ANNALS OF THE MISSOURI BOTANICAL GARDEN FLORA OF PANAMA! BY ROBERT E. Woopson, JR. AND ROBERT W. SCHERY and Collaborators Part IX FAMILY 179. RUBIACEAE—PART 1. JOHN D. DWYER? 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, occasionally 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, ! Assisted by National Science Foundation Grant DEB 77-04300, (Principal Investigator) W. G. D'Arc ssouri Botanical Garden, pid Office Box 299, St. Louis, Missouri 63166, and St. Louis University, 221 North Grand, St. Louis, Missouri 63103. Part 2 will appear in Vol. 67, no. 2 of the ANNALS of the Missouri Botanical Garden After this paper went to press, several T in the Panamanian Rubiaceae were waad These will be published in a supplement in this journ ANN. Missouri Bor. GARD. 67: 1-256. à 0026-6493/80/1—256/$25.75/0 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 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. 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). 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. s 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 full show little interspecific variability. Randia, like Hoffmannia except for a few species, is poorly represented in herbaria and then usually only in fruit. rom 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 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 3 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- 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. /sertia 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 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 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 buds, 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- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 5 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. Illustrations for this treatment were all prepared from material at Missouri Botanical Garden (MO). Literature: 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. k eae A 1845. Prodromus Systematis Naturalis Regni Vegetabilis 4: 341— 622. Par Dempster, E T. 1973. The Fleshy-fruited Galiums (Rubiaceae) of Mexico and Central America. Brittonia 25: 15. Duke, J. 1968. Darien Ethnobotanical Dictionary. Battelle Mem. Inst., Colum- bus, Ohio. . 1970. Manual Dendrólogico Para 1000 Especies Arboreas en la Republica de Panamá (Organización 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. 969. The genus Hoffmannia (Rubiaceae) in Panama. Ann. Missouri Bot. Gard. 56: 269—286. & 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 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 seeds (Rubioideae sensu Bremekamp). Unpublished Ph.D. Dissertation, St. Louis University, St. Louis, MO. & 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. . 1916. New species of Rubiaceae of several genera chiefly from Panama. Contr. U.S. Natl. Herb. 18: 133-142. . 1918. Rubiaceae, Part I. N. Am. Flora 32: 1-86. . 1928. Flora of the Panama Canal Zone—Rubiaceae. Contr. U.S. Natl. Herb. 27: 351-365. . 1930. Rubiaceae of Colombia. Field Mus. Nat. Hist., Bot. Ser. 7: 1-75. —. 1932. Rubiaceae, Part II. N. Am. Flora 32: 87-158. . 1933. Flora of Barro Colorado Island, Panama. Contr. Arnold Arbor. 5: 148—150. . 1934a. Rubiaceae, Part III. N. Am. Flora 32: 159-228. . 1934b. Rubiaceae, Part IV. N. Am. Flora 32: 229-300. . 1938. Rubiaceae. Flora of Costa Rica. Field Mus. Nat. Hist., Bot. Ser. 18(4): 1264-1380. & L. O. 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. . 1967. Rubiaceae, Part VII. in B. Maguire and Collaborators, The Botany of the Guayana Highland. Mem. New York Bot. Gard. 17: 230—436. . 1972. Rubiaceae, Part IX. in B. Maguire and Collaborators, The Botany of the Guayana Highland. Mem. New York Bot. Gard. 23: 227-832. . 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. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 7 KEYS TO THE TRIBES AND GENERA OF THE RUBIACEAE a. Ovules more than | in each cell (subfamily Cinchoneae). b. Fruits Lx berrylike. Dee valvatein bud |... 2. 1. Mussaendeae cc. Corolla = imbricate or contorted in bud. rolla lobes imbricate in bud UU l U a. 2. Hamelieae d Corolla lobes contorted in bud... 1. 3. Gardenieae bb. Fruits dry, capsular wers arranged i in dense, compact, d ei E TEENS 8. Naucleeae ee. Flowers variously disposed, never in spherical hea eds winged or appendaged, vertically rc Seeds not winged, or if winged, horizontal. g. Corolla lobes ibus or contorted in bud ___________- 6. Rondeletieae gg. Corolla lobes valvate Seeds horizontal usually very numerous; stipules entire or bifid; trees or s S eee ee ASA 5. Condamineae hh. Seeds imbricate, vencal, usually few; stipules ee setose- laciniate; subshrubs or herbs. 1. 7. Oldenlandieae aa. Ovules solitary in the cells (subfamily Rubioideae. except Naucleeae). i. de ate ue the radicle superior; shrubs, trees, or woody vines. F eh 3 COSTEA 4. Cinchoneae (q lowers in compact globose heads -.. 8. Naucleeae i Flowers never in globose mens inserted in the throat of the corolla |. 1 9. Guettardeae 10. Chiococceae ii. Seeds ascending, the radicle inferior: plants woody « or herbaceous. l. Corolla lobes contorted in bud: shrubs or tre l. ak rolla lobes valvate in bod. habit various. Ovules basally attached; ead shrubs or trees. lled 14. Dressleriopsideae E ATES 11. Ixoreae 3 < l o e ° nn. Ovary 1-2 celled. o. D l-celled or 2-celled with a thin septum; fruits 1-seed- oe EE P PPI ys Sana E RS 12. ze 00. bene: 2-celled, or the cells rarely more numerous, the septu thick; fruits 2 or more seede p. Stamens usually inserted in the throat of the edid flowers perfec 3. Psychotrieae pp. Stamens usually Modden at the base of the a. tube; flowers often unisexual |... . . I Anthospermeae mm. a lee attached to the Moni “of the cell; habit various. q pules neither foliaceous nor setiferous; trees or large shrubs; un congested by the hypanthium to form a dense head; fruit a syncarp, the base of the hypanthiums united 1. 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 ener. or swollen d. ea s large, 14-1 m long, often pinnatifid-lobed; calycine cup sical o ibo ili glands within -222222222 5. Pentagon dd. Leaves small, less than '4 m n long, not pinnatifid-lobed; : calycine cup w - rows of glands DM RETELEI AAS 79. pM cc. Petioles tumescent, swolle bb. Leaves AA intervenal areas Du lineolate ——Á—— —— n 47. Hippotis e Inflor ee rminal. f Pan drying black: leaves and flowers succulent; inflorescence capitate eS on 77. Schradera £e £e £ ec ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 ff. Plants not drying black: leaves chartaceous, Seea stiffly so; inflo- rescence spreading (except subcapitate in Amphidays g. Stipules with conspicuous fimbriate processes _._..__..._ 5. Amphidaysa gg. Stipules without fimbriate processes. Inflorescences spikelike; shrubs . ei ra ie hh. Inflorescences cymose- -corymbose c or r cymose pan culate; i. Corolla 3 or more cm long; anthers WAREN ane a U tae oe seek ie TX 49. Isertia il. Corolla less than 1.5 cm long; anthers not transversely lo- cellate — UUY I . 69. Raritebe ee. Inflorescences axillar j- Plants pr Pepe fruits bright china blue — O .- 21. Coccocypselum jj. Plants erect or climbing: fruits not bright china blue --------------- "m Sabir cea Leaves alternate 2. HAMELIEAE Ovary 4-5 celled; ro regularly terminal; stamens with anther connectives reg- ularly produced at the apex b. Corolla lobes poder in bud; raphides absent from 1s FEPTEM hà Bertiera bb. Corolla lobes imbricate in bud: raphides di in leaves -------------------- . Hamelia Ovary 2-3 celled; inflorescences always axillary; stamens in the anther pma: not produced at the apex (few species of Hoffmannia excepte c. Ovary 2 celled; calyx with lobes ca. 8 mm long; corolla lobes valvate |... HENCE DTE 87. Xerococcus cc. Ovary 2-3 celled: ee with lobes less than 5 mm: corolla lobes imbricate |... MENS epey RES PME |. 48. Hoffmannia 3. GARDENIEAE Plants armed with spines on stems and short shoots Plants unarme Inflorescences not spikelike; fruits not gourdlike nor on an elongate curved pedicel. c. Flowers perfect. Inflorescences with flowers several to numerous, flowers disposed in a A raa Flow PRESE «669, Randia ra r buds curved at the apex; seeds with testa cells more than E. as long as wide Ll Loss. 66. Posoqueria ee. Flow wer UA not t curved. at the apex: seeds with testa cells less t tw s long as wide |... ee 3. Tocoyena dd. «urn. with pd few or solitary, not disposed in a candelabra array f. Leaves drying black... oss sss. 40. Genipa ; iie not drying black. cc. Flowers unisexu g. Fruits Bui terminal stipules forming a conic Pistillate flowers capitate or cymose; ita: e densely nsa Pew meee meee o . Amaioua Pistillate flowers usually solitary: fruits ‘densely id E E" Duroia gg. Fruits due terminal stipules not forming a conic ca i. Fru Lj by several persistent bracts, a; large as a child's haud. superficially rough - sence eles ee eas il. ieee not e by several persistent bracts, to 2.5 is erficially sm OPEP ea YAlibertia supe bb. Staminate inflorescences s pikelike; fruit 2 a gourdlike capsule c on an n donat € curved pedicel -eers IRR D r Sa DEMNM . Stachyarrhena ATI RNC NER TD Z u u REN Ec AE 57. Manettia Shrubs or trees. b. Stamens with anthers dimorphic or trimorphic bb. Stamens with anthers all alike. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 9 c. Calyx with one lobe expanded as a blade. d. Inflorescence cymose or corymbiform; stamens equal and exserted . Calycophyllum dd. Inflorescence spicate; stamens unequal, not exserted ____ 86. Wittmac kanthus Sa with all lobes similar Plants apdo leaves fleshy, the lateral veins often evanescent. f. Flowers pink or red, the corolla inflate 70. Ravnia ff. iin ice or dis eee g. Seeds winged terminally gg. Seeds with a tuft of ee at o Plants terrestrial; leaves n veins evident to the margin Inflorescences spikelike: stamens with filaments villose, attached at the base of the corolla tube 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 jj Flowers 5-merous; capsule k. Capsule splitting from below toward the apex . REN B Cinchona Capsule splitting from above toward the bas l. rolla gn split at the apex; fruits formine 4 coiled v . Joosia Corolla lobes not lobed at the apex; E not — ud alves. psule loculicidal C [e] 24. wow Hillia oO oO pole : ot fleshy, nat ye papery to coriaceous, lseis hh. guess 13. Bouvardia = = VEM 56. Macrocnemum mm. Capsule septicidal "E 52. Ladenbergia Corolla lobes imbricate or contorted in bud. Stamens conspicuously exserted; Tog not inflated; fruits neither prn dd. ssed nor lenticellate |... . Exostema nn. Stamens not conspicuously exserted; gium inflat ed; fruits penne strongly lewicebon HU NEN 26. Coutarea 5. CONDAMINEAE aa. Leaves not glandular- dpi anthers longitudinally dehiscent. b m lon a Flowers ca. 3 m ; inflorescences axillary ------------------------------ 16. Chimarrhias bb. e more t nee 3 m long: inflorescences terminal. Calycine lobes unequal, one expanded into a large colored limb ..... 65. Pogonopus cc. Calycine lobes equal of nearly so -------------------------------------- 23. Condaminea 6. RONDELETIEAE e Plants woody. b Corolla nus imbricate in the bud. ne lobes unequal, often one expanded into a foliose blade Calyx with foliose blade a eee red color |... Warscewiczia d. Calyx with foliose blade white cc. cu lobes equal or nearl Fruit s large, nutlike, dehiscing septicidally; seeds winged; wood on ex osure to air turning a purple red; filaments geniculate at nun of wack: 78. — u eR ee Or Ke Simira ee. Fruits small, dehiscing loculicidally or septicidally; seeds not winged; w oo n exposure to air not turning red; filaments not geniculate a point of attachment. Calycine lobes often unequal; capsule with valves not woody, not bifid atthe apex. —.. uuu 2 u uu sss ees es ff. = lobes usually equal; capsule with valves woody, bifid a muU M eee >. -» thysa 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 bb. Corolla ie contorted in bud. Trees, terrestrial; corolla iu short, the tube shorter than the lobes; fruits with aie s not coiled at m . Corolla glabrous are 4-merous ____._____---- eee 29. Deppea hh. Corolla villous within, 5-merous _______---_------------------------- 35. Em gg. Shrubs, growing in water or on bank of streams; corolla elongate, the tu least 5 times the length of the lobes, fruits with valves coiled at maturity |... "uM rM 4. Lin a aa. Plants herbaceous _____.______.______------------------------------------------------- 53. nos anis 7. OLDENLANDIEAE a. Plants herbaceous, often prostrate or subprostrate, not ericoid in habit; seeds angular winge - Oldentandia aa. Plants subshrubs, ericoid in habit, usually upright; seeds carinate or plano-convex |... 8. ph ytophyllum 8. NAUCLEEAE Vine with recurved spines and inflorescences axillary and globose 84. Uncaria 9. GUETTARDEAE a. aga armed with rigid spines on twigs and s eee v lobes imbricate in bud; fruits Pa Tm into 2 indehiscent mericarps Sees 55. Mac haonia bb. Corolla lobes valvate in bud; fruits nidi not splitting into 2 indehiscent mericarp .. ]9. Chomelia aa. Plants unarmed. c. C up pup imbricate in bud; leaves with intervenal areas not lineolate-striate. d ts dry, separating into 2 indehiscent mericarps or cocci; flowers not secundl dis osed 55. Machaonia dd. Fruits fleshy; flowers secundly dispose diss covered with a fine feltlike pubescence CERES 43. Guettarda ee. glabrate ntirhea cc. Corolla lobes se in bud: leaves of some species with intervenal areas lineolat te- striate |... 19. Chomelia 10. CHIOCOCCEAE a. Flowers 4-merous; fruits wi nged - 2. Allenanthus aa. Flowers 5-merous; fruits not winged. b. Corolla dew imbricate in bud; fruits not strongly compressed; shrubs (Panama) .... rl S ode he ee eee ee ed 18. Chione bb. Corolla lobes valvate in bud; fruits strongly compressed; trees, shrubs, or wood 17. Chiococca 11. IXOREAE a. Bractlets connate, calyx-like or involucral; inflorescences axillary; cultivated as a tree OD on ee, S s: 22. Coffea aa. Bractlets distinct, not calyx-like; inflorescences terminal; some species cultivated as or- mentals 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 _ EN 37. Faramea 13. PSYCHOTRIEAE a. Plants prostrate, woody herbs; leaves cordate 41. Geophila Plants erect subshrubs, ee or trees (except Declieuxia, a low woody herb [Panama a|); leaves various at the base, not cordate. Lobes of calyx, if a subequal, not polymorphic at the apex. £e e 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 11 c. Inflorescence an His dap e pass sa heads solitary, sessile or pedunculate, or with 3 heads arising from peduncle 15. Cephaelis cc. Inflorescence bracts not ae oe an involucre, flowers not congested into a head. , “beg fruits posar compressed 28. Declieuxia dd. shrubs, or subshrubs, sia epiphytes; fruits terete in cross section. "Seeds with. an incurved ventral surface; stipules pectinate ___________- 4. Rudgea ee m not iin an incurved ventral surface; stipules rarely pectinate, if so, mem f pose lla pes short, to 0.7 cm long (P. chiapensis and _ ostaurea excepted), not swollen at the base; flowers white, green, or yel- low; stipules of twiglets deciduous or persistent . 68. Psychotria ff sips dos ongate, more than 0.7 mm long, ually swollen e: flo owers ii apes orange and purple; sew of twig- let sata pers 64. Palicourea bb. Lobes of calyx u i pasas d ea "n pe 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 Fruits longitudinally soi or separating into open or closed valves or cocci, lacking circumscissile dehisc b. ° e 72. Richardia Cocci € SN with sheath in more than 3 setae; plants not fetid. d. occi indehisce e. lyx Ca aem cocci separating from a central axis or carpophore ____ . Crusea ee. ies alyx dentate; cocci not separating from a central axis or [RR RON 32. Diodia dd Cocci aa t, often only 1 of the 2 opening. occi opening only at the base; Paane axillary .......... 45. Hemidiodia s occi opening at the apex; flowers axillary or terminal. E > uits with cells similar, both dehiscent ---------------- 12. Borreria uits with cells dissimilar, only one opening .... 80. F arie oce cc. Stipules wale pers with only 3 setae; plants fetid 2. Tabagoa 18. RUBIEAE a. Flowers with the hypanthium immediately subtended by an involucre of dms bracts Relbunium aa. Flowers bid the hypanthium not immediately subtended by an involucre of Ter ie bracts . Leaves opposite .. . Didyi maea bb. gem verticillate 39. 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Ju e ur pouoejie 'əqn1 ej[0402 ay) uiojj 991J *3uo| WW ç] “ed sluəuue|U uoure]S “Cp 'seSpu 12urnsip YUM JO uloouus a»gpins JY) *1e[nsdeo 10 ƏWI|Á-uƏq SUNIA ^, pb smjunualy ^c IJU ‘posurm simi ‘pp "Suo[ WW ç uey} a1oui 9qn) e[[O102 Əy} YUM noe “Ip sipA4puti[) ^9] Jejnsdes sjmaj :umnidas ay) 01 A[uerpour paysene *snojouinu sə[nAO ` DIAJOYIAS ^89 speay ajejonjoAur ur jou saouoosaJopur uado ui siƏA0|4 a sijopida `ç| speəu IIBIONIOAU! UI SIOMOL "£p `səuəiÁd ¿ç Jo simuj :poyseye ÁAj[eseq ‘Z SIMAO “Tp `ñuo[ WW c-c Əqn1 e[[0105 ‘aINUIW SIMOJ “LP `ç AÁ[qeureAur səqo| SUIDA]eD juojsisJad uəljo aui 'Suo| ww ç uey} 3aJoui A[[ensn 'e[nsde2 sima Ji :səuəiÁd +—¿ JO 10 2xXr[ÁLI9q simu :291e2uquir JO JIVATLA 10 A[ISIOIISIUIS pa1101uoo e[[010)) * Or vaddaq °6Z səqo| JUNA juajsis1od p YIM *Suo| WU ç o] '1e[nsdeo sliniuj :A[9SO11x9p poijojuoo eooo “Op DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 1980] DIAS] `6f P2]199-9-Z AIBAO :oje[[920| A[ASJOASURI] SIOYIUY `Q/ "Suo| WW Qc-p v[[0102 JY) `Ə8Ie| SiSAO|4 `,9/ snanum) 79] POPIAIPUN saA[eA IYI ‘NLALA Á[[epiondos sima "¿LL DIuJOD[] “SE pyiq Á[o1eurnn saA[eA ay) *a1eA[PA!Q Á[[epror[noO| SUNIA 744 "Suo| WwW ç 0] e[[O102 əy} ‘pews S19^0[4 '9/ '||[9» sod snoiəumu so[nA() “SZ 'sno[npuad jou 3[nAO J0 SƏ|nAO ^,p 2uonj) gl psaylog 6 — sno|npuəd sə[nAO "p, `luəsqe 10 PNIJ Əy} uo [puruii9] dnd xÁ[e°O `,£¿ xəde nay Əy} Buuimosqo pue jin jo £4 Joddn uo Juise ‘pnay ur jusumuoud dno xA[e) 'EL 'Xe| Áj[ensn S120] əy} *uinaqejopueo e se posod -SIp JOU *uLiojruro2eJ 10 uliojiids Á[[euorse2oo *2je1idvo-osoqo[8 “WAOJISAIY} *oje[norued soouoosojopu[ :,0/ DUIKOIO] “ER puanbosoq ‘99 ~~ u3no4 sini] :A[[e11u25 pojuauo səqo| ej[o102 YUM spng "cL EPIRI E uloouis S]InJj ‘SSEW 3LlJu9339 ue se pojuaLio saqo| ej[o102 YIM spng ‘ZZ 'saioads pim SOPIM WU ç 0} *Suo| WD c [-8 əqn1 ej[o102 :21qA s19^0]p :umnqe[apueo e oxi] pasod “AL DAOX[ "(6 ID DI paieAn[no :ƏpIA ww ç'[ 0} 'Suo| WD c'p-('c qN} e[[0102 INYM J0 Pad SIIMOP :21e1ideo-asoquiÁ105 adUadSaIOYU] '[/ ‘1Yysudn siəmoj ay) *e1e1idvo-osoquiÁA105 10 urnaqe[opueo e oxi pesodsip 22uoosoajogu[ `0¿ 'Jeuru.a] sa»uaosa1opgu[ ',p9 DIAJOYIAS °Q9 speəu IIONIOAUI! ui JOU 'soouoosouogur uədo ui SIƏA0|4 `,69 syapyda) $1 speəu Əled3n|oAUI UI SIMOJ 69 `əsn1qo 10 aynoe ‘sde ay) je IPALI jou səqo|l aur A[e;) `,89 SuDUIDJUOJA ^66 xode ay) 1? 3AL3 possouduioo *auios ISVLA je 'səqo| outoAq[e?) ^89 'sai»ads ppm :d[nd suneidaquisip uoos v Aq popunouuns sauaiAd z Jo SUNIA ^,/9 paffO} "cc pa1eAn[no :dno axrxÁ[e5 & OU! ojeuuoo so[oojoeuq IVIL SUNIA 'L9 `HƏ3 dad | sə[nAO `,ç9 piuupufoH “gp Suo| WW 0:'[—ç'0 S9qo| aUIdATeD *,99 $H2202042 X "[8 DPADHIAND) "Cp sisdouo|ssoa(] “EE peces aouadsaqnd JJJ Əuu e UHA paj2402 SUNIA "£9 `(g—)ç—Z s[9due» :aseq ay) 1? 21e»unu A[[euorseooo 10 ISNIQO ‘ANIL SIALIT ',C9 ~~~ g spadues :aseq ay) 1? o1e[noune 10 2]epJOoO SIALIT “7g '"ejeuns-aje[ooui| JOU SVAL FRUIAIAIUT IYI YUM SAARI] `,gç DIaMoy) ^61 xade əy} wouy juopued *[[99 Jad | so[n40 :oje1uop soqo| aUIDATeD ^,[9 D42UMIOS "6L [esaye] ejua5e|[d əy} 'snoiəuinu sə|nAo *snoooei[oJ səqo| aUIDATeED `[9 'Suo| WD çZ—01 ‘ews soAe2T ',09 DIUOSDIUdad ^69 SUC] W 00`[—ç£`0 ‘231e soAe2'] '09 `ƏltuA SUIAIP JOU 'ən[q x1ep JO *xoe[q 'uə318 ‘UMOIG *s[dand ‘aduRI1O 'xəAuo2-oug|d jou *3s0qo[8 sima `,ç6 [Vol. 67 DIIOIOIY) "L| š ƏluA du!