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(Last Modified On 8/19/2013)
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(Last Modified On 8/19/2013)
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Family
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RUBIACEAE
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Contributor
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JOHN D. DWYER
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Description
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Trees, shrubs or herbs, occasionally vines. Leaves simple, entire, opposite, occasionally ternate or whorled, the margin regular, usually petiolate; stipules very variable, frequently deciduous, interpetiolar or intrapetiolar, often connate, truncate or more usually with teeth or lobes, often connate to the petiole, leaflike in the Rubieae, the leaves and stipules occasionally united as a whorl of four or more segments. Inflorescences typically cymose, usually cymose-paniculate, small dichasia frequent, the flowers rarely solitary; bracts and bracteoles fre- quently deciduous. Flowers usually bisexual, regular, usually radially symmet- rical, epigynous, usually 4-5-merous; hypanthium mostly oblong or rotund in outline, the calycine cup usually well defined, the teeth or lobes usually present, occasiQnally with interdental glands or with glands within the cup, the lobes equal or unequal, occasionally with one or more lobes foliaceous; corolla usually sal- verform, rotate or funnel shaped, the lobes usually 4-5, valvate, imbricate or twisted, occasionally cucullate; stamens usually 4-5, epipetalous or rarely free of the corolla, the anthers mostly oblong and longitudinally dehiscent, the con- nective occasionally produced at the apex, the filaments usually slender, rarely pubescent; ovary inferior, rarely half-inferior, usually surmounted by a conspic- uous disc, 2(-4-8) loculed, the ovules 1 to many per locule, erect, pendulous or horizontal, mostly adnate to peltate placentas affixed to the median septum, the latter occasionally very thin, rarely evanescent; style simple, usually glabrous,
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Habit
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Trees, shrubs or herbs
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Description
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the stigmas mostly 2, occasionally capitate. Fruits capsular, baccate or drupa- ceous, occasionally a syncarp; drupes with 2 (occasionally 4-5) pyrenes, a fibrous endocarp containing single seeds per carpel; cocci when present dehiscent or indehiscent, the exocarp fleshy, deciduous or occasionally persistent; capsules dehiscing septicidally or loculicidally; berries frequently large and often with a fleshy pulp, the calyx persistent or deciduous, the ovarian disc occasionally con- spicuous; seeds winged or not, rarely with a tuft of hairs, when many then-usually oriented vertically or horizontally.
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Note
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The Rubiaceae is the largest family of dicotyledons in Panama with about 425 species in 87 genera; some new species were encountered after this paper went to press. The family occurs at all altitudes and in all kinds of habitats, the aquatic one excepted, although it may be argued that Lindenia rivalis is a semi-aquatic. Few genera are cultivated horticulturally in Panama, e.g., Ixora. Coffea (coffee) is grown extensively as a food crop. Very few trees yield timber of commercial value. Few medicinally important Rubiaceae occur in Panama, Cephaelis ipe- cacahuanha being noteworthy. One species of Cinchona, the source of quinine bark, has been collected in Panama, and then only twice. Cinchona pubescens, well distributed throughout tropical South America, is native to Panama, but it has not been exploited commercially in Panama. The Rubiaceae of the New World have been studied intensively in this century by only a few workers, in contrast, for example, to the study of African Rubi- aceae. From World War I to World War II, Dr. Paul Standley dominated the study of tropical American Rubiaceae as is evident from the literature cited throughout this paper. Another worker of special note is Dr. Julian Steyermark, who was co-author with Standley of many New World species as well as authoring many of his own. Following Standley's death in 1963, Steyermark became the foremost student of tropical American Rubiaceae. In recent years he has devoted much of his time to the Rubiaceae of the Guyana Highlands and Venezuela. In recent years, several revisions or monographs of individual genera, some covering many countries and others in the form of revisions restricted to Panama, have appeared, e.g., Bouvardia (Blackwell); Cephaelis (Molina); Crusea (Anderson); Galium (Dempster); Hamelia (Elias); Hoffmannia (Dwyer); Richardia (Lewis),; Rondeletia (Kirkbride). The vast majority of Rubiaceae in Panama are trees or shrubs with the genus Psychotria representing almost one-quarter of the species. Subshrubs or rarely herbs like Borreria, Diodia, Spermacoce, etc., are particularly difficult to identify as to species. Despite its size, species of Psychotria in Panama are, for the most part, easier to identify than those of the smaller genera, Hoffmannia and Randia. Hoffmannia is taxonomically difficult, as its stipules are soon deciduous and these rarely present on herbarium material. Added to this is the fact that the flowers and fruits show little interspecific variability. Randia, like Hoffmannia except for a few species, is poorly represented in herbaria and then usually only in fruit. From a taxonomic viewpoint, the leaf characters which are especially note- worthy, apart from the general shape, size, and texture of the blade, as well-as the apex and the base, are the number of lateral veins and whether or not these unite submarginally to form what may be described as an undulate vein; blades which are subsessile or sessile tend to be auriculate or cordate at the base. The stipules of Rubiaceae are of great taxonomic value, although frequently deciduous. Often it is difficult to decide whether the stipules are free or connate and this may lead to an arbitrary decision. There is a wide range of variability in the inflorescences of the Rubiaceae. In terminal inflorescences, the occasional absence of a peduncle may bring the pri- mary branches into a sessile position, thus creating the impression of several independent inflorescences. The capitate inflorescence, while easy to describe in terms of length or width, usually has a complex arrangement of internal bracts and bracteoles. The flowers of the Rubiaceae, despite obvious variability in size and texture of the perianth parts, are rather uniform in general structure. Pollination mech- anisms in the Rubiaceae are in need of intensive study. The calyx usually possesses a cup which is occasionally truncate, but more usually