
Anthers in age spirally twisted; corolla usually pink Centaurium.
Anthers unchanged in age, or sometimes recurved.
Inflorescence spicate Coutoubea.
Inflorescence not spicate.
Corolla foveolate within near the base below each lobe, often calcarate
or saccate, green or greenish yellow Halenia.
Corolla not foveolate, never calcarate or saccate, usually blue, pink or
purplish, rarely green (if yellow, see Halenia).
Corolla usually pink, rarely yellowish; plants annual; calyx conspicu-
ously costate or winged Schultesia.
Corolla blue or bluish, rarely white; plants annual or perennial; calyx
not at all or not conspicuously costate, not winged.
Style very short; plants perennial or annual; flowers large or small;
lobes of the calyx shorter than the tube or but little exceeding it.
Gentiana.
Style slender and much elongate; plants annual; flowers large; lobes of
the calyx several times as long as the short tube Eustoma.
CENTAURIUM Hill
Slender annual herbs, glabrous, erect or diffusely and dichotomously branched;
leaves opposite, small, sessile or amplexicaul; inflorescence a dichotomously
branched, determinate or indeterminate few-to-many-flowered arrangement; flow-
ers small to rather large, usually pink, sometimes white or yellowish; calyx 4-5-fid,
the lobes usually carinate; corolla with a short tube or one as long as or longer than
than the lobes, usually diaphanous and marcescent, the corolla lobes usually 5,
spreading at an thesis, contorted in the bud and often after an thesis; stamens 5,
attached near the throat of the corolla tube, the anthers usually oblong, often spi-
rally twisted in age, or sometimes unchanged; ovary 1-celled, the placentae often
strongly intruded; seeds small, suborbicular to ovoid, numerous.
304 FIELDIANA: BOTANY, VOLUME 24
There are perhaps 50 or more species in temperate and tropical
regions of both hemispheres. The genus is in need of revision and
should provide a most interesting problem in distribution.
The generic name Erythraea has often been used for this group of
plants. The proper name seems to be Cenlarium of Hill.
Calyx lobes ovate or lance-ovate, shorter than the capsule or corolla tube.
C. strictum.
Calyx lobes linear-lanceolate, equaling or longer than the ovary or corolla tube.
Leaves mostly linear to oblong or elliptic.
Plants diffusely dichotomously branched; flowers at an thesis 7-9 mm. long.
C. quitense.
Plants much branched above; flowers at an thesis 10-13 mm. long. . . C. rosans.
Cauline leaves subulate or narrowly linear.
Plants to about 10 cm. tall; basal leaves broadly elliptic or ovate; corolla
6-7 mm. long C. setaceum.
Plants 15-35 cm. tall; basal leaves subulate, minute; corolla about 9 mm. long.
C. pringleanum.
Centaurium pringleanum (Wittr.) Robinson, Proc. Am. Acad.
45: 397. 1910; Erythraea pringleana Wittr. Bot. Gaz. 16: 85. 1891.
Grassy slopes or meadows, often in the pine forest, 900-1,400 m.
To be expected in Guatemala. Mexico; Honduras.
Strict, slender annual herbs 15-35 cm. tall. Stems about 1 mm. in diameter,
with 4 narrow wings or ridges originating from the bases of the leaves, these less
prominent below; leaves 5-8 opposite pairs, subfiliform, thickened but somewhat
flattened, acute, 2-18 mm. long and 0.5-0.8 mm. broad, largest pair subtending the
inflorescence; inflorescence few-20-flowered, an indeterminate dichotomous di-
chasioid arrangement or ultimately a simple dichasium, the bracts similar to the
leaves and often quite large; flowers long pedicellate, the pedicels bibracteate or
bractless, to about 1 cm. long; calyx 5-lobed, free to the base, the lobes linear-
subulate, carinate, margins scarious below, about 5 mm. long and 0.7 mm. broad;
corolla tube about 5 mm. long, diaphanous and marcescent, the lobes elliptic-
oblong, obtuse, somewhat cucullate, obscurely apiculate, about 4 mm. long and
2 mm. broad, probably spreading at anthesis; stamens alternate with the lobes and
attached in the throat, the anther oblong and about 1.2 mm. long and 0.6 mm.
broad, somewhat twisted with age, the filaments about as long as the anthers, very
slender; maturing ovary ellipsoid, about 5 mm. long; style about 1.5 mm. long, the
stigma obscurely bilamellate; mature capsule exceeded by or about as long as the
calyx lobes; seeds minute, ovoid, about 0.12 mm. in diameter.
It is curious that this species jumps from the Mexican plateau to
Honduras but I find no difference in the material available. The
species should be found in Guatemala.
Centaurium quitense (HBK.) Robinson, Proc. Am. Acad. 45:
397. 1910. Erythraea quitensis HBK. Nov. Gen. & Sp. 3: 178. 1818.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 305
E. tetramera Schiede, Bot. Zeit. 13: 919. 1855. E. divaricata Schaff-
ner ex Schiede, I.e. 920.
Grassy, usually moist fields, often in marshy places or on sand-
bars along streams, rarely in dry rocky places, frequently in oak-pine
forest, 400-2,500 m.; Alta Verapaz; Baja Verapaz; El Progreso; Chi-
quimula; Jalapa; Santa Rosa; Guatemala; Sacatepe"quez ; Chimalte-
nango; Quiche". Mexico; Honduras to Panama; West Indies; western
South America.
An annual, 30 cm. high or less, usually erect and diffusely branched, the stems
slender, angulate; lower leaves oblong to elliptic, the principal cauline leaves oblong
or lanceolate, the lower ones obtuse, the upper acute to attenuate, mostly 1-3 cm.
long, sessile; flowers usually very numerous, the slender pedicels 1-3.5 cm. long,
spreading or ascending; flowers at an thesis 7-9 mm. long; calyx 4.5-5 mm. long, the
segments linear-lanceolate, long-attenuate; corolla dull pink or dirty pink, the tube
equalling the calyx, the lobes obtuse, much shorter than the tube; capsule linear-
oblong, 5-6 mm. long.
A common and often rather weedy plant at middle elevations or
lower. The available material is slightly variable, but it is believed
that it represents a single species. Plants growing in wet places often
are very lax and have weak stems, while their leaves are relatively
broader and unusually thin. Most remarkable of the Guatemalan
collections is one (Standley 80599) from high up on the active cone
of Volcan de Pacaya. The plants are only 5 cm. high, very densely
and compactly branched, and many-flowered, the flowers on short
pedicels. It was believed at first that this represented a new species,
but apparently it is only a much stunted plant, influenced by the
cold and dry soil that prevail in this locality. This is the more prob-
able because the normal form of C. quitense has been collected at the
same place. This species was reported from Guatemala by Bentham
in Plantae Hartwegianae as Erythraea chilensis Pers.
Centaurium rosans Standl. & Steyerm. Field Mus. Bot. 23: 75.
1944.
Open, moist or rather dry banks or open forest, most often in pine-
oak forest, 1,600-3,000 m.; Alta Verapaz; Sacatepe"quez ; Chimalte-
nango; Solola; Quezaltenango (type from Volcan de Zunil, Steyer-
mark 34609); Huehuetenango; Santa Rosa. Honduras; possibly
Nicaragua.
An erect annual, densely and rather laxly branched from the base, 10-30 cm.
high, the stems slender, often diffusely branched, angulate; leaves spreading,
broadly linear or oblanceolate, 1-2 cm. long, sessile but usually attenuate to the
base, acute; flowers in anthesis 11-13 mm. long, long-pedicellate, the pedicels
FIG. 83. Centaurium rosans. A, habit, X H; B, flower dissected, X 3. Curtia
tenella. C, three plants, natural size, to show variation; D, corolla dissected open
to show pistil and stamens, X 10; E, calyx with subtending bract, X 10.
306
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 307
1-2 cm. long, straight, ascending, not erect; calyx 6 mm. long, the segments linear-
lanceolate, long-attenuate; corolla tube equalling the calyx, the lobes broadly ovate,
spreading, usually bright pink, obtuse, slightly longer than the tube; seeds very
numerous, minute, brown.
Centaurium setaceum (Benth.) Robinson, Proc. Am. Acad. 45:
397. 1910. Erythraea setacea Benth. Bot. Voy. Sulph. 128. 1846.
Grassy open places in pine forest, 1,200-1,400 m.; Chiquimula
(Caracol Mountain, 1.5 miles northwest of Quezaltepeque, Steyer-
mark 31384). Mexico (Jalisco).
An erect annual, 10-20 cm. high, laxly and rather sparsely branched above the
base, the branches widely ascending; radical leaves 4 mm. long, obovate, soon with-
ering; cauline leaves few pairs, linear-setaceous, 15 mm. long or shorter, spreading;
flowers few or rather numerous, on long slender pedicels, 8-9 mm. long; calyx 5 mm.
long, the segments linear-lanceolate, long-attenuate; corolla bright pink, the tube
equaling the calyx, the lobes elliptic-oblong, obtuse, slightly shorter than the tube;
capsule oblong, equaling the calyx, the seeds very numerous, minute.
The known range for the species is curious,- — from the Mexican
highlands near Guadalajara (although the type is said to be from
Acapulco, which seems unlikely) to Guatemala.
Centaurium strictum (Schiede) Druce, Kept. Exch. Club Brit.
Isles 1916: 614. 1917.
Wet to dry, open or shaded banks, grassy hillsides, sometimes in
open rocky places, or in Alnus or oak-pine forest, 1,300-3,000 m.;
Zacapa; Jalapa; Guatemala; Sacatepe'quez ; Quiche"; Chimaltenango;
Solola; Huehuetenango; Quezaltenango; San Marcos. Central and
southern Mexico; El Salvador; Costa Rica; Panama.
An erect annual, 40 cm. high or less, the stems mostly simple or with a few
erect branches; leaves spreading or the upper ones suberect or ascending, 10-18
mm. long, broadly linear to linear-lanceolate or oblanceolate, acute or obtuse, not
narrowed at the base; flowers rather few, 12-14 mm. long, pink or dull dirty pink,
on stout, erect or suberect pedicels, all the flowers produced in the upper half of the
plant or higher; calyx segments lance-ovate, much shorter than the corolla tube,
acute, conspicuously carinate; lobes of the corolla acute, slightly shorter than the
tube; capsule oblong, 8-9 mm. long.
We have not seen authentic material of this species and are not
sure that our material belongs here, nor that all the Central American
material is this species.
CHELONANTHUS Gilg
Mostly annuals, the stems erect, usually simple, often angulate; leaves penni-
nerved; inflorescence a terminal once or twice bifurcately branched dichasium, the
FIG. 84. Chelonanthus alatus. A, habit of plant, X 1A; B, flower dissected
to show pistil and anthers, X 2; C, a larger leaf showing venation, X H.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 309
ultimate branches being few-many-flowered secund racemes; flowers green, yellow-
ish or pinkish; calyx cupular or campanulate, the five lobes imbricated; corolla
funnelform, the tube swollen ventrally, the limb 5-parted; style persistent, the
stigma bilamellate; capsule globose or ovoid, twice as long as the calyx or longer.
About 15 species in South America with one species ranging north
to Mexico. Species of this genus have often been included in Lisian-
thus but they seem to be adequately distinct. See Williams, Field-
iana: Bot. 31: 406. 1968.
Chelonanthus alatus (Aubl.) Pulle, Enum. Vase. PI. Surin. 376.
1906. Lisianthus alatus Aubl. Hist. PI. Guian. 1: 204, t. 89. 1775.
L. acutangulus Ruiz & Pavon, Fl. Peruv. 2: 14. 1799. L. tetragonus
Benth. PI. Hartweg. 68. 1840. Excacahue (Suchitepe"quez) ; taba-
quillo (Izabal).
Moist or wet thickets, often on open or brushy, steep banks,
2,000 m. or less; Alta Verapaz; Baja Verapaz; Izabal; Escuintla;
Suchitepe"quez; Quezaltenango; San Marcos. Southern Mexico;
British Honduras; Honduras; Nicaragua; Costa Rica and Panama.
South America.
A coarse erect glabrous herb, sometimes 3 m. high but usually about a meter
high or lower, generally simple, the stems sharply 4-angulate below; leaves remote,
sessile, ovate, elliptic-ovate, or very broadly ovate, mostly 5-15 cm. long, acute,
rounded to attenuate at the base; inflorescence large, terminal, corymbiform, com-
posed of few or numerous, one or twice dichotomously branched, secund racemes
20 cm. long or less, the flowers numerous, remote, on pedicels longer than the calyx;
the pedicels reflexed, at least in age; calyx green, 8 mm. long, lobate to the middle,
the lobes rounded-ovate, erose-denticulate; corolla pale green, 2 cm. long, some-
what fleshy; style 1 cm. long, complanate, persistent on the fruit; capsule oblong,
1.5 cm. long, somewhat compressed, narrowed at the apex; seeds very numerous,
minute, irregularly cubical.
A very common and characteristic plant of roadside banks at
middle elevations on the Pacific slope in Quezaltenango and San
Marcos. The green flowers are rather conspicuous but not at all
decorative or handsome.
COUTOUBEA Aublet
Erect glabrous herbs, probably annual, simple or branched, with strongly
ascending branches, the stems terete; leaves opposite or ternate, sessile and often
amplexicaul; flowers small, white, sometimes tinged with blue or purple, in dense
or interrupted, terminal spikes or racemes; calyx bibracteolate, deeply 4-(5-)fid,
the segments narrow, attenuate, scarious-marginate; corolla tube short-cylindric,
the 4 (-5) lobes spreading, narrow, contorted; stamens 4 (-5), affixed to the corolla
tube, the filaments filiform, dilated at the base; anthers oblong, erect, deeply bifid,
310 FIELDIANA: BOTANY, VOLUME 24
unchanged in age or finally recurved; ovary 1-celled, the placentae strongly in-
truded, the style filiform, the stigma bilamellate; capsule bivalvate; seeds numer-
ous, globose, foveolate-reticulate.
Three species, in tropical America. Only one is found in conti-
nental North America.
Coutoubea spicata Aubl. PI. Guian. 72, t. 27. 1775.
Wet savannas, 200 m. or less; Alta Verapaz; Izabal. Mexico
(Chiapas); British Honduras; Costa Rica; Panama; northern South
America.
Plants erect, a meter high or less, the stems simple below, with few or numer-
ous, strongly ascending branches above; leaves opposite, sessile and often amplexi-
caul at the base, oblong-obovate to narrowly lanceolate, 2.5 8 cm. long, obtuse to
attenuate at the apex, penninerved; flower spikes few or numerous, sometimes only
1, as much as 25 cm. long, the flowers very numerous, fragrant, white, often with
a bluish throat, sessile, more or less verticillate, the whorls crowded or rather re-
mote; calyx 6 mm. long, parted almost to the base, the segments linear-lanceolate,
erect; corolla persistent in fruit, the tube as long as the calyx, the lobes slightly
shorter, acute or acuminate, spreading or often reflexed; filaments equalling or
slightly longer than the corolla tube, the anthers 2.5 mm. long; capsule enclosed
in the costate persistent corolla tube.
CURTIA Chamisso & Schlechtendal
Reference: E. Knoblauch, Bot. Centralbl. 60: 356-358. 1894.
Very slender, erect, small annuals with almost filiform branches; leaves oppo-
site or 3-4-nate, small and narrow, sessile; flowers small, white, pink or yellow,
corymbose-cymose or laxly paniculate; calyx usually 5-parted, the segments nar-
row, acute, carinate; corolla subfunnelform, the tube cylindric, little ampliate
above, the limb 5-lobate, the lobes short, ovate or lanceolate, contorted; stamens 5,
affixed to the corolla tube, the filaments filiform, the anthers usually included,
ovate or oblong, often coherent, the connective broad or narrow; ovary falsely 2-
celled by the strongly intruded placentae, the style filiform or very short, the stigma
capitate, short-clavate, or dilated, not divided; capsule bivalvate; seeds numerous,
small, foveolate.
About ten species, mostly in northern South America, only the
following one in North America.
Curtia tenella (Mart.) Cham. Linnaea 8: 13. 1833. Schuebleria
tenella Mart. Nov. Gen. & Sp. 2: 117. 1827.
Dry rocky slopes, 1,200-1,500 m.; Chiquimula (Montana Cas-
tilla, 3 miles southeast of Quezaltepeque, Steyermark 31218}. British
Honduras; Honduras; Costa Rica; Panama; northern South America.
Plants very slender, 5-35 cm. high, usually simple below, corymbosely branched
above, glabrous; cauline leaves opposite, linear, 5-12 mm. long, usually spreading,
FIG. 85. Coutoubea spicata. A, habit of part of plant, X 1A; B, flower, X 4.
Eustoma exaltatum. C, habit, X 1A; D, dissected flower to show stamens and pistil.
311
312 FIELDIANA: BOTANY, VOLUME 24
the radical leaves cordate-ovate, acute; inflorescence dichotomously branched, few-
many-flowered; calyx 3.5-4 mm. long, the segments linear-lanceolate, attenuate;
corolla white or yellowish, 5-6 mm. long, the lobes much shorter than the slender
tube, ovate, subacute; capsule cylindric, rostrate by the persistent style, about
equalling the calyx.
One of the characteristic savanna plants of Costa Rica and
Panama.
EUSTOMA Salisbury
Erect herbs, usually or always annual, often more or less glaucous, glabrous;
leaves opposite, sessile and often amplexicaul; flowers large, long-pedunculate, blue,
purplish or white; calyx deeply 5-6-fid, the segments narrow, acuminate, carinate;
corolla campanulate, the tube short, the limb deeply 5-6-lobate, the lobes oblong
or obovate, contorted; stamens 5-6, affixed to the throat of the corolla, the fila-
ments filiform, the anthers oblong, versatile, finally recurved; ovary 1-celled, the
placentae little intruded; style filiform, usually persistent in fruit, the stigma
broadly bilamellate; capsule ovoid or oblong, bivalvate; seeds very numerous,
small, foveolate.
Three species, in temperate and tropical North America and
northern South America. A single species occurs in Central Amer-
ica, another in Mexico.
Eustoma exaltatum (L.) Salisb. Parad. Lond., t. 34. 1806.
Gentiana exaltata L. Sp. PL ed. 2. 331. 1762.
Quiche" (Jose Ignacio Aguilar 1270}. Southern United States;
Mexico; British Honduras; Nicaragua; West Indies; Colombia and
Venezuela.
An erect, somewhat glaucous annual, sometimes a meter high but usually
about 50 cm. or less, simple or often copiously branched, the stems terete; leaves
mostly oblong, often narrowly oblong or oblanceolate, 3-8 cm. long or larger, gen-
erally rounded or very obtuse and apiculate at the apex, sessile and more or less
amplexicaul, or the lowest leaves attenuate to the base; flowers few or numerous,
on very long, slender, erect peduncles, blue or pale purplish blue, 2-3.5 cm. long;
calyx 10-15 mm. long, the segments lance-linear, narrowly long-attenuate, united
only at the base; tube of the corolla somewhat shorter than the calyx, the lobes
longer than the tube, broad at the apex and abruptly contracted into a short tip,
or sometimes acuminate; capsule oblong or oval, 1.5 cm. long, rounded at the apex.
GENTIANA L. Gentian
Glabrous herbs, annual or perennial, various in habit; leaves opposite, sessile;
flowers small or large, axillary and terminal, sessile or rarely pedunculate, bibracte-
olate or naked, usually blue, violet or purple, sometimes white or yellow or even
red; calyx tubular, 5-fid or rarely 4-fid or 6-7-fid, winged, carinate, or naked, some-
times spathaceously cleft; corolla salverform, funnelform, tubular-campanulate, or
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 313
rarely subrotate, the tube often with hairs or scales in the throat, the lobes spread-
ing, contorted, usually 5, the sinuses often plicate-appendaged; stamens as many as
the corolla lobes, affixed to the tube and included or rarely exserted; anthers ovate,
oblong, or linear, muticous or the connective apiculate, connate or free, usually
extrorsely dehiscent, sometimes versatile; ovary 1-celled, the placentae parietal;
style short or almost none, the stigmas 2, laminar, spreading or revolute; capsule
sessile or stipitate, bivalvate; seeds numerous, sometimes winged.
About 300 species, mostly in arctic and temperate regions, in the
tropics confined to the higher mountains. Only the following are
known in Central America but G. adsurgens Cerv. ex Griseb. has
been found in nearby Chiapas and may be expected in Guatemala.
In North America the species are rather few, but they include some
of the most beautiful of flowers. In the South American Andes the
species are very numerous.
Flowers 6-12 mm. long; plants annual.
Flowers 8-11 mm. long; stems elongate, mostly 6-10 cm. long, not completely
covered by the leaves G. sedifolia.
Flowers about 6 mm. long; plants 2.5 cm. high or less, the stems completely
hidden by the leaves G. pumilio.
Flowers 25-35 mm. long; plants perennial.
Flowers 2.5 cm. long; calyx lobes 4-7 mm. long G. lewisiae.
Flowers 3.5 cm. long; calyx lobes about 10 mm. long G. guatemalensis.
Gentiana guatemalensis Standl. & Steyerm. Field Mus. Bot.
23: 75. 1944. L. Wms. Fieldiana, Bot. 31: 405. 1968.
Moist alpine meadows, 3,300 m.; endemic; Huehuetenango (type
from region of Che'mal, Sierra de los Cuchumatanes, Standley 81113;
known only from this region).
Perennial from a thick woody caudex, the stout stems procumbent, numerous,
5-15 cm. long, 1-3-flowered, densely leafy; leaves spreading, sessile, oblong, 1.5-
2.5 cm. long, 4-6 mm. broad, obtuse or narrowly rounded at the apex, slightly
narrowed at the base, without obvious nerves, minutely scaberulous on the mar-
gins; flowers terminal or subterminal, sessile, 3.5 cm. long, purplish blue; calyx tube
obconic, 8 mm. long, the 5 lobes unequal, narrowly lance-oblong, about 10 mm.
long, subacute, scaberulous on the margins; corolla campanulate, 2.5 cm. long, the
lobes rounded-ovate, almost 1 cm. long, rounded at the apex, the plicae much
shorter than the lobes, bilobate, the lobes triangular, acute or acuminate, irregu-
larly and sparsely serrate.
This and the following species, G. lewisiae, are perhaps too much
alike and the distinctions tend to disappear with additional collections.
Gentiana lewisiae Standl. & Steyerm. Field Mus. Bot. 23: 76.
1944.
314
FIELDIANA: BOTANY, VOLUME 24
Wet meadows, 3,100-3,200 m.; endemic; Totonicapan (type from
mountains above Totonicapan, Standley 84-556).
FIG. 86. Gentiana lewisiae. A, natural size; B, dissected flower, natural size
Gentiana pumilio. C, habit, X 2; D, dissected corolla, X 2; E, ovary and styles,
X 3.
A prostrate perennial, the stems slender, 6-13 cm. long, simple, 1-few-flowered;
leaves sessile, spreading, oblong, 8-13 mm. long, 4-5 mm. broad, very obtuse or
rounded at the apex, subacute at the base, 1-costate, slightly paler beneath; flower
terminal, sessile, subtended by 2 leaves; calyx tube narrowly obconic, 5-6 mm. long,
the 5 lobes oblong or narrowly spatulate-oblong, 4-7 mm. long, subacute, erect;
corolla deep blue, campanulate, 2-2.5 cm. long, the lobes 3 mm. long, rounded-
ovate, very obtuse, the plicae much shorter than the corolla lobes, shallowly bilo-
bate, the lobes rounded, entire.
The species was named for Mrs. B. E. Lewis of Guatemala, a
devoted student of the plants of Guatemala, who has contributed
a great deal to our present knowledge of them.
The plant is a common one in most wet meadows on the moun-
tain summits of the Sierra Madre during December and January.
Gentiana pumilio Standl. & Steyerm. Field Mus. Bot. 23: 76.
1944.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 315
Moist subalpine slopes or meadows, 3,000-4,100 m.; Totonicapan;
Huehuetenango; San Marcos (type from Volcan de Tajumulco, Stey-
ermark 354.89). Mexico (Chiapas, Volcan de Tacana).
A dwarf annual, usually only 1-2 cm. high, simple or densely branched at the
base, the stems forming compact subglobose tufts, 1-few-flowers, very densely
leafy, the leaves usually closely imbricate and concealing the stems; radical leaves
about 4, much larger than the others, oblong-obovate, as much as 15 mm. long
and 6 mm. broad, obtuse or subacute; cauline leaves highly variable in size and
shape, the lower ones broadly obovate, the upper much smaller ones broadly ovate
or almost reniform and amplexicaul, all the cauline leaves broadly white-marginate,
2-7 mm. long, carinate, broadly obtuse and often mucronate; flowers sessile among
the leaves, or sometimes short pedicellate, 6-7 mm. long; calyx almost tubular, the
5 teeth oblong, short, very obtuse, separated by rather wide sinuses, the margins
minutely scaberulous, the teeth erect and subequal; corolla pale blue or white,
funnelform, the limb rotate, the lobes short, rounded-ovate or obovate, 3.5 mm.
long, rounded at the apex, the plicae shorter than the corolla lobes, emarginate or
shallowly bilobate; capsule narrowly oblong, protruded above the leaves, 6 mm. long.
This is probably the most plentiful of the Guatemalan gentians,
a diminutive plant that is hard to find except when its flowers are
fully open in bright sunshine. They at first remind one of tiny daisies
because of their much lobed limb, which is of a rather pale sky-blue
or white. The corolla is greenish outside, and has a pale yellow throat.
Plants have been found in flower in December, January, February
and March. It may be that G. pumilio is only an extreme form of
G. sedifolia, but the 12 collections at hand are fairly uniform, and
they are not matched by any South American material we have seen.
Gentiana sedifolia HBK. Nov. Gen. & Sp. 3: 173, t. 225. 1819.
Along rivulets in subalpine meadow, 2,500-3,700 m.; Huehuete-
nango; Totonicapan. Costa Rica; Andes of South America.
Annual or possibly sometimes perennial, usually much branched and bearing
numerous flowers, procumbent or prostrate, the stems as much as 10 cm. long,
densely or sparsely leafy; leaves sessile, subconnate, lanceolate, oblong-lanceolate,
or oblanceolate, acute or obtuse, 1-nerved, usually narrowed to the base, mostly
4-6 mm. long, white-marginate; flowers terminal at the ends of the branches, soli-
tary, erect, about 11 mm. long, purple-blue; calyx tubular-campanulate, the 5 lobes
linear-lanceolate, very short, erect, subequal; corolla funnelform, the limb almost
equally 10-lobate, the corolla lobes rounded-ovate, obtuse, the plicae rounded at
the apex; capsule oblong, acute, protruded above the leaves, bivalvate.
On general principles it might be expected that the Guatemalan
plant would constitute a distinct species, but good characters for
separating it from the highly variable South American plants have
not been discovered.
316 FIELDIANA: BOTANY, VOLUME 24
HALENIA Boerckhausen
Reference: Caroline K. Allen, A monograph of the American spe-
cies of the genus Halenia, Ann. Mo. Bot. Card. 20: 119-220, tt. 8-12.
1933.
Glabrous herbs, annual or perennial, simple or branched; leaves opposite, ses-
sile or petiolate, usually 3-5-nerved ; inflorescence terminal or axillary, cymose and
usually subumbellate, rarely racemose or spicate; calyx 4-lobate, the lobes folia-
ceous, linear to ovate or spatulate; corolla 4-lobate, white, green, yellow or purplish,
persistent, campanulate, the lobes dextrorsely convolute, often auriculate; stamens
4, included, adnate to the corolla tube, the filaments linear, the anthers ovate or
oblong, versatile; stigma sessile, composed of 2 oblong or ovate lobes; capsule com-
pressed, lanceolate to ovate, septicidally dehiscent from the apex; seeds globose or
somewhat compressed, brown or greenish tan-colored, granular or reticulate.
About 60 species, mostly in America, chiefly in the mountains of
the tropics, a few species in Asia. Several other species are known
in Mexico and southern Central America.
Corolla not calcarate at the base, merely obscurely saccate.
Leaves mostly basal, the stems scape-like; plants perennial H. alata.
Leaves mostly cauline, the stems not scape-like; plants annual . . .H. brevicornis.
Corolla conspicuously calcarate at the base, the spurs short or elongate.
Spurs of the corolla very short and somewhat spreading; plants mostly 10 cm.
high or less and usually with numerous, stout branches from the base, form-
ing dense clumps of stems H. crassiuscula.
Spurs usually well developed and directed downward or even incurved.
Stems scapiform, the cauline leaves few and reduced; radical leaves numerous,
from a rather thick, perennial caudex H. shannonii.
Stems leafy, the cauline leaves numerous pairs; radical leaves few or none.
H. decumbens.
Halenia alata (Mart. & Gal.) Hemsl. Biol. Centr. Amer. Bot. 2:
351. 1882. Exadenus alatus Mart. & Gal. Bull. Acad. Brux. 11, pt. 1:
372. 1844.
Subalpine meadows, frequently in open Juniperus forest, 3,200-
3,500 m.; Huehuetenango (Sierra de los Cuchumatanes) ; Mexico
(Veracruz, Volcan de Orizaba).
Perennial from a thick caudex, the stems several, erect, 13-22 cm. high; radical
leaves numerous, 5 cm. long or less, 3-4 mm. broad, obtuse, long-attenuate to the
slender petiole, 3-nerved; cauline leaves few pairs, linear or oblanceolate-linear;
flowers terminal, umbellate, the pedicels 4-angulate; calyx slightly shorter than the
corolla, the lobes oblong, acute; corolla pale yellow, small, about 6 mm. long, not
calcarate, the lobes ovate, obtuse; capsule broadly ovoid, 6-7 mm. long; seeds
yellow-brown, globose, granular.
An unusually well marked species, or perhaps appearing so be-
cause little material of it has been collected.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 317
Halenia brevicornis (HBK.) G. Don, Gen. Hist. 4: 177. 1838.
Swertia brevicornis HBK. Nov. Gen. & Sp. PL 3: 174. 1818. H. tuerck-
heimii Briq. Candollea 4: 317. 1931 (type from Alta Verapaz,
Tuerckheim 204.1). H. brevicornis var. tuerckheimii Allen, Ann. Mo.
Bot. Gard. 20: 145. 1933.
Brushy or grassy slopes, often in pine-oak forest, sometimes in
moist subalpine meadows, 1,000-3,200 m.; Alta Verapaz; El Pro-
greso; Zacapa; Jalapa; Guatemala; Sacatepe"quez ; Chimaltenango;
Quich^ ; Huehuetenango; Quezaltenango. Central and southern
Mexico; Honduras; Nicaragua; Costa Rica.
A slender annual, generally 15-45 cm. high, the stems simple below, usually
with a few branches above; leaves all or mostly cauline, linear or lanceolate, 3 cm.
long and 4 mm. broad or smaller, 1-nerved, the upper leaves shorter; inflorescence
racemiform or narrowly paniculate, with usually several whorls of flowers, the pedi-
cels elongate or short; calyx segments lanceolate, one-half to two-thirds as long as
the corolla; corolla green or greenish yellow, 5-8 mm. long, the lobes broadly or
narrowly ovate, acute, somewhat auriculate at the base, not calcarate; capsule
ovoid, 8 mm. long; seeds minute, brownish, finely reticulate.
Allen recognizes seven varieties of H. brevicornis in Mexico and
Central America, but they appear to have slight if any systematic
importance. Var. tuerckheimii is noteworthy for its slender stems
and elongate pedicels. This is the common unspurred species in the
highlands.
Halenia crassiuscula Robinson & Seaton, Proc. Am. Acad. 28:
113. 1893.
Moist or dry, open, rocky wind-swept alpine summits, sometimes
in alpine pine forest, 3,700-4,200 m. ; Quezaltenango (Volcan de Santa
Maria); San Marcos (Volcan de Tajumulco; Volcan de Tacana).
High volcanoes of central and southern Mexico.
Plants densely cespitose, dwarf, perennial or biennial, somewhat fleshy, the
stems numerous or very numerous, mostly 5-10 cm. high, stout; radical leaves
oblanceolate, 3-nerved, 2 cm. long, obtuse, attenuate to the petiole; cauline leaves
1-3 pairs, narrowly oblanceolate; inflorescence dense and compact, umbelliform,
the flowers mostly terminal, sometimes axillary, pedicellate, often nutant after an-
thesis; calyx lobes lanceolate or oblanceolate, 4.5-6 mm. long, obtuse; corolla
15 mm. long or shorter, the lobes oblong-elliptic, acute, the spurs 4 mm. long,
slender, slightly curved upward; capsule lance-oblong, acute, 14 mm. long; seeds
globose, light yellow-brown, granular.
This reduced plant is common about the summits of the volca-
noes of western Guatemala. Most of the plants are dry and dead
during the winter months, when we have visited those localities, but
318
FIELDIANA: BOTANY, VOLUME 24
the withered stems with the persistent capsules often are seen in
abundance.
FIG. 87. Halenia crassiuscula. A, habit, natural size; B, flower partially dis-
sected, X 3; C, another flower from the same collection to show variation, X 2.
Halenia decumbens Benth. PI. Hartw. 67. 1840. H. guatema-
lensis Loes. Verhandl. Bot. Vereins Brandenb. 55: 182. 1913 (type
Huehuetenango, Seler 2728). H. plantaginea var. latifolia Loes. I.e.
(type from Huehuetenango, Seler & Seler 3086). H. guatemalensis
var. latifolia Allen, Ann. Mo. Bot. Card. 20: 180. 1933. H. caleoides
Allen, I.e. 173 (type from Sacatepe'quez, Maxon & Hay 3675). H.
platyphylla Allen, I.e. 173 (type from Sacatepe'quez, Smith 2170).
Common in open moist forests and in the subalpine meadows of
the highlands 2,300-4,200 m.; Sacatepe'quez ; Chimaltenango; Solola;
Quiche* ; Huehuetenango; Totonicapan ; Quezaltenango; San Marcos.
Mexico (whence the type).
Plants erect or decumbent, perennial or apparently often annual or biennial,
mostly 20-40 cm. high, the stems simple or often sparsely branched, slender, bear-
ing usually numerous pairs of thin leaves; basal leaves oblanceolate, 3-6 cm. long,
mostly 1-1.5 cm. broad, acute and often apiculate; cauline leaves mostly 3-nerved,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 319
obtuse or acute, the upper ones elliptic-lanceolate or lanceolate, sessile; flowers ter-
minating the stems or branches, umbellate, the pedicels 3 cm. long or often much
shorter; calyx lobes obovate-spatulate, two-thirds as long as the corolla, mucro-
nate; corolla pale green or yellow-green, 12-20 mm. long, the lobes ovate, subacute,
the spurs about half the length of the whole corolla or shorter, incurved at the tip ;
capsule ovoid, 15-18 mm. long; seeds globose, granular.
This is common Halenia of the highlands of Guatemala that has
the corollas obviously spurred. There is considerable variation in
the species but it is easy to recognize. We have not followed Miss
Allen's treatment for we feel that both the keys and the systematic
account are quite unrealistic.
Halenia shannonii Briq. Candollea 4: 321. 1931. H. shannonii
f. compacta Allen, Ann. Mo. Bot. Card. 20: 178. 1933 (type from
Huehuetenango, Cook |5).
Moist subalpine meadows or open, pine or Juniperus forest, some-
times on limestone, 2,000 (?) -4,000 m.; Sacatepe"quez (type, Shannon
3613 or f3630}; Totonicapan; Huehuetenango; San Marcos; so far as
known, endemic.
Plants perennial from a rather thick caudex, the stems cespitose, mostly 12-
20 cm. high, erect, usually simple; basal leaves somewhat fleshy, narrowly oblan-
ceolate, 7 cm. long and 3-6 mm. broad or smaller, acute or obtuse, 3-nerved; cauline
leaves 2-3 pairs or none, oblanceolate to linear-lanceolate, acute, sessile; inflores-
cences mostly terminal, several-flowered, umbelliform, the pedicels 2.5 cm. long or
much shorter; calyx one-half to two-thirds as long as the corolla, the lobes oblong-
elliptic, 6-9 mm. long; corolla greenish, almost 15 mm. long, the lobes ovate, obtuse
or subacute, the spurs divaricate, slightly incurved at the tip.
This has been reported from Guatemala as H. gracilis Griseb. It
probably represents little more than small specimens of H. decumbens.
LEIPHAIMOS Schlechtendal & Chamisso
Small saprophytic herbs, without chlorophyll, simple-stemmed, 1 -several-flow-
ered, the leaves reduced to opposite scales; flowers small or rather large, mostly
white, yellow, blue or purple, solitary or in cymes; calyx campanulate or almost
tubular, with 4-5 teeth or lobes; corolla salverform or funnelform, the lobes spread-
ing, obtuse or acute, contorted, persistent in fruit; stamens 4-5, inserted in the
upper half of the tube, included, the filaments filiform, usually very short; anthers
oval, cordate, or elongate, free or connivent by their margins, introrse, obtuse or
acute at the base, sometimes appendaged; ovary 1-celled, with 2 parietal placentae,
the style filiform, the stigma capitate; capsule usually elongate, septicidally dehis-
cent by slits between (but not reaching to) the top and bottom; seeds fusiform or
winged; endosperm very scant.
About 30 species, mostly in tropical America, two in tropical
Africa. About six additional ones are known from southern Central
320 FIELDIANA: BOTANY, VOLUME 24
America. The following list includes all species known from Mexico.
The plants are often rather handsome, in spite of their usually small
size, for the flowers are conspicuously colored. In habit they remind
one much of the Burmanniaceae of similar habit.
Stems terminated by a cyme of several flowers L. parasitica.
Stems, or their branches, each terminated by a single flower.
Corolla blue, about 1.5 cm. long L. simplex.
Corolla yellow, about 3 cm. long L. aphylla.
Leiphaimos aphylla (Jacq.) Gilg, Nat. Pflanzenfam. IV. 2: 104.
1895. Gentiana aphylla Jacq. Enum. PI. Carib. 17. 1760.
Wet mixed forest, 1,500 m. or less; Izabal (Punta Palma, across
the bay from Puerto Barrios); Chiquimula (Cerro Tixixi, north of
Jocotan). Mexico (Oaxaca); Costa Rica(?); West Indies; South
America.
Stems simple, very slender and weak, white; leaf scales few pairs, remote and
very inconspicuous, linear-lanceolate, narrowly attenuate; calyx almost tubular,
about 5 mm. long, the lobes narrow, attenuate; corolla tube very slender, yellow
or whitish, about 25 mm. long, the lobes bright yellow, 5 mm. long, acute or acumi-
nate; capsule narrowly oblong, 10-15 mm. long.
It is probable that L. lutea Morton, described from Costa Rica,
is synonymous with L. aphylla.
Leiphaimos parasitica Schlecht. & Cham. Linnaea 6: 387. 1831.
Voyria mexicana Griseb. Gen. & Sp. Gent. 208. 1838.
Among decaying leaves in wet mixed forest, 350 m.; Pete"n; Alta
Verapaz; Izabal. Southern Mexico; British Honduras; Nicaragua.
Plants very slender, white throughout, 6-20 cm. high, the stems weak; leaf
scales few pairs, very small and inconspicuous, acute or obtuse, more or less ap-
pressed; flowers small, few or numerous, in a compact small terminal cyme, this
usually bifurcate from the base, with a single flower between the branches, the
white flowers secund along the branches cf the cyme, sessile or nearly so, about
9 mm. long, the calyx narrow, 3 mm. long, with narrow acute lobes; corolla lobes
scarcely 1 mm. long, rounded; capsule ellipsoid, 5-6 mm. long.
Leiphaimos simplex (Griseb.) Standl. Contr. U. S. Nat. Herb.
20: 199. 1919. Voyria simplex Griseb. in Seem. Bot. Herald 170.
1854.
Wet forest, among rotting leaves, 1,000 m. or less; Alta Verapaz;
Izabal; Huehuetenango. British Honduras; Costa Rica; Panama.
Plants very slender and delicate, the stems white, mostly 6-15 cm. long, 1-
flowered; leaf scales very small and inconspicuous, closely appressed to the stem,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 321
few; calyx narrow, 3 mm. long, appressed to the corolla base, the lobes narrow,
entire; corolla pale blue, the tube 8 mm. long, the lobes spreading, narrow, obtuse,
3-4 mm. long; capsule 5 mm. long.
LISIANTHUS L.
Reference: J. Perkins, Monog. Ubersicht der Gattung Lisianthus,
Bot. Jahrb. 31: 489-498. 1902; Louis 0. Williams, Fieldiana, Bot.
31:406-411. 1968.
Erect, often rather tall, tricotomously, diffusely branched herbs, occasionally
suffrutescent. Leaves reduced upward but present in the inflorescences, often with
connate bases similar to interpetiolar stipules; inflorescence essentially thrysoid, or
dichasioid, sometimes tending to be scorpioid; flowers small or large, mostly yellow
but sometimes almost black; calyx 5-lobate, the lobes longer than the short tube;
corolla funnelform or salverform, the 5 lobes spreading; stamens 5, inserted on the
lower part of the corolla tube, the filaments filiform, the anthers oblong; ovary
1-celled, the style filiform, the stigma subcapitate; capsule enclosed in the persist-
ent corolla, bivalvate; seeds commonly tuberculate or muricate.
About 15 species in Mexico, Central America and the West Indies
with one species extending to Colombia, South America. Most South
American species referred to this genus belong in Chelonanthus, or to
other related genera.
Flowers single [rarely 2-3] in the axils of leaves; corolla dull red L. axillaris.
Flowers few to many in the axils of reduced leaves or bracts; corolla yellow, red
or "black."
Corolla lobes widely flaring; flowers less than 20 mm. long; flowers probably
yellow or reddish L. saponarioides.
Corolla lobes mostly erect, not flaring; flowers 25 mm. or mostly much longer.
Calyx lobes ovate, acute or obtuse.
Corolla lobes obtuse; corolla 3-3.5 cm. long; leaves sessile . . . L. viscidiflorus.
Corolla lobes acute; corolla 4-6 cm. long; leaves petiolate L. skinneri.
Calyx lobes linear to lanceolate, acuminate.
Corolla 3-4.5 cm. long, the lobes long acuminate; flowers "black."
L. nigrescens & var. cuspidatus.
Corolla 2.5-4 cm. long, the lobes acuminate; flowers yellowish or greenish.
Corolla lobes about 5 mm. long L. brevidentatus.
Corolla lobes about 10 mm. long L. auratus.
Lisianthus auratus Standl. Trop. Woods 37: 29. 1934.
Mixed woods and pine savannas, 50-1,000 m. Chiquimula. Brit-
ish Honduras; Honduras (type from Siguatepeque, Edwards P-556);
Nicaragua.
Annual herbs to about 2.5 m. tall, sometimes suffrutescent, terete or obscurely
angles above, nearly simple or often profusely tricotomously branched, branching
is indeterminate; leaves elliptic to elliptic-lanceolate or lanceolate-ovate, acumi-
322 FIELDIANA: BOTANY, VOLUME 24
nate, with 2 pairs of lateral nerves, membranaceous, 2-9 cm. long and 1-2.5 cm.
broad, short petiole 3-5 mm. long, clasping the stems and joined; inflorescence a
single axillary flower to axillary few-flowered cymes, these sometimes short pedunc-
ulate, the pedicels 6-16 mm. long; flowers yellow, subdiaphanous; calyx divided
to near the base, the lobes linear-lanceolate, acute, about 4-8 mm. long; corolla
constricted above the ovary then expanding and tubular above, about 25-35 mm.
long, the lobes lanceolate-ovate, acuminate, about 5-10 mm. long; stamens ex-
serted, the anther oblong-ovate, cordate, about 2 mm. long; style as long as or
longer than the corolla, the stigma capitate; capsule narrowly ovoid, vernicose, 6-
12 mm. long at maturity.
The species is known from two collections from British Honduras
and one from Guatemala. The collections known from Nicaragua
are with smaller flowers than others. The areolar markings in the
membranaceous leaves, due to the veinlets, are distinctive.
Lisianthus axillaris (Hemsl.) 0. Kuntze, Rev. Gen. PI. 2: 429.
1891 (type British Honduras, Barlee); L. francisiae Sprague, Kew
Bull. 1929: 8. 1929 (type British Honduras, Francis 1).
Often in pine savannas or in forest clearings, usually at 600 m.
or lower; Pete"n. British Honduras where apparently frequent.
An erect herb, or sometimes suffrutescent below, stem simple or branched,
slender; leaves sessile or short-petiolate, lance-ovate to elliptic-lanceolate or ob-
long-lanceolate, 5-11 cm. long, 2-4 cm. broad, long-acuminate, acute to somewhat
rounded at the base, the lateral nerves generally 2 pairs; flowers axillary, solitary
in each axil or very rarely in 2-3-flowered dichasia, the pedicels mostly 1.5-2 cm.
long; calyx segments lance-linear, 9-10 mm. long, long-attenuate, carinate; corolla
3.5-4 cm. long, the tube very slender, the throat abruptly dilated, 5-6 mm. broad,
dull red, the lobes ovate-oblong, caudate-acuminate, green, 8-10 mm. long; sta-
mens slightly longer than the corolla tube but shorter than the lobes; style about
equalling the corolla lobes; capsule ellipsoid-oblong, 1 cm. long.
Called "red chilar" in British Honduras.
Lisianthus brevidentatus (Hemsl.) 0. Kuntze, Rev. Gen. PL 2:
429. 1891. Leianthus brevidentatus Hemsl. Biol. Cent. Am. Bot. 2: 344.
1882 (type from Izabal, Bernoulli 921>). Lisianthus quichensis Donn.-
Sm. Bot. Gaz. 52: 51. 1911 (type from Quiche", Heyde & Lux 2921).
Lisianthus collinus Standl. Carnegie Inst. Wash. Publ. 461: 81. 1935
(type from British Honduras, Schipp 1205}. Lisianthus calciphilus
Standl. & Steyerm. Field Mus. Bot. 22: 267. 1940 (type from Alta
Verapaz, Wilson 356). L. elatus Standl. & Steyerm. I.e. (type from
Izabal, Steyermark 38562). Lisianthus petenensis Standl. & Steyerm.
Bull. Torr. Bot. Club 84: 46. 1957 (type from Pete"n, Lundell 3153).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 323
Banks, grassy slopes and open woods, mostly 500-1,500 m.; Pe-
te"n; Alta Verapaz; Chiquimula; Huehuetenango. Mexico (Chiapas) ;
British Honduras.
Simple or usually diffusely branched herbs, the stems stout, terete, to 1.5 m.
tall. Leaves sessile to subamplexicaul on the lower part of the stem, elliptic to
lanceolate-oblong, the upper ones attenuate to the base, the lower ones broader at
the base and subamplexicaul, acute to long acuminate, usually with two pairs of
lateral nerves, principal ones 9-15 cm. long and 2-4 cm. broad, much reduced and
becoming bract-like above; inflorescence few-flowered to many-flowered, diffusely
branched, densely flowered to open few-flowered dichasia or cymes; flowers yellow-
green or yellow, borne on short (2 mm.) to long (30 mm.) pedicels; calyx deeply
5-lobed, the lobes lance-attenuate, acuminate, the margins scarious, 6-10 mm. long;
corolla 27-40 mm. long, the tube strong costate at the base and marcescent in age,
the throat 4-6 mm. broad, lobes erect, ovate to broadly ovate, acuminate and apic-
ulate, 2.5-5 mm. long; stamens about as long as or slightly exceeding the corolla;
stigma globose; capsule ovoid to subglobose, vernicose, apparently viscid.
We have not seen authentic material of L. brevidentatus but the
description indicates that it is this most common of the Guatemalan
species. The several synonyms indicated above seem to be based
mostly on growth forms. The plant when it begins to flower is quite
a simple one, becoming diffusely branched and coarse with age. There
are also differences in the compactness of the inflorescences as well
as differences in the size of the flowers and especially in length of the
corolla lobes. Monographic studies may indicate a more liberal view
of these plants.
Lisianthus nigrescens Schlecht. & Cham. Linnaea 6: 388. 1831.
Leianthus nigrescens Griseb. Gen. & Sp. Gent. 199. 1838.
On limestone along river (Cook 65). Huehuetenango. Eastern
and southern Mexico.
Coarse, erect herbs, usually about a meter high, branched above, the stems
terete; leaves sessile, lance-oblong to narrowly lanceolate, broad and somewhat
amplexicaul at the base or cuneately narrowed to the base; inflorescence cymose-
paniculate, large and many-flowered, open, the bracts leaf-life but reduced, the
flowers on short or much elongate pedicels, these usually recurved in age; calyx
green, 8-10 mm. long, the segments lanceolate, long-attenuate; corolla purple-
black, 3-3.5 cm. long, cuspidate in bud, the tube 3-4 mm. broad near the apex, the
lobes 7-10 mm. long, cuspidate-acuminate; capsule ellipsoid, 10-12 mm. long, very
lustrous.
A most unusual plant, notable for its almost black corollas — a
yellow-flowered variety occurs in Chiapas, Mexico.
Lisianthus nigrescens var. cuspidatus (Bertoloni) L. Wms.
Fieldiana, Bot. 31: 408. 1968. Lisianthus cuspidatus Bertoloni,
324 FIELDIANA: BOTANY, VOLUME 24
Comm. Acad. Bonon. 4: 408, t. 37. 1840 (type collected in Guate-
mala, the locality not indicated but possibly between Esquintla and
Guatemala, Velasquez).
Open dry or moist slopes or plains, 1,000-1,800 m.; El Progreso
(Volcan Siglo) ; Guatemala; Chimaltenango; Quiche" ; Huehuetenango.
Mexico.
Similar to the species but the flowers larger and more showy. Corolla purple-
black, 4-5.5 cm. long, the tube 5-6 mm. broad at the throat, the lobes 12-18 mm.
long, long attenuate or cuspidate-acuminate; capsule ellipsoid, about 12 mm. long.
The flowers are the nearest to black of any known to us in Guate-
mala. The plant is a handsome one when in full flower.
Lisianthus saponarioides Schlecht. & Cham. Linnaea 6: 389.
1831. Leianthus saponarioides Griseb. Gen. & Sp. Gent. 198. 1838.
Petasostylis saponarioides Griseb. in DC. Prodr. 9: 71. 1845. Lisian-
thus meianthus Donn.-Sm. Bot. Gaz. 52: 51. 1911 (type from Alta
Verapaz, Tuerckheim 1436). Lisianthus congestus Standl. Carnegie
Inst. Wash. Publ. 461: 82. 1935 (type from Pete"n, Lundell 2479).
