UNIVtRSi'i t OF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
JUL 2^19841
FLORA OF GUATEMALA
PAUL C. STANDLEY
and
LOUIS O. WILLIAMS
7 1971
FIELDIANA: BOTANY
VOLUME 24, PART XI, NUMBERS 1 to 3
Published by
FIELD MUSEUM OF NATURAL HISTORY
June 16, 1975
DEC 16 1975
FLORA OF GUATEMALA
PART XI, NOS. 1 TO 3
FLORA OF GUATEMALA
PAUL C. STANDLEY
The Late Curator of the Herbarium
Field Museum of Natural History
and
LOUIS O. WILLIAMS
Curator Emeritus, Department of Botany
Field Museum of Natural History
FIELDIANA: BOTANY
VOLUME 24, PART XI, NUMBERS 1 to 3
Published by
FIELD MUSEUM OF NATURAL HISTORY
June 16, 1975
Library of Congress Catalog Card Number: 48-3076
US ISSN 0015-0746
PRINTED IN THE UNITED STATES OF AMERICA
V 2-4 : )
CONTENTS
Family in Part XI, Numbers 1 to 3
PAGE
Rubiaceae... ...1
LIST OF ILLUSTRATIONS
In systematic order with the tribes indicated
PAGE
TRIBE I. Condamineae.
Fig. 1. Portlandia guatemalensis 207
Fig. 2. Pogonopus speciosus 208
TRIBE II. Oldenlandieae
Fig. 3. Oldenlandia corymbosa 209
Fig. 4. Houstonia serpyllacea 210
TRIBE III. Rondeletieae
Fig. 5. Rondeletia buddleioides 211
Fig. 6. Lindenia rivalis 212
Fig. 7. Deppeaflava 213
Fig. 8. Deppea grandiflora 214
Fig. 9. Pinarophytton flavum 215
Fig. 10. Steyermarkia guatemalensis 216
TRIBE IV. Cinchoneae
Fig. 11. Bouvardia longiflora 217
Fig. 12. Manettia recttnata 218
Fig. 13. Alseis yucatanensis 219
Fig. 14. Hillia tetrandra 220
Fig. 15. Cosmibuena matudae 221
Fig. 16. Calycophyllum candidissimum 222
Fig. 17. Exostema mexicanum 223
Fig. 18. Hintonia standleyana 224
Fig. 19. Coutarea hexandra 225
TRIBE V. Naucleeae
Fig. 20. Uncaria tomentosa 226
Fig. 21. Cephalanthus occidentalis 227
TRIBE VI. Mussaendeae
Fig. 22. Isertia haenkeana 228
Fig. 23. Gonzalagunia thyrsoidea 229
Fig. 24. Coccocypselum cordifolium 230
Fig. 25. Sabicea villosa 231
Fig. 26. Sommera guatemalensis 232
Fig. 27. Pentagonia macrophytta 233
VI
TRIBE VII. Gardenieae
Fig. 28. Randia standleyana 234
Fig. 29. Posoqueria latifotia 235
Fig. 30. Gempa caruto 236
Fig. 31. Amaiouia corymbosa 237
Fig. 32. Alibertia edulis 238
Fig. 33. Bertiera guianensis 239
TRIBE VIII. Hamelieae
Fig. 34. Hamelia barbata 240
Fig. 35. Hoffmannia cauUflora 241
Fig. 36. Hoffmannia sessUifolia 242
TRIBE IX. Guettardeae
Fig. 37. Guettarda macrosperma 243
Fig. 38. Antirhea lucida 244
Fig. 39. Pittoniotis trichantha 245
Fig. 40. Anisomeris brachypoda 246
Fig. 41. Machaonia lindeniana 247
TRIBE X. Chiococceae
Fig. 42. Chiococca semipilosa 248
Fig. 43. Asemnantha pubescens 249
Fig. 44. Chione guatemalensis 250
TRIBE XI. Ixoreae
Fig. 45. Coffea arabica 251
Fig. 46. Ixora nicaraguensis 252
TRIBE XII. Psychotrieae
Fig. 47. Psychotria chiapensis 253
Fig. 48. Psychotria marginata 254
Fig. 49. Palicourea galeottiana 255
Fig. 50. Rudgea cornifotia 256
Fig. 51. Declieuxia fruticosa var. mexicana 257
Fig. 52. Cephaelis glomerulata 258
TRIBE XIII. Anthospermeae
Fig. 53. Mitchella repens 259
TRIBE XIV. Coussareae
Fig. 54. Coussarea imitans 260
Fig. 55. Faramea occidentatis 261
Fig. 56. Faramea standleyana 262
TRIBE XV. Morindeae
Fig. 57. Appunia guatemalensis 263
Fig. 58. Morinda yucatanensis 264
TRIBE XVI. Spermacoceae
Fig. 59. Richardia scabra 265
VII
Fig. 60. Ernodea littoralis 266
Fig. 61. Crusea calocephala 267
Fig. 62. Diodia
D. brasiliensis 268
D. sarmentosa 268
Fig. 63. Hemidiodia ocymifolia 269
Fig. 64. Borreria laevis 270
Fig. 65. Spermacoce riparia 271
Fig. 66. Mitracarpus hirtus 272
TRIBE XVII. Rubieae
Fig. 67. Relbunium hypocarpium 273
Fig. 68. Didymaea
D. australis 274
D.hispidula 274
D. microflosculosa 274
Flora of Guatemala - Part XI, Numbers 1 to 3
RUBIACEAE. Madder Family
PAUL C. STANDLEY AND Louis 0. WILLIAMS
Reference: Standley, No. Am. Fl. 32: 1-300. 1918-34 (this
treatment covers only part of the family, but is the latest account
treating the genera and species of Central America).
Herbs, shrubs or trees, erect, prostrate, or rarely scandent, sometimes armed with
spines; leaves simple, opposite or verticillate, entire (in all Guatemalan
representatives except one), rarely pinnatifid; stipules present, interpetiolar or
intrapetiolar, persistent or deciduous, entire or often lobate or dentate or setiferous,
frequently connate to form a sheath, rarely (in Galieae) foliaceous and resembling
leaves; inflorescence various, usually cymose, sometimes capitate, often paniculate,
the hypanthia rarely adnate and forming a syncarp; flowers typically perfect,
sometimes unisexual, commonly regular and symmetric, frequently dimorphous;
hypanthium adnate to the ovary; calyx cupular, tubular, or nearly obsolete,
persistent or deciduous, entire, dentate, or lobate, the lobes often unequal, one of
them sometimes foliaceous; corolla gamopetalous and funnelform, salverform,
campanula* e, rotate, or rarely urceolate or tubular, glabrous or pubescent within, the
limb usually symmetric, its lobes valvate, imbricate, or contorted in bud; stamens
usually as many as the corolla lobes and alternate with them, inserted in the tube or
throat of the corolla; filaments short or elongate, often suppressed; anthers usually
oblong-linear, 2-celled, dehiscent by anterior or lateral slits or rarely by pores,
dorsifixed or basifixed; disk annular, pulvinar, hemispheric, or conic; ovary 1-10-
celled, the style short or elongate, simple or 2-10-fid, the branches filiform, linear, or
spatulate; placentae affixed to the septum or to the interior angle of the cell, or
basilar, or pendulous from the apex of the cell; ovules solitary, geminate, or
numerous, superficial or immersed in the placentae, erect, horizontal, ascending, or
pendulous; fruit capsular, baccate, or drupaceous, or of dehiscent or indehiscent
cocci, 2-10-celled, rarely 1 -celled; seeds variable in form and size, the testa usually
membranaceous or coriaceous, smooth or roughened, often winged or appendaged.
One of the largest families of plants, represented in almost all
parts of the earth, but most abundantly in the tropics. In most
countries of Central and South America it stands among the first
three or four families in number of species. A very few genera
1
2 FIELDIANA: BOTANY, VOLUME 24
besides those listed here are represented in southern Central
America, particularly Costa Rica and Panama.
The senior author of this flora specialized in the family for
many years and knew a great deal about the American Rubiaceae.
He never found the time to study the tribal arrangement of the
genera and, in fact, had little interest in the tribes other than as
convenient receptacles for the genera as he knew them.
The key to the tribes presented here is much like that in the
preliminary manuscript that Standley wrote 30 years ago. Names of
tribes, in two cases, have been changed to make them conform to
correct nomenclature and the Hamelieae has been separated out of
the Gardenieae. Several genera have been added to the flora since
Standley's day and many additional species are added to the known
flora, based upon our intensive collections in Guatemala over the
last 20 years.
KEY TO THE TRIBES AND GENERA OF RUBIACEAE
Ovules more than one in each cell. (Subfam. Cinchonoideae).
Fruit fleshy, baccate.
Corolla lobes valvate in bud VI. Mussaendeae.
Corolla lobes imbricate or contorted in bud.
Corolla lobes imbricated in the bud VIII. Hamelieae.
Corolla lobes contorted in the bud VII. Gardenieae.
Fruit dry, capsular.
Flowers arranged in very dense and compact, spherical heads; shrubs or trees.
V. Naucleeae.
Flowers variously disposed, but never in spherical heads.
Seeds winged or appendaged, vertically imbricate IV. Cinchoneae.
Seeds not winged or, if so, horizontal.
Corolla lobes imbricate or contorted in bud; shrubs or trees.
III. Rondeletieae.
Corolla lobes valvate in bud.
Seeds horizontal, usually very numerous; stipules entire or biparted; large
shrubs or trees with usually large leaves I. Condamineae.
Seeds imbricate, vertical, usually few; stipules often setose-laciniate; herbs
or very small shrubs with small leaves II. Oldenlandieae.
Ovules solitary in the cells: Subfam. Rubioideae, except Naucleeae).
Seeds pendulous in the cell, the radicle superior; shrubs or trees.
Flowers in compact globose heads V. Naucleeae.
Flowers never in globose heads.
Stamens inserted in the throat of the corolla IX. Guettardeae.
Stamens inserted at the base of the corolla tube or on disc X. Chiococceae.
Seeds ascending, the radicle inferior; plants woody or herbaceous.
Corolla lobes contorted in bud; shrubs or trees XI. Ixoreae.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 3
Corolla lobes valvate in bud.
Ovules basilar, attached at the base of the cell; mostly shrubs or trees.
Ovary 1 -celled, or 2-celled but with a very thin septum, the fruit 1 -seeded.
XIV. Coussareae.
Ovary 2-celled, or the cells rarely more numerous, the septum thick.
Stamens usually inserted in the throat of the corolla; flowers perfect.
XII. Psychotrieae.
Stamens usually inserted at the base of the corolla tube; flowers often
unisexual XIII. Anthospermeae.
Ovules lateral, attached to the septum of the cell.
Stipules neither foliaceous nor setiferous; trees or large shrubs; flowers
confluent by the hypanthia to form a dense head XV. Morindeae.
Stipules either foliaceous or setiferous; herbs or very small shrubs; flowers
never confluent by the hypanthia.
Stipules setiferous XVI. Spermacoceae.
Stipules foliaceous, similar to the leaves XVII. Rubieae.
I. CONDAMINEAE
Calyx lobes all similar Portlandia.
Calyx lobes dissimilar, one of them expanded into a large red blade Pogonopus.
II. OLDENLANDIEAE
Seeds angulate; plants (in Guatemalan species) annual Oldenlandia.
Seeds concavo-convex, not angulate; plants (in Guatemalan species) perennial.
Houstonia.
III. RONDELETIEAE
Corolla lobes imbricated in bud.
Plants herbaceous or nearly so, small and low; corolla almost rotate.
PinarophyUon.
Plants tall shrubs or small trees, corolla with an elongate tube.
Seeds very large, usually 1 cm. long or more, horizontal Sickingia.
Seeds small or minute, winged or exalate.
Flowers very large, the tube of the corolla about 5 cm. long Eizia.
Flowers relatively small, the corolla tube seldom as much as 2 cm. long.
Rondeletia.
Corolla lobes contorted in bud.
Leaves finely lineolate between the veins; flowers very small Deppea flava.
Leaves not lineolate between the veins.
Tube of the corolla short, shorter than the lobes; corolla very small.
Corolla glabrous within, 4-parted Deppea.
Corolla villous within, 5-parted Elaeagia.
Tube of the corolla 2-several times as long as the lobes; corolla 5-16 cm. long.
Plants erect shrubs with leafy branches; corolla white Lindenia.
Plants acaulescent herbs, the flowers borne on naked scapes; corolla pink.
Steyermarkia.
4 FIELDIANA: BOTANY, VOLUME 24
IV. ClNCHONEAE
Plants twining, herbaceous Manettia.
Plants erect, usually shrubs or trees.
Calyx lobes dissimilar, one of them expanded into a large leaflike, white or cream-
colored blade Calycophyllum.
Calyx lobes all similar.
Plants epiphytic; leaves fleshy when fresh, coriaceous when dry; flowers large,
white.
Seeds winged at either end Cosmibuena.
Seeds with a tuft of hair at one end Hillia.
Plants usually terrestrial; leaves herbaceous.
Inflorescence spikelike, greatly elongate and densely many-flowered; trees.
Alseis.
Inflorescence never spikelike.
Corolla lobes valvate in bud.
Flowers 4-parted; capsule subglobose Bouvardia.
Flowers 5-parted; capsule oblong to clavate Cinchona.
Corolla lobes imbricate or contorted in bud.
Anthers sessile Blepharidium.
Anthers borne on evident, often elongate filaments.
Stamens exserted Exostema.
Stamens not exserted.
Corolla symmetric; capsule not or scarsely compressed.
Corolla funnelform without a narrow tube Hintonia.
Corolla salverform, the tube narrow Balmea.
Corolla conspicuously asymmetric; capsule strongly compressed.
Coutarea.
V. NAUCLEEAE
Ovules solitary in the cell; plants erect, unarmed Cepkalanthus.
Ovules numerous in the cell; plants scandent by recurved spines Uncaria.
VI. MUSSAENDEAE
Leaf tissue finely lineolate between the veins.
Calyx spathaceous or lobate, the lobes not foliaceous; leaves very large and
leathery Pentagonia.
Calyx deeply lobate, the lobes foliaceous, green; leaves relatively small,
membranaceous Sommera.
Leaf tissue not lineolate between the veins.
Plants erect shrubs or small trees.
Inflorescence a spikelike panicle; corolla white or pink Gonzalagunia.
Inflorescence not a spikelike panicle.
Inflorescence a terminal thyrsiform panicle; flowers orange-red Isertia.
Inflorescence an axillary, subcapitate or short pedunculate cymose panicle
hardly longer than the petioles Sabicea.
Plants herbaceous or nearly so, sometimes scandent.
Ovary 2-celled; plants herbaceous, prostrate Coccocypselum.
Ovary 3-5-celled; plants herbaceous or suffrutescent, scandent Sommera.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
VII. GARDENIEAE
Plants armed with spines (in Guatemalan species) Randia.
Plants unarmed.
Inflorescence a many-flowered open thyrsiform panicle; seeds minute, foveolate.
Bertiera.
Inflorescence various but never paniculate, the flowers few or solitary; seeds large,
the testa smooth or fibrous.
Corolla somewhat irregular, curved in bud, the tube greatly elongate, 12-16 cm.
long Posoqueria.
Corolla regular or nearly so, the tube much shorter.
Flowers unisexual.
Stipules united to form a conic cap, this deciduous above a circular slit.
Pistillate flowers capitate or cymose Amaioua.
Pistillate flowers usually solitary Duroia.
Stipules not united to form a cap -Alibertia.
Flowers perfect.
Corolla tube villous in both throat and base; ovary 2-celled; native trees.
Genipa.
Corolla tube villous in the throat or the base but not in both; ovary 1-celled;
cultivated shrubs Gardenia.
VIII. HAMELIEAE
Ovary 4-5-celled Hamelia.
Ovary 2-celled Hoffmannia.
IX. GUETTARDEAE
Fruit separating at maturity into 2-cocci; calyx lobes 4-5, persistent; corolla lobes
imbricate „ Machaonia.
Fruit drupaceous, not separating into cocci at maturity.
Corolla lobes valvate in bud or but obscurely imbricate.
Corolla lobes corniculate-appendaged outside near the apex; basal lobes of the
anthers acute or attenuate Chomelia.
Corolla lobes not appendaged; basal lobes of the anthers obtuse Anisomeris.
Corolla lobes strongly imbricate in bud, 1 or 2 of them exterior.
Anthers long exserted; cymes paniculate; calyx persistent Pittoniotis.
Anthers included; inflorescence not cymose paniculate.
Calyx deciduous; plants usually abundantly pubescent Guettarda.
Calyx persistent upon the fruits; plants glabrous Antirhea.
X. CHIOCOCCEAE
Flowers 4-parted.
Inflorescence an axillary fascicle; plants abundantly pubescent A.semnantha.
Inflorescence a cymose-corymbose usually terminal panicle; plants sparsely
pubescent AUenanthus.
Flowers 5-parted, usually in racemes, cymes or corymbs.
Corolla lobes imbricate in bud; fruit not compressed Chione.
Corolla lobes valvate in bud; fruit usually strongly compressed Chiococca.
6 FIELDIANA: BOTANY, VOLUME 24
XI. IXOREAE
Bractlets connate and calyx-like; calyx usually truncate Coffea.
Bractlets distinct, not calyx-like; calyx lobate Ixora.
XII. PSYCHOTRIEAE
Plants prostrate and creeping, herbaceous; leaves cordate; flowers capita ie ..Geophila.
Plants never prostrate and creeping, usually woody; leaves rarely if ever cordate.
Inflorescence an involucrate, usually solitary head, or the heads sometimes
branched, the outer bracts large and often brightly colored Cephaelis.
Inflorescence not of involucrate heads.
Carpels of the fruit laterally compressed, the fruit didymous; low herbs.
Declieuxia.
Carpels of the fruit not laterally compressed, the fruit not didymous; trees or
shrubs.
Seeds with an incurved ventral surface; stipules pectinate- lobate or with
setiform dorsal appendages Rudgea.
Seeds not with an incurved ventral surface; stipules entire or bilobate, not
appendaged.
Corolla tube straight, not gibbous at the base; branches of the inflorescence
usually green; corolla usually white, dull greenish yellow, or greenish.
Psychotria.
Corolla tube elongate, often more or less curved, gibbous at the base;
branches of the inflorescence usually pale red or yellow; corolla often
bright yellow or red Palicourea.
XIII. ANTHOSPERMEAE
Stamens inserted in the throat of the corolla; style branches 4; fruits connate by
pairs Mitchella.
Stamens inserted at or near the base of the corolla tube; style branches 2; fruits
distinct Nertera.
XIV. COUSSAREAE
Seeds vertical; ovules connate, borne upon a common basal column Coussarea.
Seeds horizontal; ovules separate in a 1-celled ovary, collateral, basal Faramea.
XV. MORINDEAE
Flowers confluent in fruit to form a syncarp Morinda.
Flowers distinct Appunia.
XVI. SPERMACOCEAE
Fruit an indehiscent drupe; shrubs of the seashore Ernodea.
Fruit dehiscent, separating into cocci or circumscissile.
Fruit not separating into cocci, top of fruit circumscissile Mitracarpus.
Fruit separting into cocci, these not circumscissile.
Cocci 3-4 Richardia.
Cocci 2.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 7
Cocci indehiscent.
Calyx lobate, the lobes elongate-subulate; cocci separating from a persistent
axis Crusea
Calyx 2-10-dentate; fruit without a persistent axis Diodia.
Cocci, at least some of them, dehiscent.
Cocci opening only at the base; flowers axillary Hemidiodia.
Cocci opening at the apex; flowers axillary or in terminal heads.
Cells of the fruit alike, both opening; flowers axillary or in terminal heads.
Borreria
Cells of the fruit dissimilar, one opening, the other remaining closed;
flowers all or chiefly axillary Spermacoce.
XVII. RUBIEAE
Flowers each surrounded by a calyx-like involucre of leaflike bracts Relbunium
Flowers not involucrate.
Leaves opposite, broad, mostly deltoid-ovate Didymaea.
Leaves verticillate, linear to ovate Galium.
ALIBERTIA A. Richard
Unarmed shrubs or trees, usually glabrous or nearly so; leaves mostly coriaceous,
sessile or petiolate; stipules interpetiolar, connate at the base, acute; flowers rather
small, unisexual, terminal, sessile, the staminate fasciculate, the pistillate usually
solitary; hypanthium hemispheric or globose, the calyx short or tubular, truncate or
dentate; corolla coriaceous, salverform, the tube cylindric, sometimes contracted at
the throat, the throat glabrous or villous, the limb 4-8-lobate, the lobes short or
elongate, obtuse or acuminate, contorted; stamens 4-8, inserted in the corolla tube,
the filaments short or none, the anthers linear, dorsifixed, included; ovary 2-8-celled,
the style with 2-8 branches or with an elongate fusiform stigma; ovules few or
numerous, pauciseriate or multiseriate, often immersed, the placentae affixed to the
interior angle of the cell; fruit baccate, usually large, 2-8-celled, with thin septae, the
placentae pulpy; seeds usually numerous, large, compressed, vertical, the testa
coriaceous or fibrous.
Numerous species occur in South America, but only one other
is known from North America, A. garapatica Schum., of Panama.
Alibertia edulis (L. Rich.) A. Rich, ex DC. Prodr. 4: 443. 1830.
Genipa edulis L. Rich. Act. Soc. Hist. Nat. Paris 1: 107. 1792.
Albondiga (Jutiapa); guayaba de monte; guayabillo; guabillo.
Figure 32.
Common in the tierra caliente of both slopes, ascending rarely
to 1,500 m., usually at much lower elevations, in thickets or wet
forest, often on rocky stream banks; Peten; Izabal; Chiquimula;
8 FIELDIANA: BOTANY, VOLUME 24
Jutiapa; Santa Rosa; Escuintla. Southern Mexico; British
Honduras to Panama; ranging southward to the Amazon Basin.
Usually a shrub of 1-4 m. but sometimes as much as 7.5 m. tall and tree-like,
almost glabrous throughout; stipules lance-triangular or deltoid, 8-18 mm. long, acute
or attenuate-acuminate, striate; petioles 2-12 mm. long; leaf blades lance-oblong to
ovate-oblong or oval-ovate, 6-20 cm. long, 2-7 cm. broad, usually short-acuminate, at
the base acute to rounded, somewhat coriaceous, lustrous above, somewhat paler
beneath and commonly short-barbate in the axils of the nerves; staminate
inflorescence usually 6-8-flowered, the flowers white, fragrant, sessile; hypanthium
turbinate, the calyx denticulate, glabrous or puberulent, the corolla 2.5 cm. long,
minutely sericeous outside, the normally 5 lobes ovate or lance-oblong, acute or
acuminate, sericeous within, half as long as the tube or longer; pistillate flowers
solitary, sessile or subsessile, the ovary 4-celled, the calyx 4 mm. long, puberulent or
glabrous, denticulate, the corolla 2-3 cm. long, the lobes about as long as the tube;
fruit subglobose, about 2.5 cm. in diameter, green or yellowish, smooth; seeds
suborbicular, brownish, 5 mm. broad, finely striate.
Called "torolillo" in El Salvador; "lirio" (Honduras); "wild
guava" (British Honduras); "costarrica" (Tabasco).
Leaves of small seedling plants, often found abundantly in the
forest, are narrower than those of adult plants and usually
handsomely tinted with pink or purple. The fruit has been reported
as edible, but it cannot be particularly palatable. Boys sometimes
thrust a sharp stick through the fruits and use them as spinning
tops, hence the name "trompillo" given to the shrub in Costa Rica.
ALLENANTHUS Standley
Small trees with terete branches, the internodes elongated; stipules persistent or
deciduous, ovate, cuspidate. Leaves opposite, short petiolate, membranaceous;
inflorescence a compound cymose-corymbose terminal panicle, or in axils of upper
leaves; flowers small, pedicellate; hypanthium truncate, obovoid, laterally
compressed and sometimes very narrowly winged; calyx 4-lobate, small, erect,
persistent; corolla small, tubular-campanulate, 4-lobate; stamens 4, inserted in the
throat of corolla, anthers ovate-oblong, nearly sessile; style bifid, as long as the
corolla; ovary bilocular, each cell uniovulate; fruit dry, the locules parallel, central,
surrounded by a spongy wing; seeds pendulous, compressed.
A small genus, of two known species, first discovered in 1939 in
Panama. Since one species is known from Honduras and a variety of
it from Chiapas, the genus is included from Guatemala where it
may be expected.
Allenanthus hondurensis Standl. Ceiba 1: 45. 1950.
Moist valleys and hillsides in deciduous forests, about 800 m.,
Honduras.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 9
Leaves ovate-oblong to lanceolate, long attenuate-acuminate, 11-14 cm. long and
5-6 cm. broad.
Allenanthus hondurensis var. parvifolia L. Wms.,
Phytologia 25: 461. 1973.
Moist hillsides, alt. 800 m., Mexico (Chiapas).
Leaves lanceolate or ovate-lanceolate, acuminate, 3.5-5.5 cm. long and 1.5-2.5 cm.
broad.
This Mexican variety has smaller leaves, and is smaller in most
parts, except fruits.
ALSEIS Schott
Trees or large shrubs, more or less pubescent, with terete branches; leaves
opposite, petiolate, membranaceous; stipules interpetiolar, short or elongate;
inflorescence spicate or paniculate, the spikes simple or branched, axillary and
terminal; flowers small, white or yellowish; hypanthium obconic, the calyx 5-lobate,
the lobes broad or narrow, deciduous; corolla cylindric, short, pilose within, the limb
5-lobate, the lobes valvate; stamens 5, inserted at the base of the corolla tube; the
filaments elongate, villous, the anthers versatile, oblong, exserted; ovary 2-celled, the
style elongate, exserted, bipartite, the obtuse branches recurved; ovules numerous,
imbricate, the placentae pendulous from the apex of the cell; capsule oblong-
turbinate, 2-celled, septicidally bivalvate from the apex; seeds linear- fusiform, the
testa produced at each end.
The genus is a small one, most of the species being South
American. A. blackiana Hemsl. occurs in Panama.
Rachis of the inflorescence densely hirtellous; leaves hirtellous beneath, at least
along the costa; capsules mostly more or less recurved at maturity.
A. yucatanensis.
Rachis of the inflorescence puberulent; leaves almost glabrous, puberulent beneath
on the costa; capsules apparently erect or ascending at maturity ..A. hondurensis.
Alseis hondurensis Standl. Trop. Woods 16: 48. 1928. A
schippii Lundell, Lloydia 4: 56. 1941 (type from Machaca, British
Honduras, Schipp 1230). Sapote de montaha.
Moist or wet, mixed forest, at or little above sea level; Peten;
Izabal; Retalhuleu (?). British Honduras.
A tree about 10 m. high, the trunk 13 cm. in diameter, the stout branchlets
subterete, minutely puberulent when young; stipules narrowly lanceolate, 1 cm. long;
petioles 1.5-2 cm. long, the blades oblong-oblanceolate or narrowly oblong-obovate,
10-20 cm. long, 3-7.5 cm. broad, abruptly acute or short-acuminate, gradually long-
attenuate to the acute or acuminate base, glabrous above, minutely puberulent
beneath on the nerves; flower spikes 11-20 cm. long, many-flowered, forming a
terminal panicle, the flowers subsessile; hypanthium minutely puberulent, 2-2.5 mm.
10 FIELDIANA: BOTANY, VOLUME 24
long; calyx lobes ovate, 1 mm. long, obtuse, unequal, glabrous; capsules clavate, 1.5
cm. long or less.
The plant of Retalhuleu is represented only by sterile material
and may belong to a distinct species.
Alseis yucatanensis Standl. Field Mus. Bot. 8: 50. 1930.
Dzon; son; palo son. Figure 13.
Chiefly in secondary forest, Peten (Chimah; Monte Polol).
British Honduras; Mexico (Yucatan; Tabasco).
A tree 20-30 m. tall, the trunk 40 cm. or more in diameter, the young branchlets
hirtellous; stipules triangular, 5-9 mm. long, glabrous outside or nearly so, caducous;
petioles 1-2.5 cm. long, the blades obovate or oblong-obovate, 10-30 cm. long, 4-12.5
cm. broad, acuminate or rounded and abruptly short-acuminate, cuneate-attenuate
to the base, almost glabrous above, hirtellous beneath, especially on the costa, or
finally glabrate; inflorescence spicate or paniculate, 7-40 cm. long; flowers sessile or
nearly so; hypanthium hirtellous, 3 mm. long, the calyx lobes ovate, obtuse, almost 2
mm. long, puberulent; corolla broadly campanulate, 2.5 mm. long, puberulent
outside; filaments long-exserted, villous below; capsules clavate, about 1.5 cm. long
and 3 mm. thick, glabrate, brownish.
In British Honduras the tree is called "wild mamee"; from
Yucatan the name "cacao-che" is reported. In Tabasco it is said to
be called "papelillo."
AMAIOUA Aublet
Unarmed trees or shrubs, the branchlets usually sericeous; leaves opposite or
ternate, short-petiolate; stipules interpetiolar, triangular or ovate, united at first into
a conic cap, deciduous; inflorescence terminal cymes or fascicles; flowers white,
unisexual; hypanthium oblong to hemispheric; calyx cupular or short-tubular,
deciduous, truncate or 6-dentate, the teeth short, subulate; corolla salverform,
sericeous outside, the tube terete or ventricose, the throat tomentose, the limb 6-
lobate, the lobes contorted, oblong, spreading; stamens 6, inserted in the corolla tube,
the filaments very short; anthers dorsifixed, linear, included; ovary 2-celled, the style
short, the 2 branches coherent; ovules numerous, biseriate, the placenta affixed to the
septum; fruit baccate, small, oblong, areolate at the apex, 1-celled, the cortex thin-
coriaceous; seeds numerous, embedded in pulp, horizontal, compressed, suborbicular,
the testa fibrous.
A small genus, chiefly South American; only one of the species
extending into North America.
Amaioua corymbosa HBK. Nov. Gen. & Sp. PL 3: 419. 1820.
Figure 31.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 11
Wet forest, Izabal, at or near sea level. Tabasco and British
Honduras to Panama (Atlantic slope); Cuba; the Guianas.
A shrub or small tree 2-6 m. high, the trunk sometimes 15 cm. in diameter;
stipules lance-oblong, 1.5-2.5 cm. long, ferruginous-sericeous outside; petioles 3-15
mm. long, the blades oval or oval-elliptic, 5-20 cm. long, 3-10 cm. broad, short-
acuminate, at the base broadly rounded to short-acuminate, somewhat coriaceous,
glabrous above, more or less sericeous beneath on the nerves and veins; staminate
inflorescence corymbose, lax, often long-pedunculate, the calyx cupular, 3-4 mm.
long, sericeous, denticulate, the corolla about 1.8 cm. long, the tube retrorse-sericeous,
the lobes lanceolate, about equaling the tube; pistillate inflorescence capitate or
corymbose, few-flowered, on a long or short peduncle, the corolla about 1 cm. long;
fruit oval or obovoid, 12-15 mm. long, lustrous, brown-purple with pale yellow-green
at the apex; seeds reddish brown, lustrous, striate, 3 mm. long.
Called "wild coffee" and "bastard coffee" in British Honduras.
The bark is said to have the odor of pea pods. The name "tarro de
venado" is reported from Tabasco.
ANISOMERIS Presl
Shrubs or small trees, rarely scandent, often with spinose branchlets; leaves
sessile or petiolate, membranaceous or coriaceous; stipules interpetiolar, deciduous,
acute or acuminate; flowers small, white or yellowish, usually in axillary pedunculate
cymes, rarely solitary; hypanthium oblong or turbinate; calyx 4-lobate, the lobes
commonly elongate, equal or unequal, persistent; corolla salverform or funnelform,
with a slender, usually elongate tube, the throat glabrous or pilose, the limb 4-lobate,
the lobes usually short, not appendaged, valvate or with slightly imbricate margins;
stamens 4, inserted in the corolla throat; anthers linear-oblong or sagittate,
dorsifixed, sessile, included or subexserted; ovary 2-celled, the style filiform, with 2
short branches; ovules solitary, pendulous from the apex of the cell; fruit drupaceous,
small, oblong or ovoid, osseous, 2-celled; seeds pedulous, cylindric.
The majority of the species are South American. About five are
known from Central America, and two more from Mexico.
Corolla tube glabrous outside; calyx lobes minute, as broad as long A. brachypoda.
Corolla tube pilose outside.
Flowers in pedunculate cymes; calyx lobes minute, as broad as long A. protracta.
Flowers sessile at or near the ends of the branches; calyx lobes oblong-linear.
A. recordii.
Anisomeris brachypoda (Donn.-Sm.) Standl. Field Mus. Bot.
4: 293. 1929. Chomelia brachypoda Donn.-Sm. Bot. Gaz. 47: 255.
1909. Figure 40.
Wet forest along streams or in quebradas, 900-1,600 m.; Alta
Verapaz (type from banks of Rio Ogewaj near Sasis, Tuerckheim II.
2253); Chimaltenango; Huehuetenango; sterile specimens from
Quezaltenango and San Marcos may represent the same species.
12 FIELDIANA: BOTANY, VOLUME 24
An unarmed shrub or small tree, sometimes 12 m. high, with slender branches;
stipules narrowly triangular, 6-8 mm. long, subulate-attenuate, sericeous outside;
petioles 3-6 mm. long; leaf blades lance-oblong or elliptic-lanceolate, 6-10 cm. long,
1.5-3.5 cm. broad, long-acuminate, acute or short-acuminate at the base, glabrous
above or nearly so, appressed-pilose beneath along the nerves and barbate in the
nerve axils; cymes axillary, 4-8-flowered, on peduncles 5-8 mm. long, the flowers
sessile; calyx and hypanthium 2.5-3 mm. long, minutely strigillose, the lobes minute,
triangular; corolla dull dark red, the tube 11-12 mm. long, glabrous outside, the oval
lobes 2 mm. long, sparsely strigillose outside.
An inconspicuous shrub, in appearance suggestive of some
species of Rondeletia.
Anisomeris protracta (Bartl.) Standl. Field Mus. Bot. 4: 293.
1929. Guettarda protracta Bartl. ex DC. Prodr. 4: 457. 1830.
Chomelia protracta Standl. Contr. U. S. Nat. Herb. 23: 1384. 1926.
Chechem (Coban).
Common in wet, pine or mixed, limestone forest, or on open
limestone slopes, 1,500 m. or lower; Alta Verapaz; Izabal;
Chiquimula; Huehuetenango. Southern and western Mexico;
British Honduras; Honduras.
A shrub or small tree, commonly 3-6 m. high but said to attain sometimes a
height of 10 m.; branches densely sericeous; stipules 5-9 mm. long, triangular-
lanceolate, long-attenuate, sericeous outside, brown; leaves short-petiolate, narrowly
lanceolate to oblong-elliptic, 7-15 cm. long, 2.5-5.5 cm. broad, long-acuminate, with a
narrow, often falcate tip, attenuate to subobtuse at the base, green above and
appressed-pilose or glabrate, paler beneath and sericeous, especially along the nerves,
barbate in the nerve axils; cymes pedunculate, 1-2.5 cm. long, sericeous, the flowers
sessile, secund; calyx and hypanthium 1.5-2.2 mm. long, the hypanthium densely
short-pilose the calyx lobes minute, rounded; corolla white, densely sericeous outside,
the tube 7-10 mm. long, the lobes rounded, about 1.5 mm. long; fruit dark blue at
maturity, oval, 4-5 mm. long, thinly sericeous.
In Tabasco and British Honduras the species is reported as
growing in savanna forest and doubtless it occurs in similar
locations in Peten.
Anisomeris recordii Standl. N. Am. Fl. 32: 227. 1934.
Chomelia recordii Standl. Trop. Woods 7: 9. 1926 (high forest
between Los Andes and Entre Rios, Izabal, Record G31). Clavo.
Collected also on southern slopes of Cerro San Gil, Izabal, at 30-500
m. Nicaragua and Panama.
A shrub or small tree, about 5 m. high, the trunk somewhat fluted, the
branchlets hirtellous or glabrate; stipules 2.5-5 mm. long, triangular or ovate,
acuminate or cuspidate, appressed-pilose; petioles 2-3 mm. long; leaf blades broadly
ovate or ovate-elliptic, 4-8.5 cm. long, 2-5 cm. broad, obtuse to abruptly acute, with
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 13
obtuse tip, rounded or obtuse at the base, green above and almost glabrous, copiously
short-pilose beneath with spreading or subappressed hairs; flowers clustered and
sessile at the ends of the branches; calyx and hypanthium 5-6 mm. long, densely
pilose with long appressed white hairs; calyx lobes oblong-linear, equaling the
hypanthium, corolla white, densely white-sericeous, the very slender tube 2 cm. long,
the lobes oblong, obtuse, 4 mm. long, glabrous within; fruit green, turning dull
reddish, oval, 7-8 mm. long.
ANTIRHEA Commerson
Trees or shrubs, usually glabrous and resinous- viscid; leaves opposite, mostly
coriaceous and lustrous; stipules deciduous or persistent; flowers small, perfect or
polygamous, sessile or short-pedicellate, secund along the branches of a bifid,
scorpioid, axillary, pedunculate or sessile cyme; hypanthium ovoid or obovoid; calyx
lobes persistent, often unequal; corolla funnelform, glabrous or sericeous, the tube
elongate, cylindric, the throat naked or pilose, the limb 4-5-lobate, the lobes obtuse,
imbricate, 2 of them exterior; stamens 4 or 5, inserted in the corolla throat, included
or subexserted, the filaments short; anthers oblong, dorsifixed; ovary with 2-10 or
sometimes more numerous cells, the style filiform, the stigma capitate or 2-3-fid;
ovules solitary, pendulous; fruit drupaceous, small, oblong, with thin flesh, the stone
ligneous or osseous; seeds cylindric.
About 40 species, chiefly West Indian, but occurring also in
Asia and the islands of the Indian Ocean. One other species is
known from Central America, A. panamensis Standl., endemic in
Panama.
Antirhea lucida (Swartz) Benth. & Hook. Gen. PL 2: 100.
1873. Laugeria lucida Swartz, Prodr. 48. 1788. Figure 38.
Known in this region only from the Jacinto Hills of British
Honduras, at 60 m. (Schipp S616); and at Tikal, Pet£n (Tun 1000).
Bahamas, through the Greater Antilles to Trinidad.
A shrub or tree, reported to attain in British Honduras a height of 18 m. with
trunk diameter of 60 cm., the bark smooth; stipules ovate-deltoid, 5-8 mm. long,
acuminate, minutely sericeous outside, caducous; petioles 3-8 mm. long, the blades
elliptic to elliptic-oblong, obtuse or acutish, acute to rounded and short-decurrent at
the base; cymes usually once bifid, the slender branches 3-8 cm. long, the peduncle 2-
3 cm. long, the numerous flowers distant, sessile or nearly so; calyx and hypanthium
2-3 mm. long, glabrous or minutely puberulent; calyx 5-lobate, the lobes
semiorbicular, ciliolate; corolla white, 5-7 mm. long, the obtuse lobes half as long as
the tube; fruit oval or oblong, red or black, 5-7 mm. long.
The single British Honduras specimen is in fruit, and flowering
material may show that it represents a distinct species, although
this is rather improbable.
14 FIELDIANA: BOTANY, VOLUME 24
APPUNIA Hooker f.
Shrubs, glabrous or puberulent, with terete branches; leaves opposite, short-
petiolate, membranaceous, acuminate; stipules interpetiolar, broadly triangular or
annular, connate with the petiole, persistent; flowers small, white, capitate, the heads
long-pedunculate, few-flowered, axillary, bracteate at the base; hypanthium
turbinate or hemispheric, the calyx cupular, truncate, obscurely 5-dentate; corolla
tube cylindric, puberulent or glabrous in the throat, the limb 5-lobate, the lobes
valvate, linear-oblong, about equaling the tube; stamens 5, inserted in the corolla
tube, the filaments very short; anthers dorsifixed, included; ovary perfectly or
imperfectly 4-celled, the style filiform, the stigma capitate; ovules solitary, peltately
attached to the middle of the septum; fruit baccate, subglobose, containing four 2-
celled nutlets, one of the cells empty, the other 1 -seeded.
The genus is a small one, one species being found in Panama,
the others distributed in the Guianas.
Appunia guatemalensis Donn.-Sm. Bot. Gaz. 48: 294. 1909.
Morinda mesochora Standl. Field Mus. Bot. 4: 345. 1929 (type from
Honey Camp, British Honduras, Lundell 19). Figure 57.
Wet mixed forest, sometimes in thickets or hillside pine forest,
at or little above sea level; Izabal (type from Livingston,
Tuerckheim 11.1230). British Honduras; Atlantic coast of Honduras
and Nicaragua.
A glabrous shrub 1-2.5 m. tall; stipules apiculate or forming a short bicuspidate
sheath; leaves short-petiolate or almost sessile, elliptic-oblong or obovate-oblong, 10-
16 cm. long, 5-7 cm. broad, acuminate or abruptly acuminate, acute or acuminate at
the base, with 5-6 pairs of lateral nerves; peduncles axillary, as much as 4 cm. long
but usually shorter, the heads about 7-flowered, without the corollas 5-6 mm. in
diameter; corolla white, often tinged with purplish outside, 1.5 cm. long; fruit globose,
about 6 mm. in diameter, becoming dark purple.
ASEMNANTHA Hooker f.
Slender pubescent shrubs with terete branches; leaves opposite, short-petiolate,
subcoriaceous; stipules interpetiolar, small, persistent; flowers small, in few-flowered
axillary fascicles, yellow, short-pedicellate; hypanthium ovoid; calyx 4-lobate, the
lobes elongate, persistent; corolla urceolate-tubular, puberulent outside, the throat
slightly contracted, the limb 4-lobate, the lobes small, erect-patent, obtuse, valvate in
the bud; stamens 4, the filaments joined at the base and attached to the disk, not to
the corolla, free above, pilose; anthers linear-oblong, basifixed, included; ovary 2-
celled, the style filiform, the stigma obtuse, exserted; ovules solitary, pendulous from
the apex of the cell; fruit drupaceous, suborbicular, laterally compressed, chartaceous
when dry; seeds laterally compressed.
The genus consists of a single species.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 15
Asemnantha pubescens Hook. f. in Bentham & Hooker, Gen.
PL 2: 107. 1873; in Hook. Icon. 12: 40, t. 1145. 1873. Figure 43.
Collected at Honey Camp, British Honduras. Peten. Mexico
(Yucatan), whence originally described.
A slender shrub, commonly about 1 m. high, the branchlets hirtellous; stipules 3-
5 mm. long, subulate-acuminate from a broad base; petioles 2-4 mm. long, the blades
lanceolate to broadly ovate or elliptic, 3-7 cm. long, 1-2.5 cm. broad, gradually
narrowed to the acute or attenuate apex, rounded to acute at the base, short-pilose
on both surfaces, at least when young; flowers few in each fascicle, the pedicels very
short; calyx and hypanthium short-pilose, the calyx lobes lance-linear, attenuate, 4
mm. long, spreading in fruit; corolla half longer than the calyx lobes, short-pilose, the
lobes rounded-deltoid, acutish, about one-sixth as long as the tube; fruit short-pilose,
4-4.5 mm. long.
BALMEA Martinez
Shrubs, terrestrial or sometimes epiphytic, with thick branches; stipules
interpetiolar, ovate-acuminate, caducous; leaves opposite, large and broad, petiolate,
membranaceous, deciduous, mostly shallowly cordate at the base; inflorescence
terminal, cymose-corymbose, pedunculate, the peduncle recurved and the flowers
thus pendulous, the flowers few or rather numerous, mostly ternate, slender-
pedicellate; hypanthium narrowly turbinate; calyx 5-parted, the segments linear,
more or less persistent; corolla tube cylindric or slightly dilated upward, the lobes
broad, contorted, reflexed in anthesis; stamens inserted on the upper part of the
corolla tube, the filaments complanate, the anthers dorsifixed; fruit capsular, erect,
oval-oblong, 2-celled, septicidally bivalvate at the apex; seeds numerous, imbricate,
broadly winged.
The genus consists of a single species. It was named for
Professor Juan Balme of Mexico.
Balmea stormae Martinez, Bull. Torr. Bot. Club 69: 438, /. 1-
1L 1942; Anal. Inst. Biol. Mex. 13: 37, /. 1-4. 1942.
Moist or wet, mixed or oak mountain forest, 1,400-2,300 m.;
Zacapa (Sierra de las Minas); Jalapa (Potrero Carrillo);
Huehuetenango (northwest of Cuilco). Mexico (Michoacan).
A glabrous shrub or tree, sometimes 7 m. high, terrestrial or sometimes epiphytic
(on Quercus), rather sparsely branched, sometimes from the base, the trunk as much
as 20 cm. in diameter; bark smooth, greenish purple, peeling off in thin shreds; wood
hard, whitish; internodes very short, the leaf scars conspicuous; leaves often clustered
at the ends of the branches, on long slender petioles, broadly ovate or rounded-ovate,
mostly 9-13 cm. long and 6-11 cm. broad, acute or abruptly short-acuminate, usually
cordate or subcordate at the base; stipules about 1 cm. long; corymbs generally 9-13-
flowered, borne on peduncles 5 cm. long or shorter, the pedicels 2 cm. long or less;
flowers sweet-scented, bright red tinged with purple, or when fully open dark purple;
calyx lobes 8-10 mm. long; corolla tube 22-28 mm. long, the 5 lobes subovate, short;
16 FIELDIANA: BOTANY, VOLUME 24
anthers fusiform, 9-10 mm. long, not exserted, the filaments only 4 mm. long; style
20-23 mm. long, the stigma bilamellate; capsule 23-28 mm. long; seeds, including the
wing, 3.5-4 mm. long.
In Michoac£n the tree is known by the vernacular name
"ayuque." The senior author first became acquainted with this tree
in 1941, when a collection of it was made near Uruapam, Michoacan
by William C. Leavenworth and H. Hoogstraal. It was recognized
that it was something new to the Mexican flora, but study and
description of the plant was delayed because the material was
incomplete. Not long afterward complete material from the same
state was forwarded by Professor Martinez, who had made a
detailed study of it, prepared a full description, and had made
several handsome drawings showing all the details. He believed the
plant represented a new genus, a belief that was fully justified. In
December, 1939, Dr. Steyermark obtained in the Department of
Jalapa sterile material of a Rubiaceous plant that could not be
determined, until finally it was associated with the Mexican
Balmea. Two other sterile collections were made by him in Zacapa
and Huehuetenango in 1942. It thus happens that a tree growing at
or near a locality which various botanists have worked (Uruapam)
was overlooked until quite recently, when it has been found in no
less than five widely separated localities. Since the Guatemalan
collections are sterile, there is a possibility that they represent a
distinct species, but the foliage, which is rather distinctive, is
exactly like that of the Mexican plant.
BERTIERA Aublet
Shrubs or small trees, glabrous or pubescent, with terete branchlets; leaves
opposite, petiolate or sessile; stipules intrapetiolar, connate into a bifid sheath;
inflorescence terminal, cymose, the cymes arranged in terminal pedunculate panicles;
flowers small, white or greenish; hypanthium globose or turbinate, the calyx 5-
dentate or truncate, persistent; corolla funnelform, the tube terete, usually sericeous
outside, the throat glabrous or villous, the limb 5-lobate, the lobes short, ovate, acute,
contorted in bud; stamens 5, inserted in the corolla throat; filaments very short, the
anthers dorsifixed, included; ovary 2-celled, the style filiform, glabrous, the stigma
fusiform, entire or bilobate; ovules numerous, the placentae affixed to the septum;
fruit baccate, globose, fleshy, 2-celled; seeds numerous, very small, angulate,
foveolate or granulate.
A small genus of tropical North and South America. Two other
North American species are known, in Costa Rica and Panama.
Bertiera guianensis Aubl. PI. Guin. 180, t. 69. 1775. B. tenuis
Lundell, Wrightia 4: 49. 1968 (type, from Alta Verapaz, Contreras
4318). Figure 33.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 17
Wet forests and thickets, near or little above sea level; Alta
Verapaz; Izabal. Southern Mexico; British Honduras; Costa Rica
and Panama south to Bolivia and Brazil; West Indies.
A slender shrub or small tree 2-4 m. high, the trunk sometimes 8 cm. in diameter,
the branchlets appressed-pilose or glabrate; stipules 5-10 mm. long, the lobes
subulate-acuminate, brown, sericeous; petioles 3-7 mm. long, the blades lance-oblong
or ovate-oblong, membranaceous, 6-17 cm. long, 2-7 cm. broad, short-acuminate or
attenuate, acute or acutish at the base, glabrous above or nearly so, appressed-pilose
beneath, especially along the veins; panicle narrow and elongate, the flowers sessile,
secund, the bracts linear; calyx and hypanthium 1.5-2.5 mm. long, short-pilose, the
calyx lobes triangular; corolla greenish white, 4.5-7 mm. long, appressed-pilose, the
lobes deltoid-ovate, acuminate, half as long as the tube; fruit globose, 3.5-4 mm. in
diameter, blue, 10-costate; seeds 1-2 mm. long, coarsely granulate, orange-brown.
We assume that this lowland species may be found on the
Atlantic side of Honduras and Nicaragua, but we have seen no
specimens.
BLEPHARIDIUM Standley
Shrubs or small trees, pubescent or almost glabrous, with subterete branchlets;
leaves opposite, petiolate, large; stipules intrapetiolar, large, thin, acuminate,
caducous; flowers large, pedicellate, bibracteolate, in 3-flowered axillary long-
pedunculate cymes; hypanthium obovoid, the large calyx 4-lobate, the very broad
lobes imbricate, ciliolate; corolla salverform, coriaceous, the slender tube elongate,
densely villous within except near the base, the 4 lobes imbricate in bud, broad,
spreading, one of them exterior; stamens 4, inserted in the corolla throat, the anthers
sessile, linear, dorsifixed, included; ovary 2-celled, the style filiform, the stigma
bilobate, with linear elongate lobes; ovules numerous, imbricate, winged, the
placentae attached to the septum; capsule large, somewhat ligneous, 2-celled,
loculicidally bivalvate; seeds broadly winged.
One other species is known, B. mexicanum Standl., described
from Palenque, Chiapas (Mexico).
Blepharidium guatemalense Standl. Journ. Wash. Acad. Sci.
8: 59. 1918. Popiste; polo de estricnina; irayol bianco; irayol.
Usually in hilly pine forest, sometimes in savannas, 300 m. or
lower; endemic; Pet£n; Alta Verapaz (type from forest along Rio
Saclac below Secanqulm, Pittier 266); Izabal (Quiriqua; Cristina;
Santa Cruz); Huehuetenango (Ixcan).
A shrub or small tree, usually 1.5-5.5 m. tall, sometimes a tree of 6-18 m., the
branchlets glabrous; stipules ovate-triangular, 2-2.5 cm. long, acuminate, brown,
glabrous outside, sericeous within at the base; petioles stout, 2.5-5 cm. long, glabrous;
leaf blades oval-oblong to oblanceolate or oblong-obovate, 15-35 cm. long, 4-21 cm.
broad, obtuse to acute or abruptly short-acuminate, obtuse to attenuate at the base,
glabrous above, sparsely short-pilose beneath along the costa, short-barbate in the
18 FIELDIANA: BOTANY, VOLUME 24
nerve axils; peduncles 6-20 cm. long, glabrous, the stout pedicels 2 cm. long or less;
bractlets broadly ovate, 3-4 mm. long, deciduous; calyx glabrous, 4-5 mm. long, 7-8
mm. broad, half as long as the hypanthium, the broad lobes rounded or truncate;
corolla tube about 6 cm. long, 4-5 mm. thick, glabrous outside, the oval lobes 1 cm.
long; capsule oblong-oval, 1.5-3 cm. long, obtuse at the base and apex.
The shrub is plentiful on the brushy pine hillsides above
Quiriagua Hospital. It is probable that B. mexicanum also is to be
found in northern Peten. It differs in having smaller flowers, the
corolla tube only 4 cm. long, and sparse pubescence over the lower
leaf surface. The wood is used for rafters, beams, and supports of
lowland houses. In Huehuetenango the leaves and bark are reputed
poisonous, hence the name "estricnina" given the tree, but it is
improbable that poisonous properties are found in this genus.
BORRERIA G. F. W. Meyer
Annual or perennial herbs, sometimes woody at the base or almost throughout,
the stems more or less tetragonous; stipules united with the petioles to form a sheath,
this bearing few or numerous setae; leaves small and usually narrow, opposite or
appearing whorled, often with fascicles of smaller leaves in their axils; flowers small,
in sessile or terminal heads, the terminal heads subtended by leaflike bracts;
hypanthium turbinate or cylindric, the calyx 2-or 4-dentate, the lobes usually narrow,
sometimes with small interposed teeth; corolla funnelform, the 4 lobes valvate in
bud; stamens 4, inserted in or below the mouth of the corolla tube, the anthers often
exserted; ovary 2-celled, the style simple, the stigma shallowly bilobate; ovules
solitary in the cell, affixed to the septum; capsule coriaceous or membranaceous,
crowned by the persistent calyx, septicidally dehiscent or rarely both septicidal and
loculicidal, the valves coherent at the base; seeds minute, oblong, convex dorsally,
sulcate ventrally.
Perhaps as many as 100 species, pantropical in distribution but
chiefly in South America. A few others are found in Central
America. The species found in Guatemala are inclined to be weedy
and their altitudinal range is often greater than might be expected.
It is sometimes difficult to distinguish Borrerias from other genera
of the tribe Spermacoceae, unless fruits are present.
Seeds transversely striate; plants annual; calyx 4-lobate.
Flower heads 2-3 cm. broad; sepals oval, rounded at the apex B. vegeta.
Flower heads less than 1.5 cm. broad; sepals triangular-subulate B. laevis.
Seeds not transversely striate, either smooth or reticulate; calyx 2- or 4-lobate.
Leaves linear or nearly so, usually verticillate; plants perennial.
Sepals 4; plant usually pubescent B. suaveolens.
Sepals 2; plant usually glabrous B. verticillata.
Leaves mostly elliptic or broader; plants annual.
Sepals 4, fruits pubescent.
Plants usually prostrate, yellowish when dry; flower heads axillary.
B. latifolia.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 19
Plants usually erect, not yellowish when dry; flowers heads terminal and
axillary B. ocymoides.
Sepals 2; fruits white-villous above the middle B. densiflora.
Borreria densiflora DC. Prodr. 4: 542. 1830. Spermacoce
spinosa Swartz, Obs. Bot. 45. 1791, not Jacq., 1760, nor L., 1762. B.
spinosa Cham. & Schlecht. Linnaea 3: 340. 1828, excluding Linnean
synonym. B. radicosa Brandegee, Univ. Calif. Publ. Bot. 10: 416.
1924 (type from Jalisco, Mexico).
A weedy plant, in thickets, on brushy hillsides, or on sandbars
along streams, chiefly in the east at 500 m. or less, but in
Huehuetenango ascending to 1,700 m.; Zacapa; Santa Rosa;
Escuintla; Huehuetenango. Southern Mexico; El Salvador;
Honduras; Costa Rica and Panama; southward to Argentina; West
Indies.
Plants annual, usually erect or nearly so and stout, often simple at the base,
almost glabrous but the stems usually sparsely scabrous on the angles; leaves linear
or linear-lanceolate, as much as 8 cm. long and 12 mm. broad but usually smaller,
grayish green when dried, acuminate, narrowed to the base, the lateral nerves usually
obvious; stipule sheath with numerous long setae; flower heads subtended by 4-8
large leaflike bracts, 1.5-2 cm. broad; the flowers very numerous and densely
congested; hypanthium densely white-pilose above; calyx lobes 2 and 1.5-2 mm. long,
linear; corolla white, slightly exceeding the calyx, glabrous; stamens equaling the
corolla lobes; capsule oblong, white-villous above the middle, 3-4 mm. long; seeds
brown-red, finely pitted.
Inclined to be weedy as are others of the genus.
Borreria laevis (Lam.) Griseb. Fl. Brit. W. Ind. 349. 1861.
Spermacoce laevis Lam. Tabl. Encycl. 1: 273. 1791. S. echioides
HBK. Nov. Gen. & Sp. 3: 269. 1820. Golondrina (Guatemala);
sanalotodo; hierba de pdjaro; ritpur (Alta Verapaz, fide
Dieseldorff); palis (Peten, fide Lundell); zic-chichibe (Peten, Maya,
fide Lundell). Figure 64.
A common weed throughout the greater part of Guatemala,
except at high elevations, usually in thickets, on gravel bars, or in
waste or cultivated ground, commonest at low elevations but
ascending to 2,000 m. or probably even higher; Peten; Izabal; Alta
Verapaz; El Progreso; Zacapa; Chiquimula; Jalapa; Santa Rosa;
Escuintla; Guatemala; Sacatepequez; Suchitepequez; Retalhuleu;
Quezaltenango; San Marcos; Huehuetenango; Quiche. Mexico to
Panama; southward through the greater part of South America;
West Indies.
20 FIELDIANA: BOTANY, VOLUME 24
Plants annual, erect or more often decumbent or procumbent, the stems glabrous
or sparsely pubescent; leaves lanceolate to ovate-lanceolate, ovate, or oblong-
lanceolate, commonly 2-5 cm. long and 1-2.5 cm. broad, acuminate, cuneate and
usually short-petiolate at the base, glabrous or beneath usually pubescent; stipule
sheath bearing numerous setae of about the same length; flower heads terminal and
axillary, up to about 1.5 cm. broad, usually smaller; hypanthium obovoid, pubescent
above; calyx teeth 4-5, small, triangular-subulate; corolla white, the lobes about
equaling the tube, pubescent; capsule ellipsoid, 2-3 mm. long, somewhat flattened
laterally, pubescent above; seeds castaneous, transversely striate.
One of the commonest weeds of Central America, often abun-
dant in banana and coffee plantations, and grain fields. According
to Dieseldorff, in his "Las plantas medicinales del departamento de
Alta Verapaz," (1940), it is used in that department in native medi-
cine.
Borreria latifolia (Aubl.) Schum. in Mart. Fl. Bras. 6(6): 61, t.
80. 1888. Spermacoce latifolia Aubl. PL Guian. 55, t. 19, f. 1. 1775.
Damp thickets or fields, sometimes on grassy banks or even in
pine forest, at low elevations, ascending to about 1,000 m.; probably
in Peten and Izabal; Alta Verapaz; Baja Verapaz; Jutiapa;
Chiquimula; Santa Rosa; Escuintla; Suchitepequez; San Marcos;
Retalhuleu. Southern Mexico; British Honduras to Panama,
southward through the warmer parts of South America; West
Indies.
Plants annual, usually prostrate or procumbent and often with elongate stems,
sometimes erect or nearly so, the whole plant yellowish in the dry state; stems
acutely quandrangular or even narrowly winged, glabrous or villosulous; leaves
almost sessile, chiefly elliptic, sometimes slightly obovate, about 2-4 cm. long and 1-
1.8 cm. broad, acute, pilose or villosulous on both surfaces, the lateral nerves
conspicuous; stipule sheath about 1 mm. long, the setae 3.5 mm. long, pilose; flower
heads dense and rather few-flowered, mostly axillary and about 1 cm. in diameter;
sepals 4, narrowly triangular, 1.5 mm. long, acute, ciliate; corolla white or pale bluish,
3 mm. long, pilose at the apex; style bilobate, with recurved lobes; capsule 3.5 mm.
long, subglobose, densely pilose above; the seed light brown, 2.5-3 mm. long, only
obscurely reticulate.
Among Central American species, this may be recognized
readily by the evident yellowish coloring of the dried plant and by
the relatively large almost smooth seeds. This species is often weedy
and as more and more forest areas are put into agriculture it may
be expected as an invader. The resemblance to Mitracarpus hirtus
is rather great.
Borreria ocymoides (Burm.) DC. Prodr. 4: 544. 1830.
Spermacoce ocymoides Burm. f. Fl. Ind. 34, t. 13, f. 1. 1768. S.
parviflora G. F. W. Meyer, Prim. Fl. Esseq. 83, t. L 1818. B.
tampicana DC. Prodr. 4: 544. 1830, in part. S. pringlei Wats. Proc.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 21
Am. Acad. 25: 152. 1890. Golondrina bianco, (Chimaltenango);
palitaria (Pete"n); catalpim (Alta Verapaz, fide Dieseldorff).
Widely distributed as a weed in thickets, pine and oak forest,
pastures, cafetales, cultivated fields, arenales, and other habitats,
chiefly in the lowlands but ascending in the interior to about 2,000
m.; Pete"n; Alta Verapaz; Izabal; Zacapa; Chiquimula; Jalapa;
Guatemala; Sacatepequez; Chimaltenango; Suchitepequez;
Retalhuleu; San Marcos; Huehuetenango. Mexico; British
Honduras to Panama, and through the greater part of South
America; West Indies; naturalized in Asia and Malaysia.
Plants annual, usually erect, sometimes spreading, 50 cm. high or less, the stems
acutely quadrangular, glabrous or sometimes scabrous or hispidulous on the angles;
stipule sheath 2.5 mm. long, the setae as long or slightly longer; leaves small,
subsessile, elliptic to ovate or lance-elliptic, mostly 1.5-2 cm. long, sometimes longer,
5-10 mm. broad, scaberulous above, especially near the margins, scaberulous beneath
on the nerves, these usually 4 on each side and rather conspicuous; flower heads
terminal and axillary, small, the crowded flowers short-pedicellate; hypanthium
turbinate, glabrous or sometimes puberulent above; calyx lobes usually 4, divided 3
and 1, linear, acute, 1 mm. long or less; corolla white, hardly 1 mm. long, minutely
puberulent outside, villous within, the lobes slightly longer than the tube; anthers
sessile, not exserted; capsule 1 mm. long, crowned by the slightly enlarged sepals;
seeds dark brown, reticulate, about 1-1.5 mm. long.
In the Coban region this species often is found in marshes.
According to Dieseldorff, in his "Las plantas medicinales del depar-
tamento de Alta Verapaz," (1940), it is used in that department in
native medicine.
The fruit is nearly or completely glabrous, when it dehisces
usually three calyx lobes are on one part and only one on the other.
Borreria suaveolens G. F. W. Meyer, Prim. Fl. Esseq. 81, t. 1.
1818. Spermacoce tenella HBK. Nov. Gen. & Sp. 3: 345. 1820. B.
tenella Cham. & Schlecht. Linnaea 3: 317. 1828. B. haenkeana DC.
Prodr. 4: 547. 1830. Hierba de toro; lengua de pa/aro.
Pine forest or savannas, sometimes in mountain meadows or
thickets, except in Peten usually growing at 900-2,000 m.; Peten;
Zacapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe'quez;
Chimaltenango; Quiche; Huehuetenango. Mexico to Panama, and
widely distributed through South America.
Usually a stiff erect perennial, often much branched from the base, 50 cm. high
or less, sometimes suffrutescent, very variable in foliage, at least in South America,
less variable in Central America, the stems pubescent or glabrous; stipule sheath
short, with few or numerous long setae; leaves mostly linear or lance-linear and 2-5
cm. long, 1 -nerved, rather stiff, often revolute, scabrous on both surfaces or glabrous;
22 FIELDIANA: BOTANY, VOLUME 24
flower heads very dense and many-flowered, 1 cm. broad or slightly larger, often all
terminal but heads frequently present in some of the upper leaf axils, the terminal
heads subtended by large leaflike bracts; hypanthium villosulous; sepals longer than
the hypanthium, linear, ciliate; corolla white, 2.5-3 mm. long; stamens exserted;
capsule oblong, pubescent at least at the apex, 2 mm. long; seeds brown, almost
smooth.
South American material usually referred to this species is
exceedingly variable, the extremes appearing to represent quite
distinct species. Attempts to subdivide the species have not been
successful heretofore, but it is quite possible that at some time in
the future other workers may be more successful. The Central
American collections exhibit relatively little variation except in
quantity of pubescence.
Borreria vegeta Standl. & Steyerm. Field Mus. Bot. 23: 21.
1943.
Weedy fields, damp thickets, or open slopes, 150-900 m.;
Jutiapa and Santa Rosa; type collected between Jutiapa and La
Burrera, northeast of Jutiapa, Standley 76007, El Salvador;
Honduras.
A stout erect annual, simple or with a few branches, the stems subterete below,
tetragonous above, laxly whitish-pilosulous; stipule sheath 5 mm. long, bearing
several setae of equal length; leaves large, sometimes contracted into a broad petiole
as much as 1 cm. long, lance-oblong to ovate or oblong-elliptic, 3.5-7.5 cm. long, 1.2-
2.8 cm. broad, attenuate-acuminate, cuneately contracted at the base, green above
and sparsely scabrous or glabrate, rough to the touch, paler beneath, scabrous or
hispidulous on the nerves and costa, the lateral nerves 6-7 pairs; heads large, chiefly
terminal but often also in the uppermost leaf axils, 2-2.5 cm. broad, the terminal
rounded-obovate, about 1.5 mm. long, green, rounded at the apex, pectinate-ciliate;
corolla white, 2.5 mm. long, glabrous outside, the lobes ovate, subacute; anthers
exserted; capsule oval or broadly oblong, 3 mm. long, membranaceous, villosulous
above or glabrate; seeds terete, fuscous-ferruginous, 2 mm. long, transversely rugose.
Borreria verticillata (L.) G. F. W. Meyer, Prim. Fl. Esseq.
83. 1818. Spermacoce verticillata L. Sp. PI. 102. 1753. B.
podocephala DC. Prodr. 4: 541. 1840. Sanalotodo (Huehuetenango);
hierba de pajaro (fide Aguilar).
A widely distributed and in many localities a common weedy
plant, usually at low elevations but ascending to about 2,000 m., in
thickets or savannas, meadows, or in waste or cultivated ground;
Peten; Alta Verapaz; Izabal; Jalapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Chimaltenango; Quiche; Suchitepequez;
Retalhuleu; Huehuetenango. Mexico; British Honduras to Panama;
southward through most of South America; West Indies.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 23
Plants perennial, glabrous or nearly so, usually erect and simple or sparsely
branched, often copiously branched from the base, usually 40 cm. high or less, the
stems tetragonous; stipule sheath very short, the setae about 1.5 mm. long; leaves
sessile or nearly so, linear or lance-linear, mostly 1.5-4 cm. long and 1.5-6 mm. broad,
commonly 1 -nerved, often with fascicles of smaller leaves in the axils; flower heads
chiefly terminal but sometimes arising also from the upper leaf axils, the terminal
heads subtended by 2 or 4 leaflike bracts; hypanthium pilose above, the 2 sepals
narrowly triangular, 1.5 mm. long or less; corolla white, 3 mm. long, hispidulous
outside at the apex, the lobes about equaling the tube; anthers exserted; capsule 2.5
mm. long; seeds brown.
The maya names used in Yucatan are "sac-muy" and "nizots";
"manzanilla de campo" (Yucatan). A century ago the plant was
collected near Acatenango by Hartweg, and reported as in use there
as a remedy for syphilis.
The species resembles B. suaveolens very much but has only
two sepals, the flower heads are smaller and the plant is usually
glabrous or glabrate.
BOUVARDIA Salisbury
References: Standley, Paul C., Bouvardia, in No. Am. Fl. 32:
100-111. 1921. Blackwell, Will H. Jr., Revision of Bouvardia
(Rubiaceae), Ann. Mo. Bot. Gard. 55: 1-30. 1968.
Herbs or shrubs, more or less pubescent, the branches terete or angulate; stipules
interpetiolar, connate with the petioles to form a sheath, entire or lacinate or
aristate; leaves opposite or verticillate, usually petiolate; inflorescence cymose or
cymose-corymbose, flowers usually large, white, yellow, or red, rarely solitary;
hypanthium subglobose or turbinate; calyx 4-(5-)lobate, the lobes short or elongate,
erect or spreading, persistent, often with intermediate teeth; corolla tubular or
salverform, the tube glabrous within or pilose, the limb 4-lobate, the lobes short or
elongate, erect or spreading, valvate in bud, glabrous or pubescent within; stamens 4,
inserted in the throat of the corolla or in the tube above the middle, the filaments
very short or elongate; anthers versatile, linear or oblong, included or exserted; ovary
2-celled, the style filiform, the stigma usually bifid; ovules numerous, crowded, the
placentae peltately affixed to the septum; capsule didymous-globose, coriaceous, 2-
celled, loculicidally bivalvate, the valves finally bifid; seeds numerous, orbicular,
imbricate, peltate, compressed, the testa membranaceous, expanded into a broad
entire wing.
About 20 species mostly in Mexico, but reaching southward to
Honduras and Nicaragua, possibly to Costa Rica. No other species
are found in Central America.
Leaves all or mostly in whorls of 3 or 4.
Corolla pink or lavender, the tube about 5 mm. long; leaves slender-petiolate.
B. bouvardioides.
Corolla bright red, the tube 12-16 mm. long; leaves sessile or nearly so.
B. leiantha.
24 FIELDIANA: BOTANY, VOLUME 24
Leaves opposite.
Corolla red or purplish, the tube 2 cm. long or less.
Flowers umbellate at the apex of the branches, few B. laevis.
Flowers cymose-corymbose or cymose-paniculate, numerous B. dictyoneura.
Corolla white, the tube usually much more than 2 cm. long (sometimes somewhat
shorter in B. multiflora).
Corolla lobes only 3-5 mm. long, the tube about 2 cm. long B. multiflora.
Corolla lobes 6-25 mm. long, the tube usually 2-8 cm. long.
Corolla glabrous outside; leaves glabrous B. longiflora.
Corolla variously pubescent outside; leaves pubescent.
B. longiflora var. induta.
Bouvardia buvardioides (Seem.) Standl. N. Am. Fl. 32: 102.
1921. Hedyotis buvardioides Seem. Bot. Voy. Herald 296. 1856.
Bouvardia pallida Standl. Journ. Wash. Acad. Sci. 14: 245. 1924.
Flor de dolores; jazmin lila.
Occasional on the Pacific slope of the central region, 1,400-1,800
m., on brushy slopes of moist banks; Guatemala; Escuintla;
Sacatepequez. Mexico; El Salvador.
A shrub about 1 m. high or more, often subscandent or pendent from banks, the
young branches minutely puberulent; stipule sheath 2-3 mm. long, subulate-
acuminate and glandular-laciniate; leaves mostly ternate, on petioles as much as 12
mm. long but usually shorter, lanceolate to oblong-ovate, 3-10 cm. long, 1-4.5 cm.
broad, acuminate or long-acuminate, at the base acute or obtuse, thin, sparsely
scaberulous or glabrate above, paler beneath and puberulent, at least along the
nerves; inflorescences cymose-corymbose, 5-12 cm. broad, dense and many-flowered,
the bracts often large and leaflike, the slender petioles 2-4 mm. long, puberulent;
hypanthium hemispheric, puberulent; calyx lobes oblong- linear, 2-3 mm. long; corolla
lavender or pale purple, glabrous outside, the stout tube 5 mm. long dilated upward,
the lobes ovate-oblong, 3 mm. long, obtuse, glabrous within; filaments exserted;
capsule subglobose, 3 mm. broad; seeds broadly winged.
In spite of its abundant flowers, the plant is not a showy one
nor very handsome, the flower color being not at all attractive. The
shrub is plentiful on roadside banks along the road between
Alotenango and Escuintla. We have followed Blackwell in reducing
B. pallida to B. buvardioides with some hesitation. The range is a
disrupted one and the plants from Guatemala and adjacent Chiapas
have some difference which Blackwell has considered minor.
Bouvardia dictyoneura Standl. N. Am. Fl. 32: 109. 1921. B.
matudae Lundell, Lloydia 2: 105. 1939. B. venosissima Lundell,
Bull. Torr. Bot. Club 66: 602. 1939. B. pachecoana Standl. &
Steyerm. Field Mus. Bot. 23: 22. 1943 (type from San Marcos,
Standley 86226).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 25
Wet mixed forest of the western highlands extending eastward
to Volcan de Atitlan, 1,200-3,000 m.; Suchitepe'quez (southern
slopes of Atitlan); Quezaltenango (western slopes of Volcan de
Zunil); San Marcos (Tacana and Tajumulco). Mexico (Chiapas, the
type from Chicharras).
A slender shrub 1-2.5 m. high, the branches glabrous or sparsely pilose at the
nodes; stipules short, subulate; leaves opposite, sessile or on very short petioles,
lance-ovate or oblong-lanceolate to oblong-ovate, 5-11 cm. long, 2-5.5 cm. broad,
narrowly long-attenuate to the apex, rounded or subcordate at the base, rather firm,
very finely and closely reticulate-veined, glabrous or practically so, usually ciliate
when young, the lateral nerves 5-6 on each side; inflorescence cymose-corymbose,
rather dense and many-flowered, the slender pedicels 3-10 mm. long; hypanthium
glabrous, 1-1.5 mm. long; calyx lobes linear-subulate, 2.5-3.5 mm. long; corolla coral-
red or dull red, glabrous outside, the tube about 11 mm. long, the lobes ovate-oval,
3.5-4 mm. long, acutish, ascending; capsule subglobose, 4-5 mm. broad, costate and
reticulate-veined.
Bouvardia laevis Mart. & Gal. Bull. Acad. Sci. Brux. 11(11):
236. 1844. B. nubigena Standl. & Steyerm. Field Mus. Bot. 22: 382.
1940 (type from Jutiapa, Steyermark 31901).
Wet thickets or forest, 1,500-2,000 m.; Jutiapa; Jalapa. Mexico.
A branched shrub 1 m. high glabrous throughout; stipule sheath short, long-
cuspidate and sometimes setiferous; leaves opposite, on petioles 4-8 mm. long, ovate
to broadly ovate or subrhombic-ovate, 3-5 cm. long, 1.5-3 cm. broad, acuminate or
long-acuminate, at the base rounded or obtuse, slightly paler beneath, the lateral
nerves 4 on each side; flowers umbellate at the ends of the branches, usually 3-4, or
arranged in small few-flowered cymes, the pedicels 1 cm. long or less; hypanthium
obovoid, 3 mm. long, obtuse at the base or acutish, the calyx lobes linear, green, 6-7
mm. long or in fruit 10 mm. long, scaberulous on the margins; corolla red, glabrous,
the tube 2 cm. long, 2 mm. broad, the lobes ovate-oblong, suberect, 4 mm. long,
obtuse; capsule subdidymous-globose, 8 mm. broad and 6 mm. high, broadly rounded
at the base.
Bouvardia leiantha Benth. PL Hartw. 85. 1841. Jacinto;
jazmin tinto; clarincillo.
Type collected near Tejar and Chimaltenango, Chimaltenango,
Hartweg 583; common and widely distributed in the mountains,
900-2,100 m., brushy slopes or open banks, often in oak or pine
forest; Baja Verapaz; Zacapa; Chiquimula (near Quezaltepeque);
Jalapa; Jutiapa; Santa Rosa; Guatemala; Sacatepe"quez; Chimal-
tenango; Huehuetenango; Solola; San Marcos. Mexico to Nica-
ragua.
Usually a shrub about 1 m. high, sometimes lower and almost wholly herbaceous,
the slender branches densely puberulent and villosulous at first; stipule sheath 5 mm.
26 FIELDIANA: BOTANY, VOLUME 24
long or less, short-cuspidate, sometimes glandular-dentate; leaves in whorls of 3 or 4,
sometimes opposite, sessile or nearly so, ovate to broadly ovate or ovate-oblong, 3-7
cm. long, 1.5-3 cm. broad, acute or acuminate, rounded at the base, bright green
above and scaberulous or villosulous, short-villous beneath or densely white-
tomentose; inflorescence cymose-corymbose, densely many-flowered, 4-9 cm. broad or
smaller, the pedicels 1-4 mm. long; hypanthium 1.5 mm. long, hirsutulous or glabrate;
calyx lobes lance-linear, 2-4 mm. long, ciliate and often hirsutulous; corolla bright red
or scarlet, glabrous outside, the tube 12-16 mm. long, the lobes rounded-ovate, 2-3
mm. long, obtuse or rounded, erect or ascending; capsule subglobose, about 4 mm.
long.
A rather pretty plant when flowering abundantly, but too often
only a few flowers appear upon a shrub, especially during the dry
months.
Bouvardia longiflora (Cav.) HBK. Nov. Gen. Sp. PL 3: 386.
1820. Aeginetia longiflora Cav. Anal. Cienc. Nat. 3: 130. 1801; Cav.
Icon, t. 572, f. 1. 1801. B. glabra Polak. Linnaea 41: 565. 1877. B.
dolichantha Loesner, Verh: Bot. Ver. Brandenb. 65: 106. 1923 (type,
Huehuetenango, Seler 2883). B. glabra var. obtusa Loesner, I.e.
(type, Seler 2920). Jazmln de monte; jazmin. Figure 11.
Moist or wet thickets or wooded rocky slopes and banks, mostly
1,800-2,000 m.; Alta Verapaz (cultivated); San Marcos;
Quezaltenango; Huehuetenango; Totonicapan; Quiche'; Solold;
Chimaltenango; Jalapa; Jutiapa. Mexico; Honduras; Costa Rica
(cultivated and escape).
A shrub, usually 1-1.5 m. high but often lower, the branches glabrous or nearly
so; stipules 3-6 mm. long, lance-triangular, lobate or laciniate; leaves opposite, on
stout petioles 2-6 mm. long, glabrous, ovate to lanceolate or linear-lanceolate, 2-4.5
cm. long, 0.6-1.8 cm. broad, acute to acuminate, acute to attenuate at the base,
concolorous or nearly so; flowers solitary and terminal, or commonly in 3-flowered
cymes in the upper leaf axils, sessile or short-pedicellate; hypanthium glabrous, 2-3
mm. long, the calyx lobes linear to lanceolate, folia ceous, 6-18 mm. long, scaberulo-
ciliate; corolla white, glabrous, the slender tube 4-8.5 cm. long, 2-3 mm. thick, the
lobes spreading, oblong or oblong-oval, 1.5-3 cm. long, 7-10 mm. broad, acute or
acuminate; capsule subglobose, 8 mm. in diameter, broadly rounded at the apex;
seeds suborbicular, 2-3 mm. long, dark brown, broadly winged.
The large white flowers are fragrant and showy. Bouvardia
glabra is a luxuriant form described from Costa Rica and probably
based on cultivated materials. The senior author, long-time
specialist in the Rubiaceae, apparently thought that the Central
American material and that from Mexico could not be reasonably
separated. The junior author is responsible for the final treatment
and has followed fairly closely Standley's thinking in the matter —
quite different from that of Dr. Blackwell.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 27
Bouvardia longiflora var. induta Robinson, Proc. Am. Acad.
45: 404. 1910. B. induta Standl. No. Am. Fl. 32: 109. 1921. Jazrmn;
jazrmn de monte.
Open wooded slopes, often in pines, 1,000-1,900 m.;
Huehuetenango. Mexico (Chiapas).
Differing from the species in being hirtellous in most parts.