Áip Udo ‘oym `xXƏAuo2-oug|d suniy `ç6 `pəuumun sjue|d :21e[nounpod ÁAj[ensn 'ə)elide2-32soqo|3 jou ‘snojnjed sə52uə5səioju *,76 D14D3u/) "vg powe sjuejd :Əəje[nəounpədə `ə1e1ideo2-əsoqo[8 sə52uə5səioju `¿6 `SƏUIA “16 `sqiƏu JO SƏUIA `,6 Dnomuy-p - esses J9joureip ui uio | Surpaooxo jou sjinaj a4njeui :d[nd snorAqo ue INOYUM BUIAIP SpƏƏS YUM SUNIA ^,pg DHOIDUM "p 070700000000 Buoy] ui» | ugy} a1oui əqn1 e[[0102 uN uo juasqe xÁ[e» :3urÁüip uo popqurue A[snorAqo *3uo[qo sunig 5,06 DIAG |] C" 0-7 Suo| ui» | ugy} SS2] əqn1 e[[0102 :1mnuj uo juasaJd xÁ[e2 :3urxuu nou BUIAIP ‘pungos sna “06 'snoooejruo SIALIT ^,/8 biman (p. 30-05-5798 tsa mene aaa Suo| ui» | uey} a1oui əqn1 e[[o405 :3urAip uo YJOOWS ‘Zuo WD 6—ç *punjol simi 7,68 DHOIDUV ^p 7 i iid RED M Suo| ui» | uey} SSI] əqn1 ejoo :3utÁip uo po[qurie *Suo[ ur? g/[70'] 'guo[qo suny “68 'Suo[ WD | 01 10 snonpioap xÁ[e» ay} 'snoJqe[3 syna 7,88 suoddig "Lp w 7 ~~ Juo WD ç o) xA[v» luəlsisiod au) *Aureu Ap[osuop SUNIA “gg "SNOIDBLIOD JO SNOIDBLIODQNS SIALIT 7/8 'SsnoaoeLIO2 0) SNOISeLIBYS ‘ULI 10 uMOJq-poa4 doop ‘UMOIG BUIAIP Áj[ensn saAe2'T ^,68 pdit? 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Rubiaceae) 1980] '2€13s ç utu] Jow UHA yeys ends `f! `€ 19307) CI DIpA4DI21M CL (9)$7t 15505 "CI "S[issiosuinour JOU 119909 JO SƏA|8A paso[» 10 uado o1ui Junesedəs JO jua2siqop ÁApj[eurpniriuo| SUNIA ILI SNdADIDANP °C SISSIOSWNIIID SYN III 19909 ç—Ç JO SIMI “LOLI dTeARAIQ Á][eprornoo] `ip[nsde35 sunig “OL DipADAnog ^el 'Suo| uio | 01 'ə|oiəd ay) 01 2jeupe pue əlgeuuo2o so[ndiis :opra uio g'£-0r| sos] ` 801 ps&pnpnjduy `ç pluupufoH ‘8p "Suo| ui» z uey} BOW 10 snoonpgo aonad ay) jo 334} Sandys :apr Wd /-p SIVIT “ROI 'peuoene Alpeseq 10u ‘RIIAS 0] T SIMAO YUM slup|d `,¿01 DIIOYIASd ^89 naupndisouurg eg so[nAo payosene Ajjeseq z YUM sjueld— “LOI ‘apou e je ç uey) BOW JO ayIsoddo saava] :(paidooxo pviuuvuffo H) yUaysisiod Aj[ensn *snondidsuod sə|ndiiS ',901 (euupupq) WU) səApə| :Ə1ə|osqo saindns 901 'snoooejeuo saA?3| ay} 'oouvuavodde ui proonio jou SURI `,r01 Dixnaj2o(q “BT injudoiso4y ^g nipupjuapy) “TY uimn]asd420220) Uc divsoziyds g jymaj :o4nuo SUIBIEW q^ so[ndijs :snorAqo SUIƏA [EJ9]e| UNA SARA] “SOT a[nsdeo yepiondas v sini :asoia K[[eurieur so[ndijs :juosqe JO snonoidsuoour SUIDA [LIIL] UNA SAARI] “SOI 'snoaoeuooqns SIAL ay) ‘aduvieadde u! prego sjue|d "Pol pcd EE AAP sunay '1uə3səiqe[8 10 snoJqe[8 :(eureueg) apre uro ç:0 Uey? 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'`,001 snowApip Á[3uoi)s sunt `001 DIDUAPIGC “OE "duo| WW p[-c ‘NNUU JO [[BWS SILI] “66 '91epJoo JOU *3]e40 JO '3uo[qo 'a1e[oooue| IIJ SIABIT `,86 ojep1oo SIABIT `Q6 ppiydoayn ` — r 'Suida245 *aje1jsoud sjue|d 16 'Ápoo^ souilmnouios *sq1oH DIJJ2UDJIN "16 ju25saJqej8 0} snouqve[8 saAv3| :1e[nsdeo siinij :poj[32-7 KieA0 pal 01 IYM SIƏA0|[4 `,96 luə5səqnd saAv3| :3]e22eq SUNIY :po[[22-c-C AIPAO INYM SIƏA0|4 96 p2)1qDpç ‘OL 'Suo| WW 0[—çS sima :pƏi 0} 21qA SIIMOY :pi3u SWIS :Ə1E||I914ƏA JOU SIAVIT WINDY) ^6 iununqjosp “VL s19ev1g jo oym g Aq papuaiqns jou wniyUedAY ay) YUM SI9^0[J `,ç6 s]0e1q p JO oym p Aq popuoiqns Ápo1erpourur uiniuluedÁAu əy} YUM SIZMOL “G6 "Suo| uiui p Ugy} SSI] SlIniJ :ƏJUA SIAMOY :Xe2A SWIS IPINIA SIAVIT “PG l 6 [Vol. 67 N MISSOURI BOTANICAL GARDE ANNALS OF THE DOSDGO] “7B 9915s c A[uo YM yeys 1e[ndng "er 23202Dui42d$ “YR guiuado ouo Áquo '1e[ruissip sj[22 YUM SUNIA 7,4 D142440g “TI IUDDSIYSP y0q '1e[nuis S[[o2 YUM SUNIA 'LII *xade ay) 1? 3uiuado 19900 91 DipoipuaaH “Sp ~~ aseq oy) W Á[uo Suruado 19900 `9Q[I `šuiuədo OM) JY) JO UO A[UO uə1Jo “JUdDSIYaP 1990D ^el pipoig `¿ç a10ydodised 10 six? penus e woy 3urjeaedos JOU 12202 ‘ayeJUaP XATeD `,STI D2$sn4) "LC euoudod.eo 10 sixe [enu e woy Zunesedəs 12202 :31eqo| xÁ|eO “SI | "Jusosiqapur 19909 "ell 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 19 1. ALIBERTIA Alibertia A. Rich. in DC., Prodr. 4: 443. 1830. TYPE: A. edulis (L. C. Rich.) A. ich. in Cordiera A. Rich. in DC., Prodr. 4: 445. 1830. TYPE: C. triflora DC. Shrubs or trees. Leaves usually coriaceous, petiolate; stipules connate at the base. Inflorescences terminal, the staminate flowers several, fasciculate, the pis- tillate flowers usually solitary. Flowers dioecious, usually sessile; hypanthium usually globose; calyx short or tubular, truncate or dentate; corolla tube salver- form, stiffly carnose, the lobes contorted, imbricate; stamens 4-8, the anthers linear; ovary 2-8-celled, the ovules few to numerous. Fruits baccate, 2—8-celled, the placentas pulpy, the seeds several, large, compressed. Alibertia is found throughout tropical America and the West Indies. It has about 35 species. a. Leaves with secondary veins 6-15; corolla tube 1-3 cm long at anthesis, 0.5-0.8 cm wide; anthers 10-13 mm long l. A. edulis aa. Leaves with secondary veins ca. 6— zs corolla tube less than 0.8 cm long at ru ca. 0.25 cm wide; anthers ca. 3 mm lon 2. A. garapatica 1. Alibertia edulis (A. Rich.) A. Rich., DC., Prodr. 4: 443. 1830.—Fic. 1. Genipa edulis A. Rich., Actes Soc. Hist ous en 107. 1792. TvPE: not seen. Gardenia edulis (A. Rich. ) Poir. in Lam., éth. Bot. is 2: 708. 1812. Garapatica edulis (A. Rich.) Karst., Fl. Colombia, Í A 1859 Cordiera edulis (A. Rich.) yii Rev. Gen. Pl. 1891. Sabicea edulis (A. Rich.) See Jackson, ie x ew. 772. 1895. Alibertia longistipulata Riley, "Kew Put 1927: 122. TYPE: Panamá, Riley 145 (K, not seen). Alibertia panamensis Riley, Kew Bull. 1927: 123. Type: Panamá, Riley 111 (K). Shrubs or small trees, presumably dioecious, the branchlets terete or often ultimately angular, smooth, glabrous, stiffly ascending. Leaves narrowly oblong, falcately oblong, oblong, ovate or rarely obovate-oblong, 6-20 cm long, 2-8 cm wide, deltoid at the apex, acuminate, the acumen to 0.1 cm long, usually acute, deltoid to obtuse at the base, the costa plane to prominulous above, prominent beneath, to 0.2 cm wide, glabrous, the lateral veins 6—15, arcuate, to 2 cm apart, thin-coriaceous, lustrous, glabrous; petioles 0.5-1.0 cm long, rarely absent, gla- brous; stipules connate or free, deltoid, narrowly deltoid or obovate-oblong, ca. 1.5(-2.0) cm long, 0.4—0.6 cm wide, attenuate-acuminate, the acumen often longer than the corpus, scarious, striate-venose, glabrous. /nflorescences of 1 to several terminal flowers subtended by broadly ovate bracts shorter than or exceeding the hypanthium in length, the buds with the apex of the corolla acute and dilated. Flowers sessile, unisexual, the pistillate flowers solitary, the staminate flowers in a cluster of about 8; hypanthium and calycine cup ca. 0.5 cm long, carnose, glabrous, the teeth 5(—4), deltoid to linear, to 1.8 mm long, acute to obtuse; corolla white, the tube 1—3 cm long, glabrous or villosulose outside, subcoria- ceous, the lobes 4-5, imbricate, oblong, to 2 cm long, often exceeding the tube; stamens usually (4—)5(-8), the anthers narrowly oblong, 5-13 mm long, ca. 13 mm long in male flowers, ca. 5 mm long in female flowers, 0.8—1.0 mm wide, often minutely apiculate, dorsifixed, the filaments short, ca. 0.5 cm long; ovarian 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Figure 1. Alibertia edulis (A. Rich.) A. Rich.—A. Flowering branch (x!5). [After Hladik 49).]—B. Flower buds (x!4). [After Lewis et al. 1866.]—C. Fruit (x). [After Lao & Holdridge 244.) disc doughnut-shaped, occasionally scalloped, 1.0-1.8 mm long, 0.75-1.50 mm wide, the style expanding toward the apex, the stigmas 3—4, crassate, oblong, to 1.8 mm long, the ovary 4-celled. Fruits terminal, sessile, solitary, rotund to com- pressed-rotund, to 3 cm in diam., the persistent calyx coroniform and cylindrical, to 0.5 cm long, rarely to 1 cm long, ligneous, smooth, glabrous; seeds numerous in a slimy pulp, suborbicular, ca. 0.5 cm in diam., striate. Alibertia edulis is one of the most common shrubs in Panama. It ranges from Mexico to the Amazon Basin, although Steyermark does not include it in his Flora de Venezuela. It occurs in the West Indies. The fruits are scarcely edible. The names ‘‘Trompo”’ and ‘‘Trompito” refer to the fact that tops are made from the fruits. ‘‘Largartillo’’; *‘Madrono’’; ‘‘Madroño de Comer”; ‘‘Trompo’’; ""Trompito"'; ‘Wild Guava.” 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 21 BOCAS DEL TORO: Bocas, Lazor et al. 2415 (MO). Columbus Island, Wedel 32 (MO). Water Valley, Wedel 798 (MO). Chiriquí Lagoon, Wedel 1311 (MO). Old Bank Island, Wedel 2009 (MO). Isla bie Wedel 2474 (MO). Water Valley, Wedel 2672 (MO). Johns Creek, Wedel 2773 (MO). CANAL ZONE: Juan Mina, Bartlett & Lasser 1552 (MO). Cruces Trail and Madden Dam Road, ea & sein 16342 (MO). Juan Mina, Bartlett & Lasser 16563 (MO). nri Croat 15145 (MO). Pan american Highway near La Chorrera, D'Arcy 9439 (MO). Fort Kobbe, Duke 3841, 4201, p Jal MO). Albrook, Dwyer 6746 (MO); Dwyer & Robyns 119 (MO). Empire, Hayden 58 (MO). Sherman, Hayden 97, 107 (both MO). 6 km W of Gamboa, Nee 7208 (MO). Road C-29, 6 km : ge Gamboa, Nee 8970 (MO). Howard Air Force Base, Oliver & MacBryde 1894 (GH, MO, US). Balboa, Standley 26050 (MO). Farfan, Stern & Chambers 153 (MO). Río Cocolí, Road K-9, Stern et al. 38 . Río Indio, Steyermark & Allen 17439 (MO). Curundú, Tyson 1316 (MO). Chiva-Chiva Trail, Tarah 1357 (MO). Miraflores, White 129, 138 (both MO). Cocolí, Wilbur et al. 12864 (MO). BARRO co ISLAND: Aviles 23 (MO): Croat 4226, 5167, 5358, 5574, 5724, 6006, 6128, 6142, 6328, 6429, 6878, 7390, 7454, 9249, 9512, 15263, 15397 (all MO); Dwyer 1434, 1438, 8467 (all MO): Ebinger 261 (MO); Hayden 3 igre Hladik 49 (MO); Shattuck 147, 643 (both F. MO), 896, 967 (both MO); Wetmore & Abbe 78 (MO), 78 A (F, MO); Woodson & Schery 977 (MO); Zetek 4295 (F, MO). CHIRIQUÍ: Boquete, p 829 (MO); Dwyer & Hayden 7625 (COL, MO, UC, VEN); Kirkbride 93 e Stern et al. 1200 (MO). Puerto Armuelles, Woodson & Schery 847 (MO). COCLÉ: Penonomé, 12 (MO El Valle, Lee 12 (MO). COLON: Salud, Lao & Holdridge 244 (MO). Nuevo Chagres, Lewis et al. 1866 (GH, MO). DARIEN: Mouth of Rio Yapé, Allen 340 (MO). Quebrada Venado and Peje Swamp, Bristan 1048 (MO). Rio Canclones and Rio Chucunaque, Duke 5/18 (MO). Cerro Pirri, Duke 6584 (MO). Perrecenico River, Duke & Bristan 236 (MO). Rio Tuira, Duke & Kirkbride 14056 (MO). Yaviza, Gentry & Mori 13504 (MO). Between Cana and Boca de Cupé, Stern et al. 613 (MO). Los SANTOS: 1-2 mi W of Candelaria, Duke A-12433 (MO). Los Asientos, Wandehake 31 (MO). PANAMA: Nuevo Emperador, Carrera 1 (MO). Cerro Campana, Castillo 22 (MO). Río Pita, Duke 4770 (MO). El Llano, Duke 5520, 5824 (both MO). Isla del Rey, Duke 9518, 10411 (both MO). Coronado Beach, Duke 11817 (MO). Isla de La Bayoneta, Dwyer 1736 (MO). Río Pasiga, Gentry 2277 (MO). San José Island, Johnston 486 (MO). Chimán, Lewis et al. 3314 (MO, UC, VEN). Chorrera, Rodriguez 51 (MO). Bella Vista, Standley 25354 (MO). Isla de Espiritu Santo, Tyson 5562 (MO). 5 mi W of Chepo, Tyson 6725 (MO). Pacora, Woodson et al. 743 (MO). SAN BLAS: Eslogandí, Duke 10196 (MO). VERAGUAS: Río Canazas headwaters, Allen 188 (F, MO). Soná, Allen 1073 (MO). 5 mi N of Santiago, Blum & Tyson 622 (MO). La Mesa, Blum & Tyson 640 (MO). 32 mi W of Santiago, Croat 10732 (MO). San Francisco, Dwyer d (MO). Mouth of Río Concepción, Lewis et al. 2821 (COL, DUKE MICH, MO, UC). Laguna La Yeguda, Luteyn 1460 (MO). Quebrada just S of Santa Fé, Nee 8035 (MO). 5 mi E of Santiago, A. et al. 4272 (M 2. Alibertia garapatica (Karst.) Schum. in Mart., Fl. Bras. 6(6): 384. 1889. Garapatica edulis Karst., Fl. Colomb. 1: 57. 1858, not A. edulis A. Rich. TYPE: not seen. Shrubs or small trees, to 4 m tall, the twiglets slender, often puberulent, the nodes usually swollen, the leaf scars large. Leaves oblong or oblong rotund, 6- 15 cm long, 2.3-5.0(—6.0) cm wide, rounded or tapering widely into an obtuse or deltoid apex, acuminate, the acumen 0.5-2.0 cm long, obtuse to cuneate at the base, the lateral veins 6-10, thickly chartaceous to subcoriaceous, smooth, gla- brous to sparsely golden-puberulent; petioles stiff, 0.5—1.0 cm long; stipules free?, triangular, to 3 mm long, acute, puberulent, usually deciduous. Inflorescences terminal, sessile or subsessile, the flowers of the female inflorescence ca. 5, oc- casionally solitary?, those of the male 7-10, radially disposed. Flowers (only male seen) ca. 1 cm long, the buds acute, the hypanthium ca. 1.5 mm long, the calycine teeth absent; corolla white, the tube cylindrical, to 8 mm long, to 2.5 mm wide, stiffly carnose, puberulent, the lobes 5, triangular, to 2.5 mm long; stamens 4(5?), narrowly oblong, ca. 3 mm long. Fruits subsessile or sessile, solitary, rotund or subrotund, occasionally obovate-rotund, to 2.5 cm long, slight- ly wider than long, drying black, minutely rugulose, glabrous, the persistent ca- lycine cup crateriform-cylindrical, to 3 mm long. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Alibertia garapatica is known from Mexico, Panama and Colombia. The species should be studied in the field as more than one species may be involved. Stern et al. 1719 shows 4 very immature fruits in one fructescence terminating a twiglet, while another twiglet shows 4 flowers in bud which measure about 1.2 cm in length and are not as acute as typical buds of the species. One flower dissected proved to be male with 4 subfacate sessile stamens and a fifth stamen which was abnormally plump. The 4 normal stamens measured 0.5 mm in width, with an acumen to 0.8 mm long. The abnormal stamen measured 0.8 mm in diameter. The unopened anther sacs of the normal stamens were linear and the connective laminar as one sees in the stamens of the Himantandraceae. Allen 1616 showed a style 6.2 mm long, which was dilated basally and minutely bifid at the apex; no ovary was evident. Dressler & Williams 3969 has leaf blades only 4.0-6.5 cm long. Several leaves of Liesner 334 are almost rotund, being 7.0-7.5 cm long, 5-6 cm wide. Croat 39706 (MO) from Mexico (Oaxaca) is A. garapatica, even though it seems out of range. CHIRIQUÍ: Quebrada Punta de Piedra, 0-100 m, Liesner 334 (MO). cocLE: Aguadulce, McDaniel & Cooke 14762 (MO). COLON: Santa Rita Ridge, E of Transisthmian Highway, 300-500 m, Gentry 6589 (MO). Santa ee Road, Dressler & Williams 2969 (MO). Santa Rita Road, Mori & Kallunki 2787 (MO). M : Ocú, limestone area, Stern et al. 33583, 33593 (both MO). PANAMA: Río Las Lajas, Allen 6( p. Villa Rosario, International Highway, Saldana 21 (MO). Guayabito, road to Laguna, a. — Smith 3463 (MO). VERAGUAS: Isla de Coiba, Dwyer 1629 (MO). 2. ALLENANTHUS Allenanthus Standley, Ann. Missouri Bot. Gard. 27: 344. 1940. TYPE: A. eryth- rocarpus Standley. Trees. Leaves ovate elliptic, acuminate, the lateral veins few; petioles present; stipules free, ovate or rotund, cuspidate. /nflorescences terminal, paniculate, elongate pedunculate; bracts and bracteoles small. Flowers with calycine lobes 4; anthers short. Fruits compressed, drying red, the central seminiferous part surrounded by a thick wing. Allenanthus is found in Costa Rica and Panama. It is among the easiest of the Rubiaceae to recognize as the fruit has the appearance of the achene of Rumex of the Polygonaceae. 1. Allenanthus erythrocarpus Standley, Ann. Missouri Bot. Gard. 27: 344. 1940. TYPE: Panama, Allen 1999 (MO, holotype).—Fic. 2. Chimarrhis decurrens Steyermark, Ceiba 3: 18-19. 1952. Type: Allen 3566 (F, holotype). Trees 10(—20) m tall, the branchlets smooth, terete, rubescent, ultimately plano-compressed, glabrate, the nodes usually 4-5 cm apart. Leaves ovate ellip- tic, 7-11 cm long, 3.8-5.0 cm wide, deltoid at the apex, acuminate, the acumen to 1.5 cm long, obtuse and scarcely inequilateral at the base, the costa promi- nulous above, prominent beneath, the lateral veins ca. 7, arcuate, usually with one subparallel vein between 2 lateral veins, the intervenal areas mosaic-like, chartaceous, scarcely discolorous, glabrous except puberulent on the costa and the veins beneath; petioles slender, to 1.5 cm long; stipules deciduous, free, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 23 FiG SURE 2. Allenanthus erythrocarpus Standley.—A. Branch with leaves (x.75).—B. Inflores- cence (x.75). [After Duke 5654.] widely ovate or compressed rotund, 3-6 mm long, abruptly cuspidate, the cusp as long as the body. /nflorescences terminal, solitary, paniculate, to 15 cm long, to 10 cm wide, rounded terminally, the peduncles terete, 4.0—6.5 cm long, pu- berulent, the branches opposite, the lowermost to 9 cm long, arcuate or strict, strongly ascending, pubescent, the hairs often present only on part of the pedun- cle and branches; bracts small, linear-subulate. Flowers with pedicels 1-5 mm long; hypanthium turbinate, glabrous, to 2 mm long; calycine lobes 4, prominent, 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 ovate to rotund, 0.5-0.9 mm long, glabrous; corolla tube to 4 mm long, pubescent outside, sparsely pilose within, the lobes oblong, ca. 1.5 mm long, almost as wide, rounded apically; anthers oblong, ca. | mm long, the filaments slightly shorter than the anthers. Fruits strongly compressed, ca. 8 mm long, to 6 mm wide, glabrous, appearing winged, drying red, the central seminiferous part to 2.5 mm long, 3-nerved, surrounded by a thick wing. Allenanthus erythrocarpus is known only from Panama and Costa Rica. cocLéÉ: El Valle de Anton, Allen 1999, 2713 (both MO), 3566 (F). PANAMA: Between El Llano and Río Mamont, Duke 5654 (MO). 3. ALSEIS Alseis Schott in Spreng., Syst. Veg. 4(2): 4-4. 1827. TYPE: A. floribunda Schott. Trees or shrubs. Stipules free, subulate. /nflorescences axillary and terminal, spiciform cylindrical, patulous. Flowers protogynous, the hypanthium obconical, the calyx 5-lobed; corolla urceolate, inflated; stamens 5, exserted, the filaments inserted at the base of the corolla; style recurved, the ovary 2-loculate, the septum thin, the ovules numerous. Fruits capsular, narrowly oblong, lignose, dehiscing from apex to base, 2-locular, the seeds many, narrowly fusiform. Alseis has about 20 species ranging from Mexico through Central America to Brazil and Peru. a. Leaves with lateral veins ca. 25; fruits before dehiscence ca. 0.2 cm wide |... 1. A. blackiana aa. Leaves with lateral veins ca. 9; fruits before dehiscence 0.4 cm wide _______- 2. P darienensis 1. Alseis blackiana Hemsl., Diag. Pl. Nov. 30. 1879. Type: Panama, Hayes 665 (K).— Trees to 75 ft tall; twigs terete, ultimately angular, the nodes well spaced or crowded, glabrous. Leaves obovate oblong, to 36 cm long, to 9.5 cm wide, cu- neate at the apex, often cuspidate, the cusp to 1 cm long, cuneate or rarely truncate at the base, equilateral, rarely subinequilateral, the costa prominulous above, prominent beneath, diffusely villose above, the hairs weak, these short and long, occasionally aggregated in the axils, the secondary veins ca. 25, the distal veins widely arcuate, the proximal veins perpendicular, chartaceous, usu- ally scarcely discolorous, glabrous except on the costa and the veins; petioles rigid, to 3 cm long, to 0.2 cm wide, angular, the blade decurrent on the upper part of petiole; stipules deciduous, subulate, to 1.5 cm long, ca. 0.12 cm wide, scarious, glabrous. Inflorescences usually several in the axils of the terminal leaves, spiciform-cylindrical, ascending, the peduncles unbranched, ascending, often falcate, shorter than or exceeding the leaves, to 12 cm long, to 0.3 cm wide, lignose, puberulent, usually lenticellate. Flowers with pedicels slender, pubes- cent, to 0.2 cm long, the bracts basal, linear, about as long as the pedicel; hy- panthium narrowly cuneate cylindrical, ca. 1.2 mm long and indistinguishable from the pedicel, minutely puberulent, the calycine tube obsolete, the teeth 5 erect, or spreading, oblong deltoid, subequal, to 0.5 mm long, obtuse; corolla white, or pale yellow, inflated, urceolate, ca. 2.2 mm long, often slightly con- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 25 TU ye SE » 477 net Ç RC ww A VN 3. Alseis blackiana Hemsl.