dentate or lobate. The usual 4-5 teeth or lobes are occasionally strongly unequal, e.g., Rondeletia spp. Occasionally one calycine lobe dominates the others in size, e.g., Calycophyllum. More than a quarter of the Panamanian Rubiaceae have glands either within the cup or between the teeth or lobes, a circumstance which is in need of investigation. A spathaceous calyx is rarely encountered, e.g., Duroia or Psychotria boquetensis. In Morinda and Appunia, the hypanthiums of adjacent flowers are connate and this fusion may extend to the calycine cup. As for the flowers of the Rubiaceae, the corolla is usually yellow or white. Faramea is noteworthy as the corolla is usually blue. Occasionally collectors confuse the color of subtending bracts or calycine lobes with that of the true corolla. The corolla varies in length from 2-3 mm to more than 80 mm in Tocoyena and Posoqueria. Whether the lobes are valvate or imbricate is one of the most important taxonomic characters in the Rubiaceae at the generic level. Many species of Psychotria and Cephaelis have the lobes cucullate at the apex. The lobation of each lobe in Joosia is unique among the Panamanian genera. The coriaceous texture of the corolla of Pentagonia, Ladenbergia and Simira is note- worthy. Stamens of the Panamanian Rubiaceae are epipetalous except for Exostema and Coutarea. The fact that the anthers of these two genera are basifixed, an uncommon situation in the Rubiaceae, suggests the possibility that these genera belong to another family. The anthers are usually oblong, although in the Sper- macoceae they tend to be round. Some genera have the anthers usually produced at the apex, e.g., Bertiera, Hamelia, and Hoffmannia spp. Ferdinandusa pan- amensis has dimorphic or possibly trimorphic anthers. Isertia has anthers which are transversely locellate, the endothecium exhibiting a tigriform pattern. The filaments are usually slender, and are terete or plano-convex. Occasion- ally they are crassate, e.g., in Chione and Simira. Filaments tend to be glabrous, although occasional exceptions are encountered, e.g., Alseis has filaments com- pletely pubescent. The point of filament attachment is usually near the middle of the corolla tube or above the middle. Attachment position is related to the little understood phenomenon of heterostyly in the Rubiaceae. Genera with very elon- gate corolla tubes tend to have the anthers on very short filaments at the orifice of the tube, e.g., Ixora spp., Posoqueria, Lindenia, Tocoyena. In describing filament attachment position, it is important that observations be made on fully developed flowers rather than b~uds, as the corolla tube may expand unevenly as the bud approaches anthesis. References made by Anderson (1972) in his recent monograph of Crusea concerning cleistogamy in the genus, as well as protandry being the rule, are noteworthy. Croat (1979) made several references to pollina- tion mechanisms. The dioecious condition is only occasional in Panamanian Rubiaceae: Aliber- tia, Amaioua, Duroia, Randia (some spp.), Stachyarrhena, Borojoa, and pre- sumably in Nertera. Frequently the number of flowers in the inflorescence of one sex may differ greatly from the number characterizing the opposite sex. The ovarian disc is rarely absent; it is variable in shape, e.g., pulvinate, annular, mound shaped; it may be lobed or unlobed, glabrous or rarely pubescent. Secre- tory cells are not evident in sections of the disc, although Anderson (1972) ob- served nectar in young flowers of Crusea. The ovarian disc may persist in the fruit, e.g., in Hamelia and Cosmibuena. The stigma or stigmas (stigmatic lobes) are usually 2, but 3, 4, 5, or more are occasionally encountered. Dressleriopsis has 8 radially disposed stigmas. Stigmas often adhere in the bud and expand later. Most ovaries are bicarpellate with numerous ovules attached to "T-shaped" or peltate placentas, the latter connate with a relatively thick median septum. The fruits of the Rubiaceae are much more versatile at the generic level than are the flowers. They range in length from a few millimeters, e.g., Chimarrhis, to several centimeters in Posoqueria, Randia, Borojoa, Simira, Tocoyena, and Stachyarrhena. Only two genera in Panama have aggregate fruits: Morinda and Appunia. In some fleshy fruits the flesh is ephemeral, falling away to expose two pyrenes, e.g., Psychotria and Cephaelis. Geophila, however, retains the fleshy portion enclosing the two pyrenes. Most fruits are rotund or oblong. The fruits of Chiococca, Coutarea, and Declieuxia are strongly laterally compressed. Ga- lium, Didymaea, and Psychotria acuminata have the didymous pericarps. Oc- casionally fruits are markedly lenticellate, e.g., Coutarea and Exostema. The calyx persists in many genera. In Coussarea and often in Faramea it is stiffly cylindrical resembling a miniature pipe organ. Occasionally the calyx, when de- ciduous, leaves a conspicuous annular scar, e.g., Condaminea, Amaioua, Ha- melia, etc. In Hamelia the persistent ovarian disc may be prominent. Two genera which can be usually identified readily by their fruits are Chione, with the pericarp usually ovate-oblong, rounded at the base and tapering at the apex, and Deppea, being conspicuously turbinate and truncate at the apex. The achenelike alate fruits of Allenanthus are unique in the Rubiaceae. Rarely are fruits wider than long, most species of Faramea being an exception, although the common F. occidentalis has rotund fruits. Elongate fruits tend to be dry; those which are cigar shaped or elongate-turbinate usually suggest the tribe Cinchoneae, but are not restricted to this tribe. Raphides are abundant in all genera of the Rubioideae except Bertiera. Fruits with deciduous calyx are occasional, e.g., Cosmibuena, Amaioua. Very thick- walled fruits are found in a few genera, e.g., Borojoa, Posoqueria, Simira, To- coyena. The majority of tribes have fruits with two seeds. In Coussarea and Faramea (tribe Coussareae) the seeds are solitary. Eight tribes of Panamanian Rubiaceae: Cinchoneae, Condamineae, Gardenieae, Hamelieae, Mussaendeae, Naucleeae, Oldenlandieae, and Rondeletieae, have numerous seeds. The seeds may be embedded in a gelatinous pulp. In addition to 13 genera of the Cinchoneae (Panama), the following genera have alate seeds: Appunia, Arcytophyllum, Bath- ysa, occasionally Rondeletia spp., Mitracarpus, Simira, and Uncaria. In Hillia, the body of the seed has a unipolar tuft of hairs. Capsules may dehisce septici- dally, e.g., Hillia and Ladenbergia, loculicidally, e.g., Condaminea or Rustia, or circumscissilly, e.g., Mitracarpus. Whether the capsule dehisces from apex to base or vice versa is of paramount importance in segregating Cinchona from Ladenbergia. A few genera have the carpels separate with each carpel (mericarp) retaining the seed, e.g., Machaonia and Richardia. In Diodia one mericarp de- -hisces while the other does not. The valves of Joosia twist at dehiscence in conspicuous fashion. The twisting is even more exaggerated in Lindenia.