The variation may be only a local and rare one. It is probable
that it is to be found also in the Mexican state of Vera Cruz.
Bouvardia multiflora (Cav.) Schult. Mant. Syst. Veg. 3: 118.
1827. B. heterophylla Standl. No. Am. Fl. 32: 107. 1921 (type from
Santa Rosa, Heyde & Lux 3137). B. latifolia Standl. 1. c. 111. B.
saluadorensis Steyerm. Ceiba 4: 302. 1955. Jazmin de monte.
Moist or dry thickets, 1,000 to 1,500 m. (in Guatemala); Santa
Rosa; Huehuetenango. Mexico; El Salvador; possibly Nicaragua.
A shrub, the slender branches pruinose-puberulent when young; stipules short,
long-cuspidate, sometimes laciniate; leaves opposite, on slender petioles 2-4 mm. long,
suborbicular to broadly ovate or lance-ovate, 2-4 cm. long, 1-3 cm. broad, acuminate
to attenuate, at the base obtuse to truncate and usually abruptly short-decurrent,
bright green and glabrous above, paler beneath, glabrous, or puberulent along the
veins; cymes 3-5-flowered, the slender pedicels 2-8 mm. long; hypanthium 1.5 mm.
long, obscurely pruinose-puberulent; calyx lobes lanceolate or lance-linear, 2.5-5 mm.
long, glabrous; corolla white, glabrous outside, the tube 18-21 mm. long, 2-3 mm.
thick above, the lobes triangular-oblong, 4-5 mm. long, obtuse or acutish, ascending,
glabrous within.
We have followed Dr. Blackwell who considers this species to
be a common one distributed from northern Mexico to Nicaragua.
It is rare in Guatemala.
CALYCOPHYLLUM de Candolle
Trees with terete branchlets; leaves opposite, petiolate; stipules interpetiolar,
narrow, caducous; flowers small, white, in branched terminal corymbiform panicles,
sessile or short-pedicellate, in bud enclosed in membranaceous bracts; hypanthium
oblong, terete, the calyx truncate, but often developing a single large, foliaceous,
petiolate white blade; corolla short-funnelform, the tube short, with villous throat,
the limb 6-8-lobate, the broad lobes imbricate in bud or contorted, one lobe exterior;
stamens inserted in the corolla throat, the filaments subulate, the anthers oblong,
versatile, exserted; ovary 2-celled, the style filiform, glabrous, the 2 branches linear-
oblong, obtuse; ovules numerous, imbricate, the placentae adnate to the septum;
capsule oblong-cylindric, truncate, 2-celled, septicidally bivalvate, the coriaceous
valves entire; seeds few or numerous, imbricate, usually minute, the testa produced at
each end into an elongate wing.
Three other species are known, all South American.
28 FIELDIANA: BOTANY, VOLUME 24
Calycophyllum candidissimum (Vahl) DC. Prodr. 4: 367.
1830. Macrocnemum candidissimum Vahl, Symb. Bot. 2: 38. 1791.
Salamo; cascara de salamo; guayabillo; canela; palo de peine (fide
Rojas); Calan (Izabal, fide Blake); madrono (Izabal); uca; chulub
(reported from Guatemala but not verified by the authors). Figure
16.
Common on the Pacific coastal plains, in forest or open fields,
also in lowland forests of western Guatemala, ascending to about
900 m.; El Progreso; Zacapa; Chiquimula; Jutiapa; Santa Rosa;
Escuintla; Retalhuleu; Huehuetenango. Southern Mexico; El
Salvador to Panama and Colombia; Cuba.
A tall tree, sometimes 20 m. high or more, with a rather slender trunk, often
supported by small buttresses, the pale bark similar to that of guava; branchlets
glabrous or nearly so, often hirsute at the nodes; stipules triangular or lanceolate, 1
cm. long or less; petioles 0.5-3.5 cm. long; the blades oval or ovate, 5-12 cm. long, 1.5-
7.5 cm. broad, abruptly acuminate or cuspidate-acuminate, acute or obtuse at the
base and abruptly short-decurrent, bright green and glabrous above, paler beneath
and usually sparsely hirsute on the costa; corymbs few- or many-flowered, the
flowers cymose-glomerate, the large brown bracts caducous; hypanthium 2.5-3 mm.
long, setose-hirsute or glabrate; calyx in many of the flowers expanded into a white
limb, this ovate-rhombic to subreniform, 1.5-3 cm. long, rounded at the apex, obtuse
to subcordate at the base and abruptly decurrent into a petiole 1-1.5 cm. long; corolla
white, the tube 3 mm. long, glabrous or sparsely puberulent outside, the lobes slightly
longer than the tube, obtuse or subacute, the throat densely white-villous; stamens
long-exserted; capsule 6-10 mm. long, 2.5-4 mm. thick, compressed and shallowly
bisulcate, costate, acute at the base; seeds produced at each end into a short acute
wing.
One of the common trees of the Pacific plains, and of the
Pacific coast generally of Central America. When in flower, at the
end of the rainy season, it is exceedingly conspicuous and handsome
because of the great abundance of enlarged calyx lobes which give
the effect of a dense mantle of white flowers. These lobes are at first
creamy, then almost pure white, and remain so for a long time, but
finally turn brown and persist upon the tree for many weeks,
sometimes after the leaves have fallen, giving it then a withered
appearance. The name "guayabillo" refers to the fact that the bark
is so much like that of Psidium guajava in general appearance. In
some parts of Honduras the tree is called "Colorado," and in Costa
Rica "surra," "salamo," and "madrono." The wood is pale brown,
hard, heavy, strong, highly elastic, fine-textured, usually straight-
grained, easy to work and finishes smoothly. Under the name
"lance wood" it is imported from Cuba into the United States for
use in making archery bows. In Guatemala, as elsewhere in Central
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 29
America, it is utilized for making fine-toothed combs, and also for
tool handles and other purposes for which a strong, fine-grained,
and hard wood is required.
CEPHAELIS Swartz
Reference: Molina R, Antonio, Revision de las Especies de
Cephaelis de Mexico, Centre America y las Antillas, Ceiba 4: 1-38.
1953.
Shrubs or small trees, rarely herbs, glabrous or pubescent; stipules free or
connate, usually persistent; leaves opposite, petiolate or sessile; flowers capitate,
subtended by an involucre of usually sessile, free or connate, often brightly colored
bracts, the heads terminal or axillary, simple or branched, sessile or pedunculate;
calyx short or elongate, persistent, 4-7-dentate; corolla funnelform or salverform, the
tube straight, normally elongate, the throat villous or naked, the lobes 4-5, valvate in
bud; stamens inserted in the corolla throat, the filaments generally short, the linear
anthers included or exserted; ovary commonly 2-celled, the cells 1-ovulate; fruit
drupaceous, containing 2 bony, smooth or costate nutlets, these mostly longitudinally
sulcate on the flat inner face.
A large genus of tropical America and Africa, most of the
species South American; about 15 are known from Central
America. C. ipecacuanha (Brot.) A. Rich., of the lowlands of
Panama and northern South America, furnishes part of the ipecac
of commerce. It has been collected as far north as Nicaragua, and is
usually given the name "raicilla." The drug is obtained from the
slender knotted roots, which are about 6 mm. in diameter. The
plant is a low simple shrub with subsessile leaves and stipules
divided into thread-like lobes. The genus Cephaelis is not a well
marked one and is difficult to separate from Psychotria, but for
practical purposes, especially in case of the very numerous South
America species, it constitutes a most convenient segregate.
Plants densely pilose or hirsute; flower heads terminal, pedunculate C. tomentosa.
Plants glabrous or practically so.
Leaves sessile or practically so C. chiapensis.
Leaves conspicuously petiolate.
Flowers heads long-pedunculate, terminal C. elata.
Flower heads sessile.
Heads all terminal C. glomerulata.
Heads mostly axillary, sometimes also terminal C. axillaris.
Cephaelis axillaris Swartz, Prodr. Veg. Ind. Occ. 45. 1788.
Wet forest, mountains of Zacapa (Sierra de las Minas) and
Chiquimula (Cerro Brujo), at 1,700-2,500 m. Extending southward
to Panama; Colombia; Venezuela; West Indies.
30 FIELDIANA: BOTANY, VOLUME 24
A glabrous shrub 1.5 m. tall or less, with green stems, simple or sparsely
branched; stipules bilobate, the lobes obtuse or acute, persistent; leaves on short
slender petioles, obovate-oblong or oblong-oblanceolate, mostly 9-13 cm. long and 2.5-
4 cm. broad, gradually or abruptly acuminate, acute or attenuate at the base, pale
beneath; flower heads chiefly or wholly axillary, closely sessile, about 1 cm. in
diameter or in fruit larger, the numerous small bracts green or purplish; corolla white
or creamy white, inconspicuous; fruits turquoise or prussian blue.
Cephaelis chiapensis Standl. Field Mus. Bot. 4: 295. 1929.
Evea chiapensis Standl. Contr. U. S. Nat. Herb. 23: 1392. 1926
(type from Cerro del Boqueron, Chiapas).
Known in Guatemala only from Bernoulli & Carlo 1710,
collected in 1877 and labeled as coming from Retalhuleu; probably
to be found on the wetter slopes of the mountains above
Retalhuleu.
A glabrous shrub with slender branches; stipule sheath very short, bearing on
each side 2 stiff setae about 3 mm. long; leaves sessile or nearly so, thin, oblong-
elliptic or oblong-lanceolate, 7-11 cm. long, 2.5-4 cm. broad, acute to long-acuminate,
narrowed to the obtuse or acute base, green and lustrous above, paler beneath, the
lateral nerves about 9 pairs; heads terminal, on peduncles 1 cm. long or less, rather
few-flowered; outer bracts few, apparently pale green, broadly ovate or deltoid-ovate,
10-15 mm. long, acute or acuminate, thin.
Cephaelis elata Swartz, Prodr. Veg. Ind. Occ. 45. 1788.
Cephaelis punicea Vahl, Eclog. Amer. 1: 19. 1796. Evea elata
Standl. Contr. U. S. Nat. Herb. 18: 123. 1916.
Wet forest, at scattered localities: Izabel and Alta Verapaz, at
or little above sea level; Chiquimula (Cerro Tixixi, 500-1,500 m.);
Suchitepe"quez (Volcan de Zunil, 1,200 m.); San Marcos (Volcan de
Tajumulco, at 1,300-1,500 m.). Southern Mexico and British
Honduras through much of Central America to West Indies;
Colombia.
A glabrous shrub or small tree, 1.5-6 m. tall; stipules short, the lobes broad,
broadly rounded at the apex; leaves slender-petiolate, oblong- lanceolate to elliptic-
oblong, 10-25 cm. long, 3.5-7 cm. broad, acuminate, acute to attenuate at the base,
with numerous pairs of lateral nerves; peduncles terminal, usually greatly elongate,
bearing 1 or sometimes 3 large heads, the 2 outer bracts rose-red or dull dark red,
rounded-ovate, 2.5-4 cm. long, broadly rounded to acutish at the apex, rarely green or
yellow (f. lutea Standl., of Costa Rica); corolla white, well exserted from the bracts
but soon deciduous, its lobes spreading.
The shrub is a handsome and showy one because of the
brightly colored bracts of the inflorescence.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 31
Cephaelis glomerulata Donn.-Sm. Bot. Gaz. 16: 12, t. 1. 1891.
Psychotria glomerulata Steyerm. Mem. N. Y. Bot. Card. 23: 670.
1972. Figure 52.
Type from swampy woods of Rio Dulce, Izabal, at sea level, J.
D. Smith 1637; wet forests of Izabal and Pete"n, at or little above sea
level. Along or near the Atlantic coast from British Honduras to
Panama; Colombia.
A glabrous shrub 1-2 m. high, sparsely branched; stipules united to form a short
truncate sheath; leaves short-petiolate, somewhat coriaceous, oblong-lanceolate or
elliptic-oblong, 9-15 cm. long, 3.5-5 cm. broad, rather abruptly acuminate, acute at
the base, with about 14 pairs of lateral nerves; flower heads terminal, sessile, about 2
cm. broad and 1.5 cm. high, the bracts pale green, coriaceous, the broad outer ones
obovate, rounded at the apex, the inner ones spatulate; calyx unequally subulate-
dentate; corolla white, exserted beyond the bracts; fruit ovoid, 6 mm. long, blue.
Cephaelis tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19. 1796.
Tapogomea tomentosa Aubl. PL Guian. 160. 1775. Evea tomentosa
Standl. Contr. U. S. Nat. Herb. 18: 123. 1916. Madre (Peten).
Common in wet forest or second-growth thickets, Izabal, at or
near sea level; Peten; Alta Verapaz (Chirriacte' and Cubilguitz, on
limestone at 300 m.); Quiche" (Finca Chaila, Zona Reina, 330 m.);
Huehuetenango (Ixcan). British Honduras. Southern Mexico to
Bolivia and Amazonian Brazil.
A slender shrub commonly 1-2 m. tall, often almost wholly herbaceous, sparsely
branched, densely pilose or hirsute throughout; stipules persistent, deeply cleft into
narrow erect lobes; leaves membranaceous, short-petiolate, lanceolate to ovate-
elliptic, mostly 8-16 cm. long, narrowly long-acuminate, acute to attenuate at the
base; heads large, terminal, on long or short peduncles; bracts broadly ovate, 3-5 cm.
long, deep dull red, long-connate above the base, acute or short-acuminate; corollas
yellow, shorter than the outer bracts; fruit blue.
Called "oropelo" in Oaxaca and "cresta de gallo" in Tabasco. A
showy and rather handsome shrub of wide range in tropical
America, and certain to attract the attention of every amateur
collector. Actually, it is perhaps showier in herbarium specimens
than when growing, for often the plants are half hidden among
other shrubs or weeds. It may be seen in abundance along the
railroad passing through the wet lowland forest between Puerto
Barrios and Gualan.
CEPHALANTHUS Linnaeus
Erect shrubs or small trees, glabrous or pubescent; leaves opposite or verticillate,
short-petiolate; stipules short, intrapetiolar; inflorescence globose, flowers small,
32 FIELDIANA: BOTANY, VOLUME 24
white or yellowish, sessile and forming very dense, many-flowered heads, intermixed
with setaceous or paleaceous bractlets; hypanthium turbinate; calyx short-tubular,
unequally 4-5-dentate or 4-lobate, often glanduliferous; corolla tubular-funnelform,
the throat glabrous or pilose, the limb 4-5-lobate, the lobes erect or spreading,
imbricate in bud, one lobe exterior, often with glands in the sinuses; stamens 4,
inserted in the corolla throat, the filaments short, the anthers dorsifixed, bicuspidate
at the base; ovary 2-celled, the style filiform, the stigma capitate or clavate; ovules
solitary, pendulous from the top of the cell; fruit turbinate, 2-4-celled, the cells
indehiscent, 1-seeded, or one or more of them empty; seeds oblong, covered at the
apex by a white aril.
The genus is a small one of about seven species in America,
Africa, and Asia. Only two are known from North America, one or
two in South America.
Calyx glabrous or sparsely pubescent at the base C. occidentalis.
Calyx densely sericeous pubescent with short hairs C. salicifolius.
Cephalanthus occidentalis L. Sp. PI. 95. 1753. Guayabillo.
Figure 21.
Swamps near sea level; Pete*n; Izabal. British Honduras
(Maskall); Honduras; through Mexico to the United States and
New Brunswick; Cuba; Old World.
In Guatemala a shrub of 1.5 m., but often much larger or even a small tree
(reported in Honduras as 5.5 m. high); stipules 2-4 mm. long, deltoid, acute or
acuminate, usually with glands along the margin; leaves opposite or ternate, the
petioles mostly less than 2 cm. long, the blades ovate to rarely narrowly lanceolate,
10-15 cm. long, 8.5 cm. broad or less, long- or short-acuminate, subcordate to acute at
the base, glabrous above or nearly so, beneath almost glabrous or sometimes densely
pubescent (var. pubescens Raf.); peduncles terminal and axillary, simple or branched,
3-10 cm. long; heads 6-12 mm. in diameter (excluding the corolla); bractlets filiform-
clavate; hypanthium and calyx together 2-3 mm. long, glabrous, or sparsely long-
pilose at the base, the calyx about 1 mm. long, shallowly dentate; corolla 5-9 mm.
long, white or cream, glabrous outside, the limb with a small black gland in each
sinus; capsule 4-8 mm. long.
Known in the United States by the name "button-bush." The
flowers are sweet-scented.
Cephalanthus salicifolius Humb. & Bonpl. PI. Aequin. 2: 63.
1809.
To be expected in the pine or pine-oak forests at 700-1,500 m. or
perhaps along river banks down almost to sea level. Mexico (north
and central); Honduras. Not now known from Guatemala or British
Honduras.
Shrubs or small trees; leaves opposite or ternate, short petiolate, glabrous, linear-
lanceolate to narrowly oblong, 4.5-12 cm. long and 1-2.5 cm. broad, mostly acute or
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 33
acuminate at the apex and acute or rounded at the base; peduncles mostly 2-4 cm.
long; the inflorescence capitate or globose, 2-3 cm. in diameter at anthesis; calyx and
hypanthium 2-2.5 mm. long, sericeous with short hairs, the calyx about 1 mm. long, 4-
5-lobate; corolla 6-8 mm. long, glabrous outside, sparsely pilose within, usually with
small black glands in the sini.
CHIOCOCCA P. Browne
Shrubs or small trees, often scandent or clambering, glabrous or pubescent,
branches terete; stipules broad, persistent, usually cuspidate. Leaves opposite,
petiolate, membranaceous to coriaceous; inflorescence of axillary racemes, simple or
paniculate, often secund; flowers small, whitish, pedicellate; hypanthium ovoid or
turbinate, compressed; calyx 5-lobate, persistent, the lobes short; corolla funnelform,
the throat glabrous, the limb 5-lobate, the lobes valvate, spreading or reflexed;
stamens 5, inserted on the disk inside the corolla, the filaments pilose; anthers linear,
basifixed, included or exserted; ovary two-celled, the style filiform with clavate or
cylindric stigma which is entire or bilobate; ovules solitary, pendulous from the apex
of the cell; fruit drupaceous, small, somewhat fleshy, usually orbicular and much
compressed, bicarpidiate.
This genus is in need of revision and most certainly there are
more species in Guatemala than the five given here. Chiococca alba
(L.) Hitchc. as treated here is certainly a composite species. The
genus is a common one through most of northern Central America,
perhaps one of the commonest members of the Rubiaceae at low
and middle elevations and is to be found in almost any moist ravine
or creek valley in Guatemala, British Honduras, or Honduras.
The genus perhaps contains 20 species or more. Only those
listed here have been proposed for Central America and Panama.
Leaves lance-linear, only 2-4 mm. broad C. steyermarkii.
Leaves lanceolate to ovate, much broader.
Leaves pilose on both surfaces C. semipilosa.
Leaves glabrous.
Anthers nearly or quite equaling the corolla lobes, sometimes longer, the
filaments exserted . C. phaenostemon.
Anthers only slightly exceeding the corolla tube, or included, the filaments
wholly included.
Limb of the corolla 8-10 mm. broad; calyx lobes semiorbicular to almost
obsolete; leaf blades mostly 3.5-6 cm. broad; fruit but little compressed.
C. pachyphylla.
Limb of the corolla 4-6 mm. broad; calyx lobes deltoid to lance-linear, acute or
acutish; leaf blades mostly narrower; fruit strongly compressed. C. alba.
Chiococca alba (L.) Hitchc. Kept. Mo. Bot. Gard. 4: 94. 1893.
Lonicera alba L. Sp. PI. 175. 1753. C. racemosa L. Syst. Nat. ed. 10.
917. 1759. C. macrocarpa Mart. & Gal. Bull. Acad. Brux. 11, pt. 1:
231. 1844. C. petenensis Lundell, Wrightia 5: 7. 1972 (type from
34 FIELDIANA: BOTANY, VOLUME 24
Peten, Contreras 2439). C. vestita Lundell, I.e. 8 (type from Pete*n
Lundell 16425). C. vestita var. glaberrima Lundell, I.e. 9 (type from
Peten, Lundell 16425a). Lagrimas de Guadalupe (fide Aguilar).
Common in thickets and forest in many regions, especially in
the tierra caliente of both coasts, ascending to about 1,500 m.;
Peten; Alta Verapaz; Izabal; Chiquimula; Jalapa; Jutiapa; Zacapa;
Santa Rosa; Esquintla; Guatemala; Sacatepequez; Suchitepe'quez;
Quezaltenango; Huehuetenango. Florida and southwestern Texas;
Mexico; British Honduras to El Salvador and Panama; West
Indies; widely distributed in South America.
Usually a shrub, rarely a small tree, often scandent, the branchlets glabrous or
obscurely puberulent; stipules 1-2 mm. long, mucronate or subulate-acuminate;
petioles 2-10 mm. long, the blades very variable, mostly ovate or oval-ovate,
sometimes lanceolate, 3-9 cm. long, 1-4.5 cm. broad, commonly short-acuminate,
rounded and short-decurrent to acute at the base, the lateral nerves inconspicuous;
inflorescence racemose or paniculate, with few or many flowers; calyx and
hypanthium 2.5 mm. long, the calyx lobes subulate to broadly deltoid; corolla 6-8
mm. long, white or cream; anthers included or the tips rarely exserted; fruit white,
juicy, orbicular, 4-8 mm. long, strongly compressed; seeds dark brown, 3-4 mm. long,
puncticulate.
Called "lagrimas de San Pedro" and "aceitillo" in El Salvador;
"cainca" and "canchacche" in Yucatan. There is a possibility that
more than one species is represented by Guatemalan specimens.
Chiococca pachyphylla Wernham, Journ. Bot. 51: 323. 1913.
C. belizensis Lundell, Am. Midi. Nat. 29: 492. 1943 (type from Cow
Pen, near Monkey River, Toledo District, British Honduras, Gentle
4115). C. rubriflora Lundell, Wrightia 5: 7. 1972 (type from Pete"n,
Contreras 9110).
Moist mountain forest, 1,500 m. or lower; Pete'n; Such-
itep^quez; San Marcos; Huehuetenango. Mexico; British Honduras;
El Salvador; Honduras; Nicaragua.
A scandent shrub (in Guatemala), sometimes erect or a small tree, glabrous
except for the inflorescence; stipules 2-4.5 mm. long, subtruncate and mucronate;
petioles 8-14 mm. long, thick, the blades elliptic to broadly oblong or ovate, 7-14 cm.
long, 2.5-6 cm. broad, short-acuminate to obtuse, rounded to acute at the base and
abruptly short-decurrent, coriaceous, the lateral nerves rather prominent beneath;
inflorescence many-flowered, paniculate, often equaling or exceeding the leaves, the
branches obscurely puberulent or glabrous, the pedicels 2-5 mm. long; calyx and
hypanthium 2.5-3 mm. long, the calyx lobes very short; corolla 7-8 mm. long, often
reddish outside, white or cream within; anthers semiexserted; fruit subglobose,
usually little compressed, 6-7 mm. in diameter, white or greenish white.
Called "snowberry" in British Honduras.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 35
Chiococca phaenostemon Schlecht. Linnaea 9: 594. 1834.
Trueno (Huehuetenango).
Widely distributed, at 1,300-3,000 m., in humid forest or
thickets, sometimes on open rocky hillsides; Alta Verapaz; Baja
Verapaz; Chiquimula; Quiche"; Sacatepe"quez; Huehuetenango; San
Marcos; Quezaltenango; Chimaltenango; Quiche". Southern Mexico
to Honduras.
A shrub or small tree, often 5-8 m. tall, commonly erect and with no tendency to
climb, glabrous or nearly so; stipules 2-3 mm. long, mucronate or subulate-cuspidate;
petioles 3-10 mm. long, the blades mostly ovate-oblong or lance-oblong, 4-12 cm. long,
1.5-5.5 cm. broad, acuminate or long-acuminate, rounded to acutish at the base and
often short-decurrent, coriaceous, the lateral nerves obscure beneath; racemes usually
panicled, often longer than the leaves, many-flowered, the pedicels 5 mm. long or less;
calyx and hypanthium 2-3 mm. long, the calyx lobes short, broadly deltoid, obtuse or
acutish; corolla 5-8 mm. long, yellowish white; anthers 3-4 mm. long; fruit white,
compressed, 5-7 mm. long.
Growing as a small tree, this is plentiful at many places of the
central region, as on the lower slopes of Volcan de Acatenango, near
Chimaltenango.
Chiococca semipilosa Standl. & Steyerm. Field Mus. Bot. 22:
279. 1940. Figure 42.
In thickets or forest, 400-1,400 m.; endemic; Chiquimula (type
from Cerro Caracol north of Quezaltepeque, Steyermark 31406; also
at several other localities); Jalapa; Huehuetenango. Honduras.
A weak subscandent shrub 1.5-2 m. long, the branchlets glabrous; stipules
minute, cuspidate; petioles 2-3 mm. long, the blades firm-membranaceous. lanceolate
to ovate, 5-8 cm. long, 1.5-3 cm. broad, long-acuminate, acute and often decurrent at
the base, sparsely spreading-pilosulous above, densely short- hirtellous beneath;
racemes mostly 5-7-flowered, half as long as the leaves or less, the rachis hirtellous,
the slender pedicels 3-7 mm. long; calyx 1 mm. long, deeply dentate, the teeth
triangular, acute; fruit oval-orbicular, somewhat compressed, 4-5 mm. long, white,
sparsely hirtellous.
Chiococca steyermarkii Standl. Field Mus. Bot. 22: 280. 1940.
Type from San Marcos, Rio Vega, near San Rafael and
Guatemala-Mexico boundary, at 2,500-3,000 m., Steyermark 36237.
An erect, rather densely branched shrub 2-3 m. high, the branchlets puberulent;
stipules subulate from a broad base; leaves small, subcoriaceous, the petioles scarcely
1.5 mm. long, the blades lance-linear, 15-23 mm. long, 2-4 mm. broad, gradually
narrowed to the obtuse apex, acute at the base, glabrous, the lateral nerves obsolete;
flowers axillary and solitary or short-racemose, the pedicels filiform, recurved, 5-8
mm. long, minutely puberulent; hypanthium glabrous; calyx teeth subulate, 1 mm.
36 FIELDIANA: BOTANY, VOLUME 24
long; corolla white, 10-11 mm. long, glabrous; anthers included; fruit white,
subcompressed, suborbicular, 7 mm. long.
The species is perhaps the best marked one of the whole genus,
on account of its small narrow leaves.
CHIONE De Candolle
Trees or shrubs, glabrous or nearly so; stipules small, caducous, commonly short-
connate; leaves opposite, coriaceous, petiolate; flowers small, white, in terminal
pedunculate cymes or corymbs; hypanthium turbinate; calyx cupular, 5-dentate, 5-
lobate, or subentire, persistent; corolla funnelform, short and broad, the throat
naked, the limb 5-6-lobate, the lobes imbricate in bud, 2 of them exterior; stamens 5-
6, inserted above the base of the corolla tube, the filaments stout, the anthers large,
exserted, linear-oblong, dorsifixed; ovary 2-celled, the style stout, the branches linear-
oblong, obtuse, exserted; ovules solitary, pendulous from the apex of the cell; fruit
drupaceous, ovoid or ellipsoid, the stone osseous, sulcate, 2-celled; seeds elongate,
terete.
About 15 species in tropical North America. Six or seven
species, Mexico to Panama, the others in the West Indies. Chione
chiapasensis Stand!., described from Palenque, Chiapas, may well
be expected in Peten.
Chione guatemalensis Standl. & Steyerm. Field Mus. Bot. 22:
174. 1940. Figure 44.
Known only from the vicinity of the type locality, banks of Rio
Dulce above Livingston, Izabal, at sea level; type, Steyermark
39375.
A shrub or small tree, glabrous throughQut; stipules caducous; leaves on stout
petioles- 2-2.5 cm. long, subcoriaceous, narrowly elliptic-oblong to elliptic, 14-17 cm.
long, 4.5-8 cm. broad, shortly obtuse-acuminate, acute at the base, with about 9 pairs
of lateral nerves; inflorescence long-pedunculate, terminal, few-flowered cyme, much
shorter than the leaves, the flowers on rather long, stout pedicels; hypanthium
clavate, the calyx 2 mm. long, its lobes triangular, acute, erect; corolla white, 5 mm.
long, the broad tube obconic, the broad rounded lobes less than half as long as the
tube, recurved; filaments short-exserted, the oblong-linear anthers 3 mm. long; fruit
oblong, obtuse at base and apex, fleshy, greenish, turning mango-orange, red in the
upper half, 2 cm. long, 8-9 mm. broad.
In C. chiapasensis the leaves are excavate and barbate beneath
in the nerve axils; in C. guatemalensis the axils are neither excavate
nor barbate.
CHOMELIA Jacquin
Shrubs or small trees, usually armed with long stout axillary spines, the
branchlets terete; leaves opposite, petiolate, commonly membranaceous; stipules
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 37
interpetiolar, deciduous, acuminate; flowers small, white, bracteolate, in pedunculate
axillary cymes; hypanthium oblong or turbinate; calyx 4-lobate, the lobes narrow
and elongate, equal or unequal, persistent; corolla salverform, with slender elongate
tube, the throat glabrous or pilose, the limb 4-lobate, the lobes valvate in bud or with
slightly imbricate margins, corniculate-appendaged outside near the apex; stamens 4,
inserted in the corolla throat; anthers linear or sagittate, dorsifixed, sessile, included
or subexserted, their basal lobes acute or attenuate; ovary 2-celled, the style filiform,
with 2 short branches; ovules solitary, pendulous from the apex of the cell; fruit
drupaceous, small, oblong, subterete, the stone osseous, 2-celled; seeds pendulous,
cylindric.
A genus of few species, all except the following one South
American. Many authors unite Anisomeris with Chomelia, here
tentatively maintained as distinct. That differs in having no
appendages on the corolla lobes.
Chomelia spinosa Jacq. Enum. PL Carib. 12. 1760; Stirp. Am.
18. 1763. Chomelia filipes Benth. ex Oerst. Vid. Medd. Kjoebenhavn
41. 1852 (type from Granada, Nicaragua). Anisomeris purpusii
Brandegee, Univ. Calif. Publ. Bot. 6: 70. 1914 (type from Chiapas).
C. purpusii Rusby, Bull. Torr. Bot. Club 52: 138. 1925.
Known in Guatemala only from brushy coastal plains of
Escuintla (San Jos6) from a sterile specimen, but doubtless may
occur elsewhere along the Pacific coast; growing in dry or moist
thickets or forest. Southern Mexico, along the Pacific coast to
Panama, southward to northern Brazil.
A shrub or small tree, sometimes 9 m. tall, the young branches mostly brown,
armed with stout spines 2.5 cm. long or less, the young branchlets appressed-pilose;
stipules 4-8 mm. long, thin, sparsely pilose, subulate-acuminate from a triangular
base; leaves often crowded on short lateral branches, the slender petioles 12 mm. long
or less, the blades very variable in shape, ovate-orbicular to elliptic or oblong-ovate,
4-8 cm. long, 2-4 cm. broad, usually abruptly short-acuminate, broadly rounded to
acute at the base, thinly pilose and green above, pilose beneath, especially along the
nerves, with appressed or spreading hairs, with 6-8 pairs of prominent lateral nerves;
cymes few-flowered, the very slender peduncles 1.5-3.5 cm. long, the flowers sessile or
subsessile; calyx and hypanthium 2-3 mm. long, appressed-pilose; calyx lobes
lanceolate or triangular-subulate, unequal, half as long as the hypanthium or shorter;
corolla white or creamy white, sericeous outside, the slender tube 12-22 mm. long, the
linear-lanceolate lobes 4-6 mm. long, each with a slender elongate horn-like
appendage at or just below the apex, the throat glabrous; fruit black or purplish
black at maturity and juicy, oval-oblong, 9-12 mm. long.
The general appearance of the shrub suggests the genus
Randia, except for the flowers and fruit.
38 FIELDIANA: BOTANY, VOLUME 24
CINCHONA Linnaeus
Reference: Popenoe, Wilson, Cinchona, The "Fever-Tree," in
Wilson, New Crops for the New World, Macmillan Co., pp. 109-125.
1945.
Trees or shrubs, the branches terete or tetragonous, the bark bitter; leaves
persistent, opposite, petiolate, coriaceous or membranaceous; stipules interpetiolar,
glandular within at the base, deciduous; flowers small, white, pink, or purplish,
fragrant, normally 5-parted but often 4- or 6-parted, in small or large, terminal
panicles; hypanthium turbinate, pubescent, the calyx persistent, cupular, 5-dentate;
corolla salverform, pubescent, the tube terete or somewhat 5-angulate, glabrous or
pilose in the throat, the limb 5-lobate, the lobes spreading, not papillose within,
valvate in bud, the margins pilose; stamens 5, inserted in the corolla tube, the
filaments short or elongate; anthers included or their apices exserted, linear,
dorsifixed; ovary 2-celled, the style slender, its branches short, obtuse, included or
subexserted; ovules numerous, the placentae linear, adnate to the septum, upwardly
imbricate; capsule ovoid, oblong, or subcylindric, bilobate, 2-celled, many-seeded,
septicidally dehiscent from the base upward to the apex; seeds peltate, the testa
broadly winged.
Perhaps 20-25 species, ranging from Costa Rica southward,
chiefly along the Andes, to Bolivia. Authors are not in agreement as
to the number of species to be recognized, principally because of the
difficulty in finding characters for their separation. Most of the
species have been based upon minute characters, some of which
appear to be of horticultural rather than systematic importance.
The genus is one of the most important of all American Rubiaceae
because it is the source of the drug quinine or cinchona, formerly
used generally in the treatment of malaria, the greatest scourge of
the tropics.
The virtues of cinchona bark as a remedy for malaria were first
made known to Europe around 1600 from the center of Loja,
Ecuador. Apparently, the Indians of the Andes had little knowledge
of the properties of the tree, except possibly in the case of those
living in the vicinity of Loja. Quina, the Spanish name of the trees
and their product is derived from the Quechua name quina-quina of
that locality. It was in 1638 that cinchona bark first attained a
reputation among the Spaniards of Peru, for in that year the
Countess of Chinchon, wife of the Viceroy, was cured of a tertian
fever by the use of the so-called Peruvian bark. When the Countess
returned to Spain in the spring of 1640, she took with her a
quantity of the bark, and from this beginning its value soon became
widely known in Europe, where it was for a time called Pulvis
Comitassae (Countess' powder), under which name it was long
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 39
known to druggists. Dr. Juan de Vega, physician of the Countess,
followed his patient to Spain, taking with him a quantity of quina
bark, which he sold for 100 reales the pound. The Countess
administered Peruvian bark to the sufferers from tertian agues on
her husband's estates in the fertile but unhealthy valleys of the
Tagus, Jarama, and Tajuna in Spain, so that her good deeds became
traditional in the region where she lived. It was therefore quite
fitting that Linnaeus dedicated to this lady the genus that yields
Peruvian bark. The source of his knowlege of her work was a foreign
and not a Spanish source, hence the name was published not as it
should have been, Chinchona, but as Cinchona. Such a garbling of
Spanish words in botanical literature of foreign countries is,
unfortunately, even more frequent today, perhaps, than in
Linnaeus' time!
Quinine was for more than three centuries the only drug
available for the control of malaria and, consequently, was
exceedingly important to people living in regions where malaria was
endemic and often epidemic. At the time of Vv^rld War II, the
discovery of synthetics, especially atabrine (quinacrine) that would
control malaria reduced reliance on quinine as the only effective
antimalarial. Cinchona plantations that had been started in
tropical America and Africa were essentially abandoned.
The story of Cinchona cultivation in Guatemala is well told by
Dr. Popenoe in the reference given above. For a more detailed
account of the history of the genus Cinchona see Standley, Field
Mus. Bot. 7: 188-197. 1931.
Corolla 14-17 mm. long; leaves mostly firm and even coriaceous, relatively small and
narrow.those of fertile branches mostly 3-4 cm. broad, glabrous or nearly so.
C. officinaUs.
Corolla 10-12 mm. long; leaves thin, large, commonly 10-20 cm. broad, often or
usually abundantly pubescent, at least beneath C. pubescens.
Cinchona officinalis L. Sp. PL 172. 1753. Quina.
A native of the South American Andes, ranging from Colombia
to Bolivia. Formerly planted extensively in the bocacosta of
Escuintla and Suchitepe"quez, in moist forests of northern
Huehuetenango, in small amounts in Alta Verapaz, and probably in
other departments.
A large or medium-sized tree, sometimes flowering when only a shrub, the bark
rugose, fuscous, the branchlets strigillose-pilosulous; stipules large lanceolate or
oblong, acute or obtuse, glabrous; leaves petiolate, comparatively small, mostly
lanceolate to oblong or elliptic-oblong but varying to elliptic or ovate, acute,
40 FIELDIANA: BOTANY, VOLUME 24
acuminate, or obtuse, at the base rounded to attenuate, coriaceous, glabrous above
and often lustrous, at least in the dry state, commonly glabrous beneath or nearly so
but sometimes puberulent or short-pilose, especially on the veins, usually about 10
cm. long and 3.5-4 cm. broad, but leaves of sterile branches often much larger, often
domatiate beneath in the nerve axils; panicles terminal, leafy, rather small and dense
or sometimes large and more open; hypanthium strigose; calyx glabrous or nearly so,
the teeth triangular, acute, reddish; corolla pink or red, sericeous outside, the tube
about 1 cm. long, the lobes ovate, acute; capsule oblong, generally 1.5-2 cm. long,
glabrate.
The form which was planted in Guatemala for exploitation is
Cinchona ledgeriana Moens (ex Trimen, Journ. Bot. 19: 323. 1881),
a native of Bolivia. This is regarded by some authors as a distinct
species, and was considered the best and principal source of the
drug quinine. Both the names C. officinalis and C. pubescens are
used here in their broad sense, C. ledgeriana thus being considered
a form or variety of the true C. officinalis.
Cinchona pubescens Vahl, Skrivt. Naturh. Selsk. 1: 19. 1790.
C. succirubra Pa von ex Klotzsch, Abh. Akad. Berl. 1857: 60. 1858.
Quina; quin (Quecchi).
Common in fincas of the Pacific bocacosta and in the Coban
region of Alta Verapaz; most of the trees doubtless planted, but in
many places seeding abundantly and becoming naturalized. Native
of the Andes, from Colombia to Bolivia, and in Costa Rica.
A medium-sized tree, often flowering when only a large shrub, the branchlets
pubescent; stipules large, ovate, obtuse or acute, sericeous or almost glabrous; leaves
large, slender-petiolate, mostly broadly ovate to orbicular, rounded to acute at the
apex, cordate to acute at the base and often decurrent, usually glabrate above,
beneath densely short-pilose or tomentose to glabrate, often domatiate in the nerve
axils; panicles usually large and many-flowered, often leafy, the flowers subsessile;
hypanthium densely sericeous; calyx appressed-pilosulous, the teeth short and broad,
acute; corolla red or pink, sericeous, the lobes half as long as the tube; capsule
lanceolate or oblong, glabrate, commonly 1.5-2.5 cm. long.
A highly variable species as here treated, but the forms are
separable by only minor characters which baffle systematic
segregation. The form occurring in Guatemala (as well as in Costa
Rica) is supposed to be C. succirubra, but in what respect, if any,
that differs from typical C. pubescens is uncertain, to say the least.
COCCOCYPSELUM P. Browne
Prostrate perennial herbs, pubescent or sometimes glabrous; leaves opposite,
petiolate, herbaceous, often purple or blue below; stipules small and inconspicuous,
often with digitiform calluses in the axils; inflorescence capitate, axillary, solitary,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 41
pedunculate or sessile; flowers small, soon falling away, white to purplish;
hypanthium ovoid or turbinate; calyx 4-lohate, the lobes narrow, persistent; corolla
4-lobate, funnelform, the lobes oblong, valvate in bud, the throat glabrous; stamens
4, inserted in the corolla tube, the filaments short, the anthers dorsifixed near the
base, oblong-linear, short-exserted or included; ovary 2-celled, the style short, with 2
short or somewhat elongate branches; ovules numerous, horizontal, the placentae
adnate to the septum; fruit baccate, ovoid or globose, 2-celled, blue; seeds numerous,
orbicular, subangulate, plano-convex, the testa granulate.
The genus Coccocypselum consists of perhaps some 20 species
in tropical America. The genus is much in need of revision and
would be a good subject for work toward a master's degree.
Apparently, no generic problem is involved to complicate the study.
The treatment of the Guatemalan species presented here will
permit one to put useable names on plants of the genus but
certainly changes may be expected when a critical study is made.
Plants glabrous throughout C. hirsutum var. glabrum.
Plants copiously pubescent.
Flowers heads sessile or nearly so; pubescence of the stems appressed.
C. herbacewn.
Flowers heads conspicuously pedunculate.
Heads many-flowered; calyx lobes somewhat obtuse C. lanceolatum.
Heads few-flowered; calyx lobes acute to attenuate.
Leaves appressed-pilose with short hairs on the upper surface C. guianense.
Leaves hirsute with spreading hairs on the upper surface.
Leaf blades mostly reniform and rounded at the apex C. cordifolium.
Leaf blades broadly ovate, rounded at the base, usually acute or acutish at
the apex C. hirsutum.
Coccocypselum cordifolium Nees & Mart. Nov. Act. Nat.
Cur. 12: 14. 1824. Geophila pleuropoda Donn.-Sm. Bot. Gaz. 52: 50.
1911 (type collected between Secanquim and Sepacuite", Alta
Verapaz, G. P. Go//). Geocardia pleuropoda Standl. Contr. U. S.
Nat. Herb. 17: 445. 1914. Tontanea pleuropoda Standl. N. Am. Fl.
32: 148. 1921. Coccocypselum rothschuhii Loesner in Engler, Bot.
Jahrb. 60: 370. 1926. Figure 24.
Moist or wet, chiefly mixed, mountain forest, 1,400-2,000 m.;
Alta Verapaz; Baja Verapaz; Zacapa; Chiquimula; Huehuetenango.
Mexico (Veracruz) to Costa Rica; eastern and southern Brazil.
Plants creeping, the stems hirsute with long, spreading, whitish or purplish hairs;
stipules filiform, 1-3 mm. long; petioles 1.5 cm. long or less, hirsute; leaf blades mostly
reniform or orbicular, 1-3.5 cm. long and broad, cordate or truncate at the base,
usually broadly rounded and apiculate at the apex, hirsute or hispid above with
spreading hairs, often bright purple beneath and sparsely or densely long-hirsute;
inflorescence capitate, mostly 2-4-flowered, the peduncles 1-3.5 cm. long, the bracts
linear or foliaceous; calyx and hypanthium long-hirsute; calyx lobes linear or oblong-
42 FIELDIANA: BOTANY, VOLUME 24
linear, acute, 2-4 mm. long; corolla 12 mm. long, purplish white or sometimes blue,
hirsute; fruit densely hirsute, bright blue or sometimes greenish white.
Coccocypselum guianense (Aubl.) Schum. in Mart. Fl. Bras.
6, pt. 6: 315. 1889. Tontanea guinensis Aubl. PI. Guian. 108. 1775.
Guatemalan material seen only from Quich£ and Chiquimula.
Northern British Honduras; southern Mexico to Brazil; Jamaica
and Cuba.
Stems often much elongate, pilose or hirsute with long or short, chiefly spreading
hairs; stipules filiform-subulate, 2-5 mm. long; petioles 3 cm. long or less, the leaf
blades oval to rounded-ovate, 2.5-7 cm. long, 1.5-4.5 cm. broad, rounded to acutish at
the apex, truncate to obtuse at the base and often short-decurrent, densely pilose on
the upper surface with long and short, mostly appressed hairs, often purplish beneath,
densely soft-pilose; inflorescence capitate, mostly 2-4-flowered, the peduncles 1-4 cm.
long, the bracts linear, 3-7 mm. long; calyx and hypanthium densely pilose, the calyx
lobes linear, 3-4 mm. long; corolla blue or lavender, 7-10 mm. long, copiously pilose,
the lobes half as long as the tube; fruit blue, 6-10 mm. long or larger, pilose.
There appears to be relatively little difference, and these
differences small, between the plants which have been called C.
guianense and C. herbaceum in our area. The quickest way to
distinguish them is by the pedunculate inflorescence in C.
guianense and the sessile inflorescence on C. herbaceum. There are,
however, mixed collections which should be looked into for they
may indicate that the two species are really not very distinct.
Coccocypselum herbaceum Lam. Encycl. 2: 56. 1786. C.
repens Swartz, Prodr. Veg. Ind. Occ. 31. 1788. Tontanea herbacea
Standl. N. Am. Fl. 32: 147. 1921.
Wet forests and Manicaria swamps, Izabal, at or near sea-level.
British Honduras to Colombia and probably extending farther
southward; Greater Antilles.
Procumbent or creeping, the young stems densely pilose with short or long,
appressed or ascending hairs; stipules linear-subulate, 2-6 mm. long; petioles slender,
2.5 cm. long or less, the blades broadly ovate to oblong, 2-5 cm. long, 1-3 cm. broad,
obtuse or subacute, rounded or obtuse at the base, sparsely or densely pilose above
with short appressed hairs, short-pilose beneath, often densely so, with subappressed
hairs; inflorescence capitate, all or mostly sessile, commonly 2-3-flowered, the bracts
linear-subulate, 3-4 mm. long; calyx and hypanthium densely appressed pilose, the
calyx lobes linear-subulate, 2-3 mm. long; corolla blue, 5-7 mm. long, short-pilose;
fruit subglobose, 5-6 mm. in diameter, bright blue, short-pilose.
See comment under C. guianense.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 43
Coccocypselum hirsutum Bartling ex DC. Prodr. 4: 396. 1830.
Tontanea hirsute Standl. N. Am. Fl. 32: 147. 1921. Pixcac (Alta
Verapaz, probably Quecchi).
Most often in moist or wet, pine forest, 1,200-1,700 m.,
sometimes in mixed forest and occasionally descending to low
elevations; Pete"n; common in the mountains of Alta Verapaz; El
Progreso; Chiquimula; Solola; Suchitepe*quez; Quiche"; Huehue-
tenango. Southern and western Mexico; British Honduras
to El Salvador and Panama; southward to Trinidad and Bolivia.
Stems procumbent or creeping, densely hirsute or hispid; stipules subulate, 3-6
mm. long; petioles 2 cm. long or less, the leaf blades oval to oblong-ovate, 2-5.5 cm.
long, 0.5-2 cm. broad, rounded to acutish at the apex, rounded at the base, hirsute or
hispid above with long spreading yellowish hairs, hirsute beneath; inflorescence
capitate, the heads 3-5-flowered, on peduncles 1-3 cm. long, the linear bracts 3-4 mm.
long; calyx and hypanthium densely hirsute, the calyx lobes linear, acute, 2.5-5 mm.
long; corolla lavender or pale blue, 12-14 mm. long, hirsute; fruit hirsute, 6-10 mm.
long, pale to bright blue.
Like most other species of the genus, this is a handsome and,
because of the abundant blue fruits, somewhat ornamental plant.
The leaves often are tinged with deep purple, at least beneath. A
decoction of the plant is said to be employed as a "remedy" for
snake bite in Alta Verapaz.
The type specimens are presumed to have been collected in
Mexico, probably by Haenke. We follow the senior author's
treatment of the species in North American Flora knowing that he
did not see the type but believe that he was correct in his
understanding of the plant. The plant is quite variable. The range
given here is an extensive one and there is a possibility that more
than one species, or perhaps several varieties, are involved.
Coccocypselum hirsutum var. glabrum (Bartling ex DC.) L.
Wms. Phytologia 25: 462. 1973. C. glabrum Bartling ex DC. Prodr.
4: 397. 1830. Tontanea glabra Standl. Journ. Wash. Acad. Sci. 15:
104. 1925.
A rare plant collected only in Pete"n and British Honduras.
Nicaragua; Panama.
Glabrous plants, but otherwise similar to the typical variety described above.
The junior author assumes that the Guatemalan collection
which was a mixed collection with the typical variety, is the same
as the type of the plant which was collected at low elevations in
Panama. The specimen from Nicaragua also was mixed with the
44 FIELDIANA: BOTANY, VOLUME 24
typical variety. It is perhaps only a minor genetic variant. C.
hirsutum and C. glabrum were described in the same publication, C.
glabrum is chosen to become the variety since Coccocypselums are
most often pubescent.
Coccocypselum lanceolatum (Ruiz & Pav6n) Pers. Syn. PI. 1:
132. 1805. Condalia lanceolata Ruiz & Pavon, Fl. Peruv. 1: 54. 1798.
C. canescens Willd. ex Cham. & Schlecht. Linnaea 4: 139. 1829.
Tontanea canescens Standl. N. Am. Fl. 32: 146. 1921. Manchupij
(Quecchi).
In Guatemala known only from the vicinity of Coban, at about
1,300 m., in wet pine or mixed forests. Guatemala, Costa Rica and
Panama to Bolivia and Brazil.
Plants rather stout, procumbent, the stems densely pilose- sericeous with
yellowish or whitish hairs when young; stipules linear-subulate, 3-10 mm. long;
petioles 2.5 cm. long or less, the blades oblong to ovate, 2-8 cm. long, 1-3.5 cm. broad,
acute or obtuse, subcordate to obtuse at the base, densely pilose above with short,
subappressed, mostly yellowish hairs, densely pilose beneath with longer, appressed or
somewhat spreading hairs; heads densely many-flowered, on peduncles 1-6 cm. long,
the bracts often large and foliaceous; calyx and hypanthium densely long-pilose, the
calyx lobes oblong or ovate, 2-3 mm. long, foliaceous, usually obtuse; corolla purplish
blue, about 5 mm. long, densely pilose; fruit densely pilose, 5-7 mm. long or more,
bright blue or sometimes violet.
COFFEA Linneaeus. Coffee
Shrubs or small trees, usually glabrous, the branchlets subterete; stipules rather
broad, persistent, acuminate; leaves opposite, membranaceous or subcoriaceous,
sessile or petiolate; flowers axillary, glomerate, sessile or short-pedicellate, white,
fragrant, the pedicels bracteolate, the bractlets often forming a cupule; hypanthium
subcylindric to turbinate, the calyx short, truncate, dentate, or lobulate, persistent,
often glandular within; corolla salverform or funnelform, the tube short or elongate,
glabrous or villous in the throat, the limb 5-8-lobate, the lobes oblong, obtuse,
spreading, contorted in bud; stamens usually 5, inserted in the throat of the corolla,
the filaments short or none; anthers dorsifixed near the base, linear, obtuse or acute,
included or exserted; ovary 2-celled, the style filiform or thickened, glabrous, the 2
branches linear or subulate; ovules solitary in the cells, affixed to the middle of the
septum; fruit baccate, globose or oval, dry or fleshy, containing 2 nutlets, these
coriaceous or chartaceous, convex dorsally, sulcate ventrally.
About 40 species, in tropical Asia and Africa, some of them now
grown in all tropical regions of the earth.
Flowers 5-parted C. arabica.
Flowers 6-8-parted C. liberica.
Coffea arabica L. Sp. PL 172. 1753. Cafe; coffee. Figure 45.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 45
Native of tropical Africa, now grown in most tropical regions of
the earth for its seeds; planted in all Guatemalan departments
except probably Totonicapan, rarely spontaneous in virgin forests.
A glabrous shrub or small tree as much as 5.5 m. tall with thin gray bark; wood
white, moderately hard and fine-grained; leaves short-petiolate, oval or elliptic, 7-20
cm. long, 3-7.5 cm. broad, acuminate, cuneate at the base, subcoriaceous, usually
persisting for three years, with 7-12 pairs of lateral nerves; flowers in clusters of 2-9 or
more, sessile or nearly so, 12-18 mm. long; bractlets ovate, the inner ones connate at
the base of the pedicel, shorter than the 5-denticulate calyx; corolla lobes equaling or
exceeding the tube; anthers exserted; fruit about 1 cm. long, at first green, then red,
finally blue-black.
Commercially coffee is the most important plant of Guatemala
and local prosperity is dependent primarily upon the coffee crop
and its market. When coffee production is good and the price is
high, Guatemala, like other Central American countries, is
prosperous. When its price in the world market is low, hard times
prevail. Foreign credit for the purchase of essential imports is
heavily dependent upon the money received from export of the
coffee crop. Coffee was introduced into Guatemala around the
middle of the 18th century, but its cultivation did not have more
than local importance until about 1875. Coffee and bananas now
make up the major portion of Guatemalan exports.
Guatemala has long been celebrated for the quality of its coffee
and it is the leading Central American producer of this product.
According to statistics of the Asociacion Nacional del Cafe* of
Guatemala, the crop in 1970-1971 amounted to 2,800,000
hundredweight or quintales de oro. All of the departments produce
coffee except Totonicapdn where the land is too high for the
production of coffee, as are the highlands of some of the other
departments. The leading departments in coffee production are
listed as: Escuintla, 6,256 metric tons; Santa Rosa, 12,604 metric
tons; Quezaltenango, 16,468 metric tons; Suchitepe"quez, 14,996
metric tons; San Marcos, 25,668 metric tons; Alta Verapaz, 6,302
metric tons. It is thus apparent that the bocacosta region of five
western departments produced much more than half of the crop,
while the Coban region, which is sometimes assumed to produce
most of the coffee of Guatemala actually produces only a small
percentage of it. The yield in some of the departments is, of course,
very small. The lowest producers are Pete"n, Izabal, and El
Progreso, all of whose land is too low for commercial cultivation.
Most Guatemalan coffee is grown on the lower or middle slopes of
the mountains, at 600 to 1,500 m., but some is planted as high as
46 FIELDIANA: BOTANY, VOLUME 24
1,800 m., near Chimaltenango. The coffee of Antiqua (1,500 m.) is
noted for its excellence. Guatemalan coffee is considered a delicacy
in many nations around the world; the resulting high price for the
coffee has encouraged an increasing cultivation of the crop to the
extent that it comprised (1972) approximately 46 per cent of
Guatemalan exports with a value of $92,000,000.
Almost all the coffee of Guatemala is grown shaded, at lower
elevations for protection from the sun, at high elevations for
protection from the cold. Several species of Inga usually are used
to provide shade. Often tall forest trees are left when the land is
cleared, and various kinds of fruit trees are planted in the cafetales.
The valley of Antigua as well as the coffee-growing regions of the
Chimaltenango uplands are unique in that the coffee shade consists
of Grevillea trees, which are said to be the best protection against
cold winds and fogs. At these high elevations the harvest begins
January 1 or even later, when all the lowland coffee has long been
gathered. In the Pacific bocacosta, especially at lower elevations, as
well as in Alta Verapaz, bananas and plantains are much used for
shade, with the production of two saleable crops on the same land.
Some of the most unusual plantations are found in the higher parts
of Quezaltenango, between San Martin Chile Verde and Colomba,
where, at about 1,500 m., the cafetales are without shade and the
soil consists of the loose white sand characteristic of this region.
On the Pacific slope, as well as in Alta Verapaz, much coffee is
planted on the exceedingly steep slopes of quebradas and
barrancos, to which it is difficult even to climb on foot. The lower
and more level land of these barrancos usually is devoted to maize,
sugar cane, and other crops. At lower elevations the coffee harvest
begins soon after the rainy season, but at high elevations the coffee
ripens much later. Therefore, taking the country as a whole, the
some 129,000 metric tons of pure coffee that was produced in 1970-
1971 was harvested throughout much of the dry season.
Traveling from one part of Guatemala to another, it is possible
in almost any season to find ripe berries on the bushes. Flowers are
another matter, and are seldom seen, unless one is in the proper
locality on just the right day. A cafetal in flower is one of the most
beautiful sights imaginable, accentuated by the delightful fragrance
pervading the air. All the bushes burst into bloom on the same day,
and in two or three days the flowers have disappeared. The date of
flowering is not constant for any locality, it depends largely on
rainfall. In Alta Verapaz, where there is constant moisture, the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 47
blooming extends over several months, and the harvest likewise is
prolonged, while in other regions the berries are gathered at one
time.
In all the markets of Guatemala coffee is offered for sale for
home consumption, and at Antigua, for instance, excellent coffee is
sold quite inexpensively. Apparently, no coffee that passes through
a beneficio is wasted, for the beans thrown out when coffee is
cleaned for export are all offered for sale in the markets for a very
low price. Cheap as it is, there are many Guatemalans who cannot
afford the beverage, but use in its place atol or other drinks
prepared from maize and other substances. Also, various seeds,
especially those of Cassias, are used for adulterating or substituting
for coffee. In times past coffee was often served in Guatemala in the
form of esencia - essence - which was obtained by boiling the
pulverized beans. The concentrated essence was then diluted by
adding hot water or milk to suit the taste of the user. Today
essence of coffee is rarely seen and coffee made in pots is the order
of the day. Instant coffees are becoming more popular and are often
served in hotels and restaurants. Where American tourists abound
coffee is made to American taste and often is no better than that
served in American hotels or cafes — or in homes, for that matter.
Some of the best coffee anywhere is served by the National Coffee
Association at the airport in Guatemala City.
Official "propaganda" for all countries from Mexico to Peru
indicates that the best coffee in the world comes from the country
being propagandized — and this may be true. The junior author,
having been Consul of Guatemala in Chicago for many years, is
quite sure that no coffee quite compares in flavor or aroma to that
of Guatemala!
The names coffee and cafe" are both derivities of the Arabic
word, kahweh, signifying wine. Coffee is a vegetable product of
relatively recent introduction into the civilized world. It is believed
that it reached Arabia from Africa during the fourteenth century,
and did not attain common use in Europe until around the middle
of the seventeenth century. It did not become a common crop in
Central America until after the middle of the nineteenth century.
Coffea liberica Bull, Retail List New, Beautif. and Rare PL
No. 97: 4. 1874. Cafe robusta; Liberian coffee.
48 FIELDIANA: BOTANY, VOLUME 24
Native of Liberia and adjacent regions of West Africa;
cultivated on a small scale in Guatemala.
A glabrous shrub or tree, sometimes 10 m. high but usually much lower, at least
in cultivation; leaves short-petiolate, coriaceous, lustrous, mostly elliptic-obovate, 12-
30 cm. long, 5-12 cm. broad, short-acuminate, acute or obtuse at the base, the lateral
nerves 8-12 pairs, domatiate in their axils; flowers several in a cluster, subsessile, 2.5
cm. long; bractlets connate, shorter than the subtruncate calyx; corolla lobes about
as long as the tube; anthers exserted; fruit 2-2.5 cm. long or even larger, yellowish
red, turning black.
This species is said to be planted in various regions of
Guatemala, but is little esteemed. We have noted but one
plantation of any size, in Retalhuleu near Chivolandia, but there
are said to be others in the Pacific bocacosta and in Alta Verapaz.
In its habit of growth Coffea liberica differs noticeably from C.
arabica. The leaves are twice as large and rather handsome.
Flowering is continued through much of the year and the berries
hang upon the bushes for a long time (in C. arabica they soon fall
if not picked). Liberian coffee is said to produce better at lower
elevations than C. arabica, and to be less susceptible to fungus
diseases. However, it never has become popular in American
countries.
It is worthy of note that in Guatemala Coffea liberica is known
among even the laborers as Cafe robusta, but it is not Coffea
robusta Linden of tropical Africa, which often is known as "robusta
coffee."
Coffea excelsa A. Chev. Rev. Cult. Col. 12: 258. 1903.
Shrubs said to be of this species were seen growing in the
grounds of the Direction de Agricultura in Guatemala years ago. It
is an African species that produces low-grade coffee, and is
cultivated in some regions of Africa.
Coffea corymbulosa Bertol. Fl. Guat. 410. 1840.
This was based upon material collected by Vel&squez at some
unspecified locality in Guatemala. Bertoloni states that "Coffea
arabica differs from this in its acuminate leaves and subsessile
flowers." It is suspected that the plant so named may be merely
Coffea arabica, but it may be a representative of some different
genus. This cannot be determined without examination of the type
specimen, in the Bertoloni herbarium in Italy.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 49
COSMIBUENA Ruiz & Pav6n
Erect or scandent, sometimes epiphytic shrubs or trees; leaves opposite, petiolate,
coriaceous or fleshy; stipules large, interpetiolar, soon deciduous; inflorescence
terminal, 1-flowered or a 3-flowered cyme (ours), sometimes paniculate; flowers large,
white; hypanthium oblong or turbinate; calyx tubular or campanulate, 5-6-dentate,
deciduous, the lobes sometimes unequal; corolla salverform or funnelform, the tube
elongate, slightly expanded at the throat, 5-6-lobate, the lobes spreading, contorted;
stamens as many as the corolla lobes, inserted below the throat; stamens included,
the anthers linear, basifixed, filaments short; ovary 2-celled; style elongate, clavate,
bifid; ovules many; capsule oblong or ovate, coriaceous, 2-celled, septicidally
bivalvate from the apex; seeds many, oblong, produced into a wing (often or always
bifid) at the ends.
A small genus of about a dozen species, mostly South
American, two others in southern Central America.
Cosmibuena matudae (Standl.) L. Wms. Fieldiana, Bot. 31:
45. 1965. Hillia matudae Stand!. Field Mus. Bot. 22: 51. 1940 (type
from Volcan de Tacana, Chiapas, Matuda 2327). Cosmibuena
holdridgei Monachino, Phytologia 3: 64. 1949 (type Holdridge s.n.
near Barillas, Huehuetenango). Figure 15.
Moist or wet mountain forests, 500-1,800 m.; Alta Verapaz;
Huehuetenango; Chiquimula; and doubtless in other western
departments. Mexico (Chiapas); Honduras; Nicaragua.
A tree of 20 m. or less, also epiphytic; stipules broadly elliptic, 2 cm. long, very
obtuse, 12 mm. broad, caducous; petioles 1.5-2.5 cm. long, the blades broadly elliptic
or oval-elliptic, 6.5-10.5 cm. long, 4-8 cm. broad, obtuse, shortly cuneate-narrowed at
the base; flowers fragrant, solitary or in three's at the apex of the branch, on short
stout pedicels; hypanthium clavate, 7-8 mm. long, narrowed to the base; calyx 9 mm.
long or less, the segments lanceolate, subulate-acuminate, erect, rigid, unequal;
corolla white, coriaceous, the slender tube 9 cm. long, 4 mm. broad at the middle, 6
mm. broad in the throat, the 5 lobes spreading, oblong, obtuse or rounded at the
apex, 3-3.5 cm. long.
COUSSAREA Aublet
Shrubs or small trees, usually glabrous; stipules commonly ovate-deltoid,
muticous, apiculate, or truncate, never subulate-aristate or connate into a long
sheath; leaves opposite or rarely verticillate, short-petiolate or subsessile, more or less
coriaceous; inflorescence terminal, usually paniculate, but variable in form;
hypanthium ovoid or turbinate, the calyx cupular and truncate, sometimes 4-dentate
or rarely 4-lobate; corolla funnelform or salverform, the throat dilated, naked within,
the 4 lobes valvate in bud, oblong or sometimes elongate; anthers subsessile in the
corolla tube, dorsifixed near the base, included or exserted; ovary 1-celled, or rarely
2-celled but with a very thin septum; the style filiform, with 2 short branches; ovules
erect, inserted on a very short basal column; fruit drupaceous, by abortion commonly
1-seeded, coriaceous, ovoid or oval, longer than broad, small or rather large; seed
erect.
50 FIELDIANA: BOTANY, VOLUME 24
A large number of species are known from South America, one
from Mexico, and about a dozen others from Central America and
Panama. The genus Dukea Dwyer, from Panama, is hardly distinct.
Leaves sessile or nearly so; corolla minutely puberulent C. imitans.
Leaves slender-petiolate; corolla glabrous C. mediocris.
Coussarea imitans L. Wms. Phytologia 26: 488, fig. 1973.
Figure 54.
Wet mixed forests at about 150 m., Pete'n; Izabal (type,
Steyermark 39013)', British Honduras.
Small trees 8-12 m. tall and to 10 cm. in diameter, the branchlets glabrous, terete
or nearly so; leaves broadly elliptic or ovate-elliptic, short acuminate, subsessile,
cuneate and abruptly terminated at the base, glabrous, the blades 12-20 cm. long and
3.5-9 cm. broad, acuminate tip about 1 cm. long, lateral veins mostly 8-10 pairs,
prominulous, petiole 2-5 mm. long; inflorescences terminal, thyrsoid-paniculate, short
pedunculate and surpassed by the subtending leaves, to about 8 cm. long, sparsely to
densely puberulent above; flowers 25-28 mm. long when mature; calyx and
hypanthium densely pubescent, about 3 mm. long, the calyx subcampanulate, about
1.5 mm. long and 2 mm. broad, bidentate or obscurely 4-dentate; corolla white,
fragrant, densely and minutely puberulent outside, 25-28 mm. long, the tube narrow,
about 18 mm. long, the lobes linear-oblong, acute, 8-9 mm. long; anthers linear, about
8 mm. long, inserted below, filaments attached near the base of anthers, about 1 mm.
long; style bifid, shorter than or as long as the corolla tube; fruits indehiscent,
subbaccate, laterally compressed, obovate, 1.5-2 cm. long and 1.2-1.5 cm. broad, 1-
seeded, the calyx persistent.
Confused with Coussarea impetiolaris Donn.-Sm., a Costa
Rican species, also with nearly sessile leaves.
Coussarea mediocris Standl. & Steyerm. Field Mus. Bot. 23:
248. 1947.
«
Known only from the type, Huehuetenango, forested ravine
near Maxbal, about 17 miles north of Barillas, Sierra de los
Cuchumatanes, 1,500 m., Steyermark 48732.
A small tree of 6-7.5 m., the branches terete, glabrous or obscurely puberulent;
stipules 3 mm. high, very broad, intrapetiolar, connate, very shallowly bilobate at the
apex; leaves firm-membranaceous, somewhat lustrous, on slender petioles 1.5-2 cm.
long, elliptic-oblong or lance-oblong, 9-14.5 cm. long, 3.5-4.5 cm. broad, rather
abruptly long-acuminate, acute at the base, glabrous, the lateral nerves about 6 pairs,
arcuate-ascending, the veins prominulous and laxly reticulate; inflorescence terminal,
cymose-paniculate, on stout peduncles 1.5-3 cm. long, 3-4 cm. long and broad,
trichotomous at the base, the branches terete, glabrous, the flowers sessile, densely
aggregate at the ends of the branches; hypanthium glabrous, campanulate, almost 1.5
mm. high, obtuse at the base; calyx barely 0.5 mm. high, obscurely undulate-dentate;
corolla white, 2.5 mm. long, glabrous outside, the tube 2 mm. broad at the base, then
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 51
slightly narrowed, then slightly dilated upward, 3 mm. broad at the apex, the lobes
linear-oblong, 7-8 mm. long, obtuse; anthers subexserted, linear, acute, 4 mm. long.
The junior author thinks that it is quite possible that the plant
is a Psychotria.
COUTAREA Aublet
Shrubs or small trees with terete branchlets; stipules interpetiolar, short, acute;
leaves opposite, petiolate, herbaceous; flowers large, terminal and axillary, solitary or
in few-flowered cymes, pedicellate; hypanthium turbinate, the calyx 6-lobate, the
lobes narrow, deciduous, subequal; corolla funnelform-campanulate, conspicuously
asymmetric, the tube slightly curved and gibbous-ventricose, the throat glabrous, the
limb 6-lobate, the lobes contorted in bud, one of them exterior; stamens 6, inserted at
the base of the corolla, the filaments filiform, much contorted; anthers basifixed,
linear, exserted; ovary 2-celled, the style filiform, the stigma clavellate-subulate,
exserted; ovules numerous, horizontal, the placentae affixed to the septum; capsule
large, obcompressed, coriaceous or ligneous, 2-celled, loculicidally bivalvate; seeds
numerous, imbricate, broadly winged.
In North America only one species is known, but probably
several species occur in South America.
Coutarea hexandra ( Jacq.) Schum. in Mart. Fl. Bras. 6, pt. 6:
196. 1889. Portlandia hexandra Jacq. Enum. PL Carib. 16. 1760. C.
speciosa Aubl. PL Guian. 314, 1. 122. 1775. Quina. Figure 19.
Brushy plains and hillsides, 850 m. or lower; probably in Pete"n;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Solold; probably in all
the Pacific coast departments. Mexico (Chiapas); British Honduras
to Panama; southward to Peru and Argentina.
A shrub or small tree, usually almost glabrous (in Central America^, the
branchlets whitish-lenticellate; stipules deltoid, 2-4 mm. long, acute and mucronate;
petioles slender, about 1 cm. long or less, the blades ovate or oval, 5-12 cm. long, 2-6
cm. broad, cuspidate-acuminate to acute, rounded to acute at the base; flowers
mostly in 3-flowered, terminal and axillary cymes, sometimes solitary, short-
pedicellate; hypanthium 4-6 mm. long; calyx lobes 6, lance-subulate or linear-
subulate, 5-12 mm. long; corolla white or yellowish, tinged below with purple, the
tube gibbous, 1-2 cm. broad in the throat, the lobes ovate, acute or obtuse, about one-
third as long as the tube or shorter, the whole corolla about 6 cm. long; stamens
usually exceeding the corolla lobes, the yellow anthers 1.5-2 cm. long; capsule oblong-
obovoid or oval-obovoid, about 2 cm. broad, shallowly bisulcate, dark brown, whitish-
lenticellate; seeds oval or orbicular, 8-10 mm. long, brown.
The leaves and branches are intensely bitter. Chiefly on this
account, perhaps, the plant is used in Guatemala and other parts of
Central America as a domestic remedy for malaria. Known in El
Salvador also by the names "quinita," "quina blanca," and "zalas"
52 FIELDIANA: BOTANY, VOLUME 24
or "salas." The bark is employed extensively there as a remedy for
fevers and a decoction of it, called agua de quina, is employed in the
treatment of wounds and sores.
CRUSEA Chamisso & Schlechtendal
Reference: Anderson, William R., A monograph of the genus
Crusea, Mem. N. Y. Bot. Card. 22 (4): 1-128, illus. 1972.
Annual or perennial herbs, erect or procumbent, glabrous or variously pubescent,
the stems 4-angulate; stipules connate with the petioles to form a ciliate sheath,
persistent; leaves opposite, ovate to linear, often conspicuously nerved; flowers small
or medium-sized, usually pink or purple, capitate, the heads commonly subtended by
4 foliaceous bracts; hypanthium ovoid or obovoid, compressed; calyx lobes 4,
persistent, elongate-subulate, sometimes alternating with small teeth; corolla
funnelform, with a slender tube, the throat glabrous, the lobes of the limb 4,
spreading, oblong or lanceolate, valvate in bud; stamens 4, inserted in the mouth of
the corolla tube, the filaments elongate, filiform; anthers dorsifixed above the base,
linear-oblong, long-exserted; ovary 2-celled, the style capillary, the stigma simple or
of 2 short branches; ovules solitary in the cells, attached at the middle of the septum;
capsule didymous, 2-coccous, the cocci indehiscent, separating from the persistent
axis.
The group consists of 13 species, in Central America, Mexico,
and southwestern United States. The illustrations in Dr. Anderson's
monograph are very good, the descriptions are not easy to follow.
Stems glabrous or puberulent or very sparsely hirsute.
Corolla 2-2.5 cm. long; leaves ovate or ovate-lanceolate C. coccinea.
Corolla less than 1 cm. long.
Leaves linear to subulate; plants to 30 cm. tall C. diversifolia.
Leaves narrowly elliptic to ovate; plants mostly much more than 30 cm. tall.
C. setosa.
Stems densely hirsute, hispid, or spreading-pilose, at least on the peduncles.
Corolla tube scarcely exceeding the calyx lobes C. parviflora.
Corolla tube much longer than the calyx lobes.
Calyx lobes lanceolate, about 1.5 mm. long; bracts subtending the flower heads
ovate or lance-ovate, 1-3 cm. long; plants annual and usually erect.
C. longiflora.
Calyx lobes linear-subulate, elongate; bracts subtending the flower heads ovate
to narrowly lanceolate, mostly 3 cm. long or more; plants chiefly perennial
and often procumbent.
Bracts at the base of the flower heads chiefly ovate, conspicuously contracted
and petiolate at the base C. hispida.
Bracts at the base of the flower heads chiefly lanceolate or narrowly
lanceolate, scarcely petiolate C. calocephala.
Crusea calocephala DC. Prodr. 4: 567. 1830. C. elata
Brandegee, Univ. Cal. Publ. Bot. 6: 68. 1914. C. guatemalensis
Gandoger, Bull. Soc. Bot. France 65: 34. 1918 (type, Tuerckheim
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 53
from Alta Verapaz). Verbena silvestre; heliotropio silvestre; hierba
de pato (fide Aguilar); chuchiim (Coban, Quecchi). Figure 61.
Common and widely distributed over much of Guatemala, at
2,500 m. or lower, chiefly in pine-oak forest, often in thickets or on
roadside banks, or even a weed in cornfields; Peten; Alta Verapaz;
El Progreso; Izabal; Chiquimula; Jutiapa; Santa Rosa; Escuintla;
Guatemala; Sacatepequez; Chimaltenango; Solola; Suchitepequez;
Quiche; Huehuetenango; San Marcos. Mexico; British Honduras;
El Salvador; Honduras.
Plants annual or apparently often perennial, copiously branched, erect or more
often procumbent and forming large mats of stems, these densely hirsute with long
spreading whitish hairs; leaves lanceolate or oblong-lanceolate, mostly 3-6 cm. long,
acute, conspicuously nerved, with very oblique nerves, hirsute on both surfaces;
inflorescence long-pedunculate, densely many-flowered, capitate, in fruit 2 cm. broad
or more, subtended at the base by usually 4 large bracts similar to the leaves; calyx
densely long-hirsute, the lobes subulate from a short triangular base; corolla purple
or rose-purple, 12-15 mm. long, the very slender tube minutely puberulent; cocci
smooth, brown, about 2 mm. long, glabrous.
The plant is a showy one and rather handsome until the
flowers and foliage begin to wither. In Guatemala it is far more
abundant than any other representative of the genus. The
nomenclature of the species is somewhat uncertain. It was based
upon one of the Sesse and Mocino drawings of Mexican plants
which agrees none too well with the material generally referred to
C. calocephala.
Crusea coccinea DC. Prodr. 4: 567. 1830. C. coccinea var.
breviloba Loes. Bot. Verh. Brandenb. 65: 115. 1923 (type collected
near San Martin, Jacaltenango, Huehuetenango, Seler 3113).
Sanalotodo (Huehuetenango).
Rocky slopes or in pine or mixed forest in the mountains, 1,000-
2,000 m.; Zacapa; Chiquimula; Suchitepe'quez; Solola;
Huehuetenango; Quezaltenango. Southern Mexico to Panama.
Plants branched, perennial, procumbent, glabrous throughout or nearly so, the
stems obtusely tetragonous; setae of the stipules filiform, elongate, glandular-
thickened at the apex; leaves evidently petiolate, mostly ovate and 3-5 cm. long,
acuminate or long-acuminate, scarcely paler beneath, conspicuously nerved, with
very oblique nerves; flowers heads long-pedunculate, dense but often with only few
flowers, subtended by 2 or 4 large bracts similar to the leaves; calyx lobes triangular-
lanceolate, ciliate; corolla lilac, about 2.5 cm. long, the throat dilated and much
broader than in other species, the broad lobes hispidulous outside; cocci smooth,
glabrous.
54 FIELDIANA: BOTANY, VOLUME 24
In Huehuetenango a decoction of the plant is applied
externally in treatment of cuts and skin afflictions.
Var. breviloba is described as having the flowers somewhat
smaller than in the typical form, with shorter corolla lobes, the
tube somewhat puberulent outside; it is known from Suchitepe"quez,
Solold, and Huehuetenango.
Crusea diversifolia (HBK.) W. R. Anderson, Mem. N. Y. Bot.
Gard. 22(4): 58. 1972. Spermacoce diversifolia HBK. Nov. Gen. &
Sp. 3: 341. 1819. Borreria subulata DC. Prodr. 4: 543. 1830.
Spermacoce subulata Hemsl. Biol. Cent. Am. Bot. 2: 60. 1881.
Diodia subulata Standl. Field Mus. Bot. 8: 386. 1931.
Open slopes in pine or oak forests in the western and central
highlands, 1,400-2,500 m.; Guatemala; Huehuetenango. New Mexico
and Arizona; widely distributed in the Mexican highlands.
Simple or diffusely branched annuals to 30 cm. tall; stems 4-angulate, glabrous
or sparsely pilosulose; leaves linear or linear-elliptic to subulate, hispidulous, acute,
mostly 7-20 mm. long and 1-3 mm. broad; stipular sheath 1-3 mm. long bearing
mostly 3-5 setae; inflorescences of terminal or lateral bracteate short-pedunculate
heads with 3-many flowers; hypanthium glabrous to scabrous, 1-1.5 mm. long; calyx
1.5-4 mm. long, the tube short, to 0.8 mm. long, the triangular to subulate lobes to 3.5
mm. long; corolla mostly 3-6 mm. long, the tube to about 4 mm. long, the triangulai
lobes to about 2 mm. long; seeds to about 2 mm. long, setulose.
The species resembles a Diodia to which it has been referred in
recent years.
Crusea hispida (Mill.) Robinson, Proc. Am. Acad. 45: 409.
1910. Crucianella hispida Mill. Gard. Diet. ed. 8. No. 4. 1768.
Spermacoce rubra Jacq. Hort. Schoenbr. 3: 3, t 256. 1798. Crusea
rubra Schlecht. & Cham. Linnaea 5: 165. 1830.
Brushy slopes or plains, 300-870 m.; Chiquimula (above El
Rincon); Escuintla; Suchitepe"quez (near Santo Domingo); Huehue-
tenango. Southern Mexico; El Salvador.
Plants annual, erect, the stems obtusely tetragonous, hirsute or hispid with
whitish, usually somewhat recurved hairs, simple or branched; setae of the stipules
slender and elongate, hispid; leaves on short or often elongate petioles, chiefly ovate
or lance-ovate, hirsute, 3.5-5(-8) cm. long, subobtuse to acuminate, thin, almost
concolorous, inconspicuously nerved; inflorescence capitate, terminal and axillary, on
long or short peduncles, rather laxly few-flowered, subtended at the base by usually 2
large bracts similar to the leaves and by numerous smaller but similar foliaceous
bracts; calyx lobes linear, green, elongate, coarsely hispid, 2-6 mm. long; corolla rose-
purple, about 6 mm. long, the very slender tube minutely puberulent, the anthers
long-exserted; cocci glabrous, smooth.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 55
Crusea longiflora (Willd. ex Roem. & Schult.) W. R.
Anderson, Mem. N. Y. Bot. Card. 22(4): 89. 1972. Spermacoce
longiflora Willd. ex R. & S. Syst. Veg. 3: 531. 1818. Crusea
brachyphylla Cham. & Schlecht. Linnaea 5: 165. 1830.