Apparently a plant of the savannas ; Pete"n ; Alta Verapaz. British
Honduras; Mexico.
An erect herb, the stems branched, terete; leaves short-petiolate, ovate or
lance-ovate, 2.5-4 cm. long, 1-1.5 cm. broad or larger, acuminate or long-acumi-
nate, obtuse or somewhat rounded at the base, the lateral nerves obsolete; flowers
yellow, in dense head-like cymes, these terminating short branches, subtended at
the base by normal leaves, very dense and many-flowered, sessile or nearly so;
calyx 8 mm. long, the segments lance-linear, long-attenuate; corolla tube slender,
striate, 9-10 mm. long, the lobes spreading, ovate, cuspidate-acuminate, 5 mm.
long; stamens and style short-exserted ; capsule oblong, 7 mm. long.
Perkins referred to this species the collection later made the type
of L. quichensis Donn.-Sm., but that, in its open lax inflorescence,
does not at all agree with the original description of L. saponarioides.
Of that we have seen no authentic representation, but its description
agrees fully with the plant described as L. congestus. Since the later
plant comes from a region far removed from Veracruz, it is still pos-
sible that it may be a valid species, but by description alone the two
described species cannot be separated.
Lisianthus skinneri (Hemsl.) 0. Kuntze, Rev. Gen. PL 2:
429. 1891; L. Wms. Fieldiana, Bot. 31: 411. 1968. Leianthus skin-
neri Hemsl. Biol. Cent. Am. Bot. 2: 345. 1882 (type: Guatemala,
Skinner). Lisianthus arcuatus Perkins in Engler, Bot. Jahrb. 31:
492. 1902.
FIG. 88. Lisianthus nigrescens var. cuspidatus. A, portion of upper part of
plant showing method of branching in inflorescence, X M; B, a leaf from lower
part of stem, X Yi', C, ultimate part of inflorescence showing a flower, natural size;
D, corolla dissected to show stamens, X 11A; E, calyx, X 3; F, pistil, X 2; G, stig-
ma, X 20.
325
326 FIELDIANA: BOTANY, VOLUME 24
Edges of clearings or in wet forests, from near sea level in north
of range to about 1,000 m. in the south. "Guatemala." Honduras;
Costa Rica; Panama. To be expected in Nicaragua.
Herbs to 1.5 m. tall or perhaps more, the stems sometimes suffruticose. Leaves
oblong-lanceolate to narrowly obovate, acute to acuminate, up to 25 cm. long and
10 mm. broad and rather long petiolate (to 6 cm.), reduced upward and those of
the inflorescence nearly sessile, usually with two pairs of prominent lateral nerves;
inflorescence corymbose, mostly with ten or fewer flowers; flowers yellow, the pedi-
cels 2 cm. long or the flowers nearly sessile; calyx 5-7 mm. long, tubular below, the
lobes 2-4 mm. long, ovate-lanceolate to ovate, acute; corolla tubular, slightly arcu-
ate, 4-6 cm. long, the lobes ovate, acute, 3-4 mm. long; stamens equalling the
corolla or slightly exserted; style exserted in mature flowers.
Lisianthus viscidiflorus Robinson, Proc. Am. Acad. 45: 398.
1910.
Open forest or brushy hillsides, on limestone, 200-1,400 m.; Alta
Verapaz (type from Coban, Tuerckheim 11.1308); collected at vari-
ous other localities in this department. Mexico (Oaxaca).
A tall and rather coarse annual, about a meter high or sometimes taller, the
stems terete, simple below, branched above; leaves sessile and amplexicaul, lance-
oblong, 7-12 cm. long and 1-2.5 cm. broad or often considerably larger, acuminate,
the lateral nerves 2 pairs; flowers numerous and forming a very large, open, much-
branched panicle, most of the bracts much reduced and lanceolate to subulate, the
dull red flowers on short or much elongate pedicels, very viscid ; calyx lobes ovate,
obtuse or acute, 4-7 mm. long; corolla 3-3.5 cm. long, the tube dull red, the lobes
green with purple at the base, the lobes broad, obtuse or rounded at the apex, about
5 mm. long, erect; stamens included, the style short-exserted.
NYMPHOIDES Seguier
Aquatic perennial herbs with rootstocks; leaves on greatly elongate petioles,
broadly ovate or orbicular, cordate at the base, entire or repand; flowers small,
yellow or white, umbellate at the ends of the stems or axillary; calyx 5-parted;
corolla subrotate, deeply 5-lobate, the lobes induplicate-valvate in bud, often with
fimbriate margins; stamens 5, inserted on the base of the corolla, the anthers sagit-
tate, versatile; ovary 1-celled, the style short or none, the stigma bilamellate;
capsule indehiscent or irregularly ruptured.
About 20 species, in tropical and temperate regions of both hemi-
spheres. Only one is found in Central America.
Nymphoides humboldtianum (HBK.) Kuntze, Rev. Gen. PL
2: 429. 1891. Villarsia humboldtiana HBK. Nov. Gen. & Sp. 3: 187.
1818. Limnanthemum humboldtianum Griseb. Gen. & Sp. Gent. 347.
1838. Cebolla de agua; Corazon de agua.
FIG. 89. Nymphoides humboldtianum. A, habit, X 1 ; B, calyx and ovary, X 3;
C, flower dissected, X 2; D, anthers, X 8; E, cross-section of ovary, X 8. B-E,
after Gilg in Engler, Pflanzenf. IV, 1: 197, fig. |«. 1891.
327
328 FIELDIANA: BOTANY, VOLUME 24
Floating on ponds or lakes or sometimes submerged or on mud,
2,000 m. or less; Pete'n; Izabal; Chiquimula; Jalapa; Jutiapa; Santa
Rosa; Esquintla; Quiche"; Huehuetenango ; probably also in other
departments. Mexico; British Honduras to Panama. West Indies.
South America.
Plants glabrous, with elongate rootstocks, fleshy, the stems stout, often spongi-
ous, mostly 40 cm. long or shorter; leaves solitary, orbicular or orbicular-reniform,
3-12 cm. broad, deeply cordate at the base; flowers white or yellow, in a sessile
umbel at the base of the petiole, the pedicels few or numerous, very unequal, 3-
10 cm. long, deflexed in fruit; calyx segments linear-lanceolate, 8 mm. long; corolla
lobes fimbriate, twice as long as the calyx, recurved; capsule somewhat shorter than
the calyx; seeds numerous, smooth, globose.
The plants sometimes are found upon mud, where the water has
receded. The leaves are often purplish beneath. In Guatemalan
plants the flowers are either white or yellow. The plants of this
alliance sometimes are placed in a separate family Menyanthaceae.
SCHULTESIA Martius
Erect annuals; leaves opposite, sessile or nearly so; flowers large or small, usu-
ally pink or white; calyx tubular, 4-costate or often 4-winged, 4-lobate, the tube
as long as the lobes or longer; corolla funnelform, the tube narrowed upward, the
limb 4-lobate, the lobes contorted in bud; stamens 4, inserted on the corolla tube,
the anthers oblong; ovary 1-celled, the style filiform, the stigma 2-lamellate; cap-
sule bivalvate; seeds small, foveolate.
About 20 species, one in tropical Africa, the others in tropical
America.
The genus Schultesia is a small one in Central America. The
names used here, except S. peckiana, are all based upon South Amer-
ican types and most of them have extensive ranges. We have fol-
lowed traditional usages here which is the best that we can do until
such time as a revision is written of the genus.
Calyx 5-10 mm. long, not winged or very obscurely so.
Calyx at an thesis 9-10 mm. long; flowers all or nearly all on slender pedicels
8-30 mm. long, the inflorescence lax and usually many-flowered.
S. peckiana.
Calyx at an thesis 6-7 mm. long; flowers, at least many or most of them, sessile,
the inflorescence generally dense and many-flowered S. lisianthoides.
Calyx 15-30 mm. long or larger, the sepals often narrowly winged dorsally.
Corolla 4-5 cm. long; calyx about 3 cm. long S. brachyptera.
Corolla 1.5-2 cm. long; calyx about 1.5 cm. long S. guianensis.
Schultesia brachyptera Cham. Linnaea 8: 8. 1833.
FIG. 90. Schultesia guianensis. A, habit of plant from subalpine region, X 3;
same plant, natural size; C, calyx, X 3; D, corolla dissected, X 3; E, pistil, X 3;
F, plant from the lowlands with 4 cm. of stem left out, natural size.
329
330 FIELDIANA: BOTANY, VOLUME 24
Savannas or open pine forest, usually in very wet soil, at or little
above sea level; British Honduras. Mexico; Honduras; Panama;
South America.
Plants slender, erect, 25-70 cm. high, simple or with a few erect branches
above; leaves sessile, linear or oblong-linear, obtuse to attenuate, 3-nerved below,
2.5-6 cm. long; flowers usually 1-5, solitary at the ends of the branches, each sub-
tended by 2 leaves, sessile or nearly so; calyx about 3 cm. long, lanceolate, attenu-
ate upward, conspicuously carinate and sometimes very narrowly winged along the
angles, the nerves between the angles none or obscure, the lobes linear-attenuate,
shorter than the tube, not appressed but slightly outcurved; corolla 4-5 cm. long,
rose-pink, the lobes rounded, inconspicuously apiculate.
This is an unusually beautiful plant when it occurs in abundance,
the flowers being large and of an exquisite shade of pink. It is typ-
ically a species of wet savannas, and often grows in shallow water.
The Central American plant has been called S. heterophylla Miq. in
recent publications. It is very close to that species, but is better
referred perhaps to S. brachyptera. It is questionable whether these
two species can be maintained, and the name used here has priority.
Schultesia guianensis (Aubl.) Malme, Arkiv. Bot. 3, no. 12:
9. 1904. Exacum guianense Aubl. PI. Guian. 68, t. 26, f. 1. 1775.
S. stenophylla Mart. Nov. Gen. & Sp. 2: 106, t. 182. 1826. S. mexi-
cana Wats. Proc. Am. Acad. 26: 144. 1891. S. chiapensis Brandeg.
Univ. Calif. Publ. Bot. 10: 413. 1924 (type from Chiapas).
Wet or moist meadows, sometimes in glades in oak forest, 1,100-
1,800 m.; Chiquimula; Jalapa; Santa Rosa; Huehuetenango. Central
and southern Mexico; British Honduras; El Salvador; Honduras;
Costa Rica; Panama; West Indies; South America.
A low annual, usually 30 cm. high or less, often only 2-3 cm. high, simple or
sparsely branched above, 1 -few-flowered; leaves sessile, linear to lanceolate or ob-
long, the lowest ones mostly shorter and broader than the upper, obtuse to attenu-
ate; flowers short-pedicellate, usually of a faded dirty pink to yellowish; calyx
about 15 mm. long, conspicuously winged on the angles, the wings green, the tube
conspicuously veined, the lobes subulate-attenuate, somewhat shorter than the
tube; corolla 1.5-2 cm. long; capsule oblong-ovoid, 12 mm. long.
Known in El Salvador as "hierba de la vida," "conchalagua" and
"sulfatillo." Although its flowers are rather large, this is not a pretty
or showy plant. A color form is f. lutescens Standl. & Steyerm. Field
Mus. Bot. 23: 77. 1944 (type from meadow in oak forest, near Jalapa,
Standley 76561], in which the corolla is pale buff or cream-colored.
It has been found also in Honduras (Siguatepeque) .
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 331
Schultesia lisianthoides (Griseb.) Benth. & Hook, ex Hemsl.
Biol. Cent. Am. Bot. 2: 348. 1882. Xestaea lisianthoides Griseb. Lin-
naea 22: 36. 1849.
Wet to dry thickets or open forest, often in stony fields, on open
banks, or in pine-oak forest, 1,000 m. or less; Alta Verapaz; Izabal;
Santa Rosa; Escuintla; Suchitepe'quez; Retalhuleu; San Marcos.
British Honduras; Southern Mexico to Panama; Colombia and
Venezuela.
Plants erect, 70 cm. high or less, usually with few or numerous suberect
branches; leaves thin, sessile, or the lowest contracted at the base into a short
petiole, the lower ones mostly obovate or obovate-oblong, the upper ones ovate-
oblong or ovate, obtuse to acuminate, the upper ones amplexicaul; flowers usually
very numerous, distributed almost throughout the length of the plant, in small open
cymes, sessile or on very short pedicels, the bracts lanceolate to subulate, small;
calyx green, at anthesis 5-7 mm. long, often longer in fruit and then distended by
the capsule, lobate almost to the base, the segments linear-lanceolate, carinate but
not winged, white-marginate, attenuate; corolla dirty pink, 10-14 mm. long; cap-
sule ellipsoid-oblong, about 8 mm. long.
Known in El Salvador by the names "sulfatillo" and "sulfato de
tierra." A rather weedy plant, with small and inconspicuous flowers.
Schultesia peckiana Robinson, Proc. Am. Acad. 45: 399. 1910.
Open places, at or little above sea level; endemic; British Hon-
duras, the type collected about plantations and in openings in forest
near Manatee Lagoon, M. E. Peck 318; collected also at All Pines,
and at Woods Bank, Sibun River.
Plants erect, similar in habit to S. lisianthoides, 60-75 cm. high, rather sparsely
and openly branched; leaves sessile, lance-ovate to elliptic, acute or the lower ones
obtuse, rounded at the base; inflorescence lax, repeatedly branched, the central
flowers of the cymes solitary, usually on long slender pedicels, these sometimes
3 cm. long but usually shorter, the pedicels mostly naked but sometimes bracteo-
late; calyx lobes narrowly lanceolate, scarious-marginate, carinate; corolla dirty
pink, the lobes ovate, acute, much shorter than the tube.
This species is evidently closely related to S. lisianthoides, perhaps
too closely so, but it appears to be distinct by the characters con-
trasted in the key.
VOYRIA Aublet
Small saprophytic plants without chlorophyll, the stems mostly simple and
1-flowered, stout; leaf scales much larger than in Leiphaimos, not appressed to the
stem but loose and somewhat spreading; flower subtended at the base by several
bractlets; calyx campanulate, 5-dentate, the teeth usually broad; corolla elongate-
cylindric, dilated at the middle, the limb 5-lobate, the lobes ovate to lanceolate,
332 FIELDIANA: BOTANY, VOLUME 24
spreading; stamens 5, inserted in the upper part of the corolla tube, the filaments
very short, the anthers elongate, obtuse; ovary sessile, with 2 parietal placentae;
style elongate and filiform ; capsule usually enclosed in the persistent corolla, elon-
gate, septicidal from base to apex; seeds globose or angulate, obscurely reticulate,
wingless; endosperm none.
Species about ten, in Central and South America. See Williams
in Fieldiana, Bot. 31: 411-415. 1968 for a discussion of this and allied
genus Leiphiamos.
Capsule truncate at the apex V. truncata.
Capsule not truncate at the apex.
Calyx 3-4 mm. long; corolla about 10 mm. long; stigma without lateral mem-
branaceous appendages V. alba.
Calyx 4-6 mm. long; corolla 10-15 mm. long; stigma with lateral membranaceous
appendages V. thalesioides.
Voyria alba (Standl.) L. Wms., Field Mus. Bot. 31: 411. 1968.
Leiphaimos alba Standl. Contr. U. S. Nat. Herb. 20: 198. 1919.
Saprophytic herbs in wet forests near sea level. British Hondu-
ras; Honduas; Nicaragua; Panama; Colombia.
Slender, leafless saprophytic herbs but with prominent cauline bracts, 6-16 cm.
tall. Inflorescence a simple to compound dichasium, 3-several usually sessile flow-
ers; flowers white, 9-12 mm. long; calyx 3-4 mm. long, cleft to the middle, the tube
campanulate, the lobes linear-lanceolate, acuminate, corolla about 10 mm. long
and 1 mm. in diameter, cylindric, lobes lanceolate, acute, ascending or spreading,
1.5-2.5 mm. long; capsule 6-8 mm. long, slender, dehiscing from base to apex;
seeds minute, trigonous to globose, not winged.
The smallest of the Voyrias of Central America and superficially
similar in aspect to Leiphaimos parasitica with which it has been
confused.
Voyria thalesioides (Standl.) L. Wms. Fieldiana, Bot. 31: 414.
1968. Leiphaimos thalesioides Standl. Contr. U. S. Nat. Herb. 20:
198. 1919.
Small saprophytic plants on the floor of wet lowland forests.
British Honduras (Gentle 5182); Panama.
Small yellow saprophytic herbs to 12 cm. tall. Leaves none, the stem with
several pairs of perfoliate bracts about 4 mm. long, the lobes triangular, interval
between nodes about 1 cm. or less; inflorescence a few-several-flowered corymb;
flowers yellow, 1-1.5 cm. long, pedicels 3-5 mm. long; calyx about 4-6 mm. long,
divided to about the middle, the tube broadly campanulate, the lobes becoming
linear or linear-lanceolate and alternated from a broad base; corolla 5-lobed, the
tube narrow, about 8-10 mm. long, persistent and becoming marcescent over the
swelling ovary, the lobes spreading, oblong-lanceolate, acute, 2-3 mm. long; sta-
mens inserted in the throat of the corolla, suborbicular, nearly sessile, about 0.75
mm. long; ovary subcylindric, style elongated and the capitate stigma reaches to
FIG. 91. Voyria thalesioides. A, habit of plant, natural size; B, corolla dis-
sected showing pistil and anthers, X 3; C, flower in natural position, X 3. Leiphai-
mos parasitica. D, habit of plant, natural size; E, corolla dissected to show pistil
and stamens, X 5; F, flower, X 5; G, anther, X 20.
333
334 FIELDIANA: BOTANY, VOLUME 24
the throat of the corolla, stigma with 5-6 small lateral membranaceous flaps; seeds
ovoid to reniform, wingless, about 0.2 mm. long.
Voyria truncata (Standl.) Standl. & Steyerm. Field Mus. Bot.
23: 78. 1944. Leiphaimos truncata Standl. Contr. U. S. Nat. Herb.
20: 196. 1919. V. allenii Steyermark, Ann. Mo. Bot. Gard. 28: 460.
1941.
Moist or wet soil among rotting leaves, 400 m. or less; Alta Vera-
paz (Cerro Chinaja) ; Izabal. Nicaragua; Panama.
Plants arising from elongate rootstocks, the stems erect, stout but somewhat
tortuous, 6-13 cm. long, simple or with 1 or 2 branches, dull red; leaf scales oppo-
site, dull red, numerous and close together, broadly ovate or rounded, rounded at
the apex, 4-5 mm. long, united by their bases; flowers 1-2, on stout pedicels 9 mm.
long or shorter, lilac or rose-lilac; calyx 7 mm. long, appressed to the base of the
corolla, the lobes rounded, minutely ciliate; corolla tube 3-4.5 cm. long, broadest
at the base around the ovary, narrowed upward, sometimes minutely pulverulent,
the lobes elliptic-ovate, obtuse, 15 mm. long or shorter; style slender, 27 mm. long,
the stigma sinuate-peltate; capsule as much as 15 mm. long and 5 mm. broad.
The species may be expected in other countries between Panama
and Guatemala.
APOCYNACEAE. Dogbane Family
References: Robert E. Woodson, Apocynaceae, N. Am. Flora 29:
103-192. 1938; The Apocynaceous Flora of the Yucatan Peninsula,
Carnegie Inst. Wash. Publ. 522: 61-102. 1940; Studies in the Apo-
cynaceae, II & IV, Ann. Mo. Bot. Gard. 15: 341-378. 1928; 20: 605-
790. 1933; 22: 153-306. 1935; 23: 169-438. 1936.
Trees, shrubs or herbs, often herbaceous or woody vines, usually with milky
latex. Leaves opposite or verticillate, rarely alternate, entire (or panduriform)
never serrate or dentate, estipulate or with stipules or stipular vestiges; flowers per-
fect, actinomorphic or slightly zygomorphic, normally pentamerous, inflorescence
of single flowers or usually racemose or cymose; calyx 5- (rarely 4-) lobed, often
with glandular appendages (squamellae) within, the lobes imbricate; corolla ga-
mopetalous, funnelform or salverform and often appendaged within, 5-lobed, the
lobes sinistrorsely or dextrorsely contorted in the bud; stamens 5 (rarely 4), epipe-
talous, alternate with corolla lobes, often connivent around the stigma, the anthers
bilocular, introrse, filaments free or sometimes united ; ovary superior, bicarpellate,
the carpels free or united, ovary 2-celled with axil placentae, or 1-celled with pari-
etal placentae, the style simple; the stigma large and various in form; ovules
1-many in each cell, anatropous or orthotropous; fruit of distinct or united carpels,
follicular, capsular, baccate or drupaceous; seeds usually with endosperm, naked,
comose at the apex, with a papery wing or sometimes arillate, embryo straight.
A large family of some 200 genera and 2,000-3,000 species with
the greater part of the species found in tropical regions of the world.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 235
The family is well represented in Guatemala. Few of the plants are
of economic importance; some are used as ornamentals; the latex of
several is known to be poisonous to man, and a few may be contact
poisons to persons with sensitive skin; cordage fibers or even textiles
may be produced from a few members of the family, especially from
the genus Apocynum.
Dr. Robert E. Woodson prepared monographs or revisionary
studies of most of the American genera of this family and has brought
them into most satisfactory order, the more convenient because of
the chaos formerly prevalent in some of the groups. Our account of
this family in Guatemala is based largely on Dr. Woodson's studies.
Members of this family usually are said to be without stipules,
however stipules or stipular vestiges are often present. Interpetiolar
stipules much like those in some Rubiaceae are to be found in some
Guatemalan species.
Plants scandent.
Leaves verticillate; fruit covered with long spines; flowers large, bright yellow.
Allamanda.
Leaves opposite.
Corolla 12-20 cm. long; cultivated plants Beaumontia.
Corolla much smaller, often very small; native plants.
Connectives of the anthers with thick obtuse basal lobes; stigma pentagonal-
umbraculiform; upper surface of the leaves glandular on the costa, at
least at its base.
Inflorescence simple, not branched Mandevilla.
Inflorescence branched, or at least obscurely dichotomous.
Corolla funnelform, dull red outside; inflorescence repeatedly branched.
Tintinnabularia .
Corolla salverform, greenish white; inflorescence obscurely dichoto-
mous, branched once or twice Mesechites.
Connectives of the anthers with slender or attenuate basal lobes or, if with
obtuse lobes (Fernaldia) the sporangia with conspicuous sterile pro-
tuberant bases; stigma fusiform or subcapitate; leaves not glandular.
Calyx without squamellae inside the lobes.
Corolla salverform, the tube usually spirally twisted, the orifice with a
callous annulus Laubertia.
Corolla funnelform, the tube not twisted, the orifice without a callous
annulus Rhabdadenia.
Calyx bearing squamellae inside the lobes.
Squamellae as many as the calyx lobes and opposite them, sometimes
deeply lacerate.
Corolla with ligular appendages within behind the stamens, the ori-
fice callous and annular; corolla funnelform or salverform.
Prestonia.
Corolla not appendaged within, without a callous annular orifice.
Corolla funnelform, the lobes arachnoid- villous within. .Fernaldia.
Corolla salverform, not arachnoid- villous within Echites.
Squamellae alternate with the calyx lobes or indefinitely distributed.
336 FIELDIANA: BOTANY, VOLUME 24
Corolla salverform, small, white Forsteronia.
Corolla funnelform, large, yellow.
Anthers with linear apical appendages; placentae of the ovary
becoming chaffy in fruit Urechites.
Anthers without linear apical appendaged; placentae not chaffy in
fruit Odontadenia.
Plants not scandent.
Leaves alternate, or a few of the lower ones sometimes opposite.
Plants low, mostly 50 cm. high or less, herbaceous throughout or merely suf-
frutescent below; flowers yellow Haplophyton.
Plants large shrubs or trees, with woody branches.
Carpels of the ovary many-ovulate.
Calyx lobes 4, the 2 outer ones connate and completely enclosing the
smaller ones Aspidosperma.
Calyx lobes 5, equal or nearly so.
Follicles of the fruit strongly compressed, ligneous; seeds surrounded
by a very broad, papery wing; flowers small Aspidosperma.
Follicles not compressed, not ligneous; seeds with a small excentric
basal wing; flowers large and showy Plumeria.
Carpels of the ovary containing only 1-6 ovules.
Calyx without squamellae; flowers small, white, the tube 7-8 mm. long;
fruit small, white, juicy Vallesia.
Calyx with squamellae within; flowers large and showy, usually yellow,
fruit large, not white Thevetia.
Leaves opposite or verticillate.
Leaves all or chiefly verticillate.
Cultivated plants, the flowers large and showy Nerium.
Native plants, with small or large flowers.
Fruit of dry follicles; leaves mostly linear-oblanceolate; flowers small,
white Tonduzia.
Fruit fleshy; leaves mostly elliptic or oval.
Ovary 1-celled; tall trees with rose-purple flowers Couma.
Ovary 2-celled; shrubs with white flowers Rauvolfia.
Leaves opposite.
Plants herbaceous.
Corolla blue or blue-purple; plants with elongate and often rooting basal
shoots Vinca.
Corolla white or pink, never bluish; plants annual, without sterile basal
shoots Lochnera.
Plants shrubs or trees.
Fruit samaroid; leaves very obtuse or rounded at the apex; flowers small,
white Cameraria.
Fruit fleshy or of dry follicles, never samaroid; leaves mostly acute or
acuminate.
Anthers connivent and agglutinated to the stigma; corolla lobes dex-
trosely convolute in bud Malouetia.
Anthers neither connivent nor agglutinated to the stigma; corolla lobes
sinistrorsely convolute in bud.
Carpels of the ovary united throughout; fruit fleshy and juicy.
Lacmellea.
Carpels of the ovary distinct; fruit dry or nearly so.
Calyx lobes large, foliaceous or petaloid; corolla funnelform or sal-
verform.. . .Stemmadenia.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 337
Calyx lobes small, corolla salverform Tabernaemonlana.
Calyx lobes linear, pubescent Haplophylon.
Calyx lobes broader, scarious, glabrous Tabernaemontana.
ALLAMANDA L.
Woody vines with copious milky sap; leaves mostly quaternate, not glandular;
inflorescence cymose, few-flowered, lateral or pseudoterminal; calyx 5-parted, the
lobes equal or nearly so, foliaceous, without squamellae; corolla large and showy,
funnelform, the limb regularly 5-lobate, the lobes sinistrorsely contorted; anthers
not connivent, wholly included, the connective not enlarged; ovary 1-celled, the
numerous ovules borne upon two linear placentae, surrounded by a low annular
nectary; fruit capsular, globose or subglobose, densely echinate; seeds numerous,
compressed, winged.
About ten species, in tropical America. Only the following is
known from North America.
Allamanda cathartica L. Mant. PI. 214. 1771. Amanda; cam-
pana.
Wooded swamps or wet forest, at or little above sea level; Izabal;
much planted for ornament in other parts of Guatemala. British
Honduras to Panama, along the Atlantic coast; northeastern South
America.
A large woody shrub or sometimes an arching vine, glabrous to villous or hir-
sute, especially on the stems and lower leaf surfaces; leaves subcoriaceous to mem-
branaceous, short-petiolate, obovate to oblong-lanceolate, 6-15 cm. long, 2.5-6 cm.
broad, acuminate, often caudate-acuminate, attenuate to the base; inflorescences
few-flowered; flowers sweet-scented, golden yellow; calyx lobes ovate to lanceolate,
acute, 5-12 mm. long, spreading; corolla 7-10 cm. long, the tube 2-3.5 cm. long,
slender, the throat 3-4 cm. long, much ampliate, the lobes obliquely obovate,
spreading, broadly rounded at the apex; capsule 4-6 cm. broad, somewhat com-
pressed, covered with numerous green spines 1 cm. long.
Called "San Jose"" in El Salvador. This handsome plant is grown
in many tropical and subtropical regions remote from its native habi-
tat, as in Florida. It is very showy when in blossom, in Guatemala
most profusely perhaps in January and February. It has every ap-
pearance of being native in wet forests along the Atlantic coast of
Central America, in Guatemala only along the coast of Izabal. In
this country it is planted commonly from the central highlands, at
about 1,500 m., down to the Pacific and Atlantic coasts. In the
Canal Zone the English speaking people give the flowers the name of
"buttercups."
338 FIELDIANA: BOTANY, VOLUME 24
ASPIDOSPERMA Martius & Zuccarini
Reference: Robert E. Woodson, Studies in the Apocynaceae. VIII.
Ann. Mo. Bot. Gard. 38: 119-204. 1951.
Large trees of the forest or rarely shrubs, with whitish or reddish latex. Leaves
alternate or approximate, rarely decussate or ternate (not ours) ; inflorescence ter-
minal or axillary, determinate, dichasially cymose or dichasial and thyrsiform;
calyx with usually 5 equal or unequal lobes, or sometimes reduced to 4, eglandular
within; corolla salverform to tubular-salverform or tubular, constricted or not at
the throat, the lobes sinistrorsely contorted in the bud; anther inserted at the
middle or higher in the tube, mucronulate; ovary superior, bicarpellate, the carpels
essentially free, fruit follicular, usually strongly compressed and more or less
woody, usually asymmetrical, from nearly circular to falciform or dolabriform;
seeds peltate, greatly compressed and with a flat, papery wing.
This genus supplies important timber trees in Guatemala, and
more especially in South America. There are about 60 species,—
only two are known in Central America.
Calyx densely pubescent; corolla lobes caudate-acuminate, the tube callose-
angulate A. megalocarpon.
Calyx nearly glabrous; corolla lobes not abruptly caudate-acuminate, the tube
essentially terete A. stegomeris.
Aspidosperma megalocarpon Muell.-Arg. Linnaea 30: 40. 1860;
Woodson, Ann. Mo. Bot. Gard. 38: 192. 1951. Macaglia megalocarpa
0. Kuntze, Rev. Gen. PI. 2: 416. 1891. Aspidosperma cruentum
Woodson, Am. Jour. Bot. 22: 634. 1935. A. matudae Lundell, Phyto-
logia 1: 339. 1939 (as Matudai). A. chiapense Matuda, Madrono 10:
172. 1950. A. chiapense forma tenax Matuda, I.e. 174.
In dense, dry forests or in river bottom forests as well; Pete"n; Alta
Verapaz; Suchitepe"quez ; Escuintla. Southeastern Mexico; British
Honduras; Honduras. South to Colombia and British Guiana.
Trees 7-30 m. tall, the trunk 2-8 dm. in diameter and with whitish, rough bark;
branches minutely gray-pilosulose when young, soon glabrate and developing a
blackish bark without apparent lenticels; leaves alternate, elliptic-obovate to nar-
rowly oblong, apex acute to broadly obtuse, base acutely or obtusely cuneate,
5-25 cm. long, 2-9 cm. broad, firmly membranaceous or subcoriaceous, wholly
glabrous, highly lustrous above, somewhat paler beneath, the secondary veins
broadly ascending to subhorizontal, very numerous and crowded; petioles 1-3 cm.
long; inflorescences terminal and axillary at the uppermost nodes, broadly corym-
bose-thyrsiform, shortly pedunculate, many-flowered, sordid brown-tomentulous
without; corolla yellowish white, glabrous without, the tube callose-angulate,
3-4 mm. long, the lobes linear, caudate-acuminate, strongly spiraled in the
bud, about 1.5-2.0 mm. long; anthers inserted about midway within the corolla
tube, about 1 mm. long; ovary globose, glabrous, about 0.7 mm. long; follicles
broadly oval to nearly circular, 8-15 cm. long and 6-11 cm. broad, woody, brown-
tomentulous, with a rather broad stipe 1-3 cm. long.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 339
FIG. 92. Aspidosperma megalocarpon, A, habit, X H; B, flower, X 5; C, co-
rolla dissected to show stamens, X 5; D, pistil, X 10; E, winged seed, X ]^.
Aspidosperma stegomeris Woodson, Ann. Mo. Bot. Card. 38:
178. 1951. Cufodontia stegomeris Woodson, Archive Bot. 10: 39.
1934. C. lundelliana Woodson, I.e. 40 (type from Pete"n, Lundell
34.08). Aspidosperma lundellianum Woodson, Am. Jour. Bot. 22:
684. 1935. Chichica; sacuallon; bayo bianco; malerio bianco.
Dry forests at low elevations, Peten ; Retalhuleu. Mexico ; British
Honduras; Costa Rica.
340 FIELDIANA: BOTANY, VOLUME 24
Trees as much as 33 m. in height, the branches rather slender, indefinitely
puberulent-papillate when very young, soon becoming glabrate and with a thin,
striate, conspicuously lenticellate, yellowish-gray bark. Leaves alternate, rather
narrowly elliptic-oblong to broadly oval, apex shortly acuminate to broadly rounded,
base broadly obtuse to rounded, 6-16 cm. long and 2.5-7 cm. broad, firmly mem-
branaceous to subcoriaceous, above dark green, glabrous, and somewhat lustrous,
beneath paler and indefinitely papillate to essentially glabrous; petioles 0.7-1.5 cm.
long; inflorescences lateral and extra-axillary near the tips of the leafy branches,
cymose, several-flowered, more or less densely yellow-papillate, the peduncles once-
to-thrice dichotomous and about 1-2 cm. long; bracts extremely inconspicuous or
absent; pedicels 1-5 mm. long; outer calyx lobes 3-6 mm. long, more or less densely
yellow-papillate; corolla yellowish-white, the tube 4-6 mm. long, about 1.5 mm.
broad, densely and appressed velutinous-papillate without, the lobes oblong-ellip-
tic, 5-6 mm. long, essentially glabrous without; stamens inserted about midway
within the corolla tube, the anthers about 1.2 mm. long; ovary ovoid, glabrous,
about 1 mm. long; follicles broadly subreniform to nearly circular, 7-11 cm. long
and 6.5-8 cm. broad, sessile, indefinitely papillate to essentially glabrous, covered
with small lenticels.
This tree was formerly not uncommon near Retalhuleu where it
was an important lumber tree.
Called "white malady" or "cojoton" in British Honduras.
BEAUMONTIA Wallich
Large woody vines; leaves opposite, membranaceous, penninerved, the axils
often glanduliferous; flowers very large, white, fragrant, in terminal cymes; bracts
mostly foliaceous and caducous; calyx 5-parted, eglandular within at the base or
bearing numerous small glands, the segments large, foliaceous; corolla funnelform,
the tube short, the throat ampliate, without squamellae, the 5 lobes dextrorsely
contorted; stamens inserted at the apex of the tube, included in the throat, the
filaments thickened at the apex; anthers sagittate, short-acuminate, connivent
about the stigma and adherent to it, the anther cells produced at the base into
acuminate appendages; disk 5-lobate; ovary 2-celled, the style filiform, the stigma
oblong-fusiform, exannulate; ovules numerous in each cell; fruit elongate, thick,
subligneous, finally separating into two follicles; seeds compressed, ovate or oblong,
attenuate to the apex and bearing a tuft of hairs; radicle short, superior.
About four species, native in the East Indies and Malaysia.
Beaumontia grandiflora Wall. Tent. Fl. Nep. 15, t. 7. 1824.
Campanula; campana blanca.
Native of eastern India, now grown in many tropical regions of
the earth as an ornamental plant; rarely in Guatemala, but cultivated
also in other parts of Central America.
A large woody vine, the stout young branches ferruginous-tomentose; leaves
short-petiolate, oblong-obovate, 10-17 cm. long or larger, rounded or obtuse at the
apex and shortly cuspidate-acuminate, acute or obtuse at the base, when young
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 341
somewhat ferruginous-tomentose beneath but in age almost glabrous; cymes ter-
minal, few-flowered, the flowers pedicellate; calyx 3-4 cm. long, ferruginous-tomen-
tose, green, the broad segments cuspidate; corolla white, 12-20 cm. long, somewhat
pubescent outside, the lobes short and broad, acutish; follicles horizontally diver-
gent, linear, with incurved obtuse tips, 15-25 cm. long, slender.
Known in El Salvador as "pomoncia," "bomoncia" and "azucena
japonesa." The vine is highly recommended for cultivation in trop-
ical and subtropical regions. It often attains a great size, and in its
native country is said to cover tall trees. The flowers are almost pure
white, exceedingly large, and in appearance almost exactly like those
of Datura arborea. In Guatemala the plant is somewhat of a rarity,
and in the smaller towns usually is closely guarded by its owners.
The flowers are sometimes employed in making funeral wreaths.
CAMERARIA L.
Glabrous shrubs or small trees; leaves opposite, eglandular; inflorescence ter-
minal, cymose, few-flowered; calyx without squamellae, 5-cleft almost to the recep-
tacle, the lobes subequal; corolla salverform, white, small, not appendaged within;
anthers not connivent, the connective not enlarged; ovary of 2 distinct carpels,
without a nectary, 1-4-ovulate; follicles 2, indehiscent, samaroid, with a broad
inequilateral wing, usually containing a single naked seed.
Two species, the other in Haiti.
Cameraria latifolia L. Sp. PI. 210. 1753. C. retusa Griseb. Fl.
Brit. W. Ind. 410. 1861. C. belizensis Standl. Trop. Woods 7: 8. 1926
(type from British Honduras, S. J. Record}. Chechem de caballo;
savanna white poisonwood; white poisonwood in British Honduras;
iquiche, chechem (Pete"n).
In low mixed forests, little above sea-level, Peten. British Hon-
duras. West Indies.
A shrub or small tree with ashy gray bark; leaves firm-membranaceous to sub-
coriaceous, on petioles 2-6 cm. long, mostly oblong-ovate or oblong-oval, 1.5-4 cm.
long, obtuse or rounded at the apex and emarginate (in British Honduras material),
obtuse or rounded at the base, very lustrous above; inflorescence terminal, usually
2-4-flowered, about equalling the subtending leaves, the pedicels 5-8 mm. long;
calyx lobes ovate, acute or obtuse, 1-1.5 mm. long, imbricate; corolla white, the
tube 5-8 mm. long, somewhat dilated above, the lobes broadly obovate, 6-15 mm.
long; stamens included, sessile, anthers short, obtuse, apex of connective produced
into a long, free filament; fruit 4-4.5 cm. long, 1.5-2 cm. broad.
The plant is highly poisonous if the latex falls upon the skin, pro-
ducing serious swelling and inflammation similar to that produced by
poison ivy (Rhus radicans). The wood is pale olive, hard, heavy,
fine- textured, finishes very smoothly, is not durable; no use is made
342
FIELDIANA: BOTANY, VOLUME 24
FIG. 93. Cameraria latifolia. A, habit, natural size; B, calyx and pistil, X 4;
C, portion of the corolla tube to show stamens in natural position, X 5.
of it. Woodson recognizes only two species of Cameraria, but other
authors commonly have recognized a larger number. The British
Honduras plant is like C. retusa Griseb., which perhaps will prove to
be a perfectly distinct species when more ample material of the genus
has been assembled for study.
COUMA Aublet
Reference: Joseph Monachino, A Revision of Couma and Para-
hancornia (Apocynaceae), Lloydia 6: 229-247. 1943.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 343
Usually large trees; leaves verticillate or sometimes opposite, eglandular; in-
florescence cymose, lateral, many-flowered, the flowers small; calyx 5-parted, the
lobes equal, imbricate, without squamellae; corolla salverform, the limb 5-parted,
the lobes equal, sinistrorsely contorted; anthers not connivent, the connective not
enlarged; ovary syncarpous, 1-celled, the ovules numerous, borne upon 2 linear
parietal placentae; fruit a large several-seeded berry.
About five or six species, in tropical America, all except the follow-
ing confined to South America.
Couma macrocarpa Barb.-Rodr. Vellosia ed. 2, 1: 32, t. l,fig. b.
1891. C. guatemalensis Standl. Trop. Woods 7: 8. 1926 (type col-
lected near Entre Rios, Izabal, S. J. Record 4-2). Palo de vaca; polo
de leche.
Low wet mixed forest, or often seen in cleared pastures, at or little
above sea level; Izabal. British Honduras; possibly Panama; Co-
lombia; Peru; and Venezuela to Amazonian Brazil.
A large or medium-sized tree with thick, dark-colored bark, the young branch-
lets thick, hirtellous with short, slender, stiff, brownish hairs; leaves ternate, on
stout petioles 8-17 mm. long, elliptic, 6.5-27 cm. long and 3.5-18 cm. broad, usually
abruptly short-acuminate, obtuse and often short-decurrent at the base, charta-
ceous at maturity, green above and glabrous or nearly so, densely and minutely
puberulent beneath; inflorescence about equalling the leaves, the flowers very
numerous, rose-purple; calyx lobes oblong-lanceolate, obtuse, 2-2.5 mm. long, mi-
nutely puberulent; corolla puberulent, the tube 7-8 mm. long, the lobes oblong,
obtuse, 4-6 mm. long; fruit subglobose, very fleshy, about 3 cm. in diameter, yellow
at maturity.
Called "barca" in British Honduras. One of the most interesting
of Central American trees, and one that has received much attention
in periodical literature of the United States, especially in newspapers.
The first known North American material was obtained by the senior
author in June, 1922, in a swamp at Puerto Barrios, but it was not
determined, because of its imperfect condition, until Professor Re-
cord obtained flowering specimens at Entre Rios in March, 1926.
When the bark of the cow tree is cut or broken, there issues from it
a rich creamy latex that is sweet and palatable. It is not very sticky
and may be drunk like cow's milk. The tree is none too abundant
in Guatemala and British Honduras, but is plentiful in some local-
ities. There are numerous trees close about Puerto Barrios, in pas-
tures or other clearings. They are 12-18 m. high, with an almost
smooth, tall trunk, and a dense rounded crown. The flowers are
borne in great abundance when the trees are leafless or nearly so, and
they make the trees conspicuous and easily spotted from considerable
distance. In British Honduras the cow tree has been collected along
FIG. 94. Couma macrocarpa. A, habit, X 1A', B, flower, X 2; C, calyx par-
tially dissected to show pistil, X 5; D, portion of corolla tube to show anthers in
position and patches of pubescence, X 4.
344
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 345
the Temash River. The best known trees of Guatemala are those
at Entre Rios not far inland from Puerto Barrios. In years past
many tourists have been taken to these by the United Fruit Com-
pany to see one of the "natural wonders" of Guatemala. The white
latex is sometimes used in British Honduras as a chicle substitute.
Little or no use is made locally of it as a beverage, although its prop-
erties are rather well known, perhaps as the result of long advertis-
ing. It is reported from British Honduras that woodsmen sometimes
use an infusion of the leaves as a beverage. The wood is dull brown,
moderately hard, of medium texture, fairly straight-grained, not
difficult to work. In Venezuela this tree is reported to attain a
height of 40 m., with a trunk as much as a meter in diameter. In
that country it is stated that the latex boiled in water gives a product
similar to guttapercha that is used for caulking canoes.
The species is to be expected in other Central American countries
in the Atlantic forests.
ECHITES P. Browne
Reference: Robert E. Woodson, Studies in Apocynaceae IV. Ann.
Mo. Bot. Gard. 23: 217-252. 1936.
Slender, woody or suffrutescent vines; leaves opposite, eglandular; inflores-
cence a more or less modified dichasium, alternate-axillary, rarely terminal or sub-
terminal, the flowers several, rarely solitary; calyx 5-parted almost to the receptacle,
the lobes subequal, somewhat imbricate, bearing within at the base a solitary, often
deeply dissected squamella; corolla salverform, the tube not appendaged or annu-
late within, the limb regularly 5-parted, dextrorsely contorted; anthers connivent
and agglutinated to the stigma, the connective enlarged, narrowly or rarely ob-
tusely bilobate; ovary of 2 distinct carpels, many-ovulate, surrounded at the base
by 5 distinct or somewhat concrescent nectaries; stigma fusiform-subcapitate; fruit
of 2 distinct follicles; seeds numerous, rostrate, comose at the apex.
Seven species are known, in tropical America. One other from
Central America has been described from Costa Rica.
Corolla about 2 cm. long E. tuxtlensis.
Corolla 3-8 cm. long.
Tube of the corolla conspicuously twisted E. umbellata.
Tube of the corolla not twisted.
Inflorescence dichasial to simple helicoid, not subumbellate; calyx lobes 3-7
mm. long; leaves rather thin, usually abruptly short-acuminate, not
lobate E. turrigera.
Inflorescence umbelliform; calyx lobes 2-3 mm. long; leaves thick, long-
acuminate, often panduriform E. yucatanensis.
Echites turrigera Woodson, Ann. Mo. Bot. Gard. 19: 381. 1932.
346 FIELDIANA: BOTANY, VOLUME 24
In thickets, 600-1,200 m.; Zacapa (type from Gualan, Deam
6376); Jutiapa (Lago de Giiija). Honduras; Nicaragua.
A somewhat woody vine; leaves long-petiolate, membranaceous, broadly ob-
long-elliptic to ovate-elliptic, 5-15 cm. long, 3-10 cm. broad, rather abruptly
short-acuminate or acute, obtuse to rounded or rarely subcordate at the base,
glabrous, green above, paler beneath, reticulate-veined on both surfaces; inflores-
cence lateral or subterminal, dichasial to simple helicoid, equalling or longer than
the leaves, 6-20-flowered, the peduncles minutely pilosulous or glabrate, the pedi-
cels 10-12 mm. long; bracts oblong-lanceolate, acuminate, somewhat foliaceous,
1-3 mm. long; calyx lobes lanceolate, acuminate, 3-7 mm. long, sparsely pilosu-
lous; corolla white or cream-colored, salverform, the tube 28-38 mm. long, 1.5-
2 mm. thick at the base, dilated below the middle, constricted toward the orifice,
the lobes obliquely obovate, 1.5-2 cm. long, spreading; young follicles slender,
pilosulous, 7 cm. long, 4 mm. thick.
Echites tuxtlensis Standl. Contr. U. S. Nat. Herb. 23: 1164.
1924.
In forest or thickets, 800-1,000 m.; Pete"n; Huehuetenango. Brit-
ish Honduras, in high ridge; Mexico (Chiapas; Yucatan); Honduras;
Costa Rica.
A small slender vine, glabrous throughout, essentially herbaceous, arising from
a small tuberous root; leaves on petioles 3-8 mm. long, thick-membranaceous, nar-
rowly lanceolate to obovate-elliptic, 6-10 cm. long, 1.5-3.5 cm. broad, acuminate,
cuneate to rounded at the base, the nerves and veins obscure; inflorescence a lax,
repeatedly compound dichasium bearing several to many, small, yellowish green
flowers, much longer than the leaves; pedicels 7-10 mm. long; calyx lobes ovate-
triangular, acute or acuminate, 1-1.5 mm. long; corolla tube 8-10 mm. long, the
lobes oblong-lanceolate to elliptic-lanceolate, acuminate, somewhat shorter than
the tube, reflexed in anthesis; nectaries half as long as the ovary.
The Maya names of Yucatan are reported as "cahuale chac can-
cel" and "ibincan."
Echites umbellata Jacq. Enum. PL Carib. 13. 1760.
British Honduras, at or little above sea level. Southern Florida;
Mexico (Yucatan); Honduras; West Indies; coast of Colombia.
A suffrutescent vine, glabrous throughout; leaves chartaceous, on petioles 3-
15 mm. long, narrowly oblong-elliptic to suborbicular, 4-12 cm. long, 2-7.5 cm.
broad, acute to usually rounded or retuse at the apex, cuneate to rounded at the
base; inflorescences lateral or subterminal, somewhat shorter than the leaves, 2-7-
flowered, pedunculate; bracts ovate or ovate-lanceolate, 1.5-3 mm. long; calyx
lobes ovate to narrowly oblong-triangular, acute or acuminate, 1.5-5 mm. long,
scarious or only slightly foliaceous; corolla cream-colored, the tube 2-5.5 cm. long,
somewhat dilated below the middle, above spirally contorted and gradually con-
stricted toward the orifice, the lobes obliquely obovate, 1-3 cm. long, spreading;
nectaries half as long as the ovary or equalling it; follicles rather stout, rigidly
divaricate, 10-25 cm. long, glabrous.
FIG. 95. Echites tuxtlensis. A, habit showing tuber, X 1A\ B, flower just before
anthesis, X 2; C, calyx and style, X 5; D, corolla tube dissected to show stamens
in natural position, X 2}/£; E, a single stamen, X 5.
347
348 FIELDIANA: BOTANY, VOLUME 24
Echites yucatanensis Millsp. ex Standl. Field Mus. Bot. 8: 35.
1930.
British Honduras (Tower Hill, Karling 28); Mexico (Yucatan
and Campeche).
A suffrutescent vine, glabrous throughout; leaves coriaceous or subcoriaceous,
on petioles 1-2 cm. long, ovate to oblong, often irregularly pandurate, 7-12 cm.
long, 2.5-7 cm. broad, acuminate, obtuse or rounded at the base, deep green and
lustrous above, the veins reticulate and very conspicuous on both surfaces; inflores-
cences lateral, subumbellate, 3-9-flowered, the pedicels 10-13 mm. long, the bracts
minute, scarious, ovate-lanceolate; calyx lobes ovate-lanceolate, 2-3 mm. long,
acuminate; corolla pale greenish yellow, the tube 4-4.5 cm. long, abruptly dilated
below the middle, gradually constricted above toward the orifice, the lobes obliquely
obovate, 2.5-3 cm. long; nectaries less than half as long as the ovary; follicles slen-
der, divaricate, 15-25 cm. long, glabrous.
FERNALDIA Woodson
Slender vines, herbaceous or suffrutescent; leaves opposite, eglandular; inflores-
cences alternate-axillary, simply scorpioid, with several large showy flowers; calyx
5-parted almost to the receptacle, the lobes equal or subequal, scarious or subfolia-
ceous, scarcely imbricate, each with a solitary squamella within; corolla funnelform,
the tube not appendaged within, the throat usually more or less closed by a dense
villous indument, the limb regularly 5-parted, dextrorsely contorted; anthers con-
nivent and agglutinated to the stigma, the connective enlarged, obtusely bilobate;
ovary of 2 distinct carpels, many-ovulate, surrounded at the base by 4 rather un-
equal, more or less concrescent nectaries; stigma fusiform-capitate; fruit of 2 dis-
tinct follicles; seeds numerous, truncate and comose at the apex.
This genus was named by Dr. Woodson for his friend and some-
times mentor, Prof. Merritt L. Fernald, who spent a lifetime studying
the flora of the northeastern United States and adjacent Canada.
