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, Petn (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 Michoacn 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; Petn; 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. 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