—A. Flowering branch (x12). [After Croat 14824.|—B. FiGuRE 3. Fruiting branch (x!'2). [After Croat 15560.] stricted medially, densely white-villose within, glabrate outside, petaloid, the orifice wide, the lobes 5, much reduced, inflexed slightly, oblong, to 0.5 mm long, obtuse: stamens 5, exserted, the anthers oblong, ca. 1 mm long, versatile, the filaments subequal, ribbonlike, ca. 6 mm long, densely villose ca. 1 mm above the base, attached at base of tube; ovarian disc prominent, mound-shaped, 2-3- lobed, 0.3-0.6 mm long, the style slender, ca. 3 mm long, expanded apically, the stigmas 2, coiled, ca. 1.3 mm long, glabrous, the ovary thin walled, the septum thin, the placentas intrusive, thick, the ovules numerous. Fruits with pedicels to 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 0.2 cm long, before dehiscence narrow-oblong, 1.2 cm long, ca. 0.2 cm wide, often falcate, penis-like or operculoid at apex, attenuate at base, lignose, drying brown, delicately longitudinally veined, puberulent and farinose, dehiscing from apex to base, the valves 2, drying lustrous yellow within; seeds numerous. This species occurs only in Panama and Colombia. It fruits readily but despite numerous collections from Panama, it has been rarely collected in flower. In his original description Hemsley says that the stamens of the lowermost flowers are long-exserted, with those of the upper flowers included. This is not the case in the few flowering specimens examined. Croat 14829 notes that protandry occurs in Alseis blackiana. CANAL ZONE: Salamanca Hydrographic Station, Río Pequení, Dodge et al. 17981 (MO). de Road, 4 km NW of Gamboa, Nee 7539 (MO). Road K-9, Stern et al. 39 (MO). BARRO CO ISLAND: Croat 5389, 6492, 8323, 9134, 9448, 9503, 10756, 14824, 14829, 16206, 15560, 16196, 16208, . (all MO); Dwyer 1439 (MO); Foster 863 (MO); Hayden 90 (MO); 1029 (GH. MO, US); Hladik 5 (MO); Shattuck 52, 822 (both MO); Wilbur & Weaver 10792 (MO); Zetek 5086, 5600 (both MO). DARIÉN: Río Tuquesa below Quebrada ir an 1087 (MO). Río Ucurgantí, Bristan 1168 I e CC Duke 8825 (MO). Río Pucro, below Pucro, Dd. or (MO). Paya, vic Río Paya, Stern et al. 187 (M O). El Real, Río Tuira, Stern et al. 284 (MO). Campamento Buena Vista, Río Chucunaque, Stern et al. 848 (MO). Cerro Piriaque, Tyson et al. 3856 (MO) 2. Alseis darienensis Dwyer.? TYPE: Panama, Mori 7001 (MO, holotype). Trees, much branching, the twiglets terete, the bark thin, scaling off, rubes- cent, glabrous, the nodes moderately crowded on new growth. Leaves ascending, oblong or lanceolate, 13 cm long, 2.0-5.5 cm wide, deltoid or acute at the apex, acuminate, the acumen widely triangular, to 1 cm long, ultimately obtuse, cuneate or acute at the base, moderately inequilateral, the costa slender, drying reddish above, somewhat prominulous beneath, the lateral veins ca. 9, arcuate, disap- pearing near the margin, not forming a conspicuous submarginal vein, with sev- eral smaller veins between a pair of laterals, these leaving the costa at right angles, soon disappearing, the smallest veins forming a reticulate mosaic beneath, the margin delicately callose, thin chartaceous, concolorous, beneath appressed or spreading hirsute on the costa, the hairs at right angles and lateral to the costa in and near the vein axils; petioles wiry, stiff, to 45 cm long, ca. 0.75 mm wide; stipules triangular or ovate triangular, to 7 mm long, acute, longitudinally striate, scarious, dan red, deg Inflorescences terminal and axillary, cymose spi- cate, to m long, to 0.1 cm wide, rubescent, glabrous. Flowers (calyx only) with the penat cup ht the lobes 5, imbricate, oblong rotund, ca. 1 mm long, about as long as wide, obtuse, venose, puberulent marginally. Fruits sessile, narrowly oblong (immature), to 1.2 cm long, to 0.4 cm wide, somewhat com- pressed, covered with a grey indumentum; seeds membranous, drying yellow, fusiform, to 5.5 mm long. * Alseis darienensis Dwyer, spec. nov. Arbores ramulis glabris. Folia oblonga vel lanceolata, 4- 13 cm longa, 2.0-5.5 cm lata, venis lateralibus ca. 9, tenui-chartacea subtus in costa hirsuta; petiolis ad 45 cm longis; stipulis triangularibus vel ovato- triangularibus, ad 7 mm longis, acutis glabris. /nflo- rescentiae terminales vel axillares, cymoso-spicatae, ad 11 cm longae, glabrae. Flores lobis calycis 5 imbricatis oblongo-rotundis, ca. | mm longis. obtusis. Fructus angusto-oblongi, ad 1.2 cm longi, conspicue griseo-puberuli. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 27 Alseis darienensis is known only from Panama. The plant bears little resem- blance to any Rubiaceae I have encountered in preparing the family for the Flora of Panama. But because of the cymose-spikelike inflorescence I have elected to place it in Alseis, but its placement is quite uncertain. DARIEN: Rio Tuquesa, middle Tuquesa Mining Co. camp called Charco Peje, 250 m, Mori 7001 O). 4. AMAIOUA Amaioua Aubl., Hist. Pl. Guiane 13. 1775. TYPE: A. guianensis Aubl. Trees or shrubs. Leaves usually crowded at the apex of the twiglets; stipules interpetiolar, connate, often irregularly cleft, finally caducous. Inflorescences terminal or with short lateral axes, eee capitate or corymbose, the pe- duncle present or absent. Flowers dioecious, 5—6-merous, the calycine cup tu- bular, the teeth reduced or conspicuous; nde cylindrical, sericeous on the outside, the lobes contorted; stamens included (male flowers), the anthers sessile, inserted near the middle or on the lower half of the tube; female flowers with the ovary 2-celled, the ovules numerous in each cell, horizontally and biseriately arranged. Fruits baccate, 2-celled; seeds numerous, horizontally disposed, tri- angular and rounded marginally, the testa fibrous. Amaioua occurs in Central and South America. There are about 25 species in the genus. a. Flowers sessile or subsessile. b. Fruits oblong rotund, to 1.5 cm long; leaves with 6-7 lateral veins _____ l. A. corymbosa bb. Fruits rotund, 1.8-2.2 cm long; leaves with ca. 14 lateral veins ________ 2. A. magnicarpa aa. Flowers pedicellate, the pedicels to 1.5 cm long 3. A. pedicellata 1. Amaioua corymbosa H.B.K., Nov. Gen. Sp. 3: 419. 1820. Type: Colombia, Humboldt & Bonpland (B, not seen, photo MO).—Fic. 4. iae hexandra = ex R. & S., Syst. Veg. 5: 243. 1819. rype: America Merid., Humboldt & land, not s Perlen. E D. Mem. Mus. Hist. Nat. 6: 14. 1820. type: French Guiana, Martin (C, not seen ijoan. ee Hemsl., Biol. Centr. Amer., Bot. 2: 39. 1881, not A. guianensis Aubl. 1775. Nomen nudum. Shrubs or trees to 40 ft tall, the branches terete, the nodes well spaced; branchlets angular, the leaves often crowded at the apex. Leaves oblong, often widely so, the large blades about twice as long as wide, 9-23 cm long, 2.5-12.5 cm wide, obtuse at the apex, shortly acuminate, the acumen to 1.2 cm long, basally inequilateral or equilateral, truncate, rounded, cuneate or acute, the costa plane to prominulous above, often appressed villose proximally, appressed villose beneath, the principal secondary veins 6-7, arcuate, prominulous above, prom- inent beneath, subcoriaceous, usually concolorous, lightly appressed villose on costa and secondary veins, often sparsely pubescent on margins, the intervenal area glabrous; petioles to 1.5 cm long, stout, to 0.3 cm wide, often arcuate; stipules oblanceolate, 1.5-2.5 cm long, ferrugineous-sericeous outside, glabrous and glandular within. Inflorescences terminal, few-flowered cymes, the peduncles 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 IGURE 4. Amaioua corymbosa H.B.K.—A. Flowering branch (x'2).—B. Opened flower (x2.5). [After Croat 16214.] arising in a fan, each usually unbranched, slender, stiff, angular, puberulent. Flowers staminate, pistillate or bisexual (functionally so?), the staminate flowers slightly larger, sessile or on pedicels shorter than the hypanthium, the buds ru- gous, turgid below the middle, tapering to an obtuse apex; hypanthium com- pressed cylindrical ca. 1 mm long, the calycine cup to 5 mm long, to 2.5 mm wide, carnose, puberulent outside, glabrous to puberulent and eglandular within, truncate, the calycine teeth 5, subulate, to 1 mm long, obtuse; corolla white, the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 29 tube cylindrical, somewhat inflated, 5-7 mm long, 1.5 mm wide, carnose, densely white villose outside, glabrous within, the lobes 5, lanceolate, 5-7 mm long, glabrous within, minutely white-puberulent adaxially; stamens 5-6, the anthers linear oblong, 4-4.5 mm long, distinctly petaloid rostrate for 1 mm, the filaments very short, inserted near middle of tube; ovarian disc doughnut shaped, 1-2 mm long, 0.9-1.8 mm wide, 0.5 mm high, the style slender, 4-9 mm long, the stigmas 2. subulate, 0.3-0.6 mm long. Fruits compactly clustered, sessile, oblong rotund, to 1.5 cm long, pruniform, marcescent, sparsely sericeous, often the vestige of the calyx forming a conspicuous annular scar; seeds compressed, angulate, ca. 2 mm long, shiny red, the surface with a series of sinuous, densely packed some- what parallel ridges. Amaioua corymbosa ranges from Mexico to northern South America. It also occurs in Cuba. The flowers should be studied critically under field conditions. Plants have been described as dioecious, but some dissected flowers have func- tional anthers, a well-defined style and a pair of stigmas. The ovary is not apparent in these flowers. One collection, Croat 11532, has fruit more oblong in shape and does not have the appearance of a small dried prune so typical of A. corymbosa. The pericarp is smoother than the typical. Croat noted on the collection that the fruits were old, which may account for the difference in appearance. CANAL ZONE: Fort Kobbe, near beach, Duke 4719, 4724 (both MO). Albrook, Dwyer & Robyns 43 (MO). Headquarters J.W.T.C., Johnston 1574 (MO). W of Gamboa, Nee 6772 (MO). BARRO MO). DARIÉN: Pico Piriaque, Duke 8137 (MO). Piñas, Duke 10598 (MO). Baia de Pinas, Stern & pee tu 192 (MO). Mouth of Rio Lara, Tyson et al. 3864 (MO). Piriaque Camp at River, Tyson et al. 4741, 4746, 4754 (all MO). PANAMA: Taboga Island, Allen 134 (BM, MO). Cerro Azul to Cerro Jefe, Austin 24 (MO); Blum et al. 18 (MO); Croat 11532 (MO). Panama Viejo, Duke 5736 ( ). San José Island, Duke 12503 (MO). Cerro Azul to Cerro Jefe, Dwyer 1487, 1501, 1871, yas g (all MO), 8041 (GH, MO); Dwyer 23 Gentry 10243 (MO); Guerra 5 (MO); Lewis et al. 268 K, MO, ; Porter et al. 4079 (MO, SCZ, US); Reece 5 (MO). Villa Rosario, Internatio ver Highway, Saldana 21 (MO). San José a Stimson 5328 (MO). Cerro Azul to Cerro Jefe, Tyson 2089 (MO): Tyson et al. 4061, 4314, 5381 (all MO). vERAGUAS: La Yeguada, 600 m, Correa 2217 (MO). Isla de Coiba, Dwyer 1629 (MO). 2. Amaioua magnicarpa Dwyer.* TYPE: Panama, Dwyer 9552 (MO). Trees 6—15 m tall, the branchlets ultimately angular, the older bark cracking, ashen grey when dry, glabrous, the nodes somewhat swollen, moderately crowd- ed. Leaves oblong lanceolate or oblong rotund, 11.5-17 cm long, 5-9 cm wide, deltoid to obtuse at the apex, acuminate, the acumen short, 0.4—1 cm long, obtuse or acute, often falcate, basally cuneate obtuse, scarcely inequilateral, the costa slender above, prominent beneath, the lateral veins ca. 13, at first strict, then arcuate toward the margins, the secondary veins perpendicular, subcoriaceous, concolorous, glabrous except appressed-white ciliate on the costa and veins, the minute raphides often abundant above, the margin lightly revolute; petioles to 1.5 * Amaioua magnicarpa — spec. nov. Arbores 6-15 m altae. Folia oblongo-lanceolata, vel oblongo- eae 11.5-17 cm longa, 5-9 cm lata, venis lateralibus ca. 13 subcoriacea glabra praeter costam venasque appresso- albido-iiatas petiolis ad 1.5 cm longis. Fructus rubri et cerasini rotundi, 1.8-2.2 cm longi, apice saepe umbona 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cm long, to 2.2 mm wide, rigid, densely appressed villose. Flowers not seen. Fruits red, cherry-like, rotund, 1.8-2.2 cm long, obtuse at the apex and base, often umbonate at the apex, puberulent, with the scars of the floral parts scarcely elevated, ca. 0.5 cm in diam. The new species is known only from Panama. It is distinguished from the relatively common A. corymbosa (Willd.) H.B.K. by having much larger fruit which resemble cultivated commercial cherries. COLON: Santa Rita Ridge, ca. 2000 ft, Dwyer 9552 (MO). 9.4 km N of Goofy Lake, Folsom et al. 1976 (MO). PANAMA: El Llano-Carti 10-12 km from junction Inter-American Highway, Mori & Kallunki 2880 (MO). El Llano-Carti Road, 1.7 km from Inter-American Highway, Mori, Gentry & Kallunki 4691 (MO). 3. Amaioua pedicellata Dwyer.? TYPE: Panama, Croat 25937 (MO, holotype; NY, isotype). Trees 5-10 m tall, to 22 d.b.h., the branchlets terete, glabrous, the bark dis- colorous, the nodes well spaced. Leaves ovate-oblong, 6-19 cm long, 2.3-10 cm wide, widely deltoid toward the apex, attenuate-acuminate, the acumen to 1 cm long, falcate, ultimately acute, basally acute or cuneate, rarely obtuse, the costa prominulous above, prominent beneath, the lateral veins 12-14, rigid, especially prominent beneath, subcoriaceous, discolorous, above dark brown, yellow brown beneath, golden pilose with the veinlets pinnately disposed; petioles rigid, woody, to 1.5 cm long, the hairs golden, spreading or appressed; stipules not seen. Zn- florescences terminal (in fruit) radially umbelliform, to 5 cm long, the peduncle absent, the pedicels to 1.5 cm long, rigid, puberulent. Flowers not seen except for a fragment of the calyx; calycine cup cylindrical, to 5 mm long, ca. 4 mm wide, truncate, ciliate without and within, the hairs golden, appressed, the teeth 5, triangular-subulate, to 0.3 mm long, these arising from a thin keel on the body of the cup. Fruits ovate-oblong or oblong-rotund, to 1.5 cm long, black when dry, purple brown, lightly puberulent. Amaioua pedicellata is known only from Panama. It is readily distinguished by the long-pedicellate fruits and presumably flowers disposed in the form of an umbel. : Lumber camp at Alto Calvaro, 7 km N of El Cope, 900 m, Folsom 1290 (MO). VERAGUAS: Road cen araja Agricola Alto Piedra and Río Dos Bocas, 730-770 m, Croat 25937 (MO). NW of Santa Fé, 8.8 km from Escuela Agricola Alio Piedra, Mori et al. 4001 (MO). 5. AMPHIDAYSA Amphidaysa Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 180. 1936. TYPE: Sabicea ambigua Standley = Amphidaysa ambigua (Standl.) Standl. 5 Amaioua pedicellata Dwyer, spec. nov. Arbores 5-10 m altae, ramulis E Folia ovato- dép. 6-19 cm longa, 2.3-10 cm lata, apicem versus lato-deltoidea attenuato-acuminata acumine ad 1 cm longo, subcoriacea venis lateralibus 12-14; petiolis rigidis, ad 1.5 cm longis. Inflorescentiae ae: radiate umbelliformes ad 5 cm longae, pedunculo nullo pedicellis ad 1.5 cm longis. Flores cupula calycis cylindrica, ad 5 mm longa, ca. 4 mm lata, truncata dentibus ad 0.3 mm longis. Fructus ovato-oblongi vel oblongo-rotundi, ad 1.5 cm longi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 31 Pittierothamnus Steyermark Bol. Soc. Venez. Cienc. Nat. 23 (101): 92. 1962. TYPE: Stevermark 86323 (holotype, VEN), 8936/ (paratype, VEN), 89763 (paratype, VEN). Herbs or subshrubs, the stems unbranched. Leaves often crowded at the apices of the stems, petiolate, the stipules interpetiolar, large, entire or more often the margin setose pectinate, the processes numerous, unequal. /nflores- cences axillary or terminal, capitate to trichotomously cymose, the bracts entire to multifid, often obscuring the flowers. Flowers with the hypanthium oblong or turbinate, the calycine cup short, the lobes 5, unequal, somewhat foliaceous, persistent; corolla tubular, 5-lobed, the lobes valvate; stamens 5, inserted in the middle of the tube, the anthers linear, dorsifixed; ovary 2-celled, the ovules nu- merous in each locule, the placentas axile, adnate to the axis. Fruits fleshy, indehiscent, the persistent calyx conspicuous. Amphidaysa is found in Panama, Colombia, Venezuela, and Brazil. The genus includes 6-7 species. Literature: Steyermark, J. 1972. Amphidaysa, in The Botany of the Guayana Highland Mem. New York Bot. Gard. 23: 316-322. a. Lobes of the calyx 5-6, 1.2-1.5 mm wide —_ .. - . A. ambigua aa. Lobes of the calyx 4, 2-4 mm wide I REEL! 7 spathulata 1. Amphidaysa a (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 181. 1931.—Fic. Sabicea ambigua Standley, Field Mus. Nat. Hist., Bot. Ser. 7: 49. 1930. rvPE: Colombia, Triana 1847, not seen. Subshrubs to ?4 m tall, the stem occasionally subprostrate or upright, lignose, often rough with petiole scars, glabrous, the leaves aggregated terminally. Leaves oblong, elliptic or occasionally obovate, 7-24 cm long, 2.2-8 cm wide, occasion- ally subfalcate, deltoid or rounded at the apex, acuminate, the acumen short or to | cm long, cuneate at the base, often slightly inequilateral, the costa grooved, plane or prominulous above, prominulous beneath, 1.2-2.5 mm wide proximally, the lateral veins 14-18, plane or prominulous above, prominulous beneath, often opposite, the intervenal areas patulous-reticulate, chartaceous, often discolorous, glabrous above, puberulent beneath, especially on the costa and veins, rarely subsessile; petioles 2-6 cm long, to 0.2 cm wide, puberulent, often flexuous; stipules free, appressed, the sheath rectangular, 4-5 mm long, or much larger (fide Standley), the lobes 6-8 subulate and flabellately disposed, terminal, 2-3 times the length of the body, acute, densely puberulent. /nflorescences terminal and axillary, the flowers densely crowded at the apex of the stem, the conglom- erate shorter than the petioles of the large leaves. Flowers with the hypanthium ca. 4 mm long, pubescent or glabrous, the calycine cup short, the calycine lobes 5-6, lanceolate, 5-12 mm long, 1.2-1.5 mm wide, acute, subcarnose, somewhat villosulose or villose especially on the margins; corolla white, the tube cylindrical, 20-30 mm long, 0.15 cm wide, petaloid to thickly carnose, puberulent on outside, glabrous or with glandular fimbriate structures within, these to 0.8 mm long, the 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Ficure 5. Amphidaysa ambigua (Standley) Standley. Habit (<2). [After Dressler & Mahler 2444.| lobes 5, narrow lanceolate or narrowly oblong, 3-5 mm long, with glandular fimbriate structures adaxially; stamens 3-5, the anthers oblong or ovate-oblong, 2-2.5 mm long, with a terminal petaloid appendage, subulate, 0.5-1 mm long, acute, the filament short: style slender, to 28 mm long, ca. 0.35 mm wide, the stigmas oblong, 4-5 mm long, 0.7 mm wide. Fruits oblong or oblong-rotund, 5-7 mm long; calycine lobes persistent, linear-subulate, unequal, to 17 mm long, acute, ca. 3 times the length of the fruit, the latter truncate, the nectary scar umbonate, the corolla tube scar annular, smooth or rugulose (the pressure of seeds against wall on drying), glabrescent to densely pubescent. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 33 Amphidaysa ambigua is found in Panama and in Colombia. It is of interest that all Panamanian collections were made during the current decade except for Terry & Terry 1489. LON: Santa Rita Ridge, D'Arcy & D'Arcy 6139 (MO); Dwyer 8583 (MO). DARIEN: Cerro Pirre, reo 511 (MO); Duke 5290, 6114, 13660 (all MO). Caná-Cuasí Trail, Chepigana, saa & Terry (MO). PANAMA: Road to Carti, 13 km N El Llano, Busey 385 (MO). Campo Tres, 5 km N Altos de Pacora, Busey 828 (MO). Road from El Llano to Carti-Tupile, Kennedy et al. Mad (MO). Eneida, Cerro Jefe, Maas & Dressler 682 (MO). Road from El Llano to Cartí-Tupile, Dressler & Mahler 2444 (MO); Liesner 652 (MO), 1142 (MO); Mori, Gentry & Kallunki 4558 (MO); Mori & Kallunki ae (MO): Mori 7710 (MO). 7 km W of Santa Fé, ca. 2900 ms Nee ae (MO). SAN BLAS: Trail from El Llano to Carti-Tupile, Kennedy 2579 (MO). COLOMBIA: CHOCO: N of es Curiche, Duke & Tok 11228 (MO). Bahia Solano (Puerto Mutis), Gentry & Forki 7228 (MO 2. Amphidaysa spathulata Dwyer, spec. nov. TYPE: Panamá, Duke 15615 (MO). Subshrubs about !⁄4 m tall, the stem erect, simple, terete, glabrous to gla- brescent, the leaves crowded toward the apex. Leaves sessile or subsessile, ob- long, 4.5-16 cm long, 1.8-8 cm wide, deltoid or obtuse at the apex, cuneate or attenuate-acute at the base, the costa slender, prominulous above or subsulcate, subprominent beneath, the lateral veins ca. 14, arcuate, forming an undulate vein beneath the margin, the pair of lateral veins including 1-2 subparallel veins, stiffly membranaceous, glabrous above, the raphides numerous, white, puberulent on the costa and veins; stipules free, puberulent, the body 2 mm long, thin, provided with glandular fimbriate structures within, with 6-8 linear setas, 0.4—0.6 mm long. Inflorescences to 4 cm long, the cymes sometimes on reduced branches. Flowers with the hypanthium narrow oblong, to 4 mm long, glabrous, the lobes of the calyx, elliptic-spathulate or narrowly obovate oblong, to 11 mm long, 2-4 mm wide, the blade 5.5-6.5 mm long, foliose, often 5-veined, acute, puberulent, the claw rigid puberulent, often longer than the blade, scarcely alate; corolla white, with the tube narrowly cylindrical, to 3 cm long, 2 mm wide, glabrous the lobes 5, lanceolate, to 6 mm long, acute; stamens 4, the anthers oblong rotund, the acumen ca. 0.4 mm long; stigmas 2, subulate, ca. 2.5 mm long. Fruits subsessile or on thick pedicels, puberulent, these provided with a stipuloid brac- teole toward the apex, this with 5 setas, linear subulate, to 8 mm long, narrowly oblong, to 1.5 cm long, 0.4 cm wide, 6-angled, lignose, drying black, with 8 slender ribs. The new species is known only from Panama but it is to be expected in Colombia. It is closely related to A. neblinae Steyermark from Brazil which has more acute leaves and narrower calycine lobes. DARIEN: Cerro Campamento, S of Cerro Pirre, Duke 15615 (MO), 15731 (US). ° Suffrutic es foliis confertis. Folia sessilia vel subsessilia oblonga, 4.5-16 cm longa, 1.8-8 cm lata, venis lateralibus 14, rigido-membranacea supra glabra; petiolis ad 0.5 cm longis: hryg liberis, corpore 2 mm longo, tenui intus fimbriis glandulis instructo setas 8 lineares ferente, eas 0.4-0.6 mm longas. /nflorescentiae ad 4 cm longae. did ^s eee ry 4 one Hoe latis vel edo obovato- oblongis, ad 11 mm Masa 2-4 mm latis na m longa, foliosa: des tubo ad 3 cm longo, 0.2 cm lato, glabro, lobis 5, ad 6 m m med DE 4, Mrs acumine ad 0.4 mm longo. Fructus subsessiles vel in n pedicellis crassis versus apicem bracteola stipulanea praediti hac 5 setis lineari-subulatis ornata, angusto-oblongi, ad 1.5 cm longi, 6-angulati costis 8 instructi. 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 6. ANTIRHEA Antirhea Commerson A. L. Juss., Gen. Pl. 204. 1789 (‘Antirhea’). TYPE: A. borbonica J. F. Gmelin. Antirhoea Endlicher, Ench. 272, 692. 15-21 Aug. pi Lo Stenostomum Gaertn. Fruct. Sem. Pl. 3: 69. 1806. TYPE: S. MU (Swartz) C. F. Gae Sturmia Gaertn. Fruct. Sem. Pl. p/. /92. 1805, not nA Hoppe (1799), not Sturmia Reichb. (1828). YPE: S. lucida Gaertn. Trees or shrubs. Leaves often coriaceous, the stipules deciduous or persistent. Inflorescences axillary, cymose, the peduncle when present with branches spreading, the flowers secundly disposed, the bracteoles present or absent. Flow- ers with the hypanthium ovoid or obovoid, the calyx lobes persistent; corolla funnelform, the 4-5 lobes imbricate, 2 of them exterior; stamens 4-5, included or scarcely exserted, the anthers slender, dorsifixed, often pubescent; style slen- der, the stigma knoblike or 2—3-lobed, included, the ovary 2-10-celled, the ovules solitary, pendulous. Fruits drupaceous, thin carnose, the shell ligneous or bony, the seeds elongate, pendulous. Antirhea, with about 40 species, occurs in the West Indies, Panama, Mad- agascar, in tropical East Asia to Australia. l. Antirhea trichantha (Griseb.) Hemsley, Biol. Centr. Amer., Bot. 2: 42. 1881.—Fic. 6 Pittoniotis trichantha Griseb., Bonplandia 6: 8. 1858. rype: Duchaissing (? P, not seen). Trees to 10 m tall, the twiglets round, smooth, drying black, glabrous or puberulent. Leaves oblong to ovate-rotund, 6-21 cm long, 2.5-8.5 cm wide, sub- equilateral, deltoid to rounded at the apex, often contracted into a narrow cusp to 2 cm long, usually acute, occasionally obtuse, basally acute to cuneate, the costa subplane above, prominulous to subprominent beneath, the lateral veins ca. 12, arcuate, the intervenal areas irregular pinnatiform, chartaceous, discolo- rous, densely strigillose above, densely pilose or villose beneath; petioles slender or stout, to 3 cm long, 0.8—1.5 mm wide, pilose; stipules persistent or decidous, appressed to the stem, triangular subulate, to 1.5 cm long, pilose. Inflorescences several at apices of the twiglets, axillary, spreading; peduncles usually wiry, 4— 7 cm long, ca. 0.8-1.5 mm wide, strict or flexuous, with a few alternate flexuous branches, these spreading, either with few flowers or occasionally with flowers densely congested, the flowers secundly disposed. Flowers with hypanthium ob- long rotund or oblong, ca. 1.5 mm long, densely pubescent, the calycine cup short, the teeth 5, oblong rotund, to 0.4 mm long, obtuse, petaloid, pubescent on the margin and outside; corolla white with the tube narrowly cylindrical or nar- rowly funnelform, 2.5-3 mm long, ca. 0.6 mm wide, petaloid, glabrous within, the lobes 5, oblong, about as long as the tube, densely villose above; stamens 5, exserted, the anthers linear oblong, 1.6-2 mm long, villose, the filaments linear, elongate, 0.5-2 mm long, attached at the corolla mouth; stigmas linear oblong, V? the length of the style, the style slender but dilated below stigmas. Fruits drupaceous, drying red, fleshy, oblong, 0.5-0.8 mm long, usually twice as long 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 35 FIGURE 6. Antirhea trichantha (Griseb.) eao s Habit (x.45).—B. Flower (x4.5).— C. SUPR cross section (x4.5). [After Stern et al. 149.] as wide, capped by a calycine cup to 0.8 mm long, drying black or brown, gla- brate, delicately longitudinally veined, seed solitary ca. 5 mm long, unilaterally groove Antirhea trichantha is known only from Panama. Few collections have been made outside the Canal Zone. L ZONE: Victoria Fill, Allen 1761 (MO). Chiva-Chiva Road, 1 mi E of Gaillard Highway, Blum 468 (MO). Albrook Tower, vicinity TTC site, Blum 551 (MO). C-15 Road to TTC Albrook T ini 1 ad K Preserve, Correa 146 (MO). Gaillard Highway near Penitentiary, Croat 9311 (MO). Road C 2 F, 42 mi NW Summit Naval Radio Station, Croat 11033 (MO). Fort Kobbe, Duke & Mussell 6667 (MO). 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Power Station near Fort Kobbe, Dwyer 2668 (MO). US Army TTC, Albrook, Dwyer 6727 (MO). Dam, Dwyer & Elias 7477 (GH, MO, US). Madden Dam, Boy Scout Road, Dwyer & Lallathin 8579, 8824 (both MO). Madden Dam, Dwyer & Robyns 24 (MO). Farfan Beach Road, Gentry 4901 (MO). Navy Pipeline Preserve, 3 mi NW Gamboa, Haines 509 (MO). Madden Dam, px 6248 (MO). Madden Dam, Boy Scout Road, Kirkbride 45 (MO). Farfan Beach, Kirkbride & Elias 74 (MO). US Navy Ammunition Depot, 2 mi NW of Balboa, Luteyn & Foster 930 (MO). Howard ^ Forc aU SE Kobbe ue rd & Mac Eo dd Sate Madden Dam, Boy Scout Road, Porter et al. 4015 (COL, DUK SCZ, UC, N). Naval Ammunition Depot, Stern et al. 38 (MO). Farfan Beach, Tyson 182 WS BARRO COLORADO ISLAND: Croat 4971, 6336, 10391, 11973, pone (all MO). DARIEN: Rio Pucro below village of Pucro, doe 13134 (MO). El Punteadero bridge, over Río Chucunaque, ya b al. 149 (MO). PANAMÁ: mbo, Croat 14906 (MO). Río Pita, DUE 4800 (MO). Chilibre, Lao & Holdridge 8 (MO). 6 2% SW Arraijan, old road to Bique, Nee 6890 (MO) 7. APPUNIA Appunia Hook. f. in Benth. & Hook. f., Gen. Pl. 2: 120. 1873. TYPE: A. tenuiflora Bellynkxia Muell.-Arg., Flora 48: 465. 1875. TYPE: B. angulata Muell.-Arg. Shrubs or small trees. Leaves usually lanceolate, opposite; stipules free, sub- ulate acuminate. Inflorescences terminal or axillary, the flowers few, congested into a small head at the end of the peduncle. Flowers with the hypanthium some- what coherent, the calycine cup truncate, the teeth absent or scarcely visible, funnelform or U-shaped, the lobes usually 5, valvate; ovary 4-celled, with an ascending, solitary ovule in each cell. Fruits sessile, aggregate, but virtually free at maturity, each fruitlet fleshy, drupaceous, with 4 pyrenes, each pyrene un- equally 2-locellate, the ventral part empty; seeds exalbuminous. Appunia is a genus of approximately 10 species in Central and South America. Literature: Hayden, M. V. & J. D. Dwyer. 1969. Seed Morphology in the Tribe Morindeae. Bull. Torrey Bot. Club 96: 704-710 — . Appunia seibertii Standley, Ann. Missouri Bot. Gard. 24: 208. 1937. TYPE: Panama, Seibert 624.—FiG. Shrubs to 1.5 m tall, the branches exfoliate, subterete, smooth, red brown and cracking when dry, glabrous or puberulent, the nodes thickened, the branchlets slender. Leaves somewhat verticillate at the apex of the branchlets, narrow el- liptic, rarely subrhombiform or obpyriform, acute or deltoid at the apex, attenuate cuneate at the base, 5-17 cm long, 1-7 cm wide, membranaceous, scarcely dis- colorous, often green when dry, glabrous or scattered granulose above, the in- tervenal areas minutely granulose beneath, the costa slender, prominulous above, subplane distally beneath, the lateral veins opposite or alternate, ca. 12, slender, prominulous, widely arcuate, the veins white puberulent, the margins glabrous or minutely puberulent; petioles 0.3-2 cm long, ca. 0.7 cm wide, puberulent; stipules triangular, 2.5-5(-7) mm long, 2-4 mm wide, smooth, glabrous, entire. Inflorescences 1-4, pseudoterminal, arising from the axils of the fasciculate leaves, the peduncles several per axil, umbellately disposed, slender, 0.5-1.5 cm 1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 37 Appunia seibertii Standley.—A. Habit (x4.5). [After Oliver & MacBryde 1896.|—B. mu e r^ unm Duke 4713.] long, ca. 0.5 mm in diam., minutely red or white puberulent, the flowers aggre- gated into a solitary terminal head ca. 4 mm in diam., with 10-11 flowers. Flowers sessile; calyx cupuliform, ca. 1.5 mm in diam., the teeth absent or sometimes as irregular points, to 0.5 mm long, fleshy, glabrous, the raphides many: corolla (in bud) with the tube ca. 7 mm long, ca. 2 mm wide, glabrous outside, pubescent within, the lobes 5, oblong to triangular, scarcely shorter than the tube, briefly cucullate; anthers 5, oblong, to 2 mm long, the filaments attached near the apex of the tube; ovary 4-loculate, the septa thick, the style short, the stigmas 2, 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 subulate ca. 1.25 mm long. Fruits sessile, usually 2-5, crowded at the apex of the peduncle, the individual fruits turbinate, to 1 cm long, black and smooth when dry, often inconspicuously 4-lobate, each pyrene unequally 2-locellate, the ventral part empty, bulliform, the endocarp replete with raphides; seeds 4, oblong, ca. 5 mm long, slightly winged. This species is known from Panama and Colombia. Additional notes may be found in Dwyer & Hayden (1969), where there is an emended description of the species. In forming fruit often several flowers of the mass fail to mature. The persistent calycine tube and floral disc, as well as the hypanthium, sim- ulate a Morinda fruit in miniature. CANAL ZONE: Fort dris Duke 3958, 4246, 4698, ed (all MO). Howard Air Force Base, SE of Kobbe Beach, Oliver & MacBryde 1896 (GH, MO, US). DARIEN: Without locality, Bristan 207 (MO). El Real to Pinogana, Duke 5126 (MO). Río Pucro to Quebrada Maskia, Duke 13082 (MO). Río Mortí, Camp Morti iac il 14118 (MO). Chep pigana, Duke & gei 267 (MO). Río Tuquesa, lower Tuquesa Mining P d Charco Chiva, Mori 6987 (MO). Periaque camp at river, Tyson et al. 4767 (MO). ae "Rio Bayano above confluence with Rio ae Duke 4000 (MO). Cermeno, Dwyer 5604 (MO). Arenoso, lower Rio Trinidad, Seibert 624 (MO). 2 mi E of El Llano, Tyson 1749 (MO). SAN BLAS: Isla Pino near Mulatupo, Elias 1718 (MO, UC, VEN) 8. ARCYTOPHYLLUM Arcytophyllum Willd. ex. Schultes in R. & S., Syst. Veg. 3: 5. 1827. TYPE: A. blaerioides Willd. ex. J. S. & J. H. Schultes. Mallostoma Karst., Fl. Colomb. 2: 9. 1862. TYPE not indicated. Rac (s T ae Prodr. 4: 433. 1833. rvPE: R. rupestris (Sw.) A. P. DeCandolle, based on Hedyotis Pseudorhachicali Benth. & Hook., Gen. Pl. 2: 60. 1873. rYPE not indicated. Ereicoctis Kuntze, Rev. Gen. Pl. 281. 1891. TYPE not indicated. Shrubs or subshrubs, erect or prostrate. Leaves small, crowded, often densely imbricate, coriaceous; stipules entire, bifid or setose. /nflorescences cymose or flowers solitary, axillary. Flowers with the hypanthium globose or turbinate, the calyx 4-5 lobed, often with glands between the lobes; corolla funnelshaped or salverform, the lobes 4; stamens with the anthers included or exserted, dorsifixed, the filaments short and adnate to the tube in riblike fashion; style slender, the stigmas 2, short, the ovary 2-celled, the placentas adnate to the septum. Fruits capsular, turbinate to globose, 2-celled, usually septicidally dehiscent to the base; seeds few, boatshaped or plano convex, oblong, punctate, winged. Arcytophyllum, a genus of about 30 species, ranges from Mexico through Central America, extending to the Andes of South America. 1. Arcytophyllum lavarum Schum. in Engl. & Prantl, Nat. Pflanzenfam. IV (4): 28. 1891. TYPE: Panamá, Maxon 5349 (US, holotype).—FIG Mallostoma lavarum (Schum.) Donnell. Smith, Enum. Pl. Guatem. 5: 36. 1899. Subshrubs, ericoid in habit, profusely branched, the branches lax, glabrous, drying silver tan or purple black, ridged, the nodes well spaced or crowded, swollen (bases of persistent stipules). Leaves oblong, 0.5-0.8 cm long, 0.2-0.4 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 39 EA í £ Bay ate Ñ of’ Q e AN Ay d "i S Ly È s Ww FIGURE 8. Arcytophyllum lavarum Schum.—A. Habit (x!2).—B. Flower, opened (x5).—C. Fruits (x 5).—D. Seeds (x5). [After Wilbur et al. 11033.) cm wide, obtuse at the apex, obtuse to truncate at the base, the costa and the lateral veins evanescent, carnose, ericoid, lustrous, scarcely bicolorous, marces- cent on drying: petioles to 1 mm long or absent; stipules connate, disposed as a short cup, to 0.25 cm long, the free portion deltoid, longer than the cup and often contiguous with the stipule above, the margin erose with a few ascending irregular subulate teeth. /nflorescences terminal, to 1.5 cm beyond the uppermost leaves, usually with 2 peduncles, to 0.3 cm long, each bearing a terminal cymule. Flowers with the hypanthium rotund, to 0.65 mm long, petaloid, glabrous, the calycine cup very short, with patches of flabellately disposed minute oblong glands and squamellae intermixed within, the teeth 4, erect, oblong or ovate oblong, often obtuse, to 1.3 mm long, often with an upright gland at the junction of 2 lobes, glabrous, petaloid; corolla white, white tinged with purple, blue or lavender pink, the tube campanulate oblong, ca. 2.5 mm long, the lobes not cucullate, villosulose on the adaxial face; stamens 4, the anthers oblong, 0.8-1 mm long, dorsifixed, 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 the filaments slender, attached above the middle of the tube, the style slender, ca. 4 mm long, slightly dilated apically, the stigmas 2, clavate when connate, ca. 0.4 mm long; ovary thin walled, the septum thin, the ovules several, arising on a distinct stipe attached almost at the base of the septum, the locule ample. Fruits short pedicellate, capsular, rotund or compressed rotund, to 1.5 mm in diam., truncate, the calycine lobes persistent, a thin annular ring marking their point of origin, the valves splitting from the apex. Arcytophyllum lavarum is restricted to high mountainous regions of Costa Rica and Chiriqui. A photo of A. lavarum at MO is labeled as a Pittier collection (unnumbered) from Costa Rica, deposited in Berlin, but this is not the type. HiRIQUÉ: Volcán de Chiriquí, arc 1293 (F, MO). Between Baru and Respinga, d m, D'Arcy 10095 (MO). Chiriqui Volcano, 3374 m, Maxon 5349 (US). W side of Volcán Baru, ca. 3000 m, Mori & Bolten 7342 (MO). Trail to summit, Volcán de s 2500-3400 m. Mori hi Kallunki 5748 (MO). Volcan de Chiriqui, W slope, Wilbur et al. 11033 (F, MO). Potrero Muleto to summit of Volcan de Chiriqui, Woodson & Schery 437 (MO). Loma Ta to summit of Volcan de Chiriquí, Woodson et al. 1087 (F, MO) 9. BATHYSA Bathysa Presl., Bot. Bemerk. 84. 1844. TYPE: B. stipulata Presl. Trees or shrubs. Leaves opposite, the stipules interpetiolar, entire, occasion- ally persistent. Inflorescences terminal, paniculate. Flowers abundant, the ca- lycine cup truncate, the teeth 4-5; corolla subrotate to infundibuliform, the lobes 4-5, imbricate; stamens 4-5, inserted on the mouth, the anthers dorsifixed, exserted; ovary 2-celled, the ovules numerous. Fruits capsular, septicidally de- hiscent, splitting into 2 valves from the apex; seeds minute, horizontal, com- pressed or angular, winged or not, the testa reticulate. Bathysa ranges from Panama through Venezuela, Brazil, and Peru. It is a genus of about 10 species. l. Bathysa veraguensis Dwyer.’ TYPE: Panamá, Lao & Gentry 531 (MO, holo- type).—FiG. 9 Trees to 5 m tall, the branchlets terete, minutely velutinous, the nodes well spaced. Leaves large, widely oblong, 10.5-31.0 cm long, 4.5-18.0 cm wide, widely deltoid toward the apex, occasionally rounded, acuminate, the acumen short, widely deltoid or narrowly deltoid, to 1.8 cm long, 0.3-0.5 cm wide in the middle, basally widely cuneate, obtuse or truncate, the costa prominulous above, prominent beneath, the lateral veins 15-19, to 2 cm apart in the middle of the 7 Bathysa veraguensis Dwyer, spec. nov. Arbores ad à m altae. Folia magna, 5-31.0 longa, 4.5-18.0 cm lata, venis lateralibus 15-19 papyracea supra in costa venisque PE E cetera minute scabridula subtus dense aureovelutina. info scentiae terminales amplae, ad 30 cm longae, cymulis plerumque 3-floribus; bracteis ovato-lanceolati 1.3 longis. Flores cupula calycis ampla ca. 2.5 mm alta, apice ca. 0.6 cm lata, lobis 4-5, ad 5 mm longis; corolla alba tubo 3.3 0 mm longo, lobis ca. 6 mm BRA pe d ca. 3.2 mm longis, filamentis crasso-subulatis. Fructus iru vel turbinati, ad lati, extus puberuli calyce persistenti magno a ongi, cm m longo, fortasse 2-plo lont: UR superanti seminibus polymorphis plano-compressis oi ad 0.8 mm longi 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 41 E9. Bathysa veraguensis Dwyer.—A. Inflorescence and leaf (x .45).—B. Leaf underside (X. 45 C. Flower, opened (x3.15). [After Lao & Gentry 531.] 