Of scattered occurrence, on grassy slopes or in pine and oak
forest, 500-2,500 m.; Chiquimula; Jalapa; Santa Rosa; Guatemala;
Chimaltenango; Huehuetenango. Mexico; Honduras and Costa
Rica.
Plants annual, erect, usually branched, generally 30 cm. high or less, the stems
terete, pilose or hirsute with rather soft, white hairs; leaves chiefly ovate or oblong-
ovate, and 1.5-5 cm. long, acute, hirsute on both surfaces, much paler beneath;
inflorescence capitate, the heads very dense and many-flowered, in fruit about 1.5 cm.
broad, the subtending bracts similar to the leaves and of about the same size, pale at
the base; calyx lobes smaller and relatively broader than in other species, 1-3.5 mm.
long, very densely covered with long soft white hairs; corolla white or purplish white,
the slender tube densely and minutely pilosulous, 6-11 mm. long; cocci minutely
papillate-roughened.
Crusea parviflora Hook. & Arn. Bot. BeecL Voy. 430, t. 99.
1840. Hierba de pajaro (fide Aguilar).
Dry or moist thickets or on rocky banks, 1,500 m. or lower;
Santa Rosa; Escuintla; Guatemala; Sacatepequez. Mexico; ranging
southward to Costa Rica.
Annual, erect or spreading, 80 cm. high or less, often much branched, the stems
obtusely tetragonous, short-pilose below or finally glabrate, more densely pilose
above with longer hairs; setae of the stipules short, appressed-pilose; leaves thin,
inconspicuously nerved, mostly ovate and 2-4 cm. long, acute, densely pilose, short-
petiolate; inflorescence capitate, the heads in fruit about 12 mm. broad, densely
many-flowered, subtended by 4 small ovate bracts similar to the leaves; calyx lobes
linear-spatulate, acute, long-ciliate, 1-2.5 mm. long; corolla white, pubescent, scarcely
exceeding the calyx lobes, pubescent; cocci subglobose, glabrous, punctate.
Crusea setosa (Mart. & Gal.) Standl. & Steyerm. Field Mus.
Bot. 23: 22. 1943. Borreria setosa Mart. & Gal. Bull. Acad. Brux. 11,
pt. 1: 131. 1844. Spermacoce setosa Hemsl. Biol. Cent.-Am. Bot. 2:
60. 1881. C. cruciata Wats. Proc. Am. Acad. 25: 152. 1890. B.
chiapensis Brandegee, Univ. Cal. Publ. Bot. 10: 417. 1924. Estrella.
Open, often gravelly or rocky slopes, sometimes in oak forest,
1,800 m. or lower; Alta Verapaz; Jalapa; Jutiapa; Guatemala;
Huehuetenango. Mexico.
A slender or stout, erect annual, the stems glabrous or nearly so, obtusely
tetragonous, the sheaths hispidulous; leaves linear-lanceolate, attenuate, often
somewhat revolute, green and scabrous above, hispidulous or scabrous beneath on the
56 FIELDIANA: BOTANY, VOLUME 24
very conspicuous nerves, paler beneath; inflorescence capitate, the heads dense and
many-flowered, long-pedunculate, subtended by 4 long bracts similar to the leaves;
calyx densely white-hirsute, the green lobes lance-subulate, short; corolla about 5
mm. long, lilac, glabrous; cocci 2 mm. long or more, minutely tuberculate.
DECLIEUXIA Humboldt, Bonpland & Kunth
Herbs or low shrubs, branched, glabrous or scaberulous, the branches terete or
angulate; stipules reduced to a line bearing 1-3 rigid setae; leaves opposite or
verticillate, sessile or short-petiolate, coriaceous; flowers small, in terminal cymes
having more or less scorpioid branches; hypanthium orbicular or obovoid,
compressed; calyx lobes 2 or 4, equal or unequal, persistent; corolla funnelform,
pilose or villous in the throat, the 4 lobes short, spreading or reflexed, valvate in bud;
stamens 4, inserted in the corolla throat, the filaments filiform, the anthers dorsifixed,
versatile, oblong, obtuse, partly or wholly exserted; ovary 2-celled, the style filiform,
with 2 short branches; ovules solitary, erect; fruit dry, laterally compressed,
didymous, the lobes orbicular, indehiscent.
A genus of tropical America with numerous species in South
America, mostly in Brazil, a single one occurring in Mexico and
Central America.
Declieuxia fruticosa (Willd.) Kuntze, var. mexicana (DC.)
Standl. Field Mus. Bot. 12: 378. 1936. Declieuxia mexicana DC.
Prodr. 4: 479. 1830. Figure 51.
Moist open pine forest, 1,000-2,000 m.; Zacapa; Huehuetenango;
reported by Hemsley as collected between Tocoy and San Jer6nimo
(Baja Verapaz?), Bernoulli 1011. Southern Mexico; savannas and
grassy pine ridges of northern British Honduras and doubtless
extending into Pet4n; ranging southward to Panama and widely
distributed in South America (the species; not the variety).
Uncommon in Guatemala.
Plants herbaceous or suffrutescent from a hard woody root, usually with several
simple or branched stems 30 cm. high or less, glabrous throughout or nearly so, the
stems somewhat angulate; leaves opposite, sessile or almost so, lance-oblong to
linear-oblong, mostly 2-3 cm. long, acute or subobtuse, narrowed to the subacute
base, lustrous above; cymes small and few-flowered, pedunculate, with conspicuous
but small, linear, persistent, spreading bracts; corolla white, 5-6 mm. long, lustrous.
The plant elsewhere in Central America grows principally in
grassy savannas. The form of Mexico and Central America differs
but little if at all from the typical one of Colombia and Venezuela.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 57
DEPPEA Schlechtendal & Chamisso
Slender shrubs, more or less pubescent; stipules small, interpetiolar, deciduous;
leaves opposite, petiolate, membranaceous, often anisophyllous; flowers small, yellow,
arranged in axillary or terminal cymes or umbels, sometimes solitary in the leaf axils,
pedicellate; hypanthium turbinate or hemispheric; calyx 4-lobate, the lobes equal or
unequal, persistent; corolla rotate or short-funnelform, the throat glabrous, the limb
4-lobate, the lobes spreading in anthesis, contorted in bud; stamens 4, inserted in the
corolla throat, the filaments short, linear, compressed; anthers exserted, dorsifixed,
oblong or linear; ovary 2-celled, the style slender, the stigma capitate or bilobate;
ovules numerous, imbricate, the placentae peltately affixed to the septum; capsule
small, turbinate or obovoid, coriaceous or chartaceous, costate, 2-celled, loculicidal
from the apex, the valves cleft; seeds numerous, minute, subglobose, obtusely
angulate, the testa granulate.
About 20 species are known, in the forests of Mexico and
Central America.
Capsule oval or ovoid, rounded at the base, very inconspicuously costate.
Calyx lobes very unequal, the larger ones lanceolate or oblong, foliaceous, and 3-8
mm. long D. inaequalis.
Calyx lobes subequal, all very small and less than 2 mm. long.
Leaves rounded or very obtuse at the apex, mostly 6-8 cm. broad or broader;
calyx lobes linear-subulate, 2 mm. long D. amaranthina.
Leaves acute or acuminate, mostly 2-3 cm. broad; calyx lobes minute, deltoid.
D. pubescens.
Capsule turbinate, acute at the base, very conspicuously costate.
Corolla 6-9 mm. long; flowers in small or large cymes D. grandiflora.
Corollas 2.5-4 mm. long; flowers in secund racemes.
Young branches and inflorescences pilosulose; corollas 2.5-3 mm. long; underleaf
surface not lineolate D. anisophylla.
Young branches and inflorescences glabrous; corolla 3-3.5 mm. long; underleaf
surface lineolate D flava.
Deppea amaranthina Standl. & Steyerm. Field Mus. Bot. 22:
281. 1940.
Known only from the fruiting type, rocky wooded slopes along
river, Chiquimula, Cerro Brujo, near village of Brujo, 1,500-2,000 m.,
Steyermark 30926.
A branched shrub, the young branchlets hirtellous-villosulous; leaves large, the
petioles 1-3.5 cm. long, the blades rounded-ovate or rhombic-ovate, sometimes almost
orbicular, 7-11 cm. long, 4-8 cm. broad, rounded or obtuse at the apex, rounded to
broadly cuneate at the base and abruptly decurrent, sparsely and minutely
villosulous above, slightly paler beneath and more densely villosulous; inflorescence
cymose-paniculate, the panicles lax and few-flowered, 4 cm. long or less, usually
shorter than the petioles, the slender pedicels 3-6 mm. long; sepals reflexed in fruit,
linear-subulate, 2 mm. long; capsule oval, 5 mm. long, 3 mm. broad, obtuse at the
base, obscurely costate, sparsely puberulent or almost glabrous.
58 FIELDIANA: BOTANY, VOLUME 24
Deppea anisophylla L. Wms., Phytologia 26: 490. 1973.
In mountain forest, 1,000-1,300 m., Suchitep<§quez (type from
Finca Moca, Skutch 2072).
Slender shrubs to 1.5 m. tall, the young branches short pilose pubescent,
becoming glabrous with age; leaves prominently anisophyllous, the large leaf of a pair
2-4 times larger than the opposite leaf, elliptic to lanceolate, acuminate, the blade
(large) 6-10 cm. long, 1.5-3 cm. broad, sparsely short pilose especially along the veins
below, the petiole up to about 2 cm. long, sparsely short pilose; inflorescences few-
several-flowered secund racemes, mostly about 1.5-3 cm. long, the peduncle extremely
slender, puberulent; hypanthium about 1 mm. long, subglabrous; the calyx divided to
the base, the lobes somewhat unequal, the longest lobe about 1 mm. long, linear,
ciliate or not; corolla rotate, divided almost to the base, about 2.5-3 mm. long, the 4
lobes lanceolate, acute, sparsely pilose dorsally, about 2.2-2.5 mm. long and to 1 mm.
broad; anthers linear-oblong, obtuse, about 1.5 mm. long, the filaments very short;
style about 2.5 mm. long, stigmas globose; capsules narrowly obovate, strongly
reticulate, 2.5-3.5 mm. long.
The smallest flowered of the Guatemalan species and the only
one with pubescence in the inflorescence and on young stems. The
very unequal leaves in each pair are distinctive.
Deppea flava (Brandegee) L. Wms. comb. nov. Plocaniophyl-
lon flavum Brandegee, Univ. Cal. Publ. Bot. 6: 69. 1914; Standley,
N. Am. Fl. 32: 139. 1921. Figure 7.
In dense mixed forests or wet thickets, 750-1,500 m.; Quezal-
tenango; San Marcos. Mexico (type from Finca Mexiquito, Chiapas,
Purpus 7019).
A slender shrub 1-3 m. tall, the branchlets slender, glabrous; leaves
anisophyllous, one of a pair often twice as large as the other, elliptic or lanceolate,
acuminate, the larger ones 6-12 cm. long and 1.5-3.5 cm. broad, tertiary veins on
lower surface prominent and lineolate, petioles slender, up to 1 cm. long;
inflorescences axillary, few-several-flowered secured racemes, often almost equaling
the leaves, the peduncles filiform, 3-5 cm. long; calyx and hypanthium about 2 mm.
long at anthesis, the lobes subequal or somewhat unequal, about 0.5 mm. long,
triangular-lanceolate; corolla yellowish, about 3-3.5 mm. long, the tube very short,
the lobes oblong, acute; capsule cylindric, acute at the base, 5-8 mm. long.
The genus Plocaniophyllum seems to differ in no substantial
way from Deppea. The senior author, in North American Flora (32:
139. 1921), placed the plant in the tribe Mussaendeae because of the
lineolate underleaf surface, the fruits being unknown. It is clear
now that the plant belongs in the tribe Rondeletieae and that the
lineolate condition is not sufficient for generic distinction.
Deppea grandiflora Schlecht. Linnaea 19: 748. 1847. D.
floribunda Hemsl. Diag. PI. Mex. 31. 1879. Figure 8.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 59
Damp mixed forest or on brushy slopes, 1,200-3,000 m.; El
Progreso; Santa Rosa; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico to Panama.
A shrub 1-2 m. high, rarely a tree of 6 m., the branches brownish, puberulent
when young; petioles 1-4 cm. long, the blades oblanceolate to ovate or ovate-oblong,
5-15 cm. long, 1.5-6.5 cm. broad, abruptly acuminate to attenuate at the apex, acute
to attenuate at the base, glabrous or sparsely puberulent above, puberulent or
villosulous beneath, especially along the veins; inflorescence cymose-corymbose,
usually many-flowered, long-pedunculate, the puberulent pedicels 1-6 mm. long;
hypanthium broadly turbinate, glabrous or obscurely puberulent; calyx lobes minute,
deltoid, acute; corolla glabrous, lemon-yellow, 6-10 mm. long, obtuse or acute in bud,
the tube very short, the lobes oblong or ovate, obtuse; anthers 2.5-3 mm. long;
capsule turbinate, 3-5 mm. long, prominently 8-costate, coriaceous, dark brown; seeds
minute, dark brown.
The species as treated here is in the broad sense and as it is in
the herbarium there is certainly more than one species involved. It
is quite probable that D. costaricensis Polak. is distinct and
probably extends northward to Honduras.
Deppea inaequalis Standl. & Steyerm. Field Mus. Bot. 22:
384. 1940.
Type from San Marcos, above Finca El Porvenir, along Rio
Cabus to within two miles of Cueva de las Palomas, southern slopes
of Volcan de Tajumulco, 1,300-1,500 m., Steyermark 37974; also no.
37947. Mexico.
A branched shrub 1-1.5 m. high, the young branchlets minutely puberulent;
petioles short, 12 mm. long or much shorter, the blades ovate-lanceolate to narrowly
lanceolate or elliptic-lanceolate, 5-12 cm. long, 2-4 cm. broad, long-acuminate, toward
the base subabruptly contracted and narrowly long-attenuate, sparsely short-pilose
above, sparsely and minutely puberulent beneath or almost glabrous; flowers cymose,
the cymes lax, few-flowered, sometimes umbelliform, the peduncles 5 cm. long or
shorter, usually 1-2 cm. long, the pedicels up to 8 mm. long, minutely and sparsely
puberulent; hypanthium broadly obovoid, 1 mm. long, minutely puberulent or almost
glabrous; calyx lobes very unequal, green, one of them much smaller and only 1-2
mm. long, the other 3 mostly 4-7 mm. long, linear or lanceolate, almost glabrous;
corolla acute in bud, sparsely and minutely puberulent, about 4 mm. long in anthesis;
anthers oblong, 2.3 mm. long; immature capsule oval, 3 mm. long.
Deppea pubescens Hemsl. Diag. PL Mex. 31. 1879.
Chiquimula (forested slopes, Volcan de Ipala, 1,510 m.);
Quezaltenango (between Santa Maria de Jesus and Calahuache,
1,200-1,300 m.). Southern Mexico.
A slender shrub 1-1.5 m. tall, the branchlets densely villosulous when young,
brown; petioles 0.5-2 cm. long, the blades ovate or lance-oblong, 3.5-7.5 cm. long, 1.2-3
60 FIELDIANA: BOTANY, VOLUME 24
cm. broad, acute or acuminate, acute at the base, puberulent above, densely
villosulous or tomentose beneath; inflorescence cymose-corymbose, few-flowered,
sometimes equaling the leaves but usually much shorter, slender-pedunculate, the
pedicels 2-5 mm. long; hypanthium 0.8 mm. long, puberulent; calyx lobes ovate-
deltoid, obtuse; corolla 2-3 mm. long, almost rotate, glabrous; capsule oval, about 3
mm. long, puberulent, very obscurely costate.
Another species may be involved. The Guatemalan material is
not good.
DIDYMAEA Hooker f.
Perennial herbs, procumbent or scandent, in habit similar to Galium, usually
much branched, glabrous to densely pubescent (rarely), with brittle stems; leaves
opposite, petiolate, herbaceous; stipules interpetiolar, geminate or a single bifid
stipule on each side, deciduous or persistent and finally recurved; flowers perfect,
axillary, minute, reported as yellowish, greenish brown, or purple, pedicellate, the
pedicel not articulate, elongating in fruit and often recurved; hypanthium turbinate-
globose, the calyx entire; corolla campanulate or rotate, glabrous, 4-lobate, the lobes
triangular, subacute, valvate in bud; stamens 4, inserted between the corolla lobes,
the short filaments subulate, the anthers dorsifixed, oblong; ovary 2-celled, the style
short, thick, with two short divaricate branches; ovules solitary, affixed to the
septum below the middle; fruits didymous, black or dark blue, lustrous, the lobes
globose, somewhat fleshy, one lobe often smaller than the other and imperfect.
Didymaea, which occurs from central Mexico southward to
Panama, was considered to consist of only two species [D.
alsinoides (Schlecht. & Cham.) Standl., divided into three varieties,
and D. linearis Standl.]. This, however, seems to be far from the
actual state of affairs. The genus is found quite commonly in
Central America and, although fruiting materials are abundant,
those with usable flowers are not. The group may be complicated
by cleistogamy and peihaps even apogamy.
Since it seems reasonably certain that D. alsinoides does not
occur in Guatemala or Central America, but that there are three
distinctive species in Guatemala (plus one which seems to be
vegetatively distinctive but lacks usable flowers or fruits), a
necessary combination and two new species are described below.
This is a departure from the usual practice of this flora but is
neccesitated by publishing deadlines.
Leaves broadly ovate to suborbicular, base obtuse or rounded D. austratis.
Leaves linear-lanceolate to lanceolate, rarely ovate, bases acute or cuneate to obtuse.
Leaves without obvious lateral nerves D. microphylla.
Leaves with obvious lateral nerves, either penni- or plinerved.
Leaves 3-(5-) plinerved, sparsely hispidulous D. hispidula.
Leaves penninerved, leaves and stems glabrous or obscurely puberulent
D. microflosculosa.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 61
Didymaea australis (Standl.) L. Wms., comb. nov. Didymaea
alsinoides var. australis Standl. Field Mus. Bot. 18: 1291. 1938.
Figure 68.
Wet mountain forests, 1,800-2,400 m.; San Marcos (Williams,
Molina & Williams 25765). Costa Rica (type, Brade & Brade 2143);
Panama.
Procumbent or scandent herbs, sparsely pilosulose or glabrous; leaves broadly
ovate to suborbicular, acute or subacute, the base rounded or subtruncate or
subdecurrent on the petiole, triplinerved, glabrous or nearly so, the blade 10-18 mm.
long and 8-12 mm. broad; inflorescences axillary, 1-flowered; pedicel slender, to 10
mm. long; fruit didymous, fleshy, black.
A single Guatemalan collection seems referable to this broad-
leaved species from the southern part of the range of the genus.
Didymaea hispidula L. Wms. sp. nov. Figure 68.
Thickets and mixed coniferous-oak forests, 1,600-3,000 m.;
Jalapa; Jutiapa; Chiquimula; Chimaltenango; Sacatepequez;
Huehuetenango; Quezaltenango; San Marcos (type, Williams,
Molina & Williams 25896).
Planta herbacea usque ad 1 m. longa; caulibus graciles 4-angulati hispiduli; folia
lanceolata acuminata breviter petiolata hispidula 3-(5-) plinervia; stipulae
interpetiolatis lobis linearibus; inflorescentiae axillares uniflores, pedicelli graciles;
flores desideratur; fructus didymus lucens nigrescens, lobis globosis carnosulis.
Small herbaceous vines perhaps to 1 m. or more long; stems slender, brittle, 4-
angulate, hispidulous becoming glabrous, internodes 4-8 cm. long; leaves lanceolate,
acuminate, short petiolate, hispidulous especially along the margins, 3-(5-) plinerved
the blades mostly 1-3(4) cm. long and 0.4-1(1.5) cm. broad, the petioles mostly 0.5
cm. long, rarely to 1.5 cm. long; stipules interpetiolar, about 1 mm. long, divided to
the base and the lobes linear, hispidulous; inflorescence axillary, 1-flowered, pedicels
slender, mostly 5-10 mm. long; mature flowers not known, the buds minute and
possibly cleistogamous (or apogamous?); fruits didymous, fleshy, black and shining, 5-
8 mm. long and about 10 mm. broad across the lobes.
Common in Guatemala and distinguished from the other
species by the plinerved hispidulous leaves, and the deeply divided
stipules. It is a species of the coniferous and oak forests at middle
and high elevations. The holotype is from "forested mountain slopes
and ravines, Sierra Madre Mountains about 6 km. (airline) north of
San Marcos, dept. San Marcos, alt. 2,700 m., Dec. 13, 1963,
Williams, Molina & Williams 25896 (F).
Didymaea microflosculosa L. Wms. sp. nov. Figure 68.
Wet forests or cloud forests, 1,500-2,000 m., Guatemala;
Escuintla; Chimaltenango; Solola; Huehuetenango; Quezaltenango;
62 FIELDIANA: BOTANY, VOLUME 24
San Marcos [type: Williams, Molina & Williams 25944 (F)].
Honduras.
Herbae parvae scandentes usque ad 1 m.; caules graciles 4-angulati glabri vel
obscure puberulenti; folia late vel anguste lanceolata acuminata penninervia lucentia
obscure puberulentia; inflorescentiae axi Hares uniflores; pedicelli graciles; flores
vinacei; corolla rotata 4-lobata, lobis ellipticis acutis trinervis; fructus didymus
nigrescens carnosus.
Herbaceous or scandent vines to perhaps a meter long; stems slender, 4-angulate,
glabrous or at first obscurely puberulent, the internodes to 10 cm. long or more;
leaves broadly to narrowly lanceolate, acuminate, acute or somewhat obtuse at the
base, penninerved, shining, glabrous or obscurely puberulent, the blades mostly 2.5-4
cm. long and 0.7-1.5 cm. broad, petioles to 0.8 mm. long; stipules interpetiolar,
lanceolate, bifid to about the middle or sometimes entire, usually deciduous;
inflorescence axillary, 1-flowered or sometime juvenile branches appear to have a
cymose, 3-flowered inflorescence; flowers small, dull red; calyx reduced to an obscure
ring; corolla rotate, about 3 mm. long, 4-lobate nearly to the base, the lobes about 3
mm. long and 1.2 mm. broad, elliptic, acute, trinerved; fruits didymous, black,
shining, somewhat fleshy, to about 7 mm. high and 10 mm. broad.
Common in the cloud forest areas of Guatemala and Honduras.
The type is: wet mountain forest near Aldea Fraternidad, between
San Rafael Pie de la Cuesta and Palo Gordo, west facing slopes of
the Sierra Madre Mountains, department of San Marcos, alt. 1,800-
2,400 m. December 10-18, 1963, Williams, Molina & Williams 25944
(F). Twenty-nine other collections from Guatemala and Honduras
are available.
Didymaea microphylla L. Wms. sp. nov.
Pine or broadleaf forested slopes of the highland volcanoes or
mountains, 2,400-4,000 m.; Sacatepequez; Chimaltenango;
Huehuetenango; Quezaltenango. Endemic.
Herbae perparvae, caulibus 4-angulatis hispidulis glabrescentes; folia lanceolata
vel elliptico-lanceolata apiculata leviter carnosa revoluta hispidula aut glabrescentes;
stipulae bifidae lobis linearibus; inflorescentiae uni- vel rariores biflorae; calyces
perparvi aut nulli; corollae parvae 4-lobatae, tubi perbreves, lobis ovatis 3-
plinervibus; fructi carnosi nigri didymi.
Small herbaceous vines to 1 m. or less long, the stems 4-angulate, retrorse
hispidulous, glabrescent with age, stramineous; leaves small, lanceolate or elliptic-
lanceolate, sometimes ovate, acute and apiculate, somewhat fleshy and the margins
revolute, hispidulous or glabrescent, blades 4-9(-14) mm. long and 3-4(-8) mm. broad
without conspicuous lateral nerves, petioles often narrowly winged, mostly less than 2
mm. long; stipules about 1 mm. long, bifid to about the middle, the lobes linear;
inflorescences axillary (or terminal), uniflorous or rarely biflorous, the peduncles
short in anthesis, to 1 cm. long in fruit and somewhat reflexed; hypanthium glabrous,
at anthesis about 0.5 mm. long; calyx none or inconspicuous; corolla small, rotate, 4-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 63
lobate, glabrous, the tube very short, about 0.2 mm. long, the lobes ovate, about 2
mm. long and 1.5 mm. broad, prominently 3-plinerved; anthers in the sini of the
corolla lobes, about 0.4 mm. long, cordate; fruit fleshy, black and shining, didymous
or only one lobe developing.
Guatemala: narrow ridge of Volcan Santo Tomas, Dept.
Quezaltenango, alt. 2,500-3,700 m., Jan. 22, 1940, Steyermark 34790
(type, F).
Distinguished from the other species in Guatemala by the very
small somewhat fleshy leaves without lateral nerves visible.
DIODIA Linnaeus
Annual or perennial herbs, rarely low shrubs, sometimes scandent, branched,
glabrous or rough-pubescent, the stems terete or 4-angulate; leaves opposite, sessile or
nearly so, mostly narrow but sometimes ovate; stipules united with the petioles to
form a setiferous sheath; inflorescence axillary, sessile, the flowers small or minute,
white or purple; hypanthium obconic or obovoid, the calyx usually 2-4-dentate,
sometimes with alternating smaller teeth, persistent; corolla funnelform, the tube
short or elongate, the throat glabrous or villous, the limb commonly 4-lobate, the
lobes ovate-triangular, valvate in bud; stamens normally 4, inserted in the corolla
throat, the filaments filiform; anthers dorsifixed, linear-oblong, exserted; ovary 2-
celled, the style filiform, exserted, with 2 very short branches or a bilobate capitate
stigma; ovules solitary, affixed to the middle of the septum; fruit 2-coccous, the cocci
crustaceous, smooth or costate dorsally, indehiscent, without an interposed axis.
About 30 species in tropical America, a few in tropical Africa. A
few species besides those listed here occur in southern Central
America.
Diodia conferta DC., D. setigera Sw. and D. prostrata Sw. are
reported from Guatemala by Hemsley but we have seen no
Guatemalan specimens of these species.
The genus Crusea can be distinguished from Diodia in no very
substantial way and certain of the species of Crusea look more like
Diodia than they do to the remaining Cruseas. A careful generic
evaluation in this tribe is much needed.
Plants erect shrubs, 30-60 cm. high, with definitely woody branches; upper leaves
greatly reduced and bract-like, the inflorescence thus spike-like.
D. brasiliensis var. angulata.
Plants herbaceous, sometimes perennial but the stems not woody; upper leaves not or
scarcely reduced.
Plants prostrate, glabrous or nearly so, growing on seashores D. maritima.
Plants erect or scandent, abundantly pubescent with harsh hairs, not confined to
seashores.
Plants usually scandent; leaves lanceolate or ovate-oblong, mostly 1-2 cm. broad.
D. sarmentosa.
64 FIELDIANA: BOTANY, VOLUME 24
Plants not scandent; leaves narrowly or broadly linear, rarely as much as 4 mm.
broad.
Cocci of the fruit 3-costate dorsally, glabrous or nearly so D. rigida.
Cocci not costate, pubescent, usually densely so.
Stems glabrous; flowers in small, chiefly terminal heads.
See Crusea diversifolia.
Stems usually densely hirsute or hispidulous; flowers axillary D. teres.
Diodia brasiliensis Spreng. var. angulata (Benth.) Standl.
Carnegie Inst. Wash. Publ. 461: 90. 1935. Triodon angulatum
Benth. PL Hartw. 70. 1840. Tomillo silvestre. Figure 62.
Peten; Alta Verapaz; 300-1,400 m., in moist meadows or
pastures or on rocky stream banks, common; British Honduras;
southern Mexico to Honduras and Costa Rica; typical form of the
species in eastern Brazil and southward.
A stiff, densely branched shrub commonly 30-40 cm. high, sparsely puberulent or
almost glabrous, the branches sharply 4-angulate, densely leafy; leaves short -
petiolate, deep green, plane, oblong to elliptic, 5-20 mm. long, often appearing
verticillate, scaberulous on the margin and costa, the nerves obsolete; flowers very
small, greenish white, densely clustered in the axils of the greatly reduced upper
leaves and forming a spiciform inflorescence; calyx minute, 4-dentate; corolla 2 mm.
long; fruit broadly turbinate, green, glabrous, the cocci not costate dorsally.
By Bentham the plant was referred to a distinct genus,
Triodon, which has been maintained by some authors, with little
reason. In general appearance it does differ greatly from other
members of the genus, but the flower and fruit characters are those
of Diodia. The North American material differs little if at all from
the Brazilian plant.
Diodia maritima Thonning ex Schumacher, Besk. Guin. PL
75. 1827.
On sea beaches, British Honduras (New Town, Schipp 809); to
be expected on the beaches of Izabal. Honduras; Nicaragua; Costa
Rica; West Indies; Colombia; tropical Africa.
Plants prostrate, the stems coarse and stout, elongate, tetragonous, usually much
branched, glabrous or nearly so, 1.5 m. long or less, usually densely leafy and often
forming mats; leaves oblong or oblong- lanceolate, 1.5-3 cm. long, subsessile, acute or
obtuse, thick and when dry somewhat coriaceous, narrowed at the base; setae of the
sheaths about 3 mm. long; flowers solitary, sessile in the leaf axils; calyx lobes 4,
lanceolate, 2 mm. long; corolla white, 6-7 mm. long; fruit ellipsoid, glabrous, 5-7 mm.
long.
Diodia rigida (Willd.) Schlecht. & Cham. Linnaea 3: 341. 1828.
Spermacoce rigida Willd. ex Roem. & Schult. Syst. Veg. 3: 531.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 65
1818. D. pulchella Brandegee, Univ. Calif. Publ. Bot. 10: 418. 1924
(type from Chiapas). Mielillon; pegajosa; pelosa (fide Aguilar).
Common and widely distributed in hills and mountains, 2,400
m. or lower, usually in pine or oak forest, grassy fields and hillsides,
rocky banks, and other situations. Chiquimula; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Chimaltenango; Quiche";
Huehuetenango. British Honduras, and to be expected in Pete"n.
Southern Mexico to Panama, and widely dispersed in South
America.
Plants usually perennial, often from a somewhat woody base, procumbent, much
branched, at least from the base, the stems stout, rarely more than 30 cm. long,
hirsute or hispid with whitish spreading hairs, the internodes short; leaves linear,
mostly 1.5-3 cm. long, thick and stiff, broadest near the base, attenuate to a
setiferous apex, hispidulous or hirsute, usually revolute; calyx lobes short, subulate-
lanceolate, cilia te, erect, green; corolla pale purple, 8-10 mm. long, hispidulous outside
on the lobes, the tube glabrous; fruit subglobose, closely sessile, 3 mm. long, glabrous
or nearly so, conspicuously tricostate dorsally, at least when fully developed.
Very closely allied to D. teres and perhaps only varietally
distinct.
Diodia sarmentosa Swartz, Prodr. Veg. Ind. Occ. 30. 1788.
Figure 62.
Pine forest, wet thickets, and brushy slopes, chiefly at 600-1,400
m., sometimes at lower elevations, most abundant in Alta Verapaz;
Peten; Izabal; Chiquimula; Suchitepequez; Solola; San Marcos.
Southern Mexico to Panama, Guianas, and West Indies.
Plants sometimes procumbent but usually more or less scandent over shrubbery,
often forming dense tangles, the stems often 2 m. long or even more, 4-angulate,
brown, hispidulous or puberulent; stipule sheaths conspicuous, bearing numerous long
brown slender bristles; leaves lanceolate or oblong-lanceolate, 3.5-5 cm. long,
acuminate, obtuse at the base, aculeolate-serrulate, scabrous above, rather densely
hispidulous beneath, the lateral nerves very conspicuous, oblique, impressed above;
flowers sessile, 6-8 in the leaf axils; sepals 2-4, ovate, acuminate, ciliate; corolla white,
about 2 mm. long; capsule puberulent, the cocci 5 mm. long, not costate dorsally.
A common weedy plant of the Coban region, especially on cut-
over land, where it frequently is associated with Dicranopteris and
other scrambling plants.
Diodia teres Walt. Fl. Carol. 87. 1788. D. prostrata Swartz,
Prodr. Veg. Ind. Occ. 30. 1788.
Moist or dry, oak or pine forest, grassy plains, or cornfields,
200-1,800 m.; Zacapa; Jalapa; Guatemala; Chimaltenango;
66 FIELDIANA: BOTANY, VOLUME 24
Huehuetenango. Widely distributed in the United States; Mexico;
ranging southward to Costa Rica and South America.
Plants usually stiffly erect, annual, strict or often copiously branched from or
above the base, the stems slender and wiry, hispidulous and hirsute with whitish
hairs; stipule sheaths conspicuous, with numerous long slender whitish setae; leaves
linear or lance-linear, attenuate to the setiferous apex, broadest near the base,
scaberulous and hirsute, usually revolute, with thickened margins, 1-nerved; flowers
sessile, solitary or clustered in the leaf axils, the sepals short, acute, greenish; corolla
small, white or purplish, usually 3-4 mm. long; fruit about 3 mm. high, densely
covered with short, whitish, spreading or appressed hairs, the carpels rounded on the
back, not at all costate.
As this species has been subdivided by Fernald and Griscom,
the Guatemalan material would fall under var. setifera Fernald &
Griscom (Rhodora 39: 307. 1937), ranging from Michigan to Texas
and southward. The variety, however, seems to have little
taxonomic importance, and is not separated definitely from the
typical form of the Atlantic region of the United States. The
terminal bristles of the leaves which are supposed to separate the
western plant from the eastern are almost as conspicuous in plants
of the Atlantic coast as in Mexican ones, and are by no means
lacking, as claimed by those authors.
DUROIA Linnaeus f.
Shrubs or trees, unarmed, glabrous or pubescent, the branchlets terete or
tetragonous; stipules interpetiolar, oblong, deciduous; leaves opposite or verticillate,
sessile or petiolate, coriaceous to membranaceous; flowers unisexual, often large,
white or yellowish, in terminal, sessile or pedunculate fascicles or cymes, the pistillate
rarely solitary; hypanthium oblong to hemispheric; calyx cupular or tubular,
persistent, truncate or 6-9-lobate; corolla salverform, often coriaceous, sericeous
outside, the throat pilose or naked, the limb 6-9-lobate, the lobes oblong, contorted in
bud; stamens 6-9, inserted in the corolla tube, the filaments very short or none;
anthers dorsifixed, linear, acute, included; ovary 2-4-celled, the style short, with 2
broad acute coherent branches; ovules numerous, biseriate, the placentae affixed to
the septum; fruit baccate, globose to oblong, with a thick cortex, 2-4-celled or 1-
celled; seeds large, horizontal, much compressed, suborbicular, covered with pulp, the
testa thin, black.
A small genus with one species in Costa Rica, the others,
besides the one here listed, South American. The genus is hardly
distinct from Amaioua.
Duroia genipifolia Standl. & Steyerm. Field Mus. Bot. 22:
186. 1940.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 67
Type from Izabal, Rio Dulce, between Livingston and 6 miles
up river, at sea level, Steyermark 39382; also no. 39572 from the
same general region.
A large shrub or small tree with thick glabrous branchlets; stipules deciduous,
almost 1 cm. long, subulate-attenuate, densely pilose; leaves large, short-petiolate,
membranaceous, the densely pilose petiole 1.5 cm. long; leaf blades obovate-oblong or
oblanceolate-oblong, 20-30 cm. long, 8-10 cm. broad, acuminate, gradually narrowed
to the acute base, sparsely hirtellous above, rough to the touch, densely short-pilose
beneath with spreading hairs, the lateral nerves about 16 on each side; staminate
flowers capitate-congested at the ends of the branches, numerous, sessile or short-
pedicellate; hypanthium campanulate, 5-6 mm. long, densely hispidulous; calyx
truncate, the 5 lobes subulate, erect, unequal, 8 mm. long or less; corolla in bud 18
mm. long, attenuate to the apex, whitish-strigose, the lobes linear-attenuate, three
times as long as the tube, contorted.
EIZIA Standley
Almost glabrous shrubs, the branchlets subterete; stipules interpetiolar, very
short, reduced almost to a line; leaves opposite, petiolate, membranaceous;
inflorescence terminal, cymose, lax, few-flowered, pedunculate, the flowers large,
pedicellate, the bracts and bractlets inconspicuous; hypanthium clavate, attenuate to
the base, costate-angulate; calyx deeply 4-lobate, green, the lobes broad, persistent;
corolla salverform, the very long tube slender, slightly dilated upward, glabrous
within, the 4 lobes slightly imbricate (or convolute?) in bud, spreading, oblong,
glabrous within; stamens 4. inserted at the apex of the tube, the filaments very short;
anthers dorsifixed, erect, obtuse, subexserted; ovary 2-celled, the style very long and
slender, included, shortly bifid at the apex; ovules numerous; capsule obovoid,
costate, produced above the persistent calyx, subcoriaceous, septicidally bivalvate at
the apex, the valves biparted at the apex; seeds numerous, angulate, tuberculate-
rugose.
The genus consists of a single species.
Eizia mexicana Standl. Field Mus. Bot. 22: 50. 1940.
Type from Volcdn de Tacand, Chiapas, at 1,000-2,000 m.,
Matuda 2402; doubtless occurring on the Guatemalan side of the
volcano.
Petioles glabrous, slender, 1-2.5 cm. long; leaf blades ovate or oblong-ovate, 7.5-
10 cm. long, 2.5-5.5 cm. broad, narrowly long-acuminate, obtuse or acute at the base
and often somewhat oblique, glabrous above, barbate beneath in the nerve axils, the
lateral nerves about 6 on each side; peduncles 1.5 cm. long or less, the inflorescence
sometimes sessile, few-flowered, as much as 7 cm. long and broad, the stiff pedicels
1.5-2.5 cm. long; hypanthium glabrous, 4-5 mm. long, the calyx 2 mm. long, its lobes
broadly triangular, acutish; corolla puberulent outside, at least above, the tube 5 cm.
long, 4 mm. broad at the throat, the lobes 8 mm. long; anthers 6 mm. long; capsule
about 12 mm. long and 8 mm. broad, obtuse or subacute at the base; seeds irregularly
angulate, 0.6-0.8 mm. in diameter, dark brown.
68 FIELDIANA: BOTANY, VOLUME 24
The genus was named for its collector, Eizi Matuda, who made
extensive collections in Chiapas near the Guatemalan border,
especially upon the Volcano of Tacana, over whose summit the
international boundary passes.
ELAEAGIA Weddell
Trees or large shrubs, pubescent, with terete branches; stipules interpetiolar, free
or connate at the base; leaves opposite, petiolate, coriaceous to membranaceous;
flowers small, in terminal racemes or panicles; hypanthium small, hemispheric,
sulcate; calyx somewhat ampliate, 5-lobate, persistent; corolla short-funnelform, the
throat villous, the limb 5-lobate, the lobes oblong, recurved, contorted in bud;
stamens 5, inserted between the corolla lobes, the filaments exserted; anthers
dorsifixed, oblong; ovary 2-celled, the style short, bifid, with short obtuse branches;
ovules numerous, crowded, the placentae peltately attached to the septum; capsule
very small, crustaceous, crowned by the persistent calyx, 2-celled, loculicidally
bivalvate, the valves finally bifid; seeds numerous, minute, elongate, with
membranaceous testa.
One other species is known from Costa Rica, and a small
number of others are confined to South America, chiefly in
mountainous areas.
Elaeagia auriculata Hemsl. Diag. PI. Mex. 32. 1879.
Known in Guatemala only from sterile material, almost surely
referable here, collected on Cerro Tixixi, north of Jocotan,
Chiquimula, at about 1,300 m., Steyermark 31579. Mountains of
Honduras and Costa Rica.
A shrub or tree of 3-8 m., in Honduras reported to reach a height of 24 m., the
branchlets tomentulose; stipules very large, 5 cm. long or even larger; leaves sessile or
nearly so, ovate-elliptic to rounded-elliptic-obovate, as much as 40 cm. long and 23
cm. broad or even larger, but often smaller, acute or abruptly short-acuminate,
narrowed to the auriculate base, densely and softly short-pilose, especially beneath;
flowers white, in large broad sessile panicles; calyx and hypanthium puberulent,
together scarcely more than 2 mm. long; corolla 3-4 mm. long, glabrous outside,
lobate almost to the base; capsule subglobose, 3 mm. in diameter.
Said to be called "jagua" in Honduras.
ERNODEA Swartz
Small shrubs usually near or along the sea beaches; stems terete or angled, the
internodes shorter than the leaves, stipules forming a sheath, this bidentate or entire,
intrapetiolar. Leaves opposite, coriaceous, sessile or nearly so; inflorescence axillary,
usually consisting of a single flower; flowers small; calyx tubular, 4-5-lobate, the
lobes erect, subulate-lanceolate; corolla with a narrow elongated tube, the throat
naked, the 4-6 lobes spreading, linear, valvate; stamens 4-6, inserted on the tube,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 69
anthers exserted, linear; style somewhat bilobate, long and exserted; fruit a
somewhat fleshy drupe, indehiscent, obovoid, with a single seed in each cell.
A small genus of perhaps some nine species, only the following
widely distributed.
Ernodea littoralis Sw. Nov. Gen. & Sp. PL Prodr. 29. 1788.
Figure 60.
Coastal thickets near the sea; Florida and the Yucatan
Peninsula to British Honduras, the West Indies to Colombia.
Small erect or usually repent shrubs up to 1 m. tall. Leaves coriaceous,
lanceolate to linear-lanceolate, acute, glabrous, to about 4 cm. long and 1 cm. broad
(ours), sessile or nearly so; the inflorescences of single axillary flowers; the calyx 4-6-
lobate, the lobes linear to subulate-lanceolate, 2-3 mm. long; corolla usually reddish,
the tube about 1 cm. long, the lobes linear, spreading, 4-5 mm. long; fruits
subbaccate, obovoid with persistent calyx, about 5 mm. long.
Known from Central America only from British Honduras.
EXOSTEMA L. Richard
Shrubs or trees, glabrous or pubescent, the branchlets usually terete; stipules
interpetiolar, deciduous or persistent; leaves opposite, petiolate or subsessile,
membranaceous or coriaceous; flowers small or large, axillary and solitary or
arranged in terminal panicles or corymbs, pedicellate, bracteolate or ebracteolate;
hypanthium cylindric or obovoid; calyx normally 5-lobate, the lobes broad or
narrow; corolla short or elongate, the tube elongate, often very long, the throat
glabrous or pilose, the limb 5-lobate, the lobes oblong or linear, imbricate in bud, two
of them exterior; stamens 5, inserted at the base of the corolla tube, the filaments
filiform, elongate; anthers basifixed, linear, usually exserted; ovary 2-celled, the style
filiform, usually exserted, the stigma capitate, entire or bilobate; ovules numerous,
ascending, the placentae adnate to the septum; capsule 2-celled, usually oblong-
cylindric, septicidally bivalvate, the valves entire or biparted; seeds numerous,
imbricate, the body oblong, compressed, the testa produced into a wing.
About 35 species, chiefly West Indian, one or two others in
Mexico, and several in the mountains of South America.
Flowers axillary, mostly solitary E. caribaeum.
Flowers in terminal or axillary panicles E. mexicanum.
Exostema canescens Bartl. is reported from Guatemala by
Hemsley. The application of the name is uncertain, and it is
doubtful what the Guatemalan plant so reported may be, possibly
E. mexicanum.
Exostema caribaeum (Jacq.) R. & S. Syst. Veg. 5: 19. 1819.
Cinchona caribaea Jacq. Enum. PI. Carib. 16. 1760.
70 FIELDIANA: BOTANY, VOLUME 24
Dry, brushy, often rocky slopes, 400-1,300 m.; Chiquimula;
Huehuetenango. British Honduras, and to be expected in Pet6n;
southern Mexico to Honduras and Costa Rica; southern Florida
and West Indies; northwestern coast of South America.
A shrub or tree, rarely more tnan 8 m. high, the branchlets glabrous; stipules 2.5-
5 mm. long; leaves short-petiolate, the blades mostly ovate, 5-11 cm. long, 1.5-5 cm.
broad, rather abruptly acuminate, obtuse or acute at the base, glabrous above,
usually glabrous beneath except in the barbate nerve axils, rarely white-pilose with
short hairs; flowers solitary in the leaf axils, the stout pedicels 4-10 mm. long;
hypanthium glabrous, 4-5 mm. long; calyx lobes 1 mm. long or less, broad, obtuse or
acutish; corolla white, glabrous, the tube 3-5 cm. long, 2 mm. thick, often curved, the
lobes linear, recurved, about equaling the tube; stamens long-exserted, the linear
anthers 2 cm. long; capsule oval or ellipsoid, 1-1.5 cm. long, smooth, dark brown,
lustrous; seeds oval, 3-6 mm. long, with a narrow brownish wing.
Maya names of Yucatan are reported as "zabacche" and
"chactsiis." The leaves and bark are bitter, and formerly the plant
was used in some parts of its range as a substitute for quinine.
Exostema mexicanum Gray, Proc. Am. Acad. 5: 180. 1861.
Quina; melena de leon (fide Aguilar). Figure 17.
Dry or wet forest, usually along stream banks, 1,500 m. o/
lower; Peten; Escuintla; Retalhuleu; Quiche; Huehuetenango.
Southern Mexico; British Honduras; Honduras; El Salvador;
possibly Costa Rica.
A small or large tree, reported as reaching a height of 23 m. with a trunk
diameter of 40 cm., the slender branchlets glabrous or obscurely puberulent,
conspicuously lenticellate; stipules 2.5-4 mm. long, triangular, acuminate or
cuspidate; petioles 1 cm. long or less, the blades ovate to oval or oblong-ovate, 5-15
cm. long, 2-7.5 cm. broad, abruptly acuminate or attenuate, very obtuse or broadly
rounded at the base, barbate beneath in the nerve axils, otherwise glabrous or nearly
so; inflorescence cymose- corymbose, 7-10 cm. broad, densely many-flowered, the
slender pedicels 2-4 mm. long, minutely puberulent; hypanthium 2.5-3 mm. long,
sparsely and minutely puberulent; calyx lobes minute, deltoid, acute; corolla sparsely
villosulous, the tube 8-10 mm. long, the lobes oblanceolate- linear, obtuse, 1 cm. long;
anthers 3.5-4 mm. long; capsule clavate-obovoid, about 1 cm. long, dark brown,
conspicuously whitish-lenticellate.
Called "quina" in El Salvador. The Maya name "sabac-ch£" is
reported from British Honduras. The flowers are fragrant. In
Huehuetenango a decoction of the bitter bark is employed as a
remedy for malaria.
FARAMEA Aublet
Shrubs or small trees, usually glabrous throughout, the branchlets compressed,
terete, or tetragonous; stipules intrapetiolar, short-triangular and apically long-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 71
aristate or often long-sheathing, usually persistent; leaves opposite, petiolate or
subsessile, chiefly coriaceous and oblong or lanceolate; inflorescence various, terminal
or axillary, few-many-flowered; hypanthium ovoid or turbinate, terete or angulate;
calyx cupular or short-tubular, persistent, truncate or rarely 4-dentate; corolla
commonly salverform, the tube short or elongate, the throat naked, the lobes 4
(rarely 5) usually linear or lanceolate, valvate in bud, spreading or recurving; stamens
4, or rarely 5, inserted in the tube or throat of the corolla, the filaments short or long;
anthers dorsifixed near the base, included or exserted, linear; ovary 1 -celled, the style
filiform, with 2 short branches; ovules geminate, basilar and collateral, erect; fruit
coriaceous, globose or transverse-oblong, terete or costate, 1-celled and 1-seeded; seed
transverse, usually deeply excavate at the base, large, with very thin testa and
corneous endosperm.
A large genus of tropical America, most of the species South
American, a few others occurring in Mexico and southern Central
America. The group is easily recognized by the single large seed,
sometimes horizontal.
Flowers fasciculate or solitary in the leaf axils F. cobana.
Flowers in few- to many-flowered pedunculate inflorescences.
Inflorescence a short few-flowered raceme; calyx about 1 mm. long.
F. standleyana.
Inflorescence not racemose, usually many-flowered cymes or corymbs.
Stipules distinct or nearly so; corolla white F. occidentalis.
Stipules united for half or even most of their length.
Corolla pale blue; corolla lobes usually longer than the tube....F. brachysiphon.
Corolla white; corolla lobes about equaling the tube F. belizensis.
Faramea cobana Donn.-Sm. Bot. Gaz. 57: 422. 1914.
Moist or wet, mixed, mountain forest, 1,200-1,600 m.; endemic;
Alta Verapaz (type from forest near Coban, Tuerckheim 2474};
Huehuetenango (Cerro Chiblac).
A glabrous shrub 1-2.5 m. high, the branchlets angulate; stipules long-aristate
from a short base; petioles only 2-4 mm. long, the blades oblong- lanceolate, 5.5-7.5
cm. long, 1.3-2 cm. broad, narrowed to a narrowly obtuse apex, acute at the base, the
lateral nerves 9-11 on each side; pedicels solitary or fasciculate in the upper leaf axils,
about equaling the flowers, filiform, 12-25 mm. long; calyx and hypanthium 2 mm.
long, the hypanthium turbinate, the calyx minutely denticulate; corolla blue or
white, about 12 mm. long, the tube gradually dilated upward, the lobes oblong-ovate,
half as long as the tube or longer; anthers included, 4 mm. long; fruit globose, blue,
smooth, 9 mm. in diameter.
Faramea belizensis Standl. Carnegie Inst. Wash. Publ. 461:
90. 1935.
Type Schipp S-721 from Camp 36 of the Guatemala-British
Honduras boundary, Peten, alt. 810 m.
72 FIELDIANA: BOTANY, VOLUME 24
A glabrous tree, 8 m. high, the trunk 7.5 cm. in diameter, the branchlets obtusely
tetragonous; stipules 5-8 mm. long, connate half their length, the free portion short-
mucronate; leaves coriaceous, darkening when dried, on petioles 1 cm. long or less,
ovate to oblong, about 14 cm. long and 4-7.5 cm. broad, caudate-acuminate, obtuse or
acute at the base, the lateral nerves about 13 on each side; inflorescence terminal,
cymose-umbellate, many-flowered, about 5 cm. long and 7 cm. broad, long-
pedunculate, the flowers in 3-flowered cymules, the pedicels stout, 3-5 mm. long;
hypanthium obovoid, 1.5 mm. long; calyx tubular-campanulate, 3-5 mm. long,
truncate; corolla attenuate in bud, white, the rather thick tube 1 cm. long, the linear
attenuate lobes of about the same length.
The collector states that this is a "very handsome" shrub or
small tree, seen only at high altitudes on mountain slopes in dense
shade; "rare." The flowers are said to be fragrant.
Faramea brachysiphon Standl. Field Mus. Bot. 8: 62. 1930.
Tinta.
Dense wet mixed lowland forest, 500 m. or lower; Alta Verapaz;
Izabal; Huehuetenango. British Honduras, the type from Middlesex,
Schipp 345.
A glabrous shrub or small tree 2.5-6 m. high, the branchlets subterete; stipules
green, the sheath 3-4 mm. long, mucronate, the mucro about 1 mm. long; leaves on
petioles 4-7 mm. long, subcoriaceous, oblong or narrowly oblong, 11-17 cm. long, 3-5
cm. broad, subabruptly acuminate, acute at the base, the lateral nerves about 15 on
each side; inflorescence cymose-paniculate, about 3.5 cm. broad, on a peduncle 2-2.5
cm. long, the flowers few or numerous, rather densely crowded, the pedicels 1-2 mm.
long; hypanthium broadly oblong, 1.5 mm. long, the calyx teeth triangular, minute,
scarcely 0.5 mm. long; corolla pale blue, glabrous, the rather thick tube 3 mm. long,
the lobes lance-oblong, 6-7 mm. long, obtuse, spreading; anthers linear, partly
exserted.
The fruit, not seen, is described as globose and dull green.
Faramea occidentalis (L.) A. Rich. Mem. Soc. Hist. Nat.
Paris 5: 176. 1834. Ixora occidentalis L. Syst. Nat. ed. 10. 2: 893.
1759. F. odoratissima DC. Prodr. 4: 496. 1830. Cerezo de montaha
(Quezaltenango). Figure 55.
Rather widely distributed, especially in the lowlands of both
slopes, ascending in the mountains of the western highlands to 1,500
m., and descending to sea level, in dense wet mixed forest or
sometimes in dry thickets of the Pacific coastal plains; Peten;
Izabal; Suchitepequez; Retalhuleu; Quezaltenango; San Marcos.
Southern Mexico; British Honduras to Panama, southward to Peru;
West Indies.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 73
A slender glabrous shrub or tree, rarely 9 m. high, usually much lower, the
branchlets green, subterete; stipules short and small, long-aristate; leaves short-
petiolate, oblong or lance-oblong, short-acuminate or caudate-acuminate, acute to
attenuate at the base, often turning dark when dried; inflorescences chiefly terminal,
few-flowered, corymbose, sometimes umbelliform, the flowers long-pedicellate,
fragrant; hypanthium obovoid, 2 mm. long; calyx cupular, 3 mm. long, obscurely
denticulate; corolla white, glabrous, 2-2.5 cm. long, the linear-lanceolate attenuate
lobes equaling or longer than the tube, 4 or rarely 5 (as in illustration); anthers
included; fruit depressed-globose, about 1.5 cm. broad, purple or wine-purple at
maturity or almost black.
In Mexico the shrub often is known by the name "cafecillo,"
and from Oaxaca the names "huesillo" and "azucenilla" are
reported.
The species is a variable one and it is possible that there is
more than one species in the material we have placed here. The
great altitudinal range is unusual.
Faramea standleyana L. Wms. Phytologia 26: 490. 1973.
Figure 56.
Known only from rocky slopes of Cerro San Gil, along Rio Frio,
Dept. Izabal, Steyermark 39962. Endemic.
Shrubs to about 2 m. tall, entirely glabrous; the branchlets terete, the intemodes
5-7 cm. long; stipules intrapetiolar, about 5 mm. long, the long aristate tip to 1 cm.
long; leaves elliptic or elliptic-oblanceolate, abruptly and shortly acuminate,
attenuate to the base, with 8-10 pairs of lateral nerves, blades 11-15 cm. long and 2-6
cm. broad, petiole 1-1.5 cm. long; inflorescence terminal, a few-flowered raceme, the
peduncle slender, about 2 cm. long, the pedicels up to 1 cm. long; calyx suburceolate,
about 1 mm. long, lobed to the middle, the lobes triangular, acute; corolla white with
the tube pale lilac, salverform, the tube slender, 2-2.5 cm. long, the lobes lanceolate,
acute, about 8-9 mm. long; the stamens attached near the middle of the corolla tube,
the anthers linear, 8-10 mm. long, the filaments as long as the anthers; style about 15
mm. long, bifid; fruits unknown.
Named for the senior author of this flora, who has described
not fewer than 17 species of the genus from tropical North America
— and many others from South America.
ADDITIONAL SPECIES:
Faramea sp.
Stann Creek district, British Honduras, Gentle 2869.
Small tree to 6 m. and 5 cm. in diameter, completely glabrous. Leaves elliptic,
acuminate, attenute to the short petiole, 9-17 cm. long and 2.5-4 cm. broad, petioles
5-10 mm. long; inflorescence few-flowered subumbellate cymes on peduncles about 1-
1.5 cm. long; hypanthium and calyx about 3 mm. long, the calyx about 1 mm. long,
the shallow lobes about 0.5 mm. long; disc on summit of ovary exceeding the calyx;
fruits verriculose (immature), about 1 cm. wide and half as high.
74 FIELDIANA: BOTANY, VOLUME 24
Related to F. stenura Standl. but inadequate for sure
determination.
GALIUM Linnaeus
CONTRIBUTED BY LAURAMAY DEMPSTER
References: Greenman, J. M., Revision of the Mexican and
Central American species of Galiwn and Relbunium, Proc. Am.
Acad 33: 468. 1898. Dempster, L. T., The fleshy-fruited Galiums
(Rubiaceae) of Mexico and Central America, Brittonia 25: 15. 1973.
Annual or (ours) perennial herbs, or sometimes suffrutescent; stems square;
leaves entire, opposite, but seemingly in whorls or 4-8 because of the leaf-like stipular
appendages; inflorescence of simple or usually compound dichasia on lateral
branches, or sometimes terminal; flowers small, pedicellate, perfect, polygamous or
dioecious; calyx lacking; corolla rotate or sometimes a little campanulate, usually 4-
lobed, white, yellow or red; stamens 4, inserted in the corolla throat alternately with
the lobes; anthers versatile, oblong; ovary 2-celled, the stigma bifid; ovules 1 in each
locule; fruit consisting of 2 dry 1 -seeded, usually more or less hemispherical
mericarps, or sometimes baccate, often 1-seeded by abortion, glabrous, tuberculate or
variously hairy; seeds not dehiscent, the two carpels falling separately at maturity;
endosperm horny.
About 400 species, in all continents except Antarctica, 22 in
mainland Mexico and Central America.
Leaves in whorls of 6 to 8.
Flowers white; Alta Verapaz G. mexicanum.
Flowers pink or red; San Marcos and Huehuetenango to Jalapa.
G. mexicanum var. platyphyllum.
Leaves in whorls of 4.
Fruits fleshy, glabrous G. aschenbornii.
Fruits dry, not glabrous.
Tiny moss-like plants with tuberculate fruits.
G. sphagnophilum var. mazocarpum.
Larger plants; fruits with curved or uncinate hairs.
Corollas a little campanulate, usually hispid externally; plants often prostrate
or sprawling. G. uncinulatum.
Corollas rotate, spreading, glabrous; semierect plants.
Leaves more or less obviously 3-nerved, tapered at base and apex, the
surfaces hispid or pubescent.
Leaves 2-8 mm. long; plants of alpine meadows; corollas usually red.
G. nelsonii.
Leaves mostly over 8 mm. long; plants of the forest belt; corollas white or
greenish G. orizabense.
Leaves 1 -nerved, abruptly petiolate at base, round-apiculate at apex, the
surfaces nearly glabrous G. quichense.
Galium aschenbornii Schauer, Linnaea 20: 701. 1847.
Relbunium aschenbornii (Schauer) Hemsley, Biol. Cent. Am. Bot.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 75
2: 62. 1881. Rubia acuminata Mart. & Gal. Bull. Acad. Brux. 11:
127. 1844. G. geminiflorum Mart. & Gal. op. cit. 11: 126. G.
galeottianum Walp. Rep. 6: 17. 1846. G. uropetalwn Hemsley, op.
cit. 66. Pegapega; ojo de perdis.
Widespread in mountainous areas at 1,200-3,200 m. Climbing on
shrubs or trailing on the ground; thickets, banks or grassy slopes,
commonly in mixed forest of oak with pine or cypress. San Marcos
and southern Huehuetenango to Zacapa, Chiquimula and Utiapa.
Mexico; Honduras; El Salvador; Costa Rica; Panama.
Perennial trailing or climbing plants with long (to 120 cm.) slender wiry stems
arising in clumps from a small rootstock, or singly from older woody stems, often
rooting at the nodes; persistent woody stems remaining slender and wiry; young
fertile stems arising singly from the axils, much shorter than the main stems; stems
more or less scabrous with very short retrorse aculeolate hairs, or sometimes glabrous
toward the end; internodes 1-5 times as long as leaves; leaf surfaces generally
glabrous, the margins set with few to many short aculeolate retrorse hairs; leaves
ovate, lanceolate, elliptical or oblong, commonly 7-15 mm. long, apparently 1-nerved,
apiculate, narrowed more or less abruptly to a short but rather definite petiole;
flowers staminate, pistillate, or perfect, on the same plant or on different plants;
corollas glabrous, usually red or pink, but sometimes yellow, white, or greenish, the
lobes with long slender apices; fruits glabrous, fleshy, green when young, becoming
red, then black, wrinkled when mature, 3-7 mm. across when dry.
Known as bermelloncillo in Costa Rica, and mala mujer in
Jalisco.
Galium mexicanum HBK. Nov. Gen. et Sp. 3: 337. 1818.
Pegagosa de pena; trementino de cerro; flor cadena.
Plants herbaceous above ground, rooting at the nodes; stems to 240 cm. long,
trailing or climbing among shrubs, scabrous with retrorse aculeate hairs, and often
also more or less abundantly set with long straight hairs; leaves 1-nerved, 6-8 to a
node, 1-2.5 cm. long, one-fourth to two-thirds as long as nodes, variously
oblanceolate, tapered gradually to base, and abruptly to the pungent apex; lower
midrib and callous margins set with retrorse aculeate hairs, the upper surface with
short apically-directed hairs, the lower surface usually more or less hirsute with long
slender hairs, but sometimes nearly glabrous; inflorescences on short leafy lateral
branches, the pedicels and branchlets divaricate; flowers perfect; ovaries turbinate,
densely set with apically-directed, strongly arcuate, hairs; corollas campanulate, cleft
about halfway, white, externally set with long hairs, or often not, usually in
conformity with the lower leaf surfaces, the lobes relatively short and blunt; fruits
dry, set with short apically-directed hairs which are strongly curved throughout their
entire length.
Galium mexicanum s.l. is a widespread species whose taxonomy
requires much further research, including chromosome study, since
polyploidy is known to occur. In the broadest sense, it should
include not only the Central American and Mexican plants of the
76 FIELDIANA: BOTANY, VOLUME 24
name, but also those plants of western United States which are
generally called G. asperrimum Gray.
The plants are extremely adhesive, owing to the many retrorse
hairs on stems and leaves, as well as to the strongly curved hairs of
the fruits. The former are very effective in helping the plant to
climb, and the latter, or both, in seed dispersal. The readiness of
this plant to cling to clothing and animal fur accounts for most of
the common names. In addition to pegagosa, it is also called
esculona, rama del coyote, flor de pulga, and yerba de la pulga in
Mexico. It is said to be put under the bed to catch fleas, and is
sometimes sold in the market for this purpose. It is also said to
curdle milk.
Galium mexicanum var. platyphyllum Greenm. Contr. Gray
Herb. n. ser. 14: 458. 1898.
Common in the mountains, 1,300-3,100 m., in wet or moist
places in ravines, thickets and mixed forest; San Marcos;
Huehuetenango; Quezaltenango; Totonicapan; Quiche; Solola;
Chimaltenango; Sacatepequez; Guatemala; Baja Verapaz; Jalapa.
Mexico, (lectotype from Chiapas, near San Cristobal, Nelson 3165
(GH!), the first of five specimens cited by Greenman).
Corollas pink or red, usually larger than in var. mexicanum; leaves commonly
somewhat larger and often much broader, usually very abrupt at apex, often truncate
or even slightly emarginate.
The original color of dried corollas is always uncertain, but
collectors' records indicate that Guatemalan material, except in
Alta Verapaz, has consistently red or pink flowers. The leaf
characters associated with colored corollas are difficult to define
with precision, but are none the less real, as would be apparent
from a biometrical study. Outside of Guatemala, red flowers are
reported only from Chiapas, the type locality of var. platyphyllum.
Galium nelsonii Greenm. Proc. Am. Acad. 33: 460. 1898.
In alpine meadows or among limestone rocks, commonly with
Juniper, at 3,300-3,900 m. Huehuetenango. Mexico.
Stems erect, procumbent at base, 4-40 cm. long, nearly glabrous, or sometimes
pubescent; leaves 2-8 mm. long, ovate, obscurely to obviously 3-nerved, tapered to a
broad base and acute apex, more or less hispid with scattered long hairs; inflorescence
racemose, on short, few-flowered, often remote lateral branches that are ascending in
flower, divaricate in fruit; corollas rotate, red (or sometimes yellow), glabrous; fruits
dry, set with many uncinate hairs.
DEMPSTER: FLORA OF GUATEMALA 77
Galium orizabense Hemsley, Diag. PI. Nov. pars 3:54. 1878.
Steep moist slopes and canyons in mixed forest, 1,350-2,700 m.
The only Guatemalan collections seen were made by von
Tuerckheim in the vicinity of Coban, Alta Verapaz, at 1,350-1,400
m. Mountains of eastern Mexico from Tamaulipas south to Puebla
and Vera Cruz; Chiapas. Plants of Costa Rica and Panama are
doubtfully referred here.
Erect or subscandent plants, the stems 20-65 cm. long, tufted from a small
rootcrown, or commonly rooting at the nodes; stems finely pubescent with curly
hairs, the internodes 2-5 times as long as leaves; leaves 3-nerved, 8-25 mm. long,
narrowly lanceolate-oblanceolate to elliptical, drawn to a petiolar base, apiculate or
obtuse at apex, sparsely set above and below with curved or curly apically-directed
hairs; flowers perfect, minute, commonly 5-15 on short lateral branches; corollas
rotate, white or greenish; fruits dry, set with uncinate hairs.
Galium quichense Dempster, Phytologia 26: 12. 1973.
On mossy ledges, about 2,200 m.
Procumbent perennial herb, 10-30 cm. high from a slender taproot or small
rootcrown, sometimes rooting at the nodes; stems entirely glabrous, or the sides set
with minute upwardly-curved hairs; leaves 3-7 mm. long, 1-nerved, somewhat thick,
tapering rather abruptly to a petiole, the blades broadly elliptical to orbicular,
apiculate; leaf blades nearly glabrous or with few short hairs on the upper surface,
the lower half of the midrib and the petiole densely set beneath with short stout
upwardly-curving hairs; lower leaf surface densely and evenly dotted with large
glandular cells; inflorescence terminal, pyramidal; pedicels 1-3 times as long as the
flowers, very sharply flexed just below the ovary, elongating and becoming more
divaricate in fruit; flowers perfect; corollas rotate, cream-color, glabrous, the apices
obtuse; fruits dry, 1-1.25 mm. across, set with uncinate hairs.
The type collection is the only one known: near junction of
Nebaj road with road between Sacapulas and Cunen at 6,700-7,300
ft. (2,168-2,362 m.), Proctor 25139 (LL!, isotype IJ).
Galium sphagnophilum (Greenm.) Dempster var.
mazocarpum (Greenm.) Dempster, Phytologia 26: 222. 1973. R.
mazocarpum Greenm. Proc. Am. Acad. 41: 250. 1905.
Growing interfingered with the mosses and other tiny plants in
wet alpine meadows, and on moist banks near springs, 1,800-3,700
m. Huehuetenango; El Quiche". Mexico.
Tiny slender creeping perennial, rooting at the nodes; leaves and stems
essentially glabrous, or sometimes with a few hairs; leaves 4 to a node, one-sixth to
one- third as long as internodes, spindle-shaped or obovate, apparently 1-nerved,
tapering to a long petiolar base and to the acute apex, tipped with a long hair;
flowers usually solitary on short leafy lateral branches, which are largely included;
78 FIELDIANA: BOTANY, VOLUME 24
corollas rotate, hispid externally, greenish-white; fruits dry, tuberculate, the pedicels
very short.
This seldom-collected species may actually be common in wet
meadows and about springs. Since it is no larger than the moss with
which it mingles, it is probably often overlooked by collectors. The
var. sphagnophilum, which has not been collected in Guatemala,
differs only in having smooth ovaries and fruits, and it may well be
a mere trivial form. The three Guatemalan collections and three of
the four Mexican collections seen, other than the type, have
tuberculate fruits.
Galium uncinulatum DC. Prodr. 4: 600. 1830. G. obstipum
Schlecht. Linnaea 9: 592. 1834.
Sierra Madre and mountains to the north, at 1,300-3,300 m.
Hillsides, ravines, or among limestone rocks, in open mixed forest;
Alta Verapaz; El Quiche; Huehuetenango; Totonicapan;
Quezaltenango; San Marcos. Throughout Mexico, except in the
lowlands, to Texas; Costa Rica; Panama.
Lax prostrate or procumbent plants, the herbaceous stems 20-60 (100) cm. long,
many from a small rootcrown, often rooting at the nodes; stems and leaves more or
less densely pubescent; leaves 4 to a node, 3-nerved, broadly elliptical or ovate-
obovate to nearly orbicular, tapered to a more or less petiolar base, apiculate at apex,
8-12 (18) mm. long, one-fifth to one-half as long as the internodes; flowers borne on
short bracteose few-flowered lateral branches, the branchlets and pedicels at
maturity becoming elongated and widely divaricate, the pedicels often sharply bent
below the fruit; ovaries with many uncinate hairs about as long as carpel width;
corollas greenish yellow to nearly white, commonly appearing no larger than the
densely hairy ovary, usually hispid externally, more or less campanulate at base, the
lobes not widely spreading; fruit dry or sometimes a little fleshy, the body 1.5-2 mm.
wide, set with many uncinate hairs.
This is a common and variable species, ranging from Panama to
southern Texas and reappearing in Baja California.
GARDENIA Linnaeus
Shrubs or trees, glabrous or pubescent, generally unarmed, the branchlets terete;
stipules intrapetiolar, triangular, acute or acuminate, often connate at the base;
leaves opposite, membranaceous or coriaceous; flowers large, usually solitary and
axillary, rarely terminal or corymbose, white or yellow; hypanthium ovoid or
obconic; calyx tubular or spathaceous, lobed or parted, usually persistent; corolla
salverform, campanulate, or funnelform, the tube much exceeding the calyx, the
throat usually glabrous, the limb 5-9-lobate, the lobes spreading or recurved, short or
elongate, contorted in bud; stamens 5-9, inserted in the corolla throat, the filaments
very short or none; anthers sessile or subsessile, dorsifixed, linear-oblong, included or
short-exserted; ovary 1-celled or rarely falsely 2-celled, the style stout, clavate or
DEMPSTER: FLORA OF GUATEMALA 79
fusiform at the apex; ovules numerous, horizontal, the placentae parietal; fruit
oblong to ovoid or pyriform, terete or costate, coriaceous or fleshy and rupturing
irregularly, or the endocarp 2-5-valvate and indurate; seeds very numerous, imbedded
in pulp, horizontal, compressed or turgid, obtusely angulate.
A genus of 70 or more species, confined to the Old World
tropics.
Gardenia augusta (L.) Merr. Interpr. Herb. Amboin. 485.
1917. Varneria augusta L. Amoen. Acad. 4: 136. 1759. G.
jasminoides Ellis, Phil. Trans. 51, pt. 2: 935. 1761. G. florida L. Sp.
PL ed. 2. 305. 1762. Jazmin de cabo; jazrmn.
Cultivated commonly for ornament at low and middle
elevations. Native of southern China, grown for ornament in most
tropical regions, and in greenhouses in temperate countries.
A stout, densely branched shrub, usually 1-2 m. high, the branches scabrous-
puberulent; stipules thin, brown, 1 cm. long or less; leaves coriaceous, short-petiolate,
the blades obovate or oblong-obovate, mostly 5-7 cm. long, obtuse or acute, narrowed
to the base, almost glabrous; flowers white, large and showy, very fragrant, double in
cultivated forms; calyx lobes foliaceous, triangular-lanceolate, 2-2.5 cm. long.
The gardenia, a favorite flower in the florists' shops of the
United States, is also highly esteemed in Central America, where it
adorns the gardens of rich and poor. Gardenia perfume is well
known, but, although originally obtained directly from the flowers,
is probably now a synthetic product.
GENIPA Linnaeus
Unarmed, glabrous or pubescent trees, the branchlets terete; stipules
intrapetiolar, elongate, deciduous; leaves opposite, usually large, petiolate or
subsessile; inflorescence few-flowered, axillary or terminal cymes; flowers rather
large, white or yellowish, pedicellate, hypanthium turbinate or campanulate, the
calyx ampliate, truncate or 5-6-lobate, persistent; corolla salverform, the tube short
or elongate, the throat pilose or glabrous, the limb 5-6-lobate, the spreading lobes
obtuse or acute, contorted in bud, coriaceous; stamens 5-6, inserted at the top of the
corolla tube, the anthers sessile, dorsifixed, linear, exserted; ovary 1-2-celled, the style
thick, the stigma fusiform; ovules numerous, the placentae parietal; fruit baccate,
subglobose, large, with thick pericarp; seeds numerous, large, compressed, with
subfibrous testa.
A small genus of tropical America with several species in South
America and four others recorded from Central America.
Corolla glabrous outside; leaves glabrous; stipules rounded at the apex.
G. vulcanicola.
Corolla densely sericeous outside; leaves softly pubescent beneath; stipules
acuminate.... G. caruto.
80 FIELDIANA: BOTANY, VOLUME 24
Genipa caruto HBK. Nov. Gen. & Sp. 3: 407. 1820 G.
americana var. caruto Schum. in Mart. Fl. Bras. 6, pt. 6: 352. 1889.
Irayol; jagua; guall (Quezaltenango); arayol (corruption of irayol);
tihe-dientes (fide Aguilar). Figure 30.
Common on the Pacific plains and in the lower hills of eastern
Guatemala, ascending to about 900 m.; Chiquimula; Jutiapa; Santa
Rosa; Quezaltenango; Huehuetenango; doubtless in several other
departments. Southern Mexico; Honduras and El Salvador to
Panama; northern South America.
A large tree, often 14 m. high, with thick trunk and broad spreading crown, the
thick branchlets densely short-pilose or glabrate; stipules triangular, 1-2.5 cm. long,
long-acuminate; petioles very thick, 1 cm. long or shorter, the leaf blades obovate to
oblong, 15-35 cm. long, 6-19 cm. broad, acute or obtusely short-acuminate, attenuate
to an acute base, herbaceous, lustrous above and glabrous or nearly so, densely short-
pilose beneath; cymes short-pedunculate, lax or dense, 4-10 cm. long, the pedicels 4-10
mm. long; hypanthium and calyx densely short-pilose, the hypanthium 4-8 mm. long,
the calyx 5-8 mm. long, truncate or shallowly crenate, sericeous within; corolla
yellowish white, 2-4.5 cm. long, the tube glabrous near the base, sericeous above, the
5-6 lobes oblong, obtuse, longer than the tube, the throat villous; anthers 1.5-2.5 cm.
long; fruit subglobose, 6-7 mm. in diameter, smooth or sparsely tuberculate, short-
pilose or glabrate; seeds irregular, 6-12 mm. long, dark brown.
Called "guaitil" in Costa Rica, but the usual names elsewhere
in Central America are "jagua" or "irayol." In Chiapas the tree is
said to be called "mamuc." The name "jagua" is believed to be of
Antillean origin, and the specific name caruto is Venezuelan. G.
caruto often is considered synonymous with G. americana, or a
variety of it. G. americana, which in continental North America
extends northward through Panama to Costa Rica, is glabrous
almost throughout while G. caruto has abundant pubescence. The
ranges of the two forms overlap in some regions but ordinarily the
two are so distinct that it is at least convenient to treat G. caruto
as a distinct species. The tree is a well known one in Central
America. Its wood, valued for construction purposes, is strong,
resistant, and flexible, in its properties somewhat resembling
hickory (Carya) of the United States. The pulp of the fruit is edible,
but it is little eaten except by children and domestic animals as it is
dark and repulsive in appearance and not particularly pleasant in
flavor. Its dark juice leaves an indelible stain upon any object that
it touches. This juice, producing a dark blue or almost black color,
was formerly used by many of the American aborigines for painting
their bodies. In El Salvador it was much used as the source of a
blue dye that is highly prized for coloring rebozos and other
textiles. In Honduras the well-ripened fruit is the basis of an
DEMPSTER: FLORA OF GUATEMALA 81
agreeably flavored "wine" which is taken by invalids as a stimulant
to the appetite and to digestion. It is claimed that if the fruit is
eaten in sufficient quantity it expels intestinal parasites. The
expressed juice of the young leaves is one of the numerous remedies
for fevers employed by the country people of Honduras and other
parts of Central America.
Genipa caruto is one of the common and conspicuous trees of
the Pacific coast. It sheds its leaves in the middle of the dry season,
but the trees are easily recognizable then because of their abundant
large fruits, unlike those of any associated tree. The vernacular
names appear as names for at least two Guatemalan settlements,
the caserio of Irayol in Jutiapa and the aldea of Jagua in Chiquim-
ula. The fruits are sometimes on sale in the Quezaltenango market,
under the name "guali." It is said that some of the Indians make it
a practice to carry them in their hands under certain conditions, to
ward off disease and other evils.
Genipa vulcanicola Standl. Field Mus. Bot. 17: 213. 1937.
Guayabillo cimarrdn (San Marcos); cola de pavo; tinajo (fide
Aguilar).
Damp or wet, mixed, mountain forest, central and western
departments, 1,300-2,100 m.; Guatemala; Suchitepe"quez (type from
Volcdn de Atitldn, Skutch 1512); Solold; Huehuetenango;
Quezaltenango; San Marcos. Mexico (Chiapas).
A tree 15 m. high or less, the trunk about 30 cm. in diameter, glabrous almost
throughout; stipules at first conspicuous but soon deciduous, oblong, rounded at the
apex, 5 cm. long or less; petioles 1-2 cm. long, the blades elliptic to elliptic- oblong,
mostly 10-23 cm. long and 4.5-11 cm. broad, acute or obtuse, acuminate to rounded at
the base, barbate beneath in the nerve axils, the lateral nerves about 7 on each side;
inflorescence sessile, about 4 cm. long and broad, few-flowered, the branches thick,
sparsely hirtellous, the pedicels 1 cm. long or less; calyx broadly campanulate, 1 cm.
long and broad, glabrous, truncate and remotely denticulate; corolla white,
subcoriaceous, glabrous outside, the tube 1 cm. long, 4 mm. thick, the lobes rounded-
obovate, broadly rounded at the apex, sparsely ciliate, glabrous within; anthers half
exserted; fruit globose, 3 cm. in diameter or larger, crowned by the persistent calyx.
Recent collections show that the tree is rather common,
especially in the coastal plain of Quezaltenango and San Marcos.
Flowering specimens were collected in October, but during the dry
season neither flowers nor fruits have been observed.
GEOPHILA D. Don
Reference: Louis O. Williams, Geophila in North America,
Phytologia 26: 263. 1973.
82 FIELDIANA: BOTANY, VOLUME 24
Creeping perennial herbs, rooting at the nodes; stipules small, persistent and
becoming somewhat indurate, shallowly bilobate; leaves small, long-petiolate, ovate-
cordate or rounded-cordate, herbaceous, inconspicuously nerved; flowers very small,
in terminal pedunculate few-flowered heads, the head subtended by 2 small free
bracts; calyx dentate or lobate, persistent; corolla tubular-funnelform, white, pilose
in the throat, the limb 4-7-lobate, the lobes spreading or recurved, valvate in bud;
stamens 4-7, inserted in the corolla tube, the filaments filiform, the anthers dorsi fixed,
linear, half exserted; ovary 2-celled, the style slender, with 2 short branches; ovules
solitary, erect from the base of the cell; fruit drupaceous, juicy, containing 2 one-
seeded nutlets, these costate dorsally and slightly twisted.
About 20 species in tropical Asia, Africa, and America. One
other species in known from Nicaragua and Panama. Several occur
in South America.