While Prof. Fernald was pleased that this genus had been named for
him, the junior author once heard Prof. Fernald speculate that Wood-
son may have been "pulling his leg" in using the first of the specific
names below, for it described Prof. Fernald quite well.
Two other species are known, one in western Mexico, the other
in Panama.
Corolla short-pilose outside, the throat narrowly conic, 16-18 mm. long.
F. brachypharynx.
Corolla glabrous outside, the throat broadly campanulate-conic, 9-12 mm. long.
F. pandurata.
Fernaldia brachypharynx Woodson, Ann. Mo. Bot. Gard. 19:
380. 1932. Loroco.
In thickets, endemic; Escuintla (type collected along the road
between Escuintla and San Jose", Sutton Hayes); Guatemala; Sacate-
pe"quez (cultivated).
FIG. 96. Fernaldia pandurata. A, habit, X Yi; B, stamen, X 5; C, calyx dis-
sected to show corona, pistil, X 5.
349
350 FIELDIANA: BOTANY, VOLUME 24
A small or large, herbaceous vine, velutinous-puberulent throughout; leaves on
petioles 1-2 cm. long, membranaceous, broadly ovate or elliptic, 7-10 cm. long,
5-7 cm. broad, abruptly short-acuminate, broadly rounded at the base; inflores-
cences slightly shorter than the leaves, few-flowered, the pedicels 4-5 mm. long,
the bracts ovate, 1-2 mm. long; calyx lobes ovate, acuminate, 2-3 mm. long, mi-
utely pilosulous; corolla greenish white, pilosulous outside, the tube 18-20 mm.
long, the throat narrowly conic, 16-18 mm. long, 7 mm. broad, the lobes 12-14 mm.
long, arachnoid-villous within at the base.
This and the following species are well known plants of northern
Central America, the flowers and flower buds being cooked and eaten
commonly, in small meat pies or with rice or other substances. The
roots are said to be very poisonous, and in Chiquimula are employed
for poisoning noxious animals.
Fernaldia pandurata (A. DC.) Woodson, Ann. Mo. Bot. Gard.
19: 48. 1932. Echites pandurata (A. DC.) in DC. Prodr. 8: 458. 1844.
Urechites karwinskii Muell.-Arg. Linnaea 30: 440. 1860. E. pingui-
folia Standl. Field Mus. Bot. 8: 35. 1930 (type from Yucatan). Loroco.
At 900 m. or lower; Izabal; Zacapa; Chiquimula; Jutiapa. Mex-
ico; El Salvador; Honduras.
A small or large herbaceous vine, densely puberulent or shortly velutinous-
pilosulous throughout; leaves membranaceous, on petioles 1-2 cm. long, oblong-
elliptic to broadly ovate, 4-13 cm. long, 1.5-8 cm. broad, short-acuminate, the
lower ones cordate at the base, the upper obtuse to truncate, usually very densely
and softly pilose beneath; inflorescences generally somewhat shorter than the
leaves, with 8-18 flowers, the pedicels 4-6 mm. long; bracts ovate, 1-2 mm. long;
calyx lobes ovate, acute or obtuse, 2-3 mm. long; corolla white within, greenish
outside and glabrous, the tube 20-22 mm. long, the throat broadly campanulate-
conic, 9-12 mm. long, 7-9 mm. broad, the lobes ciliate, 10-13 mm. long, densely
villous-arachnoid within at the base.
A glabrous variety is found in Honduras, var. glabra Molina,
where this species is a common one.
FORSTERONIA G. Meyer
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 22: 153-224.
1935.
Woody vines; leaves normally opposite, glandular above at the base of the
costa or rarely eglandular, usually pitted beneath in the axils of the nerves; inflores-
cence terminal or terminal and lateral, aggregate-dichasial or thyrsiform, the flowers
small, very numerous; calyx 5-parted almost to the receptacle, the lobes subequal,
more or less imbricate, scarious, usually with several or numerous squamellae with-
in; corolla rotate or nearly so, the tube short, the orifice not annulate, the limb
equally 5-parted, dextrorsely or rarely sinistrorsely contorted; anthers connivent
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 351
FIG. 97. Forsteronia myriantha. A, habit, X Y^', B, flower and buds, X 5;
C, corolla dissected to show anthers in natural position, X 5; D, pistil (note alate
stigma).
and agglutinated with the stigma, the connective enlarged, rather narrowly bilo-
bate; ovary of 2 distinct carpels or very rarely syncarpous, many-ovulate, sur-
rounded by 5 distinct or more or less concrescent nectaries; stigma fusiform or
subcapitate; follicles distinct or very rarely more or less agglutinated; seeds numer-
ous, truncate and comose at the apex.
About 48 species, in tropical America. No others are known from
Central America; one is known from Panama.
Anther tips barely exserted from the corolla or wholly included; filaments free from
the style F. viridescens.
Anthers wholly exserted; filaments agglutinated to the style, at least above.
Leaves not glandular on the upper surface F. peninsularis.
Leaves glandular above at the base of the costa.
Squamellae of the calyx numerous, indefinitely distributed F. myriantha.
Squamellae alternate with the calyx lobes, solitary or infrequently in groups
of 2-3 F. spicata.
352 FIELDIANA: BOTANY, VOLUME 24
Forsteronia myriantha Donn.-Sm. Bot. Gaz. 27: 435. 1899.
Moist or wet thickets or forest, 1,800 m. or less; Pete"n; Santa
Rosa; Sacatepe"quez (type from Embaulada, Heyde & Lux 4534);
Suchitepe"quez; Quezaltenango. British Honduras; Honduras; Costa
Rica; Panama.
A small or large vine, sometimes 12 m. long and climbing over trees; leaves
thick-membranaceous, on petioles 2-4.5 mm. long, elliptic to oval or elliptic-
oblong, 4.5-10 cm. long, 2-4.5 cm. broad, acute or acuminate, obtuse to broadly
acute at the base, glabrous, or sometimes sparsely pilose beneath, inconspicuously
glandular above at the base of the costa; inflorescence terminal, thyrsiform, shorter
than the leaves, the flowers white or greenish yellow, the pedicels 1-2 mm. long;
calyx lobes ovate, subacute or obtuse, 1 mm. long, puberulent or rarely tomentu-
lose, the squamellae numerous, indefinitely distributed; corolla glabrous or mi-
nutely papillate outside, the tube 1 mm. long or less, the lobes oblong-obovate,
2.5-3 mm. long; anthers glabrous, widely exserted; immature follicles 10 cm. long
or more, very slender, glabrous.
Forsteronia peninsularis Woodson, Ann. Mo. Bot. Gard. 22:
215. 1935.
Moist or wet forest, 1,300-1,500 m.; Quezaltenango (?). British
Honduras (type from Maskall, Northern River, Gentle 1281).
A large woody vine; leaves firm-membranaceous or subcoriaceous, on petioles
5-8 mm. long, oblong-elliptic to lance-oblong, 4.5-5.5 cm. long, 1.5-2 cm. broad,
subacute to acuminate, obtuse at the base, glabrous, eglandular; inflorescence sub-
thyrsiform, broadly pyramidal, somewhat shorter than the leaves, the flowers
greenish white, the pedicels 2-2.5 mm. long; calyx lobes ovate, obtuse, 1.5 mm.
long, puberulent-papillate; corolla minutely puberulent-papillate outside, the tube
2 mm. long, the lobes oblong-elliptic, 4 mm. long; anthers glabrous, wholly exserted.
Forsteronia spicata (Jacq.) G. Meyer, Fl. Esseq. 135. 1818.
Echites spicata Jacq. Enum. PI. Carib. 13. 1760.
In thickets, about 200 m.; Zacapa (Gualan, Deam 6368). Southern
Mexico; Honduras and El Salvador to Nicaragua; Cuba; Colombia.
A large or small, woody vine; leaves firm-membranaceous, on petioles 4-10 mm.
long, broadly oval or obovate-elliptic, 6-16 cm. long, 3.5-9 cm. broad, very shortly
and abruptly subcaudate-acuminate, broadly obtuse or rounded at the base, mi-
nutely pilosulous above, inconspicuously glandular at the base of the costa, minutely
tomentulose beneath; inflorescences thyrsiform and often spike-like, terminal and
lateral, shorter than the leaves, very dense, the flowers white, sessile or nearly so;
bracts ovate, 1-5 mm. long, subfoliaceous; calyx lobes ovate, acute to acuminate,
2.5-4 mm. long, densely tomentulose outside; corolla glabrous or nearly so, the
tube 1.5-2 mm. long, the lobes oblong-ovate, 3.5-4 mm. long; anthers glabrous,
wholly exserted; ovary tomentulose; follicles essentially united, relatively stout and
rigid, 12-19 cm. long, glabrate in age.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
353
FIG. 98. Forsteronia viridescens. A, habit, X %; B, flower, X 5; C, corolla
dissected, X 10; D, stamens in natural position, X 15; E, pistil with surrounding
disk, X 15.
Forsteronia viridescens Blake, Contr. Gray Herb. 52: 80. 1917.
Wet thickets or forest, sometimes in Manicaria swamps, at or
little above sea level; Izabal. British Honduras (type from Manatee
Lagoon, Peck 450).
A small or large vine; leaves coriaceous or subcoriaceous, on petioles 3-6 mm.
long, oblong-elliptic, 9-13 cm. long, 2.5-6 cm. broad, very shortly and abruptly
354 FIELDIANA: BOTANY, VOLUME 24
acuminate, broadly obtuse or rounded at the base, glabrous, glandular above at
the base of the costa; inflorescence thyrsiform, terminal and sometimes also lateral,
shorter than the leaves, the flowers very numerous, cream-colored or greenish
white, the pedicels 1-1.8 mm. long; calyx lobes ovate to ovate-lanceolate, 1 mm.
long, minutely puberulent-papillate; corolla densely puberulent-papillate, the tube
1.5 mm. long, the lobes broadly oblong to ovate-oblong, 2 mm. long; anthers
slightly exserted at the tips, minutely and sparsely barbellate.
Called "tietie" in British Honduras.
HAPLOPHYTON A. De Candolle
Plants slender, chiefly herbaceous but usually suffrutescent near the base;
leaves mostly alternate, eglandular; flowers yellow, rather large, mostly solitary
in the upper leaf axils; calyx 5-parted, the lobes subequal, not or scarcely imbricate,
without squamellae; corolla salverform, not appendaged within, the limb equally
5-parted, sinistrorsely contorted; anthers not connivent, wholly included, the con-
nective not enlarged; ovary of 2 distinct carpels, without a nectary, many-ovulate;
follicles terete, elongate; seeds numerous, comose at the apex.
The genus consists of a single species.
Haplophyton cinereum (A. Rich.) Woodson, Ann. Mo. Bot.
Gard. 23: 231. 1936. Echites cinerea A. Rich, in Sagra, Hist. Cub.
11: 93. 1850. Haplophyton cimicidum A. DC. in DC. Prodr. 8: 412.
1844.
Moist or dry thickets, often in fence rows, 200-1,300 m.; El Pro-
greso; Huehuetenango ; Zacapa. Southwestern United States and
Mexico. The type is thought to have been from Cuba, doubtless in
error.
Plants mostly 50 cm. high or less, often much branched, the stems green; leaves
membranaceous, alternate or occasionally opposite, on very short petioles, ovate to
narrowly oblong-elliptic, 2-6 cm. long, 1-3.5 cm. broad, long-acuminate, obtuse or
rounded at the base, pilose on both surfaces with short, rather rigid, subappressed
hairs; flowers pedicellate; calyx lobes linear-lanceolate, acuminate, 5-8 mm. long,
thinly pilosulous; corolla minutely puberulent outside, pale yellow, the tube 6-
9 mm. long, the lobes broadly obovate, 10-18 mm. long, spreading; anthers inserted
near the middle of the corolla tube or somewhat lower; follicles slender, terete,
6-8 cm. long, puberulent or glabrate.
In Mexico this plant has long been known to have insecticide
properties, and its Nahuatl name, "actimpatli," signifies "flea-killer."
The roots are employed for killing flies, lice, fleas, cockroaches, and
other insects. In Guatemala the plant is abundant in many localities
about Zacapa, especially where it is protected from grazing. The
plants wither, at least in part, during the long dry season of that area,
and may be found green only when there is plenty of moisture.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
355
FIG. 99. Haplophyton cinereum. A, habit, natural size; B, flower, X 2;
C, calyx with pistil, X 6; D, stigma, X 15; E, follicles, natural size; F, stamens in
position on dissected corolla, X 10.
LACMELLEA Karsten
Reference: Joseph Monachino, A revision of Lacmellea and the
transfer of Zschokkea, Lloydia 7: 275-302. 1944.
Trees with milky latex; leaves opposite, the petioles usually glandular at the
base; inflorescences alternate-axillary or opposite-axillary, cymose, few-many-
flowered, the flowers small, white; calyx 5-parted almost to the receptacle, the
lobes subequal, imbricate, without squamellae; corolla salverform, the tube usually
long and slender, usually slightly gibbous, the limb equally 5-parted, the lobes
356 FIELDIANA: BOTANY, VOLUME 24
short, sinistrorsely contorted; anthers inserted near the orifice of the corolla tube,
not connivent, included, the connective not enlarged; ovary of 2 united carpels,
surrounded by an almost completely adnate, annular nectary, each carpel with
several biseriate ovules on an axile binate placenta; fruit a small juicy berry, con-
taining 1-several seeds.
About 19 species, in tropical America. One other is known from
Panama.
Lacmellea standleyi (Woodson) Monachino, Lloydia 7: 285.
1944. Zschokkea standleyi Woodson in Standl. Field Mus. Bot. 22:
44. 1940. L. edulis Woodson, N. Am. Fl. 29: 141. 1938, not Karst.
Palo de vaca.
Wet mixed lowland forest, sometimes in open pasture land, 300 m.
or lower; Izabal (type from Entre Rios, Standley 72587; also in Mon-
tana del Mico) ; Alta Verapaz, British Honduras.
A glabrous tree of 5-11 m.; leaves on petioles 6-10 mm. long, chartaceous, ob-
long or elliptic-oblong, 10-15 cm. long, 3-5 cm. broad, abruptly acuminate, obtuse
or rounded at the base; inflorescences axillary, peduculate, 5 cm. long or less, many-
flowered, the stout pedicels 3 mm. long or less; calyx lobes ovate-subreniform,
rounded at the apex, 2-2.5 mm. long, ciliolate; corolla white or greenish white, the
tube 2-2.5 cm. long, glabrous outside, the lobes obliquely ovate-lanceolate, acumi-
nate, 8 mm. long, spreading; anthers narrowly oblong, 6 mm. long; fruit broadly
ovoid, yellow, 1.5-2 cm. long, 1-seeded.
The fruit is said to have the odor of mangos. It probably is ed-
ible. The wood is soft and light in weight. Known in British Hon-
duras as "vaca" or "palo de vaca," "vaca tree" or "prickly vaca."
The last name alludes to the fact that the trunk is covered with
woody prickles. The abundant latex obtained from incisions in the
trunk is said to be drunk sometimes, although not altogether agree-
able in consistency.
LAUBERTIA A. De Candolle
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 23: 370-
375. 1936.
Slender vines, herbaceous or suffrutescent; leaves opposite or rarely verticillate
above, eglandular; inflorescence alternate-axillary, occasionally terminal or sub-
terminal, dichotomously or trichotomously scorpioid, bearing few-many flowers of
medium size; calyx 5-parted almost to the receptacle, the lobes equal or subequal,
somewhat foliaceous, scarcely imbricate, without squamellae within; corolla salver-
form, the tube not appendaged within, spirally contorted, the limb regularly 5-
parted, dextrorsely contorted; anthers connivent and agglutinated to the stigma;
the connective enlarged, narrowly bilobate, usually slightly exserted at the apex;
ovary of 2 distinct carpels, many-ovulate, surrounded at the base by 5 separate or
FIG. 100. Lacmellea standleyi. A, habit, X 1A; B, corolla opened to show sta-
mens, X 1H; C, calyx and style, X 1^; D, stigma, X 5; E, fruit, natural size;
F, an anther from the side and another from front, X 5; G, pseudospines from the
trunk of the tree, natural size.
357
358 FIELDIANA: BOTANY, VOLUME 24
somewhat concrescent nectaries; stigma fusiform-capitate; follicles 2, distinct,
terete; seeds numerous, truncate and comose at the apex.
Four species, distributed from Mexico to Peru. Only one is
known in Central America.
Laubertia peninsularis Woodson, Ann. Mo. Bot. Gard. 23: 374.
1936.
Known only from the type, collected somewhere along the bound-
ary between Pete"n and British Honduras, W. A. Schipp.
Leaves firm-membranaceous, on petioles 2 cm. long, ovate-elliptic, 6-13 cm.
long, 4-7 cm. broad, acuminate, rounded or very obscurely cordate at the base,
minutely puberulent or glabrate above, minutely ferruginous-puberulent beneath ;
inflorescences dichotomous or trichotomous, somewhat shorter than the leaves, 10-
20-flowered, the pedicels 8-10 mm. long; calyx lobes oblong-elliptic, acute or acu-
minate, 9-11 mm. long, subfoliaceous, minutely hirtellous; corolla cream-colored,
minutely ferruginous-hirtellous outside, the tube 13-14 mm. long, spirally con-
torted above, the lobes oblique-obovate, 9-11 mm. long, reflexed; stamens inserted
about the middle of the corolla tube, the anthers minutely puberulent-papillate
dorsally, barely included; nectaries somewhat shorter than the ovary.
LOCHNERA Reichenbach
Erect herbs, annual or sometimes more enduring, somewhat succulent; leaves
opposite, eglandular; inflorescences lateral, cymose, 1-4-flowered, sessile, the flow-
ers rather large and showy, white or pink; calyx 5-parted, the lobes almost equal,
subfoliaceous, without squamellae; corolla salverform, the limb equally 5-parted;
anthers not connivent, barely included, the connective not enlarged; ovary of 2
distinct carpels, accompanied by alternate oblong-ovoid nectaries of almost equal
size, many-ovulate; follicles terete, distinct; seeds numerous, naked, subcompressed.
Three species, natives of Madagascar, one of them cultivated and
widely naturalized in tropical regions.
Lochnera rosea (L.) Reichb. Consp. 134. 1828. Vinca rosea L.
Syst. Nat. ed. 10. 944. 1759. Catharanthus roseus G. Don, Gen. Hist.
4: 95. 1838. V. rosea ft albiflora Bertol. Fl. Guat. 411. 1840 (type from
Guatemala). Chatas; chula; chatilla; lila (Pete*n).
Planted commonly in gardens for ornament, mostly at low but
sometimes at middle elevations; thoroughly naturalized in many
localities, especially in sand close to seashores, along roadsides, in
abandoned land, or in waste places, chiefly at 1,400 m. or less; Pete"n;
Izabal; Jalapa; Santa Rosa; Retalhuleu; San Marcos; doubtless to
be found wild in other departments. Generally naturalized at lower
elevations. Mexico; British Honduras to Panama; and in tropical
America generally.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 359
Plants stout, erect, usually less than 75 cm. high; leaves on petioles 10 mm.
long or usually shorter, broadly oblong-elliptic to ovate-elliptic, 2-7 cm. long, 1.5-
3 cm. broad, very obtuse or rounded at the apex, cuneate at the base, rather succu-
lent, densely short-pilose, especially beneath, or sometimes glabrate; inflorescences
produced in alternate leaf axils, the pedicels 1.5-3 mm. long; calyx lobes narrowly
lanceolate, acuminate, 4-7 mm. long, minutely pilosulous; corolla pink, the tube
2-3 cm. long, the lobes broadly obovate, 1.5-2.5 cm. long, spreading; follicles
rather short and stout, terete, 1.5-3.5 cm. long.
Sometimes called "chuladita" in El Salvador, where the plant is
a common domestic remedy for inflammation of the throat; "clavel-
lina" (Honduras); "vicaria" (Yucatan); "paragiiita" (Oaxaca). The
plant is seen in the majority of Guatemalan gardens, and often is
planted in the parks. It is popular probably because it thrives with
little or no attention, and withstands drought. It is particularly
plentiful along the Atlantic coast of Central America, usually grow-
ing in the shade of coconut palms. It is one of the most common
flowers in cemeteries. The following color forms are frequent: L. ro-
sea f. alba (Sweet) Woodson, the corolla white throughout; L. rosea
f. ocellata (Sweet) Woodson, the corolla white with a pink or deep
red eye. The various color forms often grow together.
MALOUETIA A. De Candolle
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 22: 238-
270. 1935.
Shrubs or small trees; leaves opposite, eglandular, usually pitted beneath in
the axils of the nerves; inflorescence umbellate, terminal or lateral, the flowers small
or medium-sized; calyx 5-parted almost to the receptacle, the lobes subequal, bear-
ing within alternate, solitary or binate squamellae; corolla salverform, not append-
aged within, often somewhat thickened at the orifice, the limb regularly 5-parted,
dextrorsely contorted; anthers connivent and agglutinated to the stigma, usually
more or less exserted, the connective narrowly bilobate; ovary of 2 distinct carpels,
many-ovulate, surrounded at the base by 5 distinct or more or less concrescent nec-
taries; stigma fusiform; follicles distinct, narrowly terete to broadly fusiform; seeds
numerous, not comose.
About 20 species in tropical America and chiefly in South Amer-
ica. Only one species has been found in continental North America.
Malouetia guatemalensis (Muell.-Arg.) Standl. Journ. Wash.
Acad. Sci. 15: 459. 1925. Stemmadenia guatemalensis Muell.-Arg.
Linnaea 30: 410. 1860. M. panamensis Van Heurck & Muell.-Arg.
in Van Heurck, Obs. Bot. 185. 1871.
Wet mixed forest or thickets, sometimes in second growth, at or
little above sea level; Izabal. British Honduras to Panama.
FIG. 101. Malouetia guatemalensis. A, habit, X M>; B, flower, X 5; C, calyx
(2 lobes), pistil with surrounding corona and pistil, X 10; D, follicle, X 1A', E, sta-
mens to show attachment to corolla tube; F, seed, much enlarged.
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 361
Often flowering when only a shrub, but sometimes becoming a tree of 12 meters
or more with a trunk 35 cm. in diameter; leaves on petioles 5-10 mm. long, firm-
membranaceous, oblong-elliptic to ovate-elliptic, 6-25 cm. long, 2-10 cm. broad,
long-acuminate, acute at the base, glabrous, very lustrous above, paler beneath;
flowers slightly fragrant, white, in small many-flowered, often dense and almost
head-like, lateral and terminal umbels; pedicels 3-5 mm. long; calyx lobes ovate,
acute or obtuse, subcoriaceous, closely imbricate, 1.5-2.5 mm. long, minutely puber-
ulent-papillate; corolla glabrous or nearly so, the tube 4-5.5 mm. long, the lobes
obliquely lanceolate to ovate-oblong, acuminate, 7-12 mm. long; anthers included,
minutely puberulent-papillate dorsally; follicles stout, fusiform, divaricate, 10-
13 cm. long, 1-2 cm. thick, glabrous.
Although named guatemalensis, this species probably was based
upon material collected in Nicaragua by Friedrichsthal. The local-
ity is cited as "Mniogalpa," certainly not a Guatemalan locality, but
either Honduran or Nicaraguan. All Friedrichsthal's plants have
labels with the heading "Guatemala," but the place names prove
that they came from various Central American countries.
MANDEVILLA Lindley
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 20: 645-
777. 1933.
Slender vines, herbaceous or suffrutescent; leaves normally opposite, bearing
on the upper surface several glands clustered at the base or sparsely distributed
along the costa, rarely eglandular; inflorescences usually alternate-axillary, some-
times terminal or sub terminal, racemose or rarely obscurely compound; calyx 5-
parted almost to the receptacle, the lobes subequal, usually little imbricate, bearing
squamellae within; corolla salverform, funnelform, or tubular, the limb equally
5-parted, the lobes dextrorsely contorted; anthers connivent and agglutinated to
the stigma, the connective enlarged, obtusely bilobate or truncate; ovary of 2 dis-
tinct carpels, each carpel many-ovulate, surrounded by 5 or rarely 2, separate or
variously concrescent nectaries; stigma umbraculiform; follicles terete, acuminate,
distinct, continuous or moniliform; seeds numerous, truncate, comose.
About 100 species, in tropical America. One other Central Amer-
ican species is found in Costa Rica and in Panama.
Corolla tube more or less gibbous or arcuate; squamellae as many as the calyx
lobes and opposite them, sometimes deeply lacerate; upper surface of the
leaves sparsely glandular along the costa.
Corolla funnelform; bracts of the inflorescence foliaceous or petaloid. .M. hirsuta.
Corolla salverform.
Bracts scarious, 1-5 mm. long M. subsagittata.
Bracts foliaceous or petaloid, 10-35 mm. long M. villosa.
Corolla tube straight, not gibbous or arcuate; squamellae of the calyx usually more
numerous than the calyx lobes, or alternate with them when of the same num-
ber; upper surface of the leaves glandular at the base of the costa, or very
rarely eglandular; corolla salverform.
362 FIELDIANA: BOTANY, VOLUME 24
Stamens inserted near the orifice of the corolla tube; anthers with truncate
auricles.
Inflorescence secund; leaves broadly ovate to ovate-oblong.
M. donnell-smithii.
Inflorescence not secund; leaves lanceolate to oblong-lanceolate. .M. tubi flora.
Stamens inserted about the middle of the corolla tube; anthers with rounded
auricles.
Inflorescence secund and subscorpioid; leaves obovate to ovate-lanceolate,
densely tomentulose beneath M. scorpioidea.
Inflorescence neither secund nor subscorpioid; leaves lanceolate, glabrous.
M. rosana.
Mandevilla donnell-smithii Woodson, Ann. Mo. Bot. Gard.
19: 54. 1932. Reguilete (fide Aguilar).
Moist or dry, brushy, often rocky slopes, 1,100-2,000 m.; Baja
Verapaz; Zacapa; Chiquimula; Quiche; Jalapa; Santa Rosa; Guate-
mala; Sacatepe"quez; Huehuetenango. Mexico; Honduras; Nicaragua.
A slender woody vine; leaves on petioles 7-25 mm. long, membranaceous,
broadly ovate to ovate-oblong, 4-10 cm. long, 2-8 cm. broad, acute or acuminate,
rounded and shallowly cordate at the base, hirtellous above, pale or whitish be-
neath and densely tomentose; racemes alternate-axillary, equalling or somewhat
longer than the leaves, 10-25-flowered, the pedicels 5-7.5 mm. long; calyx lobes
ovate or ovate-lanceolate, acute or acuminate, 3-4 mm. long, minutely puberulent,
the squamellae very numerous; corolla salverform, yellow, glabrous outside, the
tube 12-15 mm. long, the lobes obliquely obovate, 2.5-4 mm. long; follicles very
slender, 8 cm. long or more, glabrous or nearly so, striate, obscurely torulose.
This has been reported from Guatemala as Echites tubiflora Mart.
& Gal. and E. triflora Mart. & Gal.
Mandevilla hirsuta (A. Rich.) Schum. in Engler & Prantl Nat.
Pflanzenfam. IV. 2: 171. 1895. Echites hirsuta A. Rich. Act. Soc.
Hist. Nat. Paris 1: 107. 1792. E. tomentosa Vahl, Symb. Bot. 3: 44.
1794. E. fluminensis A. DC. Prodr. 8: 452. 1844. M. tomentosa
Kuntze, Rev. Gen. 2: 416. 1891. M. fluminensis Donn.-Sm. Enum.
PL Guat. 2: 47. 1891. M. denticulata Blake, Contr. Gray Herb. 52:
81. 1917 (type from British Honduras, Peck 696). Bejuco de culebra.
Moist or wet, mixed forest or in thickets, sometimes in lowland
pine forest, 1,100 m. or less; Alta Verapaz; Izabal. British Honduras
to Panama; southward to Bolivia and Brazil.
A small or large vine, usually suffrutescent; leaves membranaceous, on petioles
2.5 cm. long or less, obovate-elliptic to oblong-elliptic, 5-20 cm. long, 2-8 cm.
broad, abruptly short-acuminate, obscurely auriculate at the base, strigillose above,
glandular along the costa, densely tomentulose or whitish-hirtellous beneath; ra-
cemes alternate-axillary, simple, about equalling the leaves, 5-25-flowered, the
pedicels 2-5 mm. long; bracts petaloid or foliaceous, ovate or ovate-lanceolate,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
363
FIG. 102. Mandevilla donnell-smithii. A, habit, X K; B, flower, X 2; C, corolla
dissected to show stamens in natural position, their attachment and patches of
pubescence on inner face of corolla tube, X 3; D, much enlarged under surface of
leaf to show pubescence.
caducous, 5-20 mm. long; calyx lobes lanceolate or ovate-lanceolate, acuminate,
5-10 mm. long, scarious or somewhat petaloid, each bearing a single opposite
squamella within; corolla funnelform, pilose outside, greenish yellow or yellowish
white, the throat red-purple, the tube somewhat gibbous, 2-3.5 cm. long, the throat
conic or conic-campanulate, 1.5-2 cm. long, the lobes obliquely obovate, 1.5-2 cm.
long; nectaries half as long as the ovary; follicles rather stout, 6-15 cm. long, con-
spicuously articulate, hispidulous or glabrate.
Mandevilla rosana (Donn.-Sm.) Woodson, Ann. Mo. Bot. Gard.
19: 56. 1932. Echites rosana Donn.-Sm. Bot. Gaz. 40: 6. 1905.
Type from Buena Vista, Santa Rosa, 1,000 m., Heyde & Lux 454-0.
364 FIELDIANA: BOTANY, VOLUME 24
A suffrutescent vine, glabrous or nearly so; leaves on petioles 3-5 mm. long,
firm-membranaceous, lanceolate or narrowly ovate-lanceolate, 6-12 cm. long, 1.5-
3 cm. broad, acuminate, obscurely cordate at the base; racemes alternate-axillary
or subterminal, simple, equalling or sometimes surpassing the leaves, 6-15-flowered,
the pedicels 7-10 mm. long; calyx lobes ovate-lanceolate, 3-4 mm. long, the squa-
mellae in alternate groups of 5-6; corolla salverform, glabrous outside, the tube
14-16 mm. long, the lobes obliquely obovate-oblong, 6-7 mm. long; stamens in-
serted about the middle of the corolla tube, the anthers with rounded auricles.
A specimen of this species, without locality, is in the Sesse and
Mocino Herbarium (1787-1804). It may be a part of the collection
said to have been made in Guatemala by those collectors or it may
have been gathered in Mexico.
Mandevilla scorpioidea Woodson, Ann. Mo. Bot. Gard. 19:
56. 1932. M. subscorpoidea Woodson, Ann. Mo. Bot. Gard. 20: 653.
1933, lapsus.
In thickets, about 1,350 m.; Alta Verapaz; Quezaltenango; El
Progreso (?); Zacapa (?). Mexico (type from Cerro del Boqueron,
Chiapas) ; Honduras.
Leaves membranaceous, on petioles 4-10 mm. long, ovate to ovate-lanceolate,
4-14 cm. long, 1.5-7 cm. broad, acuminate, narrowly cordate at the base, hirtellous
or hispidulous to glabrate above, densely and softly pilosulous beneath; inflores-
cence racemose secund, simple, alternate-axillary, about equalling the leaves, 15-
40-flowered, the pedicels 7-10 mm. long; calyx lobes lanceolate, acuminate, 4 mm.
long, sparsely hirtellous or glabrate, the squamellae numerous, indefinitely distrib-
uted; corolla salverform, glabrous outside, the tube 15-20 mm. long, the lobes
obliquely obovate, 4-5 mm. long; stamens inserted about the middle of the corolla
tube, the anthers 4 mm. long, with rounded auricles.
This species is perhaps not distinct from M . tubiflora (Mart. &
Gal.) Woodson.
Mandevilla subsagittata (Ruiz & Pavon) Woodson, Ann. Mo.
Bot. Gard. 19: 69. 1932. Echites subsagittata Ruiz & Pavon, Fl.
Peruv. 2: 19. 1799. E. microcalyx A. DC. in DC. Prodr. 8: 456. 1844.
E. cuspidifera Blake, Contr. Gray Herb. 52: 79. 1917 (type from
Manatee Lagoon, British Honduras, Peck 35). Bejuco pie de rana
(fide Aguilar) .
Wet to dry thickets, often in second growth, 1,800 m. or lower;
Pete*n; Alta Verapaz; Baja Verapaz; Izabal; Zacapa; Chiquimula;
Jalapa; Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez ;
Quiche" ; Huehuetenango. Southern Mexico; British Honduras to
Panama; northern South America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 365
A large or small, slender suffrutescent vine, glabrous or variously pubescent;
leaves membranaceous, on petioles 5-10 mm. long, oblong-elliptic to rarely nar-
rowly lanceolate, 2-10 cm. long, 0.5-3 cm. broad, abruptly or gradually acuminate,
rarely obtuse or rounded at the apex, auriculate at the base, glandular above along
the costa, usually (in Central American material) densely and softly pilosulous
beneath; racemes alternate-axillary, simple, more or less secund, about equalling
the leaves, 8-20-flowered, the pedicels 4-6 mm. long; bracts scarious, lanceolate,
1-5 mm. long; calyx lobes narrowly triangular, 1-1.5 mm. long, each bearing a
single squamella within; corolla salverform, bright yellow, the tube more or less
gibbous or ventricose, 2-2.5 cm. long, the lobes obliquely obovate, 1-1.5 cm. long;
nectaries 5, half as long as the ovary; follicles slender, conspicuously moniliform,
10-20 cm. long.
Mandevilla tubiflora (Mart. & Gal.) Woodson, Ann. Mo. Bot.
Gard. 19: 52. 1932. Echites tubiflora Mart. & Gal. Bull. Acad. Brux.
11, pt. 1: 358. 1844. E. cobanensis Donn.-Sm. Bot. Gaz. 40: 6. 1905
(type from Coban, Alta Verapaz, Tuerckheim 8709).
Wet to dry thickets or open forest, 500-2,000 m.; Alta Verapaz;
Chiquimula; Jalapa; Solola; Huehuetenango; Quezaltenango. South-
ern Mexico; Honduras.
Plants perennial from a thick woody root, scandent, herbaceous or suffrutes-
cent; leaves on petioles 4-10 mm. long, membranaceous, lanceolate or oblong-
lanceolate, 4-10 cm. long, 1-4 cm. broad, acuminate, obscurely cordate at the base,
minutely puberulent or glabrate above, pale and densely tomentose beneath; ra-
cemes alternate-axillary, about equalling the leaves, 8-20-flowered, the pedicels
7-10 mm. long; calyx lobes ovate or ovate-lanceolate, acute or acuminate, 1-2 mm.
long, minutely puberulent or glabrate, the squamellae in alternate groups of 5-6;
corolla salverform, lemon-yellow or greenish yellow, glabrous outside, the tube 1-
1.5 cm. long, the lobes obliquely obovate, 2.5-3.5 mm. long.
Mandevilla villosa (Miers) Woodson, Ann. Mo. Bot. Gard. 19:
70. 1932. Laseguea villosa Miers, Apocyn. S. Amer. 250. 1878.
Echites comosa Kuntze, Rev. Gen. 2: 414. 1891.
In thickets, 450 m. or less; Solola; Retalhuleu. Mexico (Chia-
pas); El Salvador to Panama. Venezuela.
A slender suffrutescent vine; leaves membranaceous, on petioles 4-25 mm. long,
elliptic or obovate-elliptic, 3-9 cm. long, 1.5-4 cm. broad, rather abruptly acumi-
nate, obscurely auriculate or almost subhastate at the base, finely and sparsely
pilose above or glabrate, glandular along the costa, finely and densely pilose be-
neath or glabrate; racemes simple, alternate-axillary, secund, about equalling the
leaves, 8-20-flowered, the pedicels 2-4 mm. long; bracts petaloid or foliaceous, de-
ciduous, lanceolate to ovate-lanceolate, 1-3.5 cm. long; calyx lobes triangular,
acute or acuminate, 1-1.5 mm. long, each bearing an opposite squamella within;
corolla salverform, finely pilosulous or glabrate, the tube somewhat gibbous or
ventricose, 1.5-2 cm. long, the lobes obliquely obovate-oblong, 1-1.5 cm. long;
366 FIELDIANA: BOTANY, VOLUME 24
nectaries 5, half as long as the ovary; follicles slender, conspicuously moniliform,
10-15 cm. long, glabrous.
This has been reported from Guatemala as M. moritziana (Muell.-
Arg.) Donn.-Sm.
Many new collections of the complex to which this species be-
longs seem to indicate that this species is hardly distinct from M.
subsagittata (Ruiz & Pavon) Woodson.
MESECHITES Mueller von Argau
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 20: 629-
645. 1933.
Slender vines, herbaceous or suffrutescent; leaves opposite, bearing 1-4 glan-
dular emergencies above at the base of the costa; inflorescences alternate-axillary,
the axis dichotomously or rarely trichotomously divided, bearing several or few,
congested, medium-sized flowers; calyx 5-parted almost to the receptacle, the lobes
subequal, imbricate, bearing several alternate or indefinitely distributed squamel-
lae within; corolla salverform, the limb regularly 5-parted, the lobes dextrorsely
contorted, reflexed; anthers connivent and agglutinated to the stigma, without
apical appendages, the connective enlarged, obtusely bilobate; ovary of 2 distinct
carpels, each carpel many-ovulate, surrounded by 5 distinct or somewhat concres-
cent nectaries; stigma fusiform-umbraculif orm ; fruit apocarpous, follicular; seeds
numerous, truncate and comose at the apex.
Ten species, all in tropical America. Only the following is found
in continental North America.
Mesechites trifida (Jacq.) Muell.-Arg. in Mart. Fl. Bras. 61:
151. 1860; Echites trifida Jacq. Enum. PL Carib. 13. 1760. Reguilete
(fide Aguilar) .
Wet to dry thickets, 1,500 m. or less; Pete"n; Alta Verapaz; Za-
capa; Chiquimula; Jutiapa; Santa Rosa; Guatemala; Huehuetenango.
Southern Mexico; British Honduras to Panama; South America.
A small or large, glabrous, suffrutescent vine; leaves on petioles 5-30 mm. long,
firm-membranaceous, ovate to oblong or oblong-lanceolate, 5-12 cm. long, 2-8 cm.
broad, acuminate to obtuse, usually mucronulate, obtuse to rounded or subcordate
at the base; inflorescences axillary or rarely sub terminal, half as long as the leaves,
few-many-flowered, the pedicels 5-10 mm. long; calyx lobes broadly oblong, obtuse
or rounded at the apex, 3-5 mm. long; corolla greenish white, the tube 1.5-2.5 cm.
long, the lobes obliquely oblong-obovate, 7-15 mm. long; follicles very slender,
continuous or nearly so, terete, 15-30 cm. long.
NERIUM L. Oleander
Shrubs or small trees; leaves usually ternate, coriaceous, eglandular; inflores-
cence thyrsiform, the flowers numerous, large and showy; calyx 5-parted almost to
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
367
FIG. 103. Mesechites trifida. A, habit, X ]/z\ B» corolla in natural position,
natural size; C, calyx and style, X 2; D, stamens in position on portion of corolla
tube, X 3; E, seed, X 3/2 ; F, node showing interpetiolar stipule, X 2.
the receptacle, the lobes equal, more or less imbricate, bearing numerous squamel-
lae within at the base; corolla funnelform, the throat with rather conspicuous,
mostly 5-cleft, petaloid appendages, the limb regularly 5-parted, dextrorsely con-
torted; anthers connivent and agglutinated to the stigma, the tips exserted, the
connective enlarged, bilobate at the base; ovary of 2 distinct cells, many-ovulate,
without a nectary, the stigma fusiform; follicles 2, distinct, rather stout; seeds
numerous, compressed, densely puberulent, comose at the apex.
368 FIELDIANA: BOTANY, VOLUME 24
Three species, natives of the Mediterranean region and of Asia.
Nerium oleander L. Sp. PI. 209. 1753. Adelfa; narciso; oleander.
Planted commonly for ornament and sometimes naturalized, in
Guatemala except at high elevations, common at low altitudes;
native of the Mediterranean region.
A shrub or small tree, usually 6 m. high or less, glabrous or nearly so; leaves
opposite or in whorls of 3-4, short-petiolate, coriaceous, oblong-lanceolate or linear-
lanceolate, 6-25 cm. long, acuminate, attenuate to the base; inflorescence much
longer than the leaves, with few or numerous flowers, these often double, white to
pink or red; calyx lobes lanceolate or ovate-lanceolate, acuminate, 4-6 mm. long,
somewhat foliaceous; corolla glabrous outside, the tube 8-12 mm. long, the throat
conic-campanulate, 10 mm. long, the lobes obliquely obovate or obovate-oblong,
20-25 mm. long; follicles stout and thick, 8-15 cm. long.
The oleander is a popular ornamental shrub in Guatemala be-
cause of its attractive sweet-scented flowers, produced at all seasons
of the year. The plant contains alkaloids that act as a powerful
cardiac stimulant, and it has been employed in medicine as a heart
stimulant and tonic. It has long been used in southern Europe for
poisoning rats, and sometimes for killing people. An infusion of the
leaves in oil has been employed as a remedy for cutaneous diseases
and to destroy insect parasites. The sap is caustic to some persons.
The wood if used as a spit to roast meat may cause the meat to be-
come toxic. The plant is reported toxic to all classes of livestock and
15-20 grams are said to be sufficient to cause death in mature cattle
or horses.
ODONTADENIA Bentham
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 22: 270-
306. 1935.
Usually large woody vines; leaves opposite, eglandular, stipulate or exstipu-
late; inflorescence opposite-axillary, sometimes also terminal, thyrsiform to simply
scorpioid, with few to numerous, usually large and showy flowers; calyx 5-parted
almost to the receptacle, the lobes subequal or conspicuously unequal, closely im-
bricate, bearing within 5-many alternate or indefinitely distributed squamellae;
corolla funnelform or rarely almost salverform, the tube not appendaged within,
the limb regularly 5-parted, dextrorsely contorted; anthers connivent and agglu-
tinated to the stigma, the connective enlarged, narrowly bilobate; ovary of 2 dis-
tinct carpels, many-ovulate, surrounded at the base by 5 nectaries, these usually
concrescent and irregularly lobate or lacerate; stigma fusiform to subcapitate; fol-
licles 2, distinct, terete or dorsally compressed; seeds numerous, truncate and comose
at the apex.
FIG. 104. Odontadenia caudigera. A, habit, X 1A', B, calyx and pistil show
digitiform squamellae, X 2; C, portion of corolla tube to show stamens, one re-
versed, X 2.
369
370 FIELDIANA: BOTANY, VOLUME 24
About 25 species, in tropical America. Two other species are
known from southern Central America. The genus is perhaps too
closely related to Mandevilla.
Inflorescence not thrysiform; calyx lobes equal or nearly so; stipules absent or
minute O. caudigera.
Inflorescence thyrsiform; calyx lobes very unequal, the outer ones shorter; stipules
present but caducous 0. schippii.
Odontadenia caudigera Woodson, Ann. Mo. Bot. Gard. 23:
384. 1936.
Wet thickets or forest, at or little above sea level; Izabal; Alta
Verapaz. British Honduras (type collected by W. A. Schipp, the
exact locality unknown) ; Costa Rica.
A large woody vine, glabrous throughout, sometimes 10 m. long or more;
leaves opposite, membranaceous, on petioles 1.5-2.5 cm. long, elliptic or oblong-
elliptic, 10-20 cm. long, 6-11 cm. broad, subcaudate-acuminate, obtuse or almost
rounded at the base, the lateral nerves prominent and very conspicuous beneath;
inflorescences terminal and opposite-axillary, obscurely compound, mostly 3-6-
flowered, the pedicels about 2 cm. long; bracts ovate, 2-3 mm. long; calyx lobes
broadly ovate, obtuse, 6-7 mm. long; corolla funnelform, greenish yellow, glabrous
outside, the tube 8-9 mm. long, the throat 16 mm. long, the lobes obovate-dolabri-
form, 20 mm. long; anthers densely hirtellous dorsally; nectaries concrescent,
deeply multifid, slightly longer than the ovary.
This has been reported from British Honduras as 0. hoffmann-
seggiana (Steud.) Woodson.
Odontadenia schippii Woodson, Ann. Mo. Bot. Gard. 22: 292.
1935.
Known only from the type, collected on the boundary between
Pete"n and British Honduras, about 820 m., Schipp S709, growing
in forest.
A large vine as much as 25 m. long, with a trunk 10 cm. in diameter; leaves
glabrous, on petioles 1-1.5 cm. long, firm-membranaceous, elliptic to oval, 7-12 cm.
long, 3-5.5 cm. broad, obtusely short-acuminate, obtuse at the base, glabrous,
somewhat lustrous above; inflorescences terminal and opposite-axillary, thyrsi-
form, equalling or somewhat longer than the leaves, several-flowered, the pedicels
2-2.5 cm. long; calyx lobes evidently unequal, the outer ones broadly ovate, obtuse
or rounded at the apex, 5-6 mm. long, coriaceous, densely and minutely puberu-
lent-papillate, the inner ones broadly oblong, 9-10 mm. long; corolla funnelform,
glabrous outside, creamy white, the tube 15-17 mm. long, the throat 2.5 cm. long,
the lobes obliquely dolabriform, 1.5 cm. long; anthers minutely hirtellous dorsally;
nectaries three times as long as the ovary, concrescent throughout.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 371
PLUMERIA L.
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 25: 202-
224. 1938.
Large shrubs or medium-sized trees, the branches usually very thick and exud-
ing abundant latex when broken; leaves alternate; inflorescence terminal or pseudo-
lateral, fastigiate-thyrsiform, often very dense and many-flowered, the flowers large
and showy, waxy; calyx 5-parted almost to the receptacle, the lobes subequal, with-
out squamellae but tipped with a glandular epithelium; corolla salverform, not ap-
pendaged within, the limb equally 5-parted; stamens wholly included, the anthers
not connivent, the connective not enlarged; ovary of 2 distinct carpels, many-
ovulate, not accompanied by a nectary; follicles 2, distinct, thick, the seeds numer-
ous, winged basally.
About 40 species have been described, all from tropical America.
Only the following are known from continental North America. The
generic name has been written Plumiera and Plumieria. It is dedi-
cated to Charles Plumier (1646-1704), a brother in the Franciscan
order, one of the first botanists to explore the flora of the New World,
particularly that of the Antilles.
Leaves acute or acuminate; inflorescence corymbose P. rubra.
Leaves rounded or emarginate at the apex; inflorescence congested and subum-
bellate P. obtusa.
Plumeria obtusa L. Sp. PI. 210. 1753. Native of Bahamas and
the Greater Antilles, also on Swan Island, Honduras; represented in
Central America by the following variety:
Plumeria obtusa var. sericifolia (C. Wright) Woodson, Ann.
Mo. Bot. Gard. 25: 214. 1938. P. sericifolia C. Wright ex Griseb.
Cat. PI. Cub. 171. 1866. P. multiflora Standl. Field Mus. Bot. 8: 33.
1930 (type from Yucatan). Flor de Mayo; flor de chombo.
In dry, open forests at or little above sea level; Pete*n. British
Honduras; Mexico (Yucatan); Bahamas; Cuba; Hispaniola.
A shrub or small tree, 6 m. high or less; leaves short-petiolate, coriaceous, obo-
vate to obovate-oblong, 5-18 cm. long, 2-8 cm. broad, rounded or emarginate at
the apex, cuneate-attenuate to the base, glabrous and lustrous above, minutely
and densely pubescent beneath; inflorescence congested and subumbellate, the
flowers few or numerous, white with a yellow center, the pedicels 7-10 mm. long;
calyx lobes ovate-triangular, 1-1.5 mm. long, rounded to truncate at the apex,
glabrous or pilosulose; corolla tube 1-2 cm. long, the lobes ovate-oblong or obovate,
obtuse or rounded at the apex, 1.5-4.5 cm. long; follicles stout, 7-24 cm. long,
1-2 cm. in diameter.
Called "zopilote" in British Honduras. This species is probably
not in cultivation in Central America. The Maya name of Yucatan is
"nichte chom," and the plant is employed there in domestic medicine.
372
FIELDIANA: BOTANY, VOLUME 24
FIG. 105. Plumeria rubra. A, habit, X 1A', B, inflorescence, X Y^', C, stamens
in place on dissected portion of corolla tube, X 3.
Plumeria rubra L. Sp. PL 209. 1753.
Widely cultivated in several color forms; native in rather dry
often rocky forest and mountain slopes, occasionally on plains or in
brushy savannas, usually at 1,500 m. or less, more commonly at 500-
1,000 m.
A large shrub or small tree, seldom more than 6 meters high, except wild trees
10-12 m.; leaves firm membranaceous, somewhat succulent when fresh, obovate to
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 373
elliptic-oblong or oblong-lanceolate, 12-50 cm. long, 3.5-15 cm. broad, acute or
acuminate, cuneate at the base, glabrous or nearly so, sometimes pubescent be-
neath, the petioles 1.5-10 cm. long; inflorescence corymbose, usually rather lax,
many-flowered, the flowers very fragrant, the pedicels 1-2 cm. long; calyx lobes
ovate-quadrate to ovate-deltoid, obtuse or truncate, 1-2 mm. long; corolla tube
1-2.5 cm. long, the lobes broadly obovate, rounded or obtuse at the apex, 2.5-6 cm.
long; follicles 9-30 cm. long, 1.5-4 cm. thick.
The following forms may be distinguished by those who wish to
do so:
f. acutifolia (Poir.) Woodson, Ann. Mo. Bot. Gard. 25: 211.
1938, as f. acutifolia (Ait.) Woodson. P. acutifolia Poir. Encycl.
Suppl. 2: 667. 1812. P. mexicana Lodd. Bot. Cab. t. 1024. 1825.
P. megaphylla A. DC. in DC. Prodr. 8: 391. 1844. Flor de la cruz;
flor de Mayo; palo de cruz; cumpap (Jacaltenango) ; nicte de monte;
matuhua (Pete"n).
Usually in rather dry, open rocky forest, brushy savannas or
plains, 1,500 m. or less, most common at 500-1,000 m.; Pete"n; Baja
Verapaz; Zacapa; Chiquimula; El Progreso; Jutiapa; Santa Rosa;
Guatemala; Sacatepe"quez ; Retalhuleu; San Marcos; Huehuetenango.
Mexico to Panama.
This is the white-flowered form, the center sometimes yellow,
usually a native tree but sometimes brought into cultivation.