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 blade, the venules irregular perpendicular, papyraceous, discolorous, dark red to brown when dry, golden villosulose above on the costa and the veins, otherwise scabridulous, beneath densely golden velutinous; petioles and stipules not seen. Inflorescences terminal, solitary, pyramidal paniculate, to 30 cm long, to 30 cm wide, velutinous; peduncle to 8 cm long, to 0.35 cm wide, thick, lignose, the branches often opposite, to 15 cm long, the pairs well spaced, often 6-7 cm apart, the cymules often 3-flowered, disposed on long or short secondary branches or on tertiary branches, each branch provided with 3 flowers, rarely with one flower; bracts ovate lanceolate, to 1.3 cm long, acute, appressed villose outside, inside with a few hairs; bracteoles ovate subulate. Flowers pedicellate, the pedicels 0.3— 1.0 cm long, hypanthium turbinate, 0.3 cm long, puberulent, the calycine cup expanded, 2.5 mm long, 0.6 mm wide at the apex, eglandular within, the lobes 4—5, oblong triangular, 5 mm long, often ascending, 0.4—0.5 cm wide at the base, obtuse, coriaceous, puberulent or appressed villose within; corolla white, the tube compressed cylindrical, 3.3-5.0 mm long, often to 0.5 cm wide, glabrous outside, within densely golden villose near the mouth, the lobes imbricate in bud, oblong rotund, 6 mm long; stamens 5, the anthers narrowly oblong, 3.2 mm long, slightly curved at anthesis, obtuse, exserted, the filaments attached near the mouth, thickly subulate, 3 mm long; ovarian disc ringlike, 3.0-4.5 mm long, 1.3- 1.6 mm wide, 0.5—0.65 mm deep, the style thick, 7.5 mm long, the stigmas 2, thick, narrowly oblong, 2.5 mm long. Fruits immature? oblong or turbinate, to 1.5 cm long (including calyx), ca. 0.5 cm wide, the persistent calyx as in flowering stage, perhaps 2 times the length of the pericarp, densely puberulent outside, densely appressed villose within; seeds polymorphic, mostly oblong, to 0.8 mm long, compressed, reticulate. Bathysa veraguensis is known only from Panama. The genus is not otherwise known in Central America. COCLÉ: Cerro Pilon, ca. 3100 ft, Dwyer & Lallathin 8641 (MO). ds Pilón, hill below summit, ca. 2500 ft, Dwyer et al. 4544 (MO). vERAGUAS: Cerro Tute, 1200 m o & Gentry 531 (MO). NW of Santa Fé, 2.1 km from Escuela Agrícola Alto de Piedra, Mori & eas 4768 (MO). 10. BERTIERA Bertiera Aubl., Hist. Pl. Guiane 180. 1775. "Berthiera" TYPE: B. guianensis Aublet. Shrubs or small trees. Leaves petiolate or sessile; stipules connate, the sheath bifid. Inflorescences terminal, cymose paniculate, pedunculate, the bracts nar- row. Flowers with the hypanthium subglobose or turbinate, the calycine cup truncate and edentate or with 5 teeth: corolla funnelshaped, the tube usually sericeous outside, the lobes 5, ovate, contorted; stamens 5, the anthers ovate or subulate, dorsifixed, the connective acuminate or produced at the apex, the fil- aments short, attached to the tube; ovarian disc cupular or annular, the style slender, glabrous, the stigmas entire or disposed as 2 narrow lobes, the ovary 2-celled, the ovules numerous, the placentas entire, affixed to the septum. Fruits baccate, fleshy; seeds numerous, small, angulate, the testa foveolate or granulate. Bertiera occurs in tropical America and in Africa. It is a genus of about 30 species. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 43 Literature: Standley, P. S. 1934. 65. Bertiera Aubl., N. Am. Fl. 32(3): 180-181. a. Leaves with 8-9 lateral veins; stipules 1.5(—2.0) cm long 1. B. angustifolia aa. Leaves with ca. 7 lateral veins; stipules 0.9-1.2 cm long 2. B. guianensis 1. Bertiera angustifolia Benth., Bot. Voy. Sulphur 103. 1844. Type: Cocos Island, Costa Rica, not seen. Shrubs or trees to 5 m tall, the branchlets terete, smooth, densely pilose, the nodes well spaced. Leaves arising as juveniles from a cylindrical bract or pair of bracts, to 2 cm long, the bract drying brown, puberulent; leaf blades ovate oblong, 9-17 cm long, 2.7—7.5 cm wide, acute toward the apex, acuminate, the acumen often inconspicuous, obtuse or rounded at the base, the costa prominulous, the lateral veins 8-9, strongly ascending, sometimes immersed above, papery when dry, rubescent above, glabrous except for the costa, glabrous beneath except for the costa and veins, the hairs appressed, elongate; petioles to 0.5 cm long, pu- bescent; stipules triangular subulate, to 1.5(—2.0) cm long, ca. 0.5 cm wide at the base, ultimately appressed to the stem. Inflorescences terminal, solitary, panic- ulate, puberulent, to 17 cm long, to 7(-10) cm wide; peduncle slender, 3.5-7.0 cm long, to 0.18 cm wide, the rachis somewhat flexuous, the branches alternate or opposite, rarely 4-verticillate; bracts and bracteoles linear subulate, equal to the nonfloriferous parts of the branchlets. Flowers small (in bud); hypanthium oblong, to 1.5 mm long, pubescent, the calycine cup small, the teeth 5, small, triangular; corolla tube 1.5-2.0 mm long, carnose, glabrous or sparsely puberulent outside, densely villose inside, the lobes deltoid, to 1.3 mm long, acute; stamens 5, the anthers oblong, to 1.5 mm long, short acuminate at the apex, the acumen ca. 0.3 mm long. Fruits sessile, rotund, 3 mm in diam., drying black, glabrate to sparsely pubescent, the hairs short, strigose, the ribs obvious, obtusely margin- ate, the calycine cup vestigial. Bertiera angustifolia is known only from Costa Rica (Cocos Island), Panama and Peru. Standley erroneously annotated Mexia 6358 from Pongo de Manse- riche, Dept. Loreto, Peru as Bertiera parviflora Spruce. This collection matches well the three Panama collections cited below. The photo (MO) of the type of Bertiera parviflora (Spruce 2774) does not match the three Panama collections; the lateral veins are only 5-6 in the type but 8 or 9 in B. angustifolia. Bentham described the leaves of B. angustifolia as "'pellucid-punctate,"" but this is not so in the Panamanian collections. The only genus of Rubiaceae in Panama with pellucid-punctate leaves is Rustia. OCAS DEL TORO: 1 Ar gin Menaco and Buena Vista, Chiriquí Trail, Kirkbride & Duke 666 (MO). RIÉN: Can 0 ft, Stern et al. 403 (MO). vERAGUAS: Río Primero Brazo, 2.5 km beyond | ee rien Pos Piedra, 700—750 m, Croat 25477 (MO). 2. Bertiera guianensis Aubl., Hist. Pl. Guiane 1: 180. 1775. Type: French Guiana, Aublet (P, holotype, not seen).—Fic. 10 B. palustris A. Rich., D.C., Prodr. 4: 392. 1830. B. aequaliramosa Steyermark, Mem. New York Bot. Gard. 17(1): 319. 1967. svyNTYPE: Panamá, Cooper 586 (NY, holotype). 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 B. guianensis var. leiophylla Steyermark, Mem. N.Y. Bot. Garden 17: 319. 1967. rvPE: Panama, Pittier 5178 (NY, holot B. tenuis Lundell, Wrightia 4: 49. 1968. TYPE: Guatemala, Contreras 4318 (LL, holotype). Shrubs or small trees to 4 m tall, the branchlets terete, slender, appressed villose, the nodes well spaced. Leaves oblong, 8—17 cm long, 2-6 cm wide, acute and often vaguely acuminate at the apex, acute or cuneate at the base, the costa prominulous above and beneath, ca. 0.1 cm wide proximally, the lateral veins ca. 7, arcuate, occasionally strict and sharply ascending, discolorous or not, glabrous above and beneath except villose along the costa and veins; petioles slender, to 0.2 cm wide, 0.2-2.0 cm long; stipules deltoid, narrowly or roundly so, 0.9-1.2 cm long, ca. 0.3 cm wide, pubescent. /nflorescences terminal, solitary, cymose thyrsoid, to 16 cm long, to 6.5 cm wide, the peduncle to 7 cm long, stiff but thin, 1.0-1.5 mm wide, densely villose, the branches well spaced, the lowermost op- posite or whorled, subperpendicular; bracts persistent, subulate, to 6 mm long, the flowers secund. Flowers with the hypanthium subrotund, ca. 1 mm long, pubescent, the calycine cup short, the teeth 5-6, vague or as distinct short points; corolla white, the tube cylindrical, to 6 mm long, 0.8 cm wide in the middle, glabrate outside and within, the lobes 5, oblong, acute, villose within; stamens 5-6, the anthers 0.8-1.5 mm long, acuminate at the apex, the acumen to 0.35 mm long, acute, sagittate at the base, attached near the mouth; stigmas 2, oblong lanceolate, ca. 2 mm long, the style slender, ca. 2.5 mm long, the ovary thick walled, replete with resinous dots. Fruits sessile, rotund, 3.5-4.0 mm in diam., the pyrenes 6-costate, drying black; seeds obtusely 3-angled, ca. 1.5 mm in diam., muricate. Bertiera guianensis ranges from Mexico to Panama, the Guianas, and south to Bolivia. It also occurs in the West Indies. BOCAS DEL TORO: Buena Vista, Almirante, Cooper 586 (NY, US). BARRO COLORADO ISLAND: Aviles 9 (MO); Croat 12503 (MO); Foster 774 (MO). CHIRIQUi: San Félix, Pittier 5178 (US). DARIEN: Airstrip, Caná, 480 m, Croat 37997 (MO). Cativo Swamp near Tusijuanda, 1 day up Río Balsa, dus 13533 (MO). San José River, Duke & Bristan 397 (MO, NY). Between Paya and Palo de Los Let Duke & Kirkbride 14030 (MO). Río Tuquesa, middle Tuquesa Mining Co. Camp called Charco Peje. Mori 7057 (MO). From Paya to Pucro, TUE et al. 225 (MO). Caná, Stern et al. 693 (MO). PANAMÁ: Chimán, Lewis et al. 3243 (MO, UC, ll. BOROJOA Borojoa Cuatrecasas, Serie Bot. Aplicada, Serv. Divulg., Secr. Agric. y Ganad. (Calí, Colombia) 2(5): 474. Aug. 1949. (Preprint of Rev. Acad. Colombia Ci. 7: 474. 1950.) TYPE: B. patinoi Cuatrecasas. Small dioecious trees. Leaves large, usually somewhat coriaceous, petiolate; stipules partly connate basally, interstipular, often persistent. Inflorescences ter- minal, pistillate flowers 6—8-merous; male flowers sessile, crowded, disposed in a terminal head; pistillate flowers usually terminal, solitary. Staminate flowers 4— 5-merous; corolla hippocrateriform or funnelform, pubescent outside and within, the lobes contorted in bud; anthers 5, linear, dorsifixed; ovary 6-8-celled, the carpels united along their margins, the ovules numerous, the stigmas 6-8. Fruits baccate, large, rotund, the pericarp wall very thick; seeds embedded in a slimy pulp, disposed in rows, horizontally attached. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 45 "wA Q> Y» O 1 i | FIGURE 10. Bertiera guianensis Aubl.—A. Habit (x1⁄4).—B. Fruit (x3.5). [After Stern et al. 93.] 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Borojoa is known from Panama, Colombia, Venezuela, and Brazil. It is a genus of about 8 species. Literature: Cuatrecasas, J. 1950. Borojoa, Nuevo Gen. Rubiaceae. Aug. 1949; Revista Acad. olomb. Ci. Exact. 7: 474—477 Patino, U. M. 1950. Noticia sobre el Borojó, una nueva especie frutal de la costa Colombiana del Pacífico. Revista Acad. Colomb. Ci. Exact. 7: 478-489 a. Lateral veins of leaves 6-12. Lateral veins of leaves 10—12; dus to 6.5 cm in diam... l. B. atlantic bb. Lateral veins of leaves ca. 7: fruits to 4.5 cm in diam. |... 2. B. panamensis aa. Lateral veins of leaves 13—20 3. B. patinoi 1. Borojoa atlantica Dwyer.’ TYPE: Panama, Kennedy & Foster 2197 (MO, ho- lotype). Trees to 10 m tall, the branchlets terete, glabrous, the nodes well spaced. Leaves elliptic or obovate elliptic, 18-27 cm long, 6-13.5 cm wide, widely deltoid toward the apex, acuminate, the acumen triangular, 1-2 cm long, basally obtuse, cuneate or truncate, the costa lightly porcate above, prominent beneath, the lateral veins 10-12, arcuate, slender, the intervenal areas spreading reticulate, concolorous, drying green, thin coriaceous, glabrous above, minutely puberulent beneath; petioles 0.3-1.2 cm long, rigid or curved, puberulent; stipules (only one seen) connate?, ovate, to 1.3 cm long, ca. 0.5 cm wide, acute, subcoriaceous, pu- nds: Inflorescences and flowers not seen. Fruits apparently sessile, rotund, o 6.5 cm in diam., lignose, lightly wrinkled on the surface, lightly puberulent, un many slender longitudinal grooves when dry, the wall thick, to 2 cm wide, the persistent calycine cup umbonate, to 1 cm long; seeds many, embedded in a fleshy pulp, subplane, often angular, to 1 cm long. Borojoa atlantica is known only from Panama. It is readily distinguished from B. panamensis by the more numerous lateral veins of its leaves and it is from B. patinoi whose leaf blades have still more lateral veins (up to 20). The fruits of the new species and those of B. patinoi are similar. Borojoa atlantica seems closely related to B. universitatis Steyermark, the type of which (Aristequieta 6466; US, isotype) is from the Parque Nacional de Guatopo, 650 m, Venezuela. The lateral veins of B. universitatis number about 18 and the wall of the large fruit (in the dried state) is only about 1 cm thick. LÓN: 2-3 miles up from the Río Guanche, Kennedy & Foster 2197 (MO). PANAMÁ: El Llano- Cartí Road, Folsom 2620 (MO); Folsom & Collins 1517 (MO). Gorgas Memorial Labs, yellow fever research camp, Campamento Quatro, 5-10 km NE, Altos de Pacora, 600 m, Mori & Kallunki 3336 MO). * Borojoa atlantica Man spec. nov. Arbores ad 10 m altae. Folia se vel igi -elliptica, 18-27 (vel p us) cm longa, 6-13.5 cm lata, apicem versus lato-deltoidea acumine triangulari, 1-2 cm | h : : sessiles rotundi, ad 6.5 cm diam., pariete crasso, ad 2 cm lato, lignoso: seminibus multis in pu ln carnosa inclusis subplanis oblongis saepe angularibus, ad 1 cm long 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 47 2. Borojoa panamensis Dwyer, Phytologia 17: 446. 1968. TyPE: Panamá, Duke & Lallathin 15014 (MO, holotype). Small trees, the branchlets subterete, somewhat rimose, ultimately angular, presumably glabrous, the nodes to 9 cm apart, the scars of the petiole prominent, subrotund, ca. 0.4 cm in diam. Leaves elliptic rotund, to 19 cm long, to 12.5 cm wide, widely deltoid at apex, short acuminate, cuneate and subequilateral at base, the lateral veins ca. 7, arcuate, stiffly chartaceous, the lamina glabrescent, the costa with many minute golden hairs and the axils of the lateral veins golden barbate; petioles to 2 cm long, to 0.2 cm wide in middle; superior stipules per- sistent, connate to the middle, the sheath cylindrical, to 1 cm long, the triangular basal prominence to 6 mm long, ca. 0.5 cm wide, at maturity torn and reflexed, the free portions obovate rotund to elliptic, to 2 cm long, to 1 cm wide, acuminate, often wider above middle, thin coriaceous, dark brown, with many small veins and with a median keel. Flowers not seen. Fruits terminal, sessile, solitary, rotund, to 4.5 cm in diam., glabrous, minutely rugulose, the persistent calycine cup like a volcano’s crater, to 0.5 cm long, the orifice ca. 0.4 cm wide. The species is known only from the type locality in Panama. Duke & Lallathin note on their collection that the fruits are "green, probably edible when ripe. Fruit, when sawed into, blackening as if possessed of genipine.”’ COCLÉ: Cerro Pilón near El Valle, ca. 2500 ft, Duke & Lallathin 15014 (MO); Lallathin 5014 O). 3. Borojoa patinoi Cuatrecasas, Nuevo Gen. Rubiaceae. Aug. 1949; Revista Acad. Colomb. Ci. Exact. 7: 474. 1950. TYPE: Cuatrecasas 21137 (F, holo- type).—Fic. 11. Trees to 7 m tall, the branchlets smooth, plane, glabrous, the bark rimose, the nodes well spaced, to 6 cm apart, the pedicel scars often prominent. Leaves elliptic, to 36 cm long, to 17 cm wide, cuneate at the apex, basally cuneate, truncate or obtuse, the costa prominulous above, prominent beneath, to 1.8 mm wide, obviously porcate distally, the lateral veins 13-15, widely arcuate, to 3 cm apart, usually 1—1.5 cm apart, the venules pinnatiform, spreading, discolorous, subcoriaceous, glabrous; petioles to 4 cm long, glabrous; stipules tending to per- sist, often reflexed at maturity, connate, appressed to stem, the sheath cylindrical, the parts ovate elliptic to ovate, 2.5—4 cm long, to 1.2 cm wide, acute at the apex, stiffly chartaceous with a slender, median keel, venose, the veins crowded, prom- inulous. Inflorescences terminal, the male flowers numerous, disposed in a ter- minal head about equal to the petiole; bracts disposed as 2, connate, decussate pairs, soon spreading or reflexed; bracts subtending the solitary female flower, the inner pair suborbicular, the outer pair similar in size and texture to the sti- pules. Male flowers 4—5-merous, the hypanthium and the calycine tube tubular, 0.7-0.8 cm long, glabrous, stiffly membranous, truncate, the teeth absent or vague; corolla white, to 25 mm long, the tube hippocrateriform, to 18 mm long, thickly petaloid, densely sericeous outside, the lobes 5, 6-7 mm long, oblong, obtuse, somewhat shorter than the tube; stamens 5, the anthers linear, ca. 12 mm long, the filaments slender, short; female flowers 6—7-merous; style to 7 mm long, the stigmatic lobes 6, linear lanceolate, slightly longer than the style, acute, pap- 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FiGURE 11. Borojoa patinoi Cuatrecasas—A. Branch with leaves and fruit (x.35). [After Duke 14181.|—B. Fruit cross section (x .35). [After Kirkbride & Duke 407.] illose. Fruits sessile, rotund, applelike, 7-8 cm in diam., subtended by the per- sistent bracts, glabrous, the remains of the calyx represented by a delicate ring bordering a terminal cavity, ca. 1 mm deep, the wall very thick, 1.2 cm in diam.; seeds subplane, subovoid, 6-7 mm long, 7-12 mm wide, embedded in a pulp. The above description has drawn freely on Cuatrecasas (1949) as the Panama collections are sterile with one exception and this is in bud. Borojoa patinoi is known only from Panama and Colombia. *‘Borojo,’’ ''Bo- rojo Hembra,” *‘Borojo Macho,” or ‘‘Burijo. Duke in his Darien Ethnobotanical Dictionary states: '' This favorite fruit tree, one fruit making a bowl of chicha, has followed the Choco (Indians) into Panama (from Colombia), in whose flora it is unreported.’’ Duke adds, speaking of him- self: “I am called Borojo among the Darien Choco, because for a while, I was distributing borojo seedlings like Johnny Appleseed.’ He adds that the fruits are edible, are aromatic, and take more than one year to ripen ÉN: Cerro Pirre, Bristan 495 (MO). Santa Fé, Duke & Bristan 310, 311 (both MO). Río Tucutí between Río Tucutí and Río Uruganti, Duke 5280 (MO). Without peux locality, Duke 8332 (MO). Finca Othon near Yape, Duke 11820 (MO). Río Mortí, Drill Site 7, ca. 250 m, Duke 14161 (MO). Río Punusa and Río Pucro, Duke 14637 (MO). Río Paya, 20 minutes ced mouth of Río Paya, Kirkbride & Duke 407 (MO). SAN BLAS: Eslogandi, 200 m, Duke 10196 (MO). 12. BORRERIA Borreria G. F. W. Meyer, Prim. Fl. Esseq. 79: 1818. nomen conserv. contra Borreria Acharius (1810, Lichen). TYPE: B. suaveolens G. F. W. Meyer, type conserv. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 49 impie Adanson, Fam. Pl. 2: 145. 1763. Type: Rheede, Ag Hortus qm Malabaricus 9, tab. 76. (design. gg 1916) = Borreria articularis (L.f.) F. N. William Borrera G. F. W. Meyer, Prim. Fl. Esseq. 79. 1818. orth. esee t Gen. 1: 93. 1830. Herbs, annual or perennial, rarely suffrutescent. Leaves with the petioles united to the stipules forming a setiferous sheath. /nflorescences capitate, usually sessile, axillary or terminating the stem. Flowers with the calycine lobes 2 or 4, equal or unequal, often with interdental teeth, the corolla reduced, funnelshaped, the lobes 4, the stamens usually attached near the mouth; ovary 2 celled with an ascending ovule in each cell. Fruits capsular, coriaceous or membranaceous, 2-valved, the valves septicidally dehiscent; seeds 1 per cell. Borreria occurs throughout the warmer parts of the world. It is a genus of about 150 species. Literature: Steyermark, J. 1972. 