Fruits black when mature; inflorescence originating from the primary stem or from a
very short usually leafless secondary stem; pyrenes not ridged G. macropoda.
Fruits red or orange when mature; inflorescence usually from an elongated leaf-
bearing secondary stem; pyrenes ridged.
Ovary, fruit and leaves pubescent G. cordifotia.
Ovary and fruits glabrous, leaves usually so G. repens.
Geophila cordifolia Miquel, Stirp. Surin. Sel. 176. 1850.
Mapouria trichogyne Muell.-Arg., Mart. Fl. Bras. 6(5): 426. 1881.
Geophila trichogyne Standl. Field Mus. Bot. 7: 423. 1931.
Izabal, near sea level, in wet forest. British Honduras (Temash
River); Costa Rica; Colombia, Guianas to Venezuela, Amazonian
Brazil, and eastern Peru.
Creeping and forming dense colonies, villous-hirsute throughout with long slender
spreading hairs; leaves long-petiolate, the blades oblong-cordate to broadly ovate-
cordate, 3-11 cm. long, 2.5-4.5 cm. broad, acute or acutish, deeply and narrowly
cordate at the base, conspicuously paler beneath; peduncles usually shorter than the
petioles, the dense heads 5-8-flowered, the linear bracts rather conspicuous; calyx
lobes lance-linear, in fruit 4-5 mm. long; corolla white; fruit ovoid, about 4 mm. long,
densely villous, red or orange-red, probably red at maturity.
Geophila macropoda (Ruiz & Pavon) DC. Prodr. 4: 537. 1830.
Psychotria macropoda Ruiz & Pavon, Fl. Peruv. 2: 63, t 211, fig. b.
1799.
Wet thickets and forests, weed in banana plantations, at 600 m.
or usually less; PetǤn; Izabal; Suchitepe'quez; Escuintla. Mexico
(Vera Cruz, Oaxaca; Puebla; Chiapas); El Salvador; Honduras;
Costa Rica; Panama; Venezuela and Colombia south in the
lowlands to Bolivia, Brazil, and Argentina (Misiones).
Small colonial repent or creeping herbs up to perhaps 1 m. long; the main stems
rooting at the nodes, the internodes 5-12 cm. long, with short erect lateral branches
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 83
and erect leaves, the lateral branches almost none to about 2 cm. long, usually
leafless but bearing a long pedunculate inflorescence at the apex; leaves 2 at the
nodes, or sometimes only one, petioles slender, 3-14 cm. long, pubescent above near
the leaf-blade, .glabrescent below, leaf blades ovate-cordate to cordate, the apex
obtuse or rounded, glabrous, with 4 secondary nerves (and the main nerve) from apex
of petiole and 4 more above, (3-)5-9 cm. long and nearly as broad; inflorescence a
long-pedunculate few-flowered head arising from a short secondary stem or from the
primary stem, peduncle when mature about as long as the petioles, head subtended
by 2 or 4 short narrowly triangular bracts; calyx about 2-3 mm. long, the lobes linear
to linear-oblong, acute or obtuse, prominently nerved, about 1-2 mm. long; corolla
white, early deciduous; fruits fleshy, shining black.
Commonly identified with G. repens but easily distinguished by
the black, not red, fruits, the inflorescence borne from the primary
stem or a short usually leafless secondary stem, not from an
elongated secondary stem. The cordate base of the leaf usually has
somewhat divaricate basal lobes not closed or sometimes
overlapping ones.
Geophila repens (L.) I. M. Johnston, Sargentia 8: 281. 1949;
Rondeletia repens L. Syst. 928. 1759. Psychotria herbacea Jacq.
Enum. PL Carib. 16. 1760. Geophila reniformis D. Don, Prodr. Fl.
Nep. 136. 1825. G. herbacea Schum. in Engler & Prantl, Pflanzenf.
IV. 4: 119. 1891. Geocardia herbacea Standl. Contr. U. S. Nat.
Herb. 17: 445. 1914.
Wet thickets and forests of the hot lowlands of both coasts at
700 m. or less, often a weed in banana plantations; Pete"n; Alta
Verapaz; Izabal; Santa Rosa; Escuintla; Suchitep6quez. Southern
Mexico to Panama, and through most of tropical South America.
Plants creeping and often forming dense close colonies in shaded places; stipules
about 1.5 mm. long; petioles slender, mostly 6-12 cm. long; leaf blades rounded-ovate
to subreniform, mostly 3-5.5 cm. long, deeply cordate at the base, almost glabrous but
often slightly puberulent; peduncles long and slender, the heads 3-5-flowered; calyx
lobes green, linear- lanceolate, acuminate; corolla white or sometimes purplish, often 1
cm. long; fruit becoming red, about 5 mm. long, with scant pulp.
Sometimes called "hierba de culebra" in Honduras.
GONZALAGUNIA West
Shrubs or small trees, more or less pubescent, with slender terete branches;
stipules small, interpetiolar; leaves opposite, petiolate or subsessile, mostly
herbaceous; flowers small, sessile or pedicellate, white or pink, arranged in slender
elongate terminal spikes or thyrsiform panicles; hypanthium globose or campanulate;
calyx normally 4-lobate, the lobes small, often green, equal or unequal, persistent;
corolla funnelform or salverform, the tube short or elongate, the throat ampliate or
contracted, villous, the limb 4-lobate, the lobes short, spreading, obtuse, valvate or
84 FIELDIANA: BOTANY, VOLUME 24
imbricate in bud; stamens 4, inserted in the corolla tube, the filaments short; anthers
dorsifixed, linear-oblong, included; ovary 2- or 4-celled, the style filiform, the small
stigma obtuse or 2- or 4-lobate, included or exserted; ovules numerous, the placentae
peltately affixed to the septum; fruit baccate, usually somewhat depressed-globose, 2-
4-sulcate, 2-4-celled, many-seeded; seeds minute, subglobose, the testa deeply
foveolate.
About 15 species in tropical America, 11 known from Central
America. In general appearance the plants simulate certain species
of Rondeletia (which have capsular fruits), and if fruits are not
present, often it is difficult to tell which genus is represented by a
flowering specimen.
Young branches appressed-pilose; corolla tube about 1 mm. thick in the throat.
G. panamensis.
Young branches pilose or hirsute with long spreading hairs; corolla tube 2-3 mm.
thick in the throat.
Leaves densely covered beneath with long and very slender, matted hairs.
G. chiapasensis.
Leaves glabrate beneath except on the veins, there puberulent or pilosulous with
very short hairs.
Petioles 5-8 mm. long, hirsute; corolla sparsely appressed-pilose outside, the lobes
almost half as long as the tube G. thyrsoidea.
Petioles 10-20 mm. long, glabrate; corolla sparsely and minutely strigillose or
almost glabrous, the lobes much less than half as long as the tube..G. rojasii.
Gonzalagunia chiapasensis (Standl.) Standl. & Steyerm.
Field Mus. Bot. 22: 283. 1940. Rondeletia chiapasensis Standl. Field
Mus. Bot 22: 54. 1940.
Mountain forests or second-growth thickets of the western
highlands, 1,600-2,500 m.; Quezaltenango; San Marcos. Mexico
(Chiapas, the type from Cascarada).
A shrub or small tree 4.5-6 m. high, the branches densely pilose with long soft
spreading hairs; stipules subulate-attenuate from a triangular base, 6-7 mm. long;
leaves on petioles 5-15 mm. long, narrowly oblong-lanceolate, 10-23 cm. long, 3-6 cm.
broad, narrowly attenuate-acuminate, acute at the base, densely pilose above with
long or short, spreading or somewhat appressed hairs, very densely pilose beneath
with very long and slender, matted and interlaced hairs, the lateral nerves about 10
on each side; inflorescence spiciform, 7.5-23 cm. long, about 2 cm. broad, pedunculate,
erect or recurved, the stout rachis densely short-pilose, the flowers in dense sessile
cymules; hypanthium and calyx densely spreading-pilose, the calyx lobes lance-
oblong, erect, acute; corolla rose outside, white within, densely strigose outside, the
tube about 10 mm. long, dilated upward and 2 mm. broad or more, the rounded lobes
3 mm. long; fruit white, tinged with rose at the apex, globose, 6 mm. in diameter,
apparently very fleshy.
Gonzalagunia panamensis (Cav.) Schum. in Martius, Fl.
Bras. 6, pt. 6: 292. 1889. Buena panamensis Cav. Anal. Hist. Nat. 2:
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 85
279. 1800. Gonzalea panamensis Spreng. Syst. Veg. 1: 417. 1825.
Duggena panamensis Standl. Contr. U. S. Nat. Herb. 18: 126. 1916.
Warmer regions and plains of both coasts, usually at 900 m. or
less, but ascending on the slopes of Santa Maria and Tajumulco to
about 1,500 m., mostly in damp thickets or moist hillside forest,
sometimes on dry slopes. Pet6n; Alta Verapaz (PancajchS); Baja
Verapaz (Panzal); Izabal; Escuintla; Santa Rosa; San Marcos.
British Honduras. Southern Mexico to Panama (type from Cerro de
Anc6n) and Colombia; Cuba.
A slender shrub 3 m. high or less, the branches brownish, densely sericeous or
strigose when young; stipules triangular, subulate-acuminate, 3-8 mm. long; petioles
mostly 1-2 cm. long, often shorter, the blades ovate to lanceolate, 7-14 cm. long, 2-6
cm. broad, gradually or abruptly acuminate or long-acuminate, acute or obtuse at
the base, green above, densely or sparsely strigillose or glabrate, sparsely or densely
appressed-pilose beneath or glabrate, the lateral nerves 5-9 on each side; inflorescence
spiciform, 8-20 cm. long, short-pedunculate, the cymules sessile or subsessile, the
pedicels 2 mm. long or shorter; calyx and hypanthium 1.5-2 mm. long, strigillose or
short-pilose, the calyx lobes deltoid, obtuse or acute; corolla white, 10-17 mm. long,
the very slender tube glabrous or sparsely appressed-pilose, the lobes 2-2.5 mm. long,
sparsely tomentose within; stamens included; fruit black at maturity, juicy, globose,
4-coccous (as in the other Guatemalan species), 3-4 mm. in diameter; seeds dark
reddish brown.
Gonzalagunia rojasii Standl. Field Mus. Bot. 22: 114. 1940.
Chichipln; cachito de venado (Quezaltenango); casahuach
(Patzulin); rugey-risis (fide Aguilar).
Moist or wet, mixed, mountain forest, 1,200-1,500 m.; endemic;
Escuintla; Guatemala; Quiche"; Quezaltenango (type collected
below El Muro, below Santa Maria de Jesus, Standley 67145).
A shrub or small tree 3-9 m. high, the branchlets stout, at first densely pilose
with very long, soft, spreading, pale hairs; stipules 8 mm. long, acuminate from a
triangular base; petioles 1-2 cm. long, hispid, the blades elliptic or obovate-elliptic to
oblong- lanceolate, 11-20 cm. long, 5-9 cm. broad, acute or short-acuminate, acute at
the base or gradually narrowed and decurrent, green above, sparsely hispidulous or
glabrate, beneath appressed-pilosulous with short hairs on the veins and nerves, the
lateral nerves about 9 on each side; inflorescence spikelike, pedunculate, 15-20 cm.
long, the cymules sessile, the rachis stout, hispid; hypanthium rounded at the base,
1.5 mm. long, almost glabrous; calyx 1 mm. high, remotely 4-dentate, glabrous;
corolla pale greenish yellow or pinkish, sparsely and very minutely strigillose outside
or almost glabrous, the slender tube 6-14 mm. long, the lobes ovate-rounded, 2.5 mm.
long; fruit 4-celled, depressed-globose, 4-5 mm. broad, pink or red.
It is now somewhat questionable whether this is distinct from
G. thyrsoidea. The isolated occurrence of G. thyrsoidea would lead
one to suppose that two distinct species are involved.
86 FIELDIANA: BOTANY, VOLUME 24
Gonzalagunia thyrsoidea (Donn.-Sm.) Robinson, Proc. Am.
Acad. 45: 405. 1910. Gonzalea thyrsoidea Donn.-Sm. Bot. Gaz. 13:
188. 1888. Duggena thyrsoidea Standl. Contr. U. S. Nat. Herb. 18:
126. 1916. Figure 23.
Moist or wet, mixed, mountain forest or thickets, 800-1,500 m.;
endemic; Alta Verapaz (type from Pansamala, Tuerckheim 1249);
Huehuetenango.
A shrub or small tree of 3-6 m., the branches stout, hirsute; stipules deltoid-
acuminate, 6-7 mm. long; petioles stout, 5-8 mm. long, hirsute, the blades elliptic or
elliptic-oblong, 20-23 cm. long, acuminate at each end, lustrous above and sparsely
pilose, pilose beneath along the veins; inflorescence about 30 cm. long, the flowers
pedicellate, in 3-7-flowered subsessile cymules; calyx lobes triangular, unequal;
corolla sparsely appressed-pilose outside, white tinged with pink, the tube 14 mm.
long, 3 mm. broad in the throat, the lobes nearly half as long, farinaceous and
arachnoid within; anthers subexserted; ovary 4-celled; fruit depressed-globose, about
8 mm. in diameter, red.
GUETTARDA Linnaeus
Unarmed trees or shrubs; stipules intrapetiolar, deciduous; leaves opposite or
ternate, petiolate or subsessile, membranaceous to rigid-coriaceous; flowers rather
large or small, bracteolate or ebracteolate, arranged in axillary, bifurcate or
congested cymes or sometimes solitary, usually secund and perfect; hypanthium
ovoid or globose; calyx tubular, cupular, or campanulate, truncate or very obscurely
2-9-dentate, deciduous; corolla funnelform or salverform, usually white or pink, the
tube elongate, with naked throat, the limb 4-9-lobate, the obtuse lobes imbricate in
bud, 2 of them exterior; stamens 4-9, inserted in the corolla tube, the filaments very
short or none, the anthers linear, dorsifixed, included; ovary commonly 2-9-celled, the
cells tubular, elongate, the style filiform, the stigma capitate or shallowly bilobate;
ovules solitary, pendulous from the apex of the cell; fruit drupaceous, globose to
ovoid or oblong, sometimes acutely angulate, the flesh very thin, the stone ligneous
or osseous, 2-9-celled, the cells straight or recurved; seeds pendulous.
The genus Guettarda is one of the larger genera of the
Rubiaceae and one which has received no critical study since
Standley's account of it for the North American Flora (32: 228-262.
1934). Some 30 species have been described from Mexico, Central
America, and Panama, or credited to the region. The total from all
North America is about 100 species. The genus is also found in
South America.
Fruit acutely 4-angulate; young branches hirsute with long ascending hairs.
G. cobanensis.
Fruit terete; young branches not long-hirsute.
Tube of the corolla pilose with appressed hairs that are directed downward; leaves
mostly 7-14 cm. broad G. combsii.
Tube of the corolla pilose with hairs that are directed upward; leaves much
smaller.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 87
Pubescence of the lower leaf surfaces of spreading hairs.
Leaves acute or subacute G. gaumeri.
Leaves rounded or very obtuse at the apex G. deamii.
Pubescence of the lower leaf surface of very closely appressed hairs.
Corolla tube 6-9 mm. long; fruit 4-8 mm. in diameter G. elliptica.
Corolla tube about 12 mm. long; fruit 10-15 mm. in diameter.
G. macrosperma.
also G. petenensis and G. tikalana.
Guettarda cobanensis Donn.-Sm. Bot. Gaz. 47: 255. 1909.
Alta Verapaz (type collected in mountain forest near Coban,
1,500 m., Tuerckheim 11.2096); collected also at Chicoj northeast of
Carcha, 1,200 m., in wet thicket; and near Tactic; endemic.
A shrub 2.5 m. high, or perhaps becoming larger, the branchlets obtusely
tetragonous, densely fulvous-pilose with long ascending hairs; stipules ovate, 1-1.5 cm.
long, attenuate, thin, brown, fulvous-pilose near the base and along the costa; leaves
mostly ternate, the slender petioles 2-4.5 cm. long, hirsute; leaf blades elliptic or
elliptic-ovate, 9-21 cm. long, 3-10 cm. broad, acute or acuminate at base and apex,
green above and short-pilose, especially along the nerves, aporessed-pilose beneath,
the lateral nerves about 11 on each side; peduncles 1.5-2 cm. long, the cymes
bifurcate, with branches 1-1.5 cm. long and 3-4-flowered; flowers 4-5-parted, the
bractlets minute; calyx and hypanthium 2 mm. long, puberulent, the calyx short,
shallowly dentate; corolla tube 22 mm. long, retrorse-pilose outside, the lobes
lacerate- crispate, one-third as long as the tube; anthers 4 mm. long; fruit acutely
quadrangular, 9 mm. long, 5-7 mm. thick, 3-4-celled, the cells straight.
Apparently a rare plant, since recently it has been collected but
twice.
Guettarda combsii Urban, Symb. An till. 6: 48. 1909. G. scabra
var. seleriana Loes. Repert. Sp. Nov. 18: 361. 1922. G. seleriana
Stand!. Contr. U. S. Nat. Herb. 23: 1384. 1926. Texpac, textop
(Peten, Maya, fide Lundell).
Dense or open, wet or dry forest, 900 m. or lower, usually on
limestone; often at edges of clearings, sometimes in second growth;
Peten; Alta Verapaz; Izabal. Mexico (Chiapas, Tabasco, and
Yucatan); British Honduras; Honduras; Nicaragua; Cuba.
A tree, said to attain sometimes a height of 18 m. but usually much smaller and
often only a shrub, the trunk reported to attain a diameter of 60 cm., the young
branchlets terete, ferruginous-pilosulous with ascending or spreading hairs; stipules
mostly oblong or lanceolate and 8-14 mm. long, filiform-acuminate, pilose outside;
leaves opposite, on slender petioles 1-6 cm. long, very variable in shape, rounded-oval
to oval, rounded-ovate, or broadly oval-obovate, 7-20 cm. long, 5-13 cm. broad,
rounded and very abruptly short-acuminate at the apex or obtuse or acutish, rounded
to deeply and narrowly cordate at the base, chartaceous or subcoriaceous, green
88 FIELDIANA: BOTANY, VOLUME 24
above, glabrous or pilosulous on the veins, paler beneath or sometimes green, usually
densely and minutely pilosulous with mostly somewhat spreading hairs, the lateral
nerves 8-11 on each side; cymes once or twice bifid, on slender peduncles 5-15 mm.
long, the branches 1-5 cm. long, bearing 5 to many flowers; bractlets linear, 3-5 mm.
long; corolla white or creamy white, the tube 16-18 mm. long, densely retrorse-pilose,
the 6-7 lobes one-fourth to one-third as long as the tube; ovary 4-5-celled; fruit
globose, 6-8 mm. in diameter, densely and minutely tomentose.
The Maya name "xtez-tab" is reported from Yucatan. In
British Honduras the tree is called "glassy wood."
Guettarda deamii Standl. Contr. U. S. Nat. Herb. 20: 209.
1919. Fruta de mico (fide Aguilar).
Dry thickets, mountain sides or along arroyos, at about 500 m.
or less or apparently ascending to about 1,500 m. in central
Guatemala; Zacapa (type from Gualan, Deam 627 7); Chiquimula
(Quebrada Shusho, above Chiquimula); Guatemala (probably
vicinity of La Aurora, or perhaps at a lower elevation). Honduras.
A shrub or small tree 3-5 m. high, the older branches dark and lenticellate, the
young branchlets densely short-pilose; stipules ovate-oblong, 3-4 mm. long, obtuse or
subacute; leaves opposite, on petioles 5-9 mm. long, mostly oval, sometimes oval-
oblong or obovate-oval, 4-9 cm. long, 2.5-6.5 cm. broad, usually broadly rounded at
the apex, varying to subacute, rounded at the base, densely short-pilose or pilose-
scaberulous above, densely velutinous- pilosulous beneath, the lateral nerves 8-10 on
each side; cymes subcapitate, 3-5-flowered, the very stout peduncles 3-10 mm. long,
the bractlets subulate, 3-4 mm. long; hypanthium and calyx densely appressed-pilose,
the calyx truncate; corolla white, the tube densely antrorse-pilose, about 7 mm. long,
the broad lobes half as long as the tube; fruit globose, 8 mm. in diameter, 3-4-celled,
densely and minutely tomentulose.
Guettarda elliptica Swartz, Prodr. Veg. Ind. Occ. 59. 1788.
In forest or thickets, near sea level; Alta Verapaz; Peten (La
Libertad, Lundell 3329}. Northern British Honduras and in
Tabasco; southern Florida; West Indies; southern Mexico;
Venezuela.
A shrub or small tree, sometimes 8 m. high but usually lower, reported as
sometimes more or less scandent, the slender branches terete, often with conspicuous
white lenticels, appressed-pilose, leafy at the ends; stipules deltoid-acuminate, 3 mm.
long; leaves opposite, on stout petioles 12 mm. long or less, usually oval or rounded-
oval, mostly 3-7.5 cm. long and 1-4.5 cm. broad, commonly rounded or obtuse and
apiculate at the apex, sometimes acute or short-acuminate, truncate to acute at the
base, membranaceous or chartaceous, green above, sparsely and minutely appressed-
pilose or glabrate, minutely appressed-pilose beneath or glabrate, the lateral nerves 5-
7 on each side; cymes axillary, dense, 1-9-flowered, usually 3-flowered, the peduncles
0.5-3 cm. long; bractlets minute or sometimes equaling the calyx; calyx and
hypanthium minutely tomentulose and often short-pilose; calyx 2-2.5 mm. long,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 89
truncate; corolla white, tinged with pink outside, the tube antrorse-sericeous, the
slender tube 6-9 mm. long, the 4 lobes rounded-obovate, 2-2.5 mm. long; fruit globose,
4-8 mm. in diameter, purplish when mature, 2-4-celled.
Called "prickle wood" in British Honduras, and the Maya name
is reported as "kiikche."
Guettarda gaumeri Standl. Field Mus. Bot. 8: 58. 1930. Zedi
(Peten, fide Lundell).
Moist forest or thickets, 200 m. or lower; Peten; Izabal. Mexico
(Yucatan and Campeche) to northern British Honduras, the type
from Yucatan.
A shrub about 3 m. high or sometimes a tree of 7 m., the branches terete, the
young branchlets densely short-pilose with spreading hairs; stipules broadly ovate,
obtuse, brown and sparsely sericeous within; leaves opposite, on petioles about 4 mm.
long, oblong or oblong-elliptic, 3-7.5 cm. long, 1-4 cm. broad, acute or obtuse and
apiculate, obtuse to broadly rounded or even obscurely cordate at the base, usually
densely velutinous-pilose above, densely pilose beneath with spreading and often
matted hairs, the lateral nerves about 8 on each side; cymes <"ibcapitate, mostly 3-
flowered, the stout peduncles 4-8 mm. long; bracts lanceolate or ovate, 3-4 mm. long;
hypanthium and calyx densely pilose with long ascending hairs, the calyx shallowly
bilobate; corolla white, 12 mm. long, densely pilose with long ascending hairs, the
lobes 2 mm. long, glabrous within; fruit oval, 5 mm. long, densely tomentose and
sparsely long-pilose, 4-celled.
Guettarda macrosperma Donn.-Sm. Bot. Gaz. 18: 204. 1893.
Crucito. Figure 37.
Dry or moist thickets, sometimes in forest, chiefly at 900 m. or
less, but perhaps ascending to about 1,500 m.; Pete"n; Izabal;
Chiquimula (?); Santa Rosa (type from Santa Rosa at 900 m.,
Heyde & Lux 3160); Escuintla; Suchitepequez; Solola; Retalhuleu.
British Honduras to Panama.
A rather slender shrub or a small tree, sometimes 8 m. high, the trunk as much
as 10 cm. in diameter, the young branchlets fulvous-pilose with short, mostly
appressed hairs; stipules ovate-deltoid, 4 mm. long, filiform-acuminate; leaves
opposite, on petioles 0.5-2.5 cm. long, oval to oval-elliptic, rhombic-elliptic, or oblong,
5-13 cm. long, 2.5-7 cm. broad, subacute to acuminate, rounded to subacute at the
base, membranaceous or chartaceous, green above, hispidulous when young with
setiform hairs, glabrate in age, minutely appressed-pilose beneath, the lateral nerves
7-9 on each side; cymes few-flowered, at first dense, the branches in fruit as much as
2 cm. long, the peduncles 1-5 cm. long; bractlets subulate, shorter than the calyx;
calyx and hypanthium tomentulose, the calyx 2-2.5 mm. long, truncate; corolla white
or creamy white, sericeous outside, the tube 12 mm. long, the 5-6 lobes oval, one-third
as long as the tube; fruit globose, 1-1.5 cm. in diameter, often dark red, densely and
finely tomentulose, 3-4-celled.
90 FIELDIANA: BOTANY, VOLUME 24
The name "correa" is reported from Honduras; "tintero" (El
Salvador). The material of this and some of the other species now
at hand from Guatemala is scant and in poor condition. When more
and better specimens have accumulated, it is probable that a
realignment of the species will be necessary.
Guettarda petenensis Lundell, Wrightia 4: 124. 1969.
Tree in forest, in lowland Peten, Contreras 6947.
Said to be a tree to 20 m. and 60 cm. in diameter but probably much smaller, the
branchlets substrigose, becoming glabrous; leaves at the end of new growths, petioles
about 1 cm. long, the blades ovate or ovate-lanceolate, acuminate, 3-7.5 cm. long and
1.5-4.5 cm. broad, sparsely strigillose, to almost glabrous, barbellate in nerve axils
below; inflorescence short, to 5 cm. long, probably few-flowered, peduncle
prominently appressed-strigillose; fruit sessile or nearly so, said by collector to be
carmen-red, oblong-elliptic, up to 2 cm. long.
This species is maintained, for the type is inadequate but does
have smaller leaves than most G. macrosperma, of which it may be
a synonym.
Guettarda tikalana Lundell, Wrightia 2: 63. 1960.
Known only from the type, near Tikal, Peten, Lundell 16519.
Woody vines, branchlets densely hirsute; stipules acuminate, 6-7 mm. long,
deciduous; leaves membranaceous, obovate or obovate-elliptic, 8-16 cm. long, 4-8.5
cm. broad, acute or short-acuminate, the base narrowed, rounded or obtuse, densely
pilose on the lower surface, pubescence on the upper surface primarily along costa
and veins, lateral nerves 8-12 pairs; petioles hirsute, 7-12 mm. long; inflorescence of
axillary or terminal cymes, hirsute, the branches as much as 2.5 cm. long; fruits
sessile, finely tomentulose, conic or obovoid-conic, when immature up to 2.8 cm. long
and 1.5 cm. in diameter.
We have seen specimens of this species which was described
from fruiting material. The scandent habit is unusual in the genus.
However, it cannot be separated in the key from G. macrosperma
by the characters available.
HAMELIA Jacquin
Shrubs or trees, glabrous or pubescent; stipules interpetiolar, deciduous; leaves
opposite or often verticillate, petiolate, herbaceous; flowers yellow or red, small or
rather large, arranged in mostly terminal and scorpioid cymes, sessile or pedicellate,
the bracts minute; hypanthium ovoid or turbinate; calyx 5-lobate, the lobes short or
elongate, persistent; corolla tubular or funnelform, the tube 5-costate, constricted at
the base, the throat glabrous, the limb 5-lobate, the lobes short, imbricate in bud, 1 or
2 of them exterior; stamens 5, inserted at the base of the corolla tube, the filaments
short; anthers linear, basifixed, included or subexserted, the connective appendaged
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 91
at the apex; ovary 5-celled, the style filiform, the stigma fusiform; ovules numerous,
the placentae affixed to the axis of the ovary; fruit small, baccate, ovoid to cylindric,
5-celled; seeds numerous, minute, angulate, the testa membranaceous, foveolate.
A genus of about 30 species in tropical America. Several others
are known from Central America, and a few additional ones occur
in Mexico.
Calyx lobes oblong to subulate, conspicuously longer than broad.
Corolla 1-1.5 cm. long, yellow, glabrous; leaves opposite H. axiUaris.
Corolla 1.8-2.5 cm. long; glabrous or villous, yellow or red; leaves mostly
verticillate.
Corolla villous; leaves villous beneath H. rovirosae.
Corolla glabrous or very minutely puberulent; leaves glabrate, merelv puberulent
beneath on the veins H. calycosa,
Calyx lobes deltoid or broadly deltoid, as broad as long.
Leaves opposite, glabrous or nearly so.
Leaves barbate beneath in the axils of the nerves H. barbata.
Leaves not barbate beneath H. longipes.
Leaves in whorls of 3-4, conspicuously pubescent and often densely so.
Plant conspicuously pubescent and often densely so H. patens.
Plant glabrous or glabrescent H. patens var. glabra.
Hamelia axillaris Swartz, Prodr. Veg. Ind. Occ. 46. 1788. H.
lutea Rohr ex Smith in Rees, Cycl. 17: no. 4. 1811. Chichipin
amarillo (fide Aguilar).
Moist or wet, usually mixed forest, 1,500 m. or lower;
Suchitepe"quez. Mexico; British Honduras, along the Atlantic slope
to Panama; southward to Peru and Brazil; West Indies.
A low weak shrub or a small tree, rarely 6-9 m. high, the branches glabrous or
when young minutely and inconspicuously puberulent; stipules deltoid, acuminate or
cuspidate, 2-6 mm. long; leaves opposite, on slender petioles 0.6-4 cm. long, elliptic-
oval to obovate-oval or oblong-elliptic, 5-18 cm. long, 2-7 cm. broad, cuspidate-
acuminate or long-acuminate, acute or abruptly short-acuminate at the base,
glabrous above, minutely puberulent beneath on the veins or glabrous; inflorescence
terminal, few- or many-flowered, lax, pedunculate, the puberulent branches short or
elongate, the flowers sessile or subsessile, secund; calyx and hypanthium 3 mm. long,
minutely puberulent or glabrate, the calyx lobes oblong or linear-oblong, obtuse,
minute, in fruit recurved and sometimes 1.5 mm. long; corolla yellow, 1-1.5 cm. long,
glabrous or nearly so, constricted at the base, tubular-campanulate above and 3-4
mm. broad, the rounded lobes 1-1.5 mm. long; fruit globose-ellipsoid, 5-6 mm. long.
This shrub grows most often in deep forest. The inflorescences
usually are few and not at all conspicuous. Rare in Guatemala.
Hamelia barbata Standl. Field Mus. Bot. 22: 115. 1940.
Chichipin. Figure 34.
92 FIELDIANA: BOTANY, VOLUME 24
Chiefly in wet mixed forest, 500-1,500 m.; Escuintla (type from
Finca Monterrey, slopes of Volc&n de Fuego, Standley 64569);
Guatemala; Chimaltenango; Suchitepe'quez; Quiche';
Quezaltenango; San Marcos. Mexico (Chiapas).
A shrub or small tree 3-8 m. high, the branchlets glabrous; stipules triangular-
acuminate, 3-4 mm. long; leaves opposite, on petioles 1.5-2.5 cm. long, oblong-
oblanceolate or oblong-elliptic, 14-20 cm. long, 4.5-6 cm. broad, narrowly long-
acuminate, narrowed to the acute or acuminate base, glabrous above, densely barbate
beneath in the nerve axils but otherwise glabrous; inflorescence terminal, cymose-
corymbose, short-pendunculate, laxly many- flowered, about 4.5 cm. high and 7 cm.
broad, the flowers secund, glabrous, the slender pedicels 2-3 mm. long or in fruit more
elongate; hypanthium oblong-turbinate, 3 mm. long, the calyx lobes scarcely more
than 0.5 mm. long; corolla orange, tubular or slightly dilated above and as much as 5
mm. broad, about 18 mm. long, the very broad lobes 2 mm. long.
A rather showy and handsome plant.
Hamelia caiycosa Donn.-Sm. Bot. Gaz. 12: 132. 1887. H.
chiapensis Brandegee, Univ. Calif. Publ. Bot. 6: 71. 1914 (type from
Finca Mexiquito, Chiapas).
Izabal; Alta Verapaz (the type from Pansamala, Tuerckheim
454); Huehuetenango. Mexico (Oaxaca to Chiapas) and British
Honduras; Honduras.
A shrub or tree, reported to reach a height of 12 m. in British Honduras, with
trunk diameter of 25 cm., but this probably a great exaggeration, the branchlets
glabrous or when young minutely puberulent; leaves mostly ternate, on petioles 4-12
mm. long, elliptic to oblong-oblanceolate, 6-8.5 cm. long, 1.5-3.5 cm. broad,
acuminate, acute or acuminate at the base, glabrous above, glabrous beneath or
minutely puberulent along the veins; inflorescence terminal, usually few-flowered and
lax, the. flowers not secund or obscurely so, the pedicels 7 mm. long or less; calyx and
hypanthium glabrous or sparsely puberulent, the hypanthium 3-4 mm. long, the calyx
lobes oblong or oblanceolate, 2.5-6 mm. long, foliaceous, obtuse, recurved; corolla
yellow or deep yellow, 18-22 mm. long, glabrous or minutely puberulent, constricted
at the base, narrowly campanulate above, the throat 8 mm. broad, the semiorbicular
lobes 2-2.5 mm. long; fruit oblong or cylindric, 7-12 mm. long, 4-6 mm. thick,
glabrous.
From Oaxaca the names "cihuapate," "panelilla," and "clavo"
are reported. The bark is dark brown, the inner bark pale pink. The
wood is yellow.
Hamelia longipes Standl. Proc. Biol. Soc. Wash. 37: 53. 1924.
Dense wet mixed lowland forest, 150 m. or lower; Izabal (type
from Escoba across the bay from Puerto Barrios, Standley 24829).
Mexico (?); Atlantic coast of Honduras.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 93
A shrub or small tree, glabrous throughout, 2-6 m. high; stipules deltoid-
aciminate, 3-4 mm. long; leaves opposite, on slender petioles 2.5-3.5 cm. long, oval to
oblong-elliptic, 15-25 cm. long, 7-11 cm. broad, acute or rather abruptly short-
acuminate, rounded to acute at the base; inflorescence terminal, cymose-corymbose,
on a long or short peduncle, sometimes 12 cm. long and 20 cm. broad, the rachis and
pedicels bright red, the flowers not or scarcely secund, the slender pedicels 6-13 mm.
long; calyx and hypanthium 3 mm. long, the calyx lobes almost obsolete, short and
broadly rounded; corolla pale buff or pale yellow, 16-17 mm. long, tubular, very
slightly ampliate above, the very short lobes ovate-deltoid, erect; fruit red, turning
black at maturity, oblong-ovoid, about 7 mm. long and 4.5 mm. broad.
The local names "uva de montana" and "coloradillo" are
reported from Honduras.
Hamelia patens Jacq. Enum. PI. Carib. 16. 1760. H. erecta
Jacq. I.e. Hierba del cancer (Escuintla); chichipln; cuetillo
(cohetillol); chac-ixcanan, xcanan, ixcanan, ixcanan amarillo
(Peten, Maya, fide Lundell); chamah, sicunken (Quecchi, fide
Pittier); sisipince, clavito, flor de cangrejo (reported from
Guatemala but not verified by the authors); canuto; hierba de
erisipela.
Usually in dry to wet thickets, often in second growth,
sometimes in rather open forest, 1,000 m. or lower; Peten; Alta
Verapaz; Izabal; Zacapa; Chiquimula; Santa Rosa; Escuintla;
Suchitepequez; Retalhuleu; Quezaltenango; San Marcos;
Huehuetenango; Quiche. Southern Florida; southern Mexico;
British Honduras to El Salvador and Panama; southward to Bolivia
and Paraguay; West Indies.
A shrub or small tree, commonly 1-3 m. high, the branches brown or grayish,
villous or puberulent when young; stipules 3-6 mm. long, triangular-acuminate;
leaves mostly ternate, on slender petioles 1-5 cm. long, lance-oblong to elliptic or
ovate, 6-20 cm. long, 2-9 cm. broad, usually short-acuminate, rounded to acuminate
at the base, puberulent or villosulous above, usually copiously villosulous beneath or
tomentose; inflorescence terminal, many-flowered, the branches ofter much elongate
in fruit, the flowers secund, sessile or nearly so; calyx and hypanthium 2.5-3 mm.
long, sparsely or densely pubescent or villosulous, the calyx lobes minute, deltoid;
corolla tubular, orange- red, 1.5-2 cm. long, sparsely or densely puberulent, villosulous,
or farinose-puberulent, the lobes minute, erect; fruit globose or oblong-ellipsoid, 6-10
mm. long, 4-6 mm. thick, villosulous or puberulent, red, turning almost black; seeds
brown or yellowish brown.
Known in El Salvador by the names "chichipince," "sisipince,"
"chichipinte," and "coralillo"; "coral" (Honduras); in Oaxaca called
"canutillo;" called "coloradillo" and "achiotillo Colorado" in
Honduras. The Maya names are reported as "xcanan" or "xcanal,"
"neanan," and "chactoc" (Yucatan and British Honduras).
94 FIELDIANA: BOTANY, VOLUME 24
Although a decidedly weedy shrub, often occurring in great
abundance along roads, in hedges, and in waste places, Hamelia
patens is rather handsome when in full flower and not too dusty. In
Guatemala it sometimes is planted for ornament in parks or
gardens of the highlands where it does not grow naturally, as at
Huehuetenango. The fruits are rather sweet and edible. The plant
finds some use in domestic medicine.
This is one of the first shrubs to appear after clearings have
been made in the forest but it rarely becomes abundant in such
places.
Hamelia patens Jacq. var glabra Oersted, Vidensk. Meddel.
Kjoenh. 1852: 42. 1853. H. nodosa Mart. & Gal. Bull. Acad. Brux.
11(1): 234. 1844.
In wet forests and thickets, Mexico to Panama and northern
South America, to be expected in Guatemala but not yet known in
the country.
Similar to the species but glabrate or glabrous.
Hamelia rovirosae Wernham, Journ. Bot. 49: 211. 1911. H.
patens var. coronata Donn.-Sm. Bot. Gaz. 40: 4. 1905 (type from
Cubilguitz, Alta Verapaz, 350 m., Tuerckheim 8532). H. purpuras-
cens Blake, Contr. Gray Herb. 52: 105. 1917 (type from Manatee
Lagoon, British Honduras).
Wet forest or thickets of the Atlantic lowlands, 350 m. or less;
Peten; Alta Verapaz; Izabal. Tabasco to Panama.
A shrub or small tree, 8 m. high or less, rarely subscandent, the branchlets
sparsely or densely villous; stipules deltoid, with subulate tip, 3-5 mm. long; leaves
ternate, on slender petioles 1.5 cm. long or less, elliptic, elliptic-oblong, or oval-
elliptic, 5-15 cm. long, 2-5 cm. broad, acute or short-acuminate, acute or acuminate at
the base, glabrous or sparsely villous above and minutely papillose, sparsely or
densely villous beneath along the nerves; inflorescence terminal, with few or many
flowers, short-pedunculate, lax, the branches short or elongate, the flowers sessile or
short-pedicellate, usually secund; calyx and hypanthium sparsely or densely villous
with crisped hairs, the hypanthium 3 mm. long; calyx lobes narrowly oblong, obtuse,
2-5 mm. long, reflexed; corolla 18-24 mm. long, dull or dark red, villous, cylindric
below, gradually ampliate upward, 6 mm. broad in the throat, the rounded lobes 1.5-2
mm. long; fruit ovoid, 8 mm. long, red or purple, sparsely or densely villous.
The crisped or flattened hairs of the inflorescence and new
growths along with the narrow calyx lobes facilitate the
determination of this species.
Called "coloradillo" in Honduras.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 95
HEMIDIODIA Schumann
Perennial herbs, sometimes slightly suffrutescent at the base, with often elongate
stems; stipules united with the petioles to form a setiferous sheath; leaves opposite,
herbaceous, narrow, conspicuously nerved; inflorescence a fascicle of very small
flowers clustered in the leaf axils, sessile; sepals 4, equal, connate at the base; corolla
white, funnelform, the 4 lobes valvate in bud; stamens inserted at the apex of the
corolla tube, exserted; fruit of 2 small carpels, these separating from the central
septum, opening on the inner surface near the base.
The genus consists of a single species.
Hemidiodia ocimifolia (Willd.) Schum. in Mart. Fl. Bras.
6(6): 29. 1888. Spermacoce ocimifolia Willd. ex Roem. & Schult.
Syst. Veg. 3: 530. 1818. Figure 63.
Wet thickets, open banks, hilly pine forest, 1,200 m. or lower;
Pete'n; Izabal; Chiquimula; Suchitepe'quez. Southern Mexico;
British Honduras to Panama; West Indies; South America. Malaya.
Plants perennial, decumbent or ascending, less often erect, reported to attain
sometimes a height of 1.5 m.; but usually scarcely half as tall, the stems puberulent
or almost glabrous, obtusely tetragonous or subterete, often ferruginous; leaves
petiolate, lanceolate or lance-oblong, 3-5 cm. long or larger, mostly 7-15 mm. broad,
acuminate, attenuate to the base, puberulent or glabrous, scaberulous on the
margins, paler beneath, the nerves often impressed on the upper surface; flowers in
dense axillary clusters; sepals ovate, acute, 0.5 mm. long; corolla white or tinged with
lavender, glabrous, 3-4 mm. long; fruit 3-4 mm. long, smooth, puberulent or almost
glabrous.
A weedy plant, often common in waste places or thickets along
the Atlantic coast of Central America, but barely reaching the
Pacific slope in Honduras and crossing over in Costa Rica and
Panama. Not known nor to be expected from Mexico to Panama
along the Pacific where there is a prolonged dry season.
HILLIA Jacquin
Glabrous epiphytic shrubs with thick, terete or subangulate branches; stipules
intrapetiolar, membranaceous, caducous; leaves opposite, petiolate, fleshy, becoming
coriaceous when dried; flowers large, white, terminal, solitary, subsessile, bracteate
and bracteolate; hypanthium obovoid or cylindric; calyx none or of 2-4 foliaceous
caducous lobes; corolla salverform, the tube elongate, the throat ampliate, naked, the
limb 3-7-lobate, the spreading lobes contorted in bud; stamens 4-7, inserted below the
throat of the corolla, the filaments very short; anthers basifixed, elongate-linear,
obtuse at each end, included; ovary 2-celled, the style filiform, clavate at the apex;
ovules numerous, ascending, the placentae adnate to the septum; capsule elongate-
oblong or cylindric, truncate, 2-celled, septicidally bivalvate; seeds numerous,
imbricate, the testa spongious, produced at the base into an appendage and at the
apex into a tuft of hairs.
96 FIELDIANA: BOTANY, VOLUME 24
About 20 species of tropical America, 19 being known from
Mexico to Panama.
Leaves small, 2 cm. long or less, rounded at the apex; corolla tube 1.5 cm. long.
H. chiapensis.
Leaves much larger, mostly 4-12 cm. long; corolla tube (where known) much longer.
Capsule 8-9 cm. long; leaves acute or short-acuminate H. macrocarpa.
Capsule 5-7 cm. long; leaves rounded or very obtuse at the apex H. tetrandra.
Hillia chiapensis Standl. Journ. Wash. Acad. Sci. 16: 16. 1926.
Epiphytic in wet forest, 900-1,500 m.; Alta Verapaz;
Chiquimula. Mexico (Chiapas); Costa Rica.
A small, densely branched, glabrous shrub; stipules oblong to obovate, 3-4 mm.
long, rounded at the apex, caducous; petioles stout, 2-5 mm. long; leaf blades elliptic
or oval to oblong-elliptic, usually 9-18 mm. long, 4-8 mm. broad, coriaceous, rounded
at the apex, obtuse or subacute at the base, the lateral nerves obscure; corolla tube
15 mm. long, the broad lobes 7 mm. long; capsule about 2 cm. long, the valves after
dehiscence 3-4 mm. broad.
Hillia macrocarpa Standl. & Steyerm. Field Mus. Bot. 23: 23.
1943. Jazmin.
Quezaltenango, the type collected on slopes of Volcan de Santa
Maria, between Patzulin and Finca Pirineos, 1,300-1,500 m.,
Steyermark 33667.
An epiphytic shrub with thick ochraceous branches; leaves on thick petioles 4-7
mm. long, coriaceous, elliptic-oblong or oblanceolate-oblong, 6-8.5 cm. long, 2-3.5 cm.
broad, acute or subacuminate, cuneately narrowed to the base, broadest at or slightly
above the middle, the lateral nerves ascending at a very narrow angle, obscure;
capsule short-pedicellate, narrowly cylindric, 8-9 cm. long, almost 1 cm. thick, shortly
narrowed at the apex, gradually narrowed to the base.
Hillia tetrandra Swartz, Prodr. Veg. Ind. Occ. 58. 1788.
Jazmin; jazmin de montaha. Figure 14.
On trees in wet forest, ranging from sea level to about 2,800 m.,
and widely distributed; Izabal; Pet£n; Baja Verapaz; Chiquimula
(Montana Tajuran); Suchitep6quez; Solola; Quezaltenango; San
Marcos. Southern Mexico to Panama; Cuba and Jamaica.
An epiphytic shrub 2 m. long or less, often pendent, sometimes growing on rocks,
the stout branches brown or ochraceous; stipules oval or oblong, 1-2.5 cm. long, thin,
obtuse or rounded at the apex; petioles stout, 5-25 mm. long; leaf blades obovate,
oval, or oval-obovate, 5.5-10 cm. long, 2.5-6 cm. broad, usually rounded or very
obtuse at the apex and often cuspidate-acuminate, acute or obtuse at the base, thick
and coriaceous, the lateral nerves almost obsolete; hypanthium cylindric, 3-5 mm.
long; calyx lobes 6, linear-lanceolate, obtuse, about 4 mm. long, sometimes wanting;
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 97
corolla tube 6-10 cm. long, 3-4 mm. thick, the 6 lobes lanceolate or lance-oblong, 2.5-4
cm. long; capsule 4-12 cm. long, 7-9 mm. thick, brown; seed body 2-3 mm. long, the
brown or yellowish hairs 8-12 mm. long.
HINTONIA Bullock
Shrubs or small trees; stipules interpetiolar, small, acute; leaves opposite,
petiolate, herbaceous, pubescent or glabrate; inflorescence a single axillary or
terminal flower, rarely a 3-flowered cyme; flowers large and showy, solitary and
axillary, pedicellate; hypanthium turbinate, 6-8-costate; calyx 6-lobate, the lobes
narrow, more or less foliaceous, deciduous; corolla funnelform, symmetric, the tube 6-
7 or 8-costate, the limb 6- or 8-lobate, the lobes in bud plicate- valvate, broadly
triangular; stamens inserted at the base of the corolla tube, the filaments elongate,
filiform; anthers linear, basifixed, not or hardly exserted; ovary 2-celled, the
placentae affixed to the septum; the style filiform, shallowly bifid at the apex;
capsule ellipsoid, only slightly compressed, conspicuously 6-costate and lenticellate,
septicidally dehiscent; seeds biseriate, pendulous, numerous, compressed, winged.
Four species are known in Mexico and Guatemala. It is
probable that a few species of Coutarea described from northern
South America are congeneric. H. octomera (Hemsl.) Bullock, with
8-parted rather than 6-parted (as in Guatemalan species) corollas,
occurs in Yucatan and may well extend into Peten. H. latiflora
(Sesse & Mocino ex DC.) Bullock has been found recently in
Chiapas about 100 km. from Guatemala.
Leaves narrowly lanceolate or oblong-lanceolate; corolla 9-11 cm. long; anthers 1.5
cm. long; pedicels 2-bracteolate H. lumaeana.
Leaves elliptic or ovate; corolla about 6 cm. long; anthers 3 cm. long; pedicels not
bracteolate H. standleyana.
Hintonia lumaeana (Baill.) Bullock in Hook. Icon. 33: t. 3295,
p. 5. 1935. Coutarea lumaeana Baill. Adansonia 12: 301. 1879.
Portlandia lumaeana Baill. Hist. PL 7: 333. 1880. Quina.
Moist or wet, mixed forest, 1,200-1,500 m.; Guatemala, perhaps
only in cultivation; described from a shrub grown in the garden of
Don Jorge Luma, the seeds brought from San Jorge,
Chimaltenango, a photograph of the type in herb. Field Museum;
Quezaltenango (along road between Finca Pirineos and
Calahuache"). Also in Mexico (Veracruz).
A large shrub or a tree 8 m. tall, glabrous throughout, with slender branches;
stipules deltoid, 2-4 mm. long; petioles only 3-6 mm. long; leaf blades mostly
narrowly lance-oblong, 8-11 cm. long, 1.5-3 cm. broad, acuminate or long-acuminate,
obtuse to acuminate at the base; pedicels 5-14 mm. long, bracteolate below the
middle, the bractlets small, green, subulate; hypanthium 6-7 mm. long, the calyx
lobes 6, linear-subulate, about 1.5 cm. long; flowers sweet-scented, the corolla
greenish white outside, pure white within, the tube 4.5 cm. broad at the throat, the
98 FIELDIANA: BOTANY, VOLUME 24
lobes about one- fourth as long as the tube; stamens partly exserted, slightly shorter
than the corolla; capsule oval-globose, about 1.5 cm. long, coarsely 6-costate,
inconspicuously lenticellate; seeds suborbicular, 4-5 mm. long, brown.
A handsome plant because of the large, pure white flowers.
Employed in Huehuetenango as a remedy for malaria.
Hintonia standleyana Bullock in Hook. Icon. 33: t 3295, p. 6.
1935. Coutarea latiflora DC. sec. Standl. N. Am. Fl. 32: 127. 1921,
non Sesse & Moc. ex DC. Figure 18.
Dry plains and hillsides, about 200-250 m.; Zacapa (Zacapa and
Gualan). Western and southern Mexico.
A shrub or small tree, sometimes 6 m. high, the trunk covered with rough
checkered bark, the branchlets densely short-pilose when young, later often glabrate;
stipules deltoid, 2-4 mm. long, acute or acuminate; petioles slender, 0.5-2 cm. long, the
blades ovate or oval, 4-12 cm. long, 1.5-6 cm. broad, obtuse to short-acuminate,
rounded to acute at the base, usually short-pilose above, whitish-tomentose beneath
when young, or pilose, rarely almost glabrous except along the veins; pedicels
ebracteolate, 5-15 mm. long; hypanthium densely or sparsely pilose, about 5 mm.
long; calyx lobes linear or subulate, 6-10 mm. long; flowers sweet-scented, the corolla
white, glabrous or pilose outside, the tube 3-4.5 cm. broad at the throat, the lobes half
as long as the tube or shorter; stamens slightly shorter than the corolla; capsule oval,
about 2 cm. long, brown, 6-costate, obscurely or conspicuously lenticellate; seeds 5-7
mm. long, brown.
The tree is a handsome one when covered with its abundant
showy white flowers. It appears to be rare in the Zacapa region. In
Oaxaca the tree is called "San Juan."
HOFFMANNIA Swartz
Herbs, shrubs, or small trees, glabrous or pubescent, often simple, the branchlets
usually terete, rarely tetragonous; stipules interpetiolar, deciduous or persistent,
small and inconspicuous; leaves opposite or verticillate, sometimes anisophyllous;
flowers small, white, red, or yellow, ebracteolate, generally cymose, the cymes
axillary, sessile or pedunculate, the flowers often secund; hypanthium oblong or
turbinate, terete or angulate; calyx 4-lobate, rarely truncate, the short lobes
persistent; corolla funnelform to subrotate, with short or elongate tube, the throat
glabrous, the limb 4-lobate, the lobes lanceolate to oblong, obtuse or acute, imbricate
in bud, sometimes only obscurely so, 2 of them exterior; stamens 4, inserted in the
corolla throat or lower in the tube, the filaments short, complanate; anthers linear-
oblong, dorsifixed, relatively large, obtuse or apiculate, exserted; ovary normally 2-
celled, the style filiform, the stigma bilobate or clavate; ovules numerous,
multiseriate, the placentae bilamellate, longitudinally affixed to the septum; fruit
baccate, small, oblong to orbicular, 2-celled; seeds numerous, minute, the testa
crustaceous, foveolate.
More than 125 species in tropical America, about 90 of these
from North America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 99
There are doubtless more species in Guatemala than the rather
large number indicated in this treatment. It is one of the largest
genera of Rubiaceae in Central America with the greatest
concentration of species in Guatemala and adjacent Mexico. The
species are distinguished by small but apparently constant
characters. The senior author thought the genus to be one of little
practical interest, however mongraphic work should uncover
interesting distributional patterns.
Stems conspicuously 4-angulate and sometimes narrowly winged on the angles.
Cymes sessile or subsessile; calyx lobes minute and deltoid; plants of the Atlantic
slope H. ghiesbreghtii.
Cymes pedunculate and flowers often appearing racemose; calyx lobes triangular;
plants of the Pacific slope H. riparia.
Stems terete, not winged.
Leaves conspicuously villous beneath, at least on the costa, with lax spreading
hairs, often villous over the whole lower surface, or sometimes densely
villosulous with very short hairs beneath along the costa.
Flowers densely clustered in the leaf axils, the inflorescences mostly much
shorter than the petioles, the flowers sessile or on very short pedicels.
Calyx glabrous; leaves acute to attenuate at the base, green beneath.
H. confertiflora.
Calyx densely villous or villosulous.
Leaves long-attenuate at the base, green beneath H. huehueteca.
Leaves obtuse or rounded at the base, purple beneath H. tuerckheimii.
Flowers in cymes or panicles, not densely clustered, the inflorescences often
elongate, lax, and much exceeding the petioles.
Leaves sessile H. phoenicopoda.
Leaves petiolate.
Leaves mostly obtuse or merely acute; plants herbaceous or at least low,
commonly 60 cm. tall or less H. bullata.
Leaves acuminate or long-acuminate; plants usually tall shrubs, a meter
high or often much taller.
Leaves villous on the upper surface with long spreading ferruginous hairs.
H. gesnerioides.
Leaves not villous on the upper surface, glabrous or pilose with short
appressed hairs.
Corolla villous H. tetrastigma.
Corolla glabrous.
Corolla 20-25 mm. long, red; inflorescence usually lax and much
branched, often equaling or exceeding the petioles ....H. cautiflora.
Corolla 10-12 mm. long, yellow or greenish yellow; inflorescence dense
and compact, usually much shorter than the petioles.
H. chiapensis.
Leaves glabrous beneath, or sometimes puberulent, especially along the costa,
never villous.
Inflorescence arising at the base of the plant; leaves large, 10 cm. broad or more.
H. rhizantha.
Inflorescences arising in the leaf axils or below the leaves, not basal.
100 FIELDIANA: BOTANY, VOLUME 24
Calyx lobes linear to linear-lanceolate, as long as to twice as long as the
hypanthium, 2.5-5 mm. long.
Calyx lobes 4-5 mm. long; corolla lobes twice as long as the tube; lowland
species H. calycosa.
Calyx lobes 2.5-3 mm. long; highland species.
Leaves petiolate; corolla 16-18 mm. long, tube and lobes subequal,
glabrous H. teruae.
Leaves sessile or nearly so; corolla about 8 mm. long, the lobes about
twice as long as the tube, sparsely pubescent with segmented hairs.
H. sessilifolia.
Calyx lobes narrowly triangular or broader, often obsolete, usually much
shorter than the hypanthium, 2 mm. or less long.
Corolla 15-23 mm. long; lobes shorter than the tube; inflorescence long-
pedunculate, usually much longer than the petioles H. macrosiphon.
Corolla about 1 cm. long or shorter, lobes about equaling or usually much
longer than the tube.
Leaves conspicuously lineolate beneath with very numerous, pale, linear
cystoliths.
Leaves opposite, large, sometimes 30 cm. long and 12 cm. broad; flowers
cymose H. lineolata.
Leaves, at least in part, verticillate; 2-5 cm. long; flowers solitary in the
leaf axils on long slender pedicels H. quadrifolia.
Leaves not lineolate beneath.
Lateral nerves of the leaves about 14-20 pairs, very conspicuous
beneath; leaves large and broad, mostly 8-10 cm. broad or broader.
H. riparia.
Lateral nerves of the leaves relatively few, usually 12 or fewer; leaves
mostly less than 8 cm. broad, often only 1-2 cm. broad.
Corolla sparsely villous; flowers all sessile or nearly so.
Branches villosulous; calyx lobes scarcely 1 mm. long H. wilsonii.
Branches glabrous; calyx lobes about 2 mm. long H. tuerckheimii.
Corolla glabrous; flowers all or mostly pedicellate.
Cymes long-pedunculate, longer than the petioles, or the flowers
sometimes racemose, the peduncles usually equaling or longer
than the cymes.
Flowers cymose; leaves broadest above the middle H. rotata.
Flowers racemose; leaves broadest at or below the middle.
H. racemifera.
Cymes sessile or short-pedunculate, the flowers never racemose, the
inflorescences usually shorter than the petioles, the cymes
rarely long-pedunculate but then evidently shorter than the
petioles, or the flowers rarely solitary in the leaf axils and long-
pedicellate.
Calyx lobes lance-subulate or lance-oblong, usually 1 mm. long or
more, conspicuously longer than broad.
Inflorescences 1-2-flowered H. uniflora.
Inflorescences many-flowered.
Lateral nerves of the leaves obscure or almost obsolete, the
blades mostly 2-2.5 broad H. cryptoneura.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 101
Lateral nerves of the leaves very conspicuous, the blades
mostly 4-6 cm. broad or larger.
Leaf blades elliptic-oblong or elliptic, mostly abruptly
acute at the base and not decurrent.
H. psychotriaefolia.
Leaf blades oblanceolate or oblanceolate-oblong, long-
attenuate to the base H. angustifoUa.
Calyx lobes obsolete or broadly deltoid or rounded, less than 1
mm. long.
Calyx truncate or essentially so H. nicotanae folia.
Calyx obviously dentate.
Tube of the corolla half as long as the lobes or less.
Leaves small, mostly 7 cm. long or shorter and 1-3 cm.
broad, on petioles 1.5 cm. long or shorter...//, montana.
Leaves large, mostly 12-20 cm. long and 3-7 cm. broad.
Calyx and hypanthium less than 1.5 mm. long at
anthesis; petioles 2-3 cm. long H. vulcanicola.
Calyx and hypanthium 2-3 cm. long at anthesis; petioles
1-2 cm. long H. culminicola.
Tube of the corolla about equaling the lobes, sometimes
longer; leaves mostly 10-15 cm. long.
Leaves 2-2.5 cm. broad; corolla in bud rounded at the apex.
H. rotata.
Leaves mostly 5-8 cm. broad: corolla in bud narrowed at
the apex H. conzattii.
Hoffmannia angustifolia Standl. Contr. U. S. Nat. Herb. 20:
206. 1919.
Occasional in damp or wet mixed forest, 1,000-1,800 m.;
Quezaltenango; Sacatepequez (type from Acatepe"que, Donnell-
Smith 2747); Chimaltenango; Zacapa; Jalapa. Mexico (Chiapas).
A slender, sparsely branched shrub, usually glabrous throughout, the branches
subterete, green; leaves opposite, membranaceous, on petioles 1-2.5 cm. long,
narrowly elliptic to narrowly lance-oblong or oblanceolate-oblong, 10-20 cm. long and
3-5 cm. broad or sometimes larger, long-acuminate or long-attenuate at the apex,
usually long-attenuate to the base and often decurrent almost to the base of the
petiole, the lateral nerves 12-14 pairs, conspicuous; cymes sessile, with few or many
flowers, dense, shorter than the petioles, the flowers sessile or nearly so; hypanthium
2 mm. long, the calyx lobes lance-oblong, 1-1.5 mm. long, obtuse, ciliolate; corolla
about 12 mm. long, pale green or greenish yellow, glabrous, the lobes linear-oblong,
obtuse, equaling or slightly shorter than the tube; anthers 3 mm. long; berries 4 mm.
long or more, oblong when dry, bright red.
The fruits of this species, like those of most other species of
Hoffmannia, shrink greatly in drying so that the measurements
given for the dry fruits are perhaps not more than half those of the
fresh, juicy fruits.
102 FIELDIANA: BOTANY, VOLUME 24
This species has been assumed to be a common one extending
in range from southern Mexico to Panama. The species actually is
rather limited in distribution, as indicated above. Several other and
quite distinct species have been determined as this species. The
genus is much in need of monographic study.
Hoffmannia bullata L. Wms. Fieldiana, Bot. 36: 52. 1973.
Frequent in dense wet mixed forests or on wet banks along
streams in the forest, occurring at or near sea level in the Atlantic
lowlands but to about 1,400 m. on the Pacific slopes; Alta Verapaz;
Izabal (type, Steyermark 38719); Suchitepequez; Huehuetenango;
Quezaltenango; San Marcos. Mexico (Chiapas); British Honduras;
Honduras; Costa Rica.
Herbaceous or somewhat woody plants 10-60 cm. tall, erect or repent. Stems
simple, subterete, or terete, densely ferrugineous-villous or glabrate, leafy above;
leaves obovate to oblong or oblanceolate, acute or obtuse, long-attenuate to the base,
short petiolate, bullate, pubescent or puberulent with segmented hairs along the
nerves below, glabrous or glabrescent above, the blade 9-20 cm. long and 3-7 cm.
broad, the lateral nerves 8-15 pairs, the petioles mostly 1-3 cm. long, stout, pubescent
or glabrescent; inflorescences axillary, subumbellate or subracemose, the peduncles
slender, 3-7 cm. long, glabrous or sparsely pilose pubescent, pedicels slender, 2-5 mm.
long; flowers 3-10, dull brick red; calyx and hypanthium 3-4 mm. long, the
hypanthium turbinate, glabrous, the calyx lobes triangular or lance triangular, acute,
sparsely villosulous; corolla about 1 cm. long, glabrous, the lobes oblong, obtuse or
acute, 2-3-times as long as the tube; stamens with short filaments, the anthers
exserted; berry bright red.
This plant has gone under the name of Hoffmannia refulgens
(Hook.) Hemsl. a plant described from horticulture and possibly of
South American origin. The plant is not uncommon but the
material in herbaria is often not good. It is possible that the
material from Costa Rica is not the same as that from Mexico and
Guatemala.
Hoffmannia calycosa Donn.-Sm. Bot. Gaz. 40: 4. 1905.
Moist or wet, mixed forest, 200-800 m.; Alta Verapaz (type from
Cubilguitz, Tuerckheim 7912); Solola. Endemic.
A shrub, the branches terete, bifariously puberulent when young; stipules
minute, deltoid-acuminate; leaves opposite, membranaceous, the slender petioles 1-2
cm. long, the blades obovate or obovate-elliptic, 9-15 cm. long, 3.5-5 cm. broad, short-
acuminate, at the base acute, acuminate or attenuate, glabrous above, puberulent
beneath along the veins or almost wholly glabrous, the lateral nerves conspicuous, 6-8
on each side; flowers sessile or nearly so and crowded in the leaf axils, the cymes 1
cm. long or less, subsessile; calyx and hypanthium sparsely villosulous, the
hypanthium 2 mm. long; calyx lobes linear, 4-5 mm. long, acute; corolla glabrous,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 103
about equaling the calyx lobes, the lobes linear-lanceolate, spreading, twice as long as
the tube.
Hoffmannia cauliflora Hemsl. Diag. PI. Mex. 30. 1879. Figure
35.
Wet mixed forest or thickets, 750-2,850 m.; endemic; type
collected in Guatemala by Skinner, the locality unknown; Solola;
Suchitep6quez; Quezaltenango; San Marcos.
A shrub or small tree, 1.5-6 m. high, often almost simple or sparsely branched,
the stout branches subterete, glabrous or nearly so; stipules caducous; leaves
opposite, large, herbaceous, short-petiolate or essentially sessile, broadly elliptic or
obovate-elliptic, 15-40 cm. long and 8-20 cm. broad or even larger, usually abruptly
acuminate, usually long-attenuate to the base of the petiole, glabrous above or nearly
so, beneath villous or villosulous, especially on the veins, the lateral nerves
conspicuous, about 20 pairs; inflorescences mostly sessile, often borne at naked nodes
below the leaves, many-flowered, 9 cm. long or less, glabrous, the flowers usually on
long slender red pedicels; calyx and hypanthium 4 mm. long or less, the calyx lobes
very short, deltoid; corolla glabrous, 20-25 mm. long, bright or dull red, the small
lobes only 2-3 mm. long, ovate; ovary 3-4-celled; berries pale red or purplish black, 5-
6 mm. long.
This is more showy than other species because of its large red
flowers. The shrub is a very common one in many of the wet
quebradas of Quezaltenango and San Marcos.
Hoffmannia chiapensis Standl. Contr. U. S. Nat. Herb. 20:
206. 1919.
Wet mixed forest of the western highlands, 1,400-3,000 m.,
common in many localities; Suchitepe"quez; Quezaltenango;
Huehuetenango; San Marcos. Mexico (Chiapas, the type from Cerro
del Boqueron).
A shrub 1.5-4.5 m. tall, simple or sparsely branched, the branches obtusely
angulate, glabrous; stipules small, deltoid, caducous; leaves opposite,
membranaceous, on slender petioles 1.5-6 cm. long, elliptic to elliptic-oblong or
oblong-oblanceolate, 10-25 cm. long and 4-10 cm. broad or larger, acuminate, at the
base acuminate to long-attenuate, bright green and glabrous above, beneath slightly
paler, glabrous or along the costa somewhat puberulent, the lateral nerves about 14
pairs; cymes sessile or short-pedunculate, with few or many flowers, 4 cm. long or
less, the slender pedicels 1-6 mm. long; hypanthium oblong, glabrous, 2 mm. long;
calyx lobes lance-triangular or oblong, obtuse, 1-2.5 mm. long, sparsely puberulent or
glabrous; corolla yellow or greenish, 10-12 mm. long, glabrous outside, the lobes
narrowly oblong, obtuse, about equaling the tube; fruit oblong, 5-6 mm. long.
Hoffmannia confer tiflor a Standl. Contr. U. S. Nat. Herb. 20:
206. 1919.
104 FIELDIANA: BOTANY, VOLUME 24
Type from San Miguel Uspantan, Quiche, at 2,100 m., Heyde &
Lux 3169. Also in San Marcos and Quezaltenango. Endemic.
A branched shrub 3 m. tall, the branches obtusely quadrangular or subterete,
glabrous; leaves opposite, membranaceous, on petioles 3 cm. long or less, elliptic or
oblong-elliptic, 6-12 cm. long, 2.5-5 cm. broad, acuminate, at the base acuminate or
long-attenuate and decurrent, deep green above, copiously ferruginous-villous, at
least when young, paler beneath, villosulous along the nerves, the lateral nerves
about 11 pairs; cymes sessile, dense, with few or many crowded flowers, the flowers
sessile or short-pedicellate; calyx and hypanthium 2-2.5 mm. long, glabrous, the calyx
lobes minute, broadly deltoid; corolla 9-10 mm. long, white, glabrous outside, the
lobes lance-oblong, acute, equaling the tube; anthers 3 mm. long.
Hoffmannia conzattii Robinson, Proc. Am. Acad. 45: 406.
1910. Coffea obovata Schlecht. & Cham. Linnaea 6: 412. 1831, not
Hoffmannia obovata Standl., 1929.
Wet mixed forest 500-1,650 m.; Chimaltenango; Quezaltenango
(near Santa Maria de Jesus); San Marcos; Huehuetenango. Mexico
(Veracruz and Chiapas).
A simple or branched shrub 1-3 m. tall, glabrous throughout, the branches
subterete; stipules ovate, caducous; leaves opposite, membranaceous, on slender or
stout petioles 2-3.5 cm. long, elliptic to oblong-obovate or oblanceolate-oblong, 10-20
cm. long, 3.5-9 cm. broad, abruptly short-acuminate, long-attenuate to the base or
abruptly decurrent, deep green above, paler and puncticulate beneath, the lateral
nerves 8 or more pairs; cymes axillary, sessile or nearly so, dense, with few or many
flowers, the pedicels equaling or shorter than the hypanthium; hypanthium
subglobose, 2.5 mm. long, the calyx lobes minute, obtuse or acutish; corolla about 6
mm. long or as much as 10 mm., glabrous outside, the lobes oblong, spreading,
slightly longer than the tube; anthers almost equaling the corolla lobes.
Hoffmannia cryptoneura Standl. Field Mus. Bot. 22: 53.
1940. H. nubium Standl. & L. Wms. Ceiba 1: 167. 1950 (type from
Honduras, Glassman 1996). Tzai-yat (Huehuetenango).
Moist forests or cloud forests at 1,300-2,100 m.; Baja Verapaz;
San Marcos; Quezaltenango; Quiche; El Progreso; Zacapa. Mexico
(Chiapas, the type from Mt. Ovando, Matuda 944); Honduras.
A branched shrub 1.5 m. high, the branches terete, glabrous; leaves opposite,
membranaceous, on petioles 6-17 mm. long, oblanceolate or oblong-oblanceolate, 7-12
cm. long, 2-3 cm. broad, narrowly acuminate or sometimes attenuate to an obtuse tip,
gradually long-attenuate to the base, glabrous, blackening when dried, paler beneath,
the lateral nerves about 8 pairs but very obscure or almost obsolete; flowers densely
clustered in the leaf axils or in small short-pedunculate cymes, the pedicels 5 mm.
long or less; hypanthium turbinate, glabrous, 1 mm. long; calyx teeth short,
triangular, acute; corolla glabrous outside, the slender tube 5-6 mm. long, the lobes of
about the same length, spreading, oblong-linear, obtuse; stamens about equaling the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 105
corolla lobes, the filaments exserted, the linear anthers 3.5 mm. long; fruit dull red,
very juicy.
The species is rare in Chiapas and in Guatemala, but it is to be
found in many cloud forests in Honduras.
Hoffmannia culminicola Standl. & L. Wms. Ceiba 1: 166.
1950.
Cloud forests, alt. 1,400-2,400 m. or perhaps more; Chiquimula;
Jalapa. El Salvador and Honduras (type, Standley 4797).
Small fragile shrubs 0.5-2 m. tall. Stems simple or usually branched, obscurely
angled at first, becoming terete, glabrous, to about 8 mm. in diameter, often with
conspicuous cystoliths; leaves oblanceolate to oblong-oblanceolate, acuminate, acute
at the base, glabrous, cystoliths common on the lower surface, short petiolate, the
blade 6-23 cm. long and 1.5-6.5 cm. broad, the lateral veins 5-8 pairs, diverging at an
obtuse angle, mostly not anastomosing toward the margins, the petiole mostly 1-2 cm.
long; inflorescences short few-flowered axillary cymes, peduncles l-6(-10) mm. long,
slender, pedicels mostly 1-2 mm. long; flowers small, yellowish, the corolla early
deciduous; hypanthium and calyx small, glabrous, 2-3 mm. long at anthesis, calyx
lobes triangular, acute, about 0.5-0.75 mm. long and as broad; corolla campanula te,
5-7 mm. long, the four oblong-lanceolate, acute lobes twice as long as the tube;
stamens 4, inserted on the corolla throat, filaments about 0.5 mm. long, attached to
the anther above the base, anthers linear-oblong, obtuse, about 3 mm. long; style
subclavate, 5-6 mm. long, the stigma subglobose; fruits fleshy, globose, about 1 cm.
long at maturity.
This species may be distinguished from the closely allied H.
angustifolia (11-13 pairs) by the few lateral nerves (5-8 pairs) and
by the triangular and acute calyx lobes, not lance-oblong, obtuse
lobes.
Hoffmannia gesnerioides (Oerst.) Kuntze, Rev. Gen. PL 285.
1891. Ophryococcus gesnerioides Oerst. Vid. Medd. Kjoebenhavn
1852: 53. 1852.
Wet mixed forest about 1,500 m.; Zacapa (summit of Sierra de
las Minas); Chiquimula (Cerro Tixixi, north of Jocotan). Mountains
of Honduras and Nicaragua.
A stout shrub about 60 cm. high, or almost herbaceous, simple or sparsely
branched, the branches subterete, densely villous with long spreading rufous hairs;
leaves opposite, on slender petioles 2-3.5 cm. long or essentially sessile, oblong to
obovate-elliptic, 9-18 cm. long, 3-8.5 cm. broad, long-acuminate, long-attenuate to the
base, often to the base of the petiole, membranaceous, when young densely villous on
both sides with long soft rufous hairs, green above and marked with numerous pale
cystoliths, the lateral nerves about 9 pairs; cymes axillary, few-flowered, the
peduncles short in anthesis but in fruit as much as 2.5 cm. long or even more, the
pedicels mostly 2-3 mm. long; hypanthium turbinate, 2 mm. long, villous, the calyx
106 FIELDIANA: BOTANY, VOLUME 24
lobes linear, 3 mm. long, acuminate, villous on the margins, corolla 6-8 mm. long,
villous, the lobes short, triangular; fruit ovoid, purple or purplish pink, 7 mm. long or
more, sparsely villous; seeds minute, coarsely foveolate.
An easily distinguished plant because of abundant pubescence
of long, soft, multicellular hairs. This species is the type of Oersted's
genus Ophryococcus, Two collections from Guatemala are fruiting
and while closely allied to H. gesnerioides, they may prove to be
distinct when better material is available. The specimens actually
may represent two species.
Hoffmannia ghiesbreghtii (Lem.) Hemsl. Biol. Centr. Am.
Bot. 2: 36. 1881. Campylobotrys ghiesbreghtii Lem. 111. Hort. 8: t.
279. 1861. Higginsia ghiesbreghtii ("gheisbechtii") Hook. Bot. Mag.
89: t. 5383. 1863.
Dense wet mixed forest, often on limestone, 1,300 m. or lower;
Peten; Alta Verapaz; Izabal. Mexico; British Honduras; described
from cultivated plants possibly of Mexican origin.
A simple or sparsely branched shrub about 1 m. high, rarely somewhat taller, the
stout branches acutely quadrangular, narrowly winged on the angles, glabrous or
nearly so; leaves opposite, large, sessile or essentially so, oblanceolate-oblong or
obovate-oblong, 20-39 cm. long, 6-9 cm. broad, acute or abruptly acuminate, long-
attenuate to the base, membranaceous, bright green and glabrous above, paler
beneath, sometimes purplish, minutely puberulent, especially on the veins, or
glabrate, the lateral nerves about 20 on each side; cymes axillary, with few or many
flowers, dense and subsessile, the flowers sessile or subsessile; calyx and hypanthium 2
mm. long, puberulent, the calyx lobes minute, deltoid; corolla yellow, sometimes
tinged with red, 6 mm. long, puberulent, the lobes narrowly oblong, acute, spreading,
equaling or slightly longer than the tube; berries red, 3-4 mm. long when dry.
The leaves often are handsomely colored with deep purple
beneath. In Alta Verapaz the shrub has been found only at lower
elevations on the characteristic forested limestone formations
approaching the plains of Peten.
Hoffmannia huehueteca Standl. & Steyerm. Field Mus. Bot.
23: 249. 1947.
Moist or wet, mixed, mountain forest, 1,500-3,300 m.; El
Progreso (Volcan de Santa Luisa); Huehuetenango (Sierra de los
Cuchumatanes, type from Cerro Negro 2 miles east of Las Palmas,
Steyermark 51697). Mexico (Chiapas).
A shrub of 1.5 m., or sometimes not more than 30 cm. high, erect, usually simple,
the stems stout, very densely ferruginous-villosulous with short hairs; stipules
triangular-acuminate, 3 mm. long or less; leaves herbaceous, on stout petioles 2 cm.
long or shorter, oblanceolate or obovate-oblong, 10-20 cm. long, 2-7.5 cm. broad,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 107
shortly cuspidate-acuminate with an obtuse tip, gradually long-attenuate to the base
and often decurrent almost to the base of the petiole, green and glabrous above,
green and paler beneath, brownish when dried, very densely and shortly ferruginous-
villosulous on the costa and nerves, puberulent between them, the lateral nerves
conspicuous, about 12 on each side, arcuate-divergent at a wide angle; flowers
numerous, densely aggregate in the leaf axils, cymulose, the cymules sessile or very
shortly pedunculate, the flowers sessile or short-pedicellate; hypanthium villosulous
or almost glabrous, oval, 2 mm. long; calyx lobes narrowly triangular or narrowly
oblong, 2 mm. long, obtuse or acute, villosulous or almost glabrous; fruit red, suboval,
glabrous or glabrate, 3.5-4 mm. long when dry; seeds numerous, minute.
Hoffmannia lineolata Donn.-Sm. Bot. Gaz. 40: 5. 1905.
Mountains of Alta Verapaz, 350-1,650 m., in dense wet mixed
forest; endemic; type from Cubilguitz, at 350 m., Tuerckheim 8227.
A slender shrub, the branches subterete, glabrous or nearly so; leaves opposite,
membranaceous, on slender petioles 2-7 cm. long, obovate- oblong or oblanceolate-
oblong, as much as 30 cm. long and 12 cm. broad but usually smaller, acuminate,
attenuate to the base, densely covered with pale linear cystoliths, minutely
puberulent beneath along the veins or almost wholly glabrous, the lateral nerves 14-
17 on each side; flowers short-pedicellate, in few-flowered dense cymes 1.5-2.5 cm.
long; calyx and hypanthium ferruginous-puberulent, 2 mm. long, the calyx lobes
minute, deltoid; corolla 6 mm. long, yellowish green, sparsely puberulent, the lobes
linear-oblong, spreading, twice as long as the tube; anthers sessile, 3 mm. long; ovary
2-celled; fruit globose, the seeds reddish.
Hoffmannia macrosiphon Standl. Field Mus. Bot. 17: 276.
1937.
Wet mixed mountain forest, 2,000-3,000 m.; endemic;
Quezaltenango (type from Volcan de Zunil, at 2,400 m., Skutch
937); San Marcos. Possibly also in Mexico (Chiapas).
A glabrous shrub 1.5-4.5 m. high, sometimes weak and reclining, sparsely
branched, the branches terete; leaves opposite, thick-membranaceous, on petioles 1-2
cm. long, narrowly oblanceolate-oblong, 7-12 cm. long, 2.5-3.5 cm. broad, rather
abruptly acute or acuminate, narrowly long-attenuate to the base, pale beneath, the
lateral nerves about 8 pairs, inconspicuous; cymes axillary, solitary, mostly 2-3-
flowered, on peduncles 3 cm. long or less, the slender pedicels about 1 cm. long;
hypanthium narrowly oblong, 4 mm. long, 1.5 mm. broad, glabrous, the calyx 2.5 mm.
broad, erect, obtusely dentate; corolla glabrous, dull yellowish or whitish, the lobes
usually tinged with red or the whole corolla pale red, the tube about 2 cm. long, 5
mm. broad in the throat, the lobes rounded-ovate, obtuse, 6 mm. long, somewhat
spreading, glabrous within; anthers half exserted, almost 4 mm. long; berries
narrowly oblong, 10-12 mm. long, 3-5 mm. thick, pale green, turning deep red.
Hoffmannia montana L. Wms. Fieldiana, Bot. 36: 55. 1973.