In El Salvador sometimes called "flor blanca" and "flor de en-
sarta"; "zacnicte" (Yucatan, Maya). This wild form is abundant
in some places in Guatemala, especially along the drier Pacific foot-
hills, and in the lower Motagua Valley and in the "Oriente." Through
most of the dry season it is leafless, and flowers before the new leaves
appear. The trunk is almost white, and the whole tree more or less
distinctive in form, so that it can be recognized at a distance. When
covered with its large clusters of white flowers it is showy and very
attractive. It is not cultivated commonly in Central America, and
it is the only form found wild. The wild trees are often 10 m. tall,
with a trunk 25 cm. in diameter or sometimes more, the branches
strongly ascending. The wood is hard, compact, very fine textured,
yellowish brown with faint purplish streaks, giving it a pleasing ap-
pearance. The wood takes a high polish and is employed in some
regions for articles of turnery. In Guatemala the Indian people
sometimes employ an infusion of the flowers as a supposed remedy
for diseases of the chest. In British Honduras the tree is sometimes
called "Spanish jasmine" and "fringipanzi."
374 FIELDIANA: BOTANY, VOLUME 24
f. lutea (Ruiz & Pavon) Woodson, I.e. P. lutea Ruiz & Pavon,
Fl. Peruv. 2: 21, t. 142. 1799.
This is the yellow-flowered form. It is uncommon and is known
only from cultivation in Guatemala.
f. rubra. P. rubra L. Sp. PI. 209. 1753.
Cultivated as an ornamental at low and middle elevations around
the tropical world. Native of America and Phil Clark has seen it in
Mexico where he believed it to be native, otherwise the native habitat
not known and not recorded in the wild state by others. The flowers
are red or pinkish and from this fact comes the specific name of the
species. It may be a derivative of the forma acutifolia.
In Guatemala the plant is known as flor de cruz; flor de mayo;
nicte and nicte chachac (Pete"n, Maya); "zabacnicte," "chacnicte,"
and "cumpap" (Yucatan, Maya). The flowers of this and other
forms of the species are fragrant and attractive. A perfume, called
frangipanni, has been made from the flowers and the same name has
often been applied to the plant itself. The common name, flor de
mayo, alludes to the fact that the tree often flowers in May but they
are in flower commonly during other months. The corollas are rather
stiff and keep their form and color long after being removed from the
plants. On this account they are made into garlands, especially for
decoration of the crosses that are put up in many places in Central
America on Dia de la Cruz (Invention of the Cross) and from this
comes the common name flor de la cruz. The names fringipanni or
frangipanni is derived from the French frangipanier, coagulated milk,
in reference to the abundance of latex that pours from a broken branch.
f. tricolor (Ruiz & Pavon) Woodson, I.e. P. tricolor Ruiz &
Pavon, Fl. Peruv. 2: 20, L 39. 1799.
An infrequent color form in Guatemala which is known only from
cultivation. The corollas are white or pale yellow, with rose-mar-
gined lobes and a yellow center.
PRESTONIA R. Brown
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 23: 276-
367. 1936.
Woody or suffrutescent vines; leaves eglandular; inflorescences alternate-axil-
lary, rarely subterminal, racemose, simple or dichotomously or trichotomously
divided, often corymbose or subumbellate, bracteate, the flowers few or numer-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 375
ous, medium-sized or large; calyx 5-parted almost to the receptacle, the lobes
subequal, somewhat foliaceous, bearing an opposite solitary squamella within;
corolla salverform or rarely funnelform, the tube usually bearing 5 epistaminal
appendages within, the orifice bearing a more or less conspicuous callose annulus,
the limb regularly 5-parted, dextrorsely contorted; anthers connivent and aggluti-
nated to the stigma, the tips somewhat exserted, the connective enlarged, narrowly
bilobate; ovary of 2 distinct carpels, many-ovulate, surrounded at the base by 5
separate, more or less concrescent nectaries; stigma fusiform to subcapitate; fol-
licles 2, distinct or more or less agglutinated, narrowly terete to subf usif orm ; seeds
numerous, comose.
About 60 species, in tropical America. Nine or ten other species
are found in southern Central America and in adjacent Mexico.
There are eight species of this small genus in Guatemala, easily
divided into two groups by flowers, fruits and indument. The first
three in the following key are much alike and difficult to separate,
the remaining five are closely related also and it is doubtful if all are
distinct.
Another species to be expected in Guatemala and included here
is P. grandiflora L. Wms. described from Chiapas. It belongs in the
first section mentioned above but has flowers twice as large as any
of those species.
Corolla densely pubescent outside; leaves densely pubescent beneath.
Corolla funnelform, the tube dilated above.
Corolla 5-6 cm. long P. grandiflora.
Corolla 3-3.5 cm. long P. speciosa.
Corolla salverform, the tube cylindric.
Corolla 3.5-4.5 cm. long P. mexicana.
Corolla 2.8-3 cm. long P. amanuensis.
Corolla glabrous or merely papillate externally.
Calyx lobes small and inconspicuous, 1.5-2 mm. long, only slightly foliaceous,
usually more or less reflexed P. acutifolia.
Calyx lobes large and conspicuous, 4-20 mm. long, foliaceous or coriaceous to
petaloid, not reflexed.
Nectaries thick and fleshy throughout; calyx lobes 4-5 mm. long. .P. concolor.
Nectaries thin and more or less diaphanous, at least on the margins; calyx
lobes 8-17 mm. long.
Epistaminal appendages deeply included in the corolla tube .... P. schippii.
Epistaminal appendages exserted, or at least reaching the orifice of the
corolla.
Calyx lobes 7-9 mm. long, scarcely or not at all suffused with purple; in-
florescence 2-3 dichotomous P. guatemalensis.
Calyx lobes 11-17 mm. long, usually deeply suffused with purple; in-
florescence once divided or simple P. portobellensis.
Prestonia acutifolia (Benth.) Schum. in Engler & Prantl, Pflan-
zenfam. 4, pt. 2: 188. 1895. Haemadictyon acutifolium Benth. ex
Muell.-Arg. in Mart. Fl. Bras. 61: 167. 1860.
376 FIELDIANA: BOTANY, VOLUME 24
Dry thickets, about 120 m.; Retalhuleu (near Nueva Linda,
Standley 6654.8). Panama; southward to Argentina.
A woody vine, appearing glabrous; leaves on petioles 6-20 mm. long, thick-
membranaceous, elliptic to oblong-elliptic or ovate-elliptic, 6-16 cm. long, 2-8 cm.
broad, abruptly acute or acuminate, obtuse or rounded at the base, often veined
with red or purple when young, glabous above, minutely papillose-puberulent be-
neath; inflorescence racemose, simple, about equalling the leaves, 6-40-flowered,
the pedicels 5-12 mm. long, glabrous or minutely papillate; bracts ovate-lanceolate,
1-2 mm. long, subfoliaceous; calyx lobes narrowly ovate-lanceolate, acuminate,
1.5-2 mm. long, subfoliaceous, reflexed; corolla glabrous or minutely papillate,
greenish yellow, the tube 15-20 mm. long, the epistaminal appendages 0.5-2 mm.
long, wholly included, the lobes obliquely obovate, 7-10 mm. long; anthers mi-
nutely pubescent dorsally, the tips somewhat exserted; nectaries about equalling
the ovary; follicles slender, obscurely moniliform, agglutinated and united at the
apex, 20-40 cm. long, glabrous or nearly so.
Prestonia amanuensis Woodson, Ann. Mo. Bot. Gard. 23: 359.
1936.
Type from Stann Creek Railway, British Honduras, near sea
level, growing in open places, Schipp S7.
A woody vine 3 m. long, densely fulvous-hirtellous or somewhat tomentose
throughout, the pubescence rather harsh; leaves thick-membranaceous, almost
sessile, broadly ovate, 6-9 cm. long, 4-7 cm. broad, abruptly and shortly cuspidate-
acuminate from an often rounded apex, broadly and shallowly cordate at the base;
inflorescence dense, subumbellate, 6-12-flowered, much shorter than the leaves,
the pedicels 3-5 mm. long; bracts ovate-lanceolate, 3-7 mm. long; calyx lobes
ovate-lanceolate, acuminate, 11-17 mm. long, foliaceous, appressed-hirtellous; co-
rolla cream-colored, ferruginous-villous, the tube 20-22 mm. long, the epistaminal
appendages replaced by subquadrate protuberances, the lobes obliquely obovate,
8-9 mm. long; anthers glabrous, partly exserted; nectaries concrescent, annular,
essentially entire, about as long as the ovary.
This has been reported from British Honduras as P. mexicana,
from which it is not very distinct.
Prestonia concolor (Blake) Woodson ex Standl. & Record, Field
Mus. Bot. 12: 327. 1936. Belandra concolor Blake, Contr. Gray Herb.
52: 78. 1917.
Wet mixed forest, at or near sea level; Izabal (Rio Dulce, above
Livingston, Steyermark 39434). British Honduras (type from Rio
Grande, Peck 953}.
A somewhat woody vine, almost glabrous throughout; leaves subcoriaceous,
on petioles 6-9 mm. long, elliptic-oblong or oblong, 9-13 cm. long, 3-5 cm. broad,
short-acuminate, obtuse or rounded at the base, somewhat lustrous above, con-
spicuously reticulate- veined ; inflorescences racemose, simple, 30-40-flowered, the
FIG. 106. Prestonia concolor. A, habit, X Vi\ B, calyx and pistil, X 2; C, stig-
ma, X 5; D, stamens in place on corolla tube, one reversed to show back, X 5.
377
378 FIELDIANA: BOTANY, VOLUME 24
peduncles equalling or often much exceeding the leaves, the pedicels 7-10 mm. long;
bracts linear-lanceolate, minute; calyx lobes ovate-elliptic, acute or acuminate,
4-5 mm. long, membranaceous or subcoriaceous, deeply suffused with purple, mi-
nutely papillate outside; corolla greenish yellow, glabrous or very minutely papil-
late, the tube 15-18 mm. long, the epistaminal appendages wholly included, the
lobes obliquely obovate, 8-10 mm. long; anthers glabrous or very minutely puberu-
lent dorsally, slightly exserted; nectaries concrescent, thick and carnose, entire or
slightly undulate, somewhat surpassing the ovary.
This species was collected 170 years ago by Sesse" and Mocino,
possibly in Guatemala but more probably somewhere in southern
Mexico.
Prestonia grand iflora L. Wms. Field Mus. Bot. 31: 402. 1968.
Probably in moist thickets, type from about 2,000 m., La Gran-
doza, Chiapas, Mexico, Matuda 15570, to be expected in Guatemala.
Rampant woody vines. Branches terete, fulvous pubescent, glabrous with age,
7-8 mm. or possibly more in diameter. Leaves ovate to broadly ovate or broadly
obovate, abruptly acuminate, rounded to the base, petiole short, 1 cm. or less, with
8-10 lateral nerves, sparsely pubescent above except densely so along the nerves,
prominently fulvous-pubescent below, the blades 9-17 cm. long and 3.5-11 cm.
broad; stipules corneous, 4-6 along the base of the petiole and also interpetiolar,
2-3 mm. long; inflorescence axillary, corymbose and several times branched, bear-
ing few to several large yellow flowers, shorter than the subtending leaf, minutely
ferruginous pubescent, bracts lanceolate, to 10 mm. long; calyx divided to the base,
the lobes elliptic-lanceolate, ovate-lanceolate or lanceolate, acute or acuminate,
minutely fulvous pubescent, 12-14 mm. long and 3.5-5 mm. broad; corolla largest
of the genus, fulvous pubescent outside except on plicae, about 6 cm. long at ma-
turity, tubular-campanulate, the basal tube about 3.5-4 cm. long to the throat,
the apical portion somewhat expanded and containing the stamens, with an incon-
spicuous internal lamellate corona at the throat, the limb plicate, contorted in the
bud, when mature the lobes about 3 cm. long and up to 2 cm. broad, divided to
the throat, somewhat oblique and broadly obovate; stamens inserted below the
throat, anthers linear-lanceolate, sagittate, 8-9 mm. long and about 2 mm. broad
at the base, filaments about 2 mm. long, arched inward, attached peltately to the
base of the stamen; carpels 2, with a maniculate stigma reaching to the enclosing
anthers, subtended by 5 nearly separate nectaries; immature fruit densely fulvous
pubescent.
Prestonia guatemalensis Woodson, Ann. Mo. Bot. Gard. 23:
339. 1936.
Moist or wet thickets, 500-2,000 m.; Alta Verapaz (type from
Sepacuite", Owen 1); Santa Rosa (Volcan de Tecuamburro) ; Chimalte-
nango; Quezaltenango; San Marcos. Southern Mexico.
A suffrutescent vine, glabrous almost throughout; leaves firm-membranaceous,
on petioles 1-1.5 cm. long, broadly obovate-elliptic to oblong, 15-25 cm. long,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 379
7-10 cm. broad, abruptly short-acuminate, obtuse or rounded at the base; inflores-
cences lateral, 2-3-dichotomous, several-flowered, the peduncles 5-6 cm. long, the
pedicels 9-12 mm. long, glabrous or minutely papillate; calyx lobes elliptic, obtuse
or acute, 7-9 mm. long, subcoriaceous, pale green, minutely papillate outside;
corolla greenish yellow, glabrous outside, the tube 18-20 mm. long, the epistaminal
appendages somewhat exserted, the lobes obliquely obovate, 12-13 mm. long, often
purplish outside; anthers glabrous, their tips exserted; nectaries concrescent, mem-
branaceous, completely concealing the ovary; follicles slender, 30-50 cm. long,
glabrous.
This has been reported from Guatemala as P. macrocarpa Hemsl.
Prestonia mexicana A. DC. in DC. Prodr. 8: 429. 1844.
Moist or dry thickets or forest, 200-1,400 m.; Pete"n; Jutiapa;
Santa Rosa; Huehuetenango ; Quezaltenango (?). Southern Mexico;
British Honduras; El Salvador.
A woody vine, the older stems covered with corky, deeply sulcate or ridged,
pale bark, the plants densely fulvous-pubescent or tomentose throughout; leaves
firm-membranaceous, on petioles 3-9 mm. long, broadly ovate to oval or obovate-
elliptic, 7-23 cm. long, 4-15 cm. broad, acute or more often rounded and abruptly
short-acuminate, rounded or subcordate at the base, densely tomentose beneath;
inflorescence densely umbellate, simple, 8-20-flowered, the peduncles much shorter
than the leaves, the pedicels 5-10 mm. long; bracts ovate-lanceolate, 6-17 mm.
long, foliaceous; calyx lobes ovate-lanceolate to oblong-lanceolate, acute or acumi-
nate, 12-13 mm. long, foliaceous, appressed-tomentulose; corolla greenish yellow,
fulvous-villous, the tube 22-30 mm. long, the epistaminal appendages replaced by
linear callose ridges, the lobes obliquely obovate, 13-15 mm. long; anthers gla-
brous, partly exserted; nectaries concrescent, annular, broadly 5-lobate to essen-
tially entire, equalling or slightly surpassing the ovary; follicles thick and rigid,
divergent, 6-12 cm. long, densely fulvous-hispid.
Called "cacho de chivo" in El Salvador.
Prestonia portobellensis (Beurl.) Woodson, Ann. Mo. Bot.
Gard. 18: 553. 1931. Echites portobellensis Beurl. Svensk. Vet. Akad.
Handl. 1854: 137. 1856. P. macrocarpa Hemsl. Biol. Cent. Am. Bot.
2: 311. 1881.
Moist forest, 500-700 m.; Escuintla. Southern Mexico; Hondu-
ras and El Salvador to Panama.
A suffrutescent vine, glabrous throughout or nearly so; leaves nrm-membrana-
cous to subcoriaceous, on petioles 6-30 mm. long, oblong-elliptic, 10-30 cm. long,
3-18 cm. broad, obtuse or usually abruptly short-acuminate, obtuse or rounded
at the base; inflorescence subcorymbose, 2-3 times divided, rarely simple, 8-30-
flowered, the peduncles much shorter than the leaves, the pedicels 6-18 mm. long;
bracts ovate-lanceolate, 4 mm. long or less, scarious or slightly foliaceous; calyx
lobes oblong-elliptic, acuminate, 11-17 mm. long, coriaceous or subcoriaceous, more
or less tinged with purple, glabrous or obscurely papillate; corolla greenish yellow,
380 FIELDIANA: BOTANY, VOLUME 24
sometimes tinged with purple outside, glabrous or obscurely papillate, the tube
11-17 mm. long, the epistaminal appendages slightly exserted or at least reaching
the orifice, the lobes obliquely obovate, 10-15 mm. long; anthers puberulent-
papillate dorsally, the tips exserted; nectaries concrescent, membranaceous, con-
spicuously surpassing the ovary; follicles rather slender, 33-35 cm. long, glabrous.
Prestonia schippii Woodson, Ann. Mo. Bot. Card. 23: 337.
1936.
Vine in mixed forest; Alta Verapaz (Contreras 4527). British
Honduras (type from Eldorado, Schipp S388).
A somewhat woody vine 6-7 m. long, the stems as much as 2.5 cm. in diameter,
glabrous throughout or nearly so; leaves firm-membranaceous, on petioles 12-
16 mm. long, elliptic-oblong, 10-17 cm. long, 4-7 cm. broad, abruptly short-acumi-
nate, obtuse or rounded at the base, conspicuously reticulate-veined; inflorescence
subcorymbose, simple, 10-12-flowered, the peduncles much shorter than the leaves,
the pedicels 12-15 mm. long; bracts ovate-lanceolate, 1-2 mm. long, scarious; calyx
lobes narrowly ovate-elliptic, acute, 10-12 mm. long, subcoriaceous, glabrous,
slightly tinged with purple; corolla cream-colored, minutely papillate outside, the
tube 13-14 mm. long, the epistaminal appendages wholly included, the lobes
obliquely obovate, 9-10 mm. long; anthers minutely hirtellous dorsally, partly
exserted; nectaries concrescent, thin and somewhat diaphanous, conspicuously
surpassing the ovary.
Closely related to the preceding species, and perhaps not distinct.
Prestonia speciosa Donn.-Sm. Bot. Gaz. 27: 435. 1899.
Known in Guatemala only from the type, Buena Vista, Santa
Rosa, 1,700 m., Heyde & Lux 4497. Mexico (Chiapas) ; Honduras.
A woody vine, densely fulvous-pubescent throughout; leaves firm-membrana-
ceous, on petioles 5-12 mm. long, ovate to broadly ovate-elliptic, 10-17 cm. long,
7-11 cm. broad, abruptly short-acuminate, broadly obtuse or rounded at the base,
sparsely and minutely hispidulous-strigillose above, tomentulose beneath along
the nerves; inflorescence subumbellate, 4-8-flowered, the peduncles scarcely longer
than the petioles, the pedicels 1-1.5 cm. long; bracts narrowly lanceolate, 2-5 mm.
long, subfoliaceous; calyx lobes ovate to ovate-oblong, acute, 12-15 mm. long,
foliaceous, minutely hirtellous, corolla yellow, appressed-villosulous, the tube 15-
17 mm. long, the throat conic-campanulate, 15 mm. long, 8-9 mm. broad, the
epistaminal appendages replaced by inconspicuous obtriangular-foveolate protu-
berances, the lobes obliquely obovate, 20-25 mm. long, spreading; anthers gla-
brous, included; nectaries concrescent, rather obscurely and irregularly lobate,
somewhat longer than the ovary.
RAUVOLFIA [Plumier] Linnaeus
Reference: Aragula Sathyanarayana Rao, Ann. Mo. Bot. Gard.
43: 253-354, ittus. 1956.
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 381
Shrubs or small trees; leaves verticillate, the petioles glandular on the upper
surface, at least at the very base, interpetiolar stipules deciduous; inflorescence
cymose, terminal or lateral, the flowers small, white or greenish; calyx 5-parted,
the lobes equal, without squamellae; corolla salverform or tubular-sal verform, the
limb equally 5-lobate, sinistrorsely contorted; anthers not connivent, the con-
nective not enlarged; ovary of 2 distinct carpels, each carpal 1-2-ovulate, sur-
rounded by a low annular nectary; fruit syncarpous, drupaceous; seeds 1-2, naked.
About 100 species, in the tropics of both hemispheres, 35 in
America. Two other species are known from Panama.
Inflorescence about as long as the largest associated leaf; leaves 3 at each node,
rarely 4 R, ligustrina.
Inflorescence 2-4 times shorter than the largest associated leaf; leaves 4 at each
node, rarely 3 or 5 R. tetraphylla.
Rauvolfia ligustrina Roem. & Schult. Syst. Veg. 4: 805. 1819.
Dry scrub or savannas along the Pacific coastal area, mostly near
sea level to 1,000 m.; Escuintla; Suchitepe'quez. West Indies; Mex-
ico through Central America to northern South America.
Shrubs 1-3 m. tall, the branches glabrous, with glands ascending the petioles;
leaves ternate, slightly unequal, short petiolate, ovate to ovate-elliptic, acute to
acuminate, glabrous or pubescent along the midrib below, 1-5 cm. long and 0.5-
3 cm. broad, the largest ones at the nodes; inflorescences terminal or lateral, few
to many-flowered; peduncles slender, 1-3 cm. long and 2-3-branched, secondary
peduncles congested, glabrous or puberulent; flowers small; calyx 5-lobed, the lobes
lanceolate, acuminate, 1.5-2 mm. long, glabrous; corolla urceolate, 2-3.5 mm. long,
pilose near the throat within, otherwise glabrous, the lobes ovate to subrotate,
1-1.5 mm. long; stamens included, inserted near the throat; ovary bicarpellate,
subglobose, 1.5-2 mm. in diameter; fruits round, 5-7 mm. in diameter, 2-seeded,
glabrous.
Rauvolfia tetraphylla L. Sp. PI. 208. 1753. R. hirsuta Jacq.
Enum. PI. Carib. 14. 1760. R. tomentosa Jacq. I.e. R. canescens L.
I.e. ed. 2, 303. 1762. R. heterophylla Roem. & Schult. Syst. Veg. 4:
805. 1819, possibly. R. canescens var. glabra Muell.-Arg. Linnaea
30: 395. 1860. R. hirsuta var. glabra (Muell.-Arg.) Woodson, Ann.
Mo. Bot. Card. 26: 299. 1939. Chalchupa; curarina (Suchitepe'quez).
Wet to dry thickets, often in second growth or in waste ground
or old fields, 2,000 m. or less, most common at 500 m. or lower;
Peten; Izabal; Baja Verapaz; El Progreso; Zacapa; Jutiapa; Santa
Rosa; Escuintla; Suchitepe'quez; Chimaltenango (probably planted);
Retalhuleu; San Marcos; Huehuetenango. West Indies; Mexico;
British Honduras to El Salvador and Panama; northern South Amer-
ica. Introduced into India.
382
FIELDIANA: BOTANY, VOLUME 24
FIG. 107. Rauvolfia tetraphylla. A, habit, X 1A; B, fruit, X 2; C, pistil with
disk at base, X 10; D, flower, X 5; E, corolla opened out with stamens in place,
X 5.
Usually a shrub a meter high or less but sometimes becoming a small tree of
4 m., with copious white latex, usually much branched, finely pubescent through-
out or often almost wholly glabrous; leaves mostly in whorls of 4, sometimes of 3
or 5, very unequal, firm-membranaceous or when fresh somewhat fleshy, narrowly
oblong-elliptic to broadly ovate-elliptic or obovate-elliptic, 2-13 cm. long, 1-5 cm.
broad, acute or obtuse, broadly acute or obtuse at the base, the petioles 1-7 mm.
long, glandular; inflorescences condensed, much shorter than the subtending leaves,
few-many-flowered, the pedicels 2-3 mm. long; calyx lobes ovate or ovate-lanceo-
late, acute or subobtuse, 1.5-3 mm. long, glabrous or puberulent; corolla minutely
puberulent-papillate or glabrous, the tube 2.5-4 mm. long, somewhat constricted
at the insertion of the stamens, the lobes obliquely obovate-rounded, 1-1.5 mm.
long; stamens inserted about the middle of the corolla tube; drupes subglobose,
5-8 mm. in diameter, becoming red and at maturity almost black.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 383
Known in El Salvador as "amatillo," "hierba de San Jose"," "ma-
tacoyote," and "senorita"; "cabamuc," "chacmuc," "chacmuc-ac,"
"cabalmuc" (Yucatan, Maya).
This is a well-known plant in Guatemala, where it is employed
commonly by the country people in treating malaria, and it is also
one of the reputed remedies for snake bites. See "Estudio de la
planta llamada "chalchuapa," Revista Agric. (Guatemala) 14: 205-
215, illus. 1936. It has been found by Guatemalan investigators to
contain two alkaloids, to which the names Chalcupine A and B have
been given. The fruit is generally considered to be poisonous. The
shrub is abundant on the Pacific plains, and seldom or rarely ascends
into the foothills. The typical and original form of the species is
densely and finely pubescent throughout. It is infrequent in Guate-
mala and other parts of Central America. The common form is the
glabrous one in which the leaves are sparsely pilose along the nerves,
or almost glabrous. There are intergrading forms.
RHABDADENIA Mueller von Argau
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 23: 205-
211. 1936.
Slender woody vines, glabrous; leaves opposite, not glandular; inflorescence a
greatly reduced, dichasial cyme, alternate-axillary or subterminal, sometimes 1-
flowered; calyx 5-parted almost to the receptacle, the lobes subequal, somewhat
foliaceous, scarcely imbricate, without squamellae within; corolla funnelform, the
tube not appendaged within, straight, the limb regularly 5-parted, dextrorsely con-
torted; anthers connivent and agglutinated to the stigma, the connective enlarged,
narrowly bilobate; ovary of distinct carpels, many-ovulate, surrounded by 5 dis-
tinct or somewhat concrescent nectaries; stigma fusiform; follicles 2, distinct,
slender, terete; seeds numerous, rostrate, comose at the apex.
Three species, the others South American.
Rhabdadenia biflora (Jacq.) Muell.-Arg. in Mart. Fl. Bras. 61:
175. 1860. Echites biflora Jacq. Enum. PI. Carib. 13. 1760. E. palu-
dosa Vahl, Eclog. 2: 19. 1798. R. paludosa Miers, Apocyn. S. Amer.
119. 1878. R. macrantha Donn.-Sm. Bot. Gaz. 40: 7. 1905 (type from
Tela, Honduras).
In or near mangrove swamps; Izabal. Southern Florida; Yuca-
tan Peninsula of Mexico; British Honduras; Honduras; Nicaragua;
Panama; in Central America apparently confined to the Atlantic
coast; Greater Antilles; northern South America.
A small or large, woody vine, glabrous; leaves coriaceous or firm-membrana-
ceous, on petioles 1-2 cm. long, broadly obovate-oblong to oblong or lanceolate,
FIG. 108. Rhabdadenia biflora. A, habit, X 1A; B, stamens in natural position
on portion of dissected corolla tube, X 3; C, calyx and pistil, natural size; D, stig-
ma, X 5.
384
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 385
5-12 cm. long, 1.5-5 cm. broad, very obtuse or rounded and mucronate at the apex,
obtuse or subacute at the base, paler beneath, the nerves very slender and incon-
spicuous; inflorescences lateral or rarely subterminal, 1-5-flowered, the peduncles
equalling or longer than the leaves, the pedicels 10-13 mm. long; bracts minute,
scarious; calyx lobes broadly ovate-oblong, mucronulate, 1-9 mm. long, subfolia-
ceous; corolla white, often tinged with pink, the tube 1.5-2 cm. long, the throat
conic, 2-3 cm. long, 1-1.5 cm. broad at the orifice, the lobes broadly oblique-
obovate, 2-2.5 cm. long, spreading; anthers densely pilosulous dorsally, included;
nectaries half as long as the ovary; follicles about 8 cm. long or perhaps even longer,
straight, terete, striate, glabrous, continuous.
The flowers are showy and handsome. The plant is a characteris-
tic member of the mangrove association.
STEMMADENIA Bentham
Reference: Robert E. Woodson, A revision of the genus Stemma-
denia, Ann. Mo. Bot. Gard. 15: 341-378, ittus. 1928.
Shrubs or trees, glabrous or pubescent; leaves opposite, eglandular, membrana-
ceous; inflorescence cymose, few-flowered, the flowers large and showy, yellow or
white; calyx 5-parted, the lobes very unequal, closely imbricate, usually conspicu-
ously folia ceous, bearing numerous squamellae within; corolla salverform or fun-
nelform, the limb 5-parted, sinistrorsely convolute; anthers not connivent, the
connective not enlarged; ovary of 2 distinct carpels, many-ovulate, surrounded by
a fleshy annular nectary; fruit follicular, coriaceous or fleshy, the follicles usually
very large and thick; seeds numerous, naked, embedded among fleshy arils.
Fifteen species, in tropical America, mostly in Mexico and Cen-
tral America. Three others are known from Central America.
Corolla salverform; bracts inserted about the middle of the pedicel, not immedi-
ately subtending the calyx.
Calyx equaling the corolla tube or only slightly shorter S. donnell-smithii .
Calyx half as long as the corolla tube or shorter.
Bracts scarious or only slightly foliaceous; calyx lobes strongly imbricate.
S. decipiens.
Bracts foliaceous; calyx lobes spreading S. eubracteata.
Corolla funnelform; bracts inserted at the summit of the pedicel directly below the
calyx.
Corolla throat broadly conic, at least half as broad as long; leaves usually pubes-
cent beneath, often very densely so S. obovata.
Corolla throat tubular or narrowly conic, much longer than broad; leaves gla-
brous beneath or nearly so.
Calyx lobes 10-15 mm. long, strongly imbricate S. galeottiana.
Calyx lobes 3-6 mm. long, scarcely imbricate S. macrophylla.
Stemmadenia decipiens Woodson, Ann. Mo. Bot. Gard. 15:
363. 1928. Coyotes de coche.
Mixed forest, 100-1,100 m.; Retalhuleu (between Retalhuleu and
Nueva Linda); Huehuetenango (near Democracia). Western Mex-
ico; Nicaragua.
386 FIELDIANA: BOTANY, VOLUME 24
A shrub or small tree, 10 m. high or less, glabrous throughout or nearly so;
leaves on petioles 5-7 mm. long, elliptic to obovate-oblong, 7-10 cm. long, 2.5-
6 cm. broad, acute or acuminate, obtuse or acute at the base; inflorescence 3-9-
flowered, the pedicels 3-5 mm. long; calyx lobes imbricate, scarious or subcoriaceous,
ovate, acute or obtuse, 4-6 mm. long; corolla yellow, salverform, the tube 2-3 cm.
long, 2.5 mm. broad, the lobes obliquely obovate, somewhat reflexed.
Stemmadenia donnell-smithii (Rose) Woodson, Ann. Mo.
Bot. Gard. 15: 369. 1928. Tabernaemontana donnell-smithii Rose ex
Donn.-Sm. Bot. Gaz. 18: 206. 1893. Co/on; copal; cojon de caballo;
cojon de puerco; tonche; gutigamba (fide Aguilar).
Dry to wet thickets or open forest, often in second growth or in
roadside hedges, frequent in pastures, 1,500 m. or less, most common
at 700 m. or lower; Pete"n; Alta Verapaz; Izabal; Santa Rosa; Es-
cuintla; Guatemala; Solola; Suchitepe'quez ; Retalhuleu; Quezalte-
nango; San Marcos. Southern Mexico; British Honduras to El
Salvador and Panama.
A shrub or small tree, sometimes 15 m. high with a trunk 30 cm. or more in
diameter, usually not more than half as high, the bark almost smooth, pale; leaves
subsessile, elliptic or obovate-elliptic, 6-14 cm. long, 3-5.5 cm. broad, caudate-
acuminate, obtuse or acute at the base, glabrous above or nearly so, barbellate
beneath in the axils of the nerves; inflorescence 1-5-flowered, the pedicels 10-
13 mm. long; calyx lobes ovate to ovate-oblong, obtuse or subobtuse, 2-2.5 cm.
long, scarious; corolla salverform, buff or pale yellow, the tube 2.5-3 cm. long,
3.5 mm. broad at the base, the lobes obliquely obovate, 15-18 mm. long, somewhat
reflexed; follicles very large, hard, and heavy, 5-7 cm. long, 3-5 cm. broad, or in
the fresh state even larger, very thick, ferruginous, somewhat verrucose.
In Honduras sometimes called "cojon de mico" and "cojon de
burro"; "cojoton" (British Honduras). The name "cojon," strangely
enough, has been given to a caserio in the Department of Jutiapa.
The wood is light brown, rather light in weight but firm and strong,
fine-textured, easy to work, not durable; not utilized. The large and
very conspicuous fruits are so heavy and produced in such abundance
that they bend the branches sharply downward. When they or the
branches are cut, they exude a large amount of very sticky, white
latex that contains a substance having the properties of guttapercha
(see Standl. & Calderon, Lista PI. Salvador 175. 1925). The tree is
abundant along much of the Pacific coast of Central America, and
it has been suggested that it might be exploited as a source of this
substance, which could be obtained in substantial and probably large
amounts. The latex is employed in El Salvador for fastening the
wrappers of cigarettes, also by boys in making kites, and for other
purposes when an adhesive is needed. In Guatemala it sometimes
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 387
is chewed like chicle. The latex is said to be employed there also in
treating the bites of the spider called arana de caballo, whose poison
causes the hoofs of horses and mules to fall off.
Stemmadenia eubracteata Woodson, Ann. Mo. Bot. Gard. 15:
368, t. 49, f. 2. 1928.
Known only from the type, Volcan de Tecuamburro, Santa Rosa,
2,000 m., Heyde & Lux 4538.
A shrub or small tree, glabrous throughout or nearly so; leaves on petioles
4-5 mm. long, oblong-elliptic or oblanceolate-oblong, 6-8 cm. long, 2-3 cm. broad,
caudate-acuminate to subobtuse, acute at the base; inflorescence 2-5-flowered, the
pedicels 7-9 mm. long; bracts conspicuously foliaceous; calyx lobes foliaceous, obo-
vate-elliptic, abruptly acuminate, 8-10 mm. long, spreading; corolla yellow, salver-
form, the tube 22-25 mm. long, 3 mm. broad at the base, the lobes obliquely
obovate, 14-16 mm. long, somewhat reflexed.
This has been reported from Guatemala as S. bella Miers.
Stemmadenia galeottiana (A. Rich.) Miers, Apocyn. S. Amer.
76. 1878. Odontostigma galeottiana A. Rich, in Sagra, Hist. Cub. 11:
87. 1850. E chiles bignoniaeflora Schlecht. Linnaea 26: 372. 1854.
S. bignoniaeflora Miers, Apocyn. S. Amer. 76. 1878. S. insignis
Miers, I.e. S. bella Miers, I.e. 77. 1878. Cajon; Ixlao', ixdislan
(Pete"n, Maya, fide Lundell) ; cojon; cojon de caballo.
Moist or wet forest, 700 m. or less; Pete"n; Alta Verapaz. South-
ern Mexico.
A shrub or small tree, 1-5 m. high, glabrous throughout or nearly so; leaves on
petioles 8-11 mm. long, oval to obovate-elliptic, 9-14 cm. long, 4-6 cm. broad,
caudate-acuminate, obtuse at the base; inflorescence 1-4-flowered, the pedicels
8-13 mm. long; calyx lobes imbricate, ovate to obovate-oblong, obtuse or rounded
at the apex, 1-1.5 cm. long; corolla pure white, funnelform, the tube 8-10 mm.
long, the throat subtubular, 4-5.5 cm. long, 7-10 mm. broad at the orifice, the
lobes broadly obovate, 2-2.5 cm. long, spreading; follicles ovoid-subreniform, 2-
2.5 cm. long, 1.5 cm. broad, irregularly verrucose.
Called "xlaul" (Maya) and "laurel" in Yucatan. A small tree of
this species once grew in the park at Coban, where it attracted atten-
tion because of its abundance of handsome, large, pure white flowers.
Stemmadenia macrophylla Greenm. Proc. Am. Acad. Sci. 35:
310. 1900.
Moist or wet thickets or forest, 1,300-1,400 m.; Alta Verapaz
(type from Pansamala, Tuerckheim 981); Quezaltenango; so far as
known, endemic, but to be expected in Chiapas.
388
FIELDIANA: BOTANY, VOLUME 24
FIG. 109. Stemmadenia obovata. A, habit, X 1A\ B, calyx and pistil partially
dissected to show ovary and disk, natural size; C, one follicle of fruit, X 1A;
D, stamens in position on dissected corolla tube, X 2.
A shrub or tree, sometimes 10 m. high, glabrous throughout or nearly so;
leaves on petioles 1.5-2 cm. long, oblong-elliptic or obovate-elliptic, 15-20 cm.
long, 5.8 cm. broad, subobtuse or caudate-acuminate, obtusely cuneate at the base;
inflorescence 1-8-flowered, the pedicels 2-4 mm. long; calyx lobes scarcely imbri-
cate, ovate or oblong-obovate, acute or obtuse, 3-6 mm. long; corolla funnelform,
yellowish, the tube 1.5-2 cm. long, the throat subtubular, 2.5-3 cm. long, 7-9 mm.
broad at the orifice, the lobes obliquely ovate-oblong, 2.2-2.8 cm. long, spreading.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 389
This has been reported from Guatemala as S. bignoniaeflora Miers.
Stemmadenia obovata (Hook. & Arn.) Schum. in Engler &
Prantl, Pflanzenfam. 42: 149. 1895. Bignonia obovata Hook. & Arn.
Bot. Beechey Voy. 439. 1841. S. pubescens Benth. Bot. Voy. Sulph.
125. 1845. S. mollis Benth. I.e. S. obovata var. mollis Woodson,
Ann. Mo. Bot. Card. 15: 358. 1928. Chapopo (Zacapa); cojon; cojon
de puerco; cojon de coche.
Moist or dry thickets, often on rocky brushy slopes, sometimes
on plains or in thin forest, 1,350 m. or less; Zacapa; Chiquimula; El
Progreso; Jutiapa; Santa Rosa; Guatemala; Retalhuleu; Quiche";
Huehuetenango. Southern Mexico; El Salvador and Honduras to
Panama; southward to Bolivia.
A shrub or small tree, sometimes 10 m. high but usually 5 m. or less; leaves on
petioles 5-8 mm. long, obovate or obovate-elliptic, 10-20 cm. long, 7-10 cm. broad,
obtuse to acuminate, obtusely cuneate at the base, varying from almost glabrous
to densely and softly pubescent beneath, the upper surface glabrous or pubescent;
inflorescence pubescent, 1-6-flowered, the pedicels 9-14 mm. long; calyx lobes ovate
or oblong-ovate, obtuse, 1.5-2 cm. long; corolla bright yellow, funnelform, the tube
1.5-2.5 cm. long, the throat broadly conic, 1.5-3 cm. long, 1.5-2.5 cm. broad at the
orifice, the lobes broadly obovate, 1.5-2.5 cm. long, spreading; follicles obovoid-
subreniform, 4-4.5 cm. long and 3-3.5 cm. broad, or probably larger when fresh,
minutely verrucose.
Sometimes called "flor del dia" in El Salvador. When in flower
the shrub is a showy and handsome one, because of the large size
and brilliant coloring of the corollas, but the flowers are found for
only a short season, toward the end of the rainy season. Pubescence
on the leaves of this species is exceedingly variable, from sparse or
none to dense on both sides of the leaf, the lower surface usually
being more pubescent than the upper surface.
STEPHANOTIS FLORIBUNDA Brongn., native of Madagascar, is
planted rather frequently for ornament in Guatemalan gardens.
It is a slender glabrous vine with elliptic leaves and small, waxy,
white or cream-colored, very fragrant flowers.
TABERNAEMONTANA L.
Shrubs or trees, usually glabrous or nearly so; leaves opposite; inflorescence
compound-cymose, usually many-flowered; calyx 5-parted almost to the recep-
tacle, the lobes equal, small, scarious or only slightly foliaceous, bearing numerous
squamellae within; corolla salverform, generally small, usually white; anthers not
390 FIELDIANA: BOTANY, VOLUME 24
connivent, free from the stigma, usually tinged with bluish green, the connective
not enlarged; ovary of 2 distinct carpels, many-ovulate, with or without a basal
annular nectary; fruit follicular, the follicles often large and thick; seeds numerous,
embedded among the fleshy arils.
About 50 species, in tropical America. One other Central Amer-
ican species is found in Costa Rica.
Flowers large, commonly 6-7 cm. long, sometimes double; cultivated plants.
T. coronaria.
Flowers much smaller, usually less than 2 cm. long; native plants.
Anthers yellow throughout, not tinged with bluish green, inserted near or below
the middle of the corolla tube, wholly included T. arborea.
Anthers margined or tipped with bluish green, inserted near or above the middle
of the corolla tube, often somewhat exserted.
Anthers about half exserted T. amygdalifolia.
Anthers included or the tips barely exserted.
Corolla tube 15-16 mm. long T. laurifolia.
Corolla tube 6-9 mm. long.
Inflorescence dense and congested, much shorter than the subtending
leaves T. alba.
Inflorescence lax and open, almost equaling the subtending leaves.
T. chrysocarpa.
Tabernaemontana alba Mill. Gard. Diet. ed. 8. no. 2. 1768.
Reported from Pete"n. Southern Mexico.
A large shrub or a small or medium-sized tree, glabrous throughout, the trunk
as much as 20 cm. in diameter, the crown rather dense, spreading, the bark green-
ish or light brown; leaves membranaceous or subcoriaceous, on petioles 5-20 mm.
long, obovate-elliptic or oblanceolate-elliptic, 5-16 cm. long, 2-7 cm. broad, obtuse
or abruptly and shortly obtuse-acuminate, cuneate at the base; inflorescence
broadly corymbose, sessile or nearly so, much shorter than the subtending leaves,
many-flowered, the pedicels 2-10 mm. long; calyx lobes ovate, subobtuse, closely
imbricate, 1.5-1.8 mm. long; corolla white, the tube 6-8 mm. long, slightly con-
stricted at the insertion of the stamens, the lobes 8-10 mm. long, oblong-dolabri-
form, with an inconspicuous lateral acumen, obtuse, spreading; stamens inserted
in the upper third of the corolla tube, the anthers marginate with bluish green, the
tips barely exserted; ovary surrounded by a low annular nectary; follicles broadly
subreniform, broadly acuminate, 2-2.5 cm. long, 1-1.5 cm. broad, smooth or
nearly so.
Called "lecherillo" in Veracruz. The bark exudes a tasteless latex
when cut. The wood is whitish throughout.
Tabernaemontana amygdalifolia Jacq. Enum. PI. Carib. 14.
1760. T. deamii Donn.-Sm. Bot. Gaz. 52: 50. 1911 (type collected
along the Rio Motagua near Gualan, Zacapa, Deam 6282). Cojon de
puerco; cojon de mico; cocoguaca (Alta Verapaz) ; polo de mico (Izabal).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
391
FIG. 110. Tabernaemontana amygdalifolia. A, habit, X J^; B, corolla dissected
to show anthers in place, X 1J^; C, calyx partially dissected to show ovary and
glands, X 2; D, fruit, one follicle only, X Yi.
Moist or dry thickets, often in sandy places, frequently in second
growth, 700 m. or less; Alta Verapaz; Izabal; Zacapa; El Progreso;
Chiquimula; Santa Rosa; San Marcos. Southern Mexico; British
Honduras to El Salvador and Panama; Colombia and Venezuela.
Usually a shrub of 1-5 m., glabrous throughout; leaves membranaceous, on
petioles 5-13 mm. long, oblong-elliptic or obovate-elliptic, 5-15 cm. long, 2-5 cm.
broad, obtuse to abruptly acuminate, cuneate at the base; inflorescence corymbose,
much branched, slightly shorter than the subtending leaves, few-many-flowered,
the pedicels 1-2 cm. long; calyx lobes ovate, acute or subobtuse, somewhat imbri-
cate; corolla white, the tube 14-16 mm. long, somewhat narrowed toward the
orifice, the lobes narrowly obovate, 16-22 mm. long, somewhat reflexed, incon-
spicuously acuminate laterally; stamens inserted near the orifice of the corolla
tube, the anthers with bluish green margins, half exserted; ovary surrounded at
the base by an annular nectary; follicles narrowly ovoid-ellipsoid, 3-6 cm. long,
1.2-1.8 cm. thick, narrowly acuminate, longitudinally striate.
Known in Yucatan as "utsubpec" (Maya), "jazmin de perro,"
"olfato de perro," "chilindron," "cojon macho," "cojon de puerco,"
"leche de perra," "jazmin del monte," and "amatillo." It is reported
that in Yucatan an infusion of the leaves sometimes is administered
as a laxative. The latex is said to be used for adulterating chicle.
392 FIELDIANA: BOTANY, VOLUME 24
Tabernaemontana arborea Rose ex Donn.-Sm. Bot. Gaz. 18:
206. 1893. T. schippii Standl. Field Mus. Bot. 8: 34. 1930 (type
from Big Creek, British Honduras, Schipp 168). Peschiera arborea
Markgraf, Notizbl. Bot. Gart. Berlin 14: 173. 1938. Cojon.
Moist or wet thickets or mixed forest, often in second growth,
600 m. or less; Pete"n; Alta Verapaz; Izabal; Quezaltenango (type
from Rio Ocosito, Smith 2766); San Marcos. Southern Mexico;
British Honduras; Honduras; Panama.
A tree, commonly 6-15 m. high, glabrous throughout or nearly so, the trunk
often 30-50 cm. in diameter, often with small buttresses, the bark light brown,
somewhat scaly, the crown spreading or rounded ; leaves membranaceous, on peti-
oles 5-25 mm. long, ovate-elliptic to narrowly oblong-elliptic, 7-18 cm. long, 2-
7 cm. broad, shortly obtuse-acuminate, obtuse or acute at the base; inflorescence
broadly corymbose, much branched, usually shorter than the subtending leaves,
the flowers numerous, white, fragrant, the pedicels 5-8 mm. long; calyx lobes
ovate-lanceolate, acute or acuminate, 2-3 mm. long, scarcely imbricate at the base;
corolla tube 7-9 mm. long, conspicuously narrowed toward the orifice, the lobes
obliquely ovate-oblong, obtuse, 12-15 mm. long, spreading; stamens inserted near
the base of the corolla tube, the anthers uniformly yellow, 3-3.5 mm. long, deeply
included; ovary without a nectary; follicles broadly reniform, obtuse, 6-8 cm. long,
3-4.5 cm. broad, conspicuously verrucose.
Called "cojoton" and "cojon de caballo" in British Honduras.
A fair amount of latex exudes when the trunk is cut. The wood is
yellowish or light reddish brown, hard and heavy, fine- textured.
Tabernaemontana chrysocarpa Blake, Contr. Gray Herb. 52:
81. 1917. T. amblyblasta Blake, Contr. U. S. Nat. Herb. 24: 18, t. 6.
1922 (type from Los Amates, Izabal, Blake 7636). Chapupo, cojon,
cojon de gato, cojon de mico, cojon de caballo, lechoso (Peten), polo de
mico, huevos de mico (Izabal).
Wet or moist thickets or mixed forest, sometimes in open pine
forest, 600 m. or less, most plentiful at or near sea level; Peten; Alta
Verapaz; Izabal. Southern Mexico; British Honduras; Honduras to
Panama.
A large shrub or a tree, sometimes 15 m. high with a trunk as much as 45 cm.
in diameter, usually smaller, the crown dense, spreading, the trunk round or some-
what fluted, the bark dark grayish to light brown, with small scales, glabrous
throughout; leaves firm-membranaceous to subcoriaceous, on petioles 6-25 mm.
long, obovate to oblanceolate-elliptic, 5-25 cm. long, 2-9 cm. broad, obtuse or
shortly and abruptly acuminate, cuneate-acute at the base; inflorescence subcorym-
bose, usually lax and equalling or surpassing the leaves, many-flowered, the pedi-
cels 7-15 mm. long; calyx lobes broadly ovate, obtuse, about 2 mm. long, closely
imbricate; corolla white or yellowish white, the tube 6-9 mm. long, slightly con-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 393
stricted at the insertion of the stamens, the lobes oblong-dolabriform, obtuse, 10-
12 mm. long, spreading; stamens inserted in the upper third of the corolla tube, the
anthers with bluish green margins, barely included or a small part of the tips ex-
serted; ovary surrounded at the base by a low annular nectary; follicles subreni-
form, 2-2.5 cm. long, 1.5-2 cm. broad, obtuse or with a very inconspicuous acumen,
smooth or nearly so.
Called "cojoton" and "cojon de perro" in British Honduras,
"chanchito" (Honduras), "totoyoyote" (Veracruz, the fruit), "le-
cherillo" (Oaxaca, Veracruz). The fruit yields a large amount of
white latex that coagulates readily when rubbed between the hands.
It is reported to be used sometime for adulterating chicle, and is
chewed locally like that substance. The yellowish white wood is
sometimes utilized in construction of the poorer types of lowland
dwellings. This species has been reported from Guatemala and other
parts of Central America as T. citrifolia L., a West Indian species.
Tabernaemontana coronaria (Jacq.) Willd. Enum. Hort. Berol.
275. 1809. Nerium coronarium Jacq. Coll. Bot. 1: 138, t. 52. 1786.
T. divaricata of authors, not Nerium divaricatum L. Flor de poeta.
An ornamental plant, widely distributed in cultivation in trop-
ical regions, perhaps native in the East Indies; cultivated rather
frequently in Guatemala, chiefly in the tierra caliente.
A shrub, commonly 1-2 m. high, glabrous; leaves short-petiolate, membrana-
ceous, narrowly elliptic or elliptic-oblong, 8-12 cm. long, caudate-acuminate with
usually a long and very narrow acumination, acute at the base, paler beneath; in-
florescences 1-8-flowered, mostly in the forks of the branches; flowers fragrant,
white, often double, mostly 6-7 cm. long, the corolla limb 3-5 cm. broad, the tube
about 2 cm. long.
Known in El Salvador by the names "jazmin de Arabia" and
"jazmin de Jamaica." The most common English name is "crape
jasmine." A not very showy or attractive shrub, and scarcely worth
cultivation when so many better plants are available for tropical re-
gions. It seems not to be very highly esteemed in Central America.
Tabernaemontana laurifolia L. Sp. PI. 210. 1753. Cojon.