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Reference
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Anderson, W. R. 1972. A Monograph of the Genus Crusea (Rubiaceae). Mem. New York Bot. Gard. 22: 1-128. Blackwell, W. J. 1968. Revision of Bouvardia (Rubiaceae). Ann. Missouri Bot. Gard. 55: 1-30. Bremekamp, C. 1966. Remarks on the position, the delimitation, and the sub- division of the Rubiaceae. Acta Bot. Neerl. 15: 1-33. Croat, T. 1979. Flora of Barro Colorado Island. Stanford. Darwin, S. 1976. The Genus Lindenia (Rubiaceae). J. Arnold Arbor. 57: 426-449. DeCandolle, A. 1845. Prodromus Systematis Naturalis Regni Vegetabilis 4: 341-622. Paris. Dempster, L. T. 1973. The Fleshy-fruited Galiums (Rubiaceae) of Mexico and Central America. Brittonia 25: 15. Duke, J. 1968. Darien Ethnobotanical Dictionary. Battelle Mem. Inst., Columbus, Ohio. Duke, J. 1970. Manual Dendrologico Para 1000 Especies Arboreas en la Republica de Panamai (Organizacion de Las Naciones Unidas para Agricultura y la Al- imentacion), pp. 1-325. Panama. Dwyer, J. 1968. Borojoa and Tocoyena (Rubiaceae) in Panama. Phytologia 17:445-449. Dwyer, J. 1969. The genus Hoffmannia (Rubiaceae) in Panama. Ann. Missouri Bot. Gard. 56: 269-286. Dwyer, J. & M. V. Hayden. 1966. Notes on Coussarea (Rubiaceae) especially the Panamanian species. Ann. Missouri Bot. Gard. 53: 368-374. Escobar, N. 1972. Flora Toxica de Panama, pp. 1-279. Panama. Hayden, M. V. 1968. Systematic morphological study of New World rubiaceous seeds (Rubioideae sensu Bremekamp). Unpublished Ph.D. Dissertation, St. Louis University, St. Louis, MO. Hayden, M. V. & J. D. Dwyer. 1969. Seed morphology in the tribe Morindeae. Bull. Torrey Bot. Club 96: 704-710. Johnston, I. M. 1949. The Botany of San Jose Island (Gulf of Panama). Sargentia 8: 279-287. Kirkbride, J., Jr. 1968. A revision of the Panamanian species of Rondeletia (Rubiaceae). Ann. Missouri Bot. Gard. 55: 372-391. 1977. Index to the Rubiaceae by Julian A. Steyermark in the Botany of the Guayana Highland by B. Maguire and Collaborators. Phytologia 36: 324- 366. Lewis, W. H. & R. L. Oliver. 1974. Revision of Richardia (Rubiaceae). Brittonia 26: 271-301. The Pharmacopeia of the United States of America. 18th revision. 1970. Mack Pub. Co., Easton, Pennsylvania. Schumann, Karl. 1891. Rubiaceae in Engler & Prantl, Nat. Pflanzenfam. 4: 97- 156. Seeman, B. 1854. The Botany of the H.M.S. Herald (Flora of the Isthmus of Panama), pp. 1-468. London. Standley, P. 1916. Tardavel, a valid generic name to replace Borreria. Contr. U.S. Natl. Herb. 18: 122. Standley, P. 1916. New species of Rubiaceae of several genera chiefly from Panama. Contr. U.S. Natl. Herb. 18: 133-142. Standley, P. 1918. Rubiaceae, Part I. N. Am. Flora 32: 1-86. Standley, P. 1928. Flora of the Panama Canal Zone-Rubiaceae. Contr. U.S. Natl. Herb. 27: 351-365. Standley, P. 1930. Rubiaceae of Colombia. Field Mus. Nat. Hist., Bot. Ser. 7: 1-75. Standley, P. 1932. Rubiaceae, Part II. N. Am. Flora 32: 87-158. Standley, P. 1933. Flora of Barro Colorado Island, Panama. Contr. Arnold Arbor. 5: 148-150. Standley, P. 1934a. Rubiaceae, Part III. N. Am. Flora 32: 159-228. Standley, P. 1934b. Rubiaceae, Part IV. N. Am. Flora 32: 229-300. Standley, P. 1938. Rubiaceae. Flora of Costa Rica. Field Mus. Nat. Hist., Bot. Ser. 18(4): 1264-1380. Standley, P. & L. 0. Williams. 1975. Rubiaceae in Flora of Guatemala, Fieldiana: Botany, Vol. 24, Part XI, nos. 1-3: 1-274. Steyermark, J. A. 1965. Rubiaceae, Part VI. in B. Maguire and Collaborators, The Botany of the Guayana Highland. Mem. New York Bot. Gard. 12: 178-285. Steyermark, J. A. 1967. Rubiaceae, Part VII. in B. Maguire and Collaborators, The Botany of the Guayana Highland. Mem. New York Bot. Gard. 17: 230-436. Steyermark, J. A. 1972. Rubiaceae, Part IX. in B. Maguire and Collaborators, The Botany of the Guayana Highland. Mem. New York Bot. Gard. 23: 227-832. Steyermark, J. A. 1974. Flora de Venezuela-Rubiaceae, primera parte, 9: 1-593; secunda parte, 9: 603-1101; tercera parte, 9: 1111-2070. Caracas. Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Jard. Bot. Belg. 28: 209-290.
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Key
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a. Ovules more than 1 in each cell (subfamily Cinchoneae). b. Fruits fleshy, berrylike. c. Corolla lobes valvate in bud ...... 1. Mussaendeae cc. Corolla lobes imbricate or contorted in bud. d. Corolla lobes imbricate in bud ...... 2. Hamelieae dd. Corolla lobes contorted in bud ...... 3. Gardenieae bb. Fruits dry, capsular. e. Flowers arranged in dense, compact, spherical heads ...... 8. Naucleeae ee. Flowers variously disposed, never in spherical heads. f. Seeds winged or appendaged, vertically imbricate ...... 4. Cinchoneae ff. Seeds not winged, or if winged, horizontal. g. Corolla lobes imbricate or contorted in bud ...... 6. Rondeletieae gg. Corolla lobes valvate in bud. h. Seeds horizontal, usually very numerous; stipules entire or bifid; trees or shrubs ...... 5. Condamineae hh. Seeds imbricate, vertical, usually few; stipules often setose- laciniate; subshrubs or herbs ...... 7. Oldenlandieae aa. Ovules solitary in the cells (subfamily Rubioideae, except Naucleeae). i. Seeds pendulous, the radicle superior; shrubs, trees, or woody vines. j. Flowers in compact globose heads ...... 8. Naucleeae jj. Flowers never in globose heads. k. Stamens inserted in the throat of the corolla ...... 9. Guettardeae kk. Stamens inserted at the base of the corolla tube or on a disc ...... 10. Chiococceae ii. Seeds ascending, the radicle inferior; plants woody or herbaceous. 1. Corolla lobes contorted in bud; shrubs or trees ...... 11. Ixoreae 1. Corolla lobes valvate in bud; habit various. m. Ovules basally attached; mostly shrubs or trees. n. Ovary 7-8 celled ...... 14. Dressleriopsideae nn. Ovary 1-2 celled. o. Ovary 1-celled or 2-celled with a thin septum; fruits 1-seeded ...... 12. Coussareae oo. Ovary 2-celled, or the cells rarely more numerous, the septum thick; fruits 2 or more seeded. p. Stamens usually inserted in the throat of the corolla; flowers perfect ...... 13. Psychotrieae pp. Stamens usually inserted at the base of the corolla tube; flowers often unisexual ...... 15. Anthospermeae mm. Ovules lateral, attached to the septum of the cell; habit various. q. Stipules neither foliaceous nor setiferous; trees or large shrubs; flowers congested by the hypanthium to form a dense head; fruit a syncarp, the base of the hypanthiums united ...... 16. Morindeae qq. Stipules either foliaceous or setiferous; herbs or small shrubs; flow- ers never congested by the hypanthium; carpels free, not united. r. Stipules setiferous; fruit not a syncarp ...... 17. Spermacoceae rr. Stipules foliaceous, similar to the leaves ...... 18. Rubieae 1. MUSSAENDEAE a. Leaves opposite. b. Leaves -with intervenal areas lineolate. c. Petioles not tumescent or swollen. d. Leaves large, ?4-1 m long, often pinnatifid-lobed; calycine cup with rows of conspicuous glands within ...... 65. Pentagonia dd. Leaves small, less than ?4 m long, not pinnatifid-lobed; calycine cup with- out rows of glands within ...... 79. Sommera cc. Petioles tumescent, swollen. bb. Leaves with intervenal areas not lineolate ...... 47. Hippotis e. Inflorescences terminal. f. Plants drying black; leaves and flowers succulent; inflorescence capitate ...... 77. Schradera ff. Plants not drying black; leaves chartaceous, occasionally stiffly so; inflo- rescence spreading (except subcapitate in Amphidaysa). g. Stipules with conspicuous fimbriate processes ...... 5. Amphidaysa gg. Stipules without fimbriate processes. h. Inflorescences spikelike; shrubs ...... 42. Gonzalagunia hh. Inflorescences cymose-corymbose or cymose paniculate; trees. i. Corolla 3 or more cm long; anthers transversely locellate ...... 49. Isertia ii. Corolla less than 1.5 cm long; anthers not transversely locellate ...... 69. Raritebe ee. Inflorescences axillary. j. Plants prostrate; fruits bright china blue ...... 21. Coccocypselum jj. Plants erect or climbing; fruits not bright china blue ...... 76. Sabicea aa. Leaves alternate ...... 31. Didymochlamys 2. HAMELIEAE a. Ovary 4-5 celled; inflorescences regularly terminal; stamens with anther connectives reg- ularly produced at the apex. b. Corolla lobes contorted in bud; raphides absent from leaves ...... 10. Bertiera bb. Corolla lobes imbricate in bud; raphides present in leaves ...... 44. Hamelia aa. Ovary 2-3 celled; inflorescences always axillary; stamens with the anther connective not produced at the apex (few species of Hoffmannia excepted). c. Ovary 2 celled; calyx with lobes ca. 8 mm long; corolla lobes valvate ...... 87. Xerococcus cc. Ovary 2-3 celled; calyx with lobes less than 5 mm; corolla lobes imbricate ...... 48. Hoffmannia 3. GARDENIEAE a. Plants armed with spines on stems and short shoots ...... 69. Randia aa. Plants unarmed. b. Inflorescences not spikelike; fruits not gourdlike nor on an elongate curved pedicel. c. Flowers perfect. d. Inflorescences with flowers several to numerous, flowers disposed in a candelabra array. e. Flower buds curved at the apex; seeds with testa cells more than twice as long as wide ...... 66. Posoqueria ee. Flower buds not curved at the apex; seeds with testa cells less than twice as long as wide ...... 83. Tocoyena dd. Inflorescences with flowers few or solitary, not disposed in a candelabra array. f. Leaves drying black ...... 40. Genipa ff. Leaves not drying black. cc. Flowers unisexual. g. Fruits oblong; terminal stipules forming a conic cap. h. Pistillate flowers capitate or cymose; fruits not densely hirsute ...... 4. Amaioua hh. Pistillate flowers usually solitary; fruits densely hirsute ...... 34. Duroia gg. Fruits rotund; terminal stipules not forming a conic cap. i. Fruits subtended by several persistent bracts, as large as a child's hand, superficially rough ...... 11. Borojoa ii. Fruits not subtended by several persistent bracts, to 2.5 cm wide,superficially smooth ...... 1. Alibertia bb. Staminate inflorescences spikelike; fruit a gourdlike capsule on an elongate curved pedicel ...... 81. Stachyarrhena 4. CINCHONEAE a. Vines ...... 57. Manettia aa. Shrubs or trees. b. Stamens with anthers dimorphic or trimorphic ...... 38. Ferdinandusa bb. Stamens with anthers all alike. c. Calyx with one lobe expanded as a blade. d. Inflorescence cymose or corymbiform; stamens equal and exserted ...... 14. Calycophyllum dd. Inflorescence spicate; stamens unequal, not exserted 86. Wittmackanthus cc. Calyx with all lobes similar. e. Plants epiphytic; leaves fleshy, the lateral veins often evanescent. f. Flowers pink or red, the corolla inflated ...... 70. Ravnia ff. Flowers white or yellow. g. Seeds winged terminally ...... 24. Cosmibuena gg. Seeds with a tuft of hairs at one pole ...... 46. Hillia ee. Plants terrestrial; leaves not fleshy, mostly drying papery to coriaceous, veins evident to the margin. h. Inflorescences spikelike; stamens with filaments villose, attached at the base of the corolla tube ...... 3. Alseis hh. Inflorescences various but not spikelike; stamens attached above the base of the corolla tube (except Exostema and Coutarea). i. Corolla lobes valvate in bud. j. Flowers 4-merous; capsule rotund ...... 13. Bouvardia Jj. Flowers 5-merous; capsule oblong. k. Capsule splitting from below toward the apex ...... 20. Cinchona kk. Capsule splitting from above toward the base. 1. Corolla lobes split at the apex; fruits forming 4 coiled valves ...... 51. Joosia 11. Corolla lobes not lobed at the apex; fruits not forming coiled valves. m. Capsule loculicidal ...... 56. Macrocnemum mm. Capsule septicidal ...... 52. Ladenbergia ii. Corolla lobes imbricate or contorted in bud. n. Stamens conspicuously exserted; corolla not inflated; fruits neither compressed nor lenticellate ...... 36. Exostema nn. Stamens not conspicuously exserted; corolla inflated; fruits compressed, strongly lenticellate ...... 26. Coutarea 5. CONDAMINEAE a. Leaves glandular-punctate, the dots pellucid; anthers opening by terminal pores ...... 75. Rustia aa. Leaves not glandular-punctate; anthers longitudinally dehiscent. b. Flowers ca. 3 mm long; inflorescences axillary ...... 16. Chimarrhias bb. Flowers more than 3 mm long; inflorescences terminal. c. Calycine lobes unequal, one expanded into a large colored limb ...... 65. Pogonopus cc. Calycine lobes equal or nearly so ...... 23. Condaminea 6. RONDELETIEAE a. Plants woody. b. Corolla lobes imbricate in the bud. c. Calycine lobes unequal, often one expanded into a foliose blade. d. Calyx with foliose blade a flaming red color ...... 85. Warscewiczia dd. Calyx with foliose blade white ...... 73. Rondeletia (some spp.) cc. Calycine lobes equal or nearly so. e. Fruits large, nutlike, dehiscing septicidally; seeds winged; wood on ex- posure to air turning a purple red; filaments geniculate at point of attachment ...... 78. Simira ee. Fruits small, dehiscing loculicidally or septicidally; seeds not winged; wood on exposure to air not turning red; filaments not geniculate at point of attachment. f. Calycine lobes often unequal; capsule with valves not woody, not bifid at the apex ...... 73. Rondeletia ff. Calycine lobes usually equal; capsule with valves woody, bifid at the apex ...... 9. Bathysa bb. Corolla lobes contorted in bud. g. Trees, terrestrial; corolla very short, the tube shorter than the lobes; fruits with valves not coiled at maturity. h. Corolla glabrous within, 4-merous ...... 29. Deppea hh. Corolla villous within, 5-merous ...... 35. Elaeagnia gg. Shrubs, growing in water or on banks of streams; corolla elongate, the tube at least S times the length of the lobes, fruits with valves coiled at maturity ...... 54. Lindenia aa. Plants herbaceous ...... 53. Limnosipanea 7. OLDENLANDIEAE a. Plants herbaceous, often prostrate or subprostrate, not ericoid in habit; seeds angular or winged ...... 62. Oldenlandia aa. Plants subshrubs, ericoid in habit, usually upright; seeds carinate or plano-convex ...... 8. Arcytophyllum 8. NAUCLEEAE Vine with recurved spines and inflorescences axillary and globose ...... 84. Uncaria 9. GUETTARDEAE a. Plants armed with rigid spines on twigs and stems. b. Corolla lobes imbricate in bud; fruits dry, splitting into 2 indehiscent mericarps or cocci ...... 55. Machaonia bb. Corolla lobes valvate in bud; fruits fleshy, not splitting into 2 indehiscent mericarps ...... 9. Chomelia aa. Plants unarmed. c. Corolla lobes imbricate in bud; leaves with intervenal areas not lineolate-striate. d. Fruits dry, separating into 2 indehiscent mericarps or cocci; flowers not secundly disposed ...... 55. Machaonia dd. Fruits fleshy; flowers secundly disposed. e. Fruits covered with a fine feltlike pubescence ...... 43. Guettarda ee. Fruits glabrate ...... 6. Antirhea cc. Corolla lobes valvate in bud; leaves of some species with intervenal areas lineolate-striate ...... 19. Chomelia 10. CHIOCOCCEAE a. Flowers 4-merous; fruits winged ...... 2. Allenanthus aa. Flowers 5-merous; fruits not winged. b. Corolla lobes imbricate in bud; fruits not strongly compressed; shrubs (Panama) ...... 18. Chione bb. Corolla lobes valvate in bud; fruits strongly compressed; trees, shrubs, or woody vines ...... 17. Chiococca 11. IXOREAE a. Bractlets connate, calyx-like or involucral; inflorescences axillary; cultivated as a tree crop ...... 22. Coffea aa. Bractlets distinct, not calyx-like; inflorescences terminal; some species cultivated as ornamentals ...... 50. Ixora 12. COUSSAREAE a. Ovules connate, borne on a common basal column; seeds vertical; flowers usually white ...... 25. Coussarea aa. Ovules separate in a one-celled ovary; seeds horizontal; flowers often blue ...... 37. Faramea 13. PSYCHOTRIEAE a. Plants prostrate, woody herbs; leaves cordate ...... 41. Geophila aa. Plants erect subshrubs, shrubs or trees (except Declieuxia, a low woody herb [Panama]); leaves various at the base, not cordate. b. Lobes of calyx, if present, subequal, not polymorphic at the apex. c. Inflorescence an involucrate head, the heads solitary, sessile or pedunculate, or with 3 heads arising from a common peduncle ...... 15. Cephaelis cc. Inflorescence bracts not congested into an involucre, flowers not congested into a head. d. Herbs; fruits laterally compressed ...... 28. Declieuxia dd. Trees, shrubs, or subshrubs, rarely epiphytes; fruits terete in cross section. e. Seeds with an incurved ventral surface; stipules pectinate ...... 74. Rudgea ee. Seeds not with an incurved ventral surface; stipules rarely pectinate, if so, membranous. f. Corolla tube short, to 0.7 cm long (P. chiapensis and P. ostaurea excepted), not swollen at the base; flowers white, green, or yellow; stipules of twiglets deciduous or persistent ...... 68. Psychotria ff. Corolla tube elongate, more than 0.7 mm long, usually swollen at the base; flowers regularly orange and purple; stipules of twiglets regularly persistent ...... 64. Palicourea bb. Lobes of calyx unequal, polymorphic at the apex ...... 59. Montamans 14. DRESSLERIOPSIDEAE Ovary and fruit 7-8-carpellate ...... 33. Dressleriopsis 15. ANTHOSPERMEAE Plants forming prostrate mats ...... 61. Nertera 16. MORINDEAE a. Plants usually drying black, or at least fruit drying black; fruits succulent, 1 cm or more in diam. ...... 60. Morinda aa. Plants not drying black; fruits dry, less than 1 cm in diam. ...... 7. Appunia 17. SPERMACOCEAE a. Fruits with circumscissile dehiscence ...... 58. Mitracarpus aa. Fruits longitudinally dehiscent or separating into open or closed valves or cocci, lacking circumscissile dehiscence. b. Cocci 3-5(-6) ...... 72. Richardia bb. Cocci 2. c. Stipules with sheath with more than 3 setae; plants not fetid. d. Cocci indehiscent. e. Calyx lobate; cocci separating from a central axis or carpophore ...... 27. Crusea ee. Calyx dentate; cocci not separating from a central axis or carpophore ...... 32. Diodia dd. Cocci dehiscent, often only 1 of the 2 opening. f. Cocci opening only at the base; flowers axillary ...... 