108 FIELDIANA: BOTANY, VOLUME 24
Wet mountain forests or cloud forests, 1,400-2,700 m.; San
Marcos; Quezaltenango (type, Skutch 936); Huehuetenango;
Suchitepequez; El Progreso. Mexico (Chiapas).
Much branched slender shrubs 1.5-3 m. tall. The branches at first somewhat and
often sparsely and bifariously fulvous pubescent in the intervals or soon becoming
terete and glabrous; leaves opposite or rarely ternate, narrowly to broadly
oblanceolate, long acuminate, attenuate to a short petiole; glabrous or sparsely
pubescent on petiole and nerves of underside of leaf but soon glabrous, nerves 4-7(-8)
pairs diverging at obtuse angle, blades (3-)4-8 cm. long and (0.6-)2-3 cm. broad,
petioles mostly 0.5-1.5 cm. long; inflorescence axillary, short- pedunculate, 1-few-
flowered, peduncles subfiliform, 2-15 mm. long; pedicels filiform, mostly 3-10 mm.
long; hypanthium and calyx at anthesis turbinate, 2-3 mm. long; calyx divided to
near the base, the lobes triangular, 0.5-1 mm. long, glabrous; corolla white to pale
yellow, up to 1 cm. long, lobate to near the base, the lobes linear-lanceolate, acute,
glabrous, 5-7 mm. long, the tube 1-2 mm. long; stamens inserted in the throat of the
corolla, the filaments short, anthers linear, 2.5-3.5 mm. long, exserted; style filiform,
simple, about 5-8 mm. long, the stigmas clavate lanceiform, undivided and 4-5 mm.
long; mature fruits fleshy, to about 5-6 mm. in diameter.
This species has been confused with Hoffmannia excelsa
(HBK.) Schum. which occurs most commonly in the state of Vera
Cruz, Mexico. It has a bifid style and the corolla divided to about
the middle, not to near the base.
Hoffmannia nicotanaefolia (Mart. & Gal.) L. Wms.
Fieldiana, Bot. 36: 56. 1973. Psychotria nicotanaefolia Mart. &
Gal. Bull. Acad. Brux. 11, pt. 1: 229. 1844. Hoffmannia lenticellata
Hemsl. Diag. PI. Mex. 30. 1879. Aegiphila hoffmannioides Standl. &
Steyerm. Field Mus. Bot. 23: 227. 1947 (type from Huehuetenango,
Steyermark 48843).
Common in wet mixed forests or thickets, sometimes in pine
forest, 900-1,600 m.; Alta Verapaz; Baja Verapaz; Solold;
Huehuetenango. Mexico; British Honduras; Honduras.
A glabrous branched shrub 1-3 m. high, the branches subterete; stipules linear-
oblong, 6 mm. long or less, deciduous; leaves opposite, on stout petioles 0.5-3 cm.
long, obovate to obovate-oblong or elliptic, 10-22 cm. long, 5-8.5 cm. broad, obtusely
acuminate, long-attenuate to the base, membranaceous or subcoriaceous, the lateral
nerves conspicuous, 11-14 on each side; flowers axillary, commonly cymose-
fasciculate, about 6 mm. long, long-pedicellate; calyx and hypanthium 2 mm. long,
the calyx truncate or obscurely dentate; corolla white or yellowish green, often tinged
outside with red, the lobes ovate-oblong, about twice as long as the tube; fruit 2-
celled, purplish black, juicy, sweet, 4-8 mm. long; seeds minute, angulate, punctate.
Hoffmannia phoenicopoda Schum. Notizbl. Bot. Gart. Berlin
2: 276. 1899.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 109
Dense wet mixed forest, 1,000-2,000 m.; Alta Verapaz; Zacapa;
Huehuetenango. Described from cultivated plants of unknown
origin.
Plants herbaceous, 20-100 cm. high, the stems simple, stout, subangulate, sparsely
villous or glabrous; stipules ovate-deltoid, 4 mm. long, obtuse or subacute, green,
persistent; leaves opposite, sessile, obovate or obovate-oblong, 10-25 cm. long, 4-11
cm. broad, acute or broadly short-acuminate, or often obtuse, long-attenuate to the
rounded or slightly amplexicaul base, succulent, somewhat bullate, dark green above,
glabrous or thinly villous, reddish beneath, puberulent or villosulous along the veins,
the lateral nerves 18-20 on each side; inflorescence dense, cymose, few-flowered, the
flowers sessile or subsessile, the slender peduncles 2-8 cm. long, rufous-villous;
hypanthium turbinate, 2 mm. long, glabrous, the calyx lobes triangular, 1.5-2 mm.
long, acute, sparsely villous; corolla sparsely villous, bright red or yellowish red, 9-12
mm. long, the lobes lance-oblong, twice as long as the tube or longer; anthers 5 mm.
long; fruit deep red.
Hoffmannia psychotriaefolia (Benth.) Griseb. Fl. Brit. W.
Ind. 321. 1861. Higginsia psychotriaefolia Benth. ex Oerst. Vid.
Medd. Kjoebenhavn 1852: 50. 1852.
Baja Verapaz. Extending southward to Panama, the type from
Volcan de Barba, Costa Rica.
A branched shrub 1-2.5 m. high, glabrous throughout, the young leaves usually
ciliate, the slender branches subterete; leaves opposite, on slender petioles 1-4 cm.
long, elliptic-oblong or elliptic, 7-15 cm. long, 2.5-5 cm. broad, cuspidate-attenuate,
acute or attenuate at the base, usually abruptly differentiated from the petiole,
bright green above paler beneath, the lateral nerves 8-10 pairs, conspicuous; cymes
sessile, dense, few-flowered, usually shorter than the petioles, the flowers short-
pedicellate; calyx lobes 1.5-2 mm. long, narrowly deltoid, acute; corolla yellow, 8-11
mm. long, the lobes lance-oblong, acute, about equaling the tube; anthers yellow, 4
mm. long; style bifid; berries red, oblong, 7-8 mm. long.
This species originally described from Costa Rica is also found
in the highlands of Panama. It is possible that the single collection
known from Guatemala is another species, perhaps an undescribed
one.
Hoffmannia quadrifolia Standl. & Steyerm. Field Mus. Bot.
23: 250. 1947.
Known only from the type, Zacapa, Sierra de las Minas, around
the summit of Monte Virgen, about 2,400 m., Steyermark 42600.
A very slender and laxly branched shrub 1-2.5 m. high, the branches pale or
ochraceous, almost glabrous, the young branches sparsely pilosulous with short
curved hairs, the internodes short; leaves small, thin-membranaceous, many of them
quaternate, others ternate or opposite, on slender petioles 2-8 mm. long, oblong-
elliptic or elliptic-oblanceolate, 2-5 cm. long, 8-15 mm. broad, obtusely acute or
110 FIELDIANA: BOTANY, VOLUME 24
acuminate, long-attenuate to the base and sometimes decurrent almost to the base of
the petiole, ciliate, deep green above, fuscous when dried, much paler beneath, the
lateral nerves about 4 on each side, obscure, the blade bearing numerous short pale
linear cystoliths on both surfaces, or these often obscure or obsolete in adult leaves;
flowers axillary, solitary, few, the pedicels 2-3 mm. long; hypanthium narrowly
turbinate, glabrous, 2 mm. long; calyx lobes lance-subulate, acuminate, 0.5 mm. long;
corolla greenish white, 6 mm. long in bud, somewhat narrowed to the obtuse apex,
the tube narrowly obconic; fruit red, oval, glabrous, when dry only 3 mm. long.
The type, and only known collection of this species is
inadequate. It is possible that the plant is not a Hoffmannia. The
shining straw-colored stems and quaternate leaves are unusual.
Hoffmannia racemifera Standl. & Steyerm. Field Mus. Bot.
23: 259. 1947.
Known only from the type region, Izabal, Cerro San Gil, at 300-
1,200 m.; type, Steyermark 41910.
A simple or branched, glabrous shrub, the branches slender, the older ones
ochraceous, with short internodes; stipules inconspicuous, deltoid, 1.5 mm. long;
leaves thin-membranaceous, on slender petioles 1.5-4.5 cm. long, elliptic or ovate-
elliptic, 10-16 cm. long, 4-7.5 cm. broad, acuminate or cuspidate-acuminate with a
long narrow acumen, obtuse or acute at the base, not or scarcely decurrent, the
lateral nerves about 8 on each side, arcuate-ascending at a wide angle; inflorescences
axillary, racemose, simple, few-flowered, on long slender peduncles, lax, usually
longer than the petioles, the flowers secund, on slender pedicels 4-6 mm. long;
hypanthium obovoid-turbinate, 1.5 mm. long, acute at the base, glabrous; calyx lobes
scarcely 0.5 mm. long, triangular, acute; corolla in young bud slightly narrowed to
the obtuse apex, glabrous; fruit globose-oval, white, 4 mm. long when dried.
Hoffmannia rhizantha Standl. Carnegie Inst. Wash. Publ.
461: 90. 1935.
British Honduras, in forest along creek bank, Esperanza trail,
630 m., Schipp S731; to be expected in Pete"n.
A glabrous shrub about 60 cm. tall, simple, the thick stems obtusely tetragonous;
leaves very large, herbaceous, opposite, on petioles 3-6 cm. long, the petioles
marginate for all or most of their length; leaf blades oblong-obovate, 23-32 cm. long,
10-12 cm. broad, rather abruptly acuminate, long-attenuate to the base, paler
beneath, the lateral nerves about 14 pairs; inflorescence small and lax, arising from
the base of the stem, sessile and branched from the base, 4 cm. long, the flowers
pedicellate; hypanthium narrowly oblong, attenuate to the base, 3-5 mm. long, calyx
lobes erect, lance-oblong, acuminate; corolla red, glabrous, 5-6 mm. long, attenuate in
bud, oblong-linear, acuminate, the tube extremely short.
Hoffmannia riparia Standl. Field Mus. Bot. 17: 214. 1937.
Quinahuach del rio.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 111
Wet mixed forest of the Pacific, 550-1,400 m.; Suchitepe"quez;
Quezaltenango (type from Colomba, growing beside stream in
forest, Skutch 1291); San Marcos. Mexico (Chiapas).
A shrub 1.5-3 m. tall, simple or branched, glabrous throughout or nearly so, the
stout branches subterete; stipules short, acute; leaves large, opposite, firm-
membranaceous when dry, on stout petioles 1-5.5 cm. long, oblong-obovate or
obovate, 15-30 cm. long, 6-12 cm. broad, acute or short-acuminate, long-attenuate
toward the base and long-decurrent upon the petiole, slightly paler beneath, drying
brownish, the lateral nerves very conspicuous, about 20 pairs; cymes axillary and also
arising from naked nodes below the leaves, lax and many-flowered, about 5 cm. long,
fasciculate, sessile, glabrous, the flowers often conspicuously secund, the slender
pedicels 6 mm. long or less; calyx lobes broadly triangular, about 1 mm. long; corolla
about 7 mm. long, yellowish green, the oblong lobes longer than the tube; berries pale
purple or red, oval or subglobose, 6-8 mm. long.
The shrub is abundant in many parts of the Pacific slopes of
the Sierra Madre, usually growing in dark places along or near
stream banks.
Hoffmannia rotata Donn.-Sm. Bot. Gaz. 18: 204. 1893. H.
steyermarkii Standl. Field Mus. Bot. 22: 192. 1940 (type from San
Marcos, Steyermark 36426).
Wet forests on the Pacific slopes of the Sierra Madre, 2,000-
3,000 m.; San Marcos; Huehuetenango; Quich£ (type from
Uspantan, Heyde & Lux 3169, 3170). Mexico (Chiapas).
A slender shrub 1-2 m. high, the branches subterete, densely ferruginous-
puberulent at first; stipules minute, deltoid; leaves 3-4-verticillate or opposite, on
stout petioles 1 cm. long or shorter, narrowly oblanceolate to oblong-oblanceolate, 8-
18 cm. long, 2-4 cm. broad, acute or acuminate, long-attenuate to the base,
membranaceous, bright green above, glabrous or sparsely puberulent, paler beneath,
copiously puberulent especially along the veins, the lateral nerves 9-12 on each side;
flowers umbellate-cymose, slender-pedicellate, the pedicels usually longer than the
calyx, the cymes 6-15-flowered, on slender peduncles 1-5 cm. long; calyx and
hypanthium glabrous, 2 mm. long, the calyx lobes minute, deltoid; corolla 6 mm.
long, glabrous, the tube almost obsolete, the lobes oblong, obtuse; anthers 2.5-3 mm.
long, exserted, about twice as long as the filaments; fruit globose, red, 5 mm. in
diameter, 2-celled; seeds minute, reddish brown.
Hoffmannia sessilifolia L. Wms. Phytologia 25: 463, fig. 1973.
Figure 36.
Wet cloud forest at about 1,600 m., Baja Verapaz (Sierra de las
Minas, type from above Purulh&, Williams, Molina and Williams
41977).
Erect slender sparsely branched glabrous shrubs to 3 m. tall. The branches
slender, terete with inconspicuous longitudinal rhaphides, the internodes at maturity
112 FIELDIANA: BOTANY, VOLUME 24
6-10 cm. long; leaves opposite, equal, rather large, sessile and obtuse to subauriculate
or subcordate at the base, oblanceolate or oblong- lanceolate, acuminate, 5-16 cm.
long and 2-5.5 cm. broad, green above, bright purple beneath but apparently
becoming green with age, the lateral nerves 10-14 on each side, appearing somewhat
white-marginate with age; inflorescences short, few-flowered axillary cymes, the
peduncles 0.2-1 cm. long; flowers tinged with red; calyx and hypanthium 5-6 mm.
long, the hypanthium 2-3 mm. long, glabrous, 8-ridged, the calyx divided to the base,
the lobes linear-oblong, acute, 2.5-3 mm. long, sparsely pubescent or ciliate with
segmented hairs, with minute glandular or hairlike appendages in the sini; corolla 4-
lobate, the lobes lanceolate or lance-oblong, acute, 6-7 mm. long, sparsely pubescent
dorsally with large segmented hairs, the tube 3-4 mm. long; stamens attached below
the throat of the corolla, 4-4.5 mm. long; style 9-10 mm. long, the stigmas somewhat
enlarged and bilobate; fruits not known.
Easily distinguished from other species in Central America and
Mexico by the sessile leaves, usually obtuse and auriculate or
subcordate at the base, and the unwinged stems. The closest ally is
H. ghiesbreghtii.
Hoffmannia teruae Williams & Molina, Fieldiana, Bot. 36: 59.
1973.
Cool and wet mountains forests, 1,700-3,000 m.; endemic but to
be expected in Chiapas; San Marcos; Solola.
Weak forest shrubs to 2 m. tall. Stems glabrous, terete, the flowering stems to
about 5 mm. in diameter, the internodes about 5 cm. long; leaves elliptic to
oblanceolate, acuminate, glabrous, with about 11-13 pairs of lateral nerves not
anastomosing, the angle rather acute, petiolate, the blade 10-22 cm. long and 2.5-6.5
cm. broad, attenuate into the 1-3 cm. long petiole; inflorescence axillary in the new
growths, few-several-flowered nearly sessile fascicles, glabrous, peduncles none,
pedicels 2-3 mm. long; hypanthium turbinate and 1.5-2 mm. long at anthesis; calyx
divided almost to the base, lobes linear-lanceolate, 2.5-3 mm. long and 0.6-0.8 mm.
broad, acute, ciliolate; corolla 16-18 mm. long at anthesis, the tube narrow and 8-9
mm. long, the lobes linear-oblong, 8-9 mm. long and about 2 mm. broad, subacute
and apiculate thickened at the apex; stamens inserted in the throat of the corolla, the
filaments rather thick, 3.5-4 mm. long, anthers linear, obtuse, about 4.5 mm. long;
style about 18 mm. long, simple, clavate thickened at the apex and the stigmatic area
about 4.5 mm. long; fruits not known.
A highland species easily distinguished by the long and narrow
calyx lobes and the rather large corolla about equally divided into
tube and lobes. The long internodes and large leaves are also
distinctive.
Hoffmannia tetrastigma Donn.-Sm. Bot. Gaz. 27: 336. 1899.
Damp mixed mountain forests, 1,600-1,800 m.; endemic; Santa
Rosa (type from Zamorora, Heyde & Lux 4490); Guatamala;
Sacatepequez; Escuintla; Chimaltenango.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 113
A shrub 1-3 m. high, the stout branches subterete, densely ferruginous-villous;
stipules triangular, caducous; leaves opposite, on stout petioles 1.5-5 cm. long,
obovate or oblong-elliptic, 15-35 cm. long, 7.5-11 cm. broad, acute or acuminate, at
the base acute to attenuate, membranaceous, bright green above, appressed-pilose or
puberulent with ferruginous hairs, slightly paler beneath, copiously ferruginous-
villous, especially along the veins, the lateral nerves 14-15 on each side; cymes
axillary or arising at naked nodes below the leaves, fasciculate, many-flowered, 9 cm.
long or less, the flowers short-pedicellate; calyx and hypanthium 3 mm. long,
villosulous, the calyx lobes minute, deltoid; corolla 11-14 mm. long, ferruginous-
villosulous, the lobes oblong-triangular, 4-5 mm. long; anthers 4 mm. long; ovary 4-
celled; fruit purple or almost black, very juicy, about 1 cm. long.
Hoffmannia tuerckheimii Donn.-Sm. Bot. Gaz. 47: 254. 1909.
Moist or wet, mixed forest, 1,400-2,300 m.; Alta Verapaz (type
from forest near Coban, Tuerckheim 11.2160); El Progreso.
An erect shrub 1 m. high or less, simple or with few branches, the branches
terete, ferruginous-villous with multicellular hairs; stipules triangular; leaves
opposite, on stout petioles 1.5-2.2 cm. long, oval-ovate to rounded-elliptic, 5-10.5 cm.
long, 2.5-6 cm. broad, short-acuminate, at the base rounded or abruptly contracted
and short-decurrent, glabrous above or villous along the costa, beneath usually dark
purple-red, ferruginous-villous, especially along the veins, the lateral nerves 5-7 on
each side; flowers few, fasciculate in the leaf axils, subsessile; calyx and hypanthium
purplish-villous, 3 mm. long, the hypanthium obovoid, the calyx lobes triangular,
subulate, half as long as the hypanthium; corolla sparsely villous, 12 mm. long, the
lobes linear, the tube very short.
The plant is frequent in the very wet mountains above Tactic,
where it is conspicuous because of the usually dark purple under
surfaces of the leaves. A notable variant of the species is H.
tuerckheimii var. glabra Standl. & Steyerm. Field Mus. Bot. 23:
250. 1947 (type from Sierra de las Minas, El Progreso, Montana
Canahui, between Finca San Miguel and summit of mountain,
1,600-2,300 m., Steyermark 43806). In this the branches and leaves
are quite glabrous, but the corolla is sparsely villous. In other
characters it is exactly like the typical form of the species.
Hoffmannia uniflora Standl. Contr. U. S. Nat. Herb. 20: 204.
1919.
Moist or wet, mixed forest, 1,100-3,300 m.; endemic; Alta
Verapaz (type from Coban, Tuerckheim 11.2107).
A shrub, the slender branches subterete, bifariously rufous-puberulent; stipules
deltoid, acutish, 1 mm. long, deciduous; leaves opposite, on slender petioles 3-6 mm.
long, narrowly oblong-elliptic or lance-elliptic, 4-8 cm. long, 1-2 cm. broad, acuminate
or long-acuminate, attenuate to the base, membranaceous, deep green and glabrous
above, paler beneath, sparsely puberulent along the veins or glabrate, the lateral
114 FIELDIANA: BOTANY, VOLUME 24
nerves about 6 on each side; flowers mostly solitary in the leaf axils, sometimes in 2-
flowered cymes, the slender pedicels 3-6 mm. long, glabrate; calyx lobes linear, acute,
in fruit 2-3 mm. long; fruit oval, 6-7 mm. long, 5-6 mm. broad, glabrous; seeds minute,
brownish, coarsely reticulate.
Hoffmannia vulcanicola Standl. & Steyerm. Field Mus. Bot.
23: 251. 1947.
Moist or wet, mixed, moutain forest, 1,000-1,700 m.; endemic;
Solola (type from slopes of Volcan de Atitlan above Finca Moca,
Steyermark 47931); San Marcos (Volcan de Tajumulco, above Finca
El Porvenir).
A glabrous shrub 2-3 m. high, the branches very slender, terete, with elongate
internodes; leaves membranaceous, on slender petioles 2-3 cm. long, elliptic-lanceolate
or elliptic, mostly 12-15 cm. long and 4.5-7 cm. broad, narrowly long-acuminate, acute
or obtuse at the base, not or scarcely decurrent, fuscescent above when dried, paler
beneath, the lateral nerves about 11 on each side, very slender, arcuate-ascending at a
wide angle; inflorescences solitary or in fascicles of 2-3, on peduncles as much as 3
cm. long, laxly cymose or cymose-paniculate, mostly few-flowered, the very slender
pedicels 3-7 mm. long; hypanthium narrowly clavate, 1.5 mm. long, glabrous; calyx
scarcely 0.4 mm. high, the teeth minute, triangular, subacute; corolla pale yellow,
glabrous, 8 mm. long, in bud almost linear and acute or acuminate, 5-lobate almost
to the base, the lobes lance-oblong, acute, the tube very short; fruit oval, 5 mm. long
when dried.
Hoffmannia wilsonii Standl. Field Mus. Bot. 22: 117. 1940.
Alta Verapaz, the type from Finca Los Alpes, 1,200 m., in virgin
forest, C. L. Wilson 344.
A shrub 2-3 m. high, the branches terete, sordid-villosulous when young; stipules
acuminate from a broadly triangular base, 2 mm. long; leaves membranaceous,
opposite, on slender petioles 3-5.5 cm. long, ovate-elliptic or oblong-elliptic, 12-18 cm.
long, 6-8 cm. broad, rather abruptly long-acuminate, acute or subobtuse at the base,
green and glabrous above, appressed-pilose beneath along the veins, the lateral nerves
about 8 on each side; flowers densely clustered in the leaf axils, sessile or short-
pedicellate; hypanthium oblong-turbinate, about 2 mm. long, glabrous or very
sparsely pilosulous; calyx lobes scarcely 1 mm. long, triangular-subulate, erect, acute;
corolla glabrous outside or sparsely pilose, 7 mm. long, the narrow lobes acutish,
much longer than the tube.
HOFFMANNIA MACROPHYLLA (Planch.) Hemsl. Biol. Cent. Am.
Bot. 2: 37. 1881. Higginsia macrophylla Planch. Fl. Serres 5: 482d.
1849. Described from plants cultivated in Belgium, grown from
Guatemalan seeds. The identification of the plant is doubtful.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 115
HOUSTONIA Linnaeus
Erect or prostrate herbs, annual or perennial, rarely low shrubs, glabrous or
pubescent; stipules entire, dentate, or laciniate, persistent; leaves opposite, sessile or
petiolate, broad or narrow; flowers small, white, purple, or blue, axillary and solitary
or in dichotomous cymes, the pedicels erect or recurved in fruit; hypanthium terete or
compressed, globose to turbinate; calyx lobes 4, sometimes with interposed glands,
remote, erect or recurved; corolla funnelform or salverform, the tube short or
elongate, glabrous or pilose within, the 4 lobes valvate in bud, glabrous or pilose
inside; stamens 4, inserted in the corolla throat, the filaments short or elongate;
anthers dorsifixed, oblong or linear; ovary 2-celled, the style slender, with 2 linear
branches; ovules few or numerous, the placentae affixed to the septum; capsule one-
fourth to three-fourths superior, with few or many seeds, loculicidally dehiscent;
seeds crateriform or concavo-convex, peltate.
About 35 species in North America, chiefly in the United States
and Mexico. Only one species is known from Central America.
Houstonia serpyllacea (Schlecht.) C. L. Smith ex Greenm.
Proc. Am. Acad. 32: 284. 1897. Hedyotis serpyllacea Schlecht.
Linnaea 9: 599. 1834. Mallostoma shannonii Donn.-Sm. Bot. Gaz.
18: 203. 1893 (type from Chichoy, Chimaltenango, 2,700 m., W. C.
Shannon 371). Arcytophyllum shannonii Standl. Contr. U. S. Nat.
Herb. 18: 128. 1916. Pasto de oveja (fide Aguilar). Figure 4.
Abundant in the highlands, in pastures or meadows, often on
roadside banks or on white-sand slopes, 1,700-4,200 m.; Jalapa
(Montana Miramundo); Guatemala; Sacatepequez; Chimaltenango;
Totonicapan; Solola; Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico.
A creeping perennial herb, rooting at the nodes, often forming dense mats,
sometimes pendent from banks, the stems often greatly elongate, hirsutulous or
glabrate; stipules minute, 1-2-cuspidate, glandular-ciliate; leaves very shortly
petiolate or subsessile, oval to ovate-elliptic, mostly 3-6 mm. long and 2-4.5 mm.
broad, acute to rounded at the apex, rounded at the base, subcoriaceous, glabrous,
scaberulous-ciliate; flowers solitary in the leaf axils, the stout pedicels equaling or
shorter than the calyx and capsule; hypanthium glabrous or scaberulous; calyx lobes
lance-linear, 2-4 mm. long; corolla funnelform, 1 cm. long or less, white or sometimes
dark reddish outside, the lobes lance-triangular, acute, about equaling the tube,
hirsutulous within; capsule oblong-obovate, 4 mm. long, costate, about three-fourths
inferior, obtuse at the apex; seeds rounded-oblong, concave-convex, coarsely
scrobiculate.
This is one of the most abundant of plants throughout the
western highlands, although an inconspicuous one. Few
Guatemalan plants are represented in the country by a greater
number of individuals. Houstonia serpyllacea grows in all parts of
116 FIELDIANA: BOTANY, VOLUME 24
the alpine meadows and in the extensive pastures where sheep feed.
If one were to shut one's eyes and touch the ground anywhere in
this region, it is probable that a plant of this species would be
touched. It remains green through the dry months, and the stems
cling so closely to the ground that apparently they are not bothered
by the sheep.
ISERTIA Schreber
Trees or shrubs, more or less pubescent, with stout terete branches; leaves
opposite or ternate, petiolate, coriaceous or herbaceous, large; stipules interpetiolar, 2
on each side, erect, persistent; flowers mostly large, short-pedicellate, red and yellow
or sometimes white, disposed in cymes arranged in thyrsiform erect panicles, the
pedicels bracteolate; hypanthium subglobose or ovoid, the calyx short, 4-6-dentate or
truncate, persistent; corolla tubular-funnelform, coriaceous, the tube often costate,
barbate in the throat, the limb 4-6-lobate, the short lobes valvate in bud; stamens 4-
6, inserted in the corolla tube, the short filaments complanate; anthers dorsifixed,
linear, acuminate, included; ovary 4-6-celled, the slender style 2-6-parted at the apex
or with a thick simple stigma; ovules numerous, multiseriate, the placentae affixed to
the axis of the ovary; fruit small, baccate, globose or ovoid, 4-6-pyrenate, the nutlets
cartilaginous, containing few or many seeds; seeds minute, subglobose, the testa
crustaceous, foveolate.
A genus of about a dozen species, only two of which occur in
continental North America (including the genus Cassupa). I.
panamensis Standl. occurs in the Atlantic lowlands of Panama.
Isertia haenkeana DC. Prodr. 4: 437. 1830. /. deamii Bartlett
in Rob. & Bartl. Proc. Am. Acad. 43: 59. 1907 (type from Puerto
Barrios, C. C. Deam 48). I. deamii var. stenophylla Donn.-Sm. Bot.
Gaz. 61: 374. 1916 (type from Costa Rica). Figure 22.
Wet thickets at or little above sea level; Izabal. Honduras,
along the Atlantic coast to Panama; West Indies; Colombia.
A shrub, usually 2-3 m. high, sparsely branched, the branchlets densely sericeous
with grayish or fulvous hairs; stipules linear to oblong-ovate, about 1 cm. long;
petioles stout, 2.5 cm. long or less, the blades obovate to oblong-obovate, 20-45 cm.
long, 8-18 cm. broad, abruptly acuminate, usually acute to attenuate at the base,
membranaceous, bright green and glabrous above, somewhat paler beneath and
densely short-pilose, in age sometimes glabrate; panicles erect on stout peduncles,
thyrsiform, 7-20 cm. long, the lowest bracts often foliaceous, the others small, lance-
linear; calyx and hypanthium 2.5-4 mm. long, puberulent or glabrous, the calyx very
short, red, truncate or shallowly dentate; corolla 17-35 mm. long, yellow, sparsely or
densely tomentulose outside, sometimes glabrous, the short lobes oblong or
lanceolate, acute, densely barbate within; anthers 4-5 mm. long; fruit purplish,
depressed-globose, 5-6 mm. broad; seeds brown, 0.8 mm. long, foveolate.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 117
The plant is a handsome and very showy one when in flower.
The inflorescences are much used about the Canal Zone for
decorations.
IXORA Linnaeus
Trees or shrubs, glabrous or pubescent, the branchlets terete or angulate; stipules
interpetiolar, persistent or deciduous, usually acuminate from a broad base; leaves
mostly opposite, sessile or petiolate, commonly coriaceous; flowers in terminal or
rarely axillary corymbs, white, pink, or red, pedicellate, the pedicels bracteolate;
hypanthium ovoid, the calyx short, normally 4-lobate, persistent, the lobes short or
elongate; corolla salverform, the tube slender, cylindric, the throat glabrous or pilose,
the limb 4-lobate, the lobes usually short, spreading, contorted in bud; stamens 4,
inserted in the throat or mouth of the corolla, the filaments very short or obsolete;
anthers linear or oblong, dorsifixed, usually acuminate, erect, exserted or
semiexserted; ovary 2-celled, the style filiform, exserted, with 2 short, spreading or
recurved branches; ovules solitary, peltately attached to the middle of the septum;
fruit baccate, globose, coriaceous or fleshy, dicarpidiate, the carpels chartaceous;
seeds concavo-convex, the testa membranaceous.
A vast genus, most of the species in tropical Asia and the East
Indies, but a large number in South America. Only three species are
known to be native in North America, including the West Indies;
and only two in Central America. One, /. floribunda (A. Rich.)
Griseb., ranges from Honduras and El Salvador to Panama.
Leaves obtuse to subcordate at the base, sessile; corolla red /. coccinea.
Leaves acute to attenuate at the base, petiolate; corolla white.
Corolla tube about 30 mm. long; cultivated species I. ftnlaysoniana.
Corolla tube about 5 mm. long; native species /. nicaraguensis.
Ixora coccinea L. Sp. PL 110. 1753. Argentina.
Cultivated commonly for ornament at low and middle
elevations. Native of India.
A densely branched shrub 1-3 m. high, glabrous or nearly so; leaves sessile,
oblong to suborbicular, commonly 4-9 cm. long, rounded to subacute at the apex,
somewhat narrowed to the obtuse or more often cordate base; inflorescence dense
and almost headlike, many-flowered, the flowers sessile or nearly so; calyx lobes
triangular, acute; corolla red or orange-red, minutely puberulent or glabrate, the very
slender tube 2.5-4 cm. long, the spreading lobes ovate-oblong, about 1 cm. long.
A favorite ornamental shrub in Central American gardens
generally. Known in Costa Rica by the names "flor de fuego" and
"jazmin."
Ixora finlaysoniana Wall. Cat. No. 6166. 1832, nomen nudum;
G. Don, Gen. Syst. 3: 512. 1832.
118 FIELDIANA: BOTANY, VOLUME 24
Cultivated commonly for ornament, chiefly in the tierra
caliente and doubtless as an escape. Native of southeast Asia.
A densely branched shrub 1-3 m. tall, glabrous throughout or nearly so; stipules
rounded-ovate, short-cuspidate; leaves short-petiolate, oblong or oblanceolate-oblong,
mostly 10-14 cm. long, narrowly rounded to acute at the apex, long-attenuate to the
base; inflorescence dense and headlike, or sometimes of several terminal heads, with
very numerous flowers, the flowers sessile; calyx lobes oblong-lanceolate, somewhat
foliaceous, 2-3 times as long as the hypanthium; corolla white, the filiform tube about
3.5 cm. long, the obtuse lobes 6-7 mm. long; anthers long-exserted.
A favorite ornamental shrub in the warmer parts of Central
America. In Costa Rica it is called "corona de la reina" and "mono
de la reina;" "corona" and "buquet de novia" (El Salvador). It has
been reported from Guatemala under the name Ixora thwaitesii
Hook. f.
Ixora nicaraguensis Wernham, Journ. Bot. 50: 243. 1912. /.
rauwolfioides Standl. Trop. Woods 11: 25. 1927. Amoco (fide
Record). Figure 46.
Moist or wet lowland forest; Izabal; probably in Peten. British
Honduras to Panama.
A shrub or tree 2.5-8 m. high, the trunk as much as 12 cm. in diameter, glabrous
almost throughout; stipules triangular, acute, 5 mm. long; petioles stout, 4-11 mm.
long, the blades elliptic-oblong or lance-oblong, 7-15 cm. long, 2-5.5 cm. broad, acute
or short-acuminate, acute or acuminate at the base and decurrent, the lateral nerves
about 10 on each side; inflorescence terminal, cymose-paniculate, sessile or on a
peduncle 3 cm. long, much branched and many-flowered, 5-9 cm. long, the slender
branches minutely puberulent, the pedicels as much as 5 mm. long; calyx and
hypanthium 2 mm. long, minutely puberulent, the calyx truncate or nearly so, much
shorter than the hypanthium; corolla white, glabrous, the tube 5 mm. long, the lobes
oblong, spreading, rounded at the apex, 3-4 mm. long; fruit subglobose, red, about 7
mm. long, juicy.
LINDENIA Bentham
Low shrubs, glabrous or pubescent, with terete branches; stipules interpetiolar,
short, connate, cuspidate, persistent; leaves opposite, narrow, short-petiolate; flowers
very large, in few-flowered terminal cymes, short-pedicellate, the bracts subulate;
hypanthium elongate-turbinate, 5-angulate or 5-costate, the calyx 5-lobate, the lobes
elongate-subulate or lanceolate, equal or unequal, persistent; corolla salverform,
white, with a much elongate, terete tube, the throat naked, the limb 5-lobate, the
lobes oblong, spreading, contorted in bud; stamens 5, inserted at the top of the
corolla tube; anthers sessile, linear-oblong, obtuse, recurved, exserted; ovary 2-celled,
the style filiform, pilose, bifid, the lobes erect; ovules very numerous, the placentae
longitudinally adnate to the septum; capsule clavate or pyriform, 2-celled,
septicidally bivalvate, the valves bifid, often elastically coiled after dehiscence; seeds
numerous, small, angulate, smooth.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 119
The genus consists of a single species.
Lindenia rivalis Benth. PL Hartw. 84. 1841. L. acutiflora
Hook. Icon. t. 475. 1842. Flor de Maria; jazmin de agua. Figure 6.
Type collected on stream banks, of (Alta?) Verapaz, Hartweg
581. Confined to rocky stream banks, at 1,300 m. or less, chiefly in
the tierra caliente of the two coasts, on the Pacific slope mostly
along streams at the base of the hills; Peten; Izabal; Alta Verapaz;
Zacapa; Escuintla; Suchitepe'quez; Quiche"; Retalhuleu; San
Marcos. Southern Mexico to Panama.
A stiff shrub, usually 1 m. or less in height, the branches dark brown or
ferruginous, usually pilosulous when young; stipules annular, 3-5 mm. long, the lobes
cuspidate; leaves often crowded at the ends of the branches, on stout petioles 2-10
mm. long, oblanceolate to linear-elliptic, 5-16 cm. long, 1-2.8 cm. broad, acute to
attenuate, long-attenuate to the base, almost glabrous or somewhat puberulent or
pilosulous; inflorescences mostly 3-flowered cymes, the peduncles short and stout, the
thick pedicels 1 cm. long or less; hypanthium 7-10 mm. long, sparsely or densely
hirsutulous; calyx lobes 1-2 cm. long, green; corolla tube usually 10-16 cm. long, 2.5-3
mm. thick, pilosulous outside, the lobes elliptic to oblong-oval, ".5-3 cm. long, acute
to very obtuse; anthers 1 cm. long; capsule pyriform, 1.2-2 cm. long, costate, thick
and woody, the exocarp finally separating from the endocarp; seeds 1.5-2 mm. long,
brownish.
Called "lirio" in Costa Rica; the name "chilca" is reported from
Honduras. A showy plant because of the large white corollas with
extraordinarily long and slender tube. Few plants are so specialized
in habitat as this, which, like some of the Cuphea and Aster species,
is confined to rocks in or near the margins of streams where often it
forms dense low thickets. During floods the shrubs often are wholly
covered with rushing water. They bloom during the rainy season
and we have seen none in flower during the dry months of the year.
MACHAONIA Humboldt & Bonpland
Shrubs or small trees, glabrous or pubescent, the branchlets terete, often spinose;
leaves mostly opposite, sometimes ternate or fasciculate, petiolate or sessile; stipules
intrapetiolar, triangular, acute or acuminate; flowers small, white or whitish, the
inflorescence terminal; hypanthium turbinate or obovoid-oblong, compressed; calyx
4-5-lobate, the lobes equal or unequal, persistent; corolla small, short-funnelform, the
tube short, with villous throat, the limb 4-5-lobate, the lobes rounded, imbricate, 1 or
2 of them exterior; stamens 4-5, inserted in the corolla throat, the filaments short or
elongate; anthers short-oblong, dorsifixed, included or exserted; ovary normally 2-
celled, the style filiform, with 2 spatulate branches; ovules solitary, pendulous from
the apex of the cell; fruit small, dry, obpyramidal or oblong, laterally compressed,
dicoccous, the cocci indehiscent, obtusely trigonous, finally pendulous from the
persistent axis; seeds cylindric, the testa membranaceous.
120 FIELDIANA: BOTANY, VOLUME 24
About 25 species in tropical America, most of them West
Indian. One other species, M. martinicensis (DC.) Standl. (M.
rotundata Griseb.), occurs in Central America, in Costa Rica and
Panama; two others are in Mexico.
Leaves copiously pilose beneath; calyx lobes oval or orbicular, shorter than the
hypanthium M. acuminata.
Leaves glabrous beneath, barbate in the nerve axils; calyx lobes spatulate-obovate,
equaling the hypanthium, much accrescent in fruit M. lindeniana.
Machaonia acuminata Humb. & Bonpl. PL Aequin. 1: 101, t.
29. 1806.
Pete"n (La Libertad). British Honduras; southern Mexico to
Panama; southward to Ecuador and southern Brazil.
A shrub or small tree, said to be sometimes 10 m. high, the branchlets densely
short-pilose or tomentulose; stipules 2-4 mm. long, setaceous-acuminate; petioles 3-10
mm. long, the blades ovate or ovate-elliptic, 5-10 cm. long, 2.5-5 cm. broad, acute or
short-acuminate, rounded to subacute at the base, membranaceous, green and almost
glabrous above, tomentulose beneath when young, becoming short-pilose;
inflorescence much branched, paniculate, 6-10 cm. broad, the flowers sessile or
subsessile, densely glomerate, the bracts very small, subulate; calyx and hypanthium
2 mm. long, the hypanthium densely white-pilose with incurved hairs; corolla white,
4-5 mm. long, sparsely puberulent, the lobes half as long as the tube; anthers
exserted; fruit turbinate, 5 mm. long, puberulent.
Machaonia lindeniana Baill. Bull. Soc. Linn. Paris 1: 204.
1879. Figure 41.
Moist thickets, 300 m. or less; Pet<§n. Mexico (Campeche and
Yucatan) to British Honduras.
A shrub or tree 2-7 m. high, the trunk to 5 cm. in diameter, unarmed, the
branches dark brown, coarsely lenticellate, the branchlets puberulent in 2 lines;
petioles 2-4 mm. long, the blades broadly ovate to ovate-oblong or elliptic-oblong, 2-4
cm. long, 1-1.8 cm. broad, acute to obtuse at the apex, acute or obtuse at the base,
short-barbate beneath in the nerve axils but elsewhere glabrous; inflorescence many-
flowered, 4-7 cm. broad, the branches puberulent or glabrate, the flowers sessile t
long-pedicellate, the linear bracts sometimes 8 mm. long, green; calyx and
hypanthium 1.5-2 mm. long, the hypanthium sparsely pilose with short spreading
hairs or glabrate, the calyx lobes spatulate-obovate, rounded at the apex, ciliolate,
accrescent in age and longer than the fruit; corolla 3 mm. long, ochroleucous,
glabrous, the lobes ovate, obtuse, shorter than the tube; fruit obpyramidal, 2-2.5 mm.
long.
Known in Yucatan by the Maya names "cuchel" and
"campocolche."
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 121
MANETTIA Mutis
Reference: Standley, Paul C., Manettia, in No. Am. Fl. 32: 96-
100. 1921.
Scandent herbs, glabrous or pubescent; stipules short and broad, acute,
sometimes adnate to the petiole; leaves opposite, petiolate, small; flowers small or
large, solitary in the leaf axils or arranged in small cymes or panicles, white, red, or
yellow; hypanthium turbinate to obovoid or campanulate; calyx usually 4-lobate,
persistent, the lobes short or elongate, broad or narrow, often alternating with minute
teeth or large lobes; corolla tubular or funnelform, with short or elongate tube,
villous in the throat, the limb 4-lobate, the lobes usually short, erect or recurved,
valvate in bud; stamens 4, inserted in the corolla throat, the filaments short, included
or exserted; anthers versatile, oblong; ovary 2-celled, the style filiform, the stigma
clavate or bifid; ovules numerous, imbricate, peltate; capsule obovoid or turbinate.
bisulcate, coriaceous or chartaceous, septicidally bivalvate from the apex; seeds
numerous, compressed, surrounded by a broad, irregularly dentate wing.
About 85 species in tropical America, chiefly in South America.
Only one other species is known from Central America, M. barbata
Oerst. of Costa Rica.
Calyx lobes 8 M. reclinata.
Calyx lobes 4 M. flexilis.
Manettia flexilis Brandegee, Univ. Calif. Publ. Bot. 6: 196.
1915.
Apparently scarce in Guatemala, in thickets at 1,100-1,650 m.;
Alta Verapaz (Pansamala'); Quiche'; Huehuetenango;
Quezaltenango (Volcan de Zunil); San Marcos (southern slopes of
Volcan de Tajumulco). Mexico (Chiapas, the type from Cerro del
Boqueron); Costa Rica and Panama; reported, probably in error,
from Ecuador.
Stems slender, puberulent or villosulous; petioles 2-6 mm. long; leaf blades ovate
to lance-ovate, 3-7 cm. long, 1-3 cm. broad, acute to attenuate, acute or obtuse at the
base, glabrous above or nearly so, usually short-pilose beneath along the veins;
inflorescences axillary, cymose and few-flowered or sometimes 1-flowered, the
pedicels 2.5 cm. long or less; hypanthium 2-3 mm. long, glabrous or pilose; the 4 calyx
lobes linear to lance-ovate, 1.5-4 mm. long, recurved, usually ciliate; corolla white or
rose, glabrous outside or sparsely pilose, the stout tube 6-7 mm. long, densely barbate
in the throat, the lobes 2-3 mm. long; anthers subexserted; capsule 4.5-7 mm. long;
seeds 2-3 mm. broad, brown.
MANETTIA LUTEO-RUBRA var. PARAGUARIENSIS (Chod.)
Chung. (M. inflata Sprague). This species, native of Paraguay and
122 FIELDIANA: BOTANY, VOLUME 24
Uruguay, is well known in cultivation, and is sometimes seen in
gardens of Guatemala. The corolla is 4-lobate, very densely
tomentose outside, and scarlet with yellow lobes.
Manettia reclinata L. Mant. 558. 1771. Nacibea coccinea
Aubl. PL Guian. 96, t. 37, f. L 1775. M. cuspidata Bert, ex Spreng.
Syst. Veg. 1: 415. 1825. M. coccinea Willd. Sp. PL 1: 624. 1797. M.
seleriana Loes. Verh. Bot. Ver. Brandenb. 65: 107. 1923 (type from
Los Amates, Izabal, Seler 3309). M. orbifera Wernham, Gen.
Manettia 41. 1919 (type from Retalhuleu, Bernoulli & Carlo 1649).
Buquet de coral (fide Aguilar). Figure 12.
Occasional in the tierra caliente of both coasts, ascending in the
Pacific bocacosta to 1,400 m., usually growing in wet thickets;
Peten; Alta Verapaz; Izabal; Santa Rosa; Escuintla; Guatamala
(Fraijanes); Chimaltenango; Suchitepequez; Retalhuleu;
Quezaltenango. Southern Mexico; British Honduras to Panama;
West Indies; Guianas.
Stems slender, angulate, glabrous or short-pilose, usually retrorse-pilose on the
angles; stipules 1-2 mm. long; petioles 2-14 mm. long; leaf blades lanceolate to ovate,
3-10 cm. long, 1-5 cm. broad, acute or acuminate, acute or obtuse at the base, bright
green above and glabrous or scaberulous, short-pilose beneath, especially along the
veins, or glabrous; peduncles axillary, mostly 1-flowered, the slender pedicels 1.5-6
cm. long; hypanthium 3-5 mm. long, contracted and slightly prolonged above the
ovary, glabrous or short-pilose, the 6-8 calyx lobes subequal, linear, attenuate, 4-14
mm. long, glabrous or sparsely short-pilose, recurved; corolla dull red, 17-28 mm.
long, more or less pilose outside, sparsely barbate in the throat or naked, the lobes 4-5
mm. long, triangular-oblong, ovate, or suborbicular, glabrous within or nearly so;
stamens included; capsule 6-10 mm. long, 6-8 mm. broad; seeds 2-3 mm. broad, the
wing pale brown; the body dark brown.
Manettia reclinata was treated as a name of doubtful
application by Wernham in his monograph of the genus and by the
senior author in North American Flora. Since, however, the plant
treated here agrees in all essential details with the original
description of M. reclinata, and is the only Mexican plant known at
present that does agree with it, it seems preferable to use the name
M. reclinata for the species that has passed in recent years as M.
coccinea. Manettia orbifera was maintained as a distinct species in
North American Flora, being separated on its orbicular corolla
lobes as distinguished from the ovate corolla lobes of M. coccinea. A
phototype of M. orbifera has been examined and the Guatemalan
specimens agree with that and the original description. M. seleriana
is exactly equal to the common form of M. reclinata. In spite of its
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 123
red flowers, this, like other Central American species, is an
inconspicuous plant, in no way comparable in beauty with the
South American members of the genus that sometimes are
cultivated.
MITCHELLA Linnaeus
Perennial creeping herbs, glabrous or puberulent; stipules interpetiolar, minute;
leaves opposite, short-petiolate, ovate-rounded; inflorescence axillary and terminal,
flowers small, white, ebracteate, geminate and connate at the apex of the peduncle;
hypanthium ovoid, the calyx 3-6-dentate, persistent; corolla funnelform, barbate in
the throat, the limb 3-6-lobate, the lobes barbate within, recurved in anthesis, valvate
in bud; stamens 3-6, inserted in the corolla throat, with stout filaments; anthers
dorsifixed near the base, oblong, exserted; ovary 4-celled, the style filiform, with 4
short filiform hirsute branches; ovules solitary, erect from the base of the cell; fruit
of 2 connate drupes, containing 8 nutlets, these osseous, obtusely trigonous.
One other species is known and that in Japan.
Mitchella repens L. Sp. PL 111. 1753. Figures 53.
Zacapa, pine forest, Sierra de las Minas, 1,700-2,000 m.
Southern Mexico; eastern Canada and central and eastern United
States. Figure 53.
Stems slender, rooting at the nodes, 30 cm. long or less, glabrous or nearly so;
leaves on rather long petioles, broadly ovate to ovate-orbicular, mostly 1-3 cm. long,
obtuse to acute, at the base rounded or shallowly cordate, dark green and lustrous
above, slightly paler beneath, glabrous; peduncles shorter than the leaves, bearing at
the apex 2 sessile white flowers; calyx teeth triangular, acute; corolla about 1 cm.
long; fruit bright red, 5-8 mm. broad, long-persistent.
Called "partridge berry" and "twin berry" in the United States,
where the plant often is used for winter holiday decorations. In the
winter state it is attractive because of the abundant bright-colored
berries. The leaves persist through the winter, retaining their green
color. The plant is exceedingly rare in tropical America. We have
seen only one collection from Mexico and two from Guatemala.
MITRACARPUS Zuccarini
Erect or prostrate, annual or perennial herbs with more or less tetragonous
stems; stipules connate with the petioles to form a setiferous sheath; leaves opposite,
herbaceous, linear to ovate; flowers very small, white, crowded in dense terminal
heads, these usually subtended by 4 leaflike bracts; hypanthium turbinate to
subglobose; calyx of 4-5 narrow teeth, persistent; corolla salverform or funnelform,
the tube usually with a ring of hairs inside, the throat naked or villous, the limb 4-
lobate, the lobes spreading in anthesis, valvate in bud; stamens 4, inserted in the
corolla throat, the anthers dorsifixed, oblong or linear, included or exserted; ovary 2-
124 FIELDIANA: BOTANY, VOLUME 24
celled, the style short or elongate, with 2 short linear branches; ovules solitary,
affixed peltately to the middle of the septum; fruit didymous, membranaceous,
circumscissile at or below the middle, the upper portion separating with the calyx and
exposing the seeds, the septum persistent with the basal portion; seeds oblong or
globose, the ventral surface plane.
About 30 species in tropical America and Africa. No other
species are known from Central America, but numerous
representatives of the group occur in South America.
Plants perennial, with numerous stems from a somewhat woody base.
M. rhadinophyllus.
Plants annual, usually simple at the base M. hirtus.
Mitracarpus hirtus (L.) DC. Prodr. 4: 572. 1830. Spermacoce
hirta L. Sp. PI. ed. 2. 148. 1762. M. breviflorus Gray, PL Wright. 2:
68. 1853. Figure 66.
Common and widely distributed, usually as a weed in waste or
cultivated ground, sometimes in pastures, on open banks, or in
thickets, ascending to 1,900 m., but most common at lower
elevations, especially on the coastal plains; Alta Verapaz; Izabal;
Zacapa; Jalapa; Jutiapa; Sacatepequez; Chimaltenango; Quiche";
Huehuetenango; Escuintla; Suchitepe"quez; doubtless in most of the
other departments as well. Southwestern United States to Panama,
and southward through the greater part of South America.
Plants annual, generally erect, simple or sometimes much branched, commonly
30-60 cm. high, the stems relatively stout, villous or puberuluous, especially at the
nodes, sometimes glabrate or almost wholly glabrous; setae about equaling the
stipular sheath; leaf blades oblong to linear-lanceolate, mostly 2-5 cm. long,
sometimes longer, obtuse or acute, narrowed at the base into a short petiole or
subsessile, rough and scaberulous on the upper furface, villosulous or glabrate
beneath; flower heads about 1 cm. in diameter, terminal and axillary, the terminal
ones subtended by large leaflike bracts; hypanthium pilose, the calyx teeth subulate,
pilose; corolla white, about 2.5 mm. long; capsule hardly 1 mm. long, circumscissile
near the middle; seeds pale brown, minutely pitted.
At least one collection of this plant was reported by Captain
Smith from Alta Verapaz under the name Spermacoce latifolia
Aubl.
Mitracarpus rhadinophyllus (Rob.) L. Wms. Fieldiana, Bot.
29: 371. 1961. M. villosus var. glabrescens Griseb. Cat. PL Cub. 143.
1866. Borreria rhadinophylla Rob. Proc. Am. Acad. 45: 409. 1910
(type from British Honduras, Peck 180). M. glabrescens Urban,
Symb. Antill. 7: 551. 1913.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 125
Pine ridges of British Honduras, and doubtless extended into
Pete"n or Alta Verapaz. Western Mexico; Nicaragua; Cuba.
Plants perennial, erect, 60 cm. high or less, glabrous almost or quite throughout,
the stems sometimes short-pilose at the nodes; stipule sheaths with usually 3 setae 6
mm. long or less; leaves narrowly lanceolate to almost linear, 3-9 cm. long, 1 cm.
broad or usually much narrower; fruiting heads about 1 cm. thick; lateral sepals 2-2.5
mm. long, narrowly lanceolate, acute; corolla white, 3.5 mm. long, the lobes half as
long as the tube, glabrous.
Mexican specimens of this species have been determined at Kew
as M. schizangium DC., which was described from Mexico. In spite
of De Candolle's description of M. schizangium as a perennial, a
photograph of the type indicates that probably it is M. hirtus, and
it is certainly not the plant here treated as M. rhadinophyllus. The
type consists of the upper part of a stem and there is no obvious
basis for ascribing a perennial habit to the plant.
MORINDA Linnaeus
Shrubs or trees, erect or often scandent, glabrous or pubescent, the branches
terete or obscurely tetragonous; stipules connate with the petioles to form a sheath;
leaves opposite or ternate, herbaceous; flowers perfect, white, connate by the calyces
to form a dense head, the heads pedunculate, axillary or terminal, commonly
solitary; hypanthium urceolate or hemispheric, the calyx short, truncate or obscurely
dentate, persistent; corolla funnelform or salverform, the tube short, the throat
glabrous or pilose, the limb 5-lobate, the lobes valvate in bud, coriaceous; stamens 5,
inserted in the corolla throat, with short filaments; anthers dorsifixed, linear or
oblong, obtuse, included or exserted; ovary normally 2-celled or 4-celled, the style
included or exserted, glabrous or pilose, with 2 short or elongate, linear branches;
ovules solitary, affixed to the septum below the middle, ascending; fruit a syncarp,
succulent, containing numerous nutlets, these cartilaginous or osseous, 1-seeded; seeds
obovoid or reniform.
About 50 species, chiefly in the Old World tropics, only a few
known from America. No other species have been found in Central
America.
Leaves short-pilose or scaberulous beneath.
Leaves short-pilose beneath, mostly 2-4 cm. broad; rough on the upper surface.
M. yucatanensis.
Leaves merely scaberulous beneath, 4-7 cm. broad, smooth on the upper surface.
M. asperula.
Leaves glabrous beneath, sometimes barbate in the axils of the nerves.
Leaves elliptic, mostly 5-9 cm. broad; corolla 15 mm. long M. panamensis.
Leaves mostly oblong or lance-oblong and 1.5-5 cm. broad; corolla 7-10 mm. long.
M. royoc.
126 FIELDIANA: BOTANY, VOLUME 24
Morinda asperula Standl. Field Mus. Bot. 22: 54. 1940.
Belicia hoffmanioides Lundell, Contr. Univ. Mich. Herb. 8: 87. 1942
(type from British Honduras, Gentle 3945).
In mixed forest or in pine forests, the type collected by
Standley 72368 near Quirigua, Izabal. Mexico (Chiapas on Atlantic
side); British Honduras.
A shrub or tree 3-8 m. high, the trunk reported to attain a diameter of 35 cm. in
Chiapas, the branchlets densely and minutely scaberulous; stipules broadly oval-
ovate, 10-14 mm. long, broadly obtuse or rounded at the apex; petioles 5-10 mm. long
or more elongate, the blades subelliptic or oblong-elliptic, 11-18 cm. long, 4-7.5 cm.
broad, acuminate or long-acuminate, acute at the base, glabrous above or nearly so
and smooth to the touch, densely and minutely scaberulous beneath, rough to the
touch, densely barbate in the nerve axils; peduncles 1.5 cm. long or shorter, the
flower heads (excluding the corollas) 8 mm. in diameter; calyx truncate, minutely
puberulent; corolla white, 1 cm. long, densely puberulent; fruit about 1.5 cm. in
diameter.
Closely related to M. panamensis, but differing in pubescence
and in the smaller corollas. The type specimen of Belicia
hoffmannioides, the type of the genus Belicia, is in fruit and also
bears flower buds. It differs somewhat from other material of the
species in having the inflorescences sessile or on very short
peduncles. It is believed, however, that it is only an individual
variant of Morinda asperula.
Morinda panamensis Seem. Bot. Voy. Herald 136. 1854.
Canche (Alta Verapaz).
Swamps, pastures, and thickets, 500 m. or lower; Alta Verapaz;
Izabal. Mexico (Oaxaca and Tabasco); British Honduras along the
Atlantic coast to Panama; Colombia.
An almost glabrous shrub or tree, 6 m. tall or less, the trunk 10 cm. or more in
diameter; stipules 1 cm. long or larger, rounded or very obtuse at the apex; leaves
short-petiolate, blackish when dried, elliptic or oblong-elliptic, 10-18 cm. long, 4-10
cm. broad, acute or acuminate, barbate beneath in the axils of the nerves, elsewhere
glabrous or nearly so, smooth to the touch; peduncles stout, 1-3 cm. long, the heads
6-8 mm. in diameter; corolla white or pinkish, 1.5 cm. long, glabrous outside or nearly
so; fruiting heads 1-1.5 cm. in diameter, greenish.
The flowers are sweet-scented. The bark is thin and dark
brown, the wood deep yellow, moderately hard. In the Isthmus of
Tehuantepec it is used for railroad ties. Vernacular names reported
are "yellow-wood" and "turkey victuals" (British Honduras); "palo
de peine" (Tabasco); "calabaza" (Oaxaca); "concha de huevo"
(Honduras).
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 127
Morinda royoc L. Sp. PL 176. 1753.
Collected at various stations in British Honduras, and
doubtless extending into Pete"n, perhaps also into Izabal. Nicaragua;
Panama; West Indies; Colombia and Venezuela.
An erect or clambering shrub, sometimes climbing to a height of 6 m., with
slender branches; stipules caudate from a broad base, about 2 mm. long; petioles 5-8
mm. long, the blades linear-lanceolate to lance-oblong, commonly 5-9 cm. long and 1-
2.5 cm. broad, sometimes larger, acute, narrowed to the base, barbate beneath in the
nerve axils, otherwise glabrous or practically so; peduncles 2-5 mm. long, the heads
about 1 cm. in diameter, few-flowered; hypanthium 3 mm. long, the calyx one-third
as long; corolla white or pinkish, about 8 mm. long; anthers short-exserted; fruit
globose, sometimes as much as 2 cm. broad but usually smaller.
Morinda yucatanensis Greenm. Field Mus. Bot. 2: 262. 1907.
Pihuela. Figure 58.
Pete"n and Izabal, at or little above sea level, in pine forest or
savannas, usually on limestone. Extending to British Honduras,
Mexico (Yucatan and Campeche).
A shrub, erect or spreading or clambering, the branchlets hirtellous; stipules 3
mm. long, cuspidate from a broad base; leaves short-petiolate, lance-oblong to
narrowly lanceolate or oblong-elliptic, mostly 6-10 cm. long and 1.5-4 cm. broad,
acute or acuminate, acute at the base, minutely scaberulous and rough on the upper
surface, hirtellous or short-pilose beneath, often barbate in the nerve axils; peduncles
short or none, the heads about 1 cm. in diameter; hypanthium and truncate calyx
densely hirtellous; corolla white, minutely hirtellous, scarcely more than 5 mm. long;
fruit yellowish at maturity, 1-1.5 cm. in diameter, juicy.
The Maya names are reported as "xhoyoc," "hoyoc,"
"xoyencab" and "xhoyac." The plant is said to have been used by
the Mayas of Yucatan for dyeing. The fruits are rubbed on warts to
remove those growths.
NERTERA Banks & Solander
Slender creeping perennial herbs, rooting at the nodes, glabrous or sparsely
pilose; stipules minute, connate with the petioles to form a sheath, 2-dentate or
entire; leaves small, opposite, sessile or petiolate, ovate-lanceolate to rounded-ovate;
flowers minute, axillary, sessile, mostly perfect; hypanthium ovoid, the calyx truncate
or short-tubular and 4-5-dentate, persistent; corolla tubular or funnelform, with
glabrous throat, the 4 lobes valvate in bud; stamens 4, inserted at the base of the
corolla, the filaments filiform; anthers basifixed, oblong, apiculate, exserted; ovary 2-
celled, the style branches 2, distinct at the base, filiform, long-exserted, pilose; ovules
solitary, erect from the base of the cell; fruit red, drupaceous, juicy, ovoid or globose,
containing 2 nutlets, these plano-convex, cartilaginous.
128 FIELDIANA: BOTANY, VOLUME 24
Half a dozen species are known, in the mountains of Hawaii,
the East Indies, Australia, New Zealand, and in America from
Mexico to southern South America. In America a single species is
known, confined to temperate regions of the tropical mountains.
The generic name Nertera has been conserved over Gomozia of
Mutis.
Nertera granadensis (L. f.) Druce, Bot. Exch. Club of British
Isles, Kept, of 1916: 637. 1917. Gomozia granadensis L. f. Suppl. PL
129. 1781. Nertera depressa Banks & Sol. in Gaertn. Fruct. et Sem.
PI. 1: 124. 1788. Hiedrilla (fide Aguilar).
Widely distributed in the higher mountains, 2,000-3,400 m.,
usually on banks in dense forest, often among mosses, most
plentiful in damp cold coniferous forests, especially those of
Cupressus, sometimes in alpine meadows; El Progreso; Zacapa
(Sierra de las Minas); Jalapa; Guatemala (Volcan de Pacaya);
Chimaltenango; Solola; Quiche; Huehuetenango; Totonicapan;
Quezaltenango; San Marcos. Central Mexico to Panama; extending
far southward along the Andes of South America.
Plants creeping and often forming dense mats, almost glabrous, the stems much
branched, rooting at the nodes; leaves mostly about 5 mm. long and often slightly
wider, slender-petiolate, ovate to ovate-rounded, obtuse, puncticulate above; corolla
very small, white, inconspicuous; berries 3-4 mm. long, orange-red or bright red, juicy.
A rather handsome plant when covering mossy banks or old
logs. In general appearance it much resembles Mitchella repens, but
is smaller in all parts.
OLDENLANDIA Linnaeus
Small, annual or perennial herbs, mostly dichotomous-branched, glabrous or
pubescent; stipules small, acute or acuminate, sometime setiferous; leaves opposite,
sessile or petiolate, herbaceous; flowers small, in dichotomous, axillary and terminal
panicles or cymes, or sometimes solitary in the leaf axils; hypanthium turbinate or
subglobose, the calyx 4-parted, the lobes usually erect and remote in fruit; corolla
rotate or funnelform, the tube short or elongate, the throat usually glabrous, the limb
4-lobate, the lobes obtuse, valvate in bud; stamens 4, inserted in the corolla throat,
with short filaments, the anthers dorsifixed, generally exserted; ovary 2-celled, the
style slender, with 2 short linear obtuse branches; ovules numerous, rarely few, the
placentae attached at the base or rarely at the middle of the septum; capsule small,
usually membranaceous, terete or angulate, globose or turbinate, loculicidally
dehiscent at the apex or through its whole length, few-seeded; seeds angulate or
subglobose, the testa smooth or minutely granulate.
Eighty or more species in tropical regions, chiefly in the Old
World. About 13 are known from North America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 129
Flowers solitary in the leaf axils on long slender pedicels O. herbacea.
Flowers in few-flowered, axillary or terminal cymes.
Corolla inconspicuous, about 1 mm. long; plants annual O. corymbosa.
Corolla evident, 5-6 mm. long; plants perennial O. microtheca.
Oldenlandia corymbosa L. Sp. PL 119. 1753. Figure 3.
Izabal (weed in lawn at Quirigua, alt. 70 m.); British Honduras,
and perhaps occuring in Pet6n. Southward to Panama and through
most of South America; tropical regions of the Old World.
An erect or decumbent annual, glabrous except on the leaf blades, usually much
branched, the branches 10-30 cm. long; stipules 1-1.5 mm. long, whitish, truncate,
bearing 1 or more long setae; leaves sessile or subsessile, linear or linear-lanceolate, 1-
3.5 cm. long, 1-5 mm. broad, acute to attenuate, usually scaberulous on the upper
surface, pale beneath, 1 -nerved; cymes mostly 3-flowered, the filiform pedicels 5-15
mm. long, often recurved; hypanthium less than 1 mm. long, the calyx lobes usually
longer, triangular-acuminate; corolla white or pale lavender, short-salverform,
equaling or slightly exceeding the calyx lobes; capsule subglobose, 1.5-2 mm. broad
and almost as long; seeds 0.2-0.3 mm. long, brown.
The plant often grows in sandy soil under coconut palms along
or near the seashore.
Oldenlandia herbacea (L.) DC. Prodr. 4: 425. 1830. Hedyotis
herbacea L. Sp. PL 102. 1753.
A weedy plant, usually growing in wet soil near streams,
sometimes on open banks or gravel bars, or along seashores, usually
at or little above sea level, but ascending in Chiquimula to 1,500 m.;
Izabal; Chiquimula (southeast of Concepcion de las Minas); Santa
Rosa; San Marcos. Southern Mexico to Panama, South America,
and West Indies; Africa; India; East Indies.
An erect or procumbent annual, glabrous throughout, usually much branched,
the stems commonly 20 cm. long or less; stipules 2-3 mm. long, 2-3-cuspidate; leaves
sessile, spreading or reflexed, usually linear, 2-7 cm. long, 1-8 mm. broad, long-
attenuate or acute, pale beneath, 1-nerved; flowers axillary, mostly solitary, on
filiform pedicels 8-25 mm. long, usually reflexed in fruit; hypanthium 1.5 mm. long,
the calyx lobes about as long, lance-acuminate, distant in fruit; corolla salverform,
white, about as long as the calyx lobes; capsule depressed- globose, 2.5-3 mm. long, 3-
4.5 mm. broad; seeds minute, pale brown, finely reticulate.
Both this and the preceding species appear to be of sporadic
occurrence in Guatemala, as elsewhere in Central America. They
are both inconspicuous and weedy plants.
130 FIELDIANA: BOTANY, VOLUME 24
Oldenlandia microtheca (Schlecht. & Cham.) DC. Prodr. 4:
428. 1830. Gerontogea microtheca Schlecht. & Cham. Linnaea 5:
169. 1830.
Moist or wet, rocky places, sometimes, at least, on limestone,
300-1,150 m.; Alta Verapaz (near Cubilgiiitz); Huehuetenango
(Finca Soledad, southeast of Barillas). Southern Mexico.
A very slender perennial, erect or decumbent, sometimes rooting at the lower
nodes, 10-40 cm. high, branched, the stems glabrous or puberulent at the nodes;
stipules 1-2 mm. long, the margins fimbriate, the lobes mostly gland-tipped; leaves on
short slender petioles, ovate or lanceolate, 1-4.5 cm. long, acute or attenuate, acute or
acuminate at the base, glabrous or scaberulous on the upper surface, slightly paler
beneath, glabrous or scaberulous; flowers in terminal and axillary cymes, the cymes
slender-pedunculate, few-flowered, lax, the pedicels 3-12 mm. long, filiform, glabrous;
hypanthium less than 1 mm. long, the calyx lobes as long or longer, triangular-
lanceolate; corolla 5-6 mm. long, white, the lobes much shorter than the tube,
papillose inside; style exserted; capsule hemispheric, 2 mm. long and slightly broader,
glabrous; seeds minute, angulate, brownish black.
PALICOUREA Aublet
Shrubs or small trees, glabrous or pubescent; stipules persistent, small or large,
usually united below to form a short sheath, bilobate, the lobes broad or narrow,
erect; leaves opposite or ternate, petiolate, usually somewhat coriaceous; flowers
small, cymose, corymbose, or paniculate, the panicles often thyrsiform, the
inflorescence terminal, its branches usually colored or pale, often yellow or reddish;
calyx small, dentate; corolla tubular, with very short lobes, the tube usually elongate
and slightly curved, generally somewhat dilated on one side at the base; ovary 2-
celled, the style simple, with 2 short branches, the cells 1-ovulate, the ovules attached
basally, erect; fruit drupaceous, commonly 2-celled, containing 2 nutlets, these
coriaceous, costate dorsally, plane or excavate on the inner surface.
A large and difficult genus with perhaps more than 250 species,
most abundant in the Guayana highlands and adjacent regions of
South America. There are about 30 species in North America, most
of them in Costa Rica.
Leaves ternate, rough to the touch on both surfaces P. triphylla.
Leaves opposite, smooth.
Corolla puberulent.
Inflorescence thyrsoid-paniculate; leaves mostly 7-14 cm. broad; corolla yellow
or orange P. guianensis.
Inflorescence corymbiform; leaves mostly 2-4 cm. broad; corolla white....P. seleri.
Corolla glabrous.
Corolla white or whitish.
Calyx usually much longer than the hypanthium; corolla 15-16 mm. long.
P. leucantha.
Calyx shorter than the hypanthium; corolla about 20 mm. long ..P. macrantha.
Corolla orange or orange-yellow.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 131
Corolla about 8 mm. long; stipular sheath very short, shorter than the lobes.
P. crocea.
Corolla 12-15 mm. long; stipular sheath well developed, usually equaling or
longer than the lobes P. galeottiana.
Palicourea crocea (Swartz) R. & S. Syst. Veg. 5: 193. 1819.
Psychotria crocea Swartz, Prodr. Ind. Occ. 44. 1788.
Wet forest or thickets of Peten and the North Coast, at or little
above sea level; Peten (La Libertad); Izabal. Mexico (Campeche,
Tabasco), and British Honduras to Panama, southward to Bolivia;
West Indies.
A shrub or small tree 1-5 m. tall, glabrous or nearly so, the branches green,
subterete or obtusely quadrangular; stipules only 2-3 mm. long; leaves opposite, on
petioles 5-15 mm. long, ovate-lanceolate to lance-oblong or elliptic-oblong, 6-17 cm.
long, 2.5-6.5 cm. broad, acuminate or long-acuminate, acute or attenuate at the base,
with 8-12 pairs of lateral nerves; panicles mostly 5-10 cm. long and 3-5 cm. broad,
thyrsiform, the branches reddish, suberect or strongly ascending, the bracts filiform,
the pedicels slender, usually longer than the calyx, dark orange; calyx slightly shorter
than the hypanthium, less than 1 mm. long, the lobes rigid, with thick margins, ovate
or lanceolate-ovate, acute; corolla red or orange-red, about 8 mm. long, the lobes 1.5
mm. long; fruit ovoid, purple-black, 4.5 mm. long.
Palicourea galeottiana Mart. Bull. Acad. Brux. 11, pt. 1: 136.
1844; L. Wms. Phytologia 26: 492. 1973. Psychotria mexicana Willd.
ex Roem. & Schult. Syst. Veg. 5: 189. 1819. Palicourea mexicana
Benth. ex Oerst. Vid. Medd. Kjoebenhavn 1852: 37. 1853, not
Psychotria mexicana Willd. Raxcuac (Alta Verapaz, Quecchi);
chalchipin (fide Aguilar). Figure 49.
Widely distributed in the mountains, usually in wet, mixed or
pine forest, sometimes in thickets, 800-2,400 m., common in the
western highlands; Alta Verapaz; Baja Verapaz; Zacapa;
Chiquimula; reported from Sacatepequez; Chimaltenango; Solola;
Suchitepequez; Quiche"; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico; Honduras to Panama.
A shrub or small tree 2-7 m. high or sometimes even larger, glabrous throughout
or nearly so, the branches slender, subterete, green; stipules short, the lobes short or
elongate, narrowly triangular to subulate, often much shorter than the tube; leaves
stiff, short-petiolate, oblong or elliptic-oblong, 7-20 cm. long, 3-6 cm. broad,
acuminate, acute at the base, usually almost glabrous but more or less pilose along
the costa; panicles thyrsiform or corymbiform, usually almost as broad as long, dense
or lax and many-flowered, the branches dull red or yellowish, most of the flowers
slender-pedicellate; calyx lobes scarcely 1 mm. long, rounded and usually with thin,
undulate or irregular margins; corolla pale yellow or bright yellow, commonly 12-15
mm. long, glabrous; fruit purple-black, ovoid, about 5 mm. long.
132 FIELDIANA: BOTANY, VOLUME 24
This is a rather handsome and showy shrub or small tree,
particularly abundant in the pine forests of Coban, and almost
equally so in the wet mixed forests of Quezaltenango and San
Marcos. Palicourea mexicana "form, vel var." angustifolia Loes.
(Verh. Bot. Brandenb. 65: 113. 1923; type collected between
Trinidad and Rosario, Distr. Nenton, Seler 3048) is described as
having lanceolate or elliptic-lanceolate leaves, but from description
does not appear to be essentially different from the usual form of P.
galeottiana.
Palicourea guianensis Aubl. PI. Guian. 173, t. 66. 1775.
Psychotria guianensis Rusby, Mem. Torr. Bot. Club 3, pt. 3: 48.
1893. Bitcul (PetSn, Maya, fide Lundell).
Wet forest or thickets, lowlands of Peten and Alta Verapaz, 350
m. or less; doubtless also in Izabal. Mexico to Brazil at low
elevations.
An almost glabrous shrub or small tree, commonly 2-3 m. high, the branches
stout, subterete, constricted at the nodes when dry; stipules 8-10 mm. long, biparted,
the lobes lanceolate or ovate, obtuse; leaves opposite, large, on short stout petioles,
rather thin, elliptic-oblong to ovate or elliptic, 16-30 cm. long, 7-14 cm. broad,
acuminate, acute to almost rounded at the base, glabrous or nearly so; panicles
thyrsiform, dense and many-flowered, long-pedunculate, usually about 10 cm. long
and 7 cm. broad, the stout branches ascending or spreading, glabrous or puberulent,
the bracts subulate, inconspicuous, the flowers pedicellate; calyx lobes minute,
deltoid, obtuse or acute; corolla deep yellow or orange- red, 10-18 mm. long, densely
and minutely furfuraceous-tomentulose or glabrate; fruit juicy, purplish black, ovoid,
4-5 mm. long.
Palicourea leucantha Donn.-Sm. Bot. Gaz. 48: 295. 1909.
Moist or wet, mountain forest, 1,500-2,000 m.; endemic; Alta
Verapaz (type collected along the road between Cob&n and Tactic,
Tuerckheim 8400); Huehuetenango (Sierra de los Cuchumatanes).
An almost glabrous shrub 3 m. high, the branches obtusely tetragonous; stipule
lobes linear-lanceolate, 4-5 mm. long, about equaling the sheath; leaves on petioles
1.5-3 cm. long, opposite, elliptic-oblong, 14-17 cm. long, 4-7 cm. broad, acuminate at
each end, minutely puberulent or glabrous beneath; inflorescence thyrsiform, 13-18
cm. long, 5-7 cm. broad at the base, the bracts and bractlets linear- lanceolate, 4-9
mm. long, the pedicels 3-9 mm. long; calyx lobes unequal, oblong-ovate, 2-4 mm. long,
3-nerved, much longer than the hypanthium, at least in anthesis; corolla white, 15-16
mm. long; anthers 2 mm. long.
Palicourea macrantha Loes. Verh. Bot. Ver. Brandenb. 65:
113. 1923.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 133
Type from Huehuetenango, wet virgin forest near
Yalambohoch, Distr. Nenton, Seler 2871.
A glabrous shrub, the branches subterete; stipu'es scarcely 2 mm. long, equaling
the sheath, deltoid, subobtuse; leaves opposite, on petioles 1.5-3 cm. long, oblong,
lance-oblong, or elliptic-oblong, 13-18 cm. long, 3.5-6.5 cm. broad, acuminate or acute,
cuneate at the base, the lateral nerves 10-14 on each side; inflorescence corymbiform,
about 10 cm. long, on a peduncle 7 cm. long, the pedicels 6-10 mm. long, pale yellow;
hypanthium 2 mm. long, the calyx cupular, 1.5 mm. long, 5-dentate; corolla whitish,
almost 2 cm. long, the lobes 3 mm. long.
Palicourea seleri Loes. Verb. Bot. Ver. Brandenb. 65: 114.
1923.
Wet mixed mountain forest, 2,800-3,000 m. or lower;
Chimaltenango (Santa Elena); Huehuetenango (types collected in
Distr. Nenton, near Yalambohoch and between Trinidad and
Rosario, Seler 2676, 3049); San Marcos (Volcan de Tacana). Mexico
(Chiapas).
A shrub or small tree, 4.5 m. high or less, apparently densely branched, the
branchlets terete, densely puberulent or glabrate; stipules 2-5 mm. long, puberulent,
the lobes subulate; leaves small, on petioles 5-17 mm. long, oblong-lanceolate to
ovate-lanceolate or lanceolate, 6-15 cm. long, 1.5-4 cm. broad, acuminate or acute,
acute at the base or attenuate, thin, puberulent above along the costa, deep green,
paler beneath, densely puberulent on the costa and veins, the lateral nerves about 10
pairs; inflorescence cymose, as broad as long or broader, lax, with few or many
flowers, the branches puberulent or pilosulous, the bracts linear, 1 cm. long or less,
the pedicels 1-2 cm. long or shorter; hypanthium scarcely 1 mm. long, minutely
puberulent, the calyx cupuliform, 0.5 mm. long, 5-dentate; corolla 9-11 mm. long,
lavender and white outside, white within, puberulent, the ovate lobes 1.5 mm. long;
fruit about 4 mm. long.
Palicourea triphylla DC. Prodr. 4: 526. 1830. Psychotria
triphylla Muell.-Arg. in Mart. Fl. Bras. 6, pt. 5: 233, t. 32. 1881.
Wet thickets at or little above sea level, north coast, sometimes
in Manicaria swamps or at the edges of savannas; Alta Verapaz;
Izabal. British Honduras, southward along the Atlantic coast to
Panama, and extending to Bolivia and Brazil.
A stout shrub 1-3 m. high, the branches terete, glabrous or nearly so; stipule
lobes linear, 6-10 mm. long; leaves large, ternate, on very short, stout petioles, oblong
or obovate-oblong, mostly 10-23 cm. long and 4-7 cm. broad, long-acuminate,
commonly acute at the base, puberulent, especially beneath, or finally glabrate,
rough to the touch; inflorescence long-pedunculate, thyrsoid-paniculate, dense and
many-flowered, 6-11 cm. long or larger, short-pilose, the bracts linear-subulate, the
branches orange or yellowish; flowers pedicellate, the calyx red, its teeth obtuse;
corolla yellow, furfuraceous-puberulent, 12-15 mm. long; fruit purple-black, ovoid, 5
mm. long.
134 FIELDIANA: BOTANY, VOLUME 24
PENTAGONIA Bentham
Shrubs or small trees simple or branched, the branchlets terete or obtusely
tetragonous, glabrous or pubescent: leaves large, opposite, sessile or petiolate,
coriaceous, entire or pinnatifid, finely striolate-lineolate between the veins, the
petioles sometimes auricula te; stipules large, elongate, deciduous; inflorescence
axillary, sessile or pedunculate, cymose-corymbose, with few to many flowers, the
flowers large, red or yellow, bracteate; hypanthium turbinate or campanulate; calyx
spathaceous or 5-6-lobate, persistent, the lobes equal or unequal, obtuse; corolla
funnelform or tubular, coriaceous, the tube glabrous within or villous at the base of
the stamens, the throat glabrous, the limb 5-6-lobate, with short valvate lobes;
stamens 5-6, inserted below the middle of the corolla tube, the filaments equal or
unequal, usually villous at the base; anthers dorsifixed, ovate or oblong, obtuse,
included; ovary 2-celled; style included, its branches linear-oblong, obtuse; ovules
numerous, the placentae adnate to the septum; fruit baccate, subglobose, thick, 2-
celled; seeds numerous, obtusely angulate, with thin testa.