Brushy mountain slopes, or wet mixed-forest, 1,200-1,600 m.;
Chimaltenango (Volcan de Fuego); San Marcos (above Finca El
Porvenir, Volcan de Tajumulco). Jamaica.
A shrub 1.5-2.5 m. high, glabrous throughout; leaves membranaceous, oval to
oblong-elliptic, 5-17 cm. long, 3.5-10 cm. broad, abruptly acuminate, obtuse or
acute at the base, the petioles 1-2.5 cm. long; inflorescence congested, much shorter
than the subtending leaves, subumbellate, the pedicels 2-8 mm. long; calyx lobes
394 FIELDIANA: BOTANY, VOLUME 24
broadly ovate, obtuse, 3 mm. long, imbricate; corolla greenish white, the tube
15 mm. long, slightly constricted at the insertion of the stamens, the lobes oblong-
dolabriform, subobtuse, 10 mm. long, spreading; stamens inserted in the upper
fourth of the corolla tube, the anthers with bluish green margins, barely included ;
ovary surrounded at the base by a low annular nectary; follicles narrowly gibbous-
ellipsoid, 3-4.5 cm. long, 1-1.2 cm. thick, narrowly acuminate, smooth or nearly so.
The single collection was determined by Woodson. The range of
the species, as here treated, is a most unusual one, and on general
principles it is rather to be expected that more ample material will
prove the Central American plant to be a species distinct from the
Jamaican one.
THEVETIA Adanson
Shrubs or small trees; leaves alternate, not glandular; inflorescence cymose,
few-several-flowered, the flowers large and showy, usually yellow; calyx 5-parted
almost to the receptacle, the lobes equal or nearly so, somewhat foliaceous, bearing
numerous squamellae within; corolla salverform, the tube much shorter than the
lobes, closed at the orifice by 5 small villous scales inserted above the stamens,
the lobes broadly obovate-dolabriform, spreading; anthers not connivent, the con-
nective not enlarged; ovary of 2 distinct carpels, 2-4-ovulate, surrounded by a
carnose annular nectary; fruit syncarpous, drupaceous, containing 2 or sometimes
4 large naked seeds.
Species 6-7, in tropical America. The following are found in
Central America.
Lateral nerves of the leaves obscure or obsolete beneath in dry state.
Leaves abruptly acuminate, 4-6 cm. broad T. ahonai.
Leaves acute or short acuminate, 0.6-3 cm. broad.
Leaves acute or short-acuminate, 1.5-3 cm. broad; fruits 2-3 cm. broad.
T. gaumeri.
Leaves acutish or obtuse, long attenuate, 13 mm. broad or less; fruits 4-4.5 cm.
broad T. peruviana.
Lateral nerves of the leaves elevated and very conspicuous beneath.
Leaves glabrous beneath T. plumeriaefolia.
Leaves densely short pubescent beneath T. ovata.
Thevetia ahouai (L.) A. DC. in DC. Prodr. 8: 344. 1844;
L. Wms. Fieldiana, Bot. 31: 403. 1968. Cerbera ahouai L. Sp. PL
ed. 2. 303. 1762. C. nitida HBK. Nov. Gen. & Sp. 3: 225. 1819.
T. nitida A. DC. I.e. 345. Plumeriopsis ahouai Rusby & Woodson,
Ann. Mo. Bot. Gard. 24: 11. 1937. Chilindron; cojon de perro.
Wet to dry thickets, sometimes in second growth or in open pine
forest, 600 m. or less; Pete"n; Alta Verapaz; Izabal; Retalhuleu; San
Marcos; Huehuetenango. Southern Mexico; British Honduras to
Panama; northern South America.
FIG. 111. Thevetia ahouai. A, habit, X ^; B, calyx and pistil, X 2; C, oblique
basal view of fruit, natural size; D, corolla partly dissected, X %', E, stamens in
position on corolla tube, one reversed, X 2.
395
396 FIELDIANA: BOTANY, VOLUME 24
Usually a shrub of 1-3 m., sometimes a small tree, glabrous throughout or
nearly so; leaves thick-membranaceous to subcoriaceous, short-petiolate, obovate
to oblanceolate, 8-20 cm. long, 3-7 cm. broad, acute to abruptly acuminate, atten-
uate to the base, deep green and very lustrous above, sometimes sparsely puberu-
lent beneath; inflorescences shorter than the leaves, the flowers few, greenish yellow
or lemon-yellow; calyx lobes ovate or ovate-lanceolate, acute or acuminate, 5-
7 mm. long, reflexed, glabrous; corolla glabrous outside, the tube 2.5-3.5 cm. long,
somewhat dilated at the insertion of the stamens, the lobes obovate, 1.5-2.5 cm.
long, reflexed; fruit baccate, pyriform or subglobose, 3.5-4.5 cm. broad, scarlet,
the seeds large, black.
Known in British Honduras by the names "cogotone," "cojoton"
and "cojon de mico," "yambigo" (Oaxaca). The fruits are showy
and rather handsome.
Thevetia gaumeri Hemsl. in Hook. Icon. t. 1517. 1886 (type
from Cozumel Island, Yucatan). T. spathulata Millsp. Field Mus.
Bot. 1: 383. 1898. T. steerei Woodson, Am. Journ. Bot. 22: 685. 1935
(type from Yucatan).
British Honduras, at or little above sea level. Mexico (Yucatan).
A shrub or small tree, reported to attain rarely a height of 10 m., glabrous
throughout, the branches thick, pale; leaves thick-membranaceous, on petioles
1-2 cm. long, linear-oblanceolate, 7-18 cm. long, 1.5-3.5 cm. broad, acute or short-
acuminate, long-attenuate to the base, green and lustrous above, paler beneath,
the lateral nerves obscure; inflorescences terminal and lateral, several-flowered;
calyx lobes ovate or ovate-lanceolate, 8-10 mm. long; corolla yellow, orange or
pinkish yellow, the tube 10-13 mm. long, the lobes broadly obovate, 2.5-4 cm.
long, spreading; fruit bright red, 2-3 cm. broad.
Known in British Honduras as "willow" and "good-luck seed";
"acitz" (Yucatan, Maya). The fruit is reputed to be poisonous.
Closely related to T. ahouai.
Thevetia ovata (Cav.) A. DC. in DC. Prodr. 8: 344. 1844. Cer-
bera ovata Cav. Icon. 3: 35, t. 270. 1796. C. cuneifolia HBK. Nov.
Gen. & Sp. 3: 224. 1819. T. cuneifolia A. DC. I.e. Cascabel; cascabel
de Lacandon; chilindron; manzana de Judas (Huehuetenango) .
Generally on dry brushy slopes or in thin rocky forest, 1,250 m.
or less; El Progreso; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Retalhuleu; Quiche"; Huehuetenango. Western and southern Mex-
ico; Costa Rica.
Usually a shrub of 2-3 m., sometimes a tree of 6 m., with thick branches;
leaves subcoriaceous, on petioles 5-10 mm. long, oblanceolate-oblong or cuneate-
oblong, 6-17 cm. long, 1-4 cm. broad, usually rounded at the apex, cuneate-
attenuate to the base, glabrous above or nearly so, paler beneath, puberulent, the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
397
FIG. 112. Thevetia ovata. A, habit, X H; B, fruits, apical and lateral views,
natural size; C, anthers in position on portion of corolla tube, X 3.
lateral nerves very numerous and prominent beneath, united to form a collective
nerve near the margin; inflorescence few-flowered; calyx lobes ovate, acuminate,
8-12 mm. long, reflexed, minutely puberulent; corolla bright yellow, glabrous or
somewhat papillate outside, the tube 1.5-2 cm. long, the throat campanula te, 12-
15 mm. long, 1-1.5 cm. broad at the orifice, the lobes obovate-dolabriform, 2.5-
3.5 cm. long, spreading; fruit 4.5-5 cm. broad, rose-colored mottled with purple.
The large seeds are light-colored, hard and bony. The name
"cascabel de Lacand6n" reported from Huehuetenango by Seler
leads one to suspect that the seeds probably are used by the Lacan-
don Indians as ornaments or rattles.
Thevetia peruviana (Pers.) Schum. in Engler & Prantl, Pflan-
zenfam. 42: 159. 1895. Cerbera thevetia L. Sp. PI. ed. 2. 304. 1762.
C. peruviana Pers. Syn. PL 1 : 267. 1805. T. neriifolia Juss. ex Steud.
398 FIELDIANA: BOTANY, VOLUME 24
Norn. Bot. 180. 1821. T. thevetia Millsp. Field Mus. Bot. 2: 83. 1900.
Chilindron, chilco, chilca, acitz (Pete"n, Maya), chirca, canjura (fide
Tejada).
Planted commonly for ornament, chiefly at low elevations, but
sometimes also at middle altitudes; sometimes more or less natural-
ized along roadsides and in pastures; not native in Guatemala. Native
of tropical America, perhaps of Mexico; now widely cultivated in the
tropics of both hemispheres.
A shrub or small tree, glabrous throughout, commonly 3-5 m. high, the trunk
short, with light gray bark, the branches often irregular; leaves thick-membrana-
ceous, short-petiolate, linear or nearly so, 7-15 cm. long, 6-13 mm. broad, long-
attenuate to each end, the lateral nerves obscure or obsolete; inflorescences lateral
or terminal, several-flowered; calyx lobes ovate or ovate-lanceolate, 4-6 mm. long;
corolla bright yellow or sometimes buff or reddish buff, the tube 1-1.5 cm. long,
the lobes broadly obovate, 3.5-4.5 cm. long, spreading; fruit becoming red at ma-
turity, fleshy, 4-4.5 cm. broad.
Sometimes called "campanilla" in Yucatan and "campanilla ama-
rilla" in El Salvador; "Yambigo" (Oaxaca). The wood is white, the
flowers fragrant. In Yucatan the latex is applied to decayed teeth
to relieve toothache, and also is used to treat chronic sores and ulcers.
This tree varies greatly in flower color, and the color variations de-
serve investigation. Unfortunately, many of the collections are either
without corollas or else their color is not described by the collector.
Most of many of the plants seem to have bright yellow flowers, but
some of those planted in Florida, as well as those of Central America,
have buff or reddish-buff corollas, giving the plants a quite different
appearance. In Florida this species usually is called ' 'yellow olean-
der"; in the West Indies the seeds are called "lucky nuts" and "luck-
seeds," and are sometimes carried in the pocket to bring good luck,
doubtless because of their curious form. The milky sap and the
seeds are reputed poisonous. A tincture of the bark has been em-
ployed in some regions as a febrifuge, and in large doses it is said to
be a violent purgative and emetic.
Thevetia plumeriaefolia Benth. Bot. Voy. Sulph. 124, t. 43.
1845. Chilindron.
Dry brushy rocky hillsides, about 850 m. ; Jutiapa (near Jutiapa,
Standley 75314) ', reported, perhaps in error, from Escuintla. Western
and southern Mexico; El Salvador; Honduras.
A shrub or small tree, glabrous throughout; leaves thick-memebranaceous or
subcoriaceous, oblanceolate-oblong or obovate-oblong, 5-12 cm. long, 1.3-3.5 cm.
FIG. 113. Tintinnabularia mortonii. A, habit, X Yi; B, flower, natural size;
C, stamens, X 2; D, style and stigma, X 2; E, inner face of anther, X 2. C, D,
and E after Woodson.
399
400 FIELDIANA: BOTANY, VOLUME 24
broad, rounded or very obtuse at the apex, cuneate-attenuate to the base, deep
green and lustrous above, paler beneath, the lateral nerves numerous and promi-
nent, the petioles 5-12 mm. long; inflorescence few-flowered, the flowers buff or
pinkish yellow; calyx lobes ovate, acute or acuminate, 4-6 mm. long, more or less
reflexed; corolla tube 15-22 mm. long, the throat conic-campanulate, 15-18 mm.
long, about 1 cm. broad at the orifice, the lobes obovate-dolabriform, 4.5-6 cm.
long, spreading; fruit broader than long, 3-3.5 cm. broad, dark purple at maturity,
the flesh white.
This is very closely related to T. ovata and rather doubtfully dis-
tinct from it. Called "chilindron" in El Salvador.
TINTINNABULARIA Woodson
Woody vines, glabrous throughout or nearly so; leaves opposite, glandular on
the upper surface at the base of the costa, usually pitted beneath in the axils of the
nerves; inflorescences alternate-axillary, corymbose-cymose, several-flowered, the
flowers large and showy; calyx 5-parted, the lobes subequal, foliaceous, bearing
groups of squamellae within; corolla funnelform, the tube straight, the throat
broadly tubular, the 5 lobes equal, dextrorsely contorted; anthers connivent and
agglutinated to the stigma, the connective enlarged, produced into 2 obtuse basal
lobes; ovary of 2 distinct carpels, each carpel many-ovulate, surrounded by 5 ovoid
nectaries; stigma fusiform-capitate; fruit unknown but doubtless follicular.
The genus is known at present only from Guatemala, and consists
of a single species.
Tintinnabularia mortonii Woodson, Ann. Mo. Bot. Gard. 23:
389, t. 7. 1936.
Wet thickets or in brushy swamps, 600-2,200 m.; Alta Verapaz,
the type from Quebradas Secas, Johnson 200; Zacapa (Sierra de las
Minas).
A large woody vine; leaves thick-membranaceous, on petioles 7-10 mm. long,
oblong-elliptic, 9-10 cm. long, 3-3.5 cm. broad, caudate-acuminate with a long
obtuse acumination, obtuse at the base, anisophyllous; inflorescences pendent, the
pedicels about 2 cm. long; calyx lobes oblong-elliptic, acuminate, 12-13 mm. long,
somewhat foliaceous, spreading; corolla pale reddish outside, cream-colored within,
puberulent-papillate, the tube 8-9 mm. long, the throat broadly tubular, 3-3.5 cm.
long, 8 mm. broad at the orifice, the lobes obliquely obovate, subobtuse, 1 cm.
long, slightly spreading; anthers wholly included, bearing a linear pilosulous api-
cal appendage.
The plant appears to be very rare in Alta Verapaz, and in the
large swamp east of Tactic only a single individual was discovered.
The large pendent flowers are exceptionally handsome. It has not
been re-collected in the more than 30 years since it was described.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 401
TONDUZIA Pittier
Shrubs or trees; leaves in whorls of 3-4, rarely opposite, eglandular; inflores-
cences lateral or subterminal, thyrsiform, the flowers numerous, small, white or
cream-colored; calyx 5-parted almost to the receptacle, the lobes subequal, slightly
imbricate, without squamellae; corolla salverform, not appendaged within, the
limb equally 5-parted, sinistrorsely convolute; anthers not connivent, wholly in-
cluded, the connective not enlarged; ovary of 2 distinct carpels, containing several
biseriate ovules, surrounded at the base by a low annular nectary, or the nectary
absent; fruit of 2 follicles, these dehiscent, terete; seeds numerous, compressed,
dry, densely ciliate on the margins.
The genus consists of only the following species. It was named
for Adolfo Tonduz, born September 18, 1862, at Fully, near Lau-
sanne, Switzerland; died in Guatemala City in 1921. He went to
Costa Rica in 1888 and was engaged in botanical exploration there
for many years, assembling a large herbarium of Costa Rican plants.
Toward the end of his life he went to Guatemala, and was employed
as a botanical collector in the Direction General de Agricultura until
his death. He collected the types of many hundreds of Central
America plants, and was a most capable and assiduous collector.
Corolla 15-18 mm. long T. macrantha.
Corolla 8-10 mm. long.
Leaves 6-7 times as long as broad; stamens inserted at about the lower third of
the corolla tube T. longifolia.
Leaves 2-4 times as long as broad; stamens inserted near or but slightly below
the middle of the corolla tube T. pittieri.
Tonduzia longifolia (A. DC.) Woodson, No. Am. Fl. 29: 122.
1938. Rauwolfia longifolia A. DC. in DC. Prodr. 8: 338. 1844. R.
stenophylla Donn.-Sm. Bot. Gaz. 44: 115. 1907. T. parvifolia Pittier,
Contr. U. S. Nat. Herb. 12: 103. 1908 (type from San Salvador, El
Salvador). T. stenophylla Pittier, I.e. 104. Chilindrillo.
Dry or wet, mixed forest or thickets, often in rocky places,
1,650 m. or less; Izabal; El Progreso; Escuintla; Guatemala; Huehue-
tenango; Quezaltenango. Southern Mexico; Honduras; El Salvador;
Costa Rica.
A shrub or tree, sometimes 15m. tall, often a shrub of only 2-3 m., glabrous
throughout or nearly so; leaves 3-4-nate, or rarely opposite above, nrm-membra-
naceous, on petioles 4-15 mm. long, narrowly oblong-lanceolate or linear-lanceo-
late, 5-20 cm. long, 1-4 cm. broad, very narrowly long-attenuate, attenuate to the
base, green above, pale beneath, the nerves obscure; inflorescence corymbose, con-
gested, many-flowered, much shorter than the leaves, the pedicels 3-4 mm. long;
calyx lobes ovate-triangular, acute, 0.5 0.7 mm. long; corolla white, the tube 4 mm.
long, the lobes broadly oblong-dolabriform, rounded at the apex, 4-5 mm. long,
402 FIELDIANA: BOTANY, VOLUME 24
spreading; stamens inserted near or slightly below the middle of the tube; follicles
7-11 cm. long, terete, longitudinally striate, attenuate at each end, 5 mm. thick.
Known in El Salvador as "amatillo" and "chilindron de montana."
Tonduzia macrantha Woodson, Ann. Mo. Bot. Gard. 24: 12.
1937 (type from Volcan de Zunil, Skutch 871). T. longipedunculata
Woodson, in Standl. & Steyerm. Field Mus. Bot. 23: 78. 1944 (type
from Solola, Steyermark 47313).
Dry thickets and forest edges or in second growth, 1,500-2,300 m.
or perhaps higher, endemic but to be expected in adjacent Mexico;
Quezaltenango; San Marcos; Solola.
A glabrous tree or shrub of 6-9 m.; leaves ternate or the upper ones opposite,
firm-membranaceous, very narrowly oblong-lanceolate, 10-20 cm. long, 2-4 cm.
broad, very narrowly long- attenuate, attenuate to the base, paler beneath, the
nerves inconspicuous, the petioles 1-2 cm. long; inflorescence terminal or sub-
terminal, irregularly cymose, much shorter than the leaves, few-many-flowered,
the pedicels 5-7 mm. long; calyx lobes ovate-deltoid, rounded at the apex, 1-
1.2 mm. long; corolla white or cream-colored, the tube 7 mm. long, the lobes oblong-
dolabriform, rounded at the apex, 9-10 mm. long, spreading; follicles stout, 10-12
cm. long, 8 mm. thick, acuminate, longitudinally striate.
Tonduzia pittieri Donn.-Sm. Bot. Gaz. 49: 456. 1910.
Moist or dry forest or thickets, often in dry rocky places, 400-
1,100 m.; endemic; Zacapa; El Progreso; Baja Verapaz; Guatemala
(type from Fiscal, Deam 6098).
A shrub or small tree, 3-4 m. high, with slender branches, glabrous throughout;
leaves 3-4-nate, or the upper rarely opposite, firm-membranaceous, narrowly ellip-
tic to oblong-lanceolate, 5-12 cm. long, 1-3 cm. broad, long-acuminate, attenuate
to the base, the petioles 4-5 mm. long; inflorescence corymbose, congested, much
shorter than the leaves, many-flowered, the pedicels 3-4 mm. long; calyx lobes
ovate-triangular, acute, 0.5-0.7 mm. long; corolla tube 4 mm. long, the lobes rather
broadly oblong-dolabriform, rounded at the apex, 4-5 mm. long, spreading; sta-
mens inserted near or slightly below the middle of the corolla tube.
This is very close to T. longifolia and questionably distinct, espe-
cially since the two do not have distinct ranges.
URECHITES Mueller von Argau
Reference: Robert E. Woodson, Ann. Mo. Bot. Gard. 23: 198-205.
1936.
Woody or suffrutescent vines; leaves opposite or rarely subverticillate, eglandu-
lar; inflorescences alternate-axillary to terminal or sub terminal, scorpioid, simple,
few-many-flowered, the flowers large, yellow; calyx 5-parted almost to the recep-
tacle, the lobes subequal, imbricate, bearing within at the base binate and alternate
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
FIG. 114. Tonduzia macrantha. A, habit, X 2^; B, fruit, with one follicle de-
hisced; C, calyx partially dissected with pistil, X 10; D, stamens in natural position
on part of dissected corolla tube, X 5; E, flower, natural size.
or numerous and indefinitely distributed squamellae; corolla funnelform, the tube
not appendaged within, the limb regularly 5-parted, dextrorsely contorted; anthers
connivent and agglutinated to the stigma, the connective enlarged, narrowly bi-
lobate, appendaged at the apex; ovary of 2 distinct carpels, many-ovulate, sur-
rounded at the base by 5 distinct or concrescent nectaries; stigma fusiform-capitate;
follicles 2, distinct; seeds numerous, dry, narrowly rostrate, comose at the apex.
404
FIELDIANA: BOTANY, VOLUME 24
FIG. 115. Urechites andrieuxii. A, habit, X 1A', B, calyx and pistil with part
of the callus-like corona dissected away to show ovary, X 4; C, anthers in place
on dissected portion of corolla tube, X 5; D, follicles, X YIL-
Two species, the other in Florida and the West Indies.
Urechites andrieuxii Muell.-Arg. Linnaea 30: 442. 1860.
Wet or dry thickets or mixed forest, often in mangrove swamps,
600 m. or less; Pete"n; Izabal; Zacapa; Chiquimula; Santa Rosa;
Retalhuleu. Southern Mexico; British Honduras. Contrayerba.
A -small or large vine, usually woody, sometimes 6 m. long; leaves opposite,
membranaceous or subcoriaceous, on petioles 8-20 mm. long, oblong-elliptic to
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 405
ovate-elliptic or obovate-elliptic, 5-12 cm. long, 2.5-7 cm. broad, short-acuminate,
obtuse or rounded at the base, glabrous above, paler beneath, glabrous or finely
puberulent; inflorescences usually somewhat longer than the leaves, few-several-
flowered, the pedicels 15-22 mm. long, glabrous or inconspicuously puberulent, the
bracts minute, scarious; calyx lobes ovate, obtuse, 3-5 mm. long, the squamellae
binate, alternate with the lobes; corolla yellow, glabrous, the tube 1-1.5 cm. long,
the throat broadly tubular, 2.5-3 cm. long, 5-8 mm. broad at the orifice, the lobes
oblique-obovate, 1.5-2.3 cm. long, spreading; nectaries completely concrescent,
twice as long as the ovary; follicles rather stout, continuous, glabrous, about
25 cm. long.
The Maya name of Yucatan is "cantibteac." The nerves and
veins, although not elevated, form a conspicuous close network on
the lower leaf surface. The species is reported by Lundell to be used
in Peten to treat headaches.
VALLESIA Ruiz & Pavon
Shrubs or small trees; leaves alternate, subtended by small dentate axillary
stipular glands; inflorescence lateral, umbellate or umbellate-cymose, simple or
dichotomous, several-many-flowered, the flowers small, white or yellowish; calyx
5-parted almost to the receptacle, the lobes equal, without squamellae; corolla sal-
verform, the limb regularly 5-parted, sinistrorsely contorted; anthers not conni-
vent, the connective not enlarged; ovary of 2 distinct carpels, without a nectary,
the carpels 1-2-ovulate; fruit drupaceous, usually a single carpel developing, juicy.
About eight species, in tropical America. One other is found in
Central America (Costa Rica).
Vallesia mexicana Muell.-Arg. Linnaea 30: 393. 1860.
Usually in dense, wet or moist, mixed forest, 1,500-2,600 m.;
Guatemala; Sacatepe"quez ; Chimaltenango; Quiche; Huehuetenango;
San Marcos. Mexico.
A glabrous shrub or tree, 3-12 m. high; leaves firm-membranaceous or sub-
coriaceous, on petioles 5-8 mm. long, narrowly oblong to oblong or lance-oblong,
6-17 cm. long, 2-5 cm. broad, acuminate or acute, obtuse or rounded at the base;
inflorescence umbelliform, simple or dichotomous, much shorter than the leaves,
many-flowered, the pedicels 2-4 mm. long, minutely puberulent-papillate; calyx
lobes ovate, acuminate, 1.5-2 mm. long; corolla white, glabrous, the tube 10-13
mm. long, somewhat dilated at the insertion of the stamens, constricted at the
orifice, the lobes oblique-oblong, spreading; drupes white, juicy, oblong-ovoid,
the seed 17-20 mm. long.
VINCA L.
Perennial herbs, erect or trailing; leaves opposite, eglandular; flowers solitary
in the leaf axils, pedicellate, usually blue or violet; calyx 5-parted. the lobes sub-
406
FIELDIANA: BOTANY, VOLUME 24
FIG. 116. Vallesia mexicana. A, habit, X 1A; B, corolla, X 2; C, calyx and
pistil, X 5; D, stigma, X 10; E, stamens, X 5; F, fruits, natural size.
equal, without squamellae; corolla funnelform, equally 5-lobate, not appendaged
within; anthers not connivent, the connective produced into a rather large, apical
appendage, not appendaged at the base; ovary of 2 distinct carpels, accompanied
by 2 alternate nectaries of almost equal size, many-ovulate; follicles 2, terete,
slender, seeds numerous, naked, subcompressed.
Species 5, in Europe, Asia, and northern Africa. Two often are
grown for ornament in temperate and subtropical regions.
Vinca major L. Sp. PL 209. 1753. Sereno, Vinca, pervinca
(Coban).
Native of southern Europe and northern Africa, often planted for
ornament in other regions; frequent in Guatemala gardens, chiefly in
the higher mountains; thoroughly naturalized in thickets or open
places in Quezaltenango and probably also San Marcos, mostly at
2,000-3,000 m.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 407
Plants stout, perennial, erect, ascending, or sometimes scandent, the sterile
shoots often elongate and rooting, almost wholly glabrous; leaves persistent, thick,
deep green, often ciliate, broadly deltoid-ovate, mostly 4-5.5 cm. long, obtuse,
rounded or subcordate at the base; pedicels shorter than the subtending leaves;
calyx lobes linear, green, ciliate; corolla blue or violet, glabrous, the tube 1.5 cm.
long, the lobes cuneate-obovate, equalling or slightly longer than the tube.
Probably introduced into Guatemala directly from Europe long
ago, and now thoroughly naturalized at many places about Quezal-
tenango, and at various localities in the department of that name.
The plant is often cultivated in patios, sometimes in pots, and at
Quezaltenango there was observed a form with white-edged leaves.
The plant is noteworthy in the western highlands because it blooms
freely during the cold dry season when flowers are very scarce.
ASCLEPIADACEAE Milkweed Family
Reference: Robert E. Woodson, Jr., The North American Asclepi-
adaceae. I. Perspective of the genera, Ann. Mo. Bot. Gard. 28:
193-244. 1941.
Mostly perennial herbs, erect or scandent and twining, sometimes prostrate,
frequently more or less woody, generally pubescent, sometimes glabrous, the sap
usually milky; leaves commonly opposite, rarely verticillate; stipules none; in-
florescence normally cymose, but the cymes usually umbelliform or racemiform,
rarely 2-3 times branched, the inflorescences commonly axillary and pedunculate
or sessile; flowers small or medium-sized, white, pink, green or brown-purple, of
very complicated structure, perfect, regular; calyx tube very short or none, the 5
lobes imbricate or open in bud; corolla sympetalous, rotate to campanulate or urce-
olate, shallowly or deeply 5-lcbate, the lobes valvate or more commonly contorted;
a corona often or usually present, simple or of 5 or more scales or lobes, adnate to
the tube of the corolla or to the staminal column, highly varied as to structure;
stamens 5, inserted at or near the base of the corolla, the filaments flat, short,
usually connate to form a tube, the stamens and stigma united to form the gyno-
stegium1; anthers introrsely 2-celled, often produced at the base, the connective
often produced at the apex into a membranaceous appendage; pollen collected in
masses called pollinia, these usually solitary in each anther cell; corpuscles 5, small,
usually colored, agglutinate between the anthers to the margin of the stigma disk,
after dehiscence of the anthers extruding the pollinia; disk none; carpels of the
ovary 2, distinct, superior, included in the stamen tube; styles 2, short, distinct to
the stigma; stigma 1, peltately dilated and forming a usually pentagonal disk, flat
at the apex or depressed, convex, umbonate, or rostrate; ovules numerous in each
cell, pendulous, imbricate upon the placentae; fruit of 2 parallel or divaricate fol-
licles, or the follicles by abortion only 1, sessile, generally ovoid or lanceolate and
acuminate, smooth or muricate, sometimes longitudinally winged, ventrally de-
hiscent; seeds rather large, compressed and often marginate, the testa thick or
membranaceous, narrowed at the apex and generally bearing a tuft of long soft
1 Somewhat similar to the column in Orchidaceae.
408 FIELDIANA: BOTANY, VOLUME 24
white silk-like hairs; endosperm cartilaginous, thin; embryo large, the cotyledons
flat, the radicle short, superior.
A group of wide distribution in most temperate and tropical re-
gions of the earth. Authors have recognized as many as 250 genera,
but if the family is aligned in accordance with the treatment used
here, the number would be probably not more than a fifth as many.
The Asclepiadaceae of North America are referred by Woodson to nine
genera, about a third as many as have been recognized by most auth-
ors. All of these genera are represented in the following treatment.
The realignment of the genera by Woodson, while revolutionary,
has done much to bring order out of the chaos previously prevailing
so far as North American genera are concerned, and while to some it
may appear too radical, it must be recognized that if deviations from
it are made, a host of others must accompany them. The complexity
of the flowers, or of the androecium, more varied and highly special-
ized probably than in any other group of plants, has offered unlimited
possibilities for proposal of new genera, and if this is once begun, there
are no limits to which an enthusiastic amateur may extend his work.
It must be confessed that this recent disposition of genera does not
make it much easier to place a plant in its genus, but the Asclepia-
daceae seldom offer vegetative characters that are useful for distin-
guishing the generic groups. The Central American plants of the
family fall naturally into two groups, those with erect stems and those
with twining ones. Unfortunately, because of flower structure, it is
necessary in case of the Guatemalan species to place two plants with
erect stems in groups whose other representatives have scandent
stems.
Plants of this family are of little economic importance. The silky
fiber of the seeds looks much like silk, but it is brittle and unsuited
for textile purposes. In Central America the young and tender seed
pods of some of the scandent Asclepiadaceae are used as a vegetable,
cooked in various ways or sometimes, it is said, eaten raw.
Pollinia pendulous, their faces uniformly flattened or rounded, uniformly fertile
up to the point of attachment of the translators; plants erect or scandent
[Asclepiadeae].
Plants erect or nearly so, herbaceous, never scandent Asdepias.
Plants scandent, herbaceous or somewhat woody.
Flowers very small, mostly 2-3 mm. long, rarely slightly larger; inflorescence
umbellate or racemose Cynanchum.
Flowers larger, usually much larger, 5 mm. long or more; inflorescence um-
bellate.
Lobes of the corolla linear, 15-20 mm. long; leaves ovate-cordate.
Oxy petal um.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 409
Lobes of the corolla not linear, or if so, much less than 10 mm. long.
Corona of 5 subvescicular sacs attached separately to the backs of the
anthers; plants glabrous; leaf blades rounded at the base.
Blepharodon.
Corona of 5 closed vesicles joined at their bases by a fleshy ring adnate
to the corolla throat; plants glabrous or usually pubescent; leaf
blades acute to cordate at the base Sarcostemma.
Pollinia horizontal or erect.
Pollinia strictly erect, their faces uniformly rounded, uniformly fertile to the
point of attachment of the translators; inflorescences usually 2-3 times
branched; plants scandent or in one species erect [Tylophoreae]. . Marsdenia.
Pollinia usually horizontal or nearly so, occasionally ascending or descending,
but one or both faces more or less excavated, and with a sterile hyaline
margin or indentation near the point of attachment of the translators;
plants usually scandent, erect in only one species [Gonolobeae].
Corolla lobes conspicuously crispate; anthers very conspicuously hypertro-
phied and vesicular throughout; plants scandent Fischeria.
Corolla lobes not crispate; anthers not conspicuously hypertrophied and
vesicular.
Anthers relatively simple, not with dorsal appendages; plants scandent,
except for one species, this an erect herb Matelea.
Anthers with spreading, more or less laminate, fleshy, dorsal appendages;
plants scandent Gonolobus.
ASCLEPIAS L. Milkweed
Reference: Robert E. Woodson, Jr., North American Species of
Asclepias, in Ann. Mo. Bot. Card. 41: 1-211. 1954.
Perennial herbs, usually erect, glabrous or pubescent, sometimes tomentose;
leaves opposite or verticillate; flowers variously colored, usually white or purplish,
sometimes bright red or orange, small or medium-sized, umbellate, the umbels
usually many-flowered, terminal or in the upper leaf axils; calyx 5-parted, 5-10-
glandular within at the base, the lobes usually acute; corolla rotate or finally re-
flexed, deeply 5-lobate, the lobes usually valvate, not barbate within; scales of the
corona 5, affixed to the stamen tube, erect, concave-cucullate, ligulate within;
stamens affixed to the base of the corolla, the filaments connate into a tube; an-
thers tipped with an inflexed membrane; pollinia solitary in each cell, oblong,
pendulous from the apex of the cell; stigma flat at the apex, pentagonal or obtusely
5-lobate; follicles usually thick, acuminate, smooth or rarely spinose-tuberculate;
seeds comose.
The genus Asclepias with more than 100 species in temperate
North America and Mexico has relatively few species in Central
America. There are ten species in Guatemala, none of which is en-
demic, and all of them extend southward from Mexico. No addi-
tional species are known in Central America. Only the ubiquitous
and often weedy A. curassavica and the rare A. woodsoniana extend
as far south as Panama. There is an additional American center of
dispersal of Asclepias in subtropical South America.
410 FIELDIANA: BOTANY, VOLUME 24
Leaves sessile or essentially so, often cordate or subcordate at the base.
Leaves linear; usually not more than 3 mm. broad; stems and lower surface of
the leaves more or less tomentose A. rosea.
Leaves much broader than linear, mostly 2 cm. broad or usually broader; plants
not at all tomentose.
Stems and leaves hirsute or villous A. contrayerba.
Stems and leaves glabrous or the stems sometimes pubescent at the apex.
Column or gynostemium sessile or nearly so, the horns poorly developed.
A. elata.
Column or gynostemium stipitate (sometimes subsessile), the horns well
developed and exserted A. glaucescens.
Leaves usually conspicuously petiolate, the blades obtuse to attenuate at the base.
Leaves linear or nearly so, less than 1 cm. broad; flowers white and pale green
or greenish white A. woodsoniana.
Leaves lanceolate or broader, usually much more than 1 cm. broad, if narrow
the flowers scarlet and yellow.
Scales of the corona more than twice as long as the stamens.
Scales of the corona 6-7 mm. long; leaves usually obtuse, rough-villosu-
lous on the upper surface A. oenotherioides.
Scales of the corona about 9 mm. long; leaves acuminate, glabrous on the
upper surface A. auriculata.
Scales of the corona not or scarcely exceeding the stamens.
Follicles glabrous.
Corolla bright red; hoods of corona yellow or orange; leaves not promi-
nently bicolored on the surfaces A. curassavica.
Corolla dull rose; hoods of corona white to rose-colored; leaves obvi-
ously lighter in color below than above A. pellucida.
Follicles pubescent; hoods of the corona white or greenish-white; corolla
greenish-white or purple A. similis.
Asclepias auriculata HBK. Nov. Gen. & Sp. PI. 3: 191, t. 228.
1819. Otaria fragrans Benth. PL Hartweg. 89. 1841 (type from Gua-
temala, Hartweg, without number, the locality not indicated). A.
fragrans Dene, in DC. Prodr. 8: 571. 1844. A. purpusii Brandegee,
Zoe 5: 251. 1908. Hierba de cantil, viborana blanca (fide Aguilar).
Usually in open pine-oak forest, 1,200-1,900 m.; Zacapa; Santa
Rosa; Guatemala; Chimaltenango; Huehuetenango. Southern
Mexico.
A stout erect herb, sometimes a meter high, usually simple, the stems terete,
more or less pubescent or glabrate; leaves opposite, on short stout petioles, lance-
oblong or lanceolate, mostly 9-12 cm. long and 2.5-3.5 cm. broad, acuminate, acute
at the base or rarely obtuse, green above, pale beneath, glabrous or nearly so; um-
bels few, terminal and in the upper leaf axils, long-pedunculate, lax, many-flowered,
the slender pedicels about 2.5 cm. long, pubescent; calyx lobes puberulent, narrowly
lanceolate, short; corolla pale green or whitish, glabrous, the lobes oblong-ovate,
subacute, reflexed; corolla white, the lobes about 9 mm. long, about twice as long
as the stamens, white, ovate-oblong, broadest at the base.
It is probable that Asclepias plumerifolia Ramirez-Goyena, Fl.
Nic. 2: 658. 1911, from Nicaragua, belongs here as a synonym.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 411
Asclepias contrayerba Sesse & Mocifio, Fl. Mex. ed. 1, 2: 79.
1887. Sicaquina; jicaca. A. setosa Benth. PL Hartw. 24. 1839,
non Forsk.
Moist or dry, open fields, or most often in grassy pine-oak forest,
1,950 m. or lower; Pete"n; Jalapa; Guatemala; Solola; Huehuete-
nango. Mexico; El Salvador.
Plants herbaceous, erect, 40 cm. high or less, the stems stout, simple or
branched, villous-hispid, usually bearing numerous leaves; leaves opposite, sessile
or nearly so, thick, oblong-lanceolate or narrowly oblong, mostly 4-6 cm. long,
acute or obtuse at the apex, rounded or subcordate at the base, rather densely
villous on both surfaces with several-celled pale hairs; umbels usually 1-2 at the
apex of the stem, on rather short, slender peduncles, the slender pedicels mostly
1.5-2 cm. long, short-villous; calyx lobes lanceolate, short, short-villous, 5-6 mm.
long; corolla pale green, glabrous, 5-10 mm. long, the lobes reflexed; gynostegium
borne on a distinct thick column, the corona hoods whitish, 4 mm. long, broad,
cucullate, the horns short-exserted ; pedicels reflexed in fruit; follicles 6-11 cm. long
and 1.2-2.5 cm. broad, densely hispid.
Called "ishcaco" in El Salvador.
Asclepias curassavica L. Sp. PL 215. 1753. Viborana, viborrana
(sometimes corrupted to vibarona); mariguana amarilla (Chimalte-
nango) ; hierba de leche; olh'en (Coban, Quecchi) ; seda (Coban) ; hierba
de cantil, hierba de culebra (fide Tejado) ; cantil.
Moist or wet thickets or fields, often along roadsides or in waste
ground about dwellings, 1,900 m. or lower; Pete"n; Alta Verapaz; El
Progreso; Izabal; Chiquimula; Jalapa; Santa Rosa; Escuintla; Gua-
temala; Sacatepe"quez; Chimaltenango ; Suchitepe"quez ; Retalhuleu;
Quezaltenango; San Marcos; Huehuetenango; doubtless in most
other departments. Florida; Mexico; British Honduras to El Sal-
vador and Panama; West Indies; South America.
An erect herb, usually 60-75 cm. high, the stems often several, mostly simple,
green, glabrous or somewhat pubescent above; leaves opposite or sometimes partly
ternate, lanceolate or linear-lanceolate, 5-16 cm. long, attenuate-acuminate, acute
or attenuate at the base, short-petiolate, glabrous or sparsely and inconspicuously
pubescent, slightly paler beneath; umbels usually several, rather few-flowered, the
pedicels 1-2 cm. long, pubescent; corolla bright red or orange-red, 6-8 mm. long,
the lobes ovate to oblong; hoods of the corona erect, broadly ovate, 4-5 mm.
long, rounded or very obtuse at the apex, bright yellow, shorter than the conspic-
uous horns of the corona; fruiting pedicels erect; follicles narrowly fusiform, 4-
10 cm. long, glabrous; seeds about 6 mm. long.
Called "Polly redhead" in British Honduras; "senorita," "flor de
agua" (El Salvador); "flor de culebra" (Veracruz); variously known
in Yucatan as "chacnich," "chililoo-xiu," "analcac," "chontalpa,"
412 FIELDIANA: BOTANY, VOLUME 24
"cabal-cumche," "cochinita" (Tabasco), "quema-casas" (seeds; Ta-
basco). About Coban the milk of the stems is collected on cotton
and when dry is placed in the nostrils. It produces sneezing, and is
believed to have a beneficial effect on head colds. The latex often
is placed in cavities in the teeth to relieve toothache, and sometimes
it is used internally as an emetic and purgative. In Guatemala it is
popularly believed to be useful in treating bites of poisonous animals.
The plant is one of the most common weeds of the lowlands of Cen-
tral America, rather showy because of its bright red and yellow
flowers. There is sometimes found in Central America (we have not
observed it in Guatemala) a form in which the flowers are bright
yellow throughout, without any red coloring, A. curassavica var.
flava Kuntze.
Asclepias elata Benth. PI. Hartw. 290. 1848; Woodson, Ann.
Mo. Bot. Gard. 41: 186, fig. 1954. A. glaucescens var. elata Fourn.
Ann. Sci. Nat. Bot. ser. VI, 14: 382. 1882.
Open slopes in oak or pine woods, spreading to cultivated ground.
Huehuetenango; Chimaltenango. Southwestern United States;
Mexico.
Perennial herbs with a stout simple stem up to 70 cm. tall. Leaves sessile,
broadly oval to oblong, obtuse or rounded, the base subcordate and the lobes often
clasping the stem, 7-14 cm. long and 2.5-7 cm. broad, glabrous and glaucous; in-
florescence usually lateral from upper leaves or terminal, exceeding the leaves, sub-
umbellate, the pedicels 1.5-3 cm. long, tomentulose; calyx lobes ovate, 3-4 mm.
long; corolla pale green, the lobes 8-12 mm. long; gynostemium sessile, the hoods
saccate at the base, truncate, 4-6 mm. long, the crest adnate and about as long as
the hood; follicles fusiform, 8-12 cm. long, pilosulose to glabrous; seeds oval, about
7 mm. long, the coma 2.5-4.5 cm. long.
The species is known from a single collection in Guatemala. The
species has been confused with A. glaucescens from which it may be
distinguished easily by its sessile gynostemium.
Asclepias glaucescens HBK. Nov. Gen. & Sp. PI. 3: 190, t. 227.
1819; Woodson, Ann. Mo. Bot. Gard. 41: 97. 1954. Polin (Huehue-
tenango).
Native in open pine or oak woods but found in disturbued ground
and old fields. Pete"n; Alta Verapaz; Guatemala; Jalapa; Chimalte-
nango; Solola; Huehuetenango; Chiquimula; perhaps to be found in
most departments. Mexico; El Salvador; Costa Rica.
Herbaceous perennials, stout and erect, to 1 m. tall but usually less, glabrous
and glaucous. Leaves sessile or nearly so, ovate to oval or oblong, often narrowly
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 413
so, obtuse or broadly acute, the base cordate or amplexicaul, 5-18 cm. long and
2-7 cm. broad; inflorescences one or few from axils of upper leaves or occasionally
terminal, flowers borne in often long pedunculate subumbellate heads, the pedicels
about 1 cm. long, glabrous; calyx lobes ovate-lanceolate, 4-5 mm. long; corolla
lobes reflexed, about 9 mm. long; gynostemium short stipitate to subsessile, the
column about 1 mm. long, hoods obovate, conduplicate, rounded and frequently
emarginate, 5-7 mm. long; follicles narrowly fusiform, erect, 8-10 cm. long and
1 cm. in diameter, smooth and glabrous; seeds oval, 6-8 mm. long, the coma 2.5-
3.5 cm. long.
Asclepias glaucescens and A. elata are difficult to separate. Wood-
son gives characters for their separation and placed them in different
subgenera but was dubious as to the systematic value of these sub-
genera.
The species is known in El Salvador by the expressive names
"matacoyote," "jicaca" and "oreja de burro."
Asclepias oenotherioides Cham. & Schlecht. Linnaea 5: 123.
1830. A. longicornu Benth. PL Hartw. 24. 1839.
Moist or dry open fields or hillsides, sometimes on disturbed soils
at 1,400 m. or less; Pete"n; Izabal; Guatemala; Zacapa; Jutiapa; El
Progreso; Retalhuleu; San Marcos; and probably in other depart-
ments. Mexico; El Salvador; Honduras; Nicaragua and Costa Rica.
Plants herbaceous, erect, stout, mostly 50 cm. high or less, the stems mostly
simple, often densely leafy, rough-villosulous; leaves short-petiolate, lance-oblong
or ovate-oblong, 5-12 cm. long, rather thick, narrowed to an obtuse or rounded
apex, sometimes subacute, acute or cuneate at the base, sparsely short-villosulous
with harsh hairs and somewhat rough to the touch, the margins generally undulate;
umbels usually several, in the upper leaf axils, short-pedunculate or almost sessile,
the flowers pale green, few or numerous, on long slender pedicels; corolla pale
green, hispidulous outside, the lobes lanceolate, reflexed, 9 mm. long; hoods of the
corona very narrow below, dilated above, 7 mm. long, fully twice as long as the
gynostegium, the horns short-exserted; corona sessile or nearly sc; pedicels reflexed
in fruit; follicles thick, 8 cm. long, tomentose or glabrate.
Known in El Salvador by the names "matacoyote," "angelito,"
and "cuchamper de zope," "cabalcunche"" (Yucatan, Maya). The
latex of this and probably all the other species is used commonly for
alleviating toothache, and it often is employed in Central America
for removing warts, a use made of milkweed sap in many parts of the
United States.
Asclepias pellucida Fourn. Ann. Sci. Nat. ser. VI, 14: 381. 1882.
Known from a single collection in Guatemala, Santa Eulalia,
Huehuetenango. Mexico.
414 FIELDIANA: BOTANY, VOLUME 24
Simple, erect perennial herbs, glabrous or somewhat pilose at the nodes, to
1 m. tall but usually much less. Leaves ovate to oblong-elliptic, acuminate, the
base rounded or obtuse, glabrous or pilosulose on the mid-vein below, 7-30 cm.
long and 2.5-9 cm. broad, petioles 2-3 cm. long; inflorescences one-few at the upper
internodes, the flowers few to many disposed in a subumbel with pedicels 2-4 cm.
long; calyx lobes oblong-lanceolate, 3-4 mm. long; corolla lobes elliptic-ovate, 6-
8 mm. long; gynostemium short stipitate, column obconic, the hood cucullate,
obovate, 2.5-3.5 mm. long, with a short falciforme horn; follicles glabrous, broadly
ovoid, 7-8 cm. long; seeds oval, naked, about 1.5 cm. long.
This and A. similis are separated easily in the key which we pre-
sent but it may be difficult to separate the species if specimens are
not good or complete.
Asclepias rosea HBK. Nov. Gen. & Sp. PI. 3: 189. 1819. Vibo-
rana, platail, cicuta (fide Aguilar).
Open slopes or grassy fields, 800-2,000 m.; Santa Rosa; Guate-
mala; Chimaltenango; Solola; Quiche; Huehuetenango(?). Central
and southern Mexico; El Salvador; Honduras; Nicaragua.
Plants low, mostly 50 cm. high or less, the roots often greatly tuberous-thick-
ened or hard and woody, the stems usually several, erect or ascending, sometimes
sparsely branched, glabrous or when young somewhat whitish-tomentose; leaves
opposite, rather thick, linear, 2-18 cm. long, ascending, 2-6 mm. broad, often with
revolute margins, narrowed at each end, glabrous above or nearly so, whitish-
tomentose beneath when young; inflorescences few, on long slender, terminal or
lateral peduncles, these usually longer than the leaves, few-flowered or many-
flowered, the pedicels pubescent or tomentose, mostly 7-20 mm. long; corolla
greenish white or tinged with pink, glabrous, the lobes broad, 4-7 mm. long;
gynostegium borne on a very short column; hoods of the corona white, scarcely
more than 3 mm. long, little exceeding the anthers, the horns not or scarcely ex-
serted; fruiting pedicels reflexed; follicles slender, 6-15 cm. long, about 7 mm.
broad, tomentulose.
This is doubtless the species reported by Loesener from Huehue-
tenango as A. michauxii Dene.
Asclepias similis Hemsl. Biol. Cent. Am. Bot. 2: 326. 1881.
A. guatemalensis Donn.-Sm. Bot. Gaz. 18: 207. 1893 (type from San-
tiago, Sacatepe"quez, Guatemala, Rosalio Gomez 809}. Mishito (Chi-
maltenango); chumimi (Huehuetenango).
Moist thickets or open forest, most often in pine-oak forest, 1,400-
2,600 m.; Alta Verapaz; Zacapa; Jalapa; Guatemala; Sacatepe"quez ;
Chimaltenango; Quiche"; Solola; Huehuetenango. Mexico; Honduras.
FIG. 117. Asclepias woodsoniana. A, habit, X %; B, flower, in natural posi-
tion, X 4; C, calyx and pistil, X 8; D, pollinia, X 10.
415
416 FIELDIANA: BOTANY, VOLUME 24
An erect, rather stout herb a meter high or less, the stems usually simple, pu-
bescent at first, glabrate in age; leaves opposite, short-petiolate, oblong-ovate to
oblong-lanceolate, mostly 7-13 cm. long, acute or acuminate, rounded to acute at
the base, green above, sparsely pubescent or glabrate, pale beneath, puberulent
beneath on the veins, or sometimes more densely pubescent or glabrate; umbels
few at the ends of the stems and in the uppermost leaf axils, long-pedunculate,
many-flowered, the slender pedicels 12-25 mm. long, puberulent; calyx lobes ob-
long-lanceolate, subacute, glabrous; corolla purplish or usually greenish white or
pale green, 7-8 mm. long; gynostegium borne on a distinct thick column; hoods
of the corona white, sessile, about 4 mm. long, the horns usually exserted and con-
spicuous; pedicels reflexed in fruit; follicles about 13 cm. long and 1.5 cm. thick,
variously pubescent or in age glabrate.
It is presumably this species that was reported from Huehuete-
nango by Loesener as A. ovata Mart. & Gal.
Asclepias woodsoniana Standl. & Steyerm. Field Mus. Bot. 23:
224. 1947; Dugand, Caldesia 9, No. 45: 403. 1966.