45. Hemidiodia ff. Cocci opening at the apex; flowers axillary or terminal. g. Fruits with cells similar, both dehiscent ...... 12. Borreria gg. Fruits with cells dissimilar, only one opening ...... 80. Spermacoce cc. Stipules with sheath with only 3 setae; plants fetid ...... 82. Tabagoa 18. RUBIEAE a. Flowers with the hypanthium immediately subtended by an involucre of leaflike bracts ...... 71. Relbunium aa. Flowers with the hypanthium not immediately subtended by an involucre of leaflike bracts. b. Leaves opposite ...... 30. Didymaea bb. Leaves verticillate ......39. Galium ARTIFICIAL KEY1. Leaves alternate ......31. Didymochlamys 1'. Leaves opposite or whorled. 2. Plants armed. 3. Vines; spines recurved. 4. Inflorescence axillary, globose capitate, epedunculate ...... 84. Uncaria 4'. Inflorescence terminal, patulous, pedunculate ...... 69. Randia 3'. Shrubs or trees; spines straight. 5. Fruits dry, splitting into dehiscent mericarps or cocci ...... 55. Machaonia 5'. Fruits juicy, not dehiscent. 6. Fruits to 5 mm long; peduncles elongate ...... 19. Chomelia 6'. Fruits 10-50 mm long; peduncles absent or scarcely measurable ...... 69. Randia 2'. Plants not armed. 7. Fruits dry, the valves strongly coiled at maturity. 8. Shrubs, located in stream beds or on banks; flowers usually more than 5 cm long; fruits 1 cm or more wide ...... 54. Lindenia 8'. Trees, not located in stream beds or on banks; flowers less than 1.5 cm long; fruits ca. 0.3 cm wide ...... 51. Joosia 7'. Fruits dry or juicy, if valvular, the valves not strongly coiled. 9. Trees, shrubs, or subshrubs. 10. Plants epiphytic. 11. Flowers and fruits over 2 cm in length; fruits dry, capsular, cigar shaped. 12. Leaves to 5 cm wide; seeds with a tuft of hairs at each end ...... 24. Cosmibuena12'. Leaves to 3.5 cm wide; seeds with a tuft of hairs at one end ...... 46. Hillia 11'. Flowers and fruits less than 2 cm long; fruits fleshy, sometimes appearing dry, of 4 pyrenes, round or oblong ...... 68. Psychotria 10'. Plants not epiphytic. 13. Plants cauliflorous, flowers arising laterally on branches ...... 69. Randia 13'. Plants not cauliflorous; inflorescences axillary or terminal. 14. Calyx spathaceous. 15. Flowers ca. 1 cm long; fruits of 2 pyrenes, each 1-seeded ...... 68. Psychotria 15'. Flowers ca. 4 cm long; fruits baccate, many-seeded ...... 47. Hippotis 14'. Calyx not spathaceous. 16. Leaves pellucid punctate; anthers porocidally dehiscent ...... 75. Rustia 16'. Leaves not pellucid punctate; anthers longitudinally dehiscent. 17. Fruits oblong and gourdlike, 5-7 cm long, on an elongate reflexed pedicel ...... 81. Stachyarrhena 17'. Fruits, if oblong, not gourdlike, 0.3-6.0 cm long, not on an elongate reflexed pedicel. 18. Flowers obviously secundly disposed. 19. Carpels 2. 20. Ovules attached basally, 1 per cell ...... 68. Psychotria 20'. Ovules axile or pendent, 1-many per cell. 21. Ovules 1 per cell, pendulous from the ovary apex ...... 19. Chomelia 21'. Ovules numerous, attached laterally to the placenta. 22. Fruits with a fine feltlike pubescence; shrubs; anthers included -43. Guettarda 22'. Fruits glabrous or glabrescent; trees; anthers exserted. 23. Fruits dry ...... 3. Alseis 23'. Fruits fleshy ...... 6. Antirhea 19'. Carpels 4 ...... 44. Hamelia 18'. Flowers not secundly disposed but arranged opposite or alternate on the axis. (Alseis excepted, filaments villose). 24. Inflorescence with the flowers or flower clusters spicate. 25. Flowers (at least some) with one calycine lobe foliaceous. 26. Foliaceous lobe of calyx white or pink but not a flaming red; seeds winged ...... 86. Wittmackanthus 26'. Foliaceous lobe of calyx a flaming red; seeds unwinged ...... 85. Warszewiczia 25'. Flowers with the calycine lobes, if present, subequal. 27. Leaves chartaceous; flowers bisexual; fruits less than 2 cm long. 28. Stamens obviously exserted, attached at the base of the corolla tube, the filaments villose; fruits secundly disposed ...... 3. Alseis 28'. Stamens barely exserted, epipetalous, the filaments glabrous; fruits alternate or opposite on the axis ...... 42. Gonzalagunia 27'. Leaves coriaceous; flowers unisexual; fruits more than 5 cm long, melonlike ...... 81. Stachyarrhena 24'. Inflorescences with the flowers paniculate, racemiform, or flowers densely aggregated in leaf axils or occasionally solitary, not spicate. 29. Corolla inflated, medianly campanulate, urceolate or ellipsoid. 30. Corolla 4-6 cm long. 31. Shrubs sprawling; leaves coriaceous when dry, the lateral veins scarcely evident or evanescent; inflorescences with the flowers 1-3, z pink or red; anthers versatile; fruits cigar shaped, scarcely lenticellate ...... 70. Ravnia 31'. Shrubs upright; leaves chartaceous, the lateral veins conspicuous; inflorescence with the flowers more than 3, white; anthers basifixed; fruits purse shaped, strongly lenticellate ...... 26. Coutarea 30'. Corolla less than 4 cm long. 32. Anthers dimorphic or trimorphic; leaves and flowers not drying black ...... 38. Ferdinandusa 32'. Anthers all alike; leaves and flowers drying black ...... 77. Schradera 29'. Corolla tubular or scarcely inflated. 33. Calyx with one or all of the lobes foliaceous. 34. Leaves with the intervenal areas not lineolate-striate, only one or no calycine lobe in some flowers foliaceous. 35. Expanded calycine lobe flaming red, 2-3 times longer than wide ...... 85. Warszewiczia 35'. Expanded calycine lobe white, pink, or dull red, less than 2 times longer than wide. 36. Expanded calycine lobe to 1.5 cm wide; fruits globose ...... 73. Rondeletia 36'. Expanded calycine lobe 1.5-6.0 cm wide; fruits oblong. 37. Inflorescence paniculate. 38. Expanded calycine lobe white; seeds winged ...... 14. Calycophyllum 38'. Expanded calycine lobe red; seeds not winged ...... 65. Pogonopus 37'. Inflorescence spicate ...... 86. Wittmackanthus 34'. Leaves with the intervenal areas lineolate-striate; all calycine lobes foliaceous ...... 