About 20 species in tropical America. Eight are known from
Central America, chiefly in Panama and Costa Rica. The genus is
noteworthy for the very large leaves, which exhibit a fine striolation
on both surfaces in the dry state. In some species the blades are
deeply pinnatifid, a condition occurring regularly in no other
American genus of the Rubiaceae. The young leaves and the large
caducous stipules often are brilliantly colored in shades of red or
purple.
Pentagonia macrophylla Benth. Bot. Voy. Sulph. 105, t. 39.
1845. Watsonamra macrophylla Kuntze, Rev. Gen. 302. 1891. W.
donnell-smithii Standl. Contr. U. S. Nat. Herb. 17: 442. 1914. P.
donnell-smithii Standl. Journ. Wash. Acad. Sci. 17: 170. 1927.
Figure 27.
Dense wet forest, at or little above sea level; Izabal. Ranging
southward to Panama.
A shrub or tree, sometimes 9 m. high, simple or branched, the branchlets stout,
finely sericeous or glabrate; stipules ovate, about 5 cm. long and half as broad,
acuminate or attenuate, sericeous outside, glabrous within; petioles naked, 5-9 cm.
long, the blades oval to oblong-elliptic, 15-55 cm. long and 10-30 cm. broad or even
larger, acute, attenuate to rounded and short-decurrent at the base, glabrous above,
minutely sericeous beneath along the veins; cymes dense, sessile or short-pedunculate,
with few to many flowers, the thick pedicels 4-6 mm. long, tuberculate, bracts
caducous; hypanthium 6-8 mm. long, densely sericeous; calyx 1-1.5 cm. long, the
lobes oval or suborbicular, rounded or obtuse at the apex; corolla yellow, minutely
sericeous outside, the tube 2.5 cm. long, 2.5-6 mm. broad, the lobes lance-oblong to
broadly ovate, 5-7 mm. long, acute or acutish; fruit globose, 2.5 cm. in diameter.
PINAROPHYLLON Brandegee
Low perennial herbs, sometimes suffrutescent at the base, pubescent; stipules
foliaceous, deciduous; leaves opposite, short-petiolate, membranaceous; flowers small,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 135
ebracteolate, the inflorescence axillary, the peduncles filiform, unilaterally few-
flowered; hypanthium turbinate; calyx 4-lobate, the lobes subequal, persistent;
corolla yellow or yellowish or purplish, subrotate, with very short tube, the throat
glabrous, the limb 4-lobate, the lobes spreading in anthesis, imbricate in bud, acute;
stamens 4, inserted at the base of the corolla tube; filaments short, compressed, the
anthers exserted, oblong-linear, dorsifixed near the base, obtuse; ovary 2-celled, the
style filiform, elongate, the stigma clavate or bilobate; ovules numerous, the
placentae peltately affixed to the septum; capsule small, turbinate or oblong-
turbinate, costate, membranaceous, dehiscent at the apex; seeds numerous, minute,
globose, punctate.
The genus consists only of the two species described here.
Inflorescence several-flowered, secund; leaf acuminate P- flavum.
Inflorescence mostly 1-flowered; leaf obtuse or acute P. bullatum.
Pinarophyllon bullatum Standl. N. Am. Fl. 32: 87. 1921.
Hoffmannia pusilla L. Wms. Fieldiana, Bot. 34: 117. 1972 (type
from Alta Verapaz, Pittier 348).
Alta Verapaz, 750-1,100 m. on limestone recks in wet forest,
endemic; type collected between Sepacuite" and Secoyocte*, H.
Pittier 348.
A small perennial, the stout stems 3-12 cm. long, erect or ascending, usually
simple, densely villous with long weak hairs, densely leafy at the apex; petioles 1 cm.
long or less; leaf blades obovate to oblanceolate, 3-8 cm. long, 1-2.5 cm. broad,
subacute to rounded at the apex, attenuate to the base, conspicuously bullate, green
above and sparsely long-villous, paler beneath and copiously villous along the veins
with weak multicellular hairs; peduncles numerous, filiform, flexuous, mostly 1-2-
flowered, the pedicels to 15 mm. long; hypanthium sparsely villous, narrowly
turbinate, 2 mm. long; calyx lobes 4 or 5, narrowly triangular, acute, about 1 mm.
long; corolla 3-4 mm. long, glabrous, described as purplish, the lobes lance-oblong,
acute; capsule narrowly oblong- turbinate, 5-7 mm. long, costate and striolate; seeds
minute, dark brown.
Pinarophyllon flavum Brandegee, Univ. Cal. Publ. Bot. 6: 71.
1914. Figure 9.
Wet and shaded ravines, 500-1,000 m. (type from Chiapas,
Purpus 6700) Mexico, to be expected in Guatemala.
Small perennial herbs 5-6 cm. long; leaves oblanceolate, acuminate or long
attenuate, 6-15 cm. long and 1.5-3 cm. broad, sparsely villous with multicellular hairs;
inflorescence a unilateral spike with several flowers, peduncle slender, to 5 cm. long;
calyx lobes 4, lance-triangular, acute, about as long as the hypanthium; corolla about
4 mm. long, yellow, glabrous or sparsely pilose, the lobes lance-oblong, acuminate,
about as long as the tube; capsule about 3 mm. long, broadly turbinate.
136 FIELDIANA: BOTANY, VOLUME 24
PITTONIOTIS Grisebach
Small unarmed trees, the branches terete or obtusely angulate; leaves
membranaceous, petiolate; stipules intrapetiolar, acuminate, persistent; inflorescence
of paniculate cymes, the flowers secund along scorpioid branches, ebracteolate;
flowers small, sessile; hypanthium oblong, the calyx 4-5-lobate, persistent; corolla
salverform, pubescent outside, the tube cylindric, pilose in the throat, the limb 5-
lobate, the lobes imbricate, 2 exterior; stamens 5, inserted in the corolla throat;
anthers dorsifixed near the base, exserted; stigma bilobate, exserted; fruits
drupaceous, elliptic-oblong, flesh very thin, 2-3-celled; the seeds cylindric, elongated,
osseous.
The genus, often referred to Antirhea, consists of the following
species.
Pittoniotis trichantha Griseb. Bonplandia 6: 8. 1858. An-
tirrhoea trichantha Hemsl. Biol. Cent. Am. Bot. 2: 42. 1881. Figure
39.
Known from a single specimen in Guatemala, in humid tropical
forest near Sayaxche, Peten, Ortiz 142. Panama; northern South
America.
Small tree to 8 m. tall and the trunk 20 cm. in diameter; stipules narrowly
triangular, 4-6 mm. long, acuminate, appressed pilose outside; leaves elliptic-oblong
to broadly ovate, abruptly short acuminate, appressed pilose becoming glabrescent
with age; inflorescence paniculate cymose, the sessile flowers secund along the
branches; hypanthium and calyx about 1.5 mm. long, sericeous; calyx lobes narrowly
triangular, about 0.5 mm. long; corolla sericeous outside, about 5 mm. long with the
lobes and tube about equal in length; anthers linear-oblong, about 1.5 mm. long,
exserted; fruit 5-6 mm. long, oblong, sparsely pilose.
The genus is to be expected in other countries between Panama
and Guatemala.
POGONOPUS Klotzsch
Trees or shrubs; stipules interpetiolar, small, cuspidate, tardily deciduous; leaves
opposite, petiolate, the blades mostly membranaceous; flowers showy, pedicellate, in
small cymes, these disposed in lax terminal panicles; hypanthium turbinate; calyx
short, 5-dentate, deciduous, one of the lobes expanded into a large petiolate
foliaceous bright-colored blade; corolla tubular, villous in the throat, the short limb
5-lobate, the lobes valvate in bud; stamens 5, inserted in the corolla tube, the slender
filaments glabrous; anthers versatile, linear-oblong, exserted; ovary 2-celled, the
slender style with 2 linear or oblong, obtuse branches; ovules numerous, the placentae
longitudinally adnate to the septum; capsule subligneous, areolate at the apex, 2-
celled, loculicidally bivalvate, many-seeded; seeds horizontal, crowded.
A genus of two or three species in tropical America, only one of
them reaching North America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 137
Pogonopus speciosus (Jacq.) Schum. in Mart. Fl. Bras. 6, pt.
6: 265. 1889. Macrocnemum speciosum Jacq. Hort. Schoenbr. 1: 19,
t 43. 1797. P. exsertus Oerst. Ame>. Centr. 17. 1863. Flor de pascua;
quina. Figure 2.
Dry or damp forest of the tierra caliente of both coasts, at 300
m. or less; often planted for ornament; Alta Verapaz (Senahu;
Pancajche); Santa Rosa; Suchitep^quez; Escuintla; San Marcos.
Mexico (Chiapas) to Panama; Colombia and Venezuela.
A shrub or small tree, about 6 m. high, the branches conspicuously lenticellate,
the branchlets puberulent or hirtellous; stipules triangular, 3-5 mm. long, cuspidate;
petioles 2 cm. long or less, the blades obovate to elliptic or ovate, 6-20 cm. long, 3-10
cm. broad, abruptly acuminate or attenuate, acute to long-attenuate at the base,
glabrous or puberulent above, sparsely or densely puberulent beneath; cymes few-
flowered, arranged in a broad leafy panicle, the bracts foliaceous or linear; pedicels
stout, 1 cm. long or shorter; hypanthium 4-5 mm. long, puberulent or densely
fulvous-sericeous; calyx about 1 mm. high, 4 of the lobes minute, cuspidate, the fifth
expanded into a large blade, this rounded-ovate, 1-5 cm. long, 1-4 cm. broad, on a
slender petiole 1-2.5 cm. long, rounded to acute at the apex, rounded and short-
decurrent at the base, purplish or bright deep red, palmately e> -.erved; corolla 2.5-3
cm. long, purple or deep red, densely or sparsely puberulent or sericeous, the tube 3-5
mm. broad, the deltoid-ovate lobes 3-4 mm. long, acute, puberulent within; stamens
short-exserted; capsule 5-7 mm. long, 5-6 mm. broad, coarsely lenticellate; seeds 0.5
mm. long, pale yellowish.
Called "chorcha de gallo" in El Salvador and "pascua
cimarron" in Mexico. A showy and handsome tree because of the
abundance of large and brightly colored calyx lobes. It is employed
in Guatemala as a domestic remedy for malaria.
PORTLANDIA P. Browne
Shrubs or trees, glabrous or pubescent, often resinous; stipules intrapetiolar,
connate with the petioles to form a sheath, usually persistent; leaves opposite or
verticillate, petiolate or sessile, the blades coriaceous or herbaceous; flowers large,
axillary or terminal, the pedicels often bracteate; hypanthium usually turbinate, the
calyx lobes 4-5, short or elongate, persistent; corolla large, subcampanulate to
funnelform or tubular-funnelform, the tube 5-angulate, glabrous in the throat, the
limb 4-5-lobate, the lobes reduplicate-valvate or subimbricate; stamens 4-5, inserted
at the base of the throat or at the base of the corolla tube, the filaments filiform,
pubescent; anthers basifixed, linear, included or short-exserted; ovary 2-celled, the
style filiform, the apex entire, bilobate or with 2 long branches; ovules numerous,
crowded on swollen placentae longitudinally adnate to the septum; capsule thick-
coriaceous, terete, costate, or angular, loculicidally bivalvate from the apex; seeds
numerous, usually compressed, angulate, the testa usually granulate.
A genus of about 18 species, chiefly in the West Indies. No
others are known from Central America.
138 FIELDIANA: BOTANY, VOLUME 24
Portlandia guatemalensis Standl. Journ. Wash. Acad. Sci.
18: 162. 1928. Figure 1.
Wet mixed forest, 750-1,500 m.; endemic; Alta Verapaz (type
from Quebrada Seca, Harry Johnson 282); Huehuetenango (Max-
bal).
A shrub or small tree, sometimes epiphytic, the branchlets glabrous; stipules 5-6
mm. long, broadly triangular, cuspidate, persistent; leaves opposite, the slender
petioles 1-2 cm. long, the blades oblong-elliptic, broadest at or near the middle, 9-16
cm. long, 3.5-6 cm. broad, acuminate, acute and decurrent at the base,
membranaceous, glabrous above, much paler beneath, domatiate and short-barbate
in the axils of the nerves, the lateral nerves about 7 on each side; inflorescences
axillary, long-pedunculate, about equaling the leaves, racemiform-paniculate, the
flowers clustered at the end of the rachis and in pedunculate lateral clusters, the
bracts leaflike, lanceolate or elliptic; pedicels 3-4 mm. long; hypanthium broadly
turbinate, 2-2.5 mm. long, the calyx lobes 5, distinct, linear-subulate, 1 cm. long,
green, glabrous; corolla white, funnelform, glabrous, 4.5 cm. long, the tube very short,
2.5 mm. broad at the base, the throat 2.5 cm. broad, the 5 lobes broadly ovate-
triangular, obtuse, 1.5 cm. long; stamens included, the linear anthers 8 mm. long;
capsules and seeds unknown.
No doubt this is a showy plant when in flower. It is somewhat
questionable whether it is properly referable to Portlandia.
Capsules are not known and might suggest other placing of the
plant. The genera Hintonia and Coutarea are possibilities.
POSOQUERIA Aublet
Trees or shrubs, glabrous or pubescent, with terete branches; stipules large,
intrapetiolar, deciduous; leaves opposite, usually large, petiolate, coriaceous;
inflorescence a terminal corymb; flowers perfect, very large, white, the pedicels
ebracteate; hypanthium obovoid; calyx short, 5-dentate, usually persistent; corolla
tube greatly elongate and slender, the throat scarcely dilated, glabrous or villous, the
limb gibbous in bud, 5-lobate, the lobes oblique, obtuse, contorted in bud; stamens 5,
inserted in the corolla throat, the filaments glabrous or pilose, erect or curved;
anthers basifixed, linear-oblong, acute, pilose, exserted, the connective produced and
bifid at the base; ovary 1-2-celled, with filiform style, the stigma small, bifid,
included; ovules very numerous, peltate, the placentae stipitate, bilamellate, parietal;
fruit baccate, often very large, globose or ovoid, fleshy, 1-2-celled; seeds large,
obtusely angulate, with membranaceous testa.
About 20 species in tropical America. The only other Central
American species is P. grandiflora Standl. of Costa Rica. The genus
is easy to recognize because of the very large, white flowers with
corolla conspicuously curved in bud.
Posoqueria latifolia (Rudge) R. & S. Syst. Veg. 5: 227. 1819.
Solena latifolia Rudge, PL Guian. 1: 26, t. 40 1806. P. coriacea
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 139
Mart. & Gal. Bull. Acad. Brux. 11, pt. 1: 240. 1844. Chintorol
(Izabal). Figure 29.
Wet forest or thickets, often along streams, on the coast of both
slopes, at or little above sea level, ascending in the Pacific slopes to
1,400 m.; probably in Pete"n; Alta Verapaz; Izabal; Suchitepequez;
Solola; Quezaltenango; San Marcos; Huehuetenango. Southern
Mexico; British Honduras to Panama; southward to Bolivia and
Brazil.
A glabrous shrub or tree, sometimes 12 m. high with a trunk 25 cm. in diameter,
the branches thick and stout; stipules oval or oblong, 8-15 mm. long, green, obtuse or
acute; petioles stout, 8-12 mm. long, the blades mostly oval to oblong, 8-25 cm. long,
4-12 cm. broad, acute or abruptly short-acuminate, obtuse or rounded at the base,
lustrous above; corymbs with few or many flowers, pedunculate, most of the flowers
pedicellate, fragrant; calyx and hypanthium 4-5 mm. long, the calyx shallowly lobate,
often ciliolate, the lobes rounded; corolla tube 12-16 cm. long, 2-4 mm. thick, the
lobes spreading or reflexed, oblong or narrowly oblong, 1.5-2.5 cm. long, obtuse;
anthers 6 mm. long; fruit globose, yellow, 4-5 cm. in diameter or larger, umbonate;
seeds irregularly angulate, about 1 cm. long, black or nearly so, dull.
Among the vernacular names recorded are "mountain guava,"
"snake-seed" (British Honduras); "cachito" (Honduras); "azucena";
"toronja" (Oaxaca); "fruta de mono"; "guayabo de mico" (Costa
Rica). The tree is a showy and very handsome one when in flower.
Some of the Guatemalan specimens, particularly those from the
west, are conspicuous for their unusually large and broad leaves,
thus resembling P. grandiflora of Costa Rica, but in that the
branchlets and leaves are pubescent. It is quite possible that when
flowers of this Guatemalan form are available, it will prove to be a
distinct species.
PSYCHOTRIA Linnaeus
Shrubs or small trees, rarely perennial herbs, the branches terete or somewhat
tetragonous; leaves chiefly opposite, membranaceous to coriaceous, usually petiolate;
stipules intrapetiolar, persistent or deciduous, often connate to form a sheath; flowers
small, perfect, usually white or yellowish, terminal or rarely axillary, small, bracteate
or ebracteate, variously arranged, the branches of the inflorescence usually green or
greenish, rarely brightly colored, the inflorescence not involucrate; hypanthium small,
the calyx persistent or deciduous; corolla funnelform, tubular, or subcampanulate,
the tube short or elongate, straight, the throat naked or barbate, the limb 5-lobate,
the lobes valvate in bud; stamens 5, inserted in the throat or mouth of the corolla,
the filaments short or somewhat elongate; anthers dorsifixed near the base, linear or
oblong, obtuse, included or exserted; ovary almost always 2-celled, the style short or
elongate, glabrous or pilose, the 2 short branches subulate or linear; ovules solitary in
the cells, erect from the base of the cell; fruit drupaceous, small, juicy, sometimes
didymous, smooth or costate, of 2 nutlets or separating into 2 cocci, 2-seeded; nutlets
smooth or costate dorsally, the ventral face plane or sulcate.
140 FIELDIANA: BOTANY, VOLUME 24
The largest genus of Rubiaceae, with probably 1,000 or more
species, widely distributed in tropical regions of both hemispheres.
Many other species are known from southern Central America. In
America the genus is best represented in South America, especially
in equatorial Brazil and in the Andes.
Different authors have treated this tribe of Rubiaceae very
differently. Mueller included in Psychotria such groups as
Palicourea, Cephaelis, and Geophila. Bremekamp in treating the
Rubiaceae of Surinam has segregated several small or large groups,
but their segregation contributes little or nothing toward
simplification of the highly complicated groups of species here taken
to constitute the genus Psychotria.
The most recent account of American Psychotrias is that of Dr.
Julian A. Steyermark (in Mem. N. Y. Bot. Card. 23: 406-717. 1972)
where an attempt is made to classify the neotropical Psychotrias
into two subgenera, subg. Psychotria and subg. Heteropsychotria.
Subgenus Heteropsychotria is further divided into some 13 sections.
Steyermark's paper covers the Guayana highland region and all
contiguous areas, perhaps the richest area of the world in
Psychotrias. The paper is especially useful since he often included
data on the entire range of species known from the Guayana
highlands.
Dr. Steyermark has placed Cephaelis, which we have
maintained, into Psychotria. Cephaelis, at least in our region, seems
to be a useful generic unit.
We are pleased to acknowledge the assistance of our colleague
Dr. Donald R. Simpson in this difficult genus.
Inflorescence axillary.
Flowers densely clustered in the leaf axils, the inflorescence equaling or shorter
than the petioles; leaves elliptic P- erecta.
Flowers in long-pedunculate panicles, the inflorescences usually much longer than
the petioles; leaves oblanceolate- linear to oblong or oblanceolate.
Leaves oblanceolate- linear, 2.5 cm. broad or less, on short petioles 1 cm. long or
shorter P. pleuropoda.
Leaves mostly oblanceolate to oblong and 3-10 cm. broad or broader, long
petiolate.
Fruit white; leaves thin and membranaceous, green beneath P. macrophylla.
Fruit red; leaves succulent, very pale beneath P- uliginosa.
Inflorescences mostly terminal or at least pseudoterminal.
Stipules caducous, entire or nearly so, brown or ferruginous, thin; plants generally
turning blackish or grayish when dried. Subgenus Psychotria.
Calyx deeply lobate, the lobes lance- linear P. horizontal*.
Calyx shallowly dentate with broad teeth, or truncate.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 141
Young branches densely pilose or villosulous with long or short, spreading
hairs.
Leaves glabrous beneath or merely puberulent along the costa, small, 2-5
cm. long P. parviflora.
Leaves hirsute or densely short pilose beneath, usually over most of the
surface, mostly 5-12 cm. long or larger.
Leaves mostly 6-10 cm. broad, pubescent P. nervosa var. rufescens.
Leaves 2-5 cm. broad.
Leaves acuminate or long acuminate, mostly 3-5 cm. broad, glabrous.
P. nervosa.
Leaves obtuse or subacute, mostly 2-3 cm. broad P. erythrocarpa.
Young branches glabrous or nearly so, sometimes minutely puberulent.
Flowers glomerate, the glomerules spicate, the spikes paniculate; leaf blades
long attenuate at the base, on very short petioles P. viridis.
Flowers glomerulate or in cymes, but never in spicate glomerules.
Panicles all or mostly sessile, often or usually becoming pseudoaxillary by
the elongation of the branch above the insertion of the panicle.
Stipules shallowly or deeply bilobate at the apex P. tenuifolia.
Stipules entire.
Lateral nerves of the leaves about 9 pairs; blades broadest above the
middle, conspicuously coriaceous P. oerstediana.
Lateral nerves of the leaves 12-20 pairs; blades mostly broadest at or
below the middle, herbaceous or subcoriaceous.
Stipules 1.5 cm. long or shorter; leaves 3-7.5 cm. broad.
Fruit conspicuously longer than broad, oblong or ellipsoid.
P. nervosa.
Fruit globose.
Leaves mostly 2-3.5 cm. broad, thin, sparsely barbate beneath
in the nerve axils P. aguilarii.
Leaves mostly 5.5-7.5 cm. wide, subcoriaceous, not barbate
beneath P. schippii.
Stipules 2-4 cm. long or larger; leaves 5-15 cm. broad or even larger.
Calyx about 2 mm. broad; leaves usually subcordate to obtuse at
the base, often abruptly contracted and decurrent, rarely
acute P. limonensis.
Calyx about 1 mm. broad; leaves acute or acuminate at the base.
P. yunckeri.
Panicles on long or short peduncles, rarely or never becoming axillary by
elongation of the branch.
Stipules rounded or very obtuse at the apex, entire; flowers all sessile or
nearly so.
Leaves sparsely barbate beneath in the nerve axils; panicles usually
with only 3 basal branches P. lundellii.
Leaves not barbate beneath; panicles usually with more than 3
verticillate basal branches P. carthaginensis.
Stipules acute, acuminate, or attenuate, sometimes bilobate at the apex.
Lateral nerves of the leaves usually 6-8 pairs; leaves small, mostly 2-3
cm. broad.
Flowers mostly slender pedicellate, leaves membranaceous.
P. graciliflora.
142 FIELDIANA: BOTANY, VOLUME 24
Flowers sessile or on short stout pedicels; leaves subcoriaceous.
P. fruticetorum.
Lateral nerves of the leaves mostly 10-15 pairs.
Flowers pedicellate.
Pedicels slender, about 2-5 mm. long: lowland species.
P. marginata.
Pedicels thicker, about 2 mm. long; montane species ...P. yunckeri.
Flowers all sessile or nearly so.
Branches of the panicles finely puberulent P. clivorum.
Branches of the panicles glabrous or hirsutulous.
Leaves mostly 2-3.5 cm. broad; branches of the panicles
glabrous P. altorum.
Leaves mostly 4-7 cm. broad; branches of the panicles
hirtellous or almost glabrous P. flava.
Stipules long persistent, usually green or greenish, variable in form, most often
deeply bilobate or connate into a truncate sheath which is produced into short
or elongate lobes, sometimes entire. Subgenus Heteropsychotria.
Plants epiphytic; leaves thick and fleshy, coriaceous when dry, the lateral nerves
obscure P. parasitica.
Plants terrestrial; leaves not fleshy, the lateral nerves usually conspicuous.
Young branches densely pilose or villous with longer or short, chiefly spreading
hairs.
Bracts of the inflorescence large and conspicuous, at least equaling the
calyx, sometimes much longer.
Leaves glabrous on the upper surface, 2-3 cm. broad P. steyermarkii.
Leaves hirsute or villous on the upper surface, mostly 4-10 cm. broad.
Inflorescence head-like or of few cymose heads; leaves densely soft
pilose or villous beneath P. purpusii.
Inflorescence paniculate-cymose; leaves thinly pilose or hirsute below.
Bracts in the cymules ovate or lanceolate-ovate P. calopogon.
Bracts in the cymules linear P. chrysocalymma.
Bracts of the inflorescence small and inconspicuous, often minute, shorter
than the calyx.
Corolla 13-14 mm. long, very densely villous with long spreading hairs.
P. heydei.
Corolla much smaller, villous with short hairs or glabrous.
Inflorescence recurved, at least in age; leaves glabrous or essentially so
on the upper surface.
Fruits and inflorescences spreading pilose pubescent with whitish hairs
P. pittieri.
Fruits and inflorescences puberulent or with short sordid spreading
hairs P. dispersa.
Inflorescence erect; leaves evidently pubescent on the upper surface.
Corolla about 1 cm. long; inflorescence corymbiform P. skutchii.
Corolla much less than cm. long; inflorescence thyrsoid-paniculate.
Lateral nerves of the leaves about 20 pairs; leaf blades narrowly
lance-oblong or oblong-oblanceolate P. pachecoana.
Lateral nerves of the leaves about 13-16 pairs; leaf blades mostly
elliptic to elliptic-oblanceolate P. orogenes.
Young branches glabrous or merely puberulent.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 143
Bracts of the inflorescence large and conspicuous, longer than the calyx;
inflorescence sometimes capitate, the heads solitary or paniculate.
Corolla 3-4 cm. long P. chiapensis.
Corolla less than 1 cm. long.
Branches of the inflorescence glabrous P. capitata.
Branches of the inflorescence hirtellous.
Primary branches of the inflorescence bracteate at the base.
Stipules bifid at the center, lobes not lateral P. brachiata.
Stipules truncate with elongated linear lateral lobes. ..P. izabalensis.
Primary branches of the inflorescence not bracteate at the base.
P. officinalis.
Bracts of the inflorescence small and inconspicuous, shorter than the calyx;
inflorescence never capitate.
Stipules large, often 2 cm. long, not bilobate P. grandis.
Stipules less, usually much less than 1 cm. long, usually bilobate.
Calyx truncate.
Stipule lobes caducous, membranaceous; calyx 1 mm. long or more;
leaves acute to very obtuse and abruptly short pointed.
P. microdon.
Stipules lobes persistent, stiff, subulate; calyx scarcely 0.5 mm. long;
leaves cuspidate-acuminate P. cuspidata.
Calyx evidently dentate.
Calyx tubular below, much exceeding the hypanthium; corolla 12-15
mm. long P. mombachensis.
Calyx cleft or dentate almost or quite to the base, usually shorter
than the hypanthium, rarely longer; corolla 8 mm. long or
usually much shorter.
Inflorescence thyrsiform, the lower and middle branches often
reflexed.
Lobes of the stipules short, ovate; leaves usually more or less
pubescent beneath; branches of the inflorescence green.
P. berteriana.
Lobes of the stipules elongate, subulate; leaves glabrose; branches
of the inflorescence reddish or purplish.
Leaves long attenuate at the base, with more than 10 pairs of
lateral nerves; petioles more than 1 cm. long; species of the
Pacific slopes P. oreodoxa.
Leaves abruptly contracted at the base, not attenuate, with
fewer than 10 pairs of lateral nerves; petioles less than 1
cm. long; species of the Atlantic lowlands P. patens.
Inflorescence corymbiform or paniculate but not thyrsiform, the
branches spreading or ascending.
Leaves mostly 7-15 cm. long; inflorescences rather large and
many-flowered.
Branches of the inflorescence glabrous or essentially so.
P. simiarum.
Branches of the inflorescence densely puberulent or hirtellous.
P. pubescens.
Leaves mostly 2-6 cm. long; inflorescences small and with very
few flowers.
144 FIELDIANA: BOTANY, VOLUME 24
Calyx lobes much longer than the hypanthium P. Ulacina.
Calyx lobes shorter than the hypanthium P. minarum.
Psychotria aguilarii Standl. & Steyerm. Field Mus. Bot. 23:
23. 1943. Pihtziquin (Aguilar).
Damp thickets or forest, at about 1,500 m.; endemic;
Guatemala; Chimaltenango (type collected between Chimaltenango
and San Martin Jilotepeque, Standley 80900).
A shrub 1-1.5 m. high, the branches glabrous; stipules caducous, about 1 cm.
long, membranaceous, ferruginous, glabrous, at first enclosing the terminal buds but
soon cleft on each side; leaves short-petiolate, membranaceous, on petioles 5-12 mm.
long, oblanceolate or oblong-oblanceolate, 8-12 cm. long, 2-3.5 cm. broad, narrowly
attenuate-acuminate, narrowly long-attenuate to the base and long-decurrent, green
and glabrous above, paler beneath, glabrous but domatiate in the nerve axils and
sparsely short-barbate, the lateral nerves about 11 on each side; inflorescence
terminal, sessile, lax and few-flowered, as much as 4.5 cm. long, few-rayed at the
base, the branches glabrous, the small bracts caducous, the flowers sessile or nearly
so, aggregate at the ends of the branches; calyx and hypanthium about 1 mm. long,
glabrous, the calyx truncate and remotely dentate, slightly shorter than the
hypanthium; corolla white, glabrous outside, 5 mm. long, not barbate in the throat,
the lobes 1.5 mm. long, obtuse, glabrous within; fruit deep red, globose, 6 mm. in
diameter, the nutlets coarsely granulate dorsally, ecostate.
Psychotria altorum Standl. & Steyerm. Field Mus. Bot. 23:
86. 1944. Huesito bianco.
Wet mixed mountain forest and thickets, 300-2,700 m.;
Chimaltenango; Solola; Suchitepe"quez; Quezaltenango (type from
Montana Chicharro, southeastern slopes of Volcan de Santa Maria,
2-4 miles south of Santa Maria de Jesus, Steyermark 34302); San
Marcos. Mexico (Chiapas); El Salvador.
A shrub 1.5-3.5 m. tall or even larger, the slender branches glabrous; stipules
caducous, distinct, about 12 mm. long, ferruginous, ovate or oblong-ovate, acuminate
or sometimes shallowly bilobate at the apex; leaves short-petiolate, herbaceous, on
petioles 5-10 mm. long, oblong-oblanceolate or narrowly oblanceolate, 9-12 cm. long,
2-3.5 cm. broad, acute or obtuse, narrowly long-attenuate to the base, glabrous, the
lateral nerves about 10 on each side, the nerve axils not barbate or domatiate;
inflorescence terminal, ovoid-paniculate, many-flowered, erect on a peduncle 1.5-5
cm. long, as much as 5.5 cm. long and 5 cm. broad, the basal branches about 4,
divergent or reflexed, glabrous, the bracts caducous, the flowers densely aggregate in
small cymes or umbels, sessile or short-pedicellate; calyx and hypanthium 2 mm.
long, the calyx campanulate, slightly longer than the thick hypanthium, remotely
dentate or subentire; corolla white, about 5 mm. long, glabrous outside, not barbate
in the throat, the lobes oblong, obtuse, recurved, a little more than half as long as the
tube, glabrous within; style short-exserted; apices of the anthers exserted; fruit pale
orange or red, subglobose, 6 mm. long, the nutlets coarsely costate dorsally.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 145
Psychotria berteriana DC. Prodr. 4: 515. 1830. P. platyphylla
DC. I.e. 517. P. crebrinervia Standl. Field Mus. Bot. 4: 343. 1929
(type from the Atlantic coast of Honduras).
Dense wet mixed forest or thickets, sometimes in pine forest,
ranging from sea level to about 1,500 m.; Pete"n; Alta Verapaz; Baja
Verapaz; Izabal; Huehuetenango. Southern Mexico; British
Honduras to Panama and Colombia; West Indies.
Usually a shrub of 2-4.5 m., sometimes a tree of 6 m., the slender brittle branches
glabrous or puberulent, sometimes hirtellous when young; stipules persistent, thin,
about 4 mm. long, bilobate, the short lobes ovate, acute; leaves large and thin, on
petioles 1.5 cm. long, lance-oblong to oblong-ovate, mostly 15-18 cm. long and 5-6.5
cm. broad, often smaller, long-acuminate, rounded to acute at the base and usually
abruptly short-decurrent, almost glabrous above but usually rough to the touch,
sparsely villosulous beneath or almost glabrous; panicles terminal, mostly long-
pedunculate, often recurving in age, generally thyrsiform but often pyramidal,
usually large, lax, and open, as much as 16 cm. long and broad, the branches densely
short-pilose, the flowers mostly short-pedicellate; bractlets small but relatively broad,
green, rather conspicuous; calyx minute and remotely dentate; corolla greenish
yellow, minutely pilose or almost glabrous, 10 mm. long or less; fruit black at
maturity, subglobose, 4-5 mm. in diameter, round cystoliths present.
The species as we have treated it here is quite variable one and
may include more than one species. In Guatemala the species is
found only on the Atlantic side but the elevations and the
ecological situations in which it occurs are rather too diverse.
Specimens from the Pacific highlands put here by the senior author
have been segregated by the junior author as Psychotria
standleyana.
Psychotria brachiata Swartz, Prodr. Veg. Ind. Occ. 45. 1788.
Moist or wet, mixed, lowland forest, 800 m. or less; Izabal; Alta
Verapaz (region of Cubilguitz); Huehuetenango. British Honduras,
and doubtless also in Peten at or little above sea level. Southern
Mexico to Panama, southward to Peru; West Indies.
A shrub 4 m. high or less, the branches green, usually glabrous; stipules
persistent, 5-8 mm. long, bilobate, the lobes obtuse or rounded; leaves on petioles 2
cm. long or less, oblong-obovate to oblong-elliptic, 10-15 cm. long, 6 cm. broad or less,
short-acuminate, acute to attenuate at the base, glabrous or nearly so, often pilose
beneath along the costa, the lateral nerves about 10 pairs, strongly curved;
inflorescence terminal, erect, long-pedunculate, thyrsoid-paniculate or racemiform,
rarely somewhat pyramidal, the branches opposite, subdivaricate, subtended at the
base by long narrow green bracts, the flowers sessile in small dense bracteate heads;
calyx remotely denticulate; corolla 6-8 mm. long, white or ochroleucous, villous in
the throat, the oblong lobes shorter than the tube; fruit blue at maturity, about 5
mm. long, crowned by the persistent calyx, the nutlets costate dorsally.
146 FIELDIANA: BOTANY, VOLUME 24
Psychotria calopogon L. Wms., Phytologia 28: 227. 1974.
Dense, rich lowland forests, 150-300 m.; endemic; Alta Verapaz;
Huehuetenango (type, Steyermark 49308).
Shrubs to 2.5 m. tall, the stems spreading pilose pubescent, probably glabrescent
with age, stipules lanceolate, bilobate, the apices scarious. Leaves elliptic to broadly
elliptic, acuminate, with about 20 pairs of secondary nerves these prominent below,
pilose on both surfaces, more so below and along the mid-vein, pale green above and
lighter below, the blade 12-30 cm. long and 3.5-12 cm. broad, the petioles 1-3 cm. long,
spreading pilose pubescent; inflorescence terminal, pedunculate, a rather loose, many-
flowered panicle with the lateral branches of capitulate, bracteate cymules; cymules
subtended by 3-5 lanceolate, acute or acuminate, pilose bracts mostly 4-5 mm. long
and 1.2-2 mm. broad; hypanthium and calyx about 2 mm. long, densely pilose, the
calyx lobes narrowly lanceolate, acute, about 0.7-0.8 mm. long; corolla white, mostly
4-5 mm. long, the tube narrowly cylindric and about 3.5 mm. long, the lobes ovate,
acute, pilose dorsally, 1-1.5 mm. long; style about as long as the tube, stigma
bipartite; stamens attached in throat of the corolla and the anthers exserted and
about 0.5 mm. long; fruit not known.
This species belongs in a complex surrounding P. pilosa R. & P.
of South America. Three or four related species are known from
Central America and Mexico.
Psychotria capitata Ruiz & Pavon, Fl. Peruv. 2: 59, t. 206, f.
a. 1799. P. inundata Benth. in Hook. Journ. Bot. 3: 229. 1841.
Palicourea stevensonii Standl. Trop. Woods 16: 42. 1928 (type from
Middlesex, British Honduras, Neil S. Stevenson).
Collected at various localities in British Honduras, and to be
expected in Izabal and Peten. Honduras; Nicaragua; Panama, and
southward to Bolivia and Brazil.
A glabrous shrub 1-3 m. high; stipules persistent, green, biparted, with long
subulate lobes, the stipules near the ends of the branches with much longer,
lanceolate lobes; leaves subcoriaceous, on short stout petioles, lanceolate to oblong-
elliptic or oval, mostly 11-16 cm. long and 5-6 cm. broad, acuminate or abruptly
short-acuminate, acute or obtuse at the base, the lateral nerves about 18 pairs,
prominent and pale beneath; inflorescence terminal, thyrsoid-paniculate, erect, rather
dense, the branches pubescent, opposite or verticillate, the primary ones without
bracts at the base, the flowers sessile or nearly so, the bracts subtending them
lanceolate or linear, pale green or whitish, conspicuous, about equaling the flowers;
calyx teeth minute, unequal; corolla white or ochroleucous, glabrous, 8-9 mm. long,
the lobes almost equaling the tube; fruit subglobose, 4 mm. long, coarsely costate,
black.
Psychotria carthaginensis Jacq. Enum. PI. Carib. 16. 1760.
P. foveolata Ruiz & Pav6n, Fl. Peruv. 2: 59, t. 207, f. b. 1799.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 147
Moist or dry forest or thickets, chiefly on the Pacific coastal
plain, at 400 m. or less; Zacapa; Escuintla; Suchitepequez; San
Marcos; probably along the whole Pacific coast. Southern Mexico
to Panama, southward to Bolivia and Argentina.
A shrub 1-2 m. high, glabrous or nearly so; stipules membranaceous, ferruginous,
caducous, less than 1 cm. long, obtuse or rounded at the apex; leaves short-petiolate,
commonly rather thick, lance-elliptic to oblong-obovate, broadest at or above the
middle, commonly 6-12 cm. long and 3-5 cm. broad, acute or acuminate or sometimes
obtuse, at the base acute to long-attenuate, glabrous beneath or rarely puberulent on
the nerves, often conspicuously domatiate in the nerve axils, the lateral nerves about
10 pairs; inflorescence terminal, erect, paniculate, usually long-pedunculate, generally
radiately branched from the base, with divergent branches, the branches glabrous, the
bracts caducous, the flowers sessile or nearly so; calyx obsoletely denticulate; corolla
white, glabrous or very minutely puberulent, 4-5 mm. long, with very short lobes;
fruit red, subglobose, 4 mm. long, the nutlets costate dorsally.
It is not altogether certain that the plants of Guatemala are
identical with those of Cartagena, Colombia. Neither is it certain
that only one species is involved in the Central American material.
The subgenus Psychotria is badly in need of revision. Half of the
species of Psychotria in Guatemala belong to this subgenus and
most of them present problems.
Psychotria chiapensis Standl. Contr. U. S. Nat. Herb. 23:
1390. 1926. Cephaelis tetragona Donn.-Sm. Bot. Gaz. 61: 376. 1916,
not Psychotria tetragona Seem. 1865-67 (type from Tuis, Costa
Rica). Palo de agua. Figure 47.
Wet forest or thickets, widely distributed, at 1,400 m. or lower;
Petdn; Alta Verapaz; Izabal; Escuintla; Guatemala; Solol6;
Suchitepe'quez; Retalhuleu; Quezaltenango; San Marcos;
Huehuetenango. Southern Mexico and British Honduras to
Panama; type from Chiapas.
A shrub or small tree, commonly 2-6 m. high, the branches glabrous or obscurely
villosulous; stipules small, persistent, bilobate, with broad obtuse lobes; leaves short-
petiolate, elliptic-oblong to oblong-oblanceolate, commonly 12-20 cm. long and 5-10
cm. broad, short-acuminate, acute or acuminate at the base, glabrous, the lateral
nerves about 12 pairs; inflorescence terminal, erect, pedunculate, the flowers
subcapitate, in large dense heads, these corymbose or cymose, the large broad green
bracts rounded at the apex; calyx 5-6 mm. long, puberulent, denticulate; corolla
white, almost glabrous, the slender tube as much as 3 cm. long, the narrow lobes 8
mm. long; fruit oval-globose, 1-1.5 cm. long, about 1 cm. broad, the 2 nutlets acutely
carinate dorsally, plane and not sulcate on the inner face.
Called "cassada" and "white wood" in British Honduras;
"yaxcanan" (British Honduras, Maya). The plant does not fall
148 FIELDIANA: BOTANY, VOLUME 24
satisfactorily in the genus Psychotria, and could be placed almost
equally well in Cephaelis. The corolla is much larger than in any
other Central American species of Psychotria.
Psychotria chrysocalymma L. Wins. Phytologia 28: 228.
1974.
Probably in the montane forest above 2,400 m.; El Progreso
(type from Volcan Sta. Luisa, Steyermark 43518). Endemic.
Shrubs to 2 m. tall or perhaps more, the branches densely and softly short pilose
pubescent; stipules persistent, with two lateral lanceiform lobes 2-3 mm. long. Leaves
short petiolate, elliptic, acuminate, pilose-pubescent or sparsely hirsute on both
surfaces, 12-15 cm. long and 4-5.5 cm. broad when mature, secondary nerves 11-13
pairs, petiole slender, 1-2 cm. long; inflorescences lateral, long pedunculate
subumbelliform cymes covered with yellowish or golden multicellular pubescence, the
peduncle about 6 cm. long, the cymes about 4 cm. long, the bracts subtending each
division of the inflorescence linear and acute 10, to 3 mm. long; flowers in each
cymule about 4-5, short pedicellate, subtended by bracts longer that the calyx;
hypanthium and calyx about 4 mm. long, densely short pilose, calyx lobes narrowly
lanceolate, acute, 1.5-2 mm. long; corolla white, tubular, widest above the middle,
pubescent outside especially above, 13-15 mm. long, the lobes short, oblong-
lanceolate, about 2 mm. long; style as long as the corolla, apex shortly bilobate;
stamens inserted in the throat of the corolla, included, anthers about 3 mm. long;
fruits narrowly ovoid, each carpel prominently 3-ribbed dorsally, sparsely to densely
pubescent, 5-6 mm. long.
Closely related to P. purpusii Standl. which is known from the
slopes of Volcan Tacumulco and in adjacent Chiapas.
Psychotria clivorum Standl. & Steyerm. Field Mus. Bot. 23:
87. 1944.
Wet mixed forest, 600-1,400 m., in the western highlands;
Suchitepe"quez (type from Finca Mocd, Skutch 2073); Quezal-
tenango (below Santa Maria de Jesus); San Marcos. Doubtless also
in Chiapas.
A shrub 1.5-4.5 m. high, the branches glabrous; stipules caducous, 1.5 cm. long or
less, oval or broadly ovate, membranaceous, ferruginous, glabrous, obtuse or rounded
at the apex and abruptly short-acuminate, the apex shallowly bilobate; leaves large,
short-petiolate, herbaceous, on petioles 1-2.5 cm. long, oblong-oblanceolate or
narrowly oblanceolate, 13-30 cm. long, 4.5-5 cm. broad, acuminate or long-acuminate,
gradually long-attenuate to the base, glabrous, the lateral nerves about 20 pairs;
inflorescence terminal, erect, on a peduncle about 5 cm. long, paniculate, many-
flowered, dense or rather lax, 4-7.5 cm. long and as much as 7 cm. broad, the basal
branches verticillate, densely and minutely puberulent, often reflexed, the small
bracts caducous, the flowers aggregate in small cymes, sessile or nearly so; calyx and
hypanthium 1-1.5 mm. long, minutely puberulent or glabrate, the calyx limb 1.5 mm.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 149
broad, remotely and shallowly dentate or subtruncate, the teeth broadly triangular;
corolla white, glabrous outside, scarcely 4 mm. long, the lobes reflexed, oblong-ovate,
obtuse, glabrous within, about half as long as the tube, the throat not barbate; fruit
red, oval-globose, 5-6 mm. long, the nutlets coarsely costate dorsally.
Psychotria cuspidata Bredem. ex R. & S. Syst. Veg. 5: 192.
1819.
Wet mixed forest, 200 m. or less; Izabal; Pete'n;
Huehuetenango. Southern Mexico and British Honduras to
Panama, southward to Peru and Brazil; West Indies.
A slender glabrous shrub, usually 1.5-2.5 m. high; stipules green, persistent, very
short, bicuspidate, usually appressed; leaves thin but rather stiff, short-petiolate,
bright green or yellowish green when dried, ovate to elliptic or oblong-elliptic,
commonly 11-16 cm. long and 6-8 cm. broad, cuspidate-acuminate, usually with a
somewhat curved acumination, acute to obtuse at the base, the lateral nerves about
14 pairs; inflorescence terminal, small, pedunculate, thyrsoid-paniculate or sometimes
corymbiform, much shorter than the leaves, rather few-flowered, commonly 1.5-3 cm.
broad, the few pale or reddish branches minutely puberulent or glabrous, not
bracteate at the base, the flowers partly sessile and partly on short stout pedicels;
calyx minute, truncate; corolla white or cream, glabrous or minutely puberulent, 5
mm. long or less; stamens exserted; fruit didymous-globose, smooth, black at
maturity.
A common shrub in forests of the Atlantic coast of Central
America.
Psychotria dispersa Standl. Journ. Wash. Acad. Sci. 18: 184.
1928.
Wet thickets and moist forest of the Pacific plains and slopes,
550-750 m.; Escuintla (between Rio Jute and Rio Pantale6n);
Suchitepequez (Pueblo Nuevo); to be expected in Peten and Izabal
since the species has been collected in British Honduras. Ranging
southward to Panama, the type collected near Tilaran, Guanacaste,
Costa Rica.
A slender shrub 1-2 m. tall, the branches green, short-pilose with spreading hairs;
stipule sheath 2.5-3 mm. long, persistent, the lobes linear, 3-6 mm. long; leaves small,
short-petiolate, oblong-elliptic to lance-oblong, 6-10 cm. long, 2-4 cm. broad, thin,
abruptly acuminate, at the base acute and often decurrent, glabrous above, paler
beneath, pilose on the nerves with short spreading hairs, the lateral nerves about 12
pairs; inflorescence terminal, paniculate, usually reflexed in fruit, pedunculate,
paniculate, 2-4 cm. long and broad, rather lax and open, the branches hirtellous, the
branches all subtended by long narrow green bracts, the flowers sessile or subsessile;
calyx less than 1 mm. long, the teeth triangular; corolla white, puberulent, 3-3.5 mm.
long; fruit blue, puberulent or glabrate, 3 mm. long, the nutlets costate dorsally.
150 FIELDIANA: BOTANY, VOLUME 24
Psychotria erecta (Aubl.) Standl. & Steyerm. Field Mus. Bot.
23: 24. 1943. Ronabea latifolia Aubl. PI. Guian. 134, t. 59. 1775, not
Psychotria latifolia Humb. & Bonpl. 1819. R. erecta Aubl. op. cit.
156. 1775. P. axillaris Willd. Sp. PL 1: 962. 1798. Appunia
parviflora Lundell, Contr. Univ. Mich. Herb. 4: 28. 1940 (type from
Silk Grass Creek Reserve, British Honduras, Gentle 2986).
Wet, mixed lowland forest or thickets, 180 m. or lower;
probably in Peten; Alta Verapaz; Izabal. British Honduras;
Nicaragua; Costa Rica; Colombia and the Guianas to Brazil and
Bolivia.
A sparsely branched shrub, usually 1-2 m. high, rarely a tree of 7 m. (?), glabrous
or nearly so; stipules persistent, subulate from a short broad base; leaves often
blackening in drying, subcoriaceous, on petioles 1.5 cm. long or less, oval to oblong-
elliptic or obovate, mostly 12-15 cm. long and 6-7 cm. broad, abruptly short-
acuminate, acute at the base, with about 9 pairs of lateral nerves, glabrous above,
very sparsely short-pilose beneath or almost glabrous; inflorescences all axillary, few-
flowered and headlike, equaling or shorter than the petioles, the short peduncle
densely pubescent; hypanthium glabrous, 1.3 mm. long; calyx cupular, 1.3 mm. long,
truncate or remotely denticulate; corolla white, glabrous outside, 6 mm. long, the
lobes shorter than the tube; fruit black, ovoid, 8 mm. long, glabrous; pyrenes sulcate
on the inner surface.
Psychotria erythrocarpa Schlecht. Linnaea 9: 595. 1834.
Cereza silvestre; pakexte (Huehuetenango); chichipin (fide
Aguilar); caquechpox (Huehuetenango, fide Seler).
Moist thickets or dry rocky slopes, 500-1,800 m.; El Progreso;
Zacapa; Chiquimula; Jalapa; Jutiapa; Guatemala (Estancia
Grande); Quiche; Huehuetenango. Southern Mexico; Honduras.
A densely branched shrub 1-3 m. high, the branches very leafy, densely pilose
with spreading hairs; stipules 2 cm. long or less, caducous, at first enclosing the
terminal buds, ferruginous, pilose, cleft along one side as the leaves unfold; leaves
herbaceous, short-petiolate, mostly oblanceolate-oblong, sometimes obovate, 3.5-10
cm. long, 1.5-3.5 cm. broad, obtuse or acute, sparsely or densely short-pilose above,
usually very densely and softly pilose beneath with slender pale hairs, the lateral
nerves about 10 pairs, conspicuous beneath; inflorescence terminal, on a long or
short peduncle, cymose-corymbose or paniculate, usually small and rather lax, the
flowers sessile or short-pedicellate, the branches densely short-pilose; calyx and
hypanthium densely short-pilose, the calyx subtruncate; corolla small, white,
puberulent or hirtellous, the lobes shorter than the tube, the throat not barbate; fruit
subglobose, 4-5 mm. long, red or deep red, pilose, the nutlets coarsely costate dorsally.
Psychotria flava Oerst. ex Standl. Journ. Wash. Acad. Sci. 17:
341. 1927.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 151
Wet thickets or mixed forest, widely distributed in Guatemala
at 1,500 m. or less; Pete'n; Izabal; Alta Verapaz; Zacapa;
Chiquimula (record doubtful); Suchitepequez (Finca Moca); Solola.
Southern Mexico and British Honduras.
A shrub or small tree, reported to attain a height of 7 m., but usually lower, the
young branches stout, glabrous; stipules caducous, thin, ovate-triangular or broadly
ovate, 1.5 cm. long or less, narrowed to the apex and shallowly bJlobate, the apical
lobes acute or acuminate, ferruginous-puberulent on the outer surface or sometimes
glabrous; leaves large, rather thick and often coriaceous, usually yellowish when
dried, on petioles 1-3 cm. long, narrowly oblanceolate-oblong to obovate-oblong,
rarely linear-oblanceolate, 10-30 cm. long, 2.5-13 cm. broad, obtuse to short-
acuminate, cuneate-attenuate or long-attenuate at the base, beneath minutely
hirtellous on the nerves and puberulent between them or often glabrous, the nerve
axils not domatiate or barbate, the lateral nerves about 20 on each side, or often
fewer; inflorescence terminal, erect, on stout peduncles 10 cm. long or less, cymose-
paniculate, the branches hirtellous or glabrate, the lower ones verticillate and often
reflexed, the flowers sessile; hypanthium and calyx usually densely and minutely
puberulent, the calyx scarcely 1 mm. long, truncate or obscurely repand-dentate, 1.5
mm. broad; corolla white or cream, 4.5 mm. long, glabrous outside, the lobes ovate,
obtuse, shorter than the tube; anthers usually exserted; fmit red, subglobose or
obovoid, 8-15 mm. long, glabrous, the nutlets plane on the inner face, coarsely costate
dorsally.
From Oaxaca, Mexico there are reported the local names
"tepecajete bianco," "marita," and "hoja lisa." The sapwood is
described as pale yellow, the heartwood as dark chocolate-brown.
Psychotria fruticetorum Standl. Journ. Arnold Arb. 11: 42.
1930. Rax-ac (Alta Verapaz, Quecchi).
Usually in wet thickets or forest, sometimes in open places,
ranging from sea level to about 1,600 m.; Peten; Izabal; Alta
Verapaz; El Progreso; Chiquimula; Chimaltenango. Mexico
(Campeche and Tabasco) to British Honduras and Honduras, the
type from Siguatepeque, Honduras.
A branched shrub 1-3 m. tall, the branches glabrous; stipules caducous, ovate or
lanceolate, ferruginous, mostly 3.5-5 mm. long, acuminate; leaves small, often
blackening when dried, short-petiolate, obovate-oblong to oblong-oblanceolate,
mostly 5-7 cm. long and 2-2.5 cm. broad, obtuse to acuminate, cuneate-attenuate to
the base, usually quite glabrous, commonly domatiate beneath in the nerve axils but
not barbate, the lateral nerves about 8 pairs; inflorescences terminal, erect, cymose-
corymbose, on short or elongate peduncles, rather few-flowered, dense in anthesis but
in fruit open, mostly 2-2.5 cm. broad, the bracts minute, deciduous, the flowers
mostly sessile; calyx minute, 0.5 mm. long, the lobes ovate, acute; corolla white or
greenish white, 3 mm. long, glabrous outside, not barbate in the throat, the lobes
about equaling the tube; anthers usually exserted; fruit red, subglobse, 3-4 mm. long,
the nutlets coarsely costate dorsally, the inner face plane.
152 FIELDIANA: BOTANY, VOLUME 24
Some of the earlier Guatemalan collections were determined as
Psychotria alba Ruiz & Pavon, a species not definitely known to
occur in northern Central America.
Psychotria graciliflora Benth. ex Oerst. Vid. Medd.
Kjoebenhavn 1852: 35. 1852.
Moist or wet, mixed forest, 1,500 m. or less; Peten; Izabal;
Huehuetenango. Ranging southward to Panama.
A rather dense, much branched, leafy shrub 1-2 m. high, glabrous almost
throughout; stipules caducous, small, thin, ferruginous, acuminate, entire or bidentate
at the apex; leaves small, blackening when dried, membranaceous, elliptic-oblong or
lance-oblong, 8 cm. long and 3 cm. broad or usually smaller, acuminate, attenuate to
the base, the lateral nerves about 7 on each side, often domatiate in the nerve axils
but not barbate; inflorescences terminal, slender-pedunculate, erect, small and mostly
few-flowered, in fruit lax and open, trichotomous at the base, the flowers sessile or
pedicellate; calyx deeply lobate, the lobes ovate or lanceolate; corolla white or pale
yellow, 4 mm. long, glabrous outside; fruit red, subglobose, 4 mm. long, glabrous, the
nutlets coarsely costate dorsally.
Psychotria grandis Swartz, Prodr. Veg. Ind. Occ. 43. 1788.
Moist or wet forest, ranging from sea level to about 1,500 m.,
sometimes growing in swampy forest; Peten; Alta Verapaz; Izabal;
Huehuetenango. British Honduras to Panama and Colombia; West
Indies.
A shrub or small tree, sometimes as much as 10 m. high with a trunk 20 cm. in
diameter, but usually much smaller; stipules broadly ovate, long-persistent, subulate-
acuminate, usually 1.5-2 cm. long, sometimes larger; leaves large, short-petiolate,
often subsessile, thick and firm, obovate to broadly oblanceolate, mostly 15-35 cm.
long and 5-13 cm. broad, acute or obtuse and often abruptly apiculate, glabrous, the
conspicuous lateral nerves about 14 pairs; inflorescence terminal, erect, long-
pedunculate, paniculate, commonly 5-7 cm. long and as broad or broader, the stout
branches usually puberulent or villosulous, the basal ones usually verticillate, often
reflexed, the bracts small and deciduous, the flowers glomerate at the ends of the
branches, sessile or short-pedicellate; calyx and hypanthium puberulent or almost
glabrous, the calyx subtruncate or remotely dentate; corolla cream or greenish white,
about 4 mm. long, glabrous or sparsely puberulent outside, the throat barbate, the
obtuse lobes much shorter than the tube; fruit subglobose, 5 mm. long, the nutlets
coarsely costate dorsally.
This species really belongs to the subgenus Psychotria, but the
very large stipules are long-persistent, rather than caducous as is
usual in that group.
Psychotria heydei Standl. Journ. Wash. Acad. Sci. 18: 184.
1928. Flor de San Antonio.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 153
Moist or wet forest, 2,000-2,600 m.; endemic; Quiche (type from
Chiul, Heyde & Lux 3173); Huehuetenango.
Young branches densely villous with short spreading multicellular pale hairs;
stipules persistent, united to form a truncate sheath 3-4 mm. long, this densely short-
villous; leaves on stout petioles 1.5-4 cm. long, elliptic-oblong, broadest at the middle,
11-26 cm. long, 6-10 cm. broad, acuminate, at the base obtuse to rounded, villous-
hirsute above with slender yellowish hairs, densely villous-hirsute beneath, the lateral
nerves about 15 pairs; inflorescence terminal, cymose-paniculate, long-pedunculate,
erect, much branched, lax, many-flowered, 6-9 cm. long, 8-17 cm. broad, the primary
branches opposite or verticillate, divaricate or reflexed, mulberry purple, densely
villous-hirsute; bracts persistent, triangular-subulate, 7 mm. long or less; flowers
mostly sessile but sometimes on pedicels as much as 8 mm. long; hypanthium 2 mm.
long, densely villous; calyx 1.5-2 mm. long, the lobes triangular, subacute; corolla 13-
14 mm. long, densely short-villous, white or tinged outside with lavender, the lobes
ovate, obtuse, 3 mm. long; anthers included.
A very distinct species because of the very dense, long
pubescence, and apparently a rare plant.
Psychotria horizontalis Swartz, Prodr. Veg. Ind. Occ. 44.
1788. Chalchupa (Santa Rosa; probably an erroneous name).
Damp thickets or mixed forest, sometimes in pine forest,
ranging from sea level to about 1,200 m.; Peten; Izabal; Santa
Rosa; Suchitepequez; Retalhuleu; Huehuetenango. Southern
Mexico and British Honduras to Panama, southward to Ecuador;
West Indies.
A shrub of 1-3 m., glabrous throughout or nearly so; stipules caducous, ovate-
triangular, acute or obtuse, 3-8 mm. long; leaves short-petiolate, often almost sessile,
herbaceous, grayish green when dry, elliptic to oblong-lanceolate, mostly 7-11 cm.
long and 2.5-5.5 cm. broad, acuminate, at the base acute to obtuse, the lateral nerves
about 9 pairs; inflorescence terminal, pedunculate, usually small and dense at first,
rather few-flowered, more lax in fruit, commonly 5-rayed at the base, the branches
puberulent, the flowers mostly sessile, the bracts caducous, small; calyx deeply 5-
lobate, the lobes linear-lanceolate; corolla white, 3-4 mm. long, glabrous outside, the
throat not barbate, the lobes half as long as the tube; anthers exserted; fruit red,
glabrous, subglobose, 4 mm. long, the nutlets coarsely costate dorsally.
Psychotria izabalensis L. Wms. Phytologia 28: 229. 1974.
Known only from wet forest or thickets near Lake Izabal,
Dept. Izabal, Jones, Proctor & Facey 3024 (type). Endemic.
Shrubs or small trees to 4 m. tall, the stems and leaves glabrous or sparsely
pubescent, the inflorescence short pilose pubescent, the stipules persistent, truncate
with the erect lateral lobes linear, densely pubescent and about 4-6 mm. long. Leaves
membranaceous, broadly elliptic or oblong-elliptic, long acuminate, glabrous except
the petioles and nerves on lower surface sparsely puberulent, the blades when mature
154 FIELDIANA: BOTANY, VOLUME 24
15-30 cm. long and 5-11 cm. broad and attenuate into a short 1-2 cm. long petiole;
inflorescence terminal, a many-flowered paniculate cyme with the cymules at most
subcapitate but usually more open, densely short pilose pubescent, pedunculate, 8-11
cm. long and 4-6 cm. broad, bracts subtending the main divisions linear- lanceolate,
about 1 cm. long, bracts in the cymules conspicuous, lanceolate to oblong-lanceolate,
acute, sparsely pubescent and ciliate, exceeding the calyx, mostly 3-4 mm. long;
hypanthium and calyx small, about 1 mm. long, pubescent, the calyx divided to the
base, the lobes triangular-ovate, acute, about 0.5 mm. long; corolla white, crisped-
pubescent externally, tube cylindric but broadened and sparsely barbate in the
throat, about 4 mm. long, the lobes oblong-lanceolate, acute, about 2 mm. long;
stamens inserted in the corolla throat, anthers barely exserted, about 1 mm. long;
style as long as the corolla, the stigma lobate; fruits not known.
Related to Psychotria bracteata Sw. and the complex of species
related to P. costaricensis Polak.
Psychotria lilacina Standl. & Steyerm. Field Mus. Bot. 23:
252. 1947.
Wet mixed mountain forest, 1,500-2,800 m.; endemic;
Huehuetenango (type from Cerro Huitz, Steyermark 48564; also on
Cerro Canana).
A densely branched shrub 1-1.5 m. high, the branches glabrous, the older ones
stout, terete, the internodes short; stipules connate into a broad sheath scarcely 0.6
mm. long, bearing on the margin 2 triangular teeth scarcely 0.5 mm. long; leaves
small, membranaceous, on petioles 2-5 mm. long, oblong-elliptic to obovate-oblong, 2-
5.5 cm. long, 7-22 mm. broad, shortly obtuse-acuminate or obtuse, obtuse to
acuminate at the base, glabrous, paler beneath, the nerves obscure on both surfaces,
the lateral ones about 6 on each side, arcuate; inflorescences terminal, corymbiform
or umbelliform, mostly 3-5-flowered, short-pedunculate, the stout pedicels 1-3 mm.
long, glabrous; hypanthium glabrous, columnar, 0.8 mm. long; calyx 3-3.5 mm. long,
5-lobate almost to the base, the lobes lanceolate, ascending or subrecurved above,
attenuate-acuminate or sometimes obtuse; corolla lilac, glabrous outside, about 1
mm. long, the tube thick, cylindric, the lobes short, oval.
Psychotria limonensis Krause, Bot. Jahrb. 54: Beibl. 119: 43.
1916. P. limonensis var. laxinervia Loes. Repert. Sp. Nov. 18: 361.
1922 (type from Palenque, Chiapas).
Wet or moist, mixed forest, or in thickets, usually at 300 m. or
less; Peten; Izabal; Escuintla; Chimaltenango; Suchitepe'quez;
Retalhuleu. Southern Mexico and British Honduras to Panama and
Colombia.
An almost glabrous shrub or small tree, commonly 2-4 m. tall; stipules caducous,
ovate, acuminate, ferruginous, mostly 2-4 cm. long, attenuate, glabrous; leaves often
very large, herbaceous, elliptic-ovate, to broadly elliptic or oblong-elliptic, often as
much as 30 cm. long, commonly 6-12 cm. broad but sometimes much broader
subobtuse to abruptly short-acuminate, at the base usually abruptly contracted and
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 155
long-decurrent, the lateral nerves 15-20 pairs, petioles 2-9 cm. long; inflorescence
terminal (often with lateral branches developing at the side of it), pedunculate,
cymose-paniculate or corymbiform, usually equaling or slightly longer than the
petioles, many-flowered, dense or lax, usually as broad as long, the branches
puberulent or almost glabrous, the bracts small, caducous, the flowers mostly
pedicellate; calyx minutely denticulate, about 2 mm. broad; corolla white, mostly
about 3 mm. long, glabrous outside, the throat white-barbate, the broad lobes
equaling or longer than the tube; fruit oval (or ellipsoid?), 7-9 mm. long, red (possibly
black at maturity), the nutlets costate dorsally.
The material placed here may represent two species but
flowering specimens are not adequate. The specimens from the
western coastal region have ellipsoidal fruits and puberulence on
the young leaves and inflorescence while those from the Atlantic
coast are glabrous or nearly so and have oval fruits.
Psychotria lundellii Standl. in Lundell, Contr. Univ. Mich.
Herb. 4: 29. 1940.
Peten (Uaxactun). British Honduras, the type from Valentin,
El Cayo District, Lundell 6260.
A shrub or tree 2-6 m. high, said to sometimes attain a height of 13 m. and a
trunk diameter of 25 cm., the branches glabrous; stipules caducous, oval, scarcely
more than 4 mm. long, broadly rounded at the apex, ferruginous, glabrous; leaves
subcoriaceous, on petioles 1-2.5 cm. long, oblanceolate or oblong-oblanceolate, 8-14
cm. long, 2-4.5 cm. broad, narrowly long-acuminate or attenuate-acuminate,
gradually long-attenuate to the base, glabrous, sparsely short-barbate beneath in the
nerve axils, the lateral nerves about 7 pairs; inflorescence terminal, densely many-
flowered, cymose-paniculate, on a peduncle 2 cm. long or less, commonly 4-6 cm. long
and 5-8 cm. broad, trichotomous at the base, the branches glabrous or sparsely short-
pilosulous, the small bracts soon deciduous, the flowers on short stout pedicels in
small dense cymules; hypanthium glabrous, scarcely 1 mm. long, the calyx of equal
length, dentate-lobulate, the teeth triangular, acute; corolla white, 2.5 mm. long,
glabrous outside, densely white-barbate in the throat, the lobes oblong, acute,
equaling the tube; anthers semiexserted; style short-exserted; fruit globose, probably
red, 5 mm. long, the nutlets depressed-costate dorsally or almost smooth.
Psychotria macrophylla Ruiz & Pa von, Fl. Peruv. 2: 56, t
202, f. a. 1799. P. anomothyrsa Schum. & Donn.-Sm. Bot. Gaz. 35: 3.
1903 (type from Cubilguitz, Alta Verapaz, Tuerckheim 7909).
Wet mixed forest, 800-2,300 m.; Alta Verapaz; Izabal; El
Progreso; Chiquimula (Cerro Tixixi); Chimaltenango; Solola;
Huehuetenango; Quezaltenango; San Marcos. Southern Mexico and
British Honduras to Panama, southward to Bolivia.
Stems usually 1 m. high or less and simple, but sometimes branched and as much
as 2.5 m. tall, glabrous or nearly so; stipules persistent or finally deciduous, thin and
156 FIELDIANA: BOTANY, VOLUME 24
pale green, broad, apiculate; leaves large, membranaceous, slender-petiolate, elliptic-
oblong to lance-oblong or oblong- oblanceolate, commonly 15-30 cm. long and 6.5-16
cm. broad, often narrower, acuminate or abruptly acute, acute to attenuate at the
base, glabrous or pruinose-puberulent beneath on the nerves, sometimes short-
hirtellous, the lateral nerves about 14 pairs; inflorescences axillary, cymose-
paniculate, usually long-pedunculate, rather few-flowered, the branches often
reflexed, glabrous or puberulent, the flowers sessile, the bracts mostly minute; calyx
small, denticulate; corolla whitish or yellowish, 4-5 mm. long, glabrous outside; fruit
4-5 mm. long, often somewhat compressed, glabrous, white, with conspicuous
cystoliths.
Psychotria marginata Swartz, Prodr. Veg. Ind. Occ. 43. 1788.
P. nicaraguensis Benth. ex Oerst. Vid. Medd. Kjoebenhavn 1852:
34. 1853. Oocan cimarron (Pete"n, fide Lundell). Figure 48.
Wet thickets or wet mixed forest, at 600 m. or less; Peten; Alta
Verapaz; Izabal; Hijehuetenango. British Honduras to Panama,
southward to Peru; West Indies.
An almost glabrous shrub, usually about 2 m. high but sometimes as much as 4.5
m., with slender branches; stipules rather large, caducous, ferruginous,
membranaceous, oblong to ovate, entire; leaves short-petiolate, firm-membranaceous,
usually darkening to a dull gray-black when dried, obovate-oblong to oblanceolate,
mostly 9-12 cm. long and 2.5-4 cm. broad, acute or abruptly acute or acuminate,
cuneately long-attenuate to the base, minutely puberulent beneath or almost
glabrous, the lateral nerves about 12 pairs; inflorescence terminal, long-pedunculate,
cymose-paniculate, very lax, sometimes 10 cm. long and 13 cm. broad, the slender
pedicels 2-5 mm. long, glabrous or minutely puberulent; calyx small, the teeth short,
obtuse; corolla ochroleucous, about 3 mm. long, glabrous outside, the lobes slightly
shorter than the tube, the throat not barbate; fruit globose, red, 3-4 mm. long, the
carpels coarsely costate dorsally.
Found only on the Atlantic drainage from Mexico to Costa
Rica where it crosses over and is also on the Pacific side.
Psychotria microdon (DC.) Urban, Symb. Antill. 9: 539. 1928.
Rondeletia microdon DC. Prodr. 4: 408. 1830. P. pinularis Sess<§ &
Moc. Fl. Mex. ed. 2. 57. 1894. Mapouria microdon Bremekamp,
Recueil Trav. Bot. NSerl. 31: 286. 1934.
Moist thickets, 900 m. or lower; Pete"n; Huehuetenango. Mexico
to Costa Rica and Panama, southward to Ecuador and the
Guianas; West Indies.
A nearly or quite glabrous shrub, usually 1-3 m. high, with pale branches;
stipules small, more or less persistent, broadly triangular, sometimes annular and
indurate in age; leaves membranaceous, short-petiolate, mostly oblong-obovate,
sometimes elliptic, 4-7 cm. long, obtuse to short-acuminate, attenuate to the base,
barbate beneath in the nerve axils; inflorescence terminal, cymose-corymbose, few-
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 157
flowered, pedunculate, the flowers pedicellate; calyx cupular, truncate, 1 mm. high;
corolla white, sometimes 1 cm. long but usually about 8 mm., glabrous; fruit ovoid-
globose, about 7 mm. long, not costate dorsally, slightly sulcate on the inner surface.
The flowers are fragrant. In British Honduras the shrub is said
to be called "night sage," "dead man's bones," and "hueso de
finado." The Maya name "baque-ac" is reported from Yucatan,
where the plant is said to be used medicinally by the Indians.
Psychotria minarum Standl. & Steyerm. Field Mus. Bot. 23:
253. 1947.
Moist or wet, mixed, mountain forest, 2,000-3,300 m.; El
Progreso (type collected near the summit of Volcan Siglo, growing
in colonies in shade of trees and shrubs, Steyermark 43106); Zacapa.
El Salvador.
A branched shrub 30-90 cm. high, the branches subterete, puberulent or glabrate,
with short internodes, the older branches sordid-ochraceous; stipules connate into a
very short, truncate sheath, this bearing on the margin at each side a triangular tooth
scarcely 0.5 mm. long; leaves small, on slender petioles 4-6 mm. long,
membranaceous, blackish when dried, elliptic, oval-elliptic, or oblong-elliptic, 3-6.5
cm. long, 1-2.7 cm. broad, acute or acuminate, often with an obtuse tip, acute or
obtuse at the base, glabrous above, paler beneath, glabrous or sometimes scaberulous,
especially on the costa and nerves, the lateral nerves 5-6 on each side, inconspicuous,
arcuate-ascending, the veins obsolete; inflorescences terminal, on peduncles 1 cm.
long or shorter, corymbiform, mostly 3-5-flowered, trichotomous at the base, glabrous
or glabrate, much shorter than the leaves, the flowers sessile or on short thick
pedicels; hypanthium obconic, 1.2 mm. long, glabrous; calyx scarcely 0.5 mm. long,
the teeth triangular, acute; fruit 4.5-5 mm. long, the pyrenes obtusely costate
dorsally, one of them often abortive.
Psychotria mombachensis Standl. Field Mus. Bot. 8: 188.
1930. Tinta de monte.
Moist mixed lowland forest, about 100 m.; Suchitepe'quez
(Tiquisate, Steyermark 47639). Advanced forest, limestone valley,
Valentin, British Honduras; to be expected in Peten. Mexico
(Tabasco); Honduras; Nicaragua, the type from Volcan de
Mombacho.
A slender shrub about 2 m. high, glabrous throughout, the branches green;
stipules persistent, green, 3 mm. long, deeply bilobate, the lobes broadly ovate, obtuse
or rounded at the apex; leaves membranaceous, on slender petioles 1-2 cm. long,
elliptic-oblong to obovate-oblong or ovate-oblong, 10-17 cm. long, 4-6.5 cm. broad,
gradually or abruptly acuminate or long-acuminate, acute to long-attenuate at the
base, bright green above, slightly paler beneath, the lateral nerves about 14 pairs;
inflorescences terminal, erect, short-pedunculate, on a peduncle about 1 cm. long,
cymose-corymbose, lax and rather few-flowered, about 2.5 cm. long and 5 cm. broad,
158 FIELDIANA: BOTANY, VOLUME 24
trichotomous at the base, the few branches bracteate at the base, the bracts subulate,
the flowers clustered, sessile; hypanthium cylindric, 1 mm. long, the calyx slightly
longer, campanulate, 5-cleft to the middle, the lobes oblong, obtuse; corolla white,
glabrous outside, not barbate in the throat, the tube about 13 mm. long, dilated
above, the lobes oblong, obtuse, about 3 mm. long.
As known at present, the distribution of this species is rather
unusual, but it is to be expected that it will be discovered in El
Salvador.
Psychotria nervosa Swartz, Prodr. Veg. Ind. Occ. 43. 1788. P.
undata Jacq. Hort. Schoenbr. 3: 5, t. 260. 1798. P. granadensis
Benth. ex Oerst. Vid. Medd. Kjoebenhavn 1852: 34. 1853. Sacxanal
(Pet6n, Maya, fide Lundell).
Damp forest or thickets of the lowlands of both slopes,
ascending on the Pacific coast to about 1,200 m., but usually at
much lower elevations; Pet<§n; Alta Verapaz; Izabal; Escuintla;
Sacatep£quez (near Las Lajas); Huehuetenango. Southern Mexico
to Panama, extending to Colombia and Venezuela; West Indies;
southern Florida.
Usually a shrub about 1 m. high, sometimes as much as 3 m., the branches
glabrous; stipules membranous, at first enclosing the terminal buds, later cleft, often
1.5 cm. long, caducous; leaves herbaceous, lanceolate to ovate or oblong-obovate,
commonly 6-11 cm. long and 2-5 cm. broad, acuminate, acuminate to attenuate at
the base, often with undulate margins, glabrous or glabrate, the lateral nerves
elevated and conspicuous beneath, about 10 pairs, the leaves often brownish when
dried; inflorescence terminal, sessile or nearly so, cymose-paniculate, mostly 3-5 cm.
long, very dense at first but becoming open in fruit, the branches glabrate, the
flowers mostly pedicellate; hypanthium narrowly ovoid, longer than the undulate
calyx; corolla white, about 6 mm. long, pilose in the throat, the lobes half as long as
the tube; fruit red, ellipsoid or oval, conspicuously longer than broad, 6-7 mm. long,
the nutlets conspicuously costate.
Psychotria nervosa var. rufescens (HBK.) L. Wms.
Phytologia 26: 492. 1973. Psychotria rufescens HBK. Nov. Gen. &
Sp. 3: 364. 1819. P. rufescens H. & B. ex R. & S. Syst. Veg. 5: 192.
1819. P. hirta H. & B. ex R. & S. Syst. Veg. 5: 191. 1819. P. nervosa
subsp. rufescens Steyerm. Mem. N. Y. Bot. Card. 23: 480. 1972. P.
nervosa subsp. rufescens var. hirta Steyerm. I.e. 481.
Damp forest or thickets at low elevations along both coasts,
perhaps reaching to 1,200 m.; Pete"n; Izabal; Sacatepe'quez;
Escuintla. Mexico and British Honduras, south to Venezuela and
Colombia.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 159
Similar to the species but with the branches rufous-hirsute, the stipules
ferrugineous, the leaves hirsute to sparsely hirsute, the fruits often hirtellous.
This variation was considered by the senior author to be only a
pubescent phase of the species.
Psychotria oerstediana Standl. Contr. U. S. Nat. Herb. 23:
1390. 1926. Mapouria obovata Oerst. Amer. Centr. 17, t. 14, f. 3, 4.
1863. P. obovata Hemsl. Biol. Cent. Am. Bot. 2: 50. 1881, not Ruiz
& Pav6n, 1799.
Dry or moist thickets, 400 m. or less; Peten; Alta Verapaz;
Retalhuleu. Southern Mexico and British Honduras to Honduras.
A rather stout shrub 1-2.5 m. tall, glabrous or nearly so; stipules caducous, about
1 cm. long or shorter, membranous, ferruginous, obtuse or rounded at the apex,
glabrous; leaves on stout petioles, broadly elliptic to obovate or oblong- obovate,
mostly 10-13 cm. long and 4-6 cm. broad, acute or abruptly short-acuminate, with
obtuse tip, at the base cuneate-attenuate or abruptly acuminate, pale grayish green
when dry, rather thick, obscurely domatiate beneath in the nerve axils, the lateral
nerves conspicuous, about 8 pairs; inflorescence terminal, usually sessile, many-
flowered, at first dense and contracted, in fruit open and lax and sometimes as much
as 7 cm. broad, the branches usually glabrous, the bracts caducous, the flowers
glomerate, sessile or nearly so; calyx subtruncate, the teeth usually obscure,
sometimes evident but broad; corolla white or greenish cream, 5-6 mm. long, glabrous
outside, the lobes reflexed, much shorter than the tube; anthers often exserted; fruit
red, 5-7 mm. long, oval or ellipsoid, glabrous, the nutlets conspicuously costate
dorsally.
Psychotria officinalis (Aubl.) Sandwith, Kew Bull. 1931: 473.
1931. Nonatelia officinalis Aubl. PL Guian. 1: 182, t. 70, fig. 1. 1775.
Psychotria involucrata Sw. Fl. Ind. Occ. 1: 413. 1797.
Wet mixed forest or thickets, mostly at less than 350 m.;
Pet6n; Alta Verapaz; Izabal; Huehuetenango. Southern Mexico and
British Honduras to Panama; Colombia and Venezuela; West
Indies.