Known in Guatemala only from the type, Jutiapa, salt flats, fields
between Trapiche Vargas and Asuncion Mita, 500 m., Steyermark
31781. Mexico; El Salvador; Honduras; Nicaragua; Costa Rica;
Colombia.
An erect perennial herb, 40 cm. high or possibly taller, the stems slender,
rather sparsely leafy, whitish-pubescent in 2 lines; leaves rather thick, on a stout
petiole 3-5 mm. long, almost linear, gradually long-attenuate from a truncate or
subhastate base 4-12 mm. broad, 5-17 cm. long, glabrous or very sparsely puberu-
lent, 1-nerved, glaucescent; inflorescences 2-4 at the apex of the stem or from the
uppermost leaf axils, on slender peduncles 1.5-3.5 cm. long, about 12-flowered, the
pedicels slender, 10-14 mm. long, grayish-puberulent; calyx lobes oblong, acute,
2 mm. long or shorter; puberulent; corolla apparently pale green or whitish, 4.5 mm.
long, glabrous, the lobes broad, obtuse, reflexed; stamen column thick, 1 mm. high;
hoods of the corona ovate, obtuse, contracted and short-unguiculate at the base,
the horns exserted; anthers 2.3 mm. long, the apex white and scarious.
The species is apparently not an uncommon one, usually in sa-
vannas, from Mexico to Colombia but is often difficult to find.
BLEPHARODON Decaisne
Scandent perennial plants, herbaceous or suffrutescent; leaves opposite, often
subcoriaceous; inflorescence cymose or subumbellate, few-many-flowered, borne
on a peduncle from the axils of leaves; flowers medium-sized to small; calyx 5-lobed
and with 5 glands within at the base; corolla valvate, 5-lobate nearly to the base,
the lobes ciliate; corona with 5 scales affixed to the stamen tube, free at the apex,
erect, concave, cymbiform or cucullate; stamens affixed to the base of the corolla,
the filaments connate into a short tube; anthers terminated by an inflexed mem-
brane, the basal auricles elongated; pollinia solitary in each cell, ovoid, pendulous;
stigma depressed or umbonate at the apex; follicles smooth.
FIG. 118. Blepharodon mucronatum. A, habit, natural size; B, corolla and
corona, X 3; C, calyx showing dactyliform squamellae and pistils, X 10; D, opened
capsule, natural size.
417
418 FIELDIANA: BOTANY, VOLUME 24
Apparently a single species in North America which may be con-
sidered typical of the genus. There are some 20 more described
from South America. Decaisne's generic description seems to be
based on a mixture of two genera. We accept the name based upon
B. mucronatum.
Blepharodon mucronatum (Schlecht.) Dene, in DC. Prodr. 8:
603. 1844. Astephanus mucronatus Schlecht. Linnaea 8: 518. 1833.
Bejuco de pescado.
Moist to wet or dry thickets, or in mixed or pine-oak forest, often
in second growth, 1,450 m. or lower; Alta Verapaz; Izabal; Chiqui-
mula; Santa Rosa; Guatemala; Suchitepe"quez; Quezaltenango; Hue-
huetenango; probably to be expected in most departments. Mexico;
British Honduras to El Salvador and Panama; South America.
A large or small, slender, branched vine, glabrous throughout; leaves subcoria-
ceous, on short or rather long, slender petioles, lance-oblong to elliptic, 3-7 cm.
long, cuspidate-acuminate, rounded or obtuse at the base, pale beneath and with
rather conspicuous but very slender nerves; umbels axillary, on long slender pe-
duncles, shorter than the leaves, few-flowered, the pedicels filiform, 1-2.5 cm. long;
calyx lobes oblong-ovate, obtuse, about 1.5 mm. long, pale-marginate; corolla
greenish white, 1 cm. broad, the lobes lance-oblong, obtuse, densely short-villous
within, along and on the margins; lobes of the corona oval, obtuse, shorter than the
gynostegium; follicles glabrous, about 9 cm. long and 2 cm. broad, smooth.
Sometimes called "tietie" in British Honduras; "cancelxiu," "hul-
kin-xiu" (Yucatan, Maya). The flowers have an odor suggestive of
lemon.
CRYPTOSTEGIA GRANDIFLORA (Roxb.) R. Br., a rubber vine, native
of India, is planted rarely in Guatemala for ornament. It is a large
woody vine, glabrous or nearly so, with petiolate, oval to elliptic-
ovate, somewhat coriaceous leaves 5-10 cm. long, and cymose, pink
or purplish flowers 5-7 cm. long. The latex yields a considerable
amount of rubber, and the vine has been exploited in the past for this
product. The plant is at home in the wet-dry regions of Central
America and produces follicles and viable seeds in abundance. It
should perhaps not be used as an ornamental, or otherwise, because
of its potential as a weed. Although we have not yet seen it natural-
ized in Central America, Drouet and Richards, in 1939, report their
number 3838 as being established on lower slopes of mountains just
east of Guaymas, Sonora, Mexico. L. H. MacDaniels number 8 from
Mazatlan, Sinaloa, was taken by the collector as a native plant.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 419
The plant is reported poisonous to livestock.
CYNANCHUM L.
Plants slender, usually scandent and herbaceous, sometimes suffrutescent,
glabrous or pubescent; leaves opposite, coriaceous to membranaceous, often very
small and narrow, acute to cordate at the base; flowers whitish or yellowish, usually
very small, the inflorescences umbelliform or racemiform, sessile or pedunculate;
calyx small, deeply 5-lobate, 5-glandular within at the base or eglandular, the lobes
most often obtuse; corolla campanulate to urceolate or subrotate, 5-lobate, the
lobes valvate, often villous within; corona of 5 scales, these distinct or connate,
laminate to filiform, sometimes variously compound or with internal processes,
rarely wholly lacking; stamens inserted near the base or middle of the corolla tube,
the filaments connate into a short or elongate tube; anthers terminated by an in-
flexed membrane; pollinia solitary in each cell, ovoid or oblong, often compressed,
pendulous; stigma flat at the apex or apiculate; follicles terete, acuminate, usually
slender, smooth; seeds bearing a coma of silky white hairs.
A very large genus, with perhaps 200 species, widely distributed
in both hemispheres, chiefly in tropical or warm regions. There are
four or five species besides those listed here that occur in southern
Central America. As here treated, the group includes plants that
have been cited previously under the names Metastelma, Roulinia,
Rouliniella, Enslenia, Vincetoxicum, Ditassa and various others.
The species referred here to the genus Cynanchum certainly are
all closely related, with the exception of C. rensonii. The flowers are
small and quite as complex as those of the larger flowered Asclepiads.
To this is due the confusion which has existed in the genus. The
genus, however, is not impossible as Woodson once hinted that it
might be for there are good characters that may be seen with a micro-
scope. The genus in North America offers a very nice doctorate
problem.
Inflorescences subracemose, with evidently elongated nodes; leaves broadly ovate,
cordate at the base, mostly 2-5 cm. broad (Cynanchum subg. Mellichampia).
C. rensonii.
Inflorescence subumbellate, the internodes suppressed or very short; leaves linear
to oval, acute to rounded to the base, usually less than 2 cm. broad (Cynan-
chum subg. Metastelma).
Leaves densely pubescent on both surfaces C. trichophyllum.
Leaves glabrous on both surfaces.
Lobes of the corona as broad or broader than long; flowers red C. rubens.
Lobes of the corona longer than broad; flowers not red.
Inflorescences sessile, not pedunculate C. miserum.
Inflorescence not sessile, pedunculate.
420 FIELDIANA: BOTANY, VOLUME 24
Lowest veins of the leaves subparallel to the margin of the leaves.
C. sepium.
Lowest veins of the leaves not parallel to the margins or the veins obsolete.
Lateral veins of the leaves obsolete; leaves mostly linear.
Lobes of the corona oblong and retuse C. palustre.
Lobes of the corona linear, not retuse C. chiapense.
Lateral veins of the leaves evident; leaves mostly broader than linear.
Inner surface of the corolla lobes nearly glabrous or obscurely puberu-
lent; peduncles of inflorescences 1.5-2 cm. long; inflorescence
few-flowered C. woodsonianum.
Inner surface of the corolla lobes densely and obviously pubescent.
Lobes of the corona as long as the corolla or nearly so ; inner face of
corolla evenly pubescent; leaf blades usually sharply reflexed
from apex of petiole; British Honduras endemic.
C. stenomeres.
Lobes of the corona much shorter than the corolla; inner face of
the corolla lobes densely and often retorsely white barbellate;
leaf blades usually not sharply reflexed from apex of petiole;
common species in Mexico, Guatemala and British Honduras.
C. schlechtendalii.
Cynanchum chiapense (Gray) Standl. & Steyerm. Field Mus.
Bot. 23: 226. 1947. Metastelma chiapense Gray, Proc. Am. Acad. 21 :
397. 1886. Metastelma collinum Blake, Contr. U. S. Nat. Herb. 24:
19. 1922 (type from Izabal, Blake 7793}. Cynanchum collinum
Standl. & Steyerm. I.e.
Twining on shrubs or grasses or rocky and brushy hillsides, 600-
1,400 m. or higher; Izabal; Pete"n; Alta Verapaz; Quezaltenango;
Huehuetenango; Jutiapa; Chiquimula; Jalapa. Mexico, the type
from Chiapas; El Salvador; Honduras.
Plants slender, often forming dense tangles of stems, sometimes low and sub-
erect, herbaceous or often woody below, the stems bifariously puberulent or almost
glabrous, subterete, often with greatly elongate internodes; leaves short-petiolate,
linear or nearly so, 1-5 cm. long, usually about 2 mm. broad, thick and firm, often
revolute-margined, 1-nerved, acute, glabrous or ciliate; umbels sessile or on very
short peduncles, mostly 3-6-flowered, the pedicels short, puberulent or glabrous;
flowers white or greenish white, 2.5 mm. long, glabrous outside; calyx lobes
ovate, obtuse; corolla in bud broadly ovoid or ovoid-globose, the lobes ovate, sub-
acute, puberulent and retrorse-barbellate within, erect or ascending; gynostegium
subsessile; scales of the corona narrowly linear, less than one-third as long as the
corolla lobes, equalling the pyramidal apex of the stigma.
This species has consistently linear leaves without visible lateral
nerves. The stems may be quite pubescent (rarely) or more often
pubescent in lines, the leaves are often obscurely ciliolate.
Cynanchum miserum L. Wms. Fieldiana, Bot. 32: 38. 1968.
Barba de leon.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 421
Vine twining over shrubs or trees, at 1,500-2,500 m. Alta Vera-
paz; Huehuetenango; possibly El Progreso. Mexico (Chiapas).
Twining and often reed-like herbaceous or suffrutescent vines. Stems slender,
striated at least when dry, internodes 2-4 cm. long, sparsely puberulent to glabrous
and somewhat vernicose, mostly 0.5-1 mm. in diameter; leaves sparse, soon decid-
uous, linear or linear-lanceolate, acute-apiculate to acuminate, with 2-3 pairs of
obscure lateral nerves, the lowest pair subparallel to the margin, glabrous or dor-
sally the mid-nerve puberulent, about 1-3 cm. long and 0.1-0.4 cm. broad; inflores-
cence sessile (not pedunculate), few-flowered (1-5) fascicles at leafless nodes, pedi-
cels of the flowers mostly about 2 mm. long, glabrous or puberulous; calyx to about
0.5 mm. long, the lobes lanceolate-triangular, acute, puberulous; corolla campanu-
late, glabrous, about 1 mm. long, the lobes subtriangular, obtuse, about 0.3 mm.
long; corona lobes lanceolate, obtuse, about 0.3 mm. long, overtopping the gyno-
stegium; gynostegium stipitate, about 0.5 mm. long; capsule slender, 4-5 cm. long.
Among the species in this flora this may be distinguished by the
sessile fascicles of flowers, the very small glabrous flowers in which
the corolla is divided only about a third its length. The species has
relatively few leaves and these soon fall away.
Cynanchum palustre (Pursh) Heller, Cat. N. Am. PL 6. 1898.
Ceropegia palustris Pursh, Fl. Am. Sept. 1: 184. 1814. Vincetoxicum
palustre Gray, Syn. Fl. N. Am. 21: 102. 1878.
A vine in salt marshes. Manatee Lagoon, British Honduras,
Peck 405. Southeastern United States. West Indies.
Slender vine to about a meter long, stems glabrous. Leaves linear, acute or
acuminate, lateral veins none, to about 45 mm. long and 1.5 mm. broad; inflores-
cence axillary, subumbellate, few-many-flowered, peduncles 2-3 cm. long, pedicels
about 0.5 cm. long; calyx lobes linear-lanceolate, acute, ciliolate, about 2 mm. long
and 0.7 mm. broad; corolla green, divided almost to the base, tube about 0.5 mm.
long, lobes lanceolate or linear-lanceolate, acuminate, 4-5 mm. long and about
1.5 mm. broad; corona lobes oblong, retuse, about 2 mm. long, slightly exceeding
the gynostegium.
The description is from the specimen cited, the only one known
from Central America. It does not fit well into the genus Cynanchum.
Cynanchum rensonii (Pittier) Woodson, Ann. Mo. Bot. Gard.
28: 210. 1941. Roulinia rensonii Pittier, Contr. U. S. Nat. Herb. 13:
101, /. 8. 1910. Cynanchum guatemalense Dugand, Caldesia 9, No. 45:
412. 1966. Cuchamper.
Moist or wet thickets or often in roadside hedges, 1,400 m. or
lower; Zacapa; Chiquimula; Santa Rosa; Guatemala; Suchitepe"quez.
El Salvador; Honduras.
A slender vine, glabrous almost throughout, the young stems sometimes hilari-
ously puberulent, the petioles and inflorescence sparsely puberulent or glabrous;
422 FIELDIANA: BOTANY, VOLUME 24
leaves slender-petiolate, membranaceous, ovate-cordate or deltoid-cordate, 4-7 cm.
long, 1.5-5.5 cm. broad, shortly cuspidate-acuminate, shallowly or deeply cordate
at the base, somewhat paler beneath; inflorescences axillary, racemose, usually
much shorter than the leaves, the peduncles 3 cm. long or shorter, the pedicels
2.5-3.5 mm. long, the flowers yellowish white, scarcely 3 mm. long; calyx glabrous
outside, the lobes oblong-elliptic, obtuse; corolla globose in bud and rounded at the
apex, glabrous, the lobes obtuse or rounded at the apex; lobes of the corona ob-
scurely hastate- trilobate, exceeding the gynostegium; follicles about 10 cm. long
and 2-3 cm. thick, attenuate to the apex, terete, smooth, glabrous.
Sometimes called "champer" in El Salvador. The young fruits
often are used as a vegetable. The status of this species is very un-
certain, and it is quite probable that an older name may be found for
it. It has been reported from Guatemala as Enslenia ligulata Benth.
and Roulinia unifaria Engelm.
Cynanchum rubens L. Wms. Fieldiana, Bot. 32: 39. 1968.
Known only from the type collected in cypress forests, Santa
Elena, Chimaltenango, Skutch 503. Endemic.
Twining vines to a meter or perhaps longer. Stems slender, crisped pubescent
in lines or glabrous, 1-1.5 mm. in diameter; leaves linear-lanceolate or lanceolate,
acuminate, obscurely ciliolate, lateral nerves 4-5 pairs, obscure, 2-5 cm. long and
0.4-0.8 cm. broad, petiole short, puberulent, 1-5 mm. long; inflorescence axillary,
subumbellate, mostly 5-8-flowered, the peduncle short, about 2-3 mm. long, the
pedicels of the flowers 2-3 mm. long, obscurely puberulent; flowers maroon; calyx
lobes ovate-lanceolate, acute, puberulent, about 1 mm. long and half as broad;
corolla rotate, completely glabrous, about 2 mm. long, deeply lobate, the lobes ob-
long-ovate, acute, about 1.3 mm. long and 1 mm. broad; corona fleshy, 5-lobate,
the lobes alternate with the corolla lobes, subcochleate, rounded, about 0.5 mm.
high; gynostegium about 1 mm. high.
Distinctive among all the species of Central America and Mexico
by the broad rounded lobes of the corona and the maroon-colored
flowers with glabrous corollas.
Cynanchum schlechtendalii (Dene.) Standl. & Steyerm. Field
Mus. Bot. 23: 226. 1947; L. Wms. Fieldiana, Bot. 32: 40. 1968.
Metastelma schlechtendalii Decaisne in A. DC. Prodr. 8: 513. 1844.
M. decipiens Pittier, Contr. U. S. Nat. Herb. 13: 98, fig. 4. 1910
(type from San Miguel Uspantan, Heyde & Lux 3060).
Moist or dry thickets, often in rocky places, 2,000 m. or less;
Pete"n; Alta Verapaz; Quezaltenango; Solola; Guatemala; Santa Rosa;
Jutiapa; Zacapa. Mexico; British Honduras.
Usually a small vine, the stems sometimes elongated, much branched and form-
ing mats over other plants; bifariously puberulent or glabrous; leaves short-petio-
STANDEE Y AND WILLIAMS: FLORA OF GUATEMALA 423
FIG. 119. Cynanchum schlechtendahlii. A, habit, natural size; B, flower, X 10;
C, calyx and gynostegium, X 10.
late, lanceolate to ovate-lanceolate, about 2-4 cm. long and 1-1.5 cm. broad,
rounded to subacute or short-acuminate at the apex and usually mucronate, rounded
at the base, glabrous or nearly so, paler beneath, penninerved; inflorescences usu-
ally numerous, umbelliform, laxly few-flowered, puberulent, on relatively short
peduncles, the flowers on short slender pedicels, white or whitish, about 2-3 mm.
long; calyx short, the lobes oblong or ovate-oblong, obtuse, puberulent or glabrous;
corolla glabrous outside, oblong or obovoid in bud, obtuse or somewhat pointed
at the apex, the lobes oblong-lanceolate, subacute, erect or ascending, densely white
barbellate within; gynostegium stipitate, the stipe about equalling the anthers;
corona scales ligulate or linear-subulate, exceeding the gynostegium; follicles slen-
der, about 5.5 cm. long and 7 mm. thick, narrowly long-acuminate, smooth, glabrous.
This is the commonest species in south Mexico and Guatemala.
It seems not to extend into Central America further. Cynanchum
424 FIELDIANA: BOTANY, VOLUME 24
chiapense is a closely related species but is usually distinguished by
the narrower leaves which do not have obvious lateral nerves.
Cynanchum sepium (Dene.) Standl. Contr. U. S. Nat. Herb.
23: 1177. 1924; L. Wms. Fieldiana, Bot. 32: 40. 1968. Vincetoxicum
sepium Dene, in DC. Prodr. 8: 526. 1844. Bejuco de San Julian.
Twining over herbs or trees in thickets. Quezaltenango; Chimal-
tenango. Mexico.
A small vine, usually herbaceous but possibly sometimes woody, glabrous or
with sparse pubescence on petioles and new growth; leaves on short slender petioles
3-5 mm. long, the blade lanceolate or narrowly lanceolate, acuminate, 2.5-6 cm.
long and 0.4-2 cm. broad, glabrous or nearly so, the lowest veins of the leaves
nearly parallel to the margin, only 3-5 pairs; inflorescence umbellate, axillary, the
peduncle 1 cm. long or less, with ten or fewer flowers; flowers greenish-white, 1.5-
2 mm. long; calyx lobes lance-ovate, obtuse or acute, puberulent, about 0.5 mm.
long; corolla lobes ovate, acute, glabrous outside, inner face except the margins
pulvinate, about 1 mm. long; follicles not known.
This is the only species in Guatemala in which the lowest veins
on the leaves follow the leaf margin, there are relatively few lateral
veins. This name has been used for a wide variety of species of
Cynanchum in Guatemala and elsewhere.
Cynanchum stenomeres Standl. & Steyerm. Field Mus. Bot.
23:224. 1947.
British Honduras, at or little above sea level, in open places or in
thickets, often or usually in open pine forest; endemic; type from
All Pines, W. A. Schipp 674-
A very slender vine, herbaceous or suffrutescent, the stems sometimes 2 m.
long, obscurely and bifariously puberulent or glabrous, sometimes purplish; leaves
thick-membranaceous, on slender petioles 4-5 mm. long, penninerved, glabrous
or sparsely and minutely puberulent on the veins, linear-lanceolate, 3-6 cm. long,
4-7 mm. broad, narrowly long-attenuate, obtuse or acute at the base, deep green
above, the costa and nerves often shallowly impressed, paler beneath, the costa
prominent, the lateral nerves obscured, 10 or more pairs; inflorescences umbelli-
form, about 10-flowered, on slender peduncles 8 mm. long or shorter, the pedicels
glabrous or sparsely puberulent, 2.5-4 mm. long; flowers white, fragrant, 4 mm.
long; calyx lobes minute, ovate-oblong, obtuse, sparsely and minutely puberulent
or glabrous; corolla in bud oblong-ovoid or oblong, obtuse, deeply lobate, the lobes
lance-oblong or oblong-linear, usually somewhat narrowed near the base, glabrous
outside, densely white-pilosulous within; gynostegium long-stipitate, the stipe more
than twice as long as the calyx; lobes of the corona with a small ovoid base, atten-
uate into a long hair-like tip, almost equalling the corolla lobes; stigma short-
rostrate, the beak little longer than the appendages of the anthers.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 425
The leaf blades are sharply reflexed from the apex of the petioles
and this shows even in many herbarium specimens. The lobes of the
corona are as long as the corolla or nearly so, the only species in this
flora with such long corona lobes.
Cynanchum trichophyllum L. Wms. Fieldiana, Bot. 32: 41.
1968. Pak-chan.
Moist or dry thickets or forests, 800-1,200 m.; Huehuetenango.
Mexico; Honduras.
Slender, often twining plants forming dense mats over trees and shrubs, stems
to several meters long and mostly about 1 mm. in diameter, herbaceous or at most
suffrutescent, entire plant soft pilose-pubescent; leaves oblong-lanceolate to ovate-
lanceolate, acuminate, apiculate, obtuse or rounded at the base, lateral nerves evi-
dent, sparsely or densely pubescent on both surfaces, about 1-3 cm. long and 0.5-
1 cm. broad, petioles slender, 2-4 mm. long; inflorescences abundant, subumbellate,
usually borne in alternate axils of each succeeding pair of leaves, sometimes in the
axils of both leaves of a pair, each bearing 4-10 flowers, from shorter to about as
long as the subtending leaf or rarely longer, peduncles 2-10 mm. long, pedicels very
slender, mostly 2-3 mm. long; flowers white or cream color; calyx deeply 5-lobed,
pilose outside, the lobes about 0.5-0.7 mm. long, lanceolate-oblong, obtuse; corolla
subcampanulate, divided almost to the base, 2.5-3 mm. long, the lobes 2-2.5 mm.
long and 0.6-0.8 mm. broad, linear-oblong or linear-lanceolate, obtuse, tip reflexed,
glabrous outside, prominently barbellate at the tip inside with two lateral lines of
pubescence extending to the base of the lobe; corona about 1 mm. long or less, the
lobes linear-lanceolate, slightly longer than the gynostegium; capsule 2.5-5 cm.
long, rostrate, densely pubescent.
Known from Guatemala from a single specimen. Abundant in
Central Honduras.
Cynanchum woodsonianum L. Wms. Fieldiana, Bot. 32: 42.
1968. Metastelma pedunculare Decaisne in DC. Prodr. 8: 514. 1844,
non Cynanchum pedunculare Lam., 1786.
Twining over shrubs or on trees, 1,500-2,000 m.; Guatemala (type
from Cuesta de Pinula, Hartweg 601); Sacatepe"quez ; Chiquimula.
Endemic.
Rampant or twining herbs or subshrubs, slender, the stems glabrous. Leaves
elliptic-lanceolate or elliptic-oblong, acute or short-acuminate, glabrous, about 1.5-
5 cm. long and 0.5-1.7 cm. broad, lateral nerves 8-10 pairs and inconspicuous,
petiole slender, puberulent or glabrous, 3-6 mm. long; inflorescence subumbellate,
few-flowered, borne on relatively long peduncles, 1.5-2 cm. long, puberulent;
flowers white or greenish-white; calyx lobes subtriangular, acute or obtuse, fleshy,
glabrous, about 1 mm. long; corolla rotate, the lobes ligulate-lanceolate, acute, the
tips reflexed in natural position, obscurely puberulent within but often appearing
glabrous, about 2 mm. long and 0.8 mm. broad, the corolla tube about 1 mm. long.
426 FIELDIANA: BOTANY, VOLUME 24
This species is easily distinguished by the long pedunculate, few-
flowered inflorescence.
FISCHERIA De Candolle
Herbaceous or suffrutescent vines, often large, setose or villous; leaves oppo-
site, usually large and long-petiolate, membranaceous; flowers medium-sized, um-
bellate or short-racemose, axillary, long-pedunculate, the inflorescence usually
blackening when dried; calyx 5-parted, 5-glandular within at the base, the lobes
narrow; corolla subrotate, deeply 5-lobate, the lobes broad, contorted; outer corona
annular, thick-carnose, adnate to the base of the corolla, the outer margin some-
what 5-sinuate; hoods of the corona 5, thick-carnose, cbtuse; stamens inserted near
the base of the corolla, the filaments connate into a short tube; anthers terminated
by an inflexed, membranaceous or thickened appendage; pollinia solitary in each
cell, oblong or subobovate, subcompressed, the caudicle short, pendulous or oblique;
stigma depressed at the apex, 5-angulate; follicles thick and hard, the seeds connate.
There are perhaps not more than a dozen species of Fischeria in
the American tropics. The genus is doubtfully distinct from Mate-
lea. One other species is known in Central America.
Fischeria martiana Dene, in DC. Prodr. 8: 601. 1844. F. mar-
tiana var. funebris Donn.-Sm. Bot. Gaz. 24: 398. 1897. F. funebris
Blake, Journ. Wash. Acad. Sci. 14: 293. 1924. F. briquetiana Standl.
Field Mus. Bot. 11: 139. 1932 (type from British Honduras, Schipp
962).
Thickets in the lowlands, 700 m. or less; Pete*n. British Hondu-
ras; Mexico; Costa Rica; Panama; probably in all the Atlantic low
lands of Central America. South America to Brazil.
A suffrutescent vine often to 10 m. long and 2.5 cm. in diameter, the young
stems densely puberulent and with sparse to dense spreading hirsute pubescence,
petioles to 7 cm. long or shorter, puberulent and hirsute; leaf blades broadly elliptic
to ovate, 7-22 cm. long, 3.5-15 cm. broad, obtuse or rounded at the apex and short-
caudate, shallowly cordate at the base, with rounded lobes, rough-hirtellous or hir-
sute-pilose above, almost concolorous beneath, densely velutinous-pilosulous or
hirsute-pilose, the hairs often longer on the costa beneath; peduncles 4.5-10 cm.
long, the umbels many-flowered, the slender pedicels 12-35 mm. long, viscid-
puberulent and sparsely hirsute; buds ovoid-globose, very obtuse; calyx lobes lance-
linear, long attenuate, 7-15 mm. long, slightly exceeding the corolla, viscid-puberu-
lent and sparsely hirsute; corolla 12-18 mm. broad, deeply 5-lobate, light green
and yellow, the lobes ovate to ovate-oblong, obtuse to subacute, crispate, densely
hirtellous on both surfaces; outer corona carnose, entire, half as long as the
gynostegium; hoods of the corona carnose, obtuse, longer than the gynnotegium.
GONOLOBUS Michaux
Plants herbaceous or more or less woody, usually scandent, variously pubes-
cent or glabrate; leaves opposite, peticlate, generally cordate; flowers rather large
FIG. 120. Fischeria martiana. A, habit, X
C, calyx, corona and pistils, X 3.
, corolla from above, X 2;
427
428 FIELDIANA: BOTANY, VOLUME 24
or small, usually in umbelliform cymes, these lax, mostly few-flowered, axillary,
pedunculate or subsessile; calyx 5-lobate, usually 5-glandular within at the base;
corolla tube short or almost none, the limb rotate or somewhat campanulate, the
lobes broad or narrow, dextrorsely contorted; corona adnate to the corolla, various
in form, the outer corona usually annular, often ciliate, a fleshy true corona borne
at the base of the stamen column; stamens inserted on the base of the corolla,
the filaments connate into a very short tube; anthers short and broad, bearing
a more or less conspicuous, fleshy, usually laminate, dorsal appendage, the ap-
pendages variable in form, sometimes reniform, often deeply bilobate or entire,
sometimes trilobate; pollinia solitary in the cells, obovate or oblong, horizontal or
somewhat pendulous; stigma depressed at the apex, conspicuously 5-angulate; fol-
licles usually thick and acuminate, smooth or tuberculate; seeds bearing a copious
coma of long white silky hairs.
There have been about 85 species described from Mexico and
Central America that may reasonably be referred to Gonolobus as
here defined. This and most of the related genera are much in need
of revision and no really good account of them can be prepared until
a revision is written.
The name "cuchamper" is applied in Guatemala and Central
America to the fruits of Gonolobus and related genera. The tender
young fruits are commonly used as vegetables, especially by the
country people and even are to be found in the big markets in Guate-
mala City. "Gatillo" is a name sometimes given in Guatemala to the
pods of these vines, in reference to the abundant "silk" of the pods.
Faucal corona erect and mostly more than 1 mm. high, ciliate at the apex; surface
variously pubescent or glabrous.
Corona deeply 5-lobate, the lobes half the length of the corona and rounded.
G. lasiostemma.
Corona entire, digitate or obscurely lobate, the lobes if present very short.
Corolla lobes pubescent dorsally with a few long, segmented or hispid hairs;
leaves attenuated to truncate or somewhat cordate at the base.
Pubescence of back of corolla lobes soft hispid, the lobes about 5 mm. long
and obtuse; pubescence of the top of the corona continuous.
Leaves deeply cordate; petioles puberulent G. roeanus.
Leaves acute or truncate at the base; petioles spreading pilose pubescent.
G. donnellsmithianus.
Pubescence of back of corolla lobes of long segmented hairs, the lobes about
10 mm. long, acute; corona glabrous.
Faucal corona adorned with flattened hairs and with similar hairs near
the apex of the inner surface; calyx lobes linear-lanceolate.
G. xanthotrichus.
Faucal corona glabrous; calyx lobes linear G. chiapensis.
Corolla lobes glabrous dorsally.
Faucal corona ciliate only at the apex, not pubescent on the sides and base.
G. calycosus.
Faucal corona ciliate at the apex and at the base, the sides sparsely to
densely pilose G. stenosepalus.
Faucal corona not erect, or if erect not pubescent.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 429
Corolla calcarate at the juncture of the lobes; leaves not cordate at the base.
G. longipetiolatus.
Corolla not calcarate at the juncture of the lobes; leaves cordate.
Corolla lobes densely barbate along one margin with long flat hairs.
G. barbatus.
Corolla lobes not barbate, glabrous, short pilose or puberulent.
Lobes of the corolla short-pilose or puberulent within, the hairs of appreciable
length.
Leaves glabrous or glabrate.
Corolla lobes linear, hispidulous in a line within G. cleniophorus.
Corolla lobes oblong or ovate, puberulous within and without . . . . G. salvinii.
Leaves pilose on both surfaces.
Corolla distinctly tubular at the base, the apex rounded in bud.
G. lasiostemma.
Corolla not at all tubular at the base, acute or acuminate in bud.
Inflorescence sessile or nearly so, peduncles mostly less than 1 cm. long.
G. lanugiflorus.
Infloresecnce conspicuously pedunculate, the peduncles 1-3 cm. long.
G. uniflorus.
Lobes of the corolla glabrous within or farinose-puberulent.
Corolla lobes farinose-puberulent within along one side.
Calyx lobes lanceolate or linear-lanceolate; corolla bicolored and reticu-
late-nerved G. uniflorus.
Calyx lobes ovate; corolla concolorous, not reticulate-nerved.
G. leianthus.
Corolla lobes glabrous within unless at the very base.
Calyx lobes oval, obtuse or rounded at the apex G. calycosus.
Calyx lobes ovate or lanceolate and acute or acuminate, rarely obtuse.
but then lanceolate.
Corolla in bud lanceolate or narrowly lanceolate, long-tapering at the
apex.
Calyx lobes linear from a somewhat broadened base; corolla about
2.5-3 cm. long G. versicolor.
Calyx lobes lanceolate; corolla 1.5-2 cm. long G. stenanthus.
Corolla in bud subglobose or ovoid, rounded or short acuminate, not
tapering to the apex.
Corolla dark purple, drying black, about 7 mm. long G. niger.
Corolla greenish or yellowish, not drying black, usually larger than
7 mm.
Corona hirtellous or short-pilose G. stenosepalus.
Corona glabrous.
Basal lobes of the leaves strongly incurved G. steyermarkii
Basal lobes of the leaves not at all incurved G. prasinanthus.
Gonolobus barbatus HBK. Nov. Gen. & Sp. PL 3: 209, t. 239.
1819. Vincetoxicum barbatum Standl. Contr. U. S. Nat. Herb. 23:
1192. 1924. Bubsaac (Pet&i, Maya).
Moist or dry thickets, often in rocky places, 200-1,600 m.; re-
ported from Pete"n; El Progreso; Zacapa; Jutiapa; Santa Rosa;
Escuintla; Huehuetenango. Western and southern Mexico; Hon-
duras; El Salvador; Nicaragua.
430 FIELDIANA: BOTANY, VOLUME 24
A small or large vine, herbaceous or usually somewhat lignescent, the stems
pale, finely puberulent, short-pilose, or glabrate; leaves firm-membranaceous, on
long slender petioles, broadly deltoid-cordate or ovate-cordate, mostly 2-5 cm.
long, acuminate, rather shallowly and openly cordate at the base, glabrous; flowers
umbellate, the umbels sessile or short-pedunculate, the pedicels very slender, mostly
2-3 cm. long, puberulent; calyx green, lobate almost to the base, the segments lan-
ceolate, attenuate-acuminate, sparsely puberulent or almost glabrous; corolla
about 1 cm. long, in bud globose and short-cuspidate, glabrous or inconspicuously
puberulent outside, green and yellow, deeply lobate, very densely yellow-barbate
with long hairs in the throat and along one side of the lobes; stigma depressed,
very deeply 5-lobate; follicles ovoid, 8 12 cm. long, glabrous, with broad longi-
tudinal wings.
Called "Matacoyote" and "cuchamper de zope" in El Salvador;
"xtuchcahoy" (Yucatan, Maya); "cuayote" (Honduras).
Gonolobus calycosus (Donn.-Sm.) Woodson, Ann. Mo. Bot.
Gard. 28: 242. 1941. Fimbristemma calycosa Donn.-Sm. Bot. Gaz.
16: 196, t. 16. 1891.
Alta Verapaz (the type from Chucaneb, 1,800 m., Tuerckheim
1500). Nicaragua.
Leaves long-petiolate, oblong-cordate, 10-25 cm. long, acuminate, deeply cor-
date at the base, more or less pubescent; peduncles shorter than the petioles, 1-2
cm. long, the 6-8 pedicels longer than the peduncle, the flowers 2.5 cm. broad;
calyx deeply 5-lobate, the segments oval, pale, obtuse or rounded at the apex,
pubescent outside; corolla 5-lobate almost to the base, the lobes dextrorsely con-
volute, deep yellow, very obtuse, oblong; outer corona elobate, densely long-fim-
bricate; lobes of the inner corona ovate, equalling the fimbriae, bidentate, naked;
anthers appendiculate below with a 2-cornute wing.
Gonolobus chiapensis (Brandegee) Woodson, Ann. Mo. Bot.
Gard. 28: 242. 1941. Vincetoxicum chiapense Brandegee, Univ. Calif.
Publ. Bot. 6: 190. 1915. Matelea chiapensis Shinners, Sida 1: 366.
1964.
Moist mixed forest, 1,300-1,500 m.; San Marcos (Volcan de Taju-
mulco). Mexico (type from Cerro del Boqueron, Chiapas). Nicaragua.
A somewhat woody vine, the stems rather densely hirsute with long spreading
brownish multicellular hairs; leaves short-petiolate, lance-oblong or oblanceolate-
oblong, mostly 9-12 cm. long and 3-4.5 cm. broad, long acuminate, subcordate at
the base, thin, copiously hirsute on both surfaces; inflorescences axillary, subum-
bellate, sessile or on very short peduncles, 3-4-flowered, the pedicels 10 mm. long
or shorter; calyx lobes lanceolate, hirsute, 4 mm. long; corolla rotate, dull green,
pilose outside, 5 mm. long, the lobes broadly lanceolate; corona simple, annular,
the margin undulate; disk of the stigma pentagonal, dark purple, flat.
FIG. 121. Gonolobus barbatus. A, habit, natural size; B, part of the calyx, disk
and pistils; C, orifice of the corolla with one corolla lobe complete, X 5; D, winged
fruit, X K; E, cross-section of fruit, X 3^>; F, pollinia, X 7)^.
431
432 FIELDIANA: BOTANY, VOLUME 24
Gonolobus cteniophorus (Blake) Woodson, Ann. Mo. Bot.
Gard. 28: 243. 1941. Vincetoxicum cteniophorum Blake, Contr. Gray
Herb. 52: 84. 1917. V. tortum Brandegee, Univ. Calif. Publ. Bot. 10:
414. 1924. V. lundellii Standl. Field Mus. Bot. 8: 148. 1930 (type
from Honey Camp, British Honduras, Lundell 540). Matelea ctenio-
phora Shinners, Sida 1 : 366. 1964.
Moist or wet thickets, sometimes in wooded swamps, 150 m. or
less; Pete"n; Izabal. Southern Mexico; British Honduras (type from
Toledo, Peck 821).
A small or rather large, slender vine, herbaceous or suffrutescent, the stems
almost glabrous, sparsely appressed-pilosulous at the nodes; leaves membrana-
ceous, on long slender petioles, oblong-ovate to deltoid-cordate, mostly 5-9 cm.
long, acuminate or long-acuminate, deeply cordate at the base, glabrous; peduncles
axillary, 2-3 cm. long, glabrous, the flowers few, umbellate, the pedicels slender,
unequal, 1-2.5 cm. long; calyx lobes ovate or lance-ovate, acuminate, ciliolate,
3 mm. long; corolla rotate, 3.5 cm. broad, lobate almost to the base, the lobes
linear or nearly so, glabrous outside, hsipidulous in a line along the middle inside;
outer corona short, densely ciliate, the inner corona short, carnose, glabrous,
broadly 5-lobate; fruit narrowly ovoid, glabrous, smooth or somewhat white-
tuberculate, about 6.5 cm. long and 2.5 cm. wide, acuminate.
Gonolobus dasystephanus (Blake) Woodson, Ann. Mo. Bot.
Gard. 28: 243. 1941. Vincetoxicum dasystephanum Blake, Contr.
Gray Herb. 52: 84. 1917.
Known only from the type, British Honduras, in forest and
thickets, Manatee Lagoon, M. E. Peck 323.
A rather large vine, the stems slender, hispid and hispidulous, the hairs sub-
retrorse, bulbous-thickened at the base; leaves short-petiolate, oblong-oval, 3.5-
5 cm. long, 1.2-2 cm. broad, short acuminate, obtuse or rounded at the base, green
above, slightly paler beneath, rather densely hispid-pilose on both surfaces with
fulvous hairs; peduncles incurved-hispidulous, 7-10 mm. long, the umbels 6-9-
flowered, the pedicels 1.5 cm. long; calyx lobes ovate, obtuse, hispidulous, 2 mm.
long; corolla rotate, green when dry, 5-lobate almost to the base, 12 mm. broad,
the lobes oval-ovate, obtuse, hispidulous outside, glabrous within; outer corona
annular, erect, obscurely 5-lobate, densely pilose-ciliate; inner corona shorter,
glabrous, carnose, few-crenulate.
Gonolobus donnellsmithianus L. Wms. Fieldiana, Bot. 32:
45, jig. 1968.
Known only from the type, Lacandon, Department of Pete"n,
Contreras 3364..
A herbaceous or suffrutescent vine with stems spreading pilose pubescent.
Leaves elliptic or elliptic-oblong, acuminate, acute to obscurely truncate at the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 433
FIG. 122. Gonclobus donnellsmithianus. A, habit, X 1A', B, flower in natural
position showing petals, corona and gynostegium, X 4; C, calyx and ovaries
(stigma removed), X 5.
base, sparsely fulvous pilose on both surfaces, the blade (2-)6-10 cm. long and
(l-)2.5-4 cm. broad, the petioles of a pair slightly unequal, spreading pilose-pubes-
cent, about 4-14 mm. long; inflorescence a subumbellate cyme 6-10-flowered, the
peduncle about 1 cm. long, the pedicels to 1.5 cm. long, puberulent; calyx pubescent
dorsally, divided nearly to the base, the lobes triangular-lanceolate or lanceolate,
acute, 2-3 mm. long; corolla sparsely pubescent dorsally, deeply lobate, the lobes
linear-oblong, obtuse, cucullate, glabrous within, about 5 mm. long and 2 mm.
broad; faucal corona about 1 mm. high, obscurely lobate and ciliate at the apex,
sides and base glabrous; staminal appendages furcate, the branches slender and
arcuate; fruit unknown.
This species is allied to G. xanthotrichus Brandegee but is easily
distinguished by floral detail.
Gonolobus lanugiflorus Woodson, Ann. Mo. Bot. Gard. 28:
282. 1941.
Moist or wet, mixed forest, 2,300-2,800 m.; known only from the
vicinity of the type locality, Volcan de Tajumulco, near Tajumulco,
the type being Steyermark 36906.
Plants scandent, herbaceous or lignescent, densely fulvous-hispid or hirsute on
the stems, leaves, and inflorescence, the hairs slightly harsh to the touch, the stems
434 FIELDIANA: BOTANY, VOLUME 24
stout; leaves on petioles 2-3 cm. long, membranaceous, ovate-cordate or broadly
oblong-ovate, 9-14 cm. long, 4.5-7.5 cm. broad, rather abruptly and narrowly long-
acuminate, deeply cordate at the base; inflorescence umbelliform, subsessile, many-
flowered, the flowers greenish, on pedicels 1 cm. long or shorter; calyx lobes
lanceolate, 1 cm. long, acuminate; corolla rotate, fulvous-hispidulous inside and
outside, the lobes ovate-lanceolate, 1.5 cm. long, narrowly acuminate; annulus of
the throat conspicuous, minutely fulvous-hispidulous, entire; corona rotate, 5-
parted, glabrous, the margin minutely crenulate; gynostegium subsessile, 4.5 mm.
in diameter.
Gonolobus lasiostemma (Hemsl.) Woodson, Ann. Mo. Bot.
Gard. 28: 243. 1941. Lachnostoma lasiostemma Hemsl. Biol. Centr.
Am. Bot. 2: 335. 1882. Gatos.
Moist or dry, often rocky thickets or in hedges, 1,200-2,300 m. ; en-
demic; type collected in Guatemala by Skinner, the locality unknown;
Santa Rosa; Guatemala; Sacatepe"quez ; Solola; Huehuetenango.
A rather large and coarse vine, herbaceous or somewhat woody, the stems
densely short-hirsute with fulvous retrorse hairs; leaves long-petiolate, membrana-
ceous, broadly oblong-ovate, mostly 6-13 cm. long and 3-7 cm. broad, acuminate
or acute, sometimes abruptly acuminate, rather shallowly and openly cordate,
densely velutinous-pilose with short spreading hairs on both surfaces, little paler
beneath; inflorescences lax and few-flowered, umbelliform, short-pedunculate, the
pedicels mostly 2 cm. long or shorter, densely pilosulous, the flowers 2-3 cm. broad;
calyx densely pilosulous, deeply 5-lobate, 5-glandular within, the segments linear-
lanceolate, acute, almost equalling the corolla; corolla green, tubular at the base,
more or less hirtellous outside and within, the tube shorter than the lobes, the lobes
narrow, obtuse; corona adnate to the corolla tube, the outer corona of 5 large sub-
galeate scales opposite the corolla lobes, these barbate on all sides, the inner corona
of 5 bifid glabrous scales alternate with the corolla lobes; gynostegium free, the
stigma depressed; ovaries glabrous; young follicles broadly ovoid, acuminate, 6 cm.
long and more than 3 cm. broad (at maturity doubtless much larger).
Gonolobus leianthus Donn.-Sm. Bot. Gaz. 48: 296. 1909.
Vincetoxicum schippii Standl. Field Mus. Bot. 8: 37. 1930 (type from
Middlesex, British Honduras, W. A. Schipp Si 9) . Sucrixcam (Co-
ban, Quecchi).
Moist or wet thickets or open forest, sometimes in roadside hedges,
500-2,400 m., or sometimes at lower elevations; Pete"n; Alta Verapaz;
Sacatepe"quez ; Quezaltenango; Huehuetenango. Mexico, British
Honduras.
A large or small, herbaceous vine, sometimes suffrutescent, rarely trailing over
the ground, the stems, petioles, and inflorescence pilose with rather stiff, spreading
or reflexed, short hairs, or the stems sometimes glabrate and the peduncles and
pedicels glabrous or nearly so; leaves firm-membranaceous, on long slender petioles,
oblong-ovate, mostly 6-12 cm. long and 2.5-5 cm. broad, acuminate, deeply cordate
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 435
at the base, rather densely short-pilose on both surfaces or often glabrate, especially
on the upper surface, slightly paler beneath; peduncles few-flowered, mostly 2-
flowered, 2-4 cm. long or often very short, the pedicels very unequal, mostly 2-5 cm.
long; flowers as much as 5 cm. in diameter but often not more than 3.5 cm.; calyx
lobes ovate or broadly ovate, green or pale, 1.5 cm. long or shorter, acute or acu-
minate; corolla puberulent or almost glabrous outside, papillose-puberulent within
along one side of each lobe, deeply lobate, olive-green within, dull yellow-green
outside, the lobes lance-oblong, obtuse or subacute; corona annular, subentire,
adnate to the very short gynostegium; gynostegium 6 mm. in diameter; follicles
about 13 cm. long and 4.5 cm. thick, longitudinally winged, smooth.
Gonolobus longipetiolatus Woodson, Ann. Mo. Bot. Gard. 28:
282. 1941.
Known only from the type, San Marcos, southern slopes of Volcan
de Tajumulco, above Finca El Porvenir, 1,300-1,500 m., Steyermark
37428.
Plants slender, suffrutescent, the stems sparsely fulvous-hirsute with short
hairs; leaves on long slender petioles, membranaceous, oblong-elliptic or oblong-
ovate, 4.5-10 cm. long, 2-5 cm. broad, narrowly long-acuminate, rounded or trun-
cate at the base, glabrous; petioles 2-5 cm. long, glabrous; inflorescence subrace-
mose, lax, few-flowered, the peduncle 2-2.5 cm. long, the pedicels of about the same
length; calyx lobes narrowly lanceolate, long-acuminate, 7 mm. long, minutely
papillose outside; corolla green, rotate, glabrous, the lobes broadly linear, 9 mm.
long, acuminate, with involute margins, short calcarate at the juncture of corolla
lobes; corona conspicuously 5-lobate, the lobes erect, broadly oblong, 2 mm. long,
truncate or subemarginate at the apex; gynostegium borne on a stipe 2 mm.
long; stigma pentagonal, 3.5 mm. broad, conspicuously rostrate.
This is the only species of Gonolobus which we know that has
obscure calcarate processes at the juncture of lobes of the corolla.
Gonolobus niger (Cav.) R. Br. Wern. Soc. 1: 35. 1809. Cynan-
chum nigrum Cav. Icon. PI. 2: 45, 1. 159. 1793. Vincetoxicum cavanil-
lesii Standl. Contr. U. S. Nat. Herb. 23: 1188. 1924. Millona.
Dry rocky thickets, about 1,600 m.; Huehuetenango (above San
Ildefonso Ixtakuacan, Steyermark 50678). Southern Mexico.
A slender vine, herbaceous or somewhat woody, the stems minutely puberulent
and short-hirsute with sparse, fulvous, spreading or reflexed hairs, sometimes gla-
brate; leaves firm membranaceous, on long slender petioles, oblong-ovate or del-
toid-ovate, 4-7.5 cm. long, acuminate or cuspidate-acuminate, rather shallowly
and openly cordate at the base and sometimes subhastate, thinly hirtellous or
almost glabrous on the upper surface, paler beneath, puberulent on the veins or
glabrate; inflorescences umbelliform or racemiform, very open and few-flowered,
often equalling the leaves, the peduncles often much longer than the petioles, the
pedicels 1-2.5 cm. long, puberulent or almost glabrous; calyx lobes ovate or ovate-
lanceolate, much shorter than the corolla, acute, greenish, glabrous or nearly so;
436 FIELDIANA: BOTANY, VOLUME 24
corolla deep dark purple, blackish when dried, puberulent outside, glabrous within,
about 6 mm. long, lobate almost to the base, the lobes oblong or ovate, obtuse,
reflexed in an thesis; corona very short and shallowly lobate.
Gonolobus prasinanthus Donn.-Sm. Bot. Gaz. 46: 114. 1908.
Vine in thickets or over shrubs, up to about 800 m. altitude (type:
Cubilquitz, Alta Verapaz, Tuerckheim 8711); Pete"n; Alta Verapaz;
Suchitepe"quez; San Marcos. Southern Mexico. British Honduras.
Vines, herbaceous or suffrutescent, the branches glabrous or pubescent. Leaves
nearly glabrous above, subglabrous or pubescent below, the blades lanceolate-
ovate, cordate, acuminate, 5-13 cm. long and 3-6 cm. broad, the petioles pubes-
cent, long and slender, 4-6 cm. long; inflorescences borne on peduncles 15-25 mm.
long; pedicels 3-6, about 12-20 mm. long; flowers 15-20 mm. across; calyx lobes
lanceolate or linear-lanceolate, acute, obscurely puberulent outside, 5-6 mm. long
and about 1.5 mm. broad; corolla deeply 5-lobed, the tube very short, the lobes
lanceolate or narrower, acuminate, glabrous, 8-12 mm. long and about 4 mm.
broad at the base; corona fleshy, lobate, less than 1 mm. broad; folicle smooth
when immature.
The specimens which we refer to this species have usually been
referred to the Mexican G. fraternus Schlecht. and G. prasinanthus
placed there as a synonym. No authentic material or type photo-
graphs of G. fraternus are available and the description is not ade-
quate for determination of the species.
Gonolobus roeanus L. Wms. Fieldiana, Bot. 32: 47. 1968.
Vine in scrub forest, alt. 1,500 m. Endemic (type Roe, Roe &
Mori 813, Alotenango, Sacatepe"quez) .