39. Borreria G. F. W. Meyer. The Botany of the Guayana Highland—Part IX. Mem. New York Bot. Gard. 23: 805-831. a. Plants more than 1 dmt b Inflorescences E heads 1.1-2.0 cm in diam. Leaves 0.1—0.25 cm wide or if wider, corolla lobes pilulose within -------------- 6. B. suaveolens cc. Leaves 0.6-3.2 cm wide. Leaves 2 at a node, corolla lobes pilulose within; stipular sheaths pubes- 2. B. laevis cent dd. jan 5-6(-8) at some of the nodes, corolla lobes not pilulose; stipular ths glabrous B. densiflora bb. igen i. with “heads (0.3—)0.5-1.0 cm wide. Inflorescences with heads axillary, or if terminal heads present, these m reduced in diam.; stems villose; leaf blades usually drying yellow ee flushed with red 3, B. tacfollo ee. Inflorescences with heads both terminal and axillary, similar in diam.; stems ee = ie sel not drying yellow green or flushed with red. f es2ata ae "Lente 0. pi 1. 0 cm wide; corolla ca. 1 mm long; fruits with one carpel with 2 calycine lobes . ocimoides gg. Leaves (0.6—)1.0-3.0 cm wide; corolla ca. 2.5 mm long; fruits with one carpel with 3 of the 4 calycinelobes -_-------------------- . B. laevis Leaves 5-6 at some of the nodes |... - l. B. densiflora aa. Plants less than 0.6 dm tall. h. Leaves to 6 mm long _________----------------------------------------------------- 5. B. pumilio hh. Leaves 10-25 mm long ________-------------------------------------------------- 4. B. ocimoides 1. Borreria densiflora DC., Prodr. 4: 542. 1830. TYPE: Jamaica, Swartz, not seen. Spena nar e spinosa L. Sp. Pl., ed 2. 148. 1762, nomen illegit. (cites S. suffruticosum Loefl.; see Steyermark 1972). S. spinosa aeree Obs. Bot. 45. 1791, not Jacq. 1760, nomen nudum. S. spinosa Cham. & Schl., Linnaea 3: 340. 1828. Tardavel e (L.) Standley, Contr. U.S. Natl. Herb. 18: 122. B. je 1 Brandegee, Univ. Calif. Publ. Bot. 10: 416. 1924. TYPE: qe Brandegee 9223 (CAL, en) Herbs to 0.5 m tall, the stems arising from a taproot, erect, decumbent or prostrate, often the branches restricted basally and strict, occasionally with short branches above, angular, smooth, glabrous or puberulent, the nodes well spaced. 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Leaves 5-8 at some of the nodes, of variable length, some 3 times longer than others, linear lanceolate to linear, to 6 cm long, to 0.8 cm wide, acute at the apex, attenuate at the base, the costa prominent beneath, the lateral veins few and scarcely distinguishable, papyraceous, stiff, smooth, the costa often with scat- tered erect calcareous bodies, the margin scabrid, revolute; petioles absent or seemingly present as the blades attenuate basally, the setiferous sheath cupuli- orm, 2-4 mm long, the setae 15—20 linear, as long as or longer than the sheath. Inflorescences subtended by 4-8 leaves, terminal or axillary in upper axils, the heads sessile, 1.5-2.0 cm in diam. with slender, white bristles among the flowers. Flowers with the hypanthium oblong, the calycine cup ca. 0.3 mm long, petaloid, glabrous, the calycine teeth 2, subulate or linear oblong, 1.3-1.8 mm long; corolla white, cupular, 1.5-2.0 mm long, strongly constricted basally, the lobes 4, lan- ceolate or oblong, to 1 mm long; stamens 4, exserted, the anthers oblong rotund, 0.3-0.5 mm long, the filaments slender, to 0.5 mm long, the style 1.3-1.7 mm long, the stigmatic lobes minute. Fruits pyriform or obovate oblong, the 2 carpels to 2.5 mm long, somewhat compressed, smooth, shiny, glabrous except pubescent to arachnoid pubescent at the apex, the 2 awns erect, shorter than the body of the fruit, pubescent. This species is widely distributed throughout tropical America. For an elab- orate discussion of ‘‘Borreria densiflora DC. (as) the earliest legitimate name for the species now passing as B. spinosa," see Steyermark (1972) p. 814. CAS DEL TORO: Junction Ríos Changuinola and Teribe, Lewis et al. 923 m MO, US). Isla Colón, Wedel 2946 (MO). CANAL ZONE: Fort Lorenzo, Baker & Oliver 1043 (GH, K, MO, UC, US). Panamerican Highway, La Chorrera, D' Äre 'y 9452 (MO). Toro Point, Fort Sherman, os 4324 (MO). Fort Sherman, Dwyer 7191 (MO). Between Rodman irs Base and Chorrera, Nowicke et al. 3606 (MO, SCZ, UC). Fort Amador, Tyson 2017 (MO). Fort Sherman, Tyson 2248 (MO). BARRO COLORADO ISLAND: Croat 4407, 6984, 7394, 12609, 12884 (all MO); p 421 (MO). cuiRiQUÉ: Methodist Camp near Nueva Swissa, Croat 13525 (MO). Burica Peninsula, 12 mi W of Puerto Armuelles, 400— 00 m, c Creat 22181 (MO). Llan nos Francia, 4 mi from Boquete. Dwyer & Hayden 7612 (MO). Cerro N : Mouth of Río Piedras. Lewis et al. 3176 6 (MO). HERRERA: Salinas de Chitre on Bahia Parita, ran 9683 (MO). Between Río Bayano Dam and Canitas, D'Arcy 9400 (MO). PANAMA: Isla Taboga, Allen 1272 (MO). Punta Chame, D'Arcy 10234 (MO). San José Island, — 838 (MO). 2 km W of El Llano, Nee 7958 (MO). Nuevo Emperador, Troetsch 8 (MO). SAN BLAS: Mandingo EU Duke 14824 (MO). vERAGUAS: Mouth of Río Concepción, Lewis et al. 2766. (COL. MO, UC, N). m NW of Atalaya, Nee 8234, 8243 (both MO). 2. Borreria laevis (Lam.) Griseb.,? Goett. Abh. 7: 231. 1857.—Fic. 12. Spermacoce laevis Lam., Tabl. is Méth. 1: 273. 1791. rvPE: Santo ges W.I., not seen. Spermacoce echioides H.B.K.. . Gen. Sp. Pl. 3: 269. 1820. TYPE: not s Tardavel laevis (Lam.) ido. Cn U. S. Natl. Herb. 18: 122. 1916 Spermacoce guianensis Brem., Recueil Trav. Bot. Néerl. 33: 714. 1936. type: Surinam, Rombouts 5 (Utrecht?) Herbs, upright or scandent, to 2⁄4 m in length, the stems smooth or rimose, glabrous to glabrate, often retrorse pilose, the nodes well spaced. Leaves ovate elliptic to elliptic, 1.5-7.0 cm long, 0.6-3.0 cm wide, acute at the apex, rounded at the base, occasionally attenuate acute, often inequilateral, the costa promi- nulous beneath, the lateral veins 5—6, strongly arcuate, chartaceous, concolorous, ° For more complete synonymy see Steyermark (1972) p. 819. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 51 FiGuRE 12. Borreria laevis (Lam.) Griseb.—A. Habit ( x1⁄4).—B. Fruit (x8). [After Duke 4902.] 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 scabrous above and beneath, glabrous above, glabrate beneath; petioles to 1 cm long; stipules connate, the base of the petiole adnate to the sheath, the latter cylindrical, appearing to be subnodal, to 4 mm long, puberulent to villosulose, the awns several, narrowly subulate, 2-7 mm long. Inflorescences axillary and terminal, globose capitate, 0.6-1.0 cm in diam. Flowers with the hypanthium glabrous or puberulent, oblong, ca. 1.5 mm long, the calycine teeth 4, unequal, subulate, to 0.5 mm long, ciliolate; corolla cylindrical, 2.5 mm long, the lobes 4, longer than the tube, pilose within; stamens 4—5, the anthers subrotund, ca. 0.3 mm long, the filaments 1.3 mm long; style to 2 mm long, the stigmas rotund capitate, ca. 0.25 mm in diam. Fruits oblong, 1-2 mm long, glabrous to minutely pubescent, dehiscent, one carpel bearing 3 of the 4 calycine lobes; seeds oblong, 1.0-1.5 mm long, 0.75 mm wide, the transverse grooves prominent. Borreria laevis is a common weed in Central America and in most of South America. It also occurs in the West Indies. ''Sab Tutu” (Cuna, Duke 14439). CAS DEL TORO: Isla Bocas del Toro, Durkee 71-155 (MO). Without other jo peti ‘hs (MO). Water Valley, Wedel 595, 99] (both MO). Isla Colon, Wedel 2833 (MO). CANAL ZO Amador, D’Arcy 9328 (MO). Between Rodman Marine Base and Chorrera, Nowicke et PE 585 a ). Ancón voy us 107 (MO). Fort San Lorenzo, Tyson & Blum 3701 (MO). Ancón Hill. Woodson et al. 1325 O). Salamanca Hydrographic Station, Woodson et al. 1617 (MO). BARRO COLORADO ISLAND: Tos 4045, 4188, 4644, 5796, 6078, 6946, 7047, 7060, 7064, 7322, 9199 (all MO); Ebinger 48 (MO); Hayden 81 (MO); McDaniel 5025 (MO); Netting 49 (F); Shattuck 478 (MO); Starry 4 (both MO). CHIRIQUÍ: N of Cerro Punta, Croat 10448, 10471, 10508 (all MO). 1 mi N of E Hato del Volcán, Croat 10703 (MO). Road to Río Serano, eee et al. 2093 (MO). Boquete to 3 mi N, 3300-4200 ft, Lewis et al. 630 (GH, K, MO, NY, US). W of Puerto Armuelles, Liesner 82 (MO). Puerto Armuelles, Liesner 140 (MO). Upper Rio Uere Viejo, Monte Lirio, Seibert 247 (MO). cocrÉ: Behind Club Campestre, Duke 13273 (MO). NE of EI Valle, pes et al. 1698 (MO, UC). Between Las Margaritas and El Valle, Woodson et al. 1288 (MO). COLON: Ridge behind Garotte, D'Arcy 9332 (MO). Achiote, Zimmerman, Folsom et al. 3322 (MO). DARIEN: Boca de Cupe, Allen 984 (MO). El Real, Burch et al. 1076 (COL, MO, UC). S of El Real, Duke 4902 (MO). From Pucro o Río Pucro, Duke 5375 (MO). 10 mi S of El Real on Río Pirre, Duke 5444 (MO). 1—4 mi N of Pucro, Duke 13019 (MO). Río Pucura between Cerro Mali and Cerro Tacarcuna, near Río Pucro base camp, 640 m, Gentry & Mori 13951 (MO). Paya, Stern et al. 259 (MO). Los sANTOS: Loma Prieta, Duke 11863, 11895 (both MO). 25 mi SW of Tonosi, Lewis et al. 2983 (MO, UC). PANAMA: Near Jenené, Mon 3831, 3847 (both MO). Piria, ca. 150 m, Duke 14439 (MO). x mi E of El Llano, Tyson 1723 (MO). Road to Cerro Campana, 7.6 km from Panamerican Highway, Witherspoon & Witherspoon 8366 a. SAN BLAS: SE of Puerto Obaldía, p 16736 (MO). Puerto Obaldia, Gentry 1555 (MO). 3. Borreria latifolia (Aubl.) Schum. in Mart., Fl. Bras. 6(6): 61. 1888. Spermacoce latifolia Aubl., Hist. Pl. Guiana 55. 1775. Type: Cayenne, French Guiana, Aublet, not seen. Borreria perrottetii DC., Prodr. 4: 548. 1830. TYPE: not indicated. Tardavel latifolia (Aubl.) Standley, Contr. U.S. Natl. Herb. v" 122. 1916. Herbs, often prostrate, often erect and more than | m tall, the stems villose, porcate, the nodes well spaced. Leaves with the blades often spreading, disposed at right angles to the stem or deflexed, trapeziform, ovate oblong, oblong rotund or oblong, 1.5-6.0 cm long, 0.7-3.2 cm wide, acute at the apex, the costa prom- inulous beneath, the lateral veins ca. 5, immersed above, prominulous beneath, chartaceous, scabrous, often with white bulbous-based hairs above, drying yellow green or flushed with red; petioles absent or to 0.5 cm long; stipules with the sheath subtriangular in outline, often split as the inflorescence expands, the setae to 5 mm long. /nflorescences with the flowers disposed in rotund heads in the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 53 axils, these usually to 1 cm wide. Flowers with the hypanthium short, ca. 1 mm long, densely appressed white villose to glabrous (fide Standley), the calycine cup much shorter than the lobes, the lobes 4, lanceolate to ovate, to 1.5 mm long, acute, villose; corolla white, the tube funnelshaped, 2.5-3.0 mm long, glabrous, the lobes 4; stamens 4, the anthers oblong or ovate, 0.35-0.50 mm long, the filaments at least twice as long as anthers, attached at the mouth; style minute, knoblike or conspicuously bifid, the ovarian disc scarcely developed. Fruits ob- long or turbinate, to 3 mm long, densely villose, opening apically and often ex- panding into 2 valves, each valve circular in top view, the smooth light-colored interior of the valve revealed; seeds oblong, ca. 2 mm long, smooth, drying orange, shiny, reticulate, the calyx tending to wither at dehiscence. Borreria latifolia is widely distributed in tropical America. It also occurs in the West Indies. ‘‘Cansa-mozo’’ (Standley, 1928). Steyermark (1972) has 2 varieties and several forms which do not hold up as far as the Panamanian material is concerned. In the synonomy of B. latifolia I have included only a minimum of synonomy as compared with Steyermark’s many synonyms. L ZONE: Panamerican Highway near La Chorrera, D’ sh : 9432, 9453 (both MO). Ft. Kobbe, Duke 3905 (MO). E of Gatun Locks, Duke 4306 (MO). 5-9 mi N of Gamboa, Gentry 2408 (MO). Chiva-Chiva Trail, Tyson 1370 (MO). Curundu, Tyson 6288 (MO). Miraflores Lake, White 253 (MO). BARRO COLORADO ISLAND: Croat 5982, 6461, 12816, 12824 (all MO). CHIRIQUi: Palo Alto, E of Bo- quete, Stern et al. 1042 (MO). COLON: Spr 7-8 mi E of cement plant, Blum & Tyson 497 (MO). 0.7 mi NE of Rio Piedras, Nee & Mori 3641 (MO). PANAMA: Arraijan, Duke 4681 (MO). Cerro Jefe, Gentry 2138 ge San José Island, ape st Ee 1085 (both MO). Goofy Lake, Lewis et al. 229 H, M US). Chimán, Lewis et al 6 (MO, PAN). El Llano-Cartí Road, 5 km N of Panamerican Highway, Nee 7879 (MO). VERAGUAS: 1-2 mi above Santa Fé, Gentry 3063 (MO). 4. Borreria ocimoides (Burm. f.)!° DC., Prodr. 4: 544. 1830. Spermacoce ocimoides Burm. f., Fl. Ind. 34. 1768. Type: Tropical Asia. Borreria parviflora G. F. W. Mey., Prim. Fl. Esseq. 83. 1818. TYPE: not seen B. tampicana DC., Prodr. 4: 544. 1830, in part. rvPE: Mexico, Berlandier 37 bis (?B, not seen; photo, MO B. pringlei Wats., Proc. Amer. Acad. Arts 25: 152. 1890. T Watson 2464 not seen. Tardavel acymoldbet (Burm.) Heirn, Cat. Pl. Welw. 2: 503. Terrestrial herbs, often procumbent, rarely epiphytic on tree roots, to 4% m tall, the stems simple or branched, angular, delicately alate, glabrous, the nodes well spaced. Leaves lance ovate or ovate trapeziform, 1.0-2.5 cm long, to 1 cm wide, acute or occasionally obtuse at the apex, basally acute to cuneate, the costa immersed, the lateral veins 4—5, strongly arcuate ascending, concolorous, sca- brous above and beneath; petioles absent or to 0.15 cm long; stipules connate and adnate to the base of the petiole, the sheath cupshaped, scarcely measurable, to 1.5 mm long, the setae 6-10, to 2.5 mm long, usually curled. /nflorescences terminal and axillary, sessile, capitate, 0.2-0.8 cm wide. Flowers minute, the calycine lobes 2—3(-4), erect, subulate, ca. 1 mm long; corolla white, to 1 mm long, the lobes 4, acute; stamens 4, the anthers minute. Fruits capsular, the ? For more extensive synonymy see Steyermark (1972). Only these names have been used for PNG s material. 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 valves 2, splitting and separating, the persistent calycine lobes glabrous or pu- berulent; seeds oblong, ca. 1 mm long, brown, reticulate. Borreria ocimoides is a widespread weed in the tropics. ANAL ZONE: Military Road, K-9, Ebinger 513 mici Gaillard Highway, /? km NW of Summit mis Nee 6929 (MO). Salamanca Hydrographic Station, Woodson et al. 1619 (MO). BARRO COL- ORADO ISLAND: Croat 8268, 8303, 8826, 10306. 11339, "2277. 12582, 12938, 13152, 17037 (all MO); Shattuck 47, 497 (both MO). cHIRIQUí: Districto Guanabano, along Quebrado Quanabano, Croat 22510 (MO). due 3300-4200 ft, Davidson 597 (MO); Lewis et al. d (GH, K, MO, NY, UC US); Partch 69-76 (MO); Woodson ^ Schery 729 (MO). Cerro Punta, 6000 ft, * son 7128 (MO). cocLÉ: El Valle, yes 2825 (MO). La Mesa, Gentry 6826 (MO). DARIEN: El Real, Río Tuira, Stern et al. 774 (MO). PANAMA: Between Pus and Chepo, Dodge et al. 16698 a Hacienda la Joya, Dodge et al. 16891 (MO). Cerro Campana, road near summit, Duke 6000 (MO). Confluence Río Pacora and Río Corso, ca. 450 m, Duke 11957 (MO). Goofy Lake, Dwyer 7137 (MO). San José Island, oy eh 902 (MO). Cerro Campana, Lewis et al. 3103 (COL, DUKE, F, K, MO, NY, UC). W slope o Campana 2500 ft, Tyson 7 al. 2349 (MO). 2 mi S of Tocumen Airport, Tyson & Clewell 5929 (MO). 5. Borreria pumilio Standley, Ann. Missouri Bot. Garden 27: 342. 1940. TYPE: Panamá, Allen 822 (F, holotype, isotypes MO, NY, ). Herbs to 6 cm tall, unbranched or with 1—2 branches, the stem with appressed white hairs. Leaves sessile, linear oblong, 4-6 mm long, the costa and veins not visible or the lateral veins 4—5, scarcely visible beneath, arcuate, granulose above and beneath; stipular sheath ca. | mm long, the setae several, to 1 mm long. Inflorescences capitate, terminal or in the uppermost axils, the heads ca. 0.5 cm in diam., the outermost bracts linear, to 9 mm long. Flowers with the corolla white, the tube somewhat longer than the calycine lobes. Fruits subrotund, each mericarp oblong, ca. 0.8 mm long, the persistent calycine lobes 1.0-1.5 mm long, glabrous to pubescent, usually slightly longer than the body of the fruit. Borreria pumilio is known only from Panama. Because of the scant material, the description of flowers is taken from Standley's diagnosis. CANAL ZONE: Fort Kobbe, Curatella Savanna, Duke & Mussell 6573 (MO). CocLE: Nata, Allen 822 (F, MO, NY, US). 6. Borreria suaveolens G. F. W. Meyer, Prim. Fl. Esseq. 81. 1818. TYPE: Surinam, in Arenosis Continentis Circa Arouabischkreck, Meyer, not seen. tenella Cham. & Schlecht., Linnaea 3: 317. 1828. rype: Brazil, Sello, not seen. 88. tenella var. suaveolens (Meyer) Schum., Mart., Fl. Bras. 6: 56. 18 . capitata var. suaveolens (Meyer) Steyermark, Mem. N.Y. Botanical Garden 23: 825. 1972. is Perennial Aerbs, smaller plants somewhat ericoid in habit, the stems often arising from a taproot, frequently unbranched to the apex, densely pubescent to glabrate, the nodes well spaced. Leaves sessile or subsessile, 2(—6) at a node, the shorter leaves linear or narrowly ovate lanceolate, 2-6(-8) cm long, to 0.25(-0.5) cm wide, occasionally falcate, acute at the apex, rounded or obtuse at the base, the costa prominent beneath, the lateral veins ca. 4, strongly ascending, usually ob- scure, stiffly papyraceous, usually scabrid, the margins revolute; stipules with or without a distinct sheath, the latter to 2 mm long, with 4-8 setae on each side, these 3-5 mm long, rubescent. /nflorescences globose capitate, the heads ter- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 55 minal, axillary in the 2 superior nodes, to 1 cm in diam., the leaflike bracts reflexed in older heads, the bracts densely aggregated in the head, linear subulate, 3.5 mm long. Flowers with the corolla funnelshaped or subcylindrical, 3.5 mm long, glabrous outside; stamens 4, the anthers oblong, ca. 1 mm long, the fila- ments 1.2-1.5 mm long, attached above the middle of the tube; style slender, 2— 3 mm long, the stigmatic area minute, capitate, ca. 0.2 mm long. Fruits oblong, 1.5-2.0 mm long, the calycine lobes often persistent, as long as the body of the fruit, the latter splitting at the pubescent apex; seeds not sulcate. Borreria suaveolens is well distributed throughout tropical America; it is a common species in savannas from low to relatively high elevations. CANAL ZONE: Río Indio, Allen 17395 (MO). Drowned Río Azote Caballo, Dodge et al. 16833 (MO). Río Indio, Dodge & Allen 17377 (MO). Ancón Hill, Standley 26206 (MO). cHIRIQUÍ: Volcán de Chiriquí, Allen 989 (MO). S of Boqueté, Allen 4691 (MO). 1 mi N of El Hato de Volcán, Croat 10700 (MO). Cerro Colorado, 20-28 mi from San Félix, 1200-1500 m, Croat 33385 (MO). Boquete, D'Arcy 9773B (MO). El Baru above Boquete, 1200-1800 m, D'Arcy 9928 (MO). A flow a El Hato and Bambito, D’Arcy 10025 (MO). Boquete, N 682 (MO). 3 km NE of El Hato del Volcán, 1500-1800 m, Davidse & D'Arcy 10383 (MO). Volcán, Duke 9153, 9184 E MO). Llanos Francia, near Boquete, Dwyer s.n. (MO). 4 mi S of Bambito, [js or & Correa A. 2700 (MO). Volcan arü, ca. 2000 m, Mori & Bolten 7385 (MO). Boquete, Partch 69-37 (MO). Llanos Francia, Stern et al. 1155 (MO). ed Punta, 6000 ft, Tyson 7128 (MO). Río Chiriquí Viejo Valley, El Volcán, White 209 (MO). AMÁ: Río Pacora and Chepo Highway, Duke 5925, 5926 (both MO). Cerro Campana, Ebinger 364 (MO); Sullivan 436 (MO); Tyson et al. 2383 (MO). Río Tartaré, Woodson & Schery 1009 (MO). vERAGUAS: Soná, Allen 1050 (MO). 13. BOUVARDIA Bouvardia Salisbury, Parad. Lond. pl. 88. 1807. TYPE: B. triphylla Salisbury. Aeginetia Cav., Anales Ci. Nat. 3: 129. 1801; Ic. 6: 51. 1801, not Aeginetia L. (1753). Shrubs or herbs. Leaves opposite or 3—4(—6) per node, membranous; stipules interpetiolar with a basal sheath. Inflorescence terminal, cymose. Flowers di- morphic, calycine lobes 4, persistent; corolla tubular or salverform, the lobes 4, valvate; stamens 4, the anthers linear or oblong, in pin flowers included and subsessile, in thrum flowers included and with filaments distinct and exserted; styles of 2 lengths, usually slender, the ovary 2-celled, the ovules numerous, attached peltately to a complete septum. Fruits capsular, globose or oblate, de- hiscing at first loculicidally, then septicidally; seeds numerous, vertically imbri- cate, winged, the wing entire. Bouvardia, a genus of about 50 species, ranges from Mexico to Panama. Literature: Blackwell. W. H. 1968. Revision of Bouvardia. Ann. Missouri Bot. Gard. 55: 1. Bouvardia glabra Polak., Linnaea 41: 565. 