An almost glabrous shrub, commonly 3 m. high or less, with slender but rigid,
green branches; stipules persistent, short, green, the lobes very short, acute or
acuminate; leaves short-petiolate, thin, bright green when dried, lanceolate to elliptic-
ovate or rhombic-ovate, commonly 9-18 cm. long and 4-8 cm. broad, abruptly long-
acuminate, contracted and long-decurrent at the base, the lateral nerves about 12
pairs, very slender; inflorescence terminal, erect or recurved, pedunculate, subcapitate
and commonly 1-2 cm. long, few-flowered, in age becoming more open and often
racemiform, the short branches usually purplish, hirtellous, the flowers sessile or
nearly so and densely capitate-glomerate at the ends of the stout branches; outer
bracts narrow and often much exceeding the flowers, purple or purplish, the inner
bracts shorter, broad, conspicuous; calyx minute, denticulate; corolla white or
160 FIELDIANA: BOTANY, VOLUME 24
ochroleucous, 4-5 mm. long, hirtellous outside, the lobes much shorter than the tube;
fruit globose, 4 mm. long, black at maturity, the nutlets coarsely costate dorsally.
Somewhat variable in the form of the inflorescence, and it is
possible that the Central American material embraces more than a
single species, although the variations perhaps represent only
different stages in the development from flower to fruit.
Psychotria oreodoxa L. Wms. Phytologia 28: 230. 1974. Chile.
Moist forests on the Pacific slopes of the western highlands,
1,000-1,500 m. or perhaps higher; Solola; Suchitepequez;
Quezaltenango. Endemic.
Shrubs to 5 m. tall, branched, the branches glabrous or obscurely puberulent,
stipules persistent, about 3 mm. long, bilobate laterally with aceriform lobes. Leaves
elliptic or elliptic-oblanceolate, long acuminate, attenuate to the slender petiole,
glabrous, with about 10 pairs of secondary nerves, the blades 10-18 cm. long and 2-5
cm. broad, petioles slender, mostly 1-3 cm. long; inflorescence terminal, thyrsiform,
pedunculate, to about 7 cm. long; flowers white, abundant; hypanthium and calyx 1-
1.5 mm. long, the calyx short, the lobes narrowly triangular, acute, about 0.5 mm.
long; the corolla small, tubular below and slightly expanded above, about 5-6 mm.
long, the lobes small, oblong; stamens inserted in the corolla throat and anthers
exserted and about 1.5 mm. long; fruit very fleshy, purple-black, the seeds about 2.5
mm. long, obscurely ridged.
Psychotria orogenes L. Wms. Phytologia 28: 231. 1974.
Wet montane cloud forests at 1,400-1,600 m.; Alta Verapaz;
Baja Verapaz (type, Williams, Molina & Williams 41924). Endemic.
Shrubs or weak trees 3-4 m. tall, the branches densely villous pubescent with
short spreading hairs, the stipules persistent, joined and surrounding the branches,
each pubescent stipule with a lateral pair of linear-lanceolate lobes 3-4 mm. long.
Leaves elliptic or elliptic-oblanceolate, acuminate, sparsely pilose above, prominently
and softly so beneath, with mostly 13-16 pairs of lateral nerves, the blade 8-15 cm.
long and 1.5-4 cm. broad, petioles short, 0.7-1.5 cm. long; inflorescence terminal, a
pedunculate panicle of cymules up to about 6 cm. long, conspicuously soft pilose
when immature but less so in fruit, bracteolate, the bracts linear-filiform, those of the
main branches 1 cm. long, those subtending flowers smaller; flowers white,
conspicuously short pilose, usually subtended by a filiform bract about as long as the
calyx; hypanthium and calyx 1-1.5 mm. long, villous; calyx very small, the lobes
lance-triangular, about 0.5 mm. long; corolla small, 4-5 mm. long, short villous
outside, the lobes oblong, acute, about 2 mm. long; fruits subglobose, pilose,
prominently ridged, 4-5 mm. long.
A montane cloud forest species somewhat related to Psychotria
pubescens Sw.
Psychotria pachecoana Standl. & Steyerm. Field Mus. Bot.
22: 205. 1940.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 161
Damp mixed forest or wet thickets, 1,400-2,500 m.; endemic;
Alta Verapaz (region of Tactic); Baja Verapaz (type from region of
Santa Rosa, Standley 69922) ; El Progreso; Zacapa (Sierra de las
Minas); Huehuetenango.
A branched shrub about 2 m. tall, the branches densely pilosulous; stipules
persistent, the sheath 4 mm. long, the lobes remote, 3-4 mm. long, triangular-
attenuate; leaves membranaceous, short-petiolate, oblong-oblanceolate or narrowly
oblong-oblanceolate, 7-15 cm. long, 2-4 cm. broad, narrowly long-attenuate, cuneately
narrowed to the base, densely pilose above with short, spreading or subappressed
hairs, somewhat paler beneath, densely velutinous-pilose, the lateral nerves about 20
pairs; inflorescence terminal, erect, on a peduncle 1.5-2 cm. long, cymose-paniculate,
in fruit 3.5-5 cm. long and as much as 7 cm. broad, in flower smaller and denser, the
branches all bracteate at the base, hispidulous-pilose, the bracts almost subulate, 4
mm. long or less, the flowers sessile, densely crowded; corolla greenish white,
hispidulous; fruit subglobose, 4 mm. long, sparsely hirtellous, the nutlets coarsely
costate dorsally.
The species was named for Don Mariano Pacheco Herrarte,
Director General de Agricultura of Guatemala for many years and
friend of the senior author.
Psychotria parasitica Swartz, Prodr. Veg. Ind. Occ. 44. 1788.
Viscoides pendulum Jacq. Sel. Stirp. Amer. 73, t. 51, f. 1. 1763. P.
pendula Urban, Symb. Antill. 1: 445. 1899, not Hook. f. 1882.
Moist or wet forest, 1,500 m. or usually much lower; Alta
Viscoides pendulum Jacq. Sel. Stirp. Amer. 73, t. 51, f. 1. 1763. P.
pendula Urban, Symb. Antill. 1: 445. 1899, not Hook. f. 1882.
A glabrous epiphytic shrub with dark terete branches, often pendent; stipules 2-3
mm. long, united to form a truncate sheath, this persistent and becoming indurate
and pale in age; leaves very thick and fleshy, short-petiolate, broadly ovate to
elliptic-oblong, mostly 2-10 cm. long and 1.5-2.5 cm. broad, obtuse to acuminate,
obtuse or rounded at the base, the costa conspicuous but the lateral nerves obsolete;
inflorescence terminal, erect, small and few-flowered, pedunculate, the flowers sessile
or on short thick pedicels; calyx dentate, small, the teeth triangular; corolla white, 6-
7 mm. long or shorter, glabrous outside; fruit obovoid-globose, red, 3.5-4 mm. long,
the nutlets smooth or nearly so dorsally.
Unlike any other Guatemalan species in its epiphytic habit and
thick fleshy leaves.
Psychotria parvifolia Benth. ex Oerst. Vid. Medd.
Kjoebenhavn 1852: 36. 1853.
Dense damp mixed forest of the slopes of the western
highlands, 1,300-2,000 m.; Quezaltenango (southern slopes of Volcan
162 FIELDIANA: BOTANY, VOLUME 24
de Santa Maria); San Marcos (Volcan de Tajumulco). Costa Rica
and Panama.
A slender, usually densely branched and very leafy shrub 1-2 m. tall, the
branches puberulent or short-pilose; stipules ovate, caducous, ferruginous, about 4
mm. long, often villosulous; leaves small, membranaceous, slender-petiolate, chiefly
oblong-lanceolate or narrowly elliptic-lanceolate, mostly 3.5-5.5 cm. long and 1-1.5
cm. broad, subobtuse to long-attenuate, usually long-attenuate and decurrent at the
base, the margins often somewhat undulate, blackening when dried, glabrous, usually
minutely domatiate beneath in the nerve axils, the lateral nerves obscure, about 7
pairs; inflorescence terminal, erect, cymose-paniculate, sessile, small and few-
flowered, the flowers mostly sessile, the bracts caducous; calyx obscurely dentate;
corolla white, 4 mm. long or less, glabrous outside, barbate in the throat; fruit
globose, red, about 4 mm. in diameter, the nutlets rather inconspicuously costate
dorsally.
Psychotria patens Swartz, Prodr. Veg. Ind. Occ. 45. 1788. P.
flexuosa Willd. Sp. PI. 1: 966. 1798.
Wet mixed forest or sometimes in second growth, 500 m. or less;
Alta Verapaz; Izabal; Huehuetenango; to be expected in Peten.
Southern Mexico and British Honduras to Bolivia and Brazil. West
Indies.
A slender glabrous shrub commonly 1-3 m. tall, with green or yellowish branches;
stipules small, persistent, green, biaristate; leaves thin but stiff, drying green and
lustrous, lanceolate to lance-oblong, mostly 10-15 cm. long and 3.5-5.5 cm. broad,
narrowly long-acuminate, obtuse or acute at the base, the lateral nerves about 7
pairs; inflorescence terminal, erect, slender-pedunculate, thyrsoid-paniculate, often
elongate, the short slender branches mostly reflexed, not bracteate at the base, the
flowers sessile or short-pedicellate; calyx minute, with triangular teeth; corolla white
or whitish, glabrous outside, 7 mm. long, the lobes very short; fruit subglobose or
often somewhat didymous, white or pale blue at maturity, 3 mm. long, the nutlets
costate dorsally.
Psychotria pittieri Standl. Contr. U. S. Nat. Herb. 18: 132.
1916.
Wet mixed forest or in thickets, 300-500 m.; Alta Verapaz
(Montana Yxocubvain, Steyermark 44994). Costa Rica; Panama.
A slender shrub 1-2 m. high, often densely branched, the branches pilose with
rather long, pale, appressed or spreading hairs; stipules persistent, 4-6 mm. long,
bilobate, the lobes subulate; leaves small, short-petiolate, elliptic-oblong, mostly 4-6
cm. long and 1.5-2.5 cm. broad, occasionally somewhat larger, green above,
hirsutulous along the costa, appressed-pilose beneath; inflorescence terminal, reflexed
in age, cymose-paniculate, slender-pedunculate, commonly about 2 cm. long, rather
few-flowered, the branches pilose, the flowers subsessile; calyx 1 mm. long, the lobes
obtuse; corolla white, 4-5 mm. long, minutely appressed-pilose; fruits small, blue,
pilose with long white hairs.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 163
The only known collection in Guatemala is not in good
condition.
Psychotria pleuropoda Donn.-Sm. Bot. Gaz. 40: 5. 1905.
Moist or wet mixed forest, often on limestone, 900 m. or lower;
Pete"n; Alta Verapaz (type from Cubilguitz, Tuerckheim 8529);
Izabal. British Honduras.
A glabrous shrub 1-4.5 m. high; stipules caducous, membranaceous, ferruginous,
more or less connate, 10-13 mm. long, bifid at the apex, the lobes linear-setaceous,
ciliate; leaves herbaceous, blackening when dried, lance-linear, 10-20 cm. long and 1.5-
2.5 cm. broad, long-attenuate to the apex, acute at the base, the lateral nerves
obscure; inflorescence axillary, long-pedunculate, cymose, usually dense and head-
like with few branches, the peduncle as much as 8 cm. long, the flowers mostly on
pedicels 1.5 mm. long or less; calyx lobes ovate, 0.5 mm. long; corolla white, 6 mm.
long, the lobes about equaling the tube, oblong, reflexed; fruit red, oval, 5-6 mm.
long, the nutlets 4-costate dorsally, not sulcate on the inner surface.
An isolated species easily recognized by the long and narrow
leaves and axillary inflorescences. Referable to the subgenus
Psychotria.
Psychotria pubescens Swartz, Prodr. Veg. Ind. Occ. 44. 1788.
P. justicioides Schlecht. Linnaea 9: 596. 1834. Chalchupa (Santa
Rosa, perhaps an erroneous name); zac-ixcanan (Peten, Maya, fide
Lundell); guayabeno (fide Aguilar).
Damp thickets or wet mixed forest, chiefly in the hot country
but ascending to about 1,500 m., but most common at low
elevations; Pete"n; Alta Verapaz; Izabal; Chiquimula; Jalapa;
Jutiapa; Santa Rosa; Escuintla; Guatemala; Sacatepe"quez; Solola;
Retalhuleu; Quezaltenango; Huehuetenango; San Marcos. Mexico
to Panama; West Indies.
A shrub mostly 1.5-3 m. high, the branches often constricted at the nodes when
dry, puberulent or short-pilose; stipules small, deeply bifid, the lobes acute or
acuminate; leaves membranaceous, usually green when dried, petiolate, elliptic to
oblong-lanceolate, mostly 7-15 cm. long and 3-7 cm. broad, acuminate, acute to
attenuate at the base, often rough to the touch on the upper surface and minutely
pilosulous or scaberulous, slightly paler beneath, usually puberulent or finely short-
pilose, the lateral nerves about 11 pairs; inflorescence terminal, cymose-corymbose,
usually pedunculate, many-flowered, the branches densely short-pilose, the flowers
sessile or nearly so; calyx lobes ovate, very small; corolla greenish white or greenish
yellow, 4-5 mm. long, usually puberulent pilose in the throat, the oblong lobes shorter
than the tube; fruit subglobose, juicy, black or dark purple, 3-4 mm. long, the nutlets
acutely costate dorsally.
164 FIELDIANA: BOTANY, VOLUME 24
Psychotria purpusii Standl. Contr. U. S. Nat. Herb. 23: 1388
1926.
San Marcos, slopes of Volcan de Tajumulco, about 2,000 m.,
wet mixed forest. Mexico (Chiapas), the type from Cerro del
Boquer6n, Purpus 7012.
A shrub or small tree 1.5-7 m. tall, the branches densely soft-villous; stipules
persistent, small, the lobes remote, ovate, obtuse; leaves large, membranaceous,
slender-petiolate, elliptic or oval, 10-21 cm. long and 4-10 cm. broad, abruptly
acuminate, acute to almost rounded at the base, soft-pilose on both surfaces, more
densely so beneath, the lateral nerves about 14 pairs; inflorescence terminal or
pseudolateral, on very long peduncles, often nutant, subtending bracts lanceolate or
sometimes broader, head-like with 3 short branches, in fruit sometimes more open,
the bracts usually about equaling the flowers, orange and green, oblong or lanceolate,'
2.5 cm. long or shorter, acute or acuminate, the flowers sessile or short-pedicellate;
corolla about 15 mm. long, white, the short lobes erect, very obtuse; fruit subglobose',
7 mm. long, densely pilose, white, the hairs mustard yellow.
An unusually well-marked species, almost equally well
referable to the genus Cephaelis, noteworthy for the very large and
brightly colored bracts subtending the headlike inflorescence, also
for the abundant soft dense pubescence of the leaves.
Psychotria schippii Standl. & Steyerm. Field Mus. Bot. 23:
24. 1943.
Pete"n (Uaxactun). British Honduras, the type from advanced
forest, limestone valley, Valentin, El Cayo District, Lundell 6193.
A glabrous shrub or tree, often 5-9 m. high, the trunk often 5-10 cm. in diameter;
stipules caducous, 1.5 cm. long or less, ferruginous, firm, attenuate; leaves rather
large, on petioles 1-2.5 cm. long, firm or subcoriaceous, elliptic or oblong-elliptic, 11-19
cm. long, 5-9.5 cm. broad, abruptly acuminate or caudate-acuminate, at the base
acute or abruptly contracted and long-decurrent, grayish green above when dried,
lustrous, the costa and nerves prominent, minutely and sparsely barbate beneath in
the nerve axils, the lateral nerves very conspicuous, about 14 on each side;
inflorescence terminal, sessile, cymose-paniculate, abundantly branched in fruiting
state, many-flowered and lax, 6 cm. long and 8 cm. broad or smaller, the bracts
caducous, the flowers sessile; fruit red, glabrous, globose, 4-5 mm. in diameter,
rounded at the base and apex, the nutlets obtusely costate dorsally; calyx teeth
minute, obtuse and rounded.
Psychotria simiarum Standl. Field Mus. Bot. 4: 344. 1929.
Wet forests or thickets, at 700 m. or less; Alta Verapaz; Izabal.
British Honduras; Honduras, the type from Lancetilla valley near
Tela.
A shrub or small tree 2-7.5 m. high, the slender branches green, glabrous,
constricted at the nodes when dry; stipules persistent, green 1-2 mm. long, connate
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 165
into a sheath, this bidentate at the apex; leaves membranaceous, on slender petioles
1-2.5 cm. long, elliptic or oblong-elliptic, 8-15 cm. long, 2.5-7 cm. broad, abruptly long-
acuminate, acute at the base, glabrous, slightly paler beneath, the lateral nerves
about 7 pairs; inflorescence terminal, erect, pedunculate, cymose-paniculate, densely
many-flc wered, about 2 cm. long and 2.5 cm. broad, but in age often larger, the
branches minutely puberulent, bracteate at the base, the bracts small, green, linear or
subulate, the flowers sessile or on very short pedicels; hypanthium 0.8 mm. long,
puberulent or glabrous, the calyx about the same length, short-dentate, the teeth
unequal, triangular or broadly ovate, obtuse or acute; corolla ochroleucous, about 3
mm. long, minutely pruinose-puberulent, the lobes oblong, obtuse, almost equaling
the tube; fruits subglobose, blue, glabrous, 5-6 mm. in diameter, the nutlets obscurely
costate dorsally, narrowly sulcate on the inner surface.
Psychotria skutchii Standl. Field Mus. Bot. 17: 283. 1937.
Moist or wet mixed forest, 1,500-2,400 m.; endemic;
Suchitepequez; Huehuetenango (type from San Juan Ixcoy, 2,400
m., in oak forest, Skutch 1074); Mexico (Chiapas).
A shrub or small tree 1-2.5 m. tall or more, the slender branches densely
pilosulous with short spreading sordid hairs; stipules persistent, the sheath 1.5 mm.
long, the lobes triangular-subulate; leaves small, on slender petioles 7-15 mm. long,
narrowly elliptic-oblong or oblanceolate-oblong, 6-9.5 cm. long, 2-3 cm. broad, long
acuminate, narrowed to the acute or acuminate base, densely puberulent above and
spreading-pilosulous along the costa, densely and softly pilosulous beneath, the
lateral nerves about 13 pairs; inflorescence terminal, erect, sessile or slender-
pedunculate, cymose-paniculate, 2.5-4.5 cm. long, trichotomous at the base, the
branches spreading or somewhat reflexed, bracteate at the base, densely pilosulous
with spreading hairs, the flowers sessile or on very short pedicels; hypanthium and
calyx densely pilose, the calyx 1 mm. long, unequally dentate, the teeth triangular,
subacute; corolla white, pilosulous, the tube 6-8 mm. long, the throat not barbate, the
lobes triangular-oblong, obtuse, shorter than the tube, glabrous within; anthers semi-
exserted; style equaling the corolla lobes.
Psychotria steyermarkii Standl. Field Mus. Bot. 22: 387.
1940.
Damp mixed forest, 1,200-2,000 m., Quezaltenango, the type
from Quebrada de San Geronimo, Finca Pirineos, Steyermark 33461.
A slender shrub about 1.5 m. high, the young branches green, at first pilose with
short spreading hairs; stipules persistent, 8-9 mm. long, bilobate almost to the base,
the lobes linear or almost subulate, attenuate, puberulent at the base; leaves
membranaceous, on slender petioles 1-2 cm. long, narrowly oblong-lanceolate to
almost linear-lanceolate, 5-14 cm. long, 1.5-2.5 cm. broad, narrowly attenuate-
acuminate, long-attenuate to the acute or acuminate base, green above, pilose along
the costa, paler beneath, sordid-pilosulous on the nerves and costa or puberulent, the
lateral nerves about 23 pairs; inflorescence terminal, sometimes apparently recurved,
on a peduncle 3.5 cm. long or shorter, cymose, trichotomous at the base, about 3 cm.
long and broad, the branches all bracteate at the base, glabrous or glabrate, the
166 FIELDIANA: BOTANY, VOLUME 24
flowers crowded in a few head-like clusters, sessile or short pedicellate, the lowest
bracts foliaceous, linear-lanceolate, about 1 cm. long, acuminate-attenuate; calyx 1.5-
2 mm. long, dentate, the teeth ovate, acute; fruit subglobose, 4 mm. long, glabrous,
the nutlets coarsely costate dorsally.
The species is known only from the valley of the Rio Samala,
in the vicinity of the type locality.
Psychotria tenuifolia Swartz, Prodr. Veg. Ind. Occ. 43. 1788.
P. sessilifolia Mart. & Gal. Bull. Acad. Brux. 11, pt. 1: 228. 1844.
Moist or wet thickets or damp mixed forest, chiefly in the
plains and foothills, ascending to about 1,200 m. but mostly at
lower elevations; Pete"n; Alta Verapaz; Santa Rosa; Escuintla;
Chimaltenango (Sibaja); Huehuetenango. Southern Mexico and
British Honduras to Honduras; West Indies.
A shrub, usually 1-2 m. high, often even lower but sometimes as much as 4 m.,
the branches green, usually puberulent; stipules caducuous, membraneous,
ferruginous, 1 cm. long or less, usually bilobate at the apex, the lobes subulate-
acuminate; leaves short petiolate, oblong-elliptic to narrowly oblong-lanceolate,
mostly 9-16 cm. long and 1.5-5.5 cm. broad, acute to acuminate or attenuate, usually
long-attenuate to the base, membranaceous, usually dark when dried, puberulent or
minutely pilose beneath or glabrate, usually glabrous above, slightly paler beneath,
the lateral nerves about 12-14; inflorescence terminal, sessile, cymose-paniculate,
often appearing axillary because of elongation of the branch, small and few-flowered
or often many-flowered and as much as 6 cm. long, rather lax in fruit, the bracts
caducous, the branches puberulent or densely short-pilose, the flowers sessile or on
very short pedicels; calyx about 1 mm. broad, very short, shallowly dentate; corolla
white, about 3 mm. long, glabrous or sparsely puberulent outside, the lobes equaling
the tube; fruit bright red, globose, 4-5 mm. long, the nutlets obtusely costate dorsally.
Although dull when dried, the leaves often are lustrous in the
fresh state.
Psychotria uliginosa Swartz, Prodr. Veg. Ind. Occ. 43. 1788.
Notopleura uliginosa Bremekamp, Recueil Trav. Bot. Neerl. 31:
290. 1934.
Wet mixed forest, 2,600 m. or less; Alta Verapaz; Izabal;
Chiquimula (Cerro Tixixi); Solola; Huehuetenango; Quezaltenango;
San Marcos. Southern Mexico and British Honduras to Panama
and the Guianas; West Indies.
A stout shrub or herb, usually simple and about 1 m. high or less, sometimes
taller, glabrous throughout or nearly so; stipules persistent, ovate-triangular, acute,
the upper portion soon decaying; leaves large, thick and succulent in the living state,
on petioles 4 cm. long or less, oblanceolate-oblong to elliptic-oblong or obovate, as
much as 25 cm. long and 10 cm. broad but usually somewhat smaller, acute or
abruptly short-acuminate, acute at the base, deep green above, pale beneath, the
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 167
lateral nerves about 15 pairs, inconspicuous; inflorescences axillary, long-pedunculate,
cymose-paniculate, the branches bracteate at the base, the flowers capitate or
glomerate at the ends of the branches; calyx cupular, irregularly dentate; corolla
white, about 9 mm. long, the lobes acute, half as long as the tube; fruit bright red,
juicy, 7-8 mm. long, the nutlets conspicuously carinate dorsally, strongly compressed.
A rather handsome plant when well grown because of the
brilliant red fruits. The plant thrives best in very wet and dense
forest at 1,200-1,500 m.
Psychotria viridis Ruiz & Pavon, Fl. Peruv. 2: 61, t. 210, f. b.
1799. P. glomerata HBK. Nov. Gen. & Sp. 3: 363. 1820. P.
microdesmia Oerst. Vid. Medd. Kjoebenhavn 1852: 36. 1853.
Moist or wet thickets or forest, on limestone, 150 m. or less;
Peten. Collected at Cocquericot, British Honduras; Costa Rica and
Panama; southward to Bolivia and the Guianas; Cuba.
A slender shrub 1.5-4.5 m. high, glabrous or nearly so; stipules caducous, 1-1.5
cm. long, membranous, ferruginous, bifid at the apex, with acute or acuminate lobes,
glabrous; leaves darkening when dried, on short petioles or almost sessile, rather thick
and firm, obovate or obovate-oblong, mostly 8-16 cm. long and 3-5 cm. broad, acute
or subacuminate, cuneately attenuate to the base; inflorescence terminal, peduncu-
late, the flowers glomerate and sessile, the glomerules spicate along the branches of a
sparsely branched, small or rather large panicle, the bracts small and inconspicuous,
caducous; calyx small and very short, truncate or obscurely dentate; corolla white
and very small; fruit red, globose, 4-5 mm. long, the nutlets costate dorsally.
Easily recognized because of the spikelike branches of the
inflorescence.
Psychotria yunckeri Standl. in Yuncker, Field Mus. Bot. 17:
397. 1938.
A forest or cloud forest species of middle to rather high
elevations, 1,000-1,800 m.; Jalapa; Quezaltenango; San Marcos.
Honduras (type, Yuncker et al 6013).
A shrub or small tree 2-8 m. tall, entirely glabrous, the stipules caducous,
connate at first, calyptriform, to 3.5 cm. long and 3.5 mm. broad; leaves elliptic or
elliptic-oblong, acuminate, attenuate or subdecurrent at the base, 10-20 cm. long and
4-9 cm. broad, lateral nerves about 15 pairs, the petiole of mature leaves to 7 cm.
long; inflorescence terminal, cymose-paniculate, many-flowered, pedunculate, to
about 10 cm. long; flowers white or greenish, short pedicellate or sessile; hypanthium
obovoid, about 1.5 mm. long, the calyx truncate or obscurely dentate, about 1 mm.
long; corolla broadly cylindric, about 5 mm. long, glabrous outside, barbate in the
throat, the lobes ovate-oblong and nearly as long as the tube; fruits ovoid or globose,
to 7-8 mm. long.
168 FIELDIANA: BOTANY, VOLUME 24
This is one of the several species often referred to P. trichotoma
Mart. & Gal, a Mexican species that may not occur in Central
America.
RANDIA Linnaeus
Trees or shrubs, usually armed with axillary or supra-axillary spines, the spines
often borne at the ends of the branches; stipules small, intra-petiolar; leaves opposite,
sessile or petiolate, membranaceous or coriaceous; inflorescences axillary or terminal;
flowers solitary or fasciculate, small or large, perfect or unisexual, usually white,
sometimes yellowish; hypanthium ovoid, obovoid, or turbinate, terete or costate;
calyx usually tubular, and lobate, the lobes often elongate or foliaceous; corolla
generally salverform, with short or elongate tube, the throat glabrous or villous, the
limb usually 5-lobate, the lobes short or elongate, acute or obtuse, contorted in bud;
stamens 5, inserted in the corolla throat, the filaments short or obsolete; anthers
dorsifixed, linear, obtuse to acuminate, included or exserted; ovary commonly 2-
celled, the style glabrous or pilose, the stigma clavate or fusiform, entire or bilobate;
ovules numerous, immersed in fleshy placentae, these affixed to the septum; fruit
baccate, globose or oval, 2-celled, the pericarp thick and hard or soft and thin,
usually smooth, sometimes tuberculate; seeds numerous or few, immersed in pulp,
commonly horizontal, compressed, the testa usually thin.
There are about 200 species of Randia distributed in the
tropics of both hemispheres. Sixty-five species have been described
from Mexico, Central America, and Panama. The genus is a
complex one easily divided into two subgenera in our region.
Basancantha Hook. f. which was distinguished by its author on the
basis of unisexual flowers is of doubtful validity. The genus in
North America is much in need of revision and until this is done no
satisfactory account of the species can be written. Much of the
material in herbaria is sterile or fragmentary so that revisionary
work would be difficult. The unisexual flowers and perhaps
dioecious plants of part of the species would not make easier
revisionary work.
Corolla 25-70 mm. long, rarely slightly shorter; fruit (unknown in R. pleiomeris) 3-8
cm. long; flowers often or always unisexual; spines often in 4's at the ends of the
branches. [Subgenus Basanacantha (Hook, f.) L. Wms.].
Leaves rounded at the apex, almost glabrous, 1-2 cm. long R. pleiomeris.
Leaves acute or acuminate, rarely obtuse or rounded at the apex but then densely
pubescent and 4-10 cm. long or larger.
Leaves densely tomentose beneath with pale matted hairs, obtuse or rounded at
the apex R. habrophlebia.
Leaves glabrous beneath or hirsute or pilose with straight, not matted, usually
somewhat appressed hairs, acute or acuminate.
Leaves mostly glabrous beneath or nearly so; corolla usually glabrous,
sometimes puberulent or sparsely short-pilose; fruit usually longer than
broad.... R- armata.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 169
Leaves densely pilose or hirsute beneath, especially when young; corolla
densely pilose with long, white, appressed or spreading hairs; fruit globose.
Corolla tube 3.5-4 cm. long or longer.
Calyx lobes linear or linear-cuneiform, about 6 mm. long R. monantha.
Calyx lobes filiform, 10-25 mm. long R- gentlei.
Corolla tube about 1.5 cm. long R- lundelliana.
Corolla 5-20 mm. long, rarely somewhat larger; fruits small, usually less than 2 cm.
long; flowers perfect; spines chiefly in pairs along the branches. [Subgenus
Randia].
Fruit or ovary pilose with spreading hairs R. standleyana.
Fruit and ovary glabrous or appressed-pilose.
Leaves densely short pilose on both surfaces; corolla 12-20 mm. long.
R. letreroana.
Leaves glabrous or sparsely pilose below; corolla 5-10 mm. long.
Corolla naked in the throat R. cookii.
Corolla white-barbate in the throat.
Corolla 4-5 mm. long, the lobes apiculate, shorter than the tube.
R. guatemalensis.
Corolla 6-8 mm. long, the lobes acute or acuminate, equaling or longer than
the tube R. aculeata.
Randia aculeata L. Sp. PL 1192. 1753. R. mitis L. I.e.
Dry hillsides and thickets at relatively low elevations, possible
along most Caribbean shores from Florida and the West Indies to
northern South America. In Guatemala possibly only in Pete"n and
Izabal. British Honduras and along the Caribbean coast to Panama.
A stout, usually densely branched shrub 1-3 m. high, rarely a small tree, the
branchlets glabrous or often scaberulous or appressed-pilose, usually bearing at the
apex 2 stout spines 1.5 cm. long or less; leaves mostly clustered at the ends of the
branches or on short lateral spurs; stipules 2 mm. long or less, ovate-deltoid, generally
acuminate, pilose within at the base; petioles very short or none, usually marginate
to the base; leaf blades very variable in shape and size, mostly obovate or obovate-
orbicular but often orbicular to elliptic-oblong or rhombic-ovate, commonly 1-6 cm.
long and 0.5-3 cm. broad but often considerably larger, acute to broadly rounded at
the apex, broadly rounded to attenuate at the base, coriaceous at maturity, usually
lustrous and glabrous above, glabrous beneath or sparsely pilose along the costa;
flowers perfect, terminal, sessile, often clustered; calyx and hypanthium 2-3 mm.
long, usually glabrous, the calyx lobes linear to ovate, generally much shorter than
the hypanthium, often ciliolate; corolla white, 6-8 mm. long, glabrous outside, the 5
lobes ovate or ovate-oblong, acute or acuminate, equaling or longer than the tube,
the throat very densely white-barbate; anthers exserted; fruit globose, 6-13 mm. in
diameter, usually smooth and glabrous; seeds commonly 5-10, brownish black, 4-5
mm. long.
It is difficult to delimit Randia aculeata and until someone
finds the time to revise the genus no satisfactory account can be
given of this entity. Herbaria contain a very large number of
collections more often than not without flowers. The name R.
170 FIELDIANA: BOTANY, VOLUME 24
aculeata has been appended to material collected at sea level up to
2,000 m. and from several ecological situations over much of Mexico
and Central America by the authors of this work, and by many
other botanists as well. Certainly some of these plants are not of
the same species and the highlands of Guatemala definitely support
allied species not accounted for in the treatment presented here.
Randia armata (Swartz) DC. Prodr. 4: 387. 1830. Mussaenda
spinosa Jacq. Sel. Stirp. Am. 70. 1763. Gardenia armata Swartz,
Prodr. Ind. Occ. 51. 1788. Randia ovata Duchass. ex Griseb.
Bonplandia 6: 8. 1858. R. spinosa Karst. Fl. Columb. 2: 128, t. 167.
1866, not R. spinosa Poir. 1811. Basanacantha spinosa var.
guatemalensis Schum. ex Loes. Verh. Bot. Ver. Brandenb. 65: 110.
1923 (type collected in forest below Alotenango, Sacatepequez, Seler
2570). Flor de cruz; palo de cruz; crucito; rosetillo; torolillo.
Dry or moist thickets or forest of the lowlands of both coasts,
ascending in the Pacific bocacosta to about 1,200 m.; Peten; Izabal;
Santa Rosa; Baja Verapaz (seen but not collected); Guatemala (?);
Escuintla; Sacatepequez; Suchitepequez; Retalhuleu; doubtless in
all the Pacific coast departments. Mexico to Panama, southward to
Argentina.
A shrub or small tree 2-9 m. high, the branchlets appressed-pilose or almost
glabrous, usually bearing at the apex 4 stout spines 1-2.5 cm. long; bark thin, brown,
fibrous; stipules broadly ovate, 3-8 mm. long, mucronate-acuminate, thin, brownish;
leaves commonly membranaceous, on slender petioles 2 cm. long or less, variable in
shape, mostly ovate, oblong-ovate, oval, or obovate, 6-20 cm. long, 2-10 cm. broad,
acute or abruptly acuminate, at the base acute or acuminate, bright green
above, glabrous or sparsely puberulent, beneath puberulent or minutely appressed-
pilose along the veins or almost glabrous; flowers perfect, slender-pedicellate,
fragrant, 2-8 at the end of each branchlet, the pedicels glabrous or puberulent; calyx
and hypanthium glabrous or puberulent, the 5 calyx lobes linear to obovate-oblong,
4-9 mm. long, acute or acuminate; corolla white or ochroleucous, glabrous outside or
sparsely pilose or short-villous, the tube 2-2.5 cm. long, the throat naked, the 5 lobes
rhombic-obovate, 1 cm. long, obtuse; fruit usually oval, 1.5-3.5 cm. long, smooth or
obscurely costate, the pulp at maturity black, sweetish; seeds numerous, suborbicular,
brown, 4-6 mm. broad.
Known in Honduras by the names "cagalera," "crucetilla," and
"jazmin cimarron," "caca de mico," "jicarillo" (El Salvador). The
flowers are sweet-scented. The ripe fruit has a hard but rather thin
shell that is readily punctured by some birds, which seem to be fond
of the pulp and seeds. This pulp sometimes is eaten by people, but
its black color makes it repulsive in appearance and the flavor is
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 171
unpleasant. The fruit is said to have been used in the Antilles for
stupefying fish and the wood is employed for fuel.
Randia cookii Standl. N. Am. Fl. 32: 176. 1934. Crucita;
naranjillo; conchitam (Huehuetenango).
Dry or moist thickets, often on rocky hillsides, 800-2,100 m.;
Baja Verapaz, the type from Santa Rosa, O. F. Cook 249;
Huehuetenango. Mexico (Chiapas); Honduras.
A very densely branched shrub 1-2 m. high, the branches appressed-pilose at first,
the spines in pairs along or at the tips of branches, 6-9 mm. long; leaves mostly
clustered on short lateral spurs, often blackening in drying; stipules 1.5-3 mm. long,
rounded-deltoid, acuminate, glabrous within; petioles 1-1.5 mm. long; leaf blades
obovate or rounded-obovate to oval-oblong or obovate-oblong, 1-3.5 cm. long, 6-12
mm. broad, acutish to rounded at the apex, obtuse to acuminate at the base,
coriaceous or subcoriaceous, glabrous and lustrous above, glabrous beneath or
sparsely scaberulous on the costa, the margin often revolute; flowers perfect, terminal
and mostly solitary, sessile; calyx and hypanthium 2-2.5 mm. long, the calyx
strigillose outside, pilose within, the lobes linear, triangular, or oblanceolate, equaling
or shorter than the hypanthium; corolla 6 mm. long, white, the throat naked, the
rounded lobes about half as long as the tube; fruit globose, greenish white, 8-12 mm.
in diameter, smooth, glabrous, with thin pericarp and few seeds.
The shrub is abundant on the dry rocky hills about Santa Rosa
and in other nearby localities in Baja Verapaz.
Randia gentlei Lundell, Wrightia 4: 125. 1969. Wild calabash.
In wet thickets or forest, Pete"n; Alta Verapaz. British
Honduras. The type from British Honduras, Gentle 7343.
Small unarmed trees or shrubs, the thick branchlets hirsute at first; leaves short
petiolate, clustered at the ends of branches, ovate-elliptic or oblanceolate, 20-45 cm.
long and 5.5-13.5 (-20) cm. broad, acuminate, sparsely hirsute above, short pilose
below, lateral nerves 10-12 pairs; flowers clustered at the ends of branches, subsessile;
hypanthium and calyx strigose, the lobes 5-6, filiform, 1-2.5 cm. long, strigose-hirsute;
corolla 4.5-8.5 cm. long, strigose-hirsute, the tube 4-4.5 cm. long, the lobes lanceolate,
acuminate, to 4 cm. long; fruits globose or ellipsoid, up to 8 cm. long, and 6.5 cm. in
diameter when immature.
We have seen no specimens of this species. The description is
taken from the original. See also R. monantha.
Randia guatemalensis Standl. Contr. U. S. Nat. Herb. 20:
202. 1919.
Type collected near Secanquim, Alta Verapaz, 550 m. Pittier
271.
172 FIELDIANA: BOTANY, VOLUME 24
Branches ferruginous, the branchlets densely puberulent when young, bearing at
the apex 2 stout spines 4-8 mm. long, the leaves fasciculate in the axils or on short
spurs; stipules ovate-deltoid, 1-1.5 mm. long, mucronate; petioles 11 mm. long or less,
the blades oblong-elliptic or broadly obovate sometimes broadly ovate or
suborbicular, 5.5 cm. long and 3 cm. broad or smaller, obtuse or acutish, often
mucronate, rounded to attenuate at the base, subcoriaceous, lustrous above,
puberulent along the costa, minutely pilose beneath along the costa, the lateral
nerves 5-8 on each side; flowers perfect, solitary, sessile; calyx and hypanthium 1.5
mm. long, scaberulous, the calyx lobes minute, triangular-subulate; corolla 4-5 mm.
long, glabrous outside, acuminate in bud, the throat densely white-barbate, the lobes
broadly ovate, apiculate, shorter than the tube.
It is doubtful whether this is distinct from R. aculeata.
Randia habrophlebia Standl. & Steyerm. Field Mus. Bot. 23:
25. 1943.
Type collected along the river near Jutiapa, Jutiapa, 850 m.,
Standley 76321; also in Huehuetenango (region of Santa Ana
Huista, 900 m.). Mexico (Chiapas).
A dense shrub 2-3 m. high or a small tree, sometimes 6 m. high, with stout
branchlets, the leaves mostly crowded at the ends of the branches; spines very few
and small on the type specimens but normally doubtless well developed and in four's
at the ends of branches; stipules glumaceous, brown, strigose outside, densely long-
sericeous within; leaves short-petiolate, herbaceous, on petioles 6-15 mm. long,
suborbicular to rounded-ovate or broadly elliptic, 6.5-9.5 cm. long, 5.5-7.5 cm. broad,
broadly rounded to obtuse at the apex, at the base broadly rounded or obtuse, very
densely velutinous-pilose above, more or less bullate, the nerves and veins impressed,
grayish or ochraceous beneath and densely tomentose with matted hairs, the lateral
nerves about 9 on each side, ascending at a very narrow angle, all the veins and
nerves very prominent and reticulate; fruit terminal, solitary, sessile, globose, 3.5 cm.
in diameter, almost smooth, densely pilose with subappressed hairs; seeds very
numerous.
Randia letreroana Lundell, Wrightia 4: 126. 1969.
The type from near Letrero, Siltepec, Chiapas, alt. 2,000 m.,
Matuda 4358, Not known in Guatemala but to be expected.
Small trees, the branchlets with small paired spines; leaves drying blackish,
obovate or obovate-elliptic, rounded at the apex and cuneate to the base, densely and
shortly pilose on both surfaces, 1-4.5 cm. long and 0.8-2.4 cm. broad; inflorescence a
1 -few-flowered axillary or terminal fascicle on the short shoots; calyx lobes linear-
subulate, to 3 mm. long, ciliate; corolla 12-20 mm. long, the tube very narrow, the
lobes lanceolate-elliptic, mostly less than 10 mm. long.
A highland species distinguished by the pilose leaves from those
known or expected in Guatemala.
Randia lundelliana Standl. in Lundell, Contr. Univ. Mich.
Herb. 4: 30. 1940.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 173
Pete"n (La Libertad). British Honduras, the type from Vaca, El
Cayo District.
A shrub or small tree, the trunk as much as 7.5 cm. in diameter, the branchlets
densely hirsute with long, spreading brownish hairs, bearing at the apex 4 stout
brown spines; stipules triangular-ovate, brown, 7-10 mm. long, attenuate-acuminate,
strigose outside; leaves short-petiolate, herbaceous, broadly ovate to oblong-ovate or
elliptic-obovate, 3-9 cm. long, 1.5-3.5 cm. broad, acute or acuminate, rounded to acute
at the base, densely puberulent and short-hirsute above, densely hirsute beneath with
straight, appressed or somewhat spreading, pale hairs, the lateral nerves about 6
pairs; flowers unisexual, the staminate ones densely aggregate at the ends of the
branches, sessile; hypanthium 4 mm. long, hirsute with subappressed hairs; calyx
lobes 6-7 mm. long or longer, filiform-attenuate from a lance-ovate base; corolla
white, densely sericeous outside with long white hairs, the tube about 1.5 cm. long,
slightly dilated in the throat, the lobes lanceolate, 1-1.5 cm. long, narrowly long-
attenuate, glabrous within; fruit globose, smooth, densely pubescent, 3.5 cm. in
diameter.
This plant has been identified incorrectly with R. watsonii
Rob. and with R. albonervia Brandegee, and has been reported
under the latter name from Peten. Both those species are Mexican,
and unknown from Central America. It was suspected by the senior
author that R. lundelliana could not be maintained as distinct from
R. monantha. Material of both species is inadequate, but it seems
possible now that R. monantha and R. lundelliana are sibling
species, the first from the Pacific side of Central America and
Mexico, the second from the Atlantic side.
Randia monantha Benth. PL Hartw. 84. 1841. Basanacantha
monantha Hook. f. ex Hemsl. Biol. Cent.-Am. Bot. 2: 39. 1881. Es-
pino; naranjillo (Santa Rosa); Jujute (Jutiapa).
Dry or wet thickets, on slopes or along streams, 1,200 m. or
lower; Peten (?); Alta Verapaz; Izabal; Chiquimula; Jutiapa; Santa
Rosa (type from Rio de los Esclavos, probably at the old bridge
near Cuilapa, Hartweg 582); Escuintla; Sacatepequez (near Las
Lajas). Reported from Veracruz.
A shrub or small tree 3-6 m. high, the thick branchlets usually bearing at the
apex 4 stout spines 1-2 cm. long; stipules glumaceous, broad, acuminate, brown,
sericeous within; leaves clustered at the ends of the branchlets, membranaceous, on
slender petioles 2 cm. long or less, obovate to ovate or elliptic, 6-17 cm. long, 3-9.5 cm.
broad, short-acuminate or obtuse and mucronate at the apex, acute at the base or
abruptly contracted and decurrent, short- hispidulous above or sometimes pilose with
long stiff hairs, short-pilose or hispidulous beneath, the hairs mostly appressed;
flowers unisexual, terminal, sessile, solitary or clustered; hypanthium short-pilose;
calyx lobes subequal, linear or linear-cuneiform, 6 mm. long, acute; corolla densely
villous outside with long white subappressed hairs, the tube 4-5 cm. long, slender,
slightly dilated above, the lobes ovate-oblong to linear-lanceolate, 2.5-3.5 cm. long,
174 FIELDIANA: BOTANY, VOLUME 24
acuminate or long-attenuate; fruit globose, 3.5-4 cm. in diameter, obscurely costate,
the shell very thick and hard, cream-colored when fresh but drying blackish,
containing very numerous seeds.
The Central American species of this alliance are poorly
understood because of the lack of ample flowering specimens. See
account of R. lundelliana. The material of this species is
inadequate— some probably does not belong here.
Randia pleiomeris Standl. Contr. U. S. Nat. Herb. 20: 202.
1919.
Collected at Santa Rosa, Dept. Santa Rosa, 900 m., Heyde &
Lux 3166. Possibly in El Salvador.
Branches slender, brownish, strigose when young, the spines in pairs along the
branches, stout, ascending, 1-1.5 cm. long, the leaves crowded on short lateral spurs;
stipules ovate-deltoid, 2 mm. long, pilose within at the base; petioles slender, 4-8 mm.
long; leaf blades cuneate-orbicular or broadly obovate, 8-18 mm. long, 7-13 mm.
broad, cuneate or abruptly decurrent at the base, rounded or truncate at the apex,
membranaceous, glabrous above, sparsely appressed-pilose beneath along the costa;
flowers solitary, terminal, sessile; hypanthium 2.5 mm. long, appressed-pilose; calyx
glabrous, 2 mm. long, the lobes usually 7, linear, 4 mm. long, sparsely ciliate; corolla
salverform, glabrous outside, the slender tube 2.5 cm. long, the 5 lobes ovate or ovate-
oblong, 1 cm. long, glabrous within, the throat naked; anthers subexserted.
Apparently a rare species, not represented in recent
Guatemalan collections. Even in the sterile state the species should
be recognizable because of its distinctive leaves. This species was
distributed by Captain Smith as R. xalapensis Mart. & Gal.
Randia standleyana L. Wms. Phytologia 24: 162, fig. 1972.
Figure 28.
Sparse forests or clearings, altitude about 200 m., endemic and
known only from near La Libertad, Pete"n (type, Lundell 3474).
Shrubs or perhaps small trees of unknown size; the branchlets opposite, mostly
about 4-5 cm. long and terminated by a pair of spines 6-10 mm. long, sparsely pilose
or glabrescent; the leaves usually 4 on very short opposite short-shoots, sessile or
nearly so, obscurely puberulent on both surfaces, the blades obovate to broadly
obovate, obtuse, attenuate to the base, mostly 7-15 mm. long and 3-10 mm. broad;
inflorescence a sessile 1-few-flowered fascicle terminal on short shoots; flowers very
small, mostly 4-5 mm. long; ovary densely white pubescent; calyx campanulate, 5-
lobate, glabrous, about 1-1.5 mm. long, the lobes lanceolate, acute, ciliate, to 0.8 mm.
long; corolla salverform, 3-4 mm. long, 5-lobate, glabrous outside, sparsely pubescent
in the throat, the tube 2-2.5 mm. long, lobes spreading, ovate or suborbicular, obtuse,
1.5-2 mm. long; stamens nearly sessile in the throat of corolla; style as long as the
corolla tube, the stigma bifid; fruit not known.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 175
The species most closely related is R. cookii Standl., possibly a
sibling species from the Pacific side of Guatemala and Chiapas.
EXCLUDED OR NOT PLACED
Randia longiloba Hemsl. Biol. Cent.-Am. Bot. 4: 101. 1887.
Dry xerophytic forests of the northern tip of the Yucatan
Peninsula; reported by Lundell (Wrightia 4: 126. 1969) from Pete"n.
We have seen no specimens of this species from the Pete"n.
Randia petenensis Lundell, Wrightia 4: 127. 1969.
Wet thickets or forests, known only from Pete"n, type Contreras
6941.
Trees to about 12 m. tall and 0.15 m. in diameter. Branchlets slender, with or
without spines and when present in pairs along the branches; stipules broadly ovate,
subulate, about 6 mm. long; the short petiolate leaves up to 14.5 cm. long and 8.5 cm.
broad, ovate or ovate-elliptic, acuminate, the base narrowed and decurrent,
secondary nerves 5 or 6 pairs; fruits clustered on older wood, sessile, glabrous, globose
and up to 1 cm. in diameter, the persistent hypanthium about 1 mm. high.
Flowers are not known and the relationship of the species is
uncertain.
RELBUNIUM (Endl.) Bentham & Hooker
Reference: F. Ehrendorfer, Revision of the Genus Relbunium,
Bot. Jahrb. 76: 516-553. 1955.
Plants usually herbaceous and perennial, in habit like Galium; leaves mostly in
whorls of 4, membranaceous or coriaceous; flowers perfect, axillary, solitary,
pedicellate, the pedicels articulate with the flower, the flowers surrounded by an
involucre of usually 4 bracts; hypanthium globose, the calyx obsolete; corolla rotate,
the 4 lobes valvate in bud; stamens 4, inserted in the corolla tube, the filaments
short, the anthers didymous, exserted; ovary 2-celled, the 2 styles short, free or
connate at the base, the stigmas capitate; ovules solitary, affixed to the septum; fruit
didymous, coriaceous or fleshy, smooth or granulate, 2-seeded.
About 30 species, ranging from the southwestern United States
to the southern extremity of South America. Distinguished readily
from Galium by the involucrate flowers. No other species are
known from Central America.
Fruits pilose; stems abundantly pilose R. hypocarpium.
Fruits and stems glabrous R, microphyllum.
176 FIELDIANA: BOTANY, VOLUME 24
Relbunium hypocarpium (L.) Hemsl. Biol. Cent. Am. Bot. 2:
63. 1881. Vaillantia hypocarpia L. Syst. Nat. ed. 10. 1307. 1759.
Rubia hypocarpia DC. Prodr. 4: 591. 1830. Hierba peluda (fide
Aguilar). Figure 67.
Common and widely distributed at middle and higher
elevations, 1,200-3,000 m., wet or moist forest or thickets, more
often on open banks or cliffs, sometimes in oak-pine forests; Alta
Verapaz; Zacapa; Chiquimula; Jalapa; Santa Rosa; Guatemala;
Sacatepequez; Quiche; Huehuetenango; Quezaltenango; San
Marcos. Southern Mexico to Panama, southward through the cooler
regions of South America.
A slender prostrate perennial, or procumbent or occasionally scandent over low
bushes, sometimes pendent from moist banks and cliffs, the stems pilose; leaves
oblong-elliptic to obovate, 5-15 mm. long, 3-8 mm. broad, rounded and usually
mucronate at the apex, membranaceous or becoming thick, deep green and often
somewhat lustrous above, pilose on both surfaces, slightly narrowed at the base;
pedicels 5-15 mm. long, the bracts ovate to oblong, acute or acutish, narrowed to a
subpetiolar base; the flowers sessile within the bracts; corolla minute, white, its lobes
ovate, ciliate, acutish; fruit orange-red, 2-3 mm. long, sparsely pilose.
A very common plant in many places in the mountains, its
abundant orange fruits sometimes rather conspicuous.
Relbunium microphyllum (Gray) Hemsl. Biol. Cent. Am.
Bot. 2: 63. 1881. Galium microphyllum Gray, PI. Wright. 1: 80. 1852.
Rubia laevigata DC. Prodr. 4: 591. 1830. Relbunium laevigatum
Hemsl. I.e.
Open forest or subalpine meadows on open banks, 2,000 m. or
m.ore, rare and local; Chimaltenango; Quezaltenango;
Huehuetenango. Ranging northward to the southwestern United
States.
A glabrous perennial, usually much branched from the base, the very slender but
rather stiff stems 30 cm. long or less, usually procumbent, the stems obtusely
tetragonous; leaves stiff, 5-15 mm. long, pungent-tipped, the margins thickened,
revolute, smooth; pedicels about as long as the leaves, the bracts similar to the leaves
but smaller, the flowers sessile; corolla greenish, the lobes ovate, subobtuse; fruits
glabrous, sometimes slightly granulose, about 3 mm. long.
RICHARDIA Linnaeus
Annual or perennial herbs, erect or prostrate, usually villous or hispid, the stems
terete; stipules united with the petioles to form a multisetose sheath; leaves opposite,
sessile or petiolate; flowers small, white or pink, arranged in dense heads, these
chiefly terminal and subtended by large leaflike bracts; hypanthium turbinate or
subglobose, the calyx 4-8-lobate, the lobes lanceolate or subulate, persistent; corolla
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 177
funnelfonn, glabrous in the throat, the limb 3-5-lobate, the lobes ovate or lanceolate,
valvate in bud; stamens 3-5, inserted in the corolla throat, with filiform filaments;
anthers dorsifixed near the base, linear or oblong, exserted; ovary 3-4-celled, the style
filiform, with 3-4 linear or spatulate branches; ovules solitary, affixed to the middle
of the septum; capsule 3-4-coccous, the apex of the capsule circumscissile, the cocci
membranaceous or crustaceous, muricate or papillose, rarely smooth, indehiscent, the
axis sometimes persistent; seeds elliptic-oblong, bisulcate ventrally.
A genus of 10 species or more, in the warmer parts of America,
chiefly in Brazil Only one species is found in Central America.
Richardia scabra L. Sp. PI. 330. 1753. Richardsonia scabra
St. Hill. PL Usu. t. 8. 1824. Golondrina blanca (Guatemala); ipeca
(fide Aguilar). Figure 59.
A common weed throughout the warmer parts of Guatemala,
especially in the plains of both coasts, ascending commonly in
settled regions to 1,500 m. or more, or in some areas to 2,000 m., and
on the Volcan de Pacaya to 2,500 m.; often a weed in cornfields,
also in waste ground generally or in pastures or thickets; Pet6n;
Alta Verapaz; Izabal; SacatepSquez; Chiquimula; Jalapa; Jutiapa;
Santa Rosa; Escuintla; Guatemala; Chimaltenango; SuchitepSquez;
Quezaltenango; Huehuetenango; Quiche". Widely distributed in
most warmer parts of America; Mexico; British Honduras to El
Salvador and Panama; south to Peru and Argentina.
Annual, usually prostrate and forming mats, sometimes erect, pilose or hispid
throughout; leaves petiolate, herbaceous, oblong or lanceolate to oblong-ovate, 2-9
cm. long, acute and mucronate, narrowed at the base, conspicuously nerved;
inflorescence capitate, with few or many flowers, usually long-pedunculate,
subtended by 2 large, broadly ovate bracts similar to the leaves; sepals lanceolate,
green; corolla white, 4-6 mm. long, glabrous outside; carpels 3, densely muriculate
dorsally, sulcate on the inner face, 2-3 mm. long.
A common weed of general distribution in most parts of
Central America.
RONDELETIA Linnaeus
Reference: Paul C. Standley, N. A. Fl. 32: 44-86. 1918.
Shrubs or trees, usually pubescent, with terete or angulate branchlets; stipules
interpetiolar, usually broad and sometimes foliaceous, obtuse to cuspidate, usually
persistent; leaves opposite or verticillate, sessile or petiolate, membranaceous or
coriaceous; inflorescence terminal or axillary, usually cymose or paniculate;
hypanthium commonly subglobose, the calyx 4-5-lobate, the lobes narrow or broad,
often unequal, persistent; corolla funnelform or salverform, white, yellowish, pink, or
red, the tube usually slender and elongate, often ampliate in the throat, the throat
often annular-thickened, sometimes barbate, the 4-5 lobes spreading, broad, obtuse,
178 FIELDIANA: BOTANY, VOLUME 24
imbricate in bud, 1 or 2 of them exterior; stamens 4-5, inserted in the corolla throat,
included or exserted, with short filaments; anthers dorsifixed, narrowly oblong or
broader, obtuse, erect; ovary 2-celled, the style filiform, obtuse or short-bifid; ovules
numerous, the placentae affixed to the septum; capsule usually small and globose,
chartaceous or coriaceous, 2-celled, loculicidally or septicidally bivalvate, the valves
commonly biparted; seeds minute, compressed or angulate, often winged or
appendaged, with thin testa.
A large genus in tropical America, about 150 species being
known from North America, and many others in South America.
Numerous other species are found in the mountains of southern
Central America, and a great many are native in Mexico.
The genus Rondeletia is a fascinating and common one in the
highlands of Guatemala. A few species are rather widely distributed
but 23 of the 31 species recognized in Guatemala, adjacent British
Honduras, and the border region in Chiapas, Mexico, are mostly
local endemics. These are often abundant in their restricted areas.
We must recognize, of course, that Guatemala and the adjacent
regions are still inadequately known and collected.
Dr. Julian A. Steyermark in his study of the Rubiaceae in
Mem. N. Y. Bot. Gard. 17: 241-261. 1967 has separated
Arachnothrix Planchon out of Rondeletia and has given reasons, in
a key, for doing this. Several of the species in this flora and, in fact,
perhaps most of them would need to be transferred to Arachnothrix
if the separation were followed.
Corolla densely yellow-barbate in the throat; stipules usually foliaceous and reflexed;
flowers 5-parted.
Corolla glabrous outside R. suffrutescens.
Corolla variously pubescent outside.
Branches sharply quadrangular; leaves usually acute at the base../?, stenosiphon.
Branches terete or nearly so; leaves subcordate to obtuse at the base.
Leaves ternate; inflorescence umbellate or subcapitate at the ends of the
branches R- strigosa.
Leaves opposite; inflorescence cymose-paniculate.
Calyx lobes linear-subulate, attenuate R. ehrenbergii.
Calyx lobes deltoid to oblong, obtuse or acutish.
Stipules subulate-deltoid from a broad base R. seleriana.
Stipules mostly oblong and foliaceous, 1 cm. long or longer, usually
obtuse.
Leaves densely soft-pilose beneath R. amoena.
Leaves glabrous beneath or sparsely strigose.
Corolla tube 8-10 mm. long R. cordata.
Corolla tube about 4 mm. long R. brachistantha.
Corolla usually naked in the throat, never yellow-barbate; stipules mostly narrow
and erect.
Inflorescence an elongate narrow spike-like panicle.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 179
Corolla arachnoid-tomentose outside, sometimes only sparsely so.
Leaves in age glabrous or glabrate, or at least never white-tomentose,
sometimes somewhat tomentose beneath when young.
Lobes of the calyx longer than the hypanthium R. septicidalis.
Lobes of the calyx shorter than the hypanthium.
Leaves tomentose beneath with loose or dense tomentum.
Corolla tube about 10 mm. long; tomentum of leaves loose and coarse.
R. skutchii.
Corolla tube about 7 mm. long; tomentum of leaves tight and fine.
R. myriantha var. armentatts.
Leaves glabrate at least in age or with sparse closely appressed arachnoid
tomentum of very slender hairs; corolla tube less than 8 mm. long.
Stipules triangular or triangular-ovate; inflorescence narrow and to 2
cm. broad or less.
Petioles of a pair of leaves unequal, one usually much longer than 2
cm.; lowland forest species R. silvicola.
Petioles of a pair of leaves subequal, mostly 1 cm. long or less; species
of wet highland forest R. gracitis.
Stipules narrowly linear-lanceolate; inflorescence mostly 2-4 cm. broad;
montane species R. rubens.
Leaves, even in age, covered beneath with a very dense and close, white
tomentum.
Corolla tube 5-8 mm. long R. buddleioides.
Corolla tube 12-15 mm. long R. laniflora.
Corolla variously pubescent outside but never tomentose.
Calyx lobes about 7 mm. long R. tacanensis.
Calyx lobes very small, rarely 2 mm. long.
Pubescence of the lower leaf surface of dense, spreading, usually rufous
hairs R. rufescens.
Pubescence of the lower leaf surface of closely appressed hairs.
Corolla tube glabrous except near the apex, there setose-pilose; leaves
thinly sericeous beneath with very long and slender hairs
R. stachyoidea.
Corolla tube pubescent throughout; leaves variously pubescent beneath
but not as above.
Hypanthium and capsule hispidulous with spreading hairs; leaves
strigose on the upper surface R. secundiflora.
Hypanthium and calyx tomentulose or strigose with appressed hairs;
leaves not strigose on the upper surface, usually glabrous.
Calyx lobes very unequal, one of them larger and foliaceous; leaves
sericeous beneath when young R. pansamalana.
Calyx lobes subequal; leaves arachnoid-tomentose to tomentose
beneath, at least when young R. myriantha & var. armentalis.
Inflorescence various but never an elongate narrow spike-like panicle.
Corolla glabrous outside R. jurgensenii.
Corolla sparsely or densely pubescent outside.
Calyx lobes, at least some of them, 4-10 mm. long or longer.
Leaves obtuse or narrowly rounded at the apex, glabrous or nearly so.
R. chinajensis.
180 FIELDIANA: BOTANY, VOLUME 24
Leaves acute or acuminate, densely pubescent or almost glabrous.
Leaves glabrous or nearly so; larger calyx lobes obtuse or rounded at the
apex.
Leaf blades oval, 4-6 cm. broad; smaller calyx lobes elliptic.
R. aetheocalymna.
Leaf blades lance-oblong, about 2 cm. broad; smaller calyx lobes linear.
R. cordovana.
Leaves sparsely or densely pubescent on both surfaces; larger calyx lobes
acuminate or attenuate.
Hypanthium appressed-pilose; calyx lobes 10 mm. long or shorter;
corolla tube 5-7 mm. long R. zolleriana.
Hypanthium hirsute with long spreading hairs; calyx lobes about 15
mm. long; corolla tube about 15 mm. long R. macrocalyx.
Calyx lobes small, 3 mm. long or shorter.
Inflorescences small, mostly 3-5-flowered; corolla very sparsely pubescent
outside R- deamii.
Inflorescence many-flowered, often large.
Stipules triangular, acuminate, about 2 mm. long R. belizensis.
Stipules oblong-spa tula te, rounded-ovate or orbicular, mostly 8-15 mm.
long or even larger.
Stipules erect, oblong-spatulate, scarcely foliaceous R. linguiformis.
Stipules reflexed, orbicular or nearly so, foliaceous R. izabalensis.
Rondeletia aetheocalymna Donn.-Sm. Bot. Gaz. 42: 298.
1906.
Alta Verapaz, the type collected between Sepacuite and
Secanquim, 550-990 m., Maxon & Hay 3275; known only from the
original collection.
A shrub 3 m. high, the branchlets sparsely strigillose or almost glabrous; stipules
narrowly oblong or spatulate-oblong, 8-10 mm. long, erect, obtuse or acute; leaves
opposite, on stout petioles 7-15 mm. long, glabrous, oval or oval-obovate, 9-15 cm.
long, 4-6.5 cm. broad, abruptly acute or acuminate, acute to abruptly attenuate at
the base, glabrous, dark green and lustrous above; inflorescences terminal or partly
axillary, cymose-corymbose, long-pedunculate, the flowers subsessile or on stout
pedicels 1 cm. long or shorter, the corymbs 5-10 cm. long and broad; bracts oblong or
linear-oblong, foliaceous, 5-12 mm. long; hypanthium densely sericeous; calyx lobes 4,
very unequal, 3 of them elliptic, acute, 2-3 mm. long, the fourth oval or elliptic, 5-8
mm. long, obtuse or rounded at the apex, green; corolla white, sericeous outside, the
stout tube 7-8 mm. long, the throat naked, the 4 lobes rounded, spreading; anthers
and style included; capsule globose, 5-7 mm. in diameter, sericeous; seeds angulate,
brown, punctate.
Rondeletia amoena (Planch.) Hemsl. Diag. PI. Mex. 26. 1879.
Rogiera amoena Planch. Fl. Serres 5: 442. 1849. Rogiera menechma
Planch. I.e. Rondeletia versicolor Hook. Bot. Mag. 77: t. 4579. 1851.
Rogiera versicolor Lindl. & Paxt. Fl. Card. 2: 69, /. 154. 1851.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 181
Rogiera latifolia Decaisne, Rev. Hort. IV. 2: 121, t. 7. 1853.
Pachatapal (fide Aguilar); xk'ac (Todos Santos, fide Seler).
Described from plants grown in Belgium from Guatemalan
seeds. Damp or wet forest or thickets, 1,200-2,250 m.; Alta Verapaz;
El Progreso; Chiquimula (Cerro Brujo); Santa Rosa (near
Oratorio); Guatemala; Quiche; Huehuetenango; Quezaltenango.
Mexico (Oaxaca) to Panama.
A shrub or small tree 1.5-9 m. high, the branches terete, densely villous-pilose
when young; stipules triangular-oblong, 10-15 mm. long, reflexed, obtuse; leaves
opposite, the stout petioles 2-10 mm. long, the blades ovate to oblong-ovate, 6-15 cm.
long, 3.5-7 cm. broad, usually abruptly short-acuminate, at the base rounded or
subcordate, glabrous or thinly pilose above, densely and softly short-pilose beneath,
petioles 5-8 mm. long, pilose; inflorescences terminal and axillary, cymose- corymbose,
5-18 cm. broad, stout-pedunculate, the cymes densely many-flowered, the flowers
sessile or short-pedicellate; bracts often large and foliaceous; hypanthium densely
fulvous-pilose, the calyx lobes 4-6, nearly unequal, oblong or obovate-oblong, 1-2 mm.
long, obtuse, erect or spreading; corolla pink, densely appressed-pilose, the stout tube
about 5 mm. long, the throat densely yellow-hirsute, the 5 lobes rounded, 2-2.5 mm.
long, puberulent within; capsule globose, 5-6 mm. in diameter, densely pilose.
A handsome shrub because of the abundant pink flowers. It
was formerly a favorite in northern hothouses, and still is grown
occasionally. The description is based on Guatemalan material.
Rondeletia belizensis Standl. Carnegie Inst. Wash. Publ. 461:
91. 1935. Bacce (Peten, Maya, fide Lundell).
Peten. British Honduras, the type collected on Jacinto Hills, 90
m., Schipp 1201.
A shrub or tree as much as 6 m. tall, the branches terete, the branchlets
puberulent when young; stipules narrowly triangular, 1.5-2 mm. long; leaves on
petioles 5-8 mm. long, the blades ovate or oblong-ovate, 3-7 cm. long, 1.5-4.5 cm.
broad, acute or subobtuse, obtuse or acute at the base, glabrous; inflorescence
terminal, cymose-corymbose, densely many-flowered, pedunculate or sessile, as much
as 5 cm. long but often much smaller and headlike, the small bracts linear, the
pedicels 2 mm. long or less; hypanthium densely whitish-puberulent, the 4 sepals
linear, erect, 1.5-2 mm. long, obtuse; corolla white, densely covered outside with
minute whitish ascending hairs, the slender tube 7-9 mm. long, glabrous in the throat,
the 4 lobes broadly rounded, 2 mm. long, villosulous within; capsule subglobose, 6
mm. broad, scaberulous-puberulent; seeds broadly winged.
Rondeletia brachistantha Standl. & Steyerm. Field Mus.
Bot. 23: 25. 1943.
San Marcos, the type collected on dry upper slopes, Volcan de
Tacana, near San Rafael and the Mexican boundary, 2,500-3,000 m.,
Steyermark 36286. Mexico (Chiapas).
182 FIELDIANA: BOTANY, VOLUME 24
A tree, the branchlets terete, appressed-pilose when young; stipules 14 mm. long,
reflexed, oblong-deltoid, brown, obtuse, glabrous within, strigose outside; leaves
subcoriaceous, on stout petioles 4 mm. long or less, ovate to oblong-ovate or elliptic-
oblong, 5-8 cm. long, 2.5-3.5 cm. broad, acute or subacuminate, rounded or narrowly
rounded at the base, glabrous or sparsely hirtellous beneath along the nerves, the
lateral nerves about 5 on each side; inflorescence terminal, cymose-corymbose,
densely many-flowered, on a peduncle 2-5 cm. long, 3-6 cm. long, 4-9.5 cm. broad, the
branches densely yellowish-strigose, the bracts minute, the flowers sessile or
subsessile; hypanthium globose, 1.5 mm. high, densely appressed-hispidulous and at
first tomentose, the calyx 0.7 mm. high; the teeth minute, remote, obtuse; corolla
pale pink, densely and minutely strigillose, the tube 4-5 mm. long, densely yellow-
barbate in the throat, the lobes rounded, 1.5 mm. long, puberulent within; style
exserted; capsule globose, 4-4.5 mm. broad, rounded at the base, glabrate,
loculicidally dehiscent; seeds compressed, dark brown, puncticulate.
Rondeletia buddleioides Benth. PL Hartweg. 69. 1840. R.
affinis Hemsl. Diag. PI. Mex. 28. 1879. Durazno (Quezaltenango).
Figure 5.
Wet mixed forest or in limestone thickets, 350-1,800 m.; Alta
Verapaz; Quiche; Huehuetenango; Quezaltenango; San Marcos.
Southern Mexico and British Honduras to Nicaragua arid possibly
to Costa Rica and Panama.
A shrub or a small tree 1.5-9 m. tall, the branches subterete, dark brownish,
densely floccose-tomentose when young but soon glabrate; stipules lanceolate or
oblong, 3-8 mm. long, obtuse to attenuate, erect; leaves opposite, on petioles 3-6 mm.
long or longer, the blades oval-elliptic to elliptic-oblong or lanceolate, 5-12 cm. long
and 2-5.5 cm. broad or larger, acute to long-attenuate, at the base obtuse to long-
acuminate, green above and sparsely pilose or usually glabrous, beneath densely
covered with a very close, white cobwebby tomentum; inflorescence terminal, the
flowers sessile or subsessile in small dense short-pedunculate cymes, these arranged in
narrow spikelike panicles 10-15 cm. long, the bractlets linear, very small; hypanthium
densely white-tomentose, the calyx lobes oblong or oval, obtuse, 1 mm. long or less,
spreading or reflexed, glabrous within; corolla variously described as yellowish, white,
or dark red, usually densely white-tomentose outside, the slender tube 4-8 mm. long,
naked in the throat, the rounded lobes 1-1.5 mm. long; capsule oblong- globose, 3-4
mm. long, glabrate; seeds minute, reticulate-puncticulate.
We presume that several species have gone under this name at
one time or another, these distributed from Mexico to Panama.
There are apparent differences between the material found from
Mexico to Nicaragua and that of Costa Rica and Panama. Careful
study may indicate a difference, or that it is all a somewhat variable
species.
Rondeletia chinajensis Standl. & Steyerm. Field Mus. Bot.
23: 254. 1947.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA
Known only from the type, Alta Verapaz, on knife edge of
limestone ridge, Cerro Chinajd, between Finca Yalpemech and
Chinaja, 150-700 m., Steyermark 45637.
A small tree of 6-9 m., the branches subterete, slender, minutely puberulent;
stipules deltoid, appressed, puberulent, 1.5 mm. long, acute or subobtuse; leaves on
petioles 8-12 mm. long, coriaceous, somewhat lustrous, elliptic-oblong or rarely oblong
or oval-ovate, 3.5-6.5 cm. long, 1.8-3 cm. broad, obtuse or narrowly rounded at the
apex, obtuse or subacute at the base, glabrous above, brownish beneath when dried,
sometimes short-barbate in the axils of the nerves, otherwise glabrous or glabrate;
inflorescences terminal, sessile or short-pedunculate, cymose but very dense and
head like, the flowers sessile or subsessile; hypanthium 1.5 mm. long, very densely
white-pubescent; calyx lobes 5, linear or narrowly spatulate-linear, mostly 4.5-5.5
mm. long, acute or subacute, sparsely puberulent below or almost glabrous,
sometimes alternating with other very short and narrow segments; corolla white,
densely hispidulous outside with spreading or subappressed hairs, the tube 14-16 mm.
long, the 4 lobes spreading, oval or obovate-oval, 6-8 mm. long, rather densely
puberulent within, the throat not barbate; capsule globose, 4.5 mm. in diameter.
Rondeletia cordata Benth. PI. Hartweg. 85. 1841. Rogiera
cordata Planch. Fl. Serres 5: 442b. 1849. R. interne Ma Hemsl. Diag.
PL Mex. 26. 1879. Huesillo (Quezaltenango); trompetal
Damp or rather dry, oak or mixed forest, often in thickets or on
cliffs, 900-2,850 m.; Santa Rosa (Volcan de Tecuamburro);
Guatemala (type collected near Guatemala, Hartweg 585);
Sacatepequez; Chimaltenango; Solola; Suchitepequez; Quiche;
Quezaltenango; San Marcos. Mexico (Chiapas); El Salvador;
Honduras.
A shrub 2-4 m. high or sometimes a tree of 9 m., the branches stout, terete,
sparsely strigose when young; stipules oblong-triangular, 6-15 mm. long, reflexed,
foliaceous, obtuse, strigose outside; leaves opposite, sessile or nearly so, the petioles 4
mm. long or less, the blades narrowly ovate-oblong to ovate or ovate-oval, 7-13 cm.
long and 3-7 cm. broad or larger, acute or acuminate, at the base rounded to
subcordate, subcoriaceous, glabrous above, sparsely strigose beneath along the veins
or glabrous; inflorescences terminal and axillary, cymose-corymbose, 5-15 cm. broad,
pedunculate, dense and many-flowered, the flowers short-pedicellate; bracts often
large and foliaceous; hypanthium strigillose, the 5 calyx lobes rounded to oblong,
unequal, 1 mm. long or less, obtuse, erect; corolla pink, thinly strigose outside, the
stout tube 8-10 mm. long, densely yellow-barbate in the throat, the 5 lobes rounded, 2
mm. long, puberulent within; anthers and style exserted or included; capsule globose,
3-4 mm. in diameter, strigillose; seeds numerous, minute, compressed, brownish
yellow.
A handsome and showy shrub, abundant in many parts of the
central highlands and in flower at the beginning of the dry season
in December and January. The species is closely related to R.
amoena.
184 FIELDIANA: BOTANY, VOLUME 24
Rondeletia cordovana Standl. & Steyerm. Field Mus. Bot. 23:
26. 1943.
Type from cloud forest in ravine bordering Quebrada
Alejandria, summit of Sierra de las Minas, vicinity of Finca
Alejandria, Zacapa, 2,500 m., Steyermark 29906.
A small, almost glabrous tree with slender terete ferruginous branchlets; stipules
subulate from a triangular base, about 3 mm. long, erect; leaves small,
membranaceous, on slender petioles 5-10 mm. long, lance-oblong, 5.5-7.5 cm. long, 1.7-
2.5 cm. broad, with a very long and narrow, attenuate acumen, glabrous above or
nearly so, much paler beneath, barbate in the axils of the nerves, elsewhere glabrous,
the lateral nerves inconspicuous, about 6 on each side; inflorescence terminal, on a
peduncle 1.5 cm. long, erect, cymose-corymbose, few-flowered, trichotomous at the
base, about 3 cm. high and 4.5 cm. broad, the flowers densely cymose-aggregate at
the apices of the primary branches, sessile or short-pedicellate, the lowest bracts
similar to the leaves but smaller; hypanthium obovoid, 1.5-2 mm. long, densely and
minutely strigillose; calyx 4-lobate, the lobes very unequal, 3 of them linear or linear-
subulate, 3 mm. long, the fourth lance-oblong, 5-6 mm. long, obtuse, glabrate; corolla
in bud broadly rounded at the apex, densely and minutely strigillose outside.
The type is a poorly developed specimen, without open corollas,
but the species represented appears to be fully distinct from R.
aetheocalymna, its nearest ally. It is dedicated to the late Alejandro
Cordova, distinguished journalist of Guatemala and proprietor of
the Finca Alejandria, where the type was collected.
Rondeletia deamii (Donn.-Sm.) Standl. N. Am. Fl. 32: 60.
1918. Bouvardia deamii Donn.-Sm. Bot. Gaz. 49: 455. 1910.
Type collected at base of a cliff near Fiscal, Guatemala, 1,110
m., Deam 6190. Usually in dry rocky thickets or along rocky stream
banks, 300-1,400 m.; Chiquimula; Guatemala; Quiche; doubtless
also in Baja Verapaz. El Salvador; Honduras.
A shrub or small tree, usually 1-2 m. high but sometimes as much as 6 m., the
branches terete, reddish brown or grayish, sparsely short-pilose when young; stipules
deltoid, acute, 1 mm. long, erect; leaves opposite, on petioles 1-2 mm. long, ovate-
orbicular or ovate, 3-6.5 cm. long, 1.5-2.5 cm. broad, abruptly acuminate or attenuate,
at the base broadly rounded to acute, glabrous above or very minutely and sparsely
pilose, minutely pilose beneath, especially along the veins; flowers mostly in terminal
3-flowered cymes, these sessile or slender-pedunculate, the slender pedicels 8 mm.
long or less, solitary long-pedicellate flowers sometimes present in the upper leaf
axils; hypanthium minutely pilose, the 4 calyx lobes linear, 1-2 mm. long, one of them
usually foliaceous, linear or linear-elliptic, and up to 4 mm. long, acute or obtuse,
erect or spreading; corolla (color not recorded) sparsely short-pilose, the slender tube
7-12 mm. long, naked in the throat, the 4 lobes rounded-oblong, 3 mm. long, minutely
papillose within; anthers and style included or the anthers partly exserted; capsule
subglobose, about 5 mm. broad, glabrate; seeds dark brown, somewhat compressed,
rather broadly and irregularly winged.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 185
The plant grows abundantly in the rocky ravines in the low
hills just northeast of Fiscal, which is presumably the original
locality.
Rondeletia ehrenbergii Schum. ex Standl. N. Am. Fl. 32: 50.
1918.
Type from Totonicapan, Ehrenberg 1033.
A shrub with terete grayish-brown branches, these densely fulvous-puberulent or
strigillose when young; stipules narrowly triangular, 5-8 mm. long, attenuate, erect;
leaves opposite, on petioles 3-5 mm. long, oval or ovate-oval, 3-5 cm. long, 1-3 cm.
broad, acute to rounded at the apex, rounded or subcordate at the base, thick-
coriaceous, densely strigillose above, densely short-pilose beneath; inflorescences
terminal and axillary, cymose-corymbose, very dense and many-flowered, 4-5 cm.
broad, the flowers short-pedicellate, the bracts elongate, linear-setaceous;
hypanthium densely fulvous-strigillose, the 5 calyx lobes linear-subulate, 2.5-4 mm.
long, erect; corolla densely strigillose outside, the slender tube 15 mm. long, the
throat densely yellow-barbate, the 5 lobes obovate-orbicular, 4 mm. long, puberulent
within; style exserted; capsule globose, 3-3.5 mm. in diameter, strigillose.
Rondeletia gracilis Hemsl. Diag. PL Nov. Mex. Cent.-Am. 53.
1880.
Wet highland forests at about 1,200 m.; Alta Verapaz (type,
Tuerckheim 404). Endemic.
Slender shrubs to 3 m. tall; branches subterete, white tomentose when young but
soon glabrous; stipules triangular or triangular-oblong, obtuse, about 5 mm. long;
leaves opposite, somewhat anisophyllous, elliptic to oblong-lanceolate, acuminate, 8-
15 cm. long and 3-6 cm. broad, almost entirely glabrous, petioles subequal, up to 1.5
cm. long; inflorescence a racemose spike about 15-20 cm. long and 2 cm. broad,
sparsely tomentose; flowers numerous, red, tetramerous, in nearly sessile heads;
hypanthium fulvous-tomentose, about 1 cm. long at anthesis; calyx lobes lance-
oblong, acute, glabrous within, erect or reflexed, about 0.5 mm. long; corolla red or
dark red, the slender tube about 7 mm. long at anthesis, thinly floccose-tomentulose,
glabrous in the throat, the lobes about 1 mm. long, anthers and style included;
capsule not known.
A rare species known only from the region of Coban, Alta
Verapaz. It is related to the lowland R. silvicola L. Wms., but is the
smallest flowered of the alliance.
Rondeletia izabalensis Standl. & Steyerm. Field Mus. Bot.
22: 286. 1940.
Wet forest at or near sea level, Izabal; type collected at Escoba,
across the bay from Puerto Barrios, Standley 72891. Endemic.
A shrub or small tree, 6 m. tall or less, the branches subterete, the young ones
sparsely strigose; stipules foliaceous, subreniform, recurved, about 1 cm. long, broadly
186 FIELDIANA: BOTANY, VOLUME 24
rounded at the apex, deeply cordate at the base; leaves firm-membranacous, on stout
petioles 6-15 mm. long, oblanceolate-oblong to oblong-obovate or elliptic- lanceolate,
11-23 cm. long, 5-9.5 cm. broad, acute to long-attenuate, cuneate at the base or
narrowly attenuate, the very base sometimes subobtuse, glabrous above, minutely
appressed-pilose beneath, the lateral nerves about 9 on each side; inflorescence
terminal, on a peduncle 7-9 cm. long, erect, cymose-corymbose, about 5 cm. long and
7 cm. broad, densely many-flowered, the basal bracts foliaceous, the upper ones
linear-attenuate and 3-10 mm. long, the branches densely sericeous-strigillose, the
flowers densely crowded, sessile; hypanthium 1 mm. long, densely appressed-pilose,
the calyx 1 mm. long, appressed-pilose, 4-lobate, the lobes ovate, obtuse or acute,
unequal; corolla white, densely pilose with subappressed hairs, the slender tube 5-6
mm. long, the throat not barbate, the 4 lobes rounded, 1.5-2 mm. long, glabrous
within.
Rondeletia jurgensenii Hemsl. Diag. PL Mex. 29. 1879;
Bullock in Hook. Icon. 34: t. 3322. 1936.
Quezaltenango, oak forest, Volcan de Santa Maria, at 1,500-
3,000 m. Southern Mexico; El Salvador; Honduras.
A shrub or small tree 2-4.5 m. high, glabrous almost throughout, the young
branchlets sometimes pilose; stipules small, deltoid, acute; leaves membranaceous, on
petioles 4-6 mm. long, narrowly ovate to elliptic-lanceolate or oblanceolate, 9.5 cm.
long and 3.5 cm. broad or smaller, acute or acuminate, at the base acute or cuneate,
almost or quite glabrous, the margins usually ciliate; flowers pink, 4-parted, the
inflorescences terminal, cymose-corymbose, lax and rather few-flowered, the pedicels
3-5 mm. long, the small bracts subulate; hypanthium 2 mm. long, pilose, the calyx
lobes subulate, 2-4 mm. long, pilose, often unequal; corolla 1.5 cm. long, glabrous, the
spreading lobes broadly elliptic to rounded, 2.5 mm. long; capsule glabrous, 4.5-6 mm.
broad; seeds numerous, prismatic.
Rondeletia laniflora Benth. PI. Hartweg. 85. 1841.
Arachnothryx laniflora Planch. Fl. Serres 5: 442. 1849. Papelillo.
Wet mixed mountain forest, 1,400-3,000 m.; El Progreso;
Zacapa; Jalapa; Guatemala (Palencia); Suchitepequez (type from
Las Nubes, Hartweg 584); San Marcos; Huehuetenango. Southern
Mexico; El Salvador.
A shrub or small tree, 2-7.5 m. high, the branches subterete, densely and closely
white-tomentose at first; stipules narrowly triangular or subulate, 3-8 mm. long,
erect; leaves opposite, crowded at the ends of the branches, on stout petioles 4-10
mm. long, obovate to narrowly lance-elliptic, 6-13 cm. long, 1-4 cm. broad, or
sometimes larger, long-attenuate or abruptly acuminate, at the base cuneate to
attenuate, subcoriaceous, dark green and glabrous above or when young thinly
floccose-tomentulose, white beneath and covered with a very dense and close
tomentum; inflorescence terminal, the flowers sessile or nearly so in dense short-
pedunculate cymes, these arranged in a dense narrow panicle 4-7 cm. long; bractlets
subulate, tomentose; hypanthium densely white-tomentose, the 4 calyx lobes
subequal, oblong, obtuse or subacute, spreading, about equaling the hypanthium;
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 187
corolla dark red, the slender tube 12-15 mm. long, densely and closely white-
tomentose outside, glabrous in the throat, the 4 lobes suborbicular, 2-3 mm. long;
capsule subglobose, 4-5 mm. long, costate, brown, glabrate; seeds minute, irregular,
brown, scrobiculate.
Rondeletia linguiformis Hemsl. Diag. PL Mex. 29. 1879.
Type collected in Guatemala by Skinner, the locality unknown,
but probably in the region of Quirigua, Izabal, frequent on forested
or brushy slopes below the pine forest, Montana del Mico and
vicinity, 40-600 m.; endemic.
A shrub or small tree 3-6 m. high, the branches subterete, glabrous; stipules
obovate-spatulate or oblong-spatulate, 6-12 mm. long, erect, rigid, mucronate; leaves
opposite, on petioles 4-8 mm. long, coriaceous, ovate-oblong or elliptic-oblong,
sometimes oval-elliptic, 7-10 cm. long, 3-6 cm. broad, obtuse to long-acuminate,
attenuate at the base, glabrous; inflorescence terminal, cymose-corymbose, open or
dense, long-pedunculate, the flowers very numerous, subsessile, the bracts linear;
hypanthium densely and minutely appressed-pilose, the 4 calyx lobes unequal,
oblong-obovate, 0.5-1 mm. long, suberect; corolla white, strigose outside, the tube 8
mm. long, the throat naked, the 4 lobes rounded, 4 mm. long or less; capsule
subglobose, 6 mm. in diameter; seeds numerous, angulate, reticulate-tuberculate.
The flowers are fragrant.
Rondeletia macrocalyx Standl. & Steyerm. Field Mus. Bot.
23: 254. 1947.
Known only from the type, Izabal, Cerro San Gil, 300-900 m.,
in dense wet forest, Steyermark 41864.
A small tree of 6-7.5 m., the branches slender, when young densely hirsute with
long, spreading, rather stiff, sordid hairs, the uppermost internodes mostly very short;
leaves very thin on hirsute petioles 6-12 mm. long, elliptic or ovate, 9-17 cm. long, 4-7
cm. broad, short-acuminate or cuspidate-acuminate, obtuse to attenuate-cuneate at
the base, rather densely hirsute on both surfaces with long slender spreading hairs,
the lateral nerves about 7 pairs; inflorescences axillary or subterminal, cymose, 3-9-
flowered, half as long as the leaves or shorter, the peduncles very slender, hirsute, 3-
5.5 cm. long, the flowers on short or elongate pedicels; calyx lobes 4, narrowly
lanceolate, subequal, about 15 mm. long, long-attenuate, foliaceous, hirsute on both
surfaces; corolla white, densely hirsute outside with long, ascending or spreading,
sordid hairs, the slender tube 18-25 mm. long, the 4 lobes spreading, obovate-oval, 7-8
mm. long, broadly rounded at the apex, glabrous within, the throat not barbate;
immature capsule oblong-ovoid, 9 mm. long, 5 mm. broad, densely hirsute; seeds very
numerous, small, angulate.
This is related to R. mexiae Standl. of Oaxaca, which has a
smaller calyx and a much less conspicuous pubescence of short
hairs.
188 FIELDIANA: BOTANY, VOLUME 24
Rondeletia myriantha Standl. & Steyerm. Field Mus. Bot. 22:
288. 1940.
Brushy slopes and banks, at about 1,000 m., Suchitepe'quez
(type from Finca Moc£, Skutch 1569). Known only from the type.
A slender shrub 3.5 m. tall or more, the branches terete, puberulent and
sometimes tomentulose when young; stipules lance-triangular, attenuate, erect;
leaves on petioles 7-15 mm. long, membranaceous, lance-oblong, 7-15 cm. long, 3.5-5
cm. broad, narrowly attenuate-acuminate, at the base acute or obtuse, green above
and glabrous or pilosulous only along the nerves, beneath at first laxly and sparsely
arachnoid-tomentulose, glabrate in age; inflorescence terminal, erect, narrowly
elongate-thyrsoid and spikelike, pedunculate, about 8 cm. long and 2.5 cm. broad,
laxly many-flowered, the flowers sessile or short-pedicellate; lowest bracts foliose but
small, the upper ones linear, green; hypanthium almost 1 mm. long, densely white-
tomentose, the calyx 4-dentate, the teeth scarcely 1 mm. long, unequal, oval or
oblong and very obtuse; corolla white, sparsely appressed-pilosulous, the slender tube
5 mm. long, the 4 lobes orbicular, 1.5 mm. long, glabrous within, the throat not
barbate; stamens included; style short-exserted; capsule subglobose, costate, glabrate,
2.5-3 mm. long.
Rondeletia myriantha var. armentalis L. Wms. Phytologia
26: 127. 1973.
In moist woods at about 2,000 m., known only from the type
from near Nebaj, Quiche, Skutch 1776.
Differs from the var. myriantha in having leaves densely tomentose below; the
hypanthium only sparsely pubescent; the calyx lobes subequal, narrowly triangular,
prominently reflexed at anthesis, and with a minute gland in the sini of the lobes; the
style included; the anthers somewhat exserted.
Rondeletia pansamalana Standl. N. Am. Fl. 32: 58. 1918.
Alta Verapaz; type from Pansamala, 1,140 m., Tuerckheim 897.
A slender shrub, the branches terete, strigillose when young; stipules triangular-
subulate, 4-6 mm. long, erect; leaves opposite, on slender petioles 3-15 mm. long,
elliptic or ovate-elliptic, 9-15 cm. long, 3-4.5 cm. broad, attenuate or long- attenuate at
the apex, abruptly acute to attenuate at the base, green and glabrous above,
strigillose beneath along the veins; inflorescence terminal, pedunculate, the flowers
short-pedicellate, in loose few-flowered pedunculate cymes, these arranged in a
narrow thrysiform panicle about 10 cm. long; bracts small, linear; hypanthium finely
strigillose, pyriform-globose, the 4 calyx lobes very unequal, 3 of them small, lance-
linear, and attenuate, the fourth elliptic-oblong, 2.5-3 mm. long, foliaceous, obtuse or
acutish; corolla thinly strigillose, the rather stout tube 5-6 mm. long, the 4 lobes
rounded, 1.5 mm. long, the throat naked; anthers and style included.
Rondeletia rubens L. Wms. Phytologia 26: 128. 1973.
Open montane forests at about 1,800 m.; Quiche (type, Skutch
1725). Endemic.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 189
Slender shrubs to about 4 m. tall; branches slender, tomentose but very soon
glabrous; stipules narrowly lanceolate, acute, subauriculate at the base, about 7-8
mm. long; leaves opposite, subequal to prominently anisophyllous, glabrous, elliptic
or elliptic-lanceolate, acuminate, with 8-10 pairs of secondary nerves, petiole to 2 cm.
long but mostly less than 1 cm. long; inflorescence a terminal circinnate-paniculate
spike, many-flowered, up to 20 cm. long and 2-4 cm. broad, the ultimate divisions and
hypanthium sparsely tomentose; hypanthium about 1 mm. long; calyx 4-lobate, the
lobes lance-oblong, acute, glabrous within, sparsely tomentose outside, slightly
unequal, 0.5-0.7 mm. long; corolla red, the tube slender, nearly glabrous outside, 7-8
mm. long, the lobes oblong-ovate, spreading, about 1.5 mm. long; style about 5 mm.
long, bifid for 1 mm. and stigmatic; anthers inserted at the throat and slightly
exserted, narrowly oblong, about 1.5 mm. long, filament very short.
Related to R. gracilis and R. silvicola. Known only from the
montane forests near Nebaj.
Rondeletia rufescens Robinson, Proc. Am. Acad. 45: 402.
1910. R. villosa f. strigosissima Donn.-Sm. Enum. PL Guat. 2: 30.
1891, nomen nudum. R. rufescens var. ovata Robinson, I.e. 403
(type from Tactic, Alta Verapaz, Tuerckheim 8401).
Wet mixed forest or in thickets, 1,400-2,000 m.; Chiquimula
(Cerro Brujo); Baja Verapaz (below Fatal); Alta Verapaz (frequent
and rather widely distributed). El Salvador; Honduras.
A shrub or small tree, 3 m. tall or more, the branches subterete, densely
ferruginous-villous when young, tardily glabrate; stipules triangular, 5-12 mm. long,
abruptly long-acuminate or cuspidate, erect; leaves opposite, on stout petioles 7-22
mm. long, elliptic-oblong to elliptic or ovate-oblong, 8-15 cm. long, 3-6 cm. broad,
acute to attenuate, at the base rounded to acute, green and minutely pilose above,
beneath densely pilose with short fulvous hairs; inflorescence terminal, short-
pedunculate, the flowers sessile or subsessile in dense cymes, these sessile or
pedunculate, arranged in a narrow thyrsiform panicle 12-20 cm. long; bracts linear-
subulate; hypanthium densely ferruginous-pilose, the 4 calyx lobes lance-oblong, 1
mm. long, acute, spreading; corolla pale pink or rather dull red, the slender tube 8-10
mm. long, short-strigose below, long-strigose above, naked in the throat, the 4 lobes
orbicular, 2 mm. long; style exserted; capsule subglobose, hirsutulous, about 3 mm.
long.
A rather pretty shrub, plentiful at some localities in the
vicinity of Tactic.
Rondeletia secundiflora Robinson, Proc. Am. Acad. 45: 403.
1910. R. vulcanicola Standl. & Steyerm. Field Mus. Bot. 22: 291.
1940 (type from Finca Pirineos, southern slopes of Volcari de Santa
Maria, 1,300-1,500 m., Steyermark 33220).
Moist or wet mixed, mountains forest, 600-1,500 m.; endemic;
Escuintla (type collected along the road between Palfn and
190 FIELDIANA: BOTANY, VOLUME 24
Escuintla, Sutton Hayes in 1860); Guatemala (?); Sacatep^quez;
Chimaltenango; Solola; Quezaltenango.
A slender shrub or small tree 1.5-6 m. high, the branches subterete, whitish-
strigose when young; stipules erect, subulate from a short triangular base, 2.5 mm.
long; leaves membranaceous, on petioles 5-15 mm. long, ovate- lanceolate to elliptic-
lanceolate, long-attenuate, 5-15 cm. long and 1.5-5 cm. broad, narrowed to the acute
or acuminate base, appressed-pilose above, beneath sparsely or rather densely
appressed-pilose, barbate in the nerve axils; inflorescence terminal, narrowly
paniculate and spikelike, 8 cm. long or less, the cymes sessile or short-pedunculate;
bracts small and inconspicuous; hypanthium 1 mm. long, densely sordid-hirtellous,
the 4 calyx lobes subequal or one lobe larger, linear or lanceolate, 1-1.5 mm. long,
hispidulous; corolla dark purplish, strigose, the slender tube 6-8 mm. long, the
suborbicular lobes 1.5 mm. long, glabrous within, the throat not barbate; capsule
didymous-globose, about 4 mm. broad and 3 mm. high, hispidulous; seeds yellowish,
obtusely angulate, coarsely punctate.
Rondeletia seleriana Loes. Verh. Bot. Ver. Brandenb. 65: 105.
1923.
In mountain forest or thickets, 2,500-2,800 m., or probably also
at lower elevations; endemic; Huehuetenango (type collected near
Chacula, Distr. Nenton, Seler 3015; a photograph of the type in
Herb. Field Mus.); also on Cerro Canana.
A shrub or small tree, 6 m. high or less, the branchlets terete, when young
sparsely strigillose; stipules subulate-deltoid, almost 1 cm. long, erect; leaves opposite,
on petioles 4-11 mm. long, coriaceous, oblong-ovate to ovate-elliptic, 6-15 cm. long, 2-
6 cm. broad, acuminate, at the base acute to almost rounded, glabrous above,
somewhat strigillose beneath along the costa, the lateral nerves 5-7 on each side;
inflorescence terminal, cymose-corymbose, the branches strigillose, the bracts minute,
the flowers sessile or subsessile; hypanthium strigillose, the calyx lobes minute;
corolla pink, strigillose and more or less fulvous-villosulous, the tube 10-12 mm. long,
the rounded lobes 2 mm. long, the throat and inner face of the tube densely yellow-
barbate; stamens included; style exserted.
Rondeletia septicidalis Robinson, Proc. Am. Acad. 45: 403.
1910.
San Marcos (volcanoes of Tajumulco and Tacana, 1,400-2,100
m., in wet mixed forest). Mexico (Chiapas), the type from
Chicharras.
A shrub or small tree 4-6 m. high, the branches slender, terete, ferruginous, thinly
arachnoid-tomentose when young but soon glabrate; stipules triangular-lanceolate, 5
mm. long, erect; leaves opposite, subcoriaceous, anisophyllous, on petioles 5-35 mm.
long, broadly ovate to lance-eliiptic or lanceolate, 5-19 cm. long, 1.5-7 cm. broad,
abruptly acuminate or long-attenuate, at the base acute or abruptly acuminate,
green and glabrous above, when young thinly and loosely arachnoid-tomentose
beneath but soon glabrous, the lateral nerves 5-9 on each side; inflorescence terminal,
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 191
slender-pedunculate, usually reflexed, the flowers subsessile in 2-3-flowered cymules,
these remote and forming a spikelike panicle or a spike 6-30 cm. long; bracts linear,
elongate; hypanthium densely floccose-tomentose, the 4 calyx lobes lance-linear,
unequal, equaling or longer than the hypanthium, deflexed; corolla red, the slender
tube 6-8 mm. long, glabrous below, floccose-tomentulose above, glabrous in the
throat, the 4 lobes suborbicular, 2-3 mm. long; stamens and style included; capsule
subglobose, 4 mm. in diameter.
Rondeletia silvicola L. Wms. Phytologia 26: 128. 1973.
Openings in wet lowland forests or in the forest near sea level,
Izabal. Honduras (type, Standley 56885).
Slender shrubs or small trees to 8 m. tall; branches slender, terete, floccose but
soon glabrescent; stipules narrowly triangular, 2.5-4 mm. long; leaves elliptic to
oblanceolate-elliptic or oblong-elliptic, long acuminate, somewhat anisophyllous,
lower surface obscurely arachnoid-floccose, soon glabrate, the blades 10-25 cm. long
and 5-10 cm. broad, the petioles of a pair unequal, the longer ones to about 3 cm.
long, obscurely floccose; the inflorescences terminal, spicate with numerous densely
cymose clusters of flowers, 15-30 cm. long and 1-2 cm. in diameter; flowers pink to
deep red; hypanthium floccose, to about 1 mm. long; calyx 4-lobate, the lobes
narrowly triangular, reflexed at anthesis, glabrous within, about 0.5-0.7 mm. long;
corolla red, arachnoid-floccose outside, the tube slender, 7-8 mm. long, the lobes
spreading, suborbicular, 2-2.5 mm. long; style 4-5 mm. long, bifid; stamens inserted in
the throat, the anthers included, linear-oblong, very small, about 0.8 mm. long;
mature capsules subglobose, about 1.5 mm. long.
An occasional shrub or small tree in the forest or in forest
openings along the north coast of Honduras and in the adjacent
Guatemalan lowlands.
Rondeletia skutchii Standl. & Steyerm. Field Mus. Bot. 22:
289. 1940.
Moist or wet, mixed, mountain forest, 1,200-2,000 m.; endemic;
Suchitepequez (type collected on forested ridge, Finca Moca, Skutch
2110)-, Solola.
A slender shrub 4.5 m. high, the branches terete, the younger ones ferruginous, at
first densely but laxly tomentose, soon glabrate; stipules subulate-attenuate from a
triangular-lanceolate base, 7 mm. long, erect; leaves membranaceous, opposite, on
petioles 1-1.5 cm. long, narrowly lance-oblong to oblong-elliptic, 10-15 cm. long, 3-6.5
cm. broad, acuminate or narrowly attenuate-acuminate, acute at the base, green
above, at first sparsely and laxly tomentose, soon glabrous, rather densely and very
laxly tomentose beneath, the lateral nerves about 11 on each side; inflorescence
reflexed and pendulous, narrowly thyrsoid and spikelike, rather laxly many-flowered,
short-pedunculate, 7-14 cm. long, about 3.5 cm. broad, the rachis densely tomentose,
the flowers in dense, few-flowered, sessile or short-pedunculate cymules, the bracts
linear, inconspicuous; hypanthium subglobose, densely white-tomentose, the 4 calyx
lobes narrowly lance-triangular, long-attentuate, 2 mm. long; corolla floccose-
192 FIELDIANA: BOTANY, VOLUME 24
tomentose, the slender tube 10-12 mm. long, the 4 lobes rounded, glabrous within, the
throat glabrous; stamens included.
Rondeletia stachyoidea Donn.-Sm. Bot. Gaz. 43: 298. 1906.
Damp or wet forest, 900 m. or lower; Peten; Alta Verapaz (type
from Semacoch, Robert Hay); Izabal; Huehuetenango (Ixcan).
British Honduras; Honduras.
A slender shrub or small tree 4.5 m. tall or less, rarely larger, the branchlets
terete, pilose-strigose when young; stipules triangular-subulate, erect, 6-10 mm. long;
leaves opposite, on petioles 3-18 mm. long, ovate-lanceolate to narrowly elliptic-
lanceolate, 6-17 cm. long, 1.5-4 cm. broad, long-attenuate, at the base attenuate or
abruptly acuminate, bright green and lustrous above, glabrous, or strigose on the
veins, densely sericeous beneath with long lustrous hairs or in age glabrate;
inflorescence terminal, short-pedunculate, often recurved, the flowers fragrant, sessile
or subsessile in small cymes, these spicate, the spikes 7-12 cm. long or longer, the
bracts linear-subulate; hypanthium sericeous, the 4 calyx lobes lance-oblong, 3-4 mm.
long, attenuate, spreading; corolla white, the slender tube 7-8 mm. long, glabrous
execpt at the top, there densely setose-pilose, naked in the throat, the 4 lobes
rounded, 1.5 mm. long; anthers and style included; capsule subglobose, 2.5 mm. long,
costate, glabrate.
Rondeletia stenosiphon Hemsl. Diag. PL Mex. 26. 1879. R.
lundelliana Standl. in Lundell, Contr. Univ. Mich. Herb. 4: 31. 1940
(type from El Cayo District, Vaca, British Honduras, Gentle 2504).
Quinillo (fide Aguilar).
Wet thickets or forest, 1,100-1,750 m.; Pet<§n; Alta Verapaz
(widely distributed and rather frequent); Baja Verapaz (Penzal);
Quich£; Huehuetenango. Mexico (Chiapas); British Honduras.
A shrub or small tree 1-3 m. high or larger, sometimes a tree of 8 m., the stout
branches acutely angulate, strigillose when young; stipules lance-triangular, 5-12 mm.
long, erect; leaves opposite, subcoriaceous, on stout petioles 4-13 mm. long, obovate
to oval-oblong or elliptic-oval, 7-15 cm. long, 4-7.5 cm. broad, usually abruptly short-
acuminate, at the base subobtuse to acuminate, dark green and lustrous above,
glabrous, beneath finely strigillose or glabrate; inflorescence terminal and axillary,
cymose-corymbose, on stout peduncles 4-6 cm. long, cymose-corymbose, 8-14 cm.
long, the cymes densely many-flowered, the flowers fragrant, sessile or short-
pedicellate; bracts and bractlets usually minute, triangular; hypanthium densely
sericeous, the 5 calyx lobes minute, ovate-triangular, acute, erect; corolla white,
densely sericeous-strigose, the slender tube 8-11 mm. long, densely yellow-barbate in
the throat, the 5 lobes oval, 2.5-4 mm. long, puberulent within; style included or
exserted.
A showy and rather handsome shrub when in flower because of
the abundant blossoms. A single specimen from near Flores, Peten
(± 200 m.) is the only one known below 1,000 m. (Contreras 5814).
It could be a mixed label.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 193
Rondeletia strigosa (Benth.) Hemsl. Diag. PL Mex. 27. 1879.
Bouvardia strigosa Benth. PL Hartweg. 75. 1841. Jazmin; arete
tinto (Guatemala).
Widely distributed and common in many localities, 1,000-3,000
m., oak, pine, or mixed forest, in thickets, or on cliffs; Alta Verapaz
(Coban); Chiquimula; Jalapa; Jutiapa; El Progreso; Santa Rosa;
Guatemala; Sacatepequez; Chimaltenango; Quiche; Solola;
Suchitep6quez; Quezaltenango (type from Zunil, Hartweg 530); San
Marcos. Mexico (Chiapas) to El Salvador and Honduras.
A shrub, sometimes 2 m. high but usually 1 m. high or less, the rather stout
branches terete, deep green, hirtellous or puberulent when young; stipules very short,
subtruncate; leaves mostly ternate, sessile or nearly so, ovate to ovate-oval, 2-5 cm.
long, acute to acuminate, rounded or subtruncate at the base, strigillose, dark green
above, paler beneath, the lateral nerves few, most of them rising from near the base
of the blade at a very narrow angle; flowers terminal, umbellate, few or numerous,
short-pedicellate; hypanthium densely strigose, the 5 calyx lobes linear or lance-
linear, 6-10 mm. long, acute, ascending; corolla deep or bright red, sometimes dull
purplish red, sparsely strigose outside, the slender tube 2-2.5 cm. long, densely yellow-
pilose in the throat, the 5 lobes obovate-orbicular, 5-10 mm. long, glabrous within;
anthers and style included; capsule globose, about 6 mm. long, strigose; seeds
numerous, rather large, densely punctate, attenuate to each end, dark brown.
A somewhat showy and handsome plant because of its
unusually colored flowers, which are larger than those of other
Guatemalan species. The yellow centers of the corollas contrast
with the deep red lobes. The deep green and rather tough and
flexible branches also are distinctive.
Rondeletia suffrutescens Brandegee, Univ. Calif. Publ. Bot.
6: 70. 1914.
Damp or wet, mixed forest or thickets of the western highlands,
1,300-3,000 m.; Suchitepe"quez; Quezaltenango; San Marcos. Mexico
(Chiapas), the type from Cerro del Boqueron.
A slender weak shrub, suberect or often scandent, 1-2 m. long, the branches
terete, the younger ones green, at first sparsely pilosulous; stipules linear or
triangular-lanceolate, 3-5 mm. long, erect; leaves opposite, membranaceous, on
slender petioles 2-6 mm. long, lanceolate to ovate, 5-13 cm. long, 1.5-5 cm. broad,
long-acuminate or attenuate, rounded or abruptly short-acuminate at the base,
sparsely setose-pilose on both surfaces, to glabrous; inflorescence terminal, cymose,
slender-pedunculate, the cymes mostly 3-flowered, the slender pedicels 5-15 mm. long;
hypanthium glabrous, subglobose, 2 mm. long, the 5 calyx lobes linear-lanceolate, 6-
14 mm. long, attenuate, green, sparsely setose-ciliate; corolla dull red or pale yellow,
sometimes spotted with purplish, glabrous, the tube 15-21 mm. long, the throat
densely yellow-barbate, the 5 lobes oblong or oval, 6-7 mm. long; anthers and style
194 FIELDIANA: BOTANY, VOLUME 24
included; capsule about 1 cm. long, conspicuously costate; seeds large, angulate,
fuscous, coarsely tuberculate.
The shrub is an inconspicuous one, often growing in the
deepest, wettest forest, and apparently is of infrequent occurrence.
Rondeletia tacanensis Lundell, Bull. Torr. Bot. Club 66: 603.
1939.
Wet mixed forest or thickets of the western highlands, 1,250-
3,000 m.; Solola; Quezaltenango; San Marcos. Mexico (Chiapas), the
type from Volcan de Tacana at 2,800 m., Matuda 2928.
A shrub or tree, sometimes 20 m. high and 15 cm. in diameter, but usually much
smaller, irregularly branched, the branches subterete, densely tomentose when young;
stipules triangular, erect, 5-9 mm. long; leaves opposite, membranaceous, on stout
petioles 3 cm. long or less, elliptic, oval, or obovate-elliptic, 11-24 cm. long, 7-13 cm.
broad, acute or abruptly short-acuminate, broadly cuneate at the base or contracted
and short-decurrent, densely pilose above with chiefly subappressed hairs, densely
hirsute beneath, the lateral nerves 12-16 on each side; inflorescence terminal, short-
pedunculate, reflexed and pendent, the flower sessile in very dense cymules, these
sessile or short-pedunculate and arranged in an interrupted spikelike panicle 15-35
cm. long; bracts linear or subulate, green, 5-7 mm. long or larger; hypanthium
densely hirsute with subappressed hairs, the 4 calyx lobes linear or subulate, 4-8 mm.
long, unequal, subfoliaceous, suberect; corolla dark dull red, sparsely pilose with
somewhat appressed hairs, naked in the throat, the 4 lobes oval or rounded, about 5
mm. long, glabrous within; anthers exserted only at the apex, the style included or
exserted.
The shrub is plentiful in many localities of the wet mountain
forests of the western highlands. It is exceptionally conspicuous
because of the many long narrow pendent panicles of rather
brilliantly colored flowers. Heterostyly apparently occurs in this
species.
Rondeletia zolleriana Standl. & Steyerm. Field Mus. Bot. 22:
292. 1940.
Wet mixed mountain forest, 1,800-2,400 m.; so far as known,
endemic, but doubtless extending into Chiapas, Mexico; San Marcos
(type collected on slopes between Finca El Porvenir and Loma
Corona, southwestern slopes of Volcan de Tajumulco, Steyermark
37738).
A slender shrub 1.5-2.5 m. high, rarely a weak tree to 10 m., the branches terete,
sparsely strigillose at first; stipules subulate-acuminate from a broad base, erect, 2.5-3
mm. long; leaves opposite, membranaceous, on slender petioles 5-13 mm. long, elliptic
or elliptic-ovate, 5-10 cm. long, 2.5-5.5 cm. broad, abruptly and narrowly long-
acuminate, acute or obtuse at the base and usually abruptly contracted, sparsely
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 195
villous above or almost glabrous, sparsely hirtellous beneath with subappressed hairs,
often barbate in the nerve axils, the lateral nerves 5-6 on each side; inflorescence
terminal, pedunculate or sessile, very lax, cymose-paniculate, few-flowered, as much
as 5 cm. long, the flowers mostly in 3-flowered cymules, the pedicels 7 mm. long or
less; bracts elongate, foliaceous, linear or linear-lanceolate; hypanthium oval, 2-2.5
mm long, densely whitish-strigose, the 4 calyx lobes very unequal, the largest
lanceolate, 10 mm. long and 3 mm. broad, acuminate, 3-nerved, the other 3 linear or
subulate, 5-7 mm. long; corolla white to ochroleucous, whitish-strigose, the tube 8-15
mm. long, the throat not barbate, the lobes suboricular, almost 4 mm. long, broadly
rounded at the apex, glabrous within.
This species was named for Mr. Erich Zoller of Finca El
Porvenir, who furnished facilities for collecting on the lower slopes
of the Volcan de Tajumulco. The species is locally abundant on the
western slopes of the volcano at middle elevations.
RONDELETIA ROEZLII (Planch.) Hemsl. Biol. Cent. Am. Bot. 2:
23. 1881. Rogiera roezlii Planch. Fl. Serres 5: 442. 1849. This was
based on cultivated specimens believed to have been of Guatemalan
origin. It is not identifiable with certainty from the description but
is probably a synonym of R. amoena.
RUDGEA Salisbury
Shrubs or small trees, glabrous or pubescent; stipules interpetiolar, bearing on
the margins or at the apex or on the dorsal surface subulate or aculeiform teeth,
these usually pale and somewhat cartilaginous, generally deciduous, the stipules
sometimes laciniate; leaves opposite, short-petiolate or sessile, generally somewhat
coriaceous; inflorescence terminal, usually paniculate or cymose, sometimes
umbellate or capitate, rarely reduced to a single flower; calyx lobes almost free or
variously connate; corolla salverform or funnelform, commonly white, the lobes
valvate in bud; stamens inserted in the corolla tube, with short or elongate filaments;
anthers dorsifixed, linear, included or exserted; ovary 2-celled, the style filiform or
columnar, the 2 lobes orbicular or oblong; ovules solitary, erect from the base of the
cell; fruit almost dry or juicy, bipyrenate, the nutlets plano-convex, smooth or
sulcate dorsally, longitudinally sulcate on the inner face.
A large group of perhaps 75 species in tropical America. Five
other Central American species have been described from Costa
Rica and Panama. The genus Rudgea, like others in the tribe
Psychotrieae, is difficult to distinguish from Psychotria and is a
genus of "convenience."
Corolla about 1 cm. long; flowers in part conspicuously pedicellate; upper leaves
usually narrowed to a narrowly obtuse base R. cornifolia.
Corolla almost 2 cm. long; flowers all sessile or nearly so; leaves narrowed to an
acute or acuminate base.... ....R. simiarum.
196 FIELDIANA: BOTANY, VOLUME 24
Rudgea cornifolia (Humb. & Bonpl.) Standl. Field Mus. Bot.
7: 432. 1931. Psychotria cornifolia Humb. & Bonpl. ex R. & S. Syst.
Veg. 5: 191. 1819. P. fimbriata Benth. in Hook. Journ. Bot. 3: 226.
1841. R. micrantha Muell. Arg. Flora 59: 454. 1876. R. fimbriata
Standl. in Standl. & Cald. Lista PL Salv. 274. 1925. R. ceratopetala
Donn.-Sm. Bot. Gaz. 35: 3. 1903 (type from Cubilgiiitz, Alta
Verapaz, Tuerckheim 7904). Canela de montaha and palito copal
(Quezaltenango; probably erroneous names). Figure 50.
Wet forest or thickets, ranging from sea level to about 1,400 m.;
Peten; Izabal; Solola; Huehuetenango; Quezaltenango (southern
slopes of Volcan de Santa Maria). Southern Mexico to Panama,
southward to Bolivia and Brazil.
A glabrous shrub 2-4 m. high; stipules deltoid-ovate, incised-laciniate; leaves
sessile or short-petiolate (especially the lower ones), elliptic to elliptic-oblong or
obovate, 8-15 cm. long, 2-7 cm. broad, gradually or rather abruptly acuminate,
commonly narrowed to an obtuse or narrowly rounded base (especially the upper
leaves); inflorescence cymose-paniculate, usually small, broad and open, many-
flowered, the lateral flowers of the cymules pedicellate, the cyme erect, pedunculate;
calyx short, minutely 5-dentate or subentire; corolla white, commonly 5-7 mm. long,
the lobes equaling or longer than the tube; fruit ellipsoid, white, 5-8 mm. long in the
dry state.
Called "membrillo" and "tapacajete" in Oaxaca.
Rudgea simiarum Standl. & Steyerm. Field Mus. Bot. 22: 389.
1940.
Izabal (type collected between Virginia and Lago de Izabal,
Montana del Mico, 50-500 m., Steyermark 38839).
A glabrous tree 6 m. high; stipules persistent, forming a truncate, obscurely
mucronate sheath 2.5-3 mm. long, this densely hirsute within, the hairs exserted;
leaves on petioles 8-15 mm. long, lance-oblong to elliptic-oblong, usually broadest
near the middle, 7-16 cm. long, 2.5-7 cm. broad, rather abruptly long-acuminate,
acute or long-decurrent at the base, the lateral nerves about 8 on each side;
inflorescence cymose-corymbose, densely few-flowered, 2-2.5 cm. broad (excluding the
flowers), the peduncle 2-3.5 cm. long, erect, the flowers sessile or nearly so;
hypanthium thick-columnar, scarcely 1 mm. high, the calyx scarcely 0.5 mm. high,
remotely and very minutely dentate; corolla white, obtuse at the apex in bud and not
appendaged, the tube 10-11 mm. long, the lobes 8 mm. long, linear-lanceolate,
spreading or recurved; anthers exserted, 3 mm. long.
The junior author believes that this plant, of which only the
type is known, may be a Psychotria.
SABICEA Aublet
Shrubs, woody or perhaps herbaceous vines (ours) or small trees, scandent or
not; stipules intrapetiolar, persistent; inflorescence axillary, the subsessile heads or
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 197
short pedunculate thyrsiform panicles hardly longer than the petioles; flowers small,
whitish; hypanthium globose; the calyx 3-6-lobate, the lobes usually elongate and
narrow, equal or not, persistent; corolla funnelform or salverform, the tube short or
elongate, the throat villous, the limb 4-5-lobate, the lobes short, valvate in bud;
stamens 4-5, inserted in the throat or tube of the corolla, the filaments long or short,
the anthers dorsifixed, linear, obtuse, included; ovary usually 4-5-celled, sometimes
only 2-celled, the style with 2-5 linear, obtuse, lobes; ovules numerous, the placentae
affixed to the axis of the ovary; fruit baccate (when mature), 2-5-celled; seeds
numerous, minute, ovoid or angulate.
A large genus of perhaps more than 100 species in Africa and
tropical America, mostly in South America. There may be other
species in Mexico and southern Central America.
Pubescence of the stems closely appressed; inflorescence when developed rather lax
and short pedunculate S. panamensis.
Pubescence of the stems of spreading hairs; inflorescence dense, sessile or not
obviously pedunculate S. villosa.
Sabicea panamensis Wernham, Monogr. Sabicea 30. 1914. S.
costaricensis Wernham, I.e. 31.
Wet thickets or forests, often in second growth, at or little
above sea level; British Honduras; to be expected in Peten; Izabal;
Baja Verapaz. Southward to Panama.
A small woody vine, climbing over shrubs, the stem ferruginous or purplish,
densely yellowish-sericeous when young; stipules ovate or ovate-oval, 7-10 mm. long
and 4-6 mm. broad, acute or obtuse; the petioles stout, 1-1.5 cm. long; leaf blades
elliptic-lanceolate to ovate-oblong, mostly 7-10 cm. long and 3-5 cm. broad, acute or
abruptly short-acuminate, the base rounded, sparsely strigose or strigillose above,
sericeous-strigose beneath along the veins; elsewhere sparsely strigose or glabrate;
inflorescence rather lax and thyrsiform, several-flowered, little exceeding the petioles,
densely yellowish-strigose, 2-3 cm. broad, the peduncle mostly 1 cm. long or shorter,
the pedicels to 8 mm. long; bracts lanceolate, connate at the base; calyx lobes linear
or oblong, 2-3.5 mm. long, green, reflexed at maturity; corolla whitish, strigose
outside, the tube 6-8 mm. long, the lanceolate lobes about 2.5 mm. long; fruits about
1 cm. long, dark purple or lavender when mature.
The species was described by Wernham as an erect shrub, and
it is often so indicated on lables, but there is little reason for
believing that any Central American member of the genus is erect.
Sabicea villosa Roem. & Schult. Syst. Veg. 5: 265. 1819. S.
hirsuta HBK. Nov. Gen. & Sp. 3: 417. 1820. Figure 25.
Wet thickets, often in second growth, 600 m. or less; Alta
Verapaz; Izabal; Retalhuleu (near city of Retalhuleu). Southern
Mexico and British Honduras along the Atlantic slope to Costa
Rica. Panama south to Brazil and Peru along both oceans.
198 FIELDIANA: BOTANY, VOLUME 24
A slender vine, either somewhat woody or wholly herbaceous, the stems densely
fulvous-hirsute; stipules broadly ovate or rounded, reflexed, 5-10 mm. long, obtuse or
acutish; leaves short petiolate, ovate to ovate-oblong or elliptic-oblong, 5-12 cm. long
and 2-2.5 cm. broad, rather abruptly acuminate, rounded or obtuse at the base and
often short-decurrent, abundantly hirsute; inflorescence sessile in the leaf axils,
flowers few and crowded; calyx and hypanthium densely hirsute, the calyx lobes
linear- lanceolate, 2-4 mm. long, reflexed at maturity; corolla white, hirsute or
strigose, the tube about 6 mm. long, the narrowly triangular lobes 1.5-2 mm. long;
fruit dark purple or dark red, about 1 cm. long, baccate and juicy at maturity; seeds
brown.
A single specimen is known along the Pacific coast in Central
America. The species is found along the coast from Panama
southward. The locality in Retalhuleu is unexpected.
SICKINGIA Willdenow
Glabrous or pubescent trees, with thick branchlets; stipules ovate or lanceolate
to triangular, caducous; leaves large, opposite, petiolate, usually membranaceous;
inflorescence paniculate, the panicles terminal or axillary, the small flowers usually
short-pedicellate, bracteolate or ebracteolate; hypanthium obconic, the calyx short,
shallowly lobate, cupular or campanulate; corolla tubular or funnelform, usually
pilose within at the insertion of the stamens, the limb 4-5-lobate, the lobes short and
broadly rounded, imbricate or open in bud; stamens inserted below the middle of the
corolla tube, exserted, the filaments villous; anthers rather large, versatile, oblong;
ovary 2-celled, the style stout, with 2 short branches; ovules very numerous, the
placentae elongate, longitudinally adnate to the septum; capsule globose, 2-celled,
bivalvate, woody; seeds large, horizontal, lunulate or semi-oblong, broadly winged.
A small genus of some 30 species mostly in South America.
Two or three others are known from Mexico and three more from
Central America and Panama. Sickingia calderoniana Standley
may be expected in eastern Guatemala, and S. rhodoclada Standley
in western Guatemala.
The descriptions, as well as the key, are based on insufficient
and often inadequate material. The senior author had placed most
of the Guatemalan and British Honduran material we have seen in
S. salvadorensis. He may have been right.
Petioles about 1 cm. long; branchlets pilose S. vestita.
Petioles longer than 1 cm., glabrous; branchlets glabrous or nearly so.
Leaves lanceolate to oblong-lanceolate, acute at the base S. lancifolia.
Leaves oblong-obovate, somewhat narrowed to the rounded or emarginate base.
S. salvadorensis.
Sickingia lancifolia Lundell, Wrightia 4: 50. 1968; S.
lancifolia var. puberula Lundell, I.e. 51.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 199
Wet forests or along streams or lakes, Peten (type Lundell
18224 and type of the variety Contreras 3514); Alta Verapaz.
Endemic.
Small trees or shrubs 1.5 m. or taller; branchlets minutely puberulent, terete,
stipules to 2 cm. long; leaves oblong-lanceolate to lanceolate, acuminate, acute to the
base, glabrous, 10-20 cm. long and 1.5-6 cm. broad, petioles mostly 1 cm. long;
inflorescence terminal, small, said to be to 7 cm. long; capsule subglobose, 1.5-2 cm. in
diameter.
This species is known only from a fruiting or sterile specimen.
Sickingia salvadorensis Standl. Field Mus. Bot. 12: 390.
1936. Calderonia salvadorensis Standl. Journ. Wash. Acad. Sci. 13:
290. 1923. Tapalcuite; colay (Alta Verapaz); polo Colorado (Peten,
fide Lundell); chacahuante; chactemuch (Pete"n, Maya); tapalcuit;
tapalcuito; palo de puntero; John Crow redwood (British
Honduras).
Frequent in forests or thickets of the plains or low hills of both
coasts, often on limestone, 250 m. or lower; Petei.; Alta Verapaz;
Santa Rosa; Escuintla; Suchitepequez; Retalhuleu. Mexico
(Yucatan); British Honduras; El Salvador.
Usually a small or medium-sized tree about 10-12 m. high, reported as sometimes
30 m. high with a trunk 30 cm. or more in diameter, the bark smooth, pale, the young
branchlets puberulent; stipules linear-lanceolate, about 2 cm. long, glabrous; petioles
slender, 2-3 cm. long, the blades mostly oblong-obovate, 12-24 cm. long, 5-12 cm.
broad, acute or short-acuminate, somewhat narrowed to the rounded or emarginate
base, puberulent beneath along the nerves, barbate in the nerve axils and domatiate,
the lateral nerves 10-12 on each side; panicles 8-15 cm. long, dense and many-
flowered, the rachis minutely puberulent, the flowers mostly sessile; hypanthium 3
mm. long, glabrous; calyx 1.5 mm. long, the lobes rounded, minutely ciliolate; corolla
whitish or yellowish, 5 mm. long; filaments about equaling the corolla lobes, the
anthers 2.5 mm. long; capsule subglobose, about 2 cm. in diameter, with numerous
large pale lenticels; seeds, including the wing, about 15 mm. long and 6 mm. broad.
Known in El Salvador by the names "campeche," "brasil,"
"palo Colorado," "drago," and "sangre de chucho." When cut, the
wood turns red, but this color fades with age and upon exposure to
bright light. It is yellowish when first cut, hard, heavy, strong, fine-
textured, easy to work, finishes smoothly, and is not very durable.
It is little used locally, but is reported as suitable for articles of
turnery and carving. In British Honduras the wood is used for
dyeing hammocks and other articles red. The tree is frequent about
Retalhuleu, as high up as San Felipe, and extends far down the
plains. Often it is seen growing in the fincas. About Chiquimulilla
and Guazacapdn also the tree is common, growing everywhere
200 FIELDIANA: BOTANY, VOLUME 24
through the towns, apparently planted. The wood is used for fuel at
Mazatenango, but is said not to be very good for this purpose. It is
easily recognized wherever found freshly cut because of the red
coloring.
Sickingia vestita Lundell, Wrightia 4: 51. 1940. S. mollis
Lundell, Contr. U. Mich. Herb. 4: 32. 1940, not Standl. Chuc
chenuch (British Honduras, Maya).
Wet lowland forest, near sea level; Peten. Mexico; British
Honduras.
Trees to 15 m. and 0.3 m. in diameter, the branchlets terete, pilose-tomentose but
soon glabrate, stipules lanceolate, to 1.7 cm. long, cilia te; leaves ovate-oblong, obtuse
to subacuminate, subcordate at the base, about 10 cm. long and 5.5 cm. broad,
glabrous above or soft pilose, petioles thick, about 1 cm. long, pilose-tomentose;
capsule subglobose, 2-3 cm. in diameter.
This, like other Sickingias in Guatemala, is inadequately
known.
SOMMERA Schlechtendal
Reference: Louis O. Williams, Sommera in North America,
Phytologia 26: 121-126. 1973.
Shrubs or small trees, more or less pubescent, with terete branchlets; stipules
interpetiolar, large, caducous; leaves opposite, large, membranaceous, conspicuously
lineolate between the veins with numerous fine parallel striations; flowers small,
whitish, in pedunculate axillary cymes, corymbs, or racemes, bracteate and
bracteolate; hypanthium turbinate; calyx 5-lobate, almost bilabiate, the lobes
subfoliaceous, equal or unequal, persistent; corolla funnelform or subcampanulate,
fleshy with the center line of lobes thickened, sericeous outside, the throat villous, the
limb 5-lobate, the lobes short or elongate, valvate in bud; stamens 5, inserted in the
corolla throat, the filaments short, barbate; anthers dorsifixed, oblong, obtuse at each
end; ovary 2-celled, the style short, with linear obtuse branches, glabrous or hirsute;
ovules numerous, the placentae adnate to the septum; fruit baccate, globose or ovoid,
2-celled; seeds numerous, minute, obtusely angulate, the testa thin, foveolate.
About 15 species in tropical America. Four others are known
from southern Central America and Panama and five additional
ones from Mexico.
Leaves sparsely pilose on the upper surface; branchlets pilose S. guatemalensis.
Leaves quite glabrous on the upper surface; branchlets glabrous or nearly so.
S. chiapensis.
Sommera chiapensis Brandegee, Univ. Calif. Publ. Bot. 6:
196. 1915.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 201
San Marcos, southern slopes of Volcan de Tajumulco, 1,300-
1,500 m., in forested ravines. Mexico (Chiapas), the type from Finca
Irlanda.
A tree of 9-12 m. with brownish branchlets, the young branchlets glabrous;
stipules linear-lanceolate, 1.5-3.5 cm. long, brown, attenuate, glabrous; petioles
slender, 2-5 cm. long, the blades obovate or obovate-oblong, 12-25 cm. long, 5-10 cm.
broad, abruptly acuminate, abruptly acuminate or attenuate at the base, bright green
above and glabrous, paler beneath, densely sericeous when young but soon glabrate;
inflorescence racemose or cymose-racemose, few-flowered, on a long or short
peduncle, the pedicels 2-3 mm. long or more; bracts small, oblong, obtuse; calyx lobes
semiorbicular or ovate-orbicular, ciliolate; corolla white, 7-8 mm. long, very densely
sericeous outside; fruit globose or oval, about 1 cm. long, glabrous.
Sommera guatemalensis Standl. Contr. U. S. Nat. Herb. 17:
436. 1914. Figure 26.
Usually in dense wet forest at 350-1,600 m.; Alta Verapaz (type
collected near Cubilgiiitz, Tuerckheim 8225); Huehuetenango;
endemic.
A shrub or small tree, sometimes 8 m. high, with reddish brown branches, these
sparsely lenticellate, appressed-pilose when young but soon glabrate; stipules
narrowly lanceolate, 3.5-4.5 cm. long; petioles 2-5 cm. long, the blades oval to oblong-
obovate, 17-30 cm. long, 8-14 cm. broad, rather abruptly acuminate, acute or obtuse
at the base, sericeous above when young but soon glabrate, appressed-pilose beneath
along the veins; inflorescence cymose-corymbose, many-flowered, the peduncles 2-5
cm. long, the flowers sessile or short-pedicellate; bracts broadly ovate or oblong,
obtuse, or abruptly short-acuminate; hypanthium 3-4 mm. long, densely sericeous;
calyx lobes 5, oval or broadly ovate, unequal, 3-6 mm. long, usually acute or
acuminate, sometimes obtuse, sparsely short-pilose; corolla pinkish white or cream-
colored, densely sericeous outside, the tube 6-8 mm. long, the lobes one-fifth to one-
third as long as the tube, triangular-ovate, acute; fruit globose-oval, 12 mm. long,
sparsely sericeous.
The oldest name in this genus is Sommera grandis (Bartling)
Standl., which was collected by Haenke in Mexico, and to it have
been referred most of the North American collections at one time or
another.
SPERMACOCE Linnaeus
Annual herbs, usually erect or suberect, glabrous or pubescent, the stems
tetragonous; stipules united with the petioles to form a multisetose sheath; leaves
opposite, petiolate or sessile, herbaceous, narrow, often blackening when dried;
flowers very small, white or pinkish, in dense axillary and terminal heads, the
terminal heads subtended by an involucre of 2 or more leaflike bracts; hypanthium
obovoid, the calyx 4-dentate, the lobes usually green; corolla funnelform, the limb 4-
lobate; stamens 4, inserted in the corolla tube; ovary 2-celled, the style slender, the
stigma capitate or shallowly bilobate; fruit dicoccous, coriaceous, one of the cocci
202 FIELDIANA: BOTANY, VOLUME 24
dehiscent ventrally, the other usually indehiscent; seeds 1 in each cell, oblong, convex
dorsally.
Perhaps a dozen species, natives of the warmer parts of
America, with some of them perhaps introduced into other parts of
the tropical world.
Plants glabrous throughout; fruit glabrous S. riparia.
Plants more or less pubescent, often densely so; fruit pubescent.
Stems densely hispid with long spreading hairs; leaves conspicuously hispid,
especially beneath S. tetraquetra.
Stems almost glabrous but usually scabrous on the angles; leaves merely scabrous
or almost glabrous S. confusa.
Spermacoce confusa Rendle, Journ. Bot. 74: 12. 1936; L.
Wms. Phytologia 26: 493. 1973. Hierba de pajaro (fide Aguilar).
Common in the east of Guatemala at low elevations, also in
Guatemala and Huehuetenango, ascending to 1,800 m.; a weedy
plant, growing in waste ground, cornfields, sand spits, or thickets;
Zacapa; Chiquimula; Jalapa; Jutiapa; Santa Rosa; Guatemala;
Huehuetenango. Generally distibuted in tropical America.
Plants slender, erect or spreading, 60 cm. high or less, often much branched, the
stems 4-angulate, usually scabrous, at least on the angles; leaves ovate-lanceolate to
linear-lanceolate, 2-7 cm. long, acuminate, scaberulous on one or both surfaces;
flower heads containing few or many flowers, about 7 mm. in diameter; sepals ovate-
triangular, very small; corolla white or purplish, glabrous, scarcely 2 mm. long; fruit
subglobose, 2 mm. in diameter, minutely hispidulous with whitish hairs.
Called "hierba de Santa Clara" in El Salvador; "taulmil"
(Yucatan, Maya).
We have called attention to the fact that the name Spermacoce
confusa is not acceptable since it has no basis. The problem of
finding or making an acceptable name for our plants would require
a revision of the group and more time than we have available to us.
The problem is left for a monographer. The plant here described,
and similar species, have often gone under the name of S. tenuior L.,
the type species of the genus Spermacoce. Dr. Rendle's name is "a
new name for S. tenuior of authors, which is to say that it may be
and probably is a "conglomerate."
Spermacoce riparia Cham. & Schlecht. Linnaea 3: 355. 1828.
Figure 65.
Of scattered occurrence and not common, usually in damp
thickets, sometimes in ditches, often in open fields, 1,500 m. or
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 203
lower; Pete'n; Alta Verapaz; El Progreso; Zacapa; Escuintla;
Suchitepe'quez. Mexico; British Honduras to Panama and through
much of South America; West Indies.
A glabrous annual, erect or ascending, 20-50 cm. high, often much branched;
leaves oblong to oblong-lanceolate or narrowly elliptic, 3-8 cm. long, acuminate,
narrowed to the base, short-petiolate, usually blackening when dried; flower heads
about 7 mm. in diameter, usually many-flowered; calyx lobes ovate or deltoid, acute;
corolla white, little exceeding the calyx lobes; capsule glabrous, 1.5-2 mm. long.
This plant usually has been referred to S. glabra Michx., a
species of southeastern United States, which differs in having much
larger flower heads.
Spermacoce tetraquetra A. Rich, in Sagra, Hist. Cub. 11: 29.
1850.
Northern British Honduras, and doubtless extending into
Peten. Mexico (Veracruz and Yucatan); Cuba and Bahamas.
Usually erect and 30-60 cm. high, hirsute or hispid throughout with harsh
spreading hairs, the stout stems acutely angulate; stipule sheath with numerous long
setae; leaves lanceolate to oblong-lanceolate, 2-8 cm. long, 2.5 cm. broad or less,
acute, narrowed to a short stout petiole, conspicuously nerved; flower heads small
and mostly few-flowered, inconspicuous; calyx lobes lanceolate, acuminate; corolla
white, twice as long as the calyx lobes, glabrous outside, densely barbate in the
throat; fruit 2 mm. long, densely hispidulous with whitish hairs.
The Maya name is reported from Yucatan as "poc-xum." The
plant is used medicinally by the Yucatecans.
STEYERMARKIA Standley
Perennial herbs, acaulescent or nearly so, densely long-hirsute; leaves opposite,
crowded at the apex of the short stem, large, membranaceous, short-petiolate;
stipules linear-lanceolate, persistent; flowers very large, 4-parted, pink, cymose, the
cymes dense, many-flowered, head-like, sometimes subracemose, long-pedunculate,
axillary, the bracts large, linear, foliaceous; hypanthium semiglobose or campanulate,
covered with very long hairs; calyx lobes 4, almost free, linear, erect, persistent;
corolla salverform, densely long-pilose outside, the tube elongate, slender, scarcely
dilated in the throat, not barbate within, the limb less than half as long as the tube,
4-lobate, the lobes subequal, broadly oval, rounded at the apex, pilose within with 2-
3-celled hairs, contorted in bud; stamens inserted in the throat of the corolla tube,
included, the filaments short; anthers dorsifixed, linear, obtuse; disk annular; ovary
2-celled, the style filform, the 2 branches short, oblong-linear, subrecurved, included;
ovules numerous, the placentae affixed to the septum, anatropous; capsule
subglobose, rather large, coriaceous, 2-celled, loculicidally 2-valvate; seeds numerous,
minute, globose, not angulate, the testa slightly reticulate.
The genus consists of a single species.
204 FIELDIANA: BOTANY, VOLUME 24
Steyermarkia guatemalensis Standl. Field Mus. Bot. 22: 216.
1940. Figure 10.
Known only from the type, Izabal, Rfo Dulce, 2-4 miles west of
Livingston, on the south side of the river, at sea level, Steyermark
39520; in flower April 16.
Stems 8 cm. long or shorter, thick, with very short nodes, densely leafy near the
apex; stipules linear-lanceolate, as much as 2 cm. long, long-attenuate, densely white-
hirsute outside, glabrous within; leaves large, membranaceous, the petioles sometimes
4 cm. long but usually much shorter, long-hirsute; leaf blades oblanceolate-oblong,
16-28 cm. long, 5-10 cm. broad, acute or acuminate, narrowly attenuate to the base,
green above and densely long-hirsute, the hairs as much as 6 mm. long, paler and
more densely long-hirsute beneath, the lateral nerves about 16 pairs; peduncles
axillary, 9-12.5 cm. long, naked, long-hirsute, the inflorescence dense, congested, head-
like, about 5 cm. broad (excluding the corollas), or the inflorescence composed of 3
head-like cymes; lowest bracts green, as much as 2 cm. long, narrowly lanceolate,
long-attenuate; flowers sessile or short-pedicellate; hypanthium short, densely
glandular-pilose; calyx lobes narrowly linear, erect, 10-14 mm. long, hirsute, with a
black digitiform gland on each side at the base; corolla densely hirsute with long
spreading white hairs, the slender tube 3.5-4 cm. long, 2 mm. broad, the lobes
spreading, about 2 cm. long and 1 cm. broad, orchid-pink, the tube glabrous within;
anthers 5-6 mm. long, the filaments 2 mm. long, glabrous; capsule 7-8 mm. long,
densely long-hirsute, rounded at the base; seeds 0.4 mm. in diameter, dark brown.
This genus is related to Sipanea, a chiefly South American
group of plants with small flowers, 5-parted corollas, and elongate
leafy stems. S. guatemalensis is a showy and conspicuous plant,
apparently of very restricted distribution. The habitat may have
been destroyed where the plant was collected more than 30 years
ago.
UNCARIA Schreber
Scandent shrubs, glabrous or pubescent, climbing by stout uncinate spines, these
formed from abortive peduncles; stipules interpetiolar, entire or bifid; leaves opposite,
pe tie late; inflorescences dense spherical pedunculate heads, sessile or pedicellate,
axillary and solitary, or paniculate; flowers yellowish, usually pubescent;
hypanthium elongate, fusiform, the calyx campanulate, funnel-form, or short-
tubular, 5-lobate or 5-parted; corolla small, tubular-funnelform, with elongate tube,
glabrous in the throat, the limb 5-lobate, the lobes valvate in bud; stamens 5, inserted
in the corolla throat, with short filaments, the anthers oblong, dorsifixed, bisetose at
the base; ovary fusiform, 2-celled, the style slender, long-exserted, the stigma
capitate; ovules numerous, ascending, the placentae affixed to the septum; capsule
small, elongate, 2-celled, septicidally bivalvate, the valves biparted; seeds numerous,
imbricate, the testa winged at each end.
About 35 species, chiefly in tropical Asia and Africa, with one
additional one in South America.
STANDLEY AND WILLIAMS: FLORA OF GUATEMALA 205
Uncaria tomentosa (Willd.) DC. Prodr. 4: 349. 1830. Nauclea
aculeata HBK. Nov. Gen. & Sp. PL 3: 382. 1819, not N. aculeata.
Willd., 1797. N. tomentosa Willd. ex R. & S. Syst. Veg. 5: 221. 1819.
Ourouparia tomentosa Schum. in Mart. Fl. Bras. 6, pt. 6: 132. 1889.
Figure 20.
Wooded swamps, Izabal, near sea level; probably also in Peten.
British Honduras to Panama, southward to eastern Peru.
A shrub or small tree, scandent or at least with long, pendent or clambering
branches, the branches tetragonous, densely or sparsely pilose or puberulent, in age
glabrate; stipules broadly oblong-ovate to deltoid-rounded, 6-11 mm. long, obtuse or
rounded at the apex, often reflexed; leaves on stout petioles 7-15 mm. long, oval or
ovate-oval, 10-15 cm. long, 5.5-8.5 cm. broad, very shortly obtuse-acuminate, at the
base broadly rounded or cordate, bright green and glabrous above, paler beneath,
whitish-tomentulose, or in age glabrate except along the pilose veins; flower heads
spherical, often in panicles, numerous, about 6 mm. in diameter (exclusive of the
corolla), the naked peduncles 3.5 cm. long or less, often compressed, the lowest ones
usually sterile and spinose, large, flat, recurved; flowers sessile; calyx and
hypanthium about 2 mm. long, the hypanthium sericeous, the calyx obscurely 5-
dentate, puberulent, the teeth rounded; corolla 6 mm. long, densely retrorse-pilose
outside with appressed yellowish hairs, the lobes oval or oblong, rounded at the apex;
fruit trigonous, 2-3 mm. long.
It is said that this plant is a water vine, i. e., the sap from
sections of the stem may be used as a substitute for drinking water.
In Honduras the vine is called "una de guara." In some parts of the
Atlantic coast of Central America it is reported to become a
troublesome weed in banana plantations.
FIG. 1. Portlandia quatemalensis. A, branch in flower, X 1 2; B, corolla, stamens,
and style, x 1 V4.
207
FIG. 2. Pogonopus speciosus. A, flowering branch, X Vr, B, flower, x 1 l/z; C and
D, flower dissected to show detail, X 2 V£; E, base of filament, much enlarged; F, an-
ther and apex of filament, X 5.
208
OA/TO
FIG. 3. Oldenlandia corymbosa. A, plant, x %; B, branch, x 2; C, flowers, x 5;
D, flower dissected, x 7 ^; E, fruits, X 5.
209
FIG. 4. Houstonia serpyllacea. A, habit and branch, X 1; B, fruiting branch, X
5; C, flower, X 4; D, flower dissected showing calyx, style, and corolla, and anthers,
X 5; E, anther, X 7; F, section of capsule, X 5; G, seed, X 20, with detail of surface.
210
FlG. 5. Rondeletia buddleioides. A, flowering branch, X lh; B, tomentum from
under leaf surface, much enlarged; C, outline of leaf from another specimen, X W, D,
segment of inflorescence, X 2 V6; E, flower at anthesis and at left apices of other
flowers in process of opening, X 5; F and G, flower dissected to show details, X 5; H,
portion of fruiting inflorescence, X 4; I, seed, X 20.
211
FIG. 6. Lindenia rivatis. A, flowering branch, x 1; B, fruiting branch, X W, C,
ovary, calyx, and style (12 cm. of style left out), x 1; D, capsule opened and
partially dissected to show seeds, X 1; E, seeds, X 5; F, circumaxile (and
intrapetiolar) stipule, X 2 Vz.
212
FIG. 7. Deppea flava. A, flowering branch, X 3A; B, cymose inflorescence showing
mature flower and buds, x 4; C, partial dissection of flower, x 4 V£; D, portion of
fruiting branch with dehisced capsules above, X 1 Vfe; E, mature capsule, x 2 V4; F,
seed, X 15. Detail of leaf shows lineolate tertiary veins on lower leaf surface.
213
FIG. 8. Deppea grandiflora. A, flowering branch, X Vfc; B, flower, X 3; C, corolla
dissected, X 2; D, hypanthium, calyx, and style, X 2; E, stigma, x 5.
214
FIG. 9. PinarophyUon flavum. A, plant, X W; B, pubescence of leaf, much
enlarged; C, portion of cymose inflorescence showing flowers at various stages, X 6;
D, corolla dissected, X 7.
215
FIG. 10. Steyermarkia guatemalensis. A, habit of plant in flower, x Vfc; B,
mature hypanthium and calyx, x 6; C, corolla partially dissected to show detail, X 1
Vz; D, outer surface of limb of corolla, X 1 Vr, E, young flower bud, X 1 Vz; F, seed,
X 20; G, cross-section of ovary, X 3. Sketch of Dr. Julian A. Steyermark for whom
the genus is named.
216
FIG. 11. Bouvardia longiflora. A, branch with flowers and a fruit, x V£; B, fruits,
X 1 W, C, flower, x 1; D, and E, flower dissected, x 1 %.
217
FIG. 12. Manettia reclinata. A, habit, X V4; B, bud and opened flower, x 1 Vfe; C,
flower dissected showing hypanthium, calyx, and corolla, x 2 Vr, D, capsule, x 6; E,
capsule dissected, X 2 Vr, F, seed, X 4.
218
FlG. 13. Alseis yucatanensis. A, flowering branch, x 3/i; B, flower, X 5; C,
hypanthium, calyx, and pistil, X 10; D, corolla dissected, x 5; E, details of fruiting
and flowering inflorescences, X 1.
219
FlG. 14. Hillia tetrandra. A, flowering branch, X Vfc; B, branch in bud showing
early caducous stipules, X W, C, flower partially dissected, X 1; D, capsule segments,
x ^; E, seed x 3.
FIG. 15. Cosmibuena matudae. A, flowering branch, X Vz; B, portion of corolla,
X W, C, fruiting branch, x Vr, D, seed, x 20; E, Dr. Eizi Matuda (for whom this
species was named) from a photograph taken in the field in Mexico in 1961 when
Prof. Matuda, one of the best known and most active Mexican botanists, was 65
years old. He is still an active botanist at 80!
221
FIG. 16. CalycophyUum candidissimum. A, flowering branch, X V4; B, flower, X
2 Vfc; C, corolla dissected, X 2 %; D, segment of inflorescence showing bracts,
hypanthium, and style, X 2 %; E, fruiting cyme, X 2 %.
222
FIG. 17. Exostema mexicanum. A, fruiting branch, x lh; B, inflorescence at
an thesis, X Vt\ C, portion of a cyme with one flower partially dissected, X 2 Vfe; D,
ovary with hypanthium and calyx, x 2 V£; E, base of corolla showing filament tube,
X 3.
FIG. 18. Hintonia standleyana. A, branch, x Vz; B, dissected corolla with style
and stamens, X 1 Vz; C, fruiting branch, x Vz; D, half of capsule showing septum, X
1 Vz; E, seeds from one locule, position same as D, X 1 Vz; F, seed, X 3.
224
FIG. 19. Coutarea hexandra. A, flowering branch, X 1; B, fruiting branch, x
C, capsules showing dehiscence, X 1; D, seed, X 1.
225
FIG. 20. Uncaria tomentosa. A, branch, x 'A; B, segment of inflorescence, x 1;
C, flower, X 9; D, flower dissected, x 9; E, fruiting inflorescence, X 1 Vr, F, capsules
showing dehiscence, x 2 W, G, seed, X 6.
226
Kj^Bt'C^ .^
FlG. 21. Cephalanthus occidentalis. A, flowering branch, X 1 *4; B,
inflorescences, one in bud the other at an thesis, x 1 V4; C, flower and another
dissected, X 5.
227
FIG. 22. Isertia haenkeana. A, flowering branch, x Vz; B, fruiting inflorescence,
X V6; C, fruits and seeds, X 2; D, segment of inflorescence in bud, x 3 Vz: E. flower
with corolla, anther with corolla removed, X 2 Vz; F, detail of apex of dissected
corolla, X 5; G, anther to show sculpture on inner face, X 10.
228
FlG. 23. Gonzalagunia thyrsoidea. A, branch, X V6; B, calyx and pistil, X 2; C,
stigmas, x 5; D, corolla dissected, X 2; E, fruit, X 3.
229
FIG. 24. Coccocypselum cordifolium. A, habit, X V2, with detail of pubescence; B,
flower and another dissected, x 2 Vz.
230
/
FIG. 25. Sabicea villosa. A, habit, X '/2; B, flower, X 4; C, flower dissected to
show detail, X 5; D, mature capsule, X 4; E, seeds, X 5.
231
FIG. 26. Sommera guatemalensis. A, branch, X Vr, B, portion of under surface of
leaf enlarged to show pubescence and veinlets; C, bud to show aestivation, calyx
dissected away to show summit of ovary, X 3; D, flower, X 3; E, corolla dissected to
show detail, X 3; F, style, X 3; G, stamen, X 5; H, diagram showing position of
calyx lobes and stamens.
232
FIG. 27. Pentagonia macrophylla. A, leaf and fruits, X ^; B, inflorescence, X
C, flower, x 1 Vz; D and E, flower dissected to show detail, X 1 V£.
233
_>^PV«^
FIG. 28. Randia standleyana. A, flowering branch, X Vz; B, flower to show
aestivation, X 7 Vz; C, flower at anthesis, x 7 Vz; D and E, flower dissected to show
detail, X 7 ¥2.
234
FIG. 29. Posoqueria latifolia. A, portion of flowering branch, X Vi; B, flower
partially dissected to show stamens and length of style, X 1; C, anther front and
side, x 5; D, mature fruit, X V4. V
235
FIG. 30. Genipa caruto. A, flowering branch, X Vz; B, flower, x 2; C, corolla
dissected with style in position, x 1 l/i; D, fruit, X l/2.
236
FIG. 31. Amaiouia corymbosa. A, flowering branch, x Vz; B. flowers and bud. x
2 ^; C, flower dissected, X 2 Vi\ D, fruits, x 1.
237
FIG. 32. Alibertia edulis. A, flowering branch, X Vr, Aa, fruiting branch, X Vfc; B,
hypanthium, calyx, and style, X 2; C, corolla dissected, X 2; D, anther, X 2 Vz.
238
FIG. 33. Bertiera guianensis. A, flowering branch, x tt; B, corolla dissected to
show anthers, x 5; C, hypanthium, calyx, and pistil, x 5; D, portion of fruiting
inflorescence, X 1.
239
FIG. 34. Hamelia barbata. A, flowering branch, X V4; B, bud, X 2 Vi; C and D,
flower dissected to show detail, x 2 V£; E, inner face of anther, x 4 V4; F, nearly
mature fruits, x 1.
240
/ \
FIG. 35. Hoffmannia cauliflora. A, apical half of plant (with leaves removed)
with inflorescences at nodes from which leaves have fallen, x Vfe; B, flower, x 2 V£;
C, corolla dissected, x 2 Vi; D, style, x 2 Vi; E, mature and immature fruits, x 2 V*.
241
FIG. 36. Hoffmannia sessilifolia. A, variation in leaves, x Vz; B, flower and
dissection of flower, x 1 ¥2. From type.
242
FIG. 37. Guettarda macrosperma. A, flowering branch, x Vr, B, portion of a
cyme with one flower dissected to show detail, X 2 Vz; C, fruits, x 1.
243
FIG. 38. Antirhea lucida. A, flowering branch, x %; B, flower and dissected
flower, x 5; C, portion of fruiting inflorescence, X 1.
244
PIG. 39. Pittoniotis trichantha. A, branch, X 1; B, fruiting inflorescence, x 1; C,
flowers, x 6; D, flower dissected to show detail, x 7 V4.
245
FlG. 40. Anisomeris brachypoda. A, flowering branch, X Vfe; B, flower and
dissection of a flower, X 5; C, fruit, X 3.
246
FlG. 41. Machaonia lindeniana. A, branch in flower, X Vfe; B, flower, X 7 V4; C,
calyx, apex of ovary, and style, X 9; D, corolla dissected showing anthers and
segmented hair, X 9; E, segment of fruiting inflorescence showing enlarged,
chartaceous calyx lobes, X 5.
247
FIG. 42. Chiococca semipilosa. A, flowering branch, X Vz; B, flower and dis-
section of flower, X 3 Vz; C, detail of fruiting inflorescence, X 1 1A; D, fleshy fruit
from side and from edge, X 3.
248
FIG. 43. Asemnantha pubescens. A, flowering branch, X 1; B, detail of fruiting
inflorescence, X 1 Vfc; C, detail of flowering inflorescence, X 1 V4; D, flower, X 5; E,
flower dessected, X 5.
249
FIG. 44. Chione guatemalensis. A, flowering branch, x ¥2; B, flower, x 2V2; C,
flower dissected, X 3; D, portion of fruiting branch, X 3. Drawn from the type.
250
FIG. 45. Coffea arabica. A, branch with immature fruits and flowers, x 1A; B,
partially dissected segment of inflorescence, X 5; C, cross-sections of fruits, X 1; D,
seedling, X V6.
251
FIG. 46. Ixora nicaraguensis. A, fruiting branch, X Vfc; B, flowering branch, x
C, flower, X 4; D and E, flower dissected to show detail, X 4.
252
FlG. 47. Psychotria chiapensis. A, flowering branch, X 1A; B, fruiting branch, X
V£; C, inflorescence in bud, x 1; D, flower, X 2 Vt; E, stylopodium with calyx and
base of style, X 2 V6; F, portion of dissected corolla to show anthers and style, X 3;
G, anthers side and front, x 6; H, fruit, x 1 Vfc.
253
FIG. 48. Psychotria marginata. A, flowering branch, X Vi\ B, flower in natural
position and one dissected to show detail, x 10; C, anthers from front and side, x
20; E, fruits, one dissected, X 4; F, the artist complains to the junior author about
the low rate of pay, x 1/20.
254
FIG. 49. Palicourea galeottiana. A, flowering branch, x W, B, stipule enlarged;
C, fruiting branch, x V£; D, flower showing aestivation, and one in anthesis, X 5; E
and F, flower dissected to show detail, x 6.
255
FIG. 50. Rudgea cornifolia. A, branch, X V6; B, interpetiolar stipule, X 2; C,
buds with flower at anthesis, X 6; D, bud, calyx, and style, X 6; E and F, flower
dissected to show detail, X 6; G, portion of fruiting inflorescence, X 1; H, fruit, x 2.
256
FIG. 51. Declieuxia fruticosa var. mexicana. A, habit, X V6; B, section of stem, X
2; C, portion of fruiting inflorescence, X 2 l/i; D, capsules from the top, side, and
edge, x 5; E, segment of inflorescence, X 4; F, hypanthium, calyx, and style, X 6 1A;
G, corolla dissected, x 6 Vi.
257
CEPHAELI5 GLOMERULATA, n.sp.
FIG. 52. Cephaelis glomerulata. Illustration number 1 from the Botanical
Gazette for 1891 used by permission of the University of Chicago Press.
Magnifications may be calculated from the text.
258
FIG. 53. Mitchella repens. A, habit, X Vz; B, fruiting branch, X 1 Vi; C, flower
dissected to show detail, X 4; D, flowers and leaf from an Arkansas specimen, X 2 VI
Figures A-C from a Guatemalan specimen.
259
FIG. 54. Coussarea imitans. A, flowering branch, X Vz; B, immature flower, X 4;
C, hypanthium, calyx, and style, x 4; D, corolla dissected, X 4; E, mature flower,
dissected, X 2 '/a; F, portion of a fruiting inflorescence, X Vfe; G, fruits from the side
and the edge, x about 1.
260
FIG. 55. Faramea occidentalis. A, fruiting branch, X ¥i; B, portion of
inflorescence at anthesis, note both 4- and 5-lobate corollas in same inflorescence, X
W, C and D, dissections to show detail, x 1 3/4; E, mature fruit, X 1 %.
261
E> I -SlAfO*
FIG. 56. Faramea standleyana. A, portion of flowering branch, X Vz; B, flower,
X 2; C and D, dissections to show detail of flower, X 2.
262
FIG. 57. Appunia guatemalensis. A, flowering branch, x Vfe; B, mature fruits, X
1; C, a flower, X 2 lfa; D, hypanthium, calyx, and style, X 4; E, dissected corolla
showing anthers, x 4.
FIG. 58. Morinda yucatanensis. A, flowering branch, X Vi\ B, inflorescence,
C, part of inflorescence to show calyx, X 2 Vz; D, corolla dissected, X 3 Vz; E,
style, X 3 Vr, F, enlargement of undersurface of leaf to show pubescence.
264
FIG. 59. Richardia scabra. A, habit, x Vfe; B, section of stem to show stipule, x
2 V£; C, an inflorescence with subtending bracts outlined, x 1 Vr, D, flower, x 4; E
and F, flower dissected to show detail, X 5; G, tricoccous ovary, X 4; H, locules of
ovary from front and back, X 4.
265
FIG. 60. Ernodea littoratis. A, fruiting branch and flowering branch of this strand
plant, X W, B, detail from fruiting branch, X 4; C, flower in natural position and
one partially dissected to show detail, X 4.
266
FIG. 61. Crusea calocephala. A, habit, X W, B, interpetiolar stipule, x 1 l/i; C,
flower, x 4; D, E, and F, flower dissected, X 4.
267
FIG. 62. Diodia. D. sarmentosa: A, branch, X Vi; B, flower, X 5; C, flower
dissected, X 5; D, capsule, capsule with one carpel removed and inner face of seed, X
4; E, node and stipule, x 2. D. brasiliensis var. angulata: F, branch, x Vfc; G, node
and stipule, X 3; H, hypanthium and calyx, X 10; I and J, flower dissected, X 7 Vfc.
268
D C
FIG. 63. Hemidiodia ocymifolia. A, branch, x Vfc; B, flower, x 7 V£; C,
hypanthium and calyx, x 6; D, inner face of one locule, X 6; E, both faces of seed,
X 6.
FIG. 64. Borreria laevis. A, habit, X Vr, B, flower, and corolla dissected, X 10; C,
two views of seed, X 7 Vz; D, stipule, X 5.
270
FIG. 65. Spermacoce riparia. A, branch, x ¥2; B, mature fruit and fruit dissected
to show both cocci, x 7; C, flower, x 10; D, flower dissected to show trilobate
corolla and bilobate calyx, x 12 Vz; E, fruits with two and three calyx lobes.
271
FlG. 66. Mitracarpus hirtus. A, branch, X Vfc; B, flower, X 8; C, corolla
dissected, x 10; D, calyx and part of ovary showing dehiscence and seeds in place, x
8; E, base of ovary showing cells and dehiscence, x 8; F, seeds showing two faces, X
10.
272
FIG. 67. Relbunium hypocarpium. A, habit, x V4; B, flowering branch, x 1 Vfc; C,
portion of fruiting branch, x 1 3/4; D, flower, x 10; E, summit of ovary, much
enlarged; F, flower dissected to show details, x 15; G, capsules, x 3; H, seeds, X 5.
273
FIG. 68. Didymaea in Guatemala. Didymaea hispidula: A, habit, x Vfe; B, fruits,
X 2 Vfc, from the type. Didymaea microflosculosa: C, flowering branch, X 2; D,
stipules and partial dissection of flowers, X 7 V6; E, corolla from above showing lobes
and stamens, X 7 Vz. Didymaea australis: F, portion of fruiting branch, natural size.
274
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