Branched vines with sparsely hirsute stems. Leaves long petiolate, the blades
cordate or oblong-cordate, acute or somewhat acuminate, sparsely and obscurely
short pubescent on both surfaces, nerves conspicuous, about 5 pairs with secondary
reticulate nerves, mature leaves 4-6 cm. long and 2-3 cm. broad, petioles slender,
puberulent, 2-4 cm. long; inflorescences subumbellate, axillary, pedunculate, few-
several-flowered, peduncles 1-2 cm. long, pedicels 0.5-1.5 cm. long; flowers pale
brown, small; calyx glabrous or nearly so, divided to the base, lobes linear or linear-
lanceolate, acute, about 5-nerved, 2.5-3 mm. long and 0.5-0.7 mm. broad; corolla
puberulent outside, glabrous within except corona, deeply lobed, subrotate or lobes
reflexed, 10-12 mm. broad, lobes oblong or oblong-ovate, obtuse, reticulate veined,
about 4 mm. long and 2 mm. broad; faucal corona fleshy, erect, margin ciliate,
about 0.5 mm. high; inner corona surrounding gynostegium, carnose, lobate or
somewhat fimbriate, glabrous, shorter than the gynostegium; gynostegium about
1 mm. high; follicles unknown.
Gonolobus salvinii Hemsl. Biol. Centr. Am. Bot. 2: 334. 1882.
Cuchamper.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 437
Moist or dry thickets or forest, sometimes in oak forest, 900-
2,400 m.; Jalapa; Santa Rosa; Escuintla; Quiche"; Quezaltenango
(type from Volcan de Zunil, Salviri). British Honduras; El Salvador;
Honduras.
A small or large vine, the stems pale, puberulent, short-pilose, or almost gla-
brous; leaves firm-membranaceous, on long slender petioles, oblong-ovate or oblong-
deltoid, mostly 5-12 cm. long, acuminate or long-acuminate, rather shallowly and
openly cordate at the base, glabrous or nearly so, pale beneath; peduncles mostly
1 cm. long or shorter, few-flowered, the flowers subumbellate, the pedicels 2 cm.
long or shorter, pubescent; calyx lobes linear or lanceolate, puberulent outside,
acute, 5-6 mm. long; corolla subrotate, about 1.5 cm. broad, green, deeply 5-lobate,
sparsely puberulent outside and within, the lobes oblong or ovate, obtuse or sub-
acute, reflexed in an thesis; outer corona 5-lobate, the lobes bifid; scales of the inner
corona obovate-oblong, recurved at the apex; ovaries glabrous.
Sometimes called "Siguamper" in El Salvador. The tender fruits
are sold there in the markets, being eaten raw when almost mature,
and cooked and eaten at almost all stages of growth.
Gonolobus stenanthus (Standl.) Woodson, Ann. Mo. Bot.
Gard. 28: 243. 1941. Vincetoxicum stenanthum Standl. Field Mus.
Bot. 4: 255. 1929.
Moist or wet thickets, at or little above sea level; British Hon-
duras (type from Tower Hill, J. S. Karling 27). Mexico (Quintana
Roo, Campeche). Atlantic coast of Honduras.
A slender, herbaceous or woody vine, the stems sparsely puberulent, especially
at the nodes, or glabrate; leaves membranaceous, on slender petioles 1-8 cm. long,
oblong-ovate or elliptic-ovate, 4-14 cm. long, 2-8.5 cm. broad, acuminate or
abruptly acuminate, shallowly or deeply and openly cordate, glabrous or some-
times with a few scattered short hairs on the upper surface, paler beneath; inflores-
cences umbelliform, 1-5-flowered, on peduncles 1-2.5 cm. long, the pedicels unequal,
very slender, 1-5 cm. long, sparsely and minutely puberulent or glabrous; calyx
lobes narrowly lanceolate, 1.5 cm. long or shorter, attenuate to the apex, glabrous;
corolla rotate, 5-lobate almost to the base, glabrous within except at the base of
the lobes, green, the lobes almost linear, 15-20 mm. long, 3 mm. broad at the base,
attenuate, spreading; corona short, 4 mm. broad, conspicuously 5-lobate, the lobes
broad, densely hirtellous.
Called "Cuchamper" in Honduras. The young pods there are
boiled or otherwise cooked with sugar, to make sweetmeats or dulces.
Gonolobus stenosepalus (Donn.-Sm.) Woodson, Ann. Mo. Bot.
Gard. 28: 243. 1941. Fimbristemma stenosepala Donn.-Sm. Bot. Gaz.
18: 208. 1893.
438 FIELDIANA: BOTANY, VOLUME 24
Occasionally common in oak forests or ciliar forests (the type from
Santa Rosa, Heyde & Lux 4004)', Huehuetenango; Quiche; Chimal-
tenango. Southern Mexico.
Large herbaceous or suffrutescent vines, hirsute with erect hairs. Leaves ob-
long-lanceolate to ovate, acuminate, truncate subcordate to cordate at the base,
the petioles pubescent with spreading hairs, half or less the length of the blades,
strigillose pubescent on both surfaces to glabrescent above, 4-15 cm. long and 2.5-
7 cm. broad; inflorescences of umbelliform cymes shorter than the leaves, peduncles
mostly 2-4 cm. long, about as long as the longest pedicels, mostly 5-10-flowered;
calyx lobes lanceolate or lanceolate-ovate, puberulent but the margins prominently
ciliolate, mostly 6-8 mm. long and 2-4 mm. broad; corolla green, as much as 3 cm.
broad at anthesis, divided nearly to the base, glabrous outside and glabrous inside
except near the faucal corona, the lobes mostly 8-15 mm. long and 2-4 mm. broad,
oblong-lanceolate, acute, one margin usually obscurely sphacellate; faucal corona
mostly 3-4 mm. high, the margin obscurely lobulate, the surface sparsely or usually
densely pilose; the gynostegium longer than the corona; the follicels about 15 cm.
long, lanceolate-ovoid and arcuate, with prominent thin wings.
This species is easily distinguished from all other species of the
genus in Mexico and Guatemala by the usually abundantly pilose
outer surface of the faucal corona. Assumed to be quite rare, it is
actually quite widespread in the area of south Mexico and Guatemala.
Gonolobus steyermarkii Woodson, Ann. Mo. Bot. Gard. 28:
283. 1941.
Moist or wet thickets or forest, 1,200-1,400 m.; so far as known,
endemic; Suchitepe"quez (Volcan de Zunil); Quezaltenango (Volcan
de Santa Maria); San Marcos (type from Volcan de Tajumulco,
along Rio Negro near Finca La Patria, Steyermark 37661).
A large vine, the stems fulvous-hispidulous with spreading hairs; leaves on
slender petioles 2-6 cm. long, membranaceous, oblong-ovate or elliptic-ovate, 7-
25 cm. long, 2.5-12 cm. broad, narrowly long-acuminate, deeply cordate at the
base and subauriculate, thinly fulvous-hispidulous on both surfaces, slightly paler
beneath; inflorescences racemiform or sometimes corymbiform, few-flowered, the
peduncles 2-4 cm. long, the pedicels 2-3 cm. long, sparsely pilosulous; calyx lobes
oblong-lanceolate or ovate-lanceolate, acuminate, 15 mm. long, 5-7 mm. broad,
folia ceous, glabrous or papillose; corolla rotate, pale green, glabrous, the lobes
broadly oblong-elliptic, ascending, subacute, about 12 mm. long and 5 mm. broad,
with revolute margins; throat of the corolla annulate, the ostiole densely pilose;
corona annular, the margin minutely crenulate, adnate to the corolla, 5-parted;
stigma pentagonal, 4 mm. broad.
This species is closely related to G. calycosus (Donn.-Sm.) Wood-
son, G. stenosepalus (Donn.-Sm.) Woodson and to G. oblongifolius
(Donn.-Sm.) Woodson. They may all eventually prove to be vari-
ants of a single species.
FIG. 123. Gonolobus uniflorus. A, a leaf, natural size; B, flower from above,
X iy2;C, calyx, X 3.
439
440 FIELDIANA: BOTANY, VOLUME 24
Gonolobus uniflorus HBK. Nov. Gen. & Sp. PL 3: 207, t. 228.
1819. Vincetoxicum uniflorum Standl. Contr. U. S. Nat. Herb. 23:
1192. 1924.
Moist thickets and open woods, 1,300-2,000 m.; Alta Verapaz;
Huehuetenango; Quiche"; Santa Rosa. Mexico; British Honduras;
Honduras; Nicaragua.
A small or large vine, the stems slender, densely fulvous-pilose with short,
spreading or reflexed hairs; leaves membranaceous, on long slender petioles, ob-
long-ovate, 5-10 cm. long, acuminate or long-acuminate, deeply cordate at the
base with usually somewhat incurved basal lobes, rough-hispidulous on the upper
surface, softly short-pilose beneath, usually sparsely so; peduncles mostly shorter
than the petioles, 1-few-flowered, the long slender petioles often longer than
the peduncle, densely hispidulous; calyx lobes pale, lanceolate to ovate-lanceo-
late, 1-1.5 cm. long, long-attenuate, pilosulous; corolla green, rotate, 3-5 cm.
broad, sparsely hirtellous outside or glabrous, deeply lobate, the lobes lance-oblong,
papillose-puberulent within along one margin, acuminate, annulate, the annulus
densely hirtellous; follicle ovoid, about 10 cm. long and 4.5 cm. broad, acuminate,
smooth, glabrous, with narrow thick longitudinal wings, the valves hard and lig-
nescent; seeds blackish, about 7 mm. long, flat.
There are a number of names which have been applied to the
material which we believe represents one rather variable species.
The names which have been most commonly used are G. macranthus
Kunze which may be a synonym and G. leianthus Donn.-Sm. used
in error. The reticulate venation in the corolla makes this one of the
more attractive of the Asclepiads in Guatemala.
Gonolobus versicolor Woodson in Standl. & Steyerm. Field
Mus. Bot. 23:80. 1944.
Known only from the type, Huehuetenango, near Maxbal, about
17 miles north of Barillas, 1,500 m., Steyermark 48768.
A large vine, climbing over trees, the stems pilose with spreading hairs; leaves
membranaceous, on petioles 1.5-3.5 cm. long, oblong-ovate, 4.5-8 cm. long, 2-
3.5 cm. broad, acuminate, truncate or subcordate at the base, sparsely pilose above
along the costa, otherwise glabrous, beneath setose-pilose on the veins or glabrate;
inflorescences extra-axillary, few-flowered, the peduncle glabrous, 3-5 cm. long,
the pedicels 2-2.5 cm. long, glabrous; calyx lobes linear-lanceolate, 2 cm. long, long-
acuminate, spreading or reflexed, glabrous, yellowish brown outside near the base,
blackish brown above; corolla rotate, the lobes lance-linear, long-acuminate, 2.5-
3 cm. long, ascending, glabrous outside, brownish yellow or brownish green, within
yellow-brown, pale green, and reddish brown, papillose, the ostiole with a shallowly
5-lobate annulus; corona patelliform, 5 mm. broad, shallowly 5-lobate, the margin
entire or obscurely crenulate, pale green; stigma pentagonal, almost flat or some-
what depressed.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 441
Gonolobus xanthotrichus Brandegee, Zoe 5: 251. 1908. Tri-
chostelma oblongifolius Donn.-Sm. Bot. Gaz. 48: 296. 1909. Vince-
toxcum xanthotrichum Standl. Contr. U. S. Nat. Herb. 23: 1193. 1924.
Gonolobus oblongifolius Woodson, Ann. Mo. Bot. Card. 28: 243. 1941.
Moist or wet forest areas or thickets, 1,000-1,500 m.; Alta Vera-
paz (type Trichostelma oblongifolius Donn.-Sm., Tuerckheim II 1747).
Mexico.
Plants scandent, usually herbaceous, the stems, petioles, leaf blades, and in-
florescence densely hispid-hirsute with long spreading stiff fulvous hairs; leaves
membranaceous or thick-membranaceous, on rather short petioles, oblong or obo-
vate-oblong, mostly 8-11 cm. long, shortly cuspidate-acuminate, rounded or shal-
lowly cordate at the narrow base; inflorescences umbelliform, few-flowered, sessile
or subsessile, the pedicels mostly 2-3 cm. long, hispid; calyx densely hirsute, the
lobes ovate-lanceolate, scarcely half as long as the corolla lobes; corolla rotate,
densely hirsute outside, about 1 cm. long, the lobes triangular-oblong, hirsute with-
in along one margin near the apex; outer corona annular, densely hirsute-barbate,
the inner corona consisting of fleshy quadrate scales; gynostegium 2 mm. high;
stigma pentagonal, umbonate.
EXCLUDED:
GONOLOBUS EDULIS Hemsl. in Godman & Salvin, Biol. Cent. Am.
Bot. 2: 331. 1882. Vincetoxicum edule Standl. Contr. U. S. Nat.
Herb. 23: 1680. 1926.
This species was based by Hemsley on two specimens without in-
dication of which was the type, a not unusual procedure at that time.
One specimen cited is Endres 213 from Costa Rica, the other Fried-
richsthal from Guatemala. Miss S. M. King at Kew has made ana-
lytical drawings from the "syn types" which indicates that they are
the same species and similar to a fairly common Costa Rican species.
Friedrichsthal's collections were made in Costa Rica, Nicaragua and
Guatemala but only the ones with Guatemalan labels have ever been
cited, to the best of my knowledge. It is almost certain that Nica-
raguan and Costa Rican specimens were distributed under a "Guate-
malan" label and Gonolobus edulis is probably one of these.
HOYA CARNOSA (L.) R. Br., the wax-plant, native of southeastern
Asia, is rather frequently grown for ornament in the higher regions
of Guatemala, especially about the capital. It is a small or large, al-
most glabrous vine with very fleshy, oval-oblong, short-acuminate
leaves, and pedunculate umbels of waxy white flowers. It is grown
occasionally as a pot plant in the United States, but is not very com-
mon. In Guatemala it is called "flor de cera."
442 FIELDIANA: BOTANY, VOLUME 24
MARSDENIA R. Brown
Reference: Walter Rothe, Ueber die Gattung Marsdenia R. Br.
und die Stammpflanze der Condurangorinde, Bot. Jahrb. 52: 354-
434. 1915.
Plants herbaceous or more or less woody, usually scandent, rarely erect, pubes-
cent or glabrous; leaves opposite, usually broad; cymes umbelliform or sometimes
branched, terminal or solitary in the leaf axils, the flowers small or medium-sized;
calyx 5-parted, 5-many-glandular within at the base, 5-squamulate, or rarely
eglandular, the segments obtuse; corolla campanulate to urceolate or subrotate,
shallowly or deeply 5-lobate, the throat usually closed by hairs or by an annulus,
rarely naked, the lobes obtuse, contorted, rarely subvalvate; hoods of the corona 5,
adnate to the stamen tube and to the dorsal surface of the anthers, erect, free at
the apex, plane, membranaceous or thickened, sometimes biauriculate at the base;
stamens inserted near the base of the corolla, the filaments connate into a short
tube, the anthers terminated by a small inflexed membrane; pollinia oblong or
ovoid, erect; stigma depressed, flat, or convex at the apex, sometimes rostrate;
follicles often very thick, acuminate, smooth or longitudinally winged, the pericarp
often hard when dried; seeds comose.
There are perhaps more than 50 species of Marsdenia in the tropics
and warmer regions of both hemispheres. We believe that there are
about 27 or 28 species in Mexico, Central America and Panama.
Plants erect; leaves linear-lanceolate, long-attenuate at the base. . . .M. nertifolia.
Plants scandent; leaves usually much broader than linear-lanceolate.
Leaves abundantly pubescent beneath.
Leaf blades acute or acuminate at the base M. trivirgulata.
Leaf blades cordate or rounded at the base.
Leaves glabrous on the upper surfade or nearly so M. propinqua.
Leaves densely pubescent on the upper surface.
Calyx lobes obtuse; pedicels with closely appressed pubescence of short
hairs M. gualanensis.
Calyx lobes attenuate-acuminate; pubescence of the pedicels of rather
long, stiff, spreading hairs M. bourgeana.
Leaves glabrous throughout or nearly so.
Leaves mostly 5-10 cm. broad or larger, broadly rounded at the base.
Tube of the corolla about half as long as the lobes M. schlechteriana.
Tube of the corolla about as long as the lobes.
Peduncles of the inflorescences much shorter than the petioles.
Calyx lobes pubescent dorsally; inflorescence pubescent.
M. marcophylla.
Calyx lobes ciliate only, glabrous dorsally; inflorescence glabrous.
M. maculata.
Peduncles of the inflorescence almost equaling the petioles.
M. stephanotidifolia.
Leaves mostly 2-4 cm. broad, if broader the blades acute or subacute at the
base.
Inflorescences sessile; leaf blades attenuate at the base.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 443
Calyx lobes glabrous dorsally or essentially so, ciliate.
Lateral nerves 6-8 pairs visible on lower surface of leaves; lowland
species from British Honduras and Yucatan M. mayana.
Lateral veins not visible or very obscure on lower surface of leaves;
highland species of Guatemala M. cuneata.
Calyx lobes densely strigose dorsally M. blepharodes.
Inflorescences conspicuously pedunculate, often long-pedunculate.
Pedicels filiform, mostly 15-25 mm. long; corolla lobate almost to the
base M. laxiflora.
Pedicels mostly less than 7 mm. long; corolla with a conspicuous tube.
Leaf blades acute or subacute at the base; calyx lobes conspicuously
ciliate M. pseudoedulis.
Leaf blades rounded at the base; calyx lobes eciliate or nearly so.
M. steyermarkii.
Marsdenia blepharodes Standl. & Steyerm. Field Mus. Bot.
23: 226. 1947.
Known only from the type, Quezaltenango, climbing on a tree
fern at border of forest, San Juan Patzulin, about 1,500 m. ; Steyer-
mark 33615.
A suffrutescent vine, the slender stems terete, when young sparsely puberu-
lent, soon glabrate; leaves on slender petioles 6-15 mm. long, oblong-lanceolate,
7.5-10.5 cm. long, 2-2.5 cm. broad, acuminate or long-acuminate, long-attenuate
to the base, glabrous, slightly paler beneath, the lateral nerves about 3 on each
side, prominent beneath; inflorescences axillary, sessile, many-flowered, dense,
about 1.5 cm. in diameter, the pedicels rather stout, 4-5 mm. long, sparsely puberu-
lent; calyx 2.5-3 mm. long, densely whitish-strigose outside, 5-lobate to the middle,
the lobes oval, rounded at the apex, densely ciliate; corolla dull purple outside,
grayish within, 4.5 mm. long, glabrous outside, the 5 lobes broadly oblong, very
obtuse, densely papillose-puberulent within, somewhat longer than the tube,
spreading; scales of the corona broadly rhombic-ovate, scarcely more than 1 mm.
long; stigma flat or nearly so at the apex.
Marsdenia bourgeana (Baill.) Rothe in Engler, Bot. Jahrb. 52:
408. 1915; L. Wms. Fieldiana, Bot. 32: 50. 1968. Pseudomarsdenia
bourgeana Baill. Hist. Plantes 10: 268. 1890. M. gymnemoides Rothe,
I.e. 409 (type from Guaxacana, Seler & Seler 2904)- M. gilgiana
Rothe, I.e. 410 (type from Mexico, Purpus 2095, or from Guatemala,
Heyde & Lux 45^2).
Woody vines in thickets or forests or on rocky slopes, 1,200-
1,900 m.; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez ; Hue-
huetenango. Southern Mexico. Possibly Honduras.
A small or large, more or less woody vine, the stems and petioles very densely
fulvous-pilose with short spreading hairs; leaves on long or short, slender petioles,
membranaceous, broadly ovate to rounded-ovate, 5-15 cm. long and 3-8 cm. broad,
very obtuse to acuminate, rounded to cordate at the base, very densely velutinous-
pilose on both surfaces or sometimes tomentose beneath; cymes pedunculate, usu-
444 FIELDIANA: BOTANY, VOLUME 24
ally lax and many-flowered, mostly longer than the petioles, repeatedly dichotomous,
densely hirtellous; calyx lobes linear to lanceolate, acute, hirtellous; corolla urceo-
late, sparsely hirtellous outside, 4.5-6 mm. long, barbate within, the lobes semi-
ovate, rounded at the apex; scales of the corona shorter than the stamens; style
with a short conic beak; follicles densely tomentose, about 9 cm. long and 3 cm.
broad, obtuse.
This has been reported from Guatemala as M . mexicana Dene.
Marsdenia cuneata L. Wms. Fieldiana, Bot. 32: 50. 1968.
Known only from the type, Steyermark 51206, Paso del Boqueron,
Huehuetenango. Endemic.
Suffruticose vines, the stems glabrous, with few lenticular lenticels. Leaves
of a pair subequal, lighter in color below, oblanceolate, acute or acuminate, cune-
ate to the base, glabrous, the blades 8-14 cm. long and 2-3 cm. broad, the lateral
veins inconspicuous, petioles to 1.5 cm. long, each provided with 2 narrowly tri-
angular stipules at the base, these about 0.7 mm. long, interpetiolar lines evident;
inflorescences axillary, fascicular, several-many-flowered, peduncles very short or
none, pedicels puberulent, 2-4 mm. long; calyx divided to the base, the lobes ovate
or oblong-ovate, obtuse, ciliate-fimbriate, 2-3 mm. long, provided with a small
horn-shaped callus in each sinus; corolla purple, glabrous outside, campanulate,
divided to about the middle, 4-5 mm. long, the lobes suborbicular, 2-2.5 mm. long
and as broad, obscurely pubescent on inner surface; gynostegium stipitate, about
1.5 mm., faucal corona obscurely lobate, fleshy, anthers terminated with an in-
flexed ovate membrane about 0.5 mm. long; follicles unknown.
Most closely related to M. blepharodes Standl. & Steyerm. but
easily distinguished superficially by the nearly nerveless leaves which
are cuneate to the base and the glabrous instead of strigillose calyx
lobes.
Marsdenia gualanensis Donn.-Sm. Bot. Gaz. 49: 456. 1910.
In thickets or dry rocky slopes, 200-600 m.; Zacapa (type from
Gualan, Deam 6333) ; Jalapa. Apparently endemic.
A small or large, coarse, woody vine, the branches and petioles densely pubes-
cent; leaves on short or elongate, slender petioles, membranaceous, oblong-ovate
to broadly ovate or rounded-deltoid, 5-10 cm. long, short-acuminate or sometimes
narrowed to an obtuse apex, rounded or truncate at the base or sometimes cordate,
very densely and softly pubescent on both surfaces, somewhat paler beneath; in-
florescences much branched, many-flowered, about equaling the petioles, the
branches and pedicels densely and finely pubescent with mostly appressed hairs,
the flowers slender-pedicellate; calyx appressed-puberulent, deeply lobate, the seg-
ments oblong, obtuse; corolla about 2.5 mm. long, glabrous outside, barbate within,
much longer than the short tube; scales of the corona minute; ovaries glabrous;
follicles glabrous, about 8 cm. long and 2.5 cm. thick, obtuse, smooth, the valves
very thick and somewhat ligneous.
FIG. 124. Marsdenia macrophylla. A, habit, X 1A'> B, flower, X 3; C, calyx
and pistil showing basal gland, X 4; D, pollinia, X 40; E, gland from base of
leaf, X 5.
445
446 FIELDIANA: BOTANY, VOLUME 24
This has usually been assumed to be a synonym of Marsdenia
coulteri Hemsl., a Mexican species that seems to be quite different.
Marsdenia laxiflora Donn.-Sm. Bot. Gaz. 40: 7. 1905.
Moist or wet thickets or forest, often on limestone, 350 m. or less;
Alta Verapaz (type from Cubilgiiitz, Tuerckheim 8558b); Pete"n; Iza-
bal. British Honduras; reported from Costa Rica.
A very slender, glabrous, herbaceous or suffrutescent vine; leaves on short
slender petioles, membranaceous or chartaceous, often lustrous, lanceolate or ob-
long-lanceolate, 7-14 cm. long, 1.5-5 cm. broad, gradually or very abruptly acumi-
nate, acute or obtuse at the base, bearing several small glands on the upper surface
at the base of the blade; inflorescences axillary, few-many-flowered, extremely lax
and open, longer or shorter than the leaves, the peduncles 4-6 cm. long or longer,
the filiform pedicels 1.5-2.5 cm. long; calyx lobes ovate, obtuse, 2 mm. long, ciliate;
corolla purple, rotate, the tube 2 mm. long, the lobes oblong or triangular-oblong,
5-6 mm. long, glabrous outside and within, ciliate; follicles very slender, 20-25 cm.
long, 1 cm. thick, smooth, glabrous; seeds brown, thin, 1 cm. long, the coma of
very numerous soft silky white hairs.
Marsdenia macrophylla (Humb. & Bonpl.) Fourn. in Mart. Fl.
Bras. 64: 321. 1885. Asclepias macrophylla Humb. & Bonpl. ex Roem.
& Schult. Syst. Veg. 6: 86. 1820. Cuchampera; cuchamperillo.
Moist or dry thickets or forest, often in rocky places, sometimes
in hedges, 2,250 m. or lower; Baja Verapaz; Escuintla; Quezaltenango.
Mexico; Central America; South America.
Usually a large and coarse, more or less woody vine, glabrous or nearly so out-
side the inflorescence; leaves on rather long, stout petioles, chartaceous or when
fresh somewhat succulent, ovate to elliptic, oval or rounded, 7-19 cm. long, mostly
7-13 cm. broad, usually rounded or very obtuse at the apex and shortly cuspi-
date-acuminate, broadly rounded to shallowly cordate at the base, paler beneath,
glandular on the upper side at the base of the blade; cymes many-flowered, short-
pedunculate or sometimes long-pedunculate, shorter or longer than the petioles,
sparsely or densely puberulent, very dense or rather open, the flowers on stout
pedicels; calyx segments broad, obtuse or rounded at the apex, ciliate, usually
pubescent outside; corolla yellowish or greenish, 5-6 mm. long, glabrous outside,
the lobes broad, somewhat shorter than the urceolate tube or about equaling it;
follicles terete, about 10-23 cm. long and about 5 cm. in diameter, smooth.
The Maya name of Yucatan is recorded as "xemtzul"; "estropajo
cimarron" (Oaxaca).
Marsdenia maculata Hook. f. Bot. Mag. 73: t. 4299. 1847;
L. Wms. Fieldiana, Bot. 32: 51. 1968. Bejuco de leche.
A vine over shrubs and trees at 600 m. or less; Pete"n. Mexico;
Honduras; Panama; Lesser Antilles. Perhaps South America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 447
A large coarse vine to 20 m. or perhaps more, the stems to 1 cm. in diameter
or possibly more. Leaves elliptic-oblong to ovate or suborbicular, the base truncate
to cordate, glabrous, with 5-6 pairs of secondary nerves, 9-25 cm. long and 5-21
cm. broad, petiole 1.5-4 cm. long, with a group of 2-5 small mammillate calluses
at its juncture with the blade; inflorescence subumbellate, many-flowered, peduncle
shorter than the petioles, to about 1.5 cm. long, pedicels 1 cm. or less long; calyx
divided to the base, the lobes ovate, obtuse, the margins sphacelate, ciliate, 3-
4 mm. long and 2-2.5 mm. broad; corolla divided to about the middle, the tube
3-4 mm. long, the lobes 3-4 mm. long and oblong-ovate, obtuse, ciliate, otherwise
glabrous except obscurely pubescent at the base within; corona scales linear-
oblong, obtuse, shorter than the styles; follicles ellipsoidal, up to 22 cm. long and
5 cm. in diameter.
This species is one of the complex often called M. macrophylla.
The group is in need of restudy.
Marsdenia mayana Limdell, Contr. Univ. Mich. Herb. 4: 23.
1940.
British Honduras, the type from Valentin, El Cayo District, on
limestone, C. L. Lundell 6192. Mexico.
A large woody vine as much as 10 m. long, glabrous except in the inflorescence;
leaves short-petiolate, the petioles 12-20 mm. long, the blades membranaceous or
chartaceous, narrowly oblong or oblong-lanceolate, 10-15 cm. long, 2.5-4.5 cm.
broad, acuminate or long-acuminate, long-attenuate to the base, green above, often
lustrous, paler beneath, the lateral nerves 6-8 on each side; inflorescences sessile,
dense, many-flowered, shorter than the petioles, the pedicels 3 mm. long or less,
sparsely puberulent or glabrate; calyx lobes broadly ovate, 2.5-3 mm. long, rounded
at the apex, ciliate, glabrous dorsally or nearly so; corolla greenish, 5 mm. long,
glabrous outside, minutely tomentulose within, the tube campanulate, slightly
longer than the lobes, these broadly ovate, rounded at the apex; scales of the corona
oblong, 3 mm. long; ovaries glabrous.
Marsdenia neriifolia (Dene.) Woodson, Ann. Mo. Bot. Gard.
28: 244. 1941. Blepharodon neriifolium Dene, in DC. Prodr. 8:
604. 1844. Nephradenia neriifolia Benth. & Hook, ex Hemsl. Biol.
Cent. Am. Bot. 2: 336. 1882. N. fruticosa Donn.-Sm. Bot. Gaz. 16:
196. 1891 (type from Rio Rubelcruz, Alta Verapaz, Tuerckheim 1251}.
Along stream banks or on rocks in streams, 200-750 m.; Alta
Verapaz. Oaxaca and perhaps elsewhere in southern Mexico.
An erect glabrous shrub 1-2 meters high, sparsely branched; leaves on short
slender petioles, linear-lanceolate, mostly 8-20 cm. long and 1-2.5 cm. broad, nar-
rowly long-acuminate, long-attenuate to the base, deep green above, somewhat
paler beneath; inflorescences umbellate, axillary, solitary, slender-pedunculate,
mostly 5 cm. long or shorter, few-flowered, the pedicels long and slender; calyx
lobes ovate or suborbicular, ciliate, rounded at the apex; corolla cream-colored,
about 1.5 cm. broad, glabrous outside, the lobes broad, obtuse; stigma convex,
not at all umbonate or rostrate.
448 FIELDIANA: BOTANY, VOLUME 24
Marsdenia propinqua Hemsl. Biol. Cent. Am. Bot. 2: 337. 1882.
Moist or dry thickets, 300-1,400 m.; Jutiapa; Escuintla; Guate-
mala; Huehuetenango. Southern Mexico.
A large, coarse, more or less woody vine, the stems somewhat puberulent or
glabrous; leaves thick, on rather short, slender petioles, oval to rounded-oval
or ovate-oval, mostly 8-17 cm. long and 7-10 cm. broad, usually rounded and
abruptly short-pointed at the apex, broadly rounded or subcordate at the base,
glabrous above or nearly so, velutinous-tomentose or densely short-pilose beneath;
cymes very dense and many-flowered, short-pedunculate, about 3 cm. in diameter,
the branches and stout pedicels tomentulose; calyx thinly tomentose, the lobes
broad, obtuse; corolla 7-8 mm. long, sometimes dark red, pubescent outside and
sparsely pilose within, the lobes broad, rounded at the apex; follicles 11-15 cm.
long, obtuse, smooth, the valves thick and hard; seeds nearly 1 cm. long, flat,
comate with long silky white hairs.
Marsdenia pseudoedulis Woodson, Ann. Mo. Bot. Card. 28:
284. 1941.
Known definitely only from the type, San Marcos, lower southern
slopes of Volcan de Santa Maria, along the great barranco between
Finca Pirineos and San Juan Patzulin, 1,300-1,500 m., Steyermark
33633.
A slender woody vine, the stems puberulent when young; leaves membrana-
ceous, on slender petioles 1-2.5 cm. long, elliptic, 4.5-11 cm. long, 1.5-5 cm. broad,
acuminate, acute or subacute at the base, glabrous, slightly paler beneath; inflores-
cences umbelliform, few-flowered, the peduncles 5 mm. long, sparsely pilosulous,
the pedicels of about the same length; calyx lobes ovate, obtuse, 2 mm. long, pilosu-
lous; corolla campanulate, white, sparsely pilosulous outside, more densely pilos-
ulous within, the lobes slightly longer than the tube, ovate-elliptic, obtuse,
spreading; stigma flat or nearly so; anthers 1 mm. long; scales of the corona
reniform, obtuse, 0.5 mm. long.
Marsdenia schlechteriana Rothe in Engler, Bot. Jahrb. 52:
418, fig. 5. 1915.
Known only from the type, Solola, La Corona, 1,500 m., J. D.
Smith 2773.
A woody vine, the branches glabrous; leaves on petioles about 3 cm. long, oval,
5-11 cm. long, 4-7 cm. broad, acuminate, truncate at the base, membranaceous,
glabrous; inflorescences umbelliform, dense, many-flowered, pedunculate, the pe-
duncle 3 cm. long, the pedicels about 1 cm. long, stout; calyx lobes spreading,
pilosulous outside, ciliate, 2.5 mm. long; corolla tube 2 mm. long, the lobes 4 mm.
long, rounded at the apex, ciliate; gynostegium 2.5 mm. high; scales of the corona
acuminate; stigma bearing a short, thick-conic beak.
This is closely related to M. macrophylla, and it may be that some
of the specimens we have referred to that species belong rather with
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 449
M. schlechteriana. The key characters used by Rothe, however,
would refer all our material rather to M. macrophylla, from which
M. schlechteriana may not be distinct.
Marsdenia stephanotidifolia Woodson in Standl. & Steyerm.
Field Mus. Bot. 23: 79. 1944.
Moist or wet, mixed forest or thickets, 2,200 m. or less; Quezal-
tenango (type collected near Santa Maria de Jesus, along Rio Sa-
mala, Standley 84-593).
A large, more or less woody vine, glabrous except in the inflorescence, the stems
stout, conspicuously lenticellate; leaves on stout petioles 2.5-4.5 cm. long, sub-
chartaceous, oblong-ovate to rounded-ovate, 10-25 cm. long, 6-17 cm. broad,
obtuse or rounded at the apex and cuspidate-acuminate, broadly rounded or sub-
cordate at the base; inflorescences extra-axillary, umbelliform or corymbiform,
repeatedly dichotomous, few-many-flowered, the peduncles almost equaling the
petioles, papillose-puberulent; the pedicels about 8 mm. long, papillose-puberulent;
calyx lobes ovate, obtuse, 5 mm. long; corolla urceolate, 11 mm. long, glabrous
outside, densely villosulous within except on the margins of the lobes, the lobes
erect or somewhat spreading, equaling the tube; gynostegium included, the beak
of the stigma exserted; anthers 5 mm. long; beak of the stigma 4 mm. long.
Marsdenia steyermarkii Woodson, Ann. Mo. Bot. Gard. 28:
285. 1941.
Moist or wet forest or thickets, 1,300-2,500 m.; San Marcos (type
collected on Volcan de Tacana, Steyermark 36019). Mexico (Chiapas).
A small or large, more or less woody vine, glabrous throughout or nearly so;
leaves on slender petioles 1-2 cm. long, oblong or elliptic-oblong, mostly 6-10 cm.
long and 2-3.5 cm. broad, abruptly short-acuminate, rounded at the base, rather
thick, deep green and often lustrous above, somewhat paler beneath; inflorescence
few-flowered, the peduncle bifid, 2.5 cm. long, the pedicels 4 mm. long, sparsely
puberulent or glabrate; calyx lobes ovate, acute, 2 mm. long, minutely papil-
lose-puberulent; corolla campanulate, white, glabrous outside, pilosulous within,
the tube 3 mm. long, the lobes ovate, obtuse, 2.5 mm. long, spreading; gynostegium
3 mm. long; scales of the corona 2 mm. long; stigma broad, conic.
Marsdenia trivirgulata Bartlett, Proc. Am. Acad. Sci. 44: 632.
1909.
Dry rocky slopes, climbing over Opuntia, about 200 m.; Zacapa
(Steyermark 29320). Southwestern Mexico; Honduras; Panama.
A very slender, somewhat woody vine, the stems puberulent in 2 lines or gla-
brate; leaves slender-petiolate, membranaceous, elliptic, narrowly elliptic, or ovate-
elliptic, usually 2.5-7.5 cm. long, acuminate or long-acuminate, acute or attenuate
at the base or rarely obtuse, rather densely puberulent on both surfaces; cymes
almost sessile, few-flowered, sparsely puberulent, the flowers on rather stout, short
450 FIELDIANA: BOTANY, VOLUME 24
pedicels; calyx lobes ovate, very obtuse, white-ciliate, sparsely puberulent; corolla
glabrous outside, white or pale pink striped with lilac, 6 mm. long, the lobes linear-
oblong, several times as long as the short tube, spreading, obtuse, sparsely and
minutely hirtellous within; stigma ending in a beak 1-1.5 mm. long.
MATELEA Aublet
Plants herbaceous or sometimes woody below, usually scandent, sometimes
prostrate or erect, pubescent or almost glabrous; leaves petiolate, various in form,
most often cordate; flowers small or medium-sized, sometimes rather large, the
cymes usually umbelliform; calyx 5-lobate, 5-glandular within at the base; corolla
usually rotate or nearly so, shallowly or deeply 5-lobate, the lobes narrow or broad,
dextrorsely contorted; corona very variable in form, the outer corona entire or
lobate, often annular or fimbriate, sometimes consisting of a large disk adnate to
the corolla throat; stamens inserted near the base of the corolla, the filaments con-
nate into a short tube, the anthers not evidently vesicular, without dorsal append-
ages; pollinia solitary in each cell, clavate or obovoid, horizontal or subpendulous;
stigma depressed at the apex, more or less pentagonal; follicles slender or broad,
smooth or costate, often tuberculate-muricate.
More than 100 species are listed for North America by Woodson,
and many others are found in South America. A few additional spe-
cies are found in southern Central America. The genus is wholly
American.
We have followed Woodson (Ann. Mo. Bot. Gard. 28: 217-238.
1941) in the treatment of Matelea with some misgivings. Looking at
the material from Mexico and Central America that has been placed
here there would seem to be several genera involved. Some of these
have been described, as Woodson points out, as distinct genera.
Certainly, some species included here are less closely related (M. bal-
bisii and M. quirosii are examples) than certain other Mateleas are
to Gonolobi or even to Fischerias.
Leaves not cordate at the base.
Plants erect, not scandent, herbaceous, hirsute M. balbisii.
Plants scandent.
Leaves glabrous beneath.
Leaves 13-18 cm. long; corolla glabrous outside M. glaberrima.
Leaves 4.5-7.5 cm. long; corolla puberulent outside.
Peduncles 3-5 mm. long; calyx lobes broadly ovate; corolla 1 cm. broad.
M. tennis.
Peduncles 8-12 mm. long; calyx lobes linear-lanceolate; corolla about
2 cm. broad M. belizensis.
Leaves pilose beneath, at least on the nerves.
Leaves long-hirsute beneath on the nerves, membranaceous; corolla about
2 cm. broad M. tuerckheimii.
Leaves sparsely puberulent beneath on the nerves, thick; corolla scarcely
1 cm. broad M. campechiana.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 451
Leaves cordate at the base.
Leaf blades broadest at or above the middle, narrowed to a narrow, shallowly-
cordate base; stems hirsute.
Calyx lobes lance-linear; inflorescence lax, few-flowered, the pedicels mostly
2-3 cm. long M. grandiflora.
Calyx lobes ovate or broadly ovate; inflorescence dense and congested, many-
flowered, the pedicels usually very short M. pleistantha.
Leaf blades broadest at the base or at least below the middle, not at all narrowed
toward the base.
Corolla densely pilose within, large, about 3.5 cm. broad M. sylvicola.
Corolla glabrous within or rarely arachnoid-tomentose.
Lobes of the corolla linear or linear-deltoid, 1-2 mm. broad or sometimes
to 5 mm. broad.
Inflorescence racemose; leaf blades deeply cordate at the base.
M. prosthecidiscus.
Inflorescence umbellate; leaf blades shallowly cordate or rounded at the
base.
Leaves glabrous beneath; pubescence of the calyx of minute appressed
hairs M. belizensis.
Leaves hirsute beneath on the nerves; pubescence of the calyx of stiff
spreading hairs.
Corolla lobes pale green, 1 mm. broad M. medusae.
Corolla lobes purple-brown, 2-2.5 mm. broad M. gonoloboides.
Lobes of the corolla triangular-lanceolate to suborbicular, usually much
more than 2 mm. broad.
Lobes of the corolla rounded at the apex, usually as broad as long, rarely
only oval.
Flowers small, the corolla 3-7 mm. long.
Calyx lobes acute, longer than broad.
Plant glabrous; calyx lobes less than 3 mm. long . . .M. pusilliflora.
Plant hirsute; calyx lobes more than 5 mm. long . . .M. molinarum.
Calyx lobes suborbicular, apex rounded M. velutina.
Flowers relatively large the corolla usually more than 1 cm. long (some-
times only 8 mm. in some flowers).
Calyx lobes rounded at the apex and subulate-cuspidate; plants
densely hirsute throughout M. guatemalensis.
Calyx lobes acute to very obtuse at the apex, not cuspidate.
Calyx lobes very obtuse.
Corolla merely puberulent outside Vincetoxicum hatchii.
Corolla hispid or hirsute outside M. velutina.
Calyx lobes acute or acuminate.
Leaves lance-oblong, shallowly cordate at the base; lobes of the
corolla much longer than broad M. steyermarkii.
Leaves rounded-ovate, deeply cordate at the base; lobes of the
corolla about as broad as long M. quirosii.
Lobes of the corolla acute or at least pointed, sometimes attenuate,
usually much longer than broad, sometimes broadly triangular.
Corolla glabrous outside.
Leaves glabrous or nearly so, 12-15 cm. long M. tikalana.
Leaves pubescent on both surfaces, 4-7 cm. long . . . .M. megacarpha.
Corolla sparsely or densely pubescent outside.
Corolla arachnoid-tomentose within the throat. . . .M. araneosa.
452 FIELDIANA: BOTANY, VOLUME 24
Corolla not arachnoid-tomentose within.
Leaves oblong-ovate.
Corolla about 14 mm. long, the lobes linear-lanceolate.
M. patalensis.
Corolla about 7 mm. long, the lobes broadly ovate. .M. violacea.
Leaves broadly ovate-cordate.
Corolla lobes broadly ovate; calyx long-hirsute; follicles glandu-
lar-puberulent M. pseudobarbata.
Corolla lobes triangular; calyx hirtellous; follicles glabrous.
Calyx lobes 4-5 mm. long; corolla about 8 mm. long.
M. diffusa.
Calyx lobes about 8 mm. long; corolla about 15 mm. long.
M. ceratopetala.
Matelea araneosa (Donn.-Sm.) Woodson, Ann. Mo. Bot. Gard.
28: 222. 1941. Gonolobus araneosus Donn.-Sm. Bot. Gaz. 47: 257.
1909.
Known only from the type, Alta Verapaz, mountains between
Tactic and Coban, 1,650 m., Tuerckheim II. 2332.
Plants scandent, suffrutescent, the branches, petioles, inflorescence and calyx
pilose with spreading hairs; leaves on petioles 1-2 cm. long, oblong-ovate or lance-
ovate, 6-8 cm. long, 2.5-3.5 cm. broad, acuminate, shallowly cordate at the base,
with a broad open acute sinus, softly pilose; inflorescence umbelliform, slightly
longer than the petioles, the peduncle 3-5 mm. long, the 4-5 pedicels 3-7 mm.
long; calyx lobes linear-lanceolate, 5 mm. long; corolla reticulate-veined, lobate to
the middle, whitish-arachnoid within below the lobes, 8 mm. long, rotate, the lobes
glabrous within, sparsely pilosulous outside, ciliate at the apex; corona cyathiform,
equalling the gynostegium and free from it, 5-denticulate, 1 mm. high, 2 mm. broad.
Matelea balbisii (Dene.) Woodson, Ann. Mo. Bot. Gard. 28:
231. 1941. Asclepias villosa Balb. Mem. Acad. Sci. Torino 7: 386.
1803, not Mill. 1768. Lachnostoma balbisii Dene, in DC. Prodr. 8:
602. 1844. Gonolobus pogonanthus Hemsl. Biol. Cent. Am. Bot. 2:
333. 1882. Pherotrichis balbisii Gray, Proc. Am. Acad. 21: 400. 1886.
Grassy open slopes, 2,000 m. ; Huehuetenango (southeast of Hue-
huetenango, Steyermark 48172). Mexico.
Plants herbaceous, erect, arising from a usually somewhat tuberous-thickened
root, the stems usually simple, stout, abundantly leafy, densely hirsute with spread-
ing yellowish hairs; leaves short-petiolate, rather thick, broadly ovate to lanceolate
or even oblanceolate, 3.5-8 cm. long, 1-5.5 cm. broad, rounded or very obtuse at
the apex, usually rounded or very obtuse at the base, rarely attenuate, sparsely or
densely hirsute on both surfaces or sometimes glabrate; flowers pale green, umbel-
late, the umbels appearing in most of the leaf axils above the middle of the stem,
sessile or nearly so, rather few-flowered, densely hirsute, the pedicels 6 mm. long
or shorter; calyx lobes short, lanceolate, hispidulous; corolla campanulate-rotate,
5-7 mm. long, densely white-barbate within; crown composed of 5 quadrate ex-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 453
cised-truncate scales, these longer than the hyaline-tipped anthers; stigma bearing
a large, globose or conic appendage; follicles rather slender, terete, narrowly long-
acuminate, not tuberculate, very densely hirsute with long spreading hairs.
In general appearance this plant resembles an Asclepias, and is
very unlike other Central American species of Matelea and perhaps
does not belong in this genus.
Matelea belizensis (Lundell & Standl.) Woodson, Ann. Mo. Bot.
Card. 28: 232. 1941. Vincetoxicum belizense Lundell & Standl. ex
Standl. Field Mus. Bot. 17: 268. 1937. Dama de la noche.
Known only from the type, British Honduras, Corozal District,
little above sea level, Gentle 299.
Stems herbaceous, scandent, appressed-pilose; leaves on petioles 5-17 mm.
long, ovate or ovate-oblong, 4.5-7.5 cm. long, 2-4.5 cm. broad, acutely acuminate,
shallowly cordate at the base or only rounded, rather thick, glabrous above except
for a few appressed hairs along the nerves, paler beneath, with a few short hairs
along the costa, otherwise glabrous; peduncles 8-12 mm. long, puberulent, the in-
florescence subumbellate, 8-12-flowered, the pedicels slender, puberulent, 6-10 mm.
long; calyx 5-parted, the segments linear-lanceolate, 3 mm. long, subobtuse, puber-
ulent outside, alternating with 5 small basal appendages; corolla campanulate,
5-lobate almost to the base, about 10 mm. long, the lobes narrowly lanceolate,
9 mm. long, 2-3 mm. broad, subacute, glabrous within, sparsely puberulent out-
side; outer corona glabrous, fleshy, deeply 5-lobate, the lobes 3-4-dentate.
Matelea campechiana (Standl.) Woodson, Ann. Mo. Bot. Gard.
28: 234. 1941. Vincetoxicum campechianum Standl. Carnegie Inst.
Wash. Publ. 461:82. 1935.
Moist or wet thickets, about 100 m.; Peten; Alta Verapaz. Mex-
ico (Campeche) ; British Honduras.
A somewhat woody vine, the stems slender, rather densely pilose with short
spreading hairs or in age glabrate; leaves on petioles 5-15 mm. long, subcoriaceous,
lance-oblong to oblong-ovate or broadly ovate, 3-7 cm. long, 1-4.5 cm. broad,
acute or short-acuminate, rounded at the base, somewhat lustrous above, ap-
pressed-pilosulous along the costa, paler beneath, glabrous; inflorescences umbelli-
form, 3-6-flowered, the peduncle 3-5 mm. long, the pedicels stout, 3-6 mm. long,
puberulent; calyx 1.5-2 mm. long, the lobes oval or broadly ovate, obtuse or
rounded at the apex, minutely appressed-pilosulous; corolla spreading or reflexed,
dull green with bronze-green reticulations, minutely hirtellous outside, 5 mm. long,
the lobes broadly elliptic or almost rounded, rounded at the apex, glabrous within
or puberulent only at the base; gynostegium pentagonal, 2 mm. broad, borne on a
short thick stipe; corolla pale, narrow, subentire; follicles narrowly lanceolate,
narrowly long-acuminate, about 7.5 cm. long and almost 2 cm. broad, glabrous,
bearing few thick fleshy tubercles 2-4 mm. long.
454 FIELDIANA: BOTANY, VOLUME 24
Matelea ceratopetala (Donn.-Sm.) Woodson, Ann. Mo. Bot.
Gard. 28: 236. 1941. Dictyanthus ceratopetalus Donn.-Sm. Bot. Gaz.
18: 208. 1893. Cochita; cochitos; chununa; corazon de low (Escuintla) ;
sombreros, sombreritos (flowers).
Moist or dry, brushy, often rocky plains and hillsides, 1,350 m.
or lower; Jalapa; Jutiapa; Santa Rosa (type from Santa Rosa,
Heyde & Lux 3999} ; Escuintla. Southern Mexico (Oaxaca, Puebla) ;
Honduras.
A small or rather large, more or less woody vine, or the stems sometimes pros-
trate, the older stems ochraceous, the stems thinly hispid with bulbous-based hairs
and minutely puberulent; leaves on long slender petioles, broadly ovate-cordate,
membranaceous, 2-7 cm. long, acuminate or short-acuminate, usually deeply cor-
date at the base, scaberulous and hispidulous above, hispidulous beneath on the
nerves and veins, the veins usually elevated and closely reticulate; inflorescence
umbellate, 2-8-flowered, sessile or short-pedunculate, the flowers slender-pedicel-
late; calyx lobes lanceolate, 12 mm. long or shorter; corolla green and brown-purple
within, about 2 cm. long, rotate-campanulate, puberulent outside, more or less
puberulent within or almost glabrous, 5-lobate to about the middle, the lobes tri-
angular, acute or with a narrow obtuse tip, the margins revolute; follicles glabrous,
6.5-10 cm. long, narrowly long-acuminate, bearing few or numerous, fleshy, long
or short tubercles.
The young fruits of this and other species are cooked and eaten.
Called "cuchampel" and "chanchitos" in Honduras.
Matelea decaisnei Woodson, Ann. Mo. Bot. Gard. 28: 232.
