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
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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.
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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.
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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.
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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.
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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.
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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.
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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.
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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.
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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
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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.
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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 culti