79. Sommera 33'. Calyx lobes, if present, dentate or lobular, not foliaceous. 39. Flowers bisexual. 40. Corolla contorted dextrosely; fruits capsular, to 5 mm long, with 4 persistent calycine lobes -29. Deppea 40'. Corolla contorted sinistrorsely or valvate or imbricate; fruits berrylike or of 2-4 pyrenes; if fruits capsular, usually more than 5 mm long, the often persistent calycine lobes invariably 5. 41. Flowers minute, corolla tube 2-3 mm long. 42. Ovules 2, basally attached; fruits of 2 pyrenes. 43. Flowers in involucrate heads ...... 15. Cephaelis 43'. Flowers in open inflorescences not in involucrate heads ...... 68. Psychotria 42'. Ovules numerous, attached medianly to the septum; fruits capsular ...... 16. Chimarrhis 41'. Flowers with the corolla tube more than 3 mm long. 44. Fruits winged, achenelike ...... 2. Allenanthus 44'. Fruits berrylike or capsular, the surface smooth or with distinct ridges. 45. Stamen filaments ca. 15 mm long, free from the corolla tube, attached in a ring at the base of the corolla tube ...... 36. Exostema 45'. Stamen filaments less than 10 mm long, epipetalous. 46. Fruits capsular. 47. Fruits 1.0-1.5 cm long, topped by a conspicuous calycine scar ...... 23. Condaminea 47'. Fruits usually more than 1.5 cm long; if less than 1.5 cm long, not topped by a persistent annular calycine scar. 48. Capsule splitting from below toward the apex; fruits ovoid or oblong ...... 20. Cinchona 48'. Capsule splitting from above toward the base; fruits cigar shaped or linear-oblong. 49. Fruits forming 4 coiled valves; corolla lobes split apically ...... 51. Joosia 49'. Fruits not forming coiled valves; corolla lobes not split at the apex. 50. Capsule loculicidal; flowers to 12 mm long ...... 56. Macrocnemum 50'. Capsule septicidal; flowers to 40 mm long ...... 52. Ladenbergia a 46'. Fruits juicy and berrylike (except Machaonia), not capsular. 51. Ovary 4-5-celled; anther connective regularly produced at the apex. 52. Corolla lobes contorted in the bud; cystoliths absent ...... 10. Bertiera 52'. Corolla lobes imbricate in the bud; cystoliths abundant ...... 44. Hamelia 51'. Ovary 2- or 8-celled (Dressleriopsis); anthers apically obtuse or emarginate (except Raritebe). 53. Corolla imbricate or contorted in bud. 54. Fruits dry, splitting into mericarps or cocci; shrubs, occasionally armed ...... 55. Machaonia 54'. Fruits baccate, indehiscent; trees or shrubs, unarmed ...... 50. Ixora 53'. Corolla valvate in bud. 55. Twigs with wood on exposure to air bright red or purple; stamen filaments basally geniculate ...... 78. Simira 55'. Twigs with wood on exposure to air not bright red or purple; stamen filaments basally straight. 56. Flowers with the hypanthiums and calycine cups of adjacent flowers partially connate; fruit a syncarp. 57. Fruits succulent; plants usually drying black ...... 60. Morinda 57'. Fruits dry; plants not drying black ...... 7. Appunia 56'. Flowers with the hypanthiums and calycine cups of adjacent flowers free; fruits with carpels free. 58. Leaves with intervenal areas lineolate-striate. 59. Leaves pinnatifid-lobed ...... 65. Pentagonia 59'. Leaves not pinnatifid-lobed. 60'. Leaves small, 10-25 cm long ...... 65. Pentagonia 61. Calycine lobes foliaceous, ovules numerous, the placenta lateral ...... 79. Sommera 61'. Calycine lobes dentate; ovules 1 per cell, pendent from the apex ...... 19. Chomelia 58'. Leaves with the intervenal areas not lineolate-striate. 62. Leaves cordate or auriculate at the base; carpels 8 ...... 33. Dressleriopsis 62'. Leaves acute, obtuse or occasionally truncate at the base; carpels 2-5(-6). 63. Fruits covered with a fine feltlike pubescence ...... 43. Guettarda 63'. Fruits glabrous or sparsely hairy. 64. Inflorescences axillary. 65. Ovules several per cell. 66. Calycine lobes 4.0-5.5 mm long ...... 87. Xerococcus 66'. Calycine lobes 0.5-1.0 mm long ...... 48. Hoffmannia 65'. Ovules 1 per cell. 67. Fruits baccate; bracteoles connate into a calyxlike cup; cultivated ...... 22. Coffea 67'. Fruits of 2 pyrenes surrounded by a soon disintegrating pulp; wild species. 68. Calycine lobes, at least some, compressed galeate at the apex ...... 59. Montamans 68'. Calycine lobes not galeate at the apices, acute or obtuse. 69. Flowers in involucrate heads ...... 15. Cephaelis 69'. Flowers in open inflorescences, not in involucrate heads ...... 68. Psychotria 64'. Inflorescences terminal. 70. Inflorescence disposed like a candelabrum or corymbose-capitate, the flowers upright. 71. Inflorescence corymbose-capitate; flowers red or white; corolla tube 2.0-4.5 cm long, to 1.5 mm wide; cultivated ...... 50. Ixora 71'. Inflorescences disposed like a candelabrum; flowers white; corolla tube 8-15 cm long, to 3 mm wide; wild species. 72. Buds with corolla lobes oriented as an eccentric mass; fruits smooth ...... 66. Posoqueria 72'. Buds with corolla lobes oriented centrally; fruits rough ...... 83. Tocoyena 70'. Inflorescences paniculate, thrysiform, globose-capitate, occasionally spiciform or racemiform, not disposed as a candelabrum, the flowers usually lax. 73. Calyx cup prominent in fruit, arising on upper 1/3 of fruit and obscuring the fruit apex ...... 9. Bathysa 73'. Calyx cup terminal on the fruit or absent. 74. Ovules pendulous ...... 18. Chione 74'. Ovules or ovule not pendulous. 75. Ovules numerous per cell. 76. Flowers small, the corolla to 3 mm long. 77. Fruits loculicidally bivalvate, the valves ultimately bifid ...... 35. Elaegnia 77'. Fruits septicidally valvate, the valves undivided ...... 16. Chimarrhis 76'. Flowers large, the corolla 4-50 mm long. 78. Anthers transversely locellate; ovary 2-6-celled ...... 49. Isertia 78'. Anthers not transversely locellate; ovary 2-celled ...... 34. Raritebe 75'. Ovules 1 per cell. 79. Fruits 1-seeded by abortion, one pyrene empty. 80. Ovules connate, attached to a basal column; flowers white; fruits longer than wide ...... 25. Coussarea 80'. Ovules separate, collateral, basal; flowers blue, occasionally white; fruits as wide or wider than long ...... 37. Faramea 79'. Fruits of 2 pyrenes, each with a single seed. 81. Fruits an involucral head, the single flowers enclosed by a pair of bracteoles ...... 15. Cephaelis 81'. Fruits not an involucral head, the single flowers not enclosed in a pair of bracteoles. 82. Stipules without teeth on the dorsal surface or margins; ovules separated by a thick septum; 2 seeds present, one in each pyrene. 83. Flowers white or yellow; corolla tube not inflated basally; stipules free, occasionally connate, usually deciduous ...... 68. Psychotria 83'. Flowers often orange or purple; corolla tube inflated basally; stipules usually connate, usually persistent ...... 64. Palicourea 82'. Stipules with subulate or prickly teeth arranged very irregularly on the dorsal surface and/or margins; ovules connate; one seed occasionally aborted ...... 74. Rudgea 39'. Flowers unisexual. 84. Fruits with seeds drying in a gelatinous pulp; mature fruits exceeding 1 cm in diameter. 85. Leaves usually drying black, chartaceous. 86. Fruits 4.5-8.0 cm in diameter, the most terminal stipules involucral, immediately subtending the fruit, tending to persist ...... 11. Borojoa 86'. Fruits to 4 cm wide, the terminal stipules not involucral, usually deciduous ...... 40. Genipa 85'. Leaves usually drying brown, deep red-brown or tan, chartaceous to coriaceous. 87. Leaves subcoriaceous or coriaceous. 88. Fruits densely hairy, the persistent calyx to 3 cm long ...... 47. Hippotis 88'. Fruits glabrous, the calyx deciduous or to 1 cm long. 89. Fruits oblong, 1.0-1.5 cm long, wrinkled on drying; corolla tube less than 1 cm long ...... 4. Amaioua 89'. Fruits rotund, 3-9 cm long, smooth on drying; corolla tube more than 1 cm long ...... 40. Genipa 87'. Leaves chartaceous. 90. Fruits rotund, drying without wrinkling; calyx present on fruit; corolla tube less than 1 cm long ...... 1. Alibertia 90'. Fruits oblong, obviously wrinkled on drying; calyx absent on fruit; corolla tube more than 1 cm long ...... 4. Amaioua 84'. Fruits with seeds drying without an obvious pulp; mature fruits not exceeding 1 cm in diameter ...... 4. Amaioua 9'. Vines or herbs. 91. Vines. 92. Inflorescences globose-capitate, epedunculate; plants armed ...... 84. Uncaria 92'. Inflorescences patulous, not globose-capitate, usually pedunculate; plants unarmed. 93. Fruits plano-convex, white, often drying white ...... 17. Chiococca 93'. Fruits globose, not plano-convex, orange, purple, brown, green, black, or dark blue, not drying white. 94. Leaves verticillate; stems )weak; flowers white; fruits less than 4 mm long.95. Flowers with the hypanthium immediately subtended by a whorl of 4 bracts ...... 71. Relbunium 95'. Flowers with the hypanthium not subtended by a whorl of bracts ...... 39. Galium 94'. Leaves not verticillate; stems rigid; flowers white to red; fruits 5-10 mm long. 96. Flowers white; ovary 2-5-celled; fruits baccate; leaves pubescent ...... 76. Sabicea 96'. Flowers white to red; ovary 2-celled; fruits capsular; leaves glabrous to glabrescent ...... 57. Manettia 91'. Herbs, sometimes woody. 97. Plants prostrate, creeping. 98. Leaves cordate ...... 41. Geophila 98'. Leaves linear, lanceolate, oblong, or ovate, not cordate. 99. Leaves small or minute, 2-14 mm long. 100. Fruits strongly didymous ...... 30. Didymaea 100'. Fruits not strongly didymous. 101. Leaves to 3 mm long; stipules not connate with petiole ...... 62. Oldenlandia 101'. Leaves 4-13 mm long; stipules connate with petiole ...... 61. Nertera 99'. Leaves relatively large, more than 15 mm long. 102. Leaves 1-5 cm wide, usually densely hairy; fruits fleshy. 103. Fruits strongly didymous, wider than long, glabrous or glabrescent, the calyx deciduous ...... 30. Didymaea 103'. Fruits subrotund, not didymous, longer than wide, pubescent, the calyx persistent ...... 21. Coccocypselum 102'. Leaves less than 0.5 cm wide (Panama); glabrous or glabrescent, fruits dry ...... 62. Oldenlandia97'. Plants erect. 104. Plants ericoid in appearance, the leaves subcoriaceous. 105. Leaves with lateral veins inconspicuous or absent; stipules marginally erose; fruits a septicidal capsule ...... 8. Arcytophyllum 105'. Leaves with lateral veins obvious; stipules with margins entire; fruit a schizocarp ...... 28. Declieuxia 104'. Plants not ericoid in appearance, the leaves chartaceous. 106. Stipules obsolete; leaves ternate (Panama) ...... 53. Limnosipanea 106'. Stipules conspicuous, usually persistent (Hoffmannia excepted); leaves opposite or more than 3 at a node. 107. Plants with 2 basally attached ovules ...... 68. Psychotria 107'. Plants with ovules 2 to several, not basally attached. 108. Leaves 4-7 cm wide; stipules free of the petiole, caducous or more than 2 cm long. 109. Stipules deciduous, rarely seen; corolla tube to 1 cm long ...... 48. Hoffmannia 109'. Stipules present, 2-3 cm long; corolla tube 2-3 cm long ...... 5. Amphidaysa 108'. Leaves 1.0-3.8 cm wide; stipules connate and adnate to the petiole, to 1 cm long. 110. Fruits capsular, loculicidally bivalvate ...... 13. Bouvardia 110'. Fruits of 2-5 cocci. 111. Fruits circumscissile ...... 58. Mitracarpus 111'. Fruits longitudinally dehiscent or separating into open or closed valves or cocci; not circumscissile. 112. Cocci 3-5(-6) ...... 72. Richardia 112'. Cocci 2. 113. Stipular sheath with more than 3 setae. 114. Cocci indehiscent. 115. Calyx lobate; cocci separating from a central axis or carpophore ...... 27. Crusea 115'. Calyx dehtate; cocci not separating from a central axis or carpophore ...... 32. Diodia114'.; Cocci dehiscent, often only one of the two opening. 116. Cocci opening only at the base ...... 45. H-emidiodia 116'. Cocci opening at the apex. 117. Fruits with cells similar, both dehiscent ...... 12. Bcrreria 117'. Fruits with cells dissimilar, only one opening ...... 80. Spermacoce 113'. Stipular sheath with only 3 setae ...... 82. Tobagoa
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