1941. Polystemma viridi 'flora Dene, in DC. Prodr. 8: 602. 1844, not
M. viridiflora Woodson, 1941. Malacate.
Moist or dry, often rocky, brushy slopes, 200-900 m.; Zacapa;
Chiquimula. Southern Mexico; Honduras.
A small woody vine, the slender stems puberulent and sparsely hispid; leaves
slender-petiolate, broadly or narrowly ovate-cordate, 2.5-6.5 cm. long, usually
long-acuminate, rather deeply and narrowly cordate at the base, densely puberu-
lent and sparsely hispidulous on both surfaces, membranaceous; umbels 1-4-flow-
ered, often equaling the leaves, the peduncles 2 cm. long or shorter, the pedicels
equaling or longer than the peduncles; sepals pale green, lanceolate, about 8 mm.
long, acuminate, puberulent and hispidulous; corolla in bud lanceolate and long-
acuminate, glabrous outside or very minutely puberulent, 1.5-2 cm. long, green
within and glabrous or nearly so, the lobes triangular-lanceolate, attenuate to an
obtuse tip; corona adnate to the gynostegium, composed of 5 ligulate scales and
20 small hair-like ones; stigma flat; follicles rather slender, smooth, about 9 cm.
long and 1.5 cm. thick, glabrous.
So far as known, the range of this species in Guatemala is narrowly
limited, and confined to the dry hills and plains of Zacapa and Chi-
quimula. Woodson has referred some of the specimens to M. decais-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 455
nei and to M. rupestris (Brandegee) Woodson, but apparently all the
material represents a single species.
Matelea diffusa Woodson, Ann. Mo. Bot. Card. 28: 236. 1941.
Dictyanthus prostratus Brandegee, Univ. Cal. Publ. Bot. 7: 329. 1920,
not M. prostrata Woodson, 1928. Chinuna.
Dry open slopes, 800-1,600 m.; Jalapa; Huehuetenango. South-
ern Mexico; El Salvador.
A small vine, the stems herbaceous or suffrutescent, sometimes prostrate,
densely puberulent and thinly hispid; leaves thick-membranaceous, very broadly
ovate-cordate, mostly 2-2.5 cm. long and about as broad, acute or short-acuminate,
deeply and narrowly cordate at the base, densely hispidulous on both surfaces, the
veins prominent beneath and closely and conspicuously reticulate; peduncles very
short, mostly 2-flowered, the peduncles and pedicels together scarcely equaling
the petiole; calyx lobes ovate, acuminate, 3 mm. long; corolla broadly campanulate
or rotate-campanulate, about 7 mm. long, puberulent outside, minutely puberulent
within, dark brown-purple, the lobes triangular, narrowed to a narrow obtuse tip,
revolute; outer corona deeply 5-lobate, the lobes narrow, wholly adnate to depres-
sions in the lower half of the corolla tube; stigma pentagonal, umbonate; follicles
about 5 cm. long and 1.5 cm. broad, long-acuminate, glabrous, somewhat lustrous,
bearing few large stout fleshy spine-like tubercles.
Here probably belongs a collection reported from Huehuetenango
by Loesener as Dictyanthus parviflorus Hemsl.
Matelea glaberrima Woodson, Ann. Mo. Bot. Gard. 28: 281.
1941.
Known only from the type, Pete"n, Uaxactun, Bartlett 12300.
A glabrous woody vine; leaves on petioles 1-3 cm. long, ovate-elliptic, 13-
18 cm. long, 5-11 cm. broad, acuminate, rounded at the base, coriaceous; inflores-
cence umbelliform, few-flowered, subsessile, the pedicels about 1 cm. long; flower
green and brown; calyx lobes ovate-lanceolate, 5 mm. long, acuminate, minutely
papillose-puberulent; corolla rotate, glabrous outside, papillose-puberulent within,
17 mm. broad, the lobes ovate-elliptic, acute, 8 mm. long, spreading; crown cup-
shaped, strongly pentagonal, the margin undulate, the sinuses bilamellate and
bearing an inconspicuous ligule; gynostegium substipitate; stigma strongly pentag-
onal, depressed.
We have not seen material of this species. The large, coriaceous
leaves should help in distinguishing this species.
Matelea gonoloboides (Rob. & Greenm.) Woodson, Ann. Mo.
Bot. Gard. 28: 222. 1941. Urostephanus gonoloboides Rob. & Greenm.
Am. Journ. Sci. III. 50: 159. 1895.
FIG. 125. Matelea diffusa. A, habit, X Y2; B, corolla, X 2; C, calyx and pistil,
X 4; D, follicle, X 1.
456
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 457
Moist thickets or forest, 1,800-2,300 m.; Quezaltenango; Solola;
also collected elsewhere (Barranco de los Condenados ?), at a local-
ity of uncertain position. Mexico (Chiapas, Morelos).
A very slender, apparently herbaceous vine, the stems hirsute with deflexed
fulvous hairs; leaves thin, ovate-cordate to oblong-ovate, 5-6.5 cm. long, acumi-
nate or long- acuminate, rather shallowly and narrowly cordate at the base, thinly
hirsute on both surfaces; inflorescence umbellate, 1-5-flowered, the peduncle 4 mm.
long, the pedicels longer than the peduncle; calyx lobes ovate-lanceolate, acute,
almost 4 mm. long, hirtellous; corolla brown-purple, about 1-2 cm. long, rotate,
5-lobate almost to the base, the lobes linear to narrowly deltoid-lanceolate, mostly
2-5 mm. broad, arachnoid-pubescent within near the base, thinly hirtellous out-
side; corona attached to the lower part of the stamen tube, almost equaling the
gynostegium, shallowly lobate at the summit with 5 internal horn-like processes
opposite the stamens and 5 outer lobes alternate with them and produced on their
outer surface into 2 filiform flexuous tails; stigma depressed.
Matelea grandiflora (Standl.) Woodson, Ann. Mo. Bot. Gard.
28: 235. 1941. Vincetoxicum grandiflorum Standl. Carnegie Inst.
Wash. Publ. 461: 83. 1935. Gonolobus fulvidus Ballard, Bot. Mag.
163: t. 9611. 1940.
Moist or wet forest or thickets, 270 m.; Pete"n; Alta Verapaz.
British Honduras, the type from Machaca, Schipp S575.
A large coarse vine, probably somewhat woody, as much as 9 m. long, the stems
stout, very densely villous or hirsute with long spreading fulvous several-celled
hairs; leaves rather thick, on petioles scarcely 1 cm. long, oblong, mostly 10-16 cm.
long and 4-7 cm. broad, shortly cuspidate-acuminate, narrowly rounded at the
base and deeply and narrowly cordate, densely setose-pilose on the upper surface
with long subappressed hairs, paler beneath, densely setose-pilose with long spread-
ing yellowish hairs; flowers large, bright yellow, umbellate, the umbels short-
pedunculate, few-flowered, the slender pedicels 3.5 cm. long or shorter; sepals
lance-linear, about 1 cm. long, long-attenuate, pilose; corolla rotate, 4-5 cm.
broad, 5-lobate to the middle or less, setose-pilose outside, the lobes ovate-orbicu-
lar, rounded at the apex and shallowly emarginate, pilose or pilosulous within,
especially near the base, conspicuously reticulate-veined; corona double, arising
from the staminal column near its base, the outer corona yellow, a short fleshy cup,
pentagonal; inner corona of 10 minute liguliform appendages; carpels of the ovary
pubescent.
Matelea guatemalensis (Schum.) Woodson, Ann. Mo. Bot.
Gard. 28: 235. 1941. Gonolobus velutinus var. calycinus Donn.-Sm.
Bot. Gaz. 13: 189. 1888 (type from Pansamala, Alta Verapaz, Tuerck-
heim 1121+). G. guatemalensis Schum. in Engler & Prantl, Pflanzenf.
4, pt. 2: 302. 1895.
Moist or wet forest or thickets, sometimes on limestone, 2,000 m.
or lower; Alta Verapaz; Izabal (?); Huehuetenango. Endemic.
458 FIELDIANA: BOTANY, VOLUME 24
A large or small, herbaceous or suffrutescent vine, densely hirsute throughout
with long, spreading, stiff, yellowish or brownish hairs; leaves slender-petiolate,
ovate or oblong-ovate, mostly 6-11 cm. long and 3.5-5.5 cm. broad, shortly cau-
date-acuminate at the broad apex, rather deeply and narrowly cordate at the base;
inflorescences umbelliform, very slender, few-flowered, equaling or somewhat
shorter than the leaves, the peduncles as much as 8 cm. long, the pedicels very
unequal, 4.5 cm. long or shorter; calyx lobes almost equaling the corolla, oval,
rounded and caudiculate at the apex, sparsely hirsute, pale; corolla 2.5 cm. broad,
flat, glabrous or nearly so, yellowish-brown, lobate almost to the base, the lobes
oval-orbicular, rounded at the apex, venose; corona dark-colored, cupular, about
3 mm. broad, carunculate, no outer corona present; young follicles slender, sparsely
fleshy-tuberculate, densely fulvous-hirsute.
Vincetoxicum hatchii Standl. Field Mus. Bot. 17: 269. 1937.
Probably a forest plant at about 1,000 m.; Alta Verapaz (type,
Hatch & Wilson 107). Apparently endemic.
A large, herbaceous or suffrutescent vine, the stems hirtellous and puberulent;
leaves on very long, slender petioles, thin, orbicular-cordate, about 20 cm. long
and of about the same breadth, acute or acuminate, or usually abruptly cuspidate-
acuminate, deeply and narrowly cordate at the base, finely and rather densely
hirtellous; calyx lobes broadly elliptic, 10-14 mm. long, obtuse; corolla brownish
or green, about 3 cm. broad, rotate, pubescent outside, glabrous within, the lobes
suborbicular, rounded at the apex; outer corona annular, the inner one thick, toru-
lose, with 5 erect rounded lobes.
The type specimen is inadequate and the species should not have
been described from it. In June, 1941 Woodson annotated the type
as "IMatelea magnifolia (Pittier) Woodson" but did not publish the
reduction. It would seem not to belong to that species.
Matelea medusae Woodson, Field Mus. Bot. 23: 78. 1944.
Rocky slopes and thickets, 1,400-1,900 m.; Huehuetenango (type
from Cerro Pueblo Viejo, Steyermark 50987). Mexico (Chiapas).
Plants very slender, scandent, herbaceous or suffrutescent, the stems very
densely and finely glandular-puberulent and sparsely short-hirsute; leaves thin-
membranaceous, on slender petioles about 1-1.5 cm. long, oblong-elliptic or oblong-
ovate, 4-9 cm. long, 1.5-3.5 cm. broad, sparsely appressed hirsute pubescent with
multicellular hairs, hirsute on the nerves, scarcely paler beneath, inconspicuously
glandular above at the very base of the blade; inflorescences extra-axillary, umbelli-
form, 5-10-flowered, the peduncles 5-10 mm. long, the pedicels about equaling the
peduncle; calyx lobes linear-lanceolate, caudate-acuminate, 3-4 mm. long, hirsute
dorsally; corolla pale green, cleft almost to the base, the lobes linear, 15-20 mm.
long, 1-2 mm. broad, revolute, glabrous within except minutely pilose at base,
pubescent outside; gynostegium long-stipitate, the corona purple, columnar, 1.3
mm. high, the 5 lobes spreading, obtuse, callous-emarginate; stigma flat, 1.3 mm.
broad.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 459
Matelea megacarpha (Brandegee) Woodson, Ann. M o. Eo
Gard. 28: 236. 1941. Vincetoxicum megacarphum Brandegee, Univ.
Cal. Publ. Bot. 4: 381. 1913. Pachystelma cordatum Brandegee, I.e.
7: 330. 1920. Dictyanthus brachistanthus Standl. Field Mus. Bot. 8:
38. 1930 (type from Atitlan, Solola, Heyde & Lux 63^6).
Moist or wet thickets, about 400-1,000 m.; El Progreso; Santa
Rosa; Solola; Quiche. Mexico (Chiapas, Veracruz); Honduras;
Nicaragua.
A slender vine, herbaceous or somewhat lignescent, the stems puberulent and
short-hirsute; leaves membranaceous, on long slender petioles, deltoid-ovate, 4-
7 cm. long, acuminate or long-acuminate, shallowly cordate at the base, green
above and rather densely short-hirsute, paler beneath, densely puberulent or short-
pilose; cymes umbelliform, few-flowered, the peduncles usually longer than the
petioles, the pedicels long and slender, very unequal; calyx deeply lobate, the lobes
ovate-lanceolate, puberulent and sparsely hirtellous, conspicuously ciliate, acumi-
nate; corolla rotate, 5-lobate to the middle, about 6 mm. long, dull brown-purple,
puberulent outside, very minutely puberulent within, the lobes deltoid, narrowed
to the obtuse apex, reticulate- veined; outer corona deeply lobate, the lobes narrow,
elongate, partially adnate to the corolla, obtuse; gynostegium sessile; stigma de-
pressed at the apex, pentagonal.
Matelea molinarum L. Wms. Field Mus. Bot. 32: 54. 1968.
A vine growing over brush or in open forests, at about 1,200 m.;
Alta Verapaz; Huehuetenango; apparently endemic.
Vines of unknown length, the stems spreading hirsute, up to about 4 mm. in
diameter. Leaves ovate, cordate, short-acuminate, 4-10 cm. long and 2.5-7 cm.
broad, sparsely fulvous-hirsute on both surfaces, especially on the nerves, and with
abundant very short puberulence, petioles slender, spreading hirsute pubescent,
3-4 cm. long, inflorescence axillary or near leaf axils, a few-flowered subumbellate
cyme, abundantly hirsute, peduncles 2-4 cm. long, the pedicels 1-3 cm. long;
flowers small, green; calyx deeply lobed, long hirsute pubescent outside, glabrous
within, lobes broadly lanceolate, acuminate, about 7 mm. long and 3 mm. broad;
corolla rotate, or the lobes reflexed, fleshy, about 1.2-1.5 cm. across, glabrous with-
in, provided with a tuft of hirsute pubescence near the apex outside and with
minute subglandular pubescence, lobes broadly ovate or suborbicular, about 6 mm.
long and 5 mm. broad; faucal corona erect, fleshy, the margin with many small
unbonate processes, about 1 mm. high; gynostegium stipitate; follicles subfusi-
form, 10-12 cm. long and 3 cm. in diameter, covered with fleshy spines 1 cm. or
less long.
This species seems to be quite closely related to M. nigrescens
(Schlecht.) Woodson, but is easily distinguished superficially by the
glabrous inner face of the corolla as well as by other detail.
460 FIELDIANA: BOTANY, VOLUME 24
Matelea patalensis (Donn.-Sm.) Woodson, Ann. Mo. Bot.
Card. 28: 223. 1941. Gonolobus patalensis Donn.-Sm. Bot. Gaz. 47:
256. 1909.
Known only from the type, Baja Verapaz, Fatal, 1,600 m., Tuerck-
heim II. 2371.
Plants scandent, suffrutescent, the stems, petioles and inflorescence pilose with
spreading hairs; leaves on petioles 1.5-3 cm. long, oblong-ovate, 7-11 cm. long,
3.5-6 cm. broad, acuminate, shallowly cordate, sparsely and minutely strigillose;
peduncles 10-13 mm. long, about 5-flowered, the pedicels 15-20 mm. long; calyx
lobes linear-lanceolate, 5 mm. long; corolla rotate, 14 mm. long, glabrous and re-
ticulate within, sparsely pilose outside, deeply lobate, the lobes oblong, obtuse;
corona cyathiform, free from the gynostegium and almost enclosing it, 2 mm. high,
3 mm. broad, the 5 scales minute, bifid.
Matelea picturata (Hemsl.) Woodson, Ann. Mo. Bot. Card. 28:
233. 1941; L. Wms. Fieldiana, Bot. 32: 55. 1968. Gonolobus pictu-
ratus Hemsl. Biol. Cent. Am. Bot. 2: 332, t. 56, B, ff. 5, 6. 1882.
Moist or wet forest, 1,300-1,500 m.; endemic; Quezaltenango
(type from Volcan de Zunil, Salvin; collected also on Volcan de Santa
Maria, between Santa Maria de Jesus and Calahuache') .
Plants scandent, herbaceous or suffrutescent, glabrous throughout or nearly so
except for the flowers, or the young parts sometimes minutely puberulent; leaves
membranaceous, on long slender petioles, cordate-ovate, mostly 7-12 cm. long,
rather abruptly long-acuminate, shallowly cordate at the base; peduncles long and
slender, often 5 cm. long, few-flowered, the flowers umbellate, about 2.5 cm. broad,
the slender pedicels mostly 1-2.5 cm. long; calyx lobes lanceolate, 3-4 mm. long;
corolla rotate, yellowish green or often with brownish lines, glabrous outside, rather
densely villous within with very slender, weak hairs, the lobes broadly ovate or
rounded-ovate, obtuse or rounded at the apex; gynostegium very short; corona
very short, fleshy, undulate; stigma depressed at the apex.
Matelea pittieri from Panama is closely related and possibly
synonymous.
Matelea pleistantha (Donn.-Sm.) L. Wms. Fieldiana, Bot. 32:
55. 1968. Macroscepis pleistantha Donn.-Sm. Bot. Gaz. 20: 543. 1895
(type, Heyde & Lux 6350). Macroscepis congestiflora Donn.-Sm. I.e.
25: 149. 1898 (type, same as preceding). Matelea congestiflora Wood-
son, Ann. Mo. Bot. Gard. 28: 224. 1941.
Most common in dry thickets or on rocky slopes, 200-1,200 m.;
Huehuetenango; Zacapa; Santa Rosa. El Salvador; Honduras; Nica-
ragua; Costa Rica.
A rampant vine with stout stems, densely setulose-hirsute with long, spreading,
fulvous or brownish hairs; leaves membranaceous, on petioles 3.5 cm. long or usu-
STANDLEY AND WILLIAMS: FLORR OF GUATEMALA 461
ally much shorter, broadly obovate or rounded-obovate, 10-20 cm. long, 7-12 cm.
broad, broad at the apex and shortly cuspidate-acuminate, narrowed to the rounded,
narrow, shallowly cordate base, densely hirtellous or puberulent on both surfaces
and long-hirsute on the nerves and veins; inflorescence short-pedunculate or almost
sessile, very dense and head-like, few-many-flowered, the bracts very small or none,
the inflorescences little longer than the petioles or sometimes shorter, the pedicels
sometimes 12 mm. long but mostly much shorter; calyx lobate almost to the base,
the lobes ovate, acuminate, about 9 mm. long; corolla glabrous outside, the tube
6 mm. long, the lobes papillose within, the limb spreading, almost 2 cm. broad,
lobate to the middle, the lobes rounded; corona scales deltoid, incurved and trun-
cate at the apex, connate below; ovaries glabrous, the stigma flat.
In El Salvador the name "cuchamper de mico" sometimes is
applied to the plant.
Matelea prothecidiscus Woodson, Ann. Mo. Bot. Gard. 28:
223. 1941. Prothecidiscus guatemalensis Donn.-Sm. Bot. Gaz. 25:
150, t. 12. 1898, not Matelea guatemalensis Woodson.
Moist forest, 1,100-1,300 m.; Santa Rosa (type from Cerro Gordo,
Heyde & Lux 384-5) ; Huehuetenango (Paso del Boqueron, below La
Libertad). Nicaragua (Managua).
A large coarse vine, probably somewhat lignescent, the stems, petioles, and
inflorescence densely glandular-pubescent and sparsely setose-hirsute; leaves mem-
branaceous, on long slender petioles, broadly rounded-ovate, 7-13 cm. long, 8-
12 cm. broad, rounded at the apex or obtuse and shortly cuspidate-acuminate, very
sparsely setose-hispidulous on the upper surface with short whitish hairs, paler
beneath, rather densely glandular-puberulent, somewhat setose-hispidulous on the
nerves and veins, the veins dark and conspicuous when dry; inflorescences short-
racemose, shorter than the leaves, few-flowered, the peduncles equaling or shorter
than the petioles, the flowers long-pedicellate, the pedicels refracted in age; calyx
deeply lobate, the lobes lanceolate, 3-5-glandular within; corolla lobate almost to
the base, abruptly reflexed, pale green, blackish when dried, glabrous within,
sparsely pubescent outside, the lobes linear, 2 cm. long; outer corona elongate-
annular, 5-crenate, the inner one reduced to 5 keels adnate to the dorsal surface
of the filaments; immature follicles 9 cm. long, densely glandular-pubescent, densely
covered with long spine-like fleshy tubercles.
Matelea pusilliflora L. Wms. Fieldiana, Bot. 32: 55. 1968.
Mosquito whist (British Honduras).
Slender vine growing over other plants, known only at low eleva-
tions; Pete"n. British Honduras (type, Gentle 6050).
Slender, glabrous, much branched, herbaceous or suffrutescent vines, stems
becoming 2-edged and finally corky, internodes on old stems short, mostly about
8 cm. long, those on lateral flowering branches mostly 2-3 cm. long. Leaves short
petiolate, glabrous or nearly so, lanceolate, acute or acuminate, truncate to sub-
cordate at the base, lateral nerves 3-5 prominent pairs, these mostly not opposite,
462
FIELDIANA: BOTANY, VOLUME 24
FIG. 126. Matelea quirosii. Photograph of flower on herbarium specimen,
Roe, Roe & Mori 494, snowing highly evolved gynostegium, X 3.
2.5-5 cm. long and 0.8-2.2 cm. broad, petioles slender, 2-10 mm. long, obscurely
puberulent, with 3-5 digitiform calluses at its juncture with the blade; inflorescence
axillary, 1-2-flowered, nearly sessile, peduncle 1-2 mm. long, pedicel 1-2 mm. long;
flowers smallest of the genus, green with petals white tipped; calyx lobed to near
the base, glabrous lobes ovate-lanceolate, acute, 1.5-2 mm. long; corolla rotate,
glabrous, prominently veined, 5-6 mm. broad, lobate to about the middle, the
lobes orbicular, obtuse, about 2 mm. long and as broad, the apex usually white;
corona fleshy, low with thin margin, 5-umbonate within; gynostegium about 1 mm.
broad, the stigma depressed; fruits unknown.
This is the smallest flowered of all the Mateleas having a rotate
corolla; it is easily distinguished by the nearly sessile 1-2-flowered
inflorescences, the small leaves and short internodes on secondary
branches.
Matelea quirosii (Standl.) Woodson, Ann. Mo. Bot. Gard. 28:
224. 1941; L. Wms. Fieldiana, Bot. 32: 56, fig. 1968. Labidostelma
guatemalense Schlechter, Bull. Herb. Boiss. II. 6: 843. 1906, not M.
guatemalensis Woodson, 1941 (type from Nenton, Huehuetenango,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 463
C. & E. Seler 3279). Vincetoxicum quirosii Standl. Field Mus. Bot.
18: 959. 1938 (type from Puntarenas, Costa Rica).
Moist or wet thickets or forest, 300-800 m. or perhaps higher;
Baja Verapaz; Chiquimula; Guatemala; Huehuetenango. Southern
Mexico; El Salvador; Nicaragua; Honduras; Costa Rica.
A large or small vine, herbaceous or lignescent, the stems slender, densely
puberulent and sparsely hirsute with long spreading slender hairs, also short-
hirtellous; leaves long-petiolate, membranaceous, broadly ovate or usually rounded-
cordate, mostly 5-9 cm. long, rather abruptly acuminate or cuspidate-acuminate,
deeply cordate at the base, green above, densely pilose with both long and short
hairs, paler beneath, densely velutinous-pilose; peduncles long and slender, mostly
1-3-flowered, or the flowers sometimes more numerous, subumbellate, the pedicels
2.5 cm. long or shorter; calyx pale green, lobate almost to the base, the lobes oblong-
lanceolate, acuminate, 8 mm. long, sparsely pilose outside, long-ciliate; corolla
rotate, about 2.5 cm. broad, puberulent outside, glabrous within, white toward the
base, reticulate-veined with brown-purple, shallowly lobate, the lobes triangular
or rounded-triangular, obtuse; corona scales 5, adnate to the base of the gynoste-
gium and to the corolla, carnose, connate only at the base, bearing near the apex
2 falcate-ligulate processes; anthers short, hyaline-appendaged; follicles very slen-
der, 7-13 cm. long, 1 cm. thick, long-attenuate to each end, smooth, glabrous.
This species is a very distinctive one because of the complicated
gynostegium. It is found mostly at low elevations near the Pacific.
It is the type species of Schlechter's genus Labidostelma.
Matelea steyermarkii Woodson, Ann. Mo. Bot. Gard. 28: 278.
1941.
Moist or wet, mixed forest, 2,500-3,500 m.; endemic; San Mar-
cos (type from Volcan de Tajumulco, southern slopes near Las
Canojas, Steyermark 35810).
A large or small, chiefly herbaceous vine, the stems slender, densely hirsute
with brownish, spreading or reflexed hairs; leaves slender- petiolate, membrana-
ceous, oblong-ovate, 5-13 cm. long, 1.5-7.5 cm. broad, narrowly long-acuminate,
shallowly cordate at the base, green above, densely short-hirsute, densely short-
hispid beneath along the nerves and veins; inflorescences umbelliform, few-flowered,
the peduncle 2.5 cm. long, the pedicels 1.5 cm. long, fulvous-hispidulous; calyx
lobes oblong-ovate, 7-8 mm. long, acuminate, brownish-pilose; corolla rotate, white
with green veins, glabrous or minutely papillose, the lobes 12 mm. long, broadly
ovate, rounded at the apex, spreading; gynostegium 4 mm. high; corona urceolate,
3 mm. high, deeply much cleft in the upper half, slightly inflated at the base and
minutely papillose.
Matelea sylvicola L. Wms. Fieldiana, Bot. 32: 57. 1968.
In forest near Pansamala, Alta Verapaz, 1,200 m. (type, Tuerck-
heim 680). Endemic.
464 FIELDIANA: BOTANY, VOLUME 24
Herbaceous vines of unknown size, the stems slender, glabrous or with obscure
pubescence in lines, internodes to about 20 cm. long; leaves oblong-ovate, abruptly
long acuminate, shallowly cordate at the base, completely glabrous, 7-12 cm. long
and 3-5.5 cm. broad, the petiole 3-4 cm. long, bearing at its apex on base of leaf
about 5 small digitiform glands; inflorescence axillary, few-flowered, subumbelli-
form, peduncles 6-8 cm. long, pedicels slender and to about 3 cm. long, both ob-
scurely puberulent; flowers relatively large, about 4 cm. in diameter; calyx 6-8 mm.
long, divided nearly to the base, the lobes lanceolate, acuminate, puberulous out-
side; corolla rotate, divided nearly to the base, obscurely puberulent outside, inside
prominently pilose toward the center and puberulent, lobes narrowly lanceolate,
acute or acuminate, about 15 mm. long and about 6 mm. broad at the base; corona
fleshy, low, about 10-lobate with principal internal lobes opposite sinuses of corolla;
stigma 2-3 mm. broad, depressed, white; follicles unknown.
It is curious that this species was not described by Donnell-Smith
or Standley since it was passed by both of them as Gonolobus pictu-
ratus Hemsl. or Matelea picturata (Hemsl.) Woodson, a quite dif-
ferent species which is beautifully illustrated in Biologia Centrali-
Americana. The species does belong in subgenus Heliostemma
(Woodson, Ann. Mo. Bot. Gard. 28: 233. 1941) and is most closely
allied to Matelea pilosa (Benth.) Woodson, a Mexican species. Super-
ficially it is easily distinguished by the much larger, glabrous leaves
and the smaller and narrow calyx lobes. The "bosses" on the corona
are not digitate.
Matelea tenuis Woodson, Ann. Mo. Bot. Gard. 28: 279. 1941.
Known only from the type, Izabal, near Quirigua, 75-225 m.,
Standley 2^036.
Plants scandent, the stems slender, inconspicuously pilosulous; leaves on peti-
cles 5-7 mm. long, elliptic-oblong, 5-7 cm. long, 1.5-2.3 cm. broad, acuminate,
obtuse at the base, thin-membranaceous, glabrous; inflorescence extra-axillary,
racemiform, few-flowered, the peduncle 3-5 mm. long, the pedicels 5 mm. long,
minutely papillose-puberulent, the flowers green; calyx lobes broadly ovate, acute,
2.5 mm. long, papillose-puberulent; corolla almost rotate, 1 cm. broad, the lobes
ovate, obtuse, minutely puberulent outside and inside; corolla patelliform, ob-
scurely 5-lobate, 4 mm. broad; gynostegium borne on a stipe 2 mm. long; stigma
pentagonal, 2 mm. broad.
Matelea tikalana Lundell, Phytologia 16: 446. 1968. Marsdenia
tikalana Lundell, Wrightia 4: 49. 1968.
Forest edges and openings, known only from around ruins at
Tikal, Pete"n (type, Lundell 15763), endemic.
Slender vines almost completely glabrous. Stems to about 2 mm. in diameter,
glabrous, internodes 20-30 cm. long; leaves oblong-lanceolate to broadly oval,
acute or short acuminate, narrowly cordate at the base, glabrous or very obscurely
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 465
puberulent at the base below, submembranaceous, with 4-6 pairs of lateral nerves,
provided with a bipartite gland with several "bosses" at juncture with the petiole,
blade 12-15 cm. long and 5-8.5 cm. broad, petioles rather stout, glabrous, 2-3.5 cm.
long; inflorescence extra axillary, l-(or 2-)flowered, peduncle none, pedicels ob-
scurely dark glandular, about 5-10 mm. long; calyx divided nearly to the base,
obscurely punctate glandular, the lobes lanceolate, acuminate, about 5 mm. long
and 1 mm. broad; corolla campanulate, or rotate at an thesis, lobate to near the
gynostegium, obscurely puberulent on the inner surface, about 17-20 mm. broad,
the lobes ovate-lanceolate, acuminate, appearing marginate due to strong lateral
nerves, 7-9 mm. long and 3-3.5 mm. broad, corona adnate to the corolla, fleshy
and five lobate, saccate (not vesicular) and with "bosses" within; stigma 5-angu-
late, the pollinia apparently erect, fusiform but flattened on the inner surface;
follicles unknown.
An unusual species in the large, essentially glabrous leaves and
the much reduced inflorescence without a peduncle.
Matelea tuerckheimii (Donn.-Sm.) Woodson, Ann. Mo. Bot.
Card. 28: 234. 1941. Gonolobus tuerckheimii Donn.-Sm. Bot. Gaz.
44: 116. 1907 (type from Alta Verapaz, Tuerckheim II 1301}. Vince-
toxicum trichoneuron Standl. in Yuncker, Field Mus. Bot. 17: 388.
1938 (type from Honduras).
Twining over shrubs in deciduous or pine forest regions, mostly
at less than 1,300 m.; Pete"n; Alta Verapaz. British Honduras;
Honduras.
A slender vine, the stems puberulent and short-hirsute with fulvous, spreading
or refracted hairs; leaves on petioles 1-2 cm. long, membranaceous, oblong or lance-
oblong, 7-12 cm. long, 3-4 cm. broad, long-acuminate, broadly rounded or sub-
truncate at the base, green above, appressed-pilose on the nerves, paler beneath,
softly pilose with short spreading brownish hairs; inflorescence umbellate, the um-
bels 1-9-flowered, the peduncle short and stout, the pedicels 1.5 cm. long or shorter;
calyx green, 7-8 mm. long, pilosulous, lobate almost to the base, the lobes oblong-
lanceolate, long-acuminate; corolla yellow and brown-reticulate within, almost
2 cm. broad, rotate, minutely pilosulous outside, glabrous within, lobate to the
middle, the lobes orbicular, rounded at the apex; corona simple, the lobes semi-
orbicular; gynostegium 1.5 mm. long; ovaries glabrous; stigma 2.5 mm. broad, the
center depressed.
Matelea velutina (Schlecht.) Woodson, Ann. Mo. Bot. Gard.
28: 234. 1941; L. Wms. Fieldiana, Bot. 32: 59. 1968. Gonolobus
velutinus Schlecht. Linnaea 8: 521. 1833. Vincetoxicum velutinum
Standl. Contr. U. S. Nat. Herb. 23: 1189. 1924. V. gentlei Lundell &
Standl. ex Standl. Field Mus. Bot. 17: 269. 1924 (type from British
Honduras, Lundell 1799). Matelea gentlei Woodson, I.e.
Usually at edges of forests from near sea level to 2,000 m.; Chi-
maltenango; Peten. Southern Mexico; British Honduras.
466 FIELDIANA: BOTANY, VOLUME 24
A large, herbaceous or suffrutescent vine, the stems, petioles, and inflorescence
densely glandular-pubescent and also hirsute with long spreading hairs; leaves
membranaceous, on long slender petioles, broadly ovate to rounded-ovate, mostly
6-16 cm. long, short- acuminate or more often abruptly and shortly cuspidate-
acuminate, deeply cordate at the base, very densely velutinous-pilosulous on both
surfaces, somewhat paler and yellowish beneath; peduncles longer or shorter than
the petioles, few-many-flowered, the pedicels slender, unequal, mostly about 2 cm.
long; calyx deeply lobate, the lobes whitish or pale green, ovate or broadly ovate,
4-10 mm. long, acute or acuminate, densely pubescent and sparsely long-pilose;
ccrolla yellowish green, about 3 cm. broad, densely short-pilose outside, glabrous
and reticulate- veined within, the lobes suborbicular, minutely retuse at the apex;
corona annuliform, the outer margin entire, bearing within 5 somewhat inflated
lobes adnate to the gynostegium; immature fruit 6-7 cm. long, densely glandular-
pubescent, bearing very numerous, slender, fleshy, spine-like tubercles 1 cm. long
or even longer.
Matelea velutina is a fairly widely distributed species covering an
unusually large altitudinal range. Lundell and his associates have
made about a dozen collections of the species around Tikal in re-
cent years.
Matelea violacea Woodson, Ann. Mo. Bot. Gard. 28: 280. 1941.
Known definitely only from the type, San Marcos, southeastern
slopes of Volcan de Tacana, 2,000-3,000 m., Steyermark 364-4.5.
A suffrutescent vine, the stems fulvous-pilose with short hairs; leaves mem-
branaceous, on petioles 3.5 cm. long or shorter, oblong-ovate, 9-13 cm. long, 4-
6 cm. broad, long-acuminate, shallowly and openly cordate at the base, densely
and softly short-pilose on both surfaces; inflorescences subsessile, few-flowered, the
peduncle 5 mm. long or less, the pedicels 1 cm. long, pilose; calyx lobes broadly
ovate, long-acuminate, fulvous-pilose; corolla rotate, pale purple and brown-reticu-
late, fulvous-pilosulous outside, minutely papillose within, the lobes broadly ovate,
obtuse, 5 mm. long, spreading; gynostegium sessile; stigma pentagonal, almost flat;
corona rotate, spreading, broadly 5-lobate, about 1 cm. in diameter, multilaciniate.
OXYPETALUM R. Brown
Mostly scandent herbs, sometimes suffrutescent, usually pubescent; leaves
opposite, generally cordate and long-petiolate; flowers medium-sized, white or
purplish, the cymes usually umbelliform, terminal or axillary; calyx 5-parted,
glandular or eglandular within at the base; corolla tube very short, campanula te
or subglobose, the limb deeply 5-lobate, the lobes usually narrow, contorted; scales
of the corona 5, shortly adnate to the corolla or free from it, distinct from the sta-
men tube or more or less connate with it, erect, usually thick, retuse, emarginate or
bifid, appendaged within or naked; stamens inserted near the base of the tube, the
filaments connate into a short tube; anthers terminated by an inflexed membrane;
pollinia solitary in each cell, oblong, pendulous; stigma with or without a bifid or
biparted beak; follicles slender cr thick, smooth or tuberculate; seeds with a coma
of soft white hairs.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 467
FIG. 127. Oxypetalum cordifolium. A, habit, X 1A\ B, flower, X 2; C, calyx,
corona and pistils, X 4; D, inner face of corona segment, X 8; E, pollinia, X20.
Fifty species or more, all American and mostly South American.
Only one has been found in Mexico and Central America.
Oxypetalum cordifolium (Vent.) Schlechter in Urban, Symb.
Antill. 1: 269. 1899. Gothofreda cordifolia Vent. Choix PL Gels. 7,
t. 60. 1803. 0. riparium HBK. Nov. Gen. & Sp. 3: 197. 1819.
468 FIELDIANA: BOTANY, VOLUME 24
Santa Rosa (Ojo de Agua, Heyde & Lux 4000); to be expected
elsewhere in Guatemala. Southern Mexico; Honduras; Costa Rica.
West Indies; South America.
A slender, chiefly herbaceous vine, finely velutinous-pilosulous on the stems,
leaves, and inflorescence; leaves slender-petiolate, soft, membranaceous, cordate-
ovate, mostly 5-10 cm. long, acuminate or cuspidate-acuminate, shallowly and
openly or deeply and narrowly cordate at the base, paler beneath; umbels few-
flowered or only 1-flowered, on long slender peduncles, often longer than the sub-
tending leaves, the very slender pedicels often longer than the peduncles; calyx
lobes linear-lanceolate or linear-subulate, densely pubescent, about equaling the
corolla tube; corolla pale yellowish, puberulent outside, glabrous or minutely papil-
lose within, the campanulate tube about 4 mm. long, the lobes linear-attenuate,
ascending or spreading, 1.5-2 cm. long; follicles about 8 cm. long, long-acuminate.
SARCOSTEMMA R. Brown
Reference: Richard W. Holm, The American species of Sarco-
stemma, Ann. Mo. Bot. Gard. 37: 477-560. 1950.
Slender vines, herbaceous or suffrutescent, pubescent or glabrous, often glau-
cous; leaves opposite; flowers rather large or small, umbellate, the umbels axillary,
usually pedunculate, generally white or greenish white, sometimes purplish, the
corolla in bud mostly depressed-globose; calyx small, 5-parted, minutely 5-glandu-
lar within, the lobes acute; corolla very broadly campanulate or subrotate, very
shallowly or rather deeply 5-lobate, the lobes contorted; outer corona submembra-
naceous, annular, adnate to the base of the corolla, its margin entire, the inner
corona of 5 scales adnate to the base of the stamen tube, their blades free, broad,
membranaceous or subcorneous, flat or concave or saccate; stamens inserted on the
base of the corolla, the filaments connate into a short tube, the anthers terminated
by an inflexed membrane; pollinia solitary in each anther cell, oblong or elongate,
pendulous from the apex; stigma flat or umbonate at the apex or bearing a short
bifid beak; follicles rather thick or often slender, acuminate, smooth.
About 35 species, in tropical and warm regions of both hemi-
spheres. Additional species are known from Mexico and Central
America.
Leaves not cordate at the base, usually obtuse or sometimes acute, linear to oblong.
S. clausum.
Leaves cordate at the base.
Corolla glabrous outside.
Leaves lanceolate-oblong to oblong-ovate; petals ciliate; highland species.
S. refractum.
Leaves ovate to orbicular; petals eciliate; lowland species S. bilobum.
Corolla densely pubescent outside S. odoratum.
Sarcostemma bilobum Hook. & Am. Bot. Beech. Voy. 438.
1841. Philibertia biloba Gray, Proc. Am. Acad. 21: 395. 1886.
FIG. 128. Sarcostemma clausum. A, habit, natural size; B, flower, X 2; C, calyx
and pistils, X 4; D, pollinia, X 25; E, seed, X 2; G, pair of leaves from another
collection to show variation, natural size.
469
470 FIELDIANA: BOTANY, VOLUME 24
Occasional in damp or dry thickets near sea level; Pete"n; Alta
Verapaz. Mexico; British Honduras; Honduras.
Trailing or twining pilosulose vines; leaves oblong-ovate to suborbicular, short
acuminate, deeply cordate with the basal lobes converging, 2-5 cm. long and 1.5-
4 cm. broad, pilosulose on both surfaces, sometimes glabrate, petiole 1-2.5 cm.
long; inflorescence umbelliform with as many as 25 flowers, peduncles 1.5-6 cm. long,
pedicels to 3 cm. long, very slender; calyx lobate to the base, the lobes linear to
ovate, about 2.5 mm. long, puberulent outside, glabrous within; corolla subrotate,
the tube about 2 mm. long, the lobes lanceolate, obtuse, reflexed, puberulent within,
ciliate, about 5-6 mm. long; corona ring fleshy, free from the vesicles, vesicles
pyramidal, widest below the middle, about 2 mm. long; follicles subfusiform, up to
8 cm. long and 2 cm. thick, puberulent, striate; seeds about 7 mm. long and coma
about 3 cm. long.
Sarcostemma clausum (Jacq.) Schult. Syst. Veg. 6: 114. 1820.
Cynanchum clausum Jacq. Stirp. Am. 1: 87, t. 60. 1763. S. cumanense
HBK. Nov. Gen. & Sp. 3: 195. 1819. S. crassifolium Dene, in DC.
Prodr. 8: 540. 1844. Philibertia crassifolia Hemsl. Biol. Cent. Am.
Bot. 2: 318. 1881. P. cumanensis Hemsl. I.e. Funastrum cumanense
Schlechter, Fedde Repert. Sp. Nov. 13: 284. 1914. F. clausum
Schlechter, I.e. 283. Bejuco de pescado; flor de Castilla (Guatemala,
fide Morales).
Wet to dry thickets or thin forest, often in thickets along streams,
1,275 m. or lower; Pete"n; Izabal; Alta Verapaz; Zacapa; Chiquimula;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Suchitepe'quez; Retal-
huleu; Solola; San Marcos. Mexico; British Honduras to El Salvador
and Panama. West Indies; South America.
A small or large, herbaceous vine, often forming large and dense tangles over
shrubs, pale and usually glaucescent, the stems glabrous or nearly so; leaves short-
petiolate, rather thick and fleshy, almost linear to oblong, 3-7 cm. long, acuminate
or cuspidate, usually obtuse or rounded at the base, glabrous, or sometimes pubes-
cent beneath; umbels long-pedunculate, few-many-flowered, the pedicels long and
slender, short-pilose; calyx densely short-pilose; corolla 10-14 mm. broad, greenish
white, rather deeply lobate, densely short-pilose outside, the lobes broadly ovate,
rounded or very obtuse and apiculate; follicles 5-6.5 cm. long, about 1 cm. broad,
long-acuminate, pubescent or glabrate.
Called "dama de noche" in British Honduras. The plant is abun-
dant in the lowlands of Guatemala, especially in rocky or gravelly
thickets along or near streams, where it often forms dense masses
that almost conceal the foliage of the supporting plants. When in
full flower it is a conspicuous vine. The flexible tough stems are used
commonly for stringing fish (hence the name bejuco de pescado) and
as a substitute for twine. Along sandbars the branches often lie flat
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 471
on the ground, sometimes running over the sand for a great distance.
Under such conditions the internodes of the stems become greatly
elongated. The leaves vary greatly in shape, being sometimes linear
or nearly so, but more often lanceolate or oblong.
Sarcostemma odoratum (Hemsl.) Holm, Ann. Mo. Bot. Gard.
37: 520. 1950. Funastrum odoratum Schlechter, Repert. Sp. Nov. 13:
286. 1914. Philibertia odorata Hemsl. Biol. Cent. Am. Bot. 2: 319.
1881. Cuchamperillo.
Moist or dry thickets, sometimes in roadside hedges, 1,500-
2,200 m.; Guatemala; Sacatepe"quez (type from Duenas, Salviri);
Huehuetenango. Endemic.
A small or large herbaceous vine, the slender stems densely pilose with short
spreading hairs; leaves membranaceous, on long slender petioles, orbicular-cordate
or ovate-cordate, 3.5-7 cm. long, 2-5.5 cm. broad, rounded or very obtuse at the
apex and cuspidate, deeply cordate at the base, densely and softly pilose on both
surfaces, sometimes white-tomentose beneath; umbels many-flowered, about equal-
ing the leaves, on long slender peduncles, the slender pedicels 1-1.5 cm. long,
densely tomentose; flowers greenish white, 10-15 mm. broad, the calyx lobes nar-
row, acute, 6 mm. long or less; corolla subrotate, deeply 5-lobate, pilose outside,
the lobes ovate-oblong, obtuse, densely puberulent within; outer corona annular,
undulate; hoods of corona 2 mm. long, inflated-cucullate, slightly longer than the
gynostegium; follicles 7-9 cm. long, about 1.5 cm. thick, densely pilose, narrowly
long-acuminate.
Sarcostemma refractum (Donn.-Sm.) L. Wms. Fieldiana, Bot.
32: 59. 1968. Philibertia refracta Donn.-Sm. Bot. Gaz. 18: 207. 1893.
Wooded slopes in the mountains, 1,800-2,300 m.; Huehuetenango;
Quiche" ; Sacatepe"quez. Southern Mexico.
Slender herbaceous vines. Stems somewhat pilose, to 2 mm. in diameter;
leaves oblong-lanceolate to oblong-ovate, shallowly cordate with basal lobes diverg-
ing, acute or acuminate at the apex, sparsely pilose or ciliate to completely gla-
brous, blade 2.5-5 cm. long and 1-2.5 cm. broad; inflorescence umbelliform, many-
flowered, peduncle 3-7 cm. long, pedicels filiform, 1.5-2.5 cm. long, crisped puberu-
lent; calyx divided to the base, pilose pubescent outside and ciliate, glabrous
within, linear-lanceolate, acute, about 4 mm. long and 1 mm. broad; corolla rotate
or the lobes reflexed, 5-7 mm. long, divided to near the base, glabrous outside,
puberulent within especially at the base of the lobes, tube short, about 1-1.5 mm.
long, lobes narrowly ovate to ovate-obtuse, obscurely ciliate on one margin, 4-
5 mm. long and 3-4 mm. broad; corona ring undulate-lobate, about 0.5 mm. high,
adnate to the base of the vesicles; vesicles about 2.5-3 mm. long; fruits unknown.
This species was included in S. bilobum by Holm in his monograph
of the genus. However the distinctively shaped leaves, difference in
472 FIELDIANA: BOTANY, VOLUME 24
corolla although not outstanding, and the highland distribution in-
dicates that it should be treated as a separate species.
Some of the species of Stapelia are cultivated rarely in Guate-
malan gardens. They are natives of South Africa, low, succulent
plants with more or less 4-angulate, leafless stems, and often very
large, luridly colored, usually very fetid flowers.
INDEX
Allamanda, 337
Ammi, 23
Anagallis, 201
Anethum, 24
Apium, 25
Apocynaceae, 334
Aralia, 2
Araliaceae, 1
Arbutus, 89
Arctostaphylos, 91
Ardisia, 136
Arracacia, 26
Asclepiadaceae, 407
Asclepias, 409
Aspidosperma, 338
Beaumontia, 340
Befaria, 94
Berula, 32
Blepharodon, 416
Buddleia, 277
Bumelia, 213
Cameraria, 341
Cavendishia, 97
Centaurium, 303
Centella, 34
Centunculus, 203
Chelonanthus, 307
Chimaphila, 83
Chrysophyllum, 216
Clethra, 74
Clethraceae, 74
Coaxana, 34
Conium, 36
Contortae, 263
Coriandrum, 37
Cornaceae, 67
Cornus, 67
Couma, 342
Coutoubea, 308
Cryptostegia, 418
Curtia, 310
Cynanchum, 419
Cynoctonum, 284
Daucus, 37
Deherainia, 128
Dendropanax, 4
Didymopanax, 8
Diospyros, 244
Dipholis, 219
Disterigma, 101
Donnellsmithia, 39
Ebenaceae, 244
Ebenales, 211
Echites, 345
Empedoclesia, 104
Enantiophylla, 41
Ericaceae, 88
Ericales, 73
Eryngium, 43
Eustoma, 312
Fernaldia, 348
Fischeria, 426
Foeniculum, 48
Forestiera, 264
Forsteronia, 350
Fraxinus, 266
Garrya, 70
Garryaceae, 70
Garryales, 69
Gaultheria, 104
Gelsemium, 286
Gentiana, 312
Gentianaceae, 302
Gentlea, 156
Gonolobus, 426
Halenia, 316
Haplophyton, 354
Hedera, 9
Hoya, 441
Hydrocotyle, 49
Jacquinia, 129
Jasminum, 269
Labidostelma, 463
Lacmellea, 355
Laubertia, 356
Leiphaimos, 319
Leucothoe, 108
Ligustrum, 271
Limonium, 208
Linociera, 273
Lisianthus, 321
Lochnera, 358
Loganiaceae, 276
Lysimachia, 205
Macleania, 110
473
474
INDEX
Malouetia, 359
Mandevilla, 361
Manilkara, 222
Marsdenia, 442
Mastichodendron, 228
Matelea, 450
Mesechites, 366
Micropholis, 232
Micropleura, 53
Monotropa, 86
Monotropaceae, 86
Myrrhidendron, 56
Myrsinaceae, 135
Nerium, 366
Nothopanax, 10
Nymphoides, 326
Odontadenia, 368
Oleaceae, 264
Oreomyrrhis, 56
Oreopanax, 11
Osmanthus, 274
Osmorhiza, 57
Ottoa, 58
Oxypetalum, 466
Parathesis, 160
Pernettya, 112
Petroselinum, 58
Pimpinella, 59
Plocosperma, 288
Plumbaginaceae, 207
Plumbaginales, 207
Plumbago, 209
Plumeria, 371
Polypremum, 290
Pouteria, 233
Prestonia, 374
Primulaceae, 200
Primulales, 127
Prionosciadium, 60
Pyrola, 84
Pyrolaceae, 81
Rapanea, 190
Rauvolfia, 380
Rhabdadenia, 383
Rhododendron, 115
Rhodosciadium, 61
Samolus, 207
Sanicula, 62
Sapotaceae, 211
Sarcostemma, 468
Satyria, 116
Schultesia, 328
Sciadodendron, 19
Spananthe, 62
Sphyrospermum, 118
Spigelia, 292
Stapelia, 472
Stemmadenia, 385
Stephanotis, 389
Strychnos, 297
Stylogyne, 192
Styracaceae, 258
Styrax, 258
Symplocaceae, 251
Symplocos, 252
Synardisia, 195
Syringa, 275
Tabernaemontana, 389
Tauschia, 64
Theophrastaceae, 127
Thevetia, 394
Tintinnabularia, 400
Tonduzia, 401
Umbelliferae, 21
Umbelliflorae, 1
Urechites, 402
Vaccinium, 120
Vallesia, 405
Vinca, 405
Vincetoxicum, 458
Voyria, 331
Yunckeria, 197
Publication 1070
UNIVERSITY OF ILLINOIS-URBANA