1877. TYPE: Polakowsky 337 (F, frag. holotype, photo F, US).—Fica. 13 B. glabra var. gracilis Polak., Linnaea 41: 566. 1877. TYPE: Polakowsky 172 (not seen; F, photo). B. glabra var. obtusa Loesner, Verh. Bot. Ver. Brandenberg 65. 106. 1923. TYPE: Seler 2920 (not seen). 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 SN S ` N [2 FiGuRE 13. Bouvardia glabra Polak. Flowering branch (x.94). [After Mori & Bolton 7356.] Shrub to 1% m tall, the branchlets smooth, glabrous, drying tan. Leaves tending to persist at the apex of the twiglets, ovate lanceolate to widely elliptic, 3.5-5.0 cm long, 1-2 cm wide, acute at the apex, acute at the base, the costa subplane above and beneath, the lateral veins 3—6, strongly ascending and ar- cuate, glabrate, chartaceous, discolorous, brown above, greenish beneath; peti- oles slender, to 3 mm long; stipules triangular with 3-5 subulate processes, the median process more elongate, often longer than the body. Inflorescences 3-7- flowered. Flowers pedicellate, the pedicels to 1.5 cm long; hypanthium glabrous, 1980] DWYER-—FLORA OF PANAMA (Family 179. Rubiaceae) 57 drying dark, the teeth 4, falcate subulate, unequal, 3-7 mm long; corolla tube cylindrical, to 2 cm long, to 0.2 cm wide in the middle, glabrous outside, sparsely pilose above the middle, the lobes ovate oblong, to 5 mm long, to 4 mm wide, obtuse; anthers (? all) subsessile, 3 mm long, ca. 1 mm wide, obtuse; style (only longer styles seen) to 2.3 cm long, the stigmas to 4.5 mm long. Fruits on pedicels to 1.7 cm long, to 0.7 mm wide, glabrous, compressed rotund, to 0.8 cm long, glabrous. Bouvardia glabra ranges from southern Chiapas, Mexico, to Guatemala, Costa Rica and Panama. CHIRIQUí: W side of Volcán Baru, oak forest, 2000-2500 ft, Mori & Bolten 7356 (MO). 14. CALYCOPHYLLUM Calycophyllum DC., Prodr. 4: 367. 1830. LECTOTYPE: C. candidissimum (Vahl) C. (Macrocnemum candidissimum Vahl). Enkylista Benth. Hooker's J. Bot. Kew Gard. Misc. 5: 230. 1850. TYPE: E. spruceana Benth. Trees. Leaves petiolate, the stipules small, deciduous. /nflorescences pa- niculate, terminal, corymbose, ebracteolate or bibracteolate. Flowers in bud en- closed by membranous bracts, sessile or subsessile, the hypanthium oblong, the calycine cup truncate, sometimes partly expanded into a large foliaceous stalked blade, the corolla with the tube short, funnelform, the mouth villous, the lobes 6-8, imbricate or contorted, 1 lobe exterior, the stamens exserted, the anthers versatile, the filaments subulate, the style slender, the stigmas 2, narrowly oblong, the ovules disposed in 2 cells, numerous, imbricate, the placentas adnate to the septum. Fruits capsular, oblong cylindrical, 2-celled, septicidally bivalvate, the valves coriaceous; seeds numerous, imbricate, usually minute, the testa mem- branous, produced at each end as an elongate wing. Calycophyllum has approximately 6 species in the West Indies and 1 in Central America and South America. 1. Calycophyllum candidissimum (Vahl) DC., Prodr. 4: 367. 1830.—Fic. 14. Macrocnemum nd Vahl, Symb. Bot. 2: 3. 1791. TYPE: not se Mussaenda candida P in Lam. Encycl. Mar Bot. aon 4: 395. 1797. TYPE: not seen. Mussaenda candissima 1 (Vahl) R . & S. Syst. Veg. 5: 1819. Trees to 15 (-40) m tall, the bark reddish brown, often exfoliate, the twiglets smooth, glabrous, rubescent. Leaves oblong rotund, obovate oblong and occa- sionally falcate, to 15 cm long, to 7 cm wide, obtuse at the apex, cuspidate, the cusp usually triangular, to 2.5 cm long, ca. 0.3-0.5 cm wide in the middle, basally attenuate cuneate or widely cuneate, often slightly inequilateral, the costa plane or immersed above, prominulous beneath, the lateral veins 8—10, arcuate, the intermediate veins parallel and branching near the margin, the intervenal areas spreading reticulate, papyraceous, often drying red brown, glabrescent or villos- ulose above and beneath, rarely with minute white raphides; petioles 3 (-4) cm long, slender, to 1.3 mm wide; stipules deciduous, oblong, ca. 2.5 cm long, obtuse, scarious, rubescent, villose. Inflorescences terminal (sometimes appearing axil- 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 QY / UNIS NY d L^ I None di APER D 7 4 YA X< = ZEB, = < See S I A aY L^ NZ Figure 14. Calycophyllum candissimum (Vahl) DC.—A. Habit (x?/:).—B. Opened flower and leaf (x2).—€C. Ovary cross section (x4). [After Miller 1962. | lary), paniculate, rarely shorter than the uppermost leaves, often flat topped or convex at the apex, to 20 cm long, usually with 3-4 peduncles arising from the tip of the twiglet, these strongly ascending, each with several strongly ascending branches, the flowers disposed in compound dichasia, the median flower of each cluster sessile. Flowers with the hypanthium turbinate, ca. 3 mm long, ca. 1.5 mm wide, truncate, glabrescent or villosulose with farinose glandular dots inter- spersed, the calycine cup less than | mm long, the teeth absent or often one expanded into a petiolate lamina, obovate spatulate, the blade to 3.3 cm long, to 2.5 cm wide, petaloid, glabrous, venose, the claw slender, ca. 0.2 cm wide; corolla white, the tube compressed cylindrical, ca. 3.3 mm long, densely villose within 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 59 above the middle, the lobes 5, strongly reflexed, oblong, 3—4 mm long, obtuse, glabrous, venose; stamens 5, the anthers oblong, ca. 1.5 mm long, subversatile, the filaments slender, elongate, to 3 mm long, exserted, attached at the mouth; ovarian disc pulviniform, pubescent, 0.75-1 mm long, 0.4—0.6 mm wide, the style 3.3-5.3 mm long, ca. 0.15 mm wide, the stigmas 2, slender, ca. 2.2 mm long, curved or erect, puberulent, the ovary rotund in cross section, 2 loculate, the septum moderately thick, the ovules numerous. Fruits sessile or subsessile, ob- long, 0.5-1 cm long, truncate or rounded apically, usually obtuse or rounded at the base, lignose, villosulose, often with minute white raphides, drying tan, with ca. 8 delicate ribs, splitting from the apex, the margins of the valves wide; seeds to 5 mm long, the wing acuminate at each pole. Calycophyllum candidissimum is widely distributed in Mexico and Central America, ranging south to Colombia and Venezuela. It also occurs in the West Indies. The hard, fine-grained wood has a variety of uses. In Panama it is used for charcoal. Standley (1928, p. 355) lists 5 common names: *'Alazano," ‘‘Gua- yabo,” ‘“‘Harino, Madroño,” and ‘‘Salamo.’ °° 66 CANAL ZONE: Albrook, Blum 2086 (MO). Chagres River, Boyd-Roosevelt Highway, Blum & di x as Madden Dam, Dodge et al. 16500 (MO); Dwyer 8380 (MO); Lewis et al. 35 (F GH, , UC, US). Gatün, Johnston 1736 (MO). Boy Scout Camp Road, Madden Dam, D Mori p (M | km S of Madden Dam, Nee 8905 (MO). Rodman Naval Ammunition Depot, 1 mi W of “hd rae ae et al. 12891 (MO). BARRO COLORADO ISLAND: Aviles 60 (F); Croat 8661, 12905 (both MO). cocLe: El le, Aguilar (MO). Rio Mata Ahogado, Allen 126 (MO). Cerro Pilon, Lal- and junction with Tuira, Gentry & Mori 13483 (MO). Campamento Buena Vista, Rio Chucunaque, Stern et al. 966 (MO). Piriaque Camp, Tyson et al. 4761 (MO). Los SANTOS: Tonosi, Duke 12486 (MO); Oliver et al. 3544 "ses Stern et al. 1816 (MO). PANAMA: Pacora, Allen 1/22 (MO). Isla El Rey, Correa 113 (MO). Cerr Azul, Nabe 13007 (MO). Old Panama, Cooper 339 (MO); Greenman & Greenman 5142 (MO). Isla Espiritu Santo, Duke 10437 (MO). Bejuco, Duke 4581 (MO). Saboga Island, Miller 1962 (MO, US); i a Loftin 5125 (MO). Chiva-Chiva Road, Smith & Smith 3340 (MO). W of Chepo, Tyson 5369 (MO). 15. CEPHAELIS Cephaelis Swartz, Prodr. 3: 45. 1788. Nomen conserv. TYPE: C. muscosa (Jacq.) Swartz type cons. Carapichea Aubl., Hist. PI. Guiane 1: 167. 1775. Nomen rejic. TYPE: C. guianensis Aubl. Tapogomea MARE Hist. Pl. Guiane 1: 367. 1775. Nomen Pu i T. violacea Aubl. Evea Aubl., Pil, Guiane 1: 103. 1775. Nomen rejic. . guianensis Au Ipecacuanha yates Diss. Pl. Brazil 44. 1810. TYPE: /. d Arruda. Shrubs, trees, or occasionally woody herbs. Leaves often glabrous and with many lateral veins; stipules free or connate, frequently persistent. /nflorescences axillary or terminal, capitate, the head often branched, subtended by an involucre of bracts, the bracts often large, often highly colored, free or connate, becoming smaller and more concave adaxially. Flowers usually sessile; hypanthium short, the calycine cup short, truncate or dentate, often irregularly so, the teeth (4—)5 (-7); corolla usually white, the indumentum variable, the tube funnelform or salverform, usually expanded basally, the lobes usually 5, valvate in bud; ovary 2-celled, the septum thick, a single ovule in each locule arising from the base of 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 67 the ovary. Fruits fleshy, the 2 pyrenes smooth or costate, the endocarp usually sulcate on the inner face. Cephaelis occurs throughout the tropics of the New and Old Worlds. It has about 200 species. I have elected to separate Cephaelis from Psychotria although there are reasons for reducing the former to the synonomy of the latter, as J. Steyermark maintains. In Panama most of the species have capituliform inflores- cences with large colored bracts which allow for easy allocation to Cephaelis. Standley & Williams in the Flora of Guatemala (1975) sum up the traditional stand: '' or practical purposes .. . it constitutes a most convenient segre- gate.” a. Inflorescences spiciform cylindrical, the rachis 5-12 cm long -------- C. Mn eue aa. Inflorescences mostly rounded capitate, not spiciform ri the ir or flowering capitulum less than 5 c ieee ng. b. i nai term ane subtended by 2 large flaring or mottled bracts d distinct in e and texture from the innermost bracts; outer bracts 2-5 cm long. Heads with the outermost bracts not mottled, uniformly red, A on only orange or yellow, rarely white. e. Plants densely hairy; heads on erect peduncles ...... 20. C. tomentosa ee. Plants glabrate; heads on nodding or erect m cles f Peduncles nodding; leaves to 29 cm long, papyraceous NE Ae aE A aa R3 . C. correae ff. Peduncles erect, the largest leaves to 15 cm long, coriaceous ER _... 10. C. elata dd. Heads with outermost bracts obviously mottled, of various s colors AEEA, EPEA C. vultusmimi olor cc. Heads of flowers with outermost bracts more or less similar in texture and co to those within, less than 2 cm long. g. Heads of flowers disposed as a rotund capitate or subglobose ma ubs Be less than 1 m tall; mass of flowers 3-5 cm wide; ia 4—7 cm i. Peduncle ca. 4 cm long |... 17. C. nana ii. Peduncle 0.7-2.0 cm long - _ 8. C. discolor hh. wu ca. 2 m tall; mass of flowers to 2.5 cm P peti- s 0.3-1.5 cm long . 12. C. glomerulata gg Heads of a ack or elliptical, not disposed as a rotund capitate or rotund m j. Heads , ca. 0.5 cm long ---------------- _. I5. C. kennedyi ï. Heads more uo 0.5 cm lon k. Leaves with 2 submarginal veins paralleling the margin; stem rhizophorous ______--------------------- === 13. C. insueta kk. Leaves with 1 vein or no vein paralleling the margin; stem not rhizophorous. l. Leaves drying red or brown beneath, the lateral veins (20—)30—-38. m. ox vade rs to 2.2 mm long _ 9. C. dressleri nn. pera 2.5.3.0 mm id "TP 19. : panchocoénsis mm. Petioles of largest leaves | ca. 4 cm long |... MERE 2; C; bristanii Leave s not drying red T brown beneath (except C. dres sleri), s lateral veins o. Peduncle prese n p. Peduncle terminated by 2 large bracts and 3 eee ae each branch terminated by a ca- pitulum of flowers; lateral veins ca. 4 ne E . C. dichroa — = 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 61 pp. Peduncle and general inflorescence not as above; lateral veins 13-25. Leaves sessile or subsessile, auriculate ries; corolla 10-14 mm long ...... 9. C. dressleri rr. Bracts not glanduliferous within; corolla ca. 30 mm long ------------ suan 2 kas 19. C. panchocoénsis Leaves with distinct petioles 0.3-1.5 cm ane stipules to 0.3 cm long. nflorescence with heads 1.5-3.0 cm wide; calycine teeth to 1. x mm long, equal; corolla tube c 0 mm long ------------ C. glomerulara ss. Inflorescence with heads to 0 long; calycine teeth absent or dis- posed as mere points or obviously unequal, to 0.3 mm long; corolla tube to 3.2 mm long |... 15. C. kennedyi oo. Peduncle absent 18. C. panamensis qq. Kel bb. de eid. axillar Leaves with lateral veins 6— u. Stipules membranous ok conspicuous marginal subulate fimbriae; plan rhizomatous 14. C. ipecac wank uu. Stipules subcoriaceous or chartaceous without marginal fimbriae; plant not rhizomatous. eaves succulent, the lateral veins si d: stipules to 1 cm lon inflorescences with peduncle to 1 c 11. C. p vv. Leaves not succulent, the lateral deos conspicuous; stipules to 7 mm long; inflorescences with peduncle absent. Leaves 1.6-5.0 cm wide; inflorescences 1.0-1.5 cm wide l. C. axillaris ww. Leaves 7-9 cm wide; inflorescences ca. 2 cm wide ` UT 16. C. latistipulata tt. "uui with lateral veins 15-38. ipules more than 1 cm lon ng: inflorescences with heads 1.5-2.2 cm wide. Leaves with uem veins evanescent; blades succulent---------------- . C. camponutans Leaves with lateral veins obvious; blades not imcculedit -. 5. C. croatii XX. aye less than 1 cm long; inflorescences with heads ca. l y wide ls. . C. bristanii 1. Cephaelis axillaris Swartz, dans Veg. Ind. Occ. 45. 1788. TYPE: not seen. ot Psychotria axillaris Willd. 1798. Tapogomea axillaris (Swartz) Poir., B Méth. Bot Suppl. aC ee 1806. Evea axillaris (Swartz) Standley, Contr. U.S. Natl. Herb. 18: 1 sychotria aubletiana Steyermark. Mem. New York Bof. Lh 23: 694. 1972. New name for Cephaelis axillaris Swartz Shrubs to 11⁄ m tall; branches few, usually arising near the middle of the stem, terete, smooth, glabrous, often attenuate toward the nodes, dilated at the node above the base of the stipule. Leaves usually salicoid, occasionally oblong to obovate oblong, occasionally falcate, 2-13 cm long, 1.6-5.0 cm wide, at least 2 times longer than wide, occasionally wider below the middle, the costa plane or grooved above, prominulous to prominent beneath, golden villosulose or gla- brous, the lateral veins ca. 8, arcuate, prominulous, the intervenal areas spreading reticulate; petioles 1.5-2.0 cm long, slender, to 0.8 mm wide or stout and to 1.8 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 mm wide, glabrous or minutely pubescent; stipules briefly connate, oblong, to 7 mm long, to 5 mm wide at the base, retuse at the apex or split almost to the base, glabrous. Inflorescences axillary or terminal with short twiglets subtending, ses- sile, the flowers conglomerate in a rotund mass, 1.0-1.5 cm in diam.; bracts purplish, 4—6, ovate, shorter than the flowers, obtuse. Flowers with the hypan- thium short, the calycine cup ca. 0.3 mm long, the teeth 5, unequal, obtuse, to 0.6 mm long, petaloid, glabrous except marginally, with a few oblong glands on the inside of the cup, to 0.2 mm long; corolla white, the tube cylindrical, some- what inflated, ca. 5 mm long, to 1.8 mm wide, petaloid, glabrous outside, villose within on the upper 2⁄4, the lobes 5, oblong, glabrous; stamens 5, the anthers oblong, ca. 2.2 mm long, 0.5 mm wide, obtuse, the filaments ca. 0.15 mm wide; style ca. 2.2 mm long, shorter than the tube, the stigmas upright, ca. 0.3 mm long. Fruits sessile, oblong rotund, 3.5—4.5 mm in diam., each pyrene compressed hemispherical in cross section, smooth, glabrate, usually with minute white ra- phides, the grooves about Cephaelis axillaris is widespread in the tropics, extending from Mexico, through Central America, to Colombia and Venezuela. It also occurs in the West Indies. Steyermark (1972), who has incorporated Cephaelis into Psychotria, anno- tated many specimens of ‘‘Cephaelis axillaris Sw.” as Psychotria aubletiana Steyermark. In the synonomy under P. aubletiana he (1974) listed: *‘Cephaelis axillaris Sw., Prodr. Veg. Ind. Occ. 45: 1788, no Psychotria axillaris Willd., Sp. Pl. 1: 962. 1798, basada sobre Ronabea erecta Aubl., ni Psychotria axillaris Vell., Fl. Flum. 67: 2. pl. 32. 1827." He recognized 7 varieties, 5 of which occur in Venezuela. í: New Switzerland, valley Río Chiriquí Viéjo, Allen 1391 (F, MO). Cerro pg Blum Ü "Dus yer 2654 (MO). Las Nubes, 5.5 km NW of Río Chiriquí Viéjo, del 674 (MO). Las Cumbres, near Cerro Punta, Croat 13712 (MO). Above Río San Félix, 13 mi N of Río San fe 800-1200 m, Croat 33424 (MO). Cerro Punta, Davidson 211 (F). Cerro Horqueta, Dwyer et al. 547 (DUKE, F, GH, K, MO, NY, PA, UC, US). Road to Bajo Grande, Cerro Punta, Pao 4013 (MO). Cerro Colorado, 1500 m, Folsom & Collins 1767 (MO). Between Quebrada Hondo and divide on Caldera Chiriquicito Trail, Kirkbride & Duke 921 (MO). Divide, Chiriquicito Trail, Kirkbride & Duke 963 (MO). Cerro Pando, 16 km NW of El Hato del Volcán, 2000-2482 m, Mori & Bolten 7266 (MO). Cerro Hornitos, 40 km NW of Gualaca, Mori & Bolten 7474 (MO). N of San Félix at Chiriqui-Bocas del Toro border, Cerro Colorado Copper Mine Road, 5000-5500 ft, Mori & Kallunki 5792 (MO). (MO). Cerro Punta, White 28 (F, MO). Upper Chiriquí Viejo, White & White 25 (F. MO). Cerro Horqueta, Wilbur et al. 13464 (MO). Bajo Chorro, Woodson & Schery 699 (F, MO). Casita Alta, Volcán de Chiriqui, Woodson et al. 853 (F, MO). DARIEN: Serrania de Pirre, 9-10 km N of Alto de Nique, 1520-1560 m, Croat 37856 (MO). Cerro Mali base camp, lower montane wet forest, 1400 m, Gentry & Mori 13824 (MO) 2. Cephaelis bristanii Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 34. 1968. TYPE: Panama, Bristan 575 (MO) Subshrubs to 30 cm tall, the stems smooth, glabrous, black, scarcely nodose, often prostrate and bearing long narrow roots. Leaves oblong, 9-16 cm long, 3.5- 6.0 cm wide, apically deltoid, ultimately obtuse, attenuate cuneate at base, the costa slender, prominulous or plano-convex beneath, the lateral veins 15-30, 0.5— 0.8 cm apart, arcuate, the intermediate veins evanescent, rigidly papyraceous, concolorous, glabrous; petioles to 4 cm long, to 1.5 mm wide; stipules connate, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 63 the sheath compressed annular, 1-2 mm long, minutely and obtusely 2-apiculate. Inflorescences axillary, the peduncles solitary, to 2 cm long, black, smooth, the flowers aggregated into 1-2 rotund capitula, ca. 1 cm in diam., the bracts many, clustered, the interior bracts becoming narrower adaxially, oblong, acute, pu- berulent. Flowers with the hypanthium funnel shaped, scarcely 2 mm long, the calycine cup petaloid in texture, ca. 1 mm long, the lobes 4(?), short, truncate or obtuse, often much reduced, the margin then erosulose, often with a few straight cilia; corolla with the tube narrow cylindrical, to 7 mm long, ca. 1 mm wide in the middle, petaloid, glabrous outside, densely white pilose within above the middle of the tube, the lobes 4, to 2 mm long, obtuse; stamens 5, the anthers ca. 0.8 mm long, the filaments linear, ca. 3 mm long; stigmas 2, erect, ca. 1 mm long. Fruits red (fide Bristan), not seen. Cephaelis bristanii is known from Costa Rica and Panama. ''Hinupichica" (Cuna, fide Duke 1243). BOCAS DEL TORO: Quebrada Lugron and Cerro Bonyik near Río Teribe, Kirkbride & Duke 641 (MO). RIÉN: Cerro Pirre, Bristan 575 (MO). Caná Cuasí Trail, Cerro Campanimento near Tres Bocas, Río Cuasí headwaters, Kirkbride & Duke 1243 (M 3. Cephaelis camponutans Dwyer & Hayden, Ann. Missouri Bot. Garden 55: 35. 1968. TYPE Panamá, Dwyer & Gauger 7346 (MO, holotype). Subshrubs to 0.7 m tall, the branchlets smooth, glabrous, the nodes somewhat prominent; roots slender. Leaves strongly ascending, to 18 cm long, to 5 cm wide, attenuate acuminate at the apex, the acumen to 3 cm long, acute at the base, scarcely inequilateral, the costa scarcely prominulous to plane beneath, ca. 2 mm wide proximally, the lateral veins 15-20, evanescent when dry, strongly ascending, the margin somewhat thickened, subcoriaceous, smooth, glabrous; petioles slender, to 3 cm long; stipules obovate rotund, to 1.5 cm long, ca. 1 cm wide, obtuse or somewhat bifid(?) at the apex, thin, with minute raphides, the margins erosulose. Inflorescences axillary, 1—3 per axil, the peduncles to 1 cm long, often nodding, the flowers conglomerate in compressed-rotund heads to 2 cm wide, blackening on drying, the exterior bracts solitary, subovate, to 1.5 cm long, coriaceous, with a median keel, the internal bracts smaller and narrower, a pair of bracteoles enclosing each flower. Flowers with the calyx tubular, the lobes 5-6, erect, subulate, longer than the tube, the margin scarious; corolla purple or white; stigmas 2, erect, digitiform, ca. 1 mm long. Fruits (immature) crowned by a persistent calyx, the pyrenes 2, elliptic; seeds with the adaxial surface plane, the endosperm smooth, the testa somewhat pigmented, perhaps bilamellate, the parenchyma cells somewhat isodiametric, 23-55 my in diam. Cephaelis camponutans Dwyer & Hayden is known only from Panama. S DEL TORO: Punta Pena near Chiriquicito, Lewis et al. 2171 (GH. MO). corów: Santa Rita Ridge, Croat 15345 (MO): Dwyer & Lallathin 9042, 9043 (both MO); Mori & Kallunki 2090 (MO). MÁ: | mi past La Eneida, Cerro Jefe, Correa & Dressler 970 (MO). Goofy Lake to Cerro Jefe, Croat 15204 (MO). El Llano-Cartí Road, 10 mi from Interamerican Highway, Croat 33766 (MO). Cerro Jefe, ca. 600 m, Croat 35871 (MO). El Llano-Cartí Road, 12 km from Panamerican Highway, D'Arcy 10609 (MO). Cerro Jefe, Dwyer & Gauger 7346 (MO); Gentry 2882, 3523 (both MO). El Llano-Cartí Road, Folsom 2589 (MO); Liesner 677, 1190 (both MO); Mori et al. 4630 (MO). VERA- GUAS: $ mi NW of Santa Fé, Liesner 974 (MO). 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 4. Cephaelis correae Dwyer & Hayden, Ann. Missouri Bot. Garden 55: 36. 1968. TYPE: Panama, Lewis et al. 1759 (MO, holotype). Small trees, to 5 m tall, the branchlets terete, smooth. Leaves oblong, to 29 cm long, to 12 cm wide, obtuse to deltoid at the apex, cuneate at the base, the costa somewhat prominulous above, ca. 2.5 mm wide, prominulous beneath, the lateral veins ca. 20, 1.0-1.5 cm apart, at first strict, becoming arcuate near the margin, the intermediate veins pinnatiform, membranaceous, smooth, glabrous; petioles 0.5-0.8 cm long, slender, ca. 0.25 cm wide, smooth, glabrate; stipules briefly connate, the sheath short, 2-5 mm long, interpetiolar, truncate, the lobes large, subrotund, longer than the sheath, obtuse, somewhat spreading, the margin scarious. Inflorescences terminal, solitary, glabrous, the peduncle to 20 cm long, at maturity nodding, ca. 0.3 cm wide in the middle, the exterior bracts 2, to 5 cm long, to 5.5 cm wide, conspicuously glistening, red or somewhat purple, venose, the interior bracts to 1.5 cm long, red to purplish, the larger exterior bracts compressed obovate rotund, much wider than long, undulate marginate, often with several shallow lobes, the smaller bracts obovate rotund and clawed, the claw short or to about the same length as the blade, glabrous, petaloid. Flowers ca. 1.5 cm long, the hypanthium and the calyx 2-3 cm long, glabrous, the cup eglandular, the calycine teeth 5, triangular, unequal, 1-1.2 mm long, each tooth often erosulose; corolla white, the tube cylindrical, to 10 mm long, inflated above, stiffly petaloid, glabrous outside, puberulent within near the filament attachment, the lobes 5, triangular oblong, ca. 3 mm long, glabrous; stamens 5, the anthers linear, ca. 3 mm long, the filaments attached near the middle of the tube; style linear, to 12 mm long, the stigmas 2, oblong, ca. 1.2 mm long. Fruits glistening blue, obovate oblong, 1.5 cm long, the calycine cup persistent; seeds subtrian- gular, flat on the ventral face, somewhat sulcate in the middle. s. correae is found only in Panama. : El Valle, Allen 1648 (MO). Cerro Pilon and La Mesa, Correa 52 (MO): Croat 14335 (MO): bus 12347 (MO); Duke & Lallathin 15010 (MO); Dwver 10534 (MO); Dwver > pai 7961 (MO); Dwyer & Lallathin 8690 A (MO); Gentry 5695 (MO); Lewis et al. 1759 (GH, , UC, US): Luteyn & Kennedy 1707 (MO); Porter et al. 4636 (COL, MO, SCZ, UC, VEN); s & Luteyn 11722 (MO). La Pintada, Hunter & Allen 541 (MO). 5. Cephaelis croatii Dwyer.!! TYPE: Panama, Croat 27204 (MO, holotype; NY, Isotype). Shrubs 1.0-1.3 m tall, the branchlets glabrous, smooth, ultimately angular, the pith porate. Leaves elliptic oblong, 14.5-29.0 cm long, 5.5-9.0 cm wide, rounded or obtuse and abruptly acuminate at the apex, basally cuneate or atten- " MALI rae Dwyer, spec. nov. Frutices ad 1.3 m alti, ramulis glabris. ~~ elliptico- oblonga, 14.5- cm longa, 5.5-9.0 cm lata, basin versus cuneata, venis lateralibus ca. 30, lat arcuatis, fe es acea minuto-puberula; petiolis 3-5 cm longis; stipulis basi ene ‘ovato-ob- longa vel oblongo-rotunda, ad 1.5 cm longa, versus apicem angusta tunc in lobos 2 subulatos vel angusto-oblongos fissa. /nflorescentiae axillares, 4-6 cm longae, 3.0-5.5 cm latae, capitibus florum 6-8 oblongis vel rotundis, ad 1.5 cm latis, bracteis exterioribus ad 0.8 cm longis. Flores cupula calycis cylindrica, 0.5 mm longa, lobis 5. 0.6-1.0 mm s corolla alba tubo cylindrico, 6.5 mm longo, extus glabro intus villoso lobis 5 lanceolatis, ca. 2.5 mm longis; antheris 1 mm longis. Fructus non visi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 65 uate acute, somewhat inequilateral, the costa prominulous above, prominent be- neath, the lateral veins ca. 30, widely arcuate with an irregular vein between a pair of lateral veins, rigidly papyraceous, slightly discolorous, minutely puberu- lent; petioles 3-5 cm long, often twisted, to 0.25 cm wide, glabrate; stipules connate, often turgid, ovate oblong or oblong-rotund, to 1.5 cm long, narrow toward the apex, then each part splitting into 2 subulate or narrowly oblong ascending lobes, these to 0.7 cm long, subcoriaceous, glabrous. Inflorescences axillary, 5-6 cm long, 3.0-5.5 cm wide, drying black, the peduncle to 3 cm long, to 0.3 cm wide, twisted, plano compressed, the flowers aggregated into heads, these 6-8, oblong or rotund, to 1.5 cm wide, sessile or on branchlets, variable in length, to 2 cm long, puberulent; bracts rotund, outermost 8-10 mm long, 8 mm wide, concave, carnose, minutely pubescent outside, minutely ciliolate within, purple. Flowers with the hypanthium short, the calycine cup cylindrical, ca. 0.5 mm long, turgid, petaloid, with a gland at the base of adjacent lobes, the lobes 5, triangular subulate or narrowly oblong, 0.6-1.0 mm long, obtuse, glabrous within, ciliate; corolla white, the tube cylindrical, ca. 6.5 mm long, somewhat dilated below the middle, glabrous outside, villose within, the lobes 5, lanceolate, ca. 2.3 mm long, somewhat cucullate; stamens 5, the anthers oblong, 1 mm long, obtuse, the filaments short; style slender, somewhat expanded toward the apex, the stigmas 2, short, rotund, wider than the style. Fruits not seen. Cephaelis croatii is known only from Panama. It is named in honor of Thomas B. Croat, distinguished student of the flora of Panama. There are several distin- guishing characters: relatively broad leaves with numerous lateral veins; large bilobed stipules; and subrotund, capitate clusters of flowers, about 5 per head. ARIEN: Upper gold mining camp of Tyler Kittredge, headwaters of Rio Tuquesa ca. 2 air km from continental divide, Croat 27204 (MO). 6. Cephaelis dichroa (Standley) Standley, Publ. Field Columbian Mus., Bot. Ser. 4: . 1929. Evea dichroa Standley, Contr. U.S. Natl. Herb. 18: 124. 1916. Type: Panama, Pittier 3218 (US, olotype). Glabrous shrubs, to 2 m tall. Leaves elliptic, 8.5—13.0 cm long, to ca. 3.2 cm wide, acuminate at the apex, cuneate at the base, the lateral veins ca. 12, stiffly papyraceous, concoloreus, often drying yellowish, glabrous; petioles to 3 cm long, wiry, often flexuous; stipules connate, ca. 3.5 mm long, the body com- pressed cylindrical, each part with 2 widely spaced, erect, obtuse lobes about as long as the sheath. Inflorescences terminal, the peduncle 5—8(—9) cm long, ter- minated by 3 heads of flowers, the trio subtended by 2 large obovate bracts, to 2.2 cm long, ca. 1.8 cm wide. Fruits subrotund to 0.8 cm wide. Cephaelis dichroa is known only from Panama. Standley's original description of the inflorescences and flowers may help in the understanding of this species: ""Peduncles terminal or rarely also axillary, 3-9 cm long; bracts 15-20 mm long, rounded ovate, acutish, slightly united at the base, green or tinged with red or yellow, subtending 3 pedunculate heads, the peduncles 3-25 mm long; each sec- ondary head subtended by 2 orbicular-rhombic bracts 12-15 mm long, these yel- 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 low or tinged with purple, each head consisting of 3 flowers, each flower sub- tended by an oblong obovate acute bract 10-12 mm long and an oblong-linear bracteole somewhat shorter; flowers sessile; calyx 3 mm long, twice as long as the ovary, somewhat oblique, yellow, deeply 5-lobed, the lobes ovate, oblong, acute; corolla yellowish white, the tube 10 mm long and 2 mm in diameter, the 5 lobes 2.5-3 mm long, oblong, obtuse; stamens included, inserted in the middle of the tube, the filaments about as long as the anthers, these linear, 2.5 mm long; fruits not seen." Another collection, Mori & Kallunki 2732 (MO), from between Fort San Lo- renzo and Fort Sherman near Pavon Road junction with Road 82 (S 2), Canal Zone, may be C. dichroa. It is 2% m tall with bracts magenta and fruits black, but its sea level locality argues against such determination. CHIRIQUi: Cerro Horqueta, Hagen & Hagen 2089 (F, US). Cerro Colorado, 50 km N of San O Felix on the continental divide. 1200-1500 m, Mori & Dressler 7843 (MO). Cuesta de Las Palmas, S slope of Cerro Horqueta, 1700-2100 m. Pittier 3218 (F, NY). Cerro Horqueta, Pittier 3266 (US). 7. Cephaelis dimorphandrioides Dwyer.!2 TYPE: Panama, Croat 27470 (MO, ho- lotype) Shrubs to 1.4 m tall. Leaves elliptic oblong, often falcate oblong, 17-30 cm long, 6-14 cm wide, deltoid or obtuse toward the apex, acute or abruptly acute toward the base, the costa slender above, prominent beneath, the lateral veins 13-17, arcuate, often with one smaller and shorter vein between 2 lateral veins, the intervenal areas spreading reticulate, rigidly chartaceous, discolorous, gla- brate above, minutely puberulent beneath; petioles lignose, to 4.5 cm long, twist- ed, puberulent; stipules (only one pair seen at tip of twig) oblong rotund, to 0.7 mm long, deeply 2-lobed, coriaceous, the lobes rounded, farinose, glabrous with- in. Inflorescences terminal, sessile, cylindrical, 5-12 cm long, to 4 cm wide, obtuse at the apex, black when dry, the flowers more than 100. Flowers perhaps sessile, the hypanthium short, swollen, puberulent, the calycine cup short, ca. 5 mm long, the teeth subulate, to 6.5 mm long, unequal, acute, puberulent outside, glabrous inside, carnose, with 3 longitudinal veins, with 2-4 oblong glands, to 0.3 mm long, at the sinuses; corolla red outside, the tube narrowly cylindrical, 10-14 mm long, puberulent outside, glabrous within, subcarnose, the lobes 5, yellow within, lance oblong, to 3 mm long, carnose, puberulent outside; stamens 5, the anthers narrow oblong, 2.8-3.0 mm long, minutely apiculate, slender, attached below the middle of the tube; ovarian disc 4-lobed, to 0.8 mm long, the style linear, 9-14 mm long, the stigmas 2, reflexed, linear oblong, to 0.8 mm long. 2 Cephaelis dimorphandrioides Dwyer, spec. nov. Frutices ad 1.4 m alti. Folia elliptico-oblonga, saepe falcato-oblonga, 17—30 m nga, 6-14 cm lata, venis pin us 13-17 rigido- isa acea ids glabrescentia infra minuto-puberula; petiolis ad 4.5 cm longis; stipulis dne. rotu ad 0.7 m longis, bilobatis glabris. oie entiae terminales ics cylindricae et ue i. 5-12 cm longae, ad 4 cm latae, floribus numerosis fortasse 100. Flores cupula calyci .5 mm longa, dentibus subulatis ad 6.5 mm longis; corolla extus rubra, tubo angusto-cylindrico, 10 14 mm longo, extus ue intus glabro lobis 5, ca. 3 mm longis; antheris ca. 3 mm longis. Fructus Pose ad 0.5 cm longi 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 67 Fruits (immature) oblong, to 5 mm long, to 0.3 mm wide, truncate, each pyrene smooth, black, 10-ribbed, the hairs weak, white, arachnoid. Cephaelis dimorphandrioides is known only from Veraguas. It is distinguished from all other Cephaelis by the cylindrical spikelike inflorescence which bears a resemblance to the inflorescence of the legume genus Dimorphandra. VERAGUAS: Valley of Rio Dos Bocas between Alto Piedra and Calovébora, 350-400 m, Croat 27470 (MO, holotype). Road from Santa Fé to Calovébora dew giia 35) 16 km from Santa Fé; on Atlantic slope near Río Caloveborita, Mori et al. 6675 (M 8. Cephaelis discolor Polak., Linnaea 41: 572. 1877. TYPE: Costa Rica, Pola- kowsky 384 (F, MO, US, photos). — tonduzii Krause, Bot. Jahrb. Syst. 54. Beibl. 119: 45. 1916. Type: Costa Rica, Tonduz F). iie guapilensis Standley, J. Wash. Acad. Sci. 15: 104. 1925. rype: Costa Rica, Standley 37025 (US, holot Cephaelis nic SE Standley, Trop. Woods 16: 46. 1928. rvPE: Nicaragua, Englesing 58 (F). C. guapilensis (Standley) Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 295. 1929. Subshrubs to 1 m tall, usually unbranched, the stem terete, smooth, glabrous. Leaves oblong, to 22 cm long, to 8.5 cm wide, subobtuse to deltoid at the apex, acuminate, the acumen to 1.5 cm long, basally cuneate or attenuate acute, slightly inequilateral, the costa prominulous above, prominent beneath, the lateral veins 12-14, arcuate, each pair with usually 1 subparallel irregular vein, soon evanes- cent, the intervenal areas spreading reticulate, delicately marginate, stiffly char- taceous, usually discolorous, glabrous; petioles to 7 cm long, slender, 0.13—0.20 cm wide, ascending, glabrous; stipules connate for 0.4-4.0 mm, each with 2 slender awns, to 8 mm long, these often reflexed, often deciduous, glabrous. Inflorescences terminal, the peduncles 0.7-2.0 cm long, lignose, glabrate, the flowers aggregated into a subrotund mass, 3-5 cm in diam.; bracts purple green or purple, obovate lanceolate, 0.8-1.0 cm long, petaloid, glabrous, venose. Flow- ers with the hypanthium subrotund or oblong, ca. 1.5 mm long, constricted below the calycine cup, the cup short, the lobes 4-5, subequal or equal, triangular, 0.8— 1.5 mm long, 2-3 times the length of the cup; corolla and sex organs not seen; ovarian disc bilobed, ca. 0.5 mm long. Fruits bright blue, ca. 1 cm in diam., the fleshy exterior often persisting, the pyrenes when united rotund, buttockslike, the central groove conspicuous, ca. 5 mm in diam., glabrate, smooth, the raphides minute, numerous, the calycine cup persistent, 0.5-0.8 mm in diam. Cephaelis discolor occurs in Costa Rica, Panama and Colombia. OCAS DEL TORO: Water Valley, Wedel 1508 (MO). CANAL ZONE: Barro Colorado Island, Croat 11870 (MO). Pipeline Road, Gentry & Dressler 1978 (MO). COLON: Santa Rita Ridge Road, 8.5 km from Boyd-Thatcher Highway, Mori & Kallunki 2089 (MO). DARIEN: La Boca de Pirre, Bristan 1273 (MO). Cerro Pirre, Duke 5307 (MO); Duke & Elias 13767 (MO). Cocolito, Dwyer 5136 (MO). PANAMA: Cerro Campana, Allen 1886 (MO). Cerro Trinidad, Allen 3774 (MO). Cerro Campana, Croat 14752 (MO). Cerro Jefe, Foster & dee 1818 (F). Cerro Campana, Gentry 5783 (MO); Kirkbride & Hayden 239 (MO); Lewis et al. 3061 (MO). vERAGUAS: N of Santa Fé, N of Escuela Agricola Altos de Piedras, Mori & Kallunki 2533, 2587, 2619 (all MO). COLOMBIA: CHOCO: Serrania del Darién, W of Unguia on Panama border, 1150 m, Gentry et al. 17038 (MO). 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 9. Cephaelis dressleri Dwyer.!3 ryPE: Panamá, Croat 16801 (MO. holotype).— Fic. 15 Subshrubs to 1.5 m tall, the stem lignose, rough, glabrous, the pith partly porous, sometimes rooting toward the apex. Leaves occasionally sessile, obovate oblong, 10-44 cm long, 6.5-22.0 cm wide, rounded or widely deltoid at the apex, short acuminate, the acumen triangular, to 1 cm long, to 1.5 cm wide at the base, cuneate toward the base finally slightly auriculate, the costa prominulous above or prominent, to 0.2 cm wide, prominent beneath, to 0.35 cm wide, dark brown and densely minutely puberulent, the lateral veins 21—25, slightly arcuate, forming an undulate marginal vein to 1 cm from the margin, the venules irregular pinnati- form and spreading, rigidly papyraceous to subcoriaceous, | of the 2 leaves drying slightly dark purple red beneath, minutely ferrugineous puberulent on the veins and venules; petioles, if present, to 4 cm long, stout; stipules (only 1 seen at the tip of the stem) ovate lanceolate, to 2.8 cm long, to 1 cm wide, subcoriaceous, glabrous, the veins somewhat prominulous, fanshaped, the awns narrowly ovate subulate, to 0.9 cm long, acute. Inflorescences terminal, 4.5-8.0 cm long, the peduncles 1-4 cm long, 0.3-0.5 cm wide, glabrous, bearing several short branch- es, the flowers disposed in a few subsessile oblong or subrotund heads, ca. 2 cm in diam., the cluster of heads compressed rotund, 3.5-5.5 cm wide; outermost bracts rotund or spathulate, to 12 mm long, brown, ultimately acute, concave, within with an irregular series of glands on each side of a median vein or with a few diffuse glands, ciliate; interior bracts several, narrower and more spathulate. Flowers subsessile; hypanthium scarcely measurable, the calycine cup to 2 mm long, petaloid, with ca. 10 unequal, narrowly oblong or widely subulate glands attached toward the base of the cup, the calycine lobes 5, subulate or triangular, to 3 mm long, slightly unequal, acute, densely ciliate; corolla white, the tube narrow cylindrical, 10-14 mm long, petaloid, glabrous or minutely puberulent outside, glabrous inside except white villose near or above the middle, the lobes 5, narrowly lanceolate, to 4.5 mm long, | mm wide, conspicuously cucullate near the middle, ultimately obtuse, carnose; stamens 5, the anthers narrowly ovate oblong or oblong, 2.5-3.0 mm long, sagittate at the base, the filaments slender, ca. 2.5 mm long, attached ca. 2 mm below the mouth; ovarian disc oblong rotund, ca. 1 mm long, the style slender, 6.5-9.0 mm long, the stigmas slender, ca. 2 mm long, densely villosulose. Fruits sessile, elliptic oblong, to 1.2 cm long, 0.4 cm wide, smooth, glabrous, each pyrene 4-sulcate, the ribs obtuse angular, the often persistent calycine cup and teeth to 3.5 mm long. Cephaelis dressleri is known from Panama and Colombia. It is named in honor of Dr. Robert Dressler, student of the flora of Panama and a specialist in the n Cephaelis dressleri Dwyer, spec. nov. Suffrutices ad 1.5 m alti, caule lignoso e glabro interdum apice rhizophoro. Folia sessilia vel subsessilia E presen , 10-44 cm longa, 6.5-22.0 cm lata, rotunda a vel vix latodeltoidea, versus basim cuneata ultime v vix auriculata venis late- ralibus ad 25, rigido-papyracea vel n bel: cuf. ovato-lanceolatis, ad 2.8 cm longis, ad 1 cm latis, setis solitariis ad 0.9 cm longis, acutis. d 4.5-8.0 cm longae, pedunculo 1-4 cm longo, caespite florum compressorotundo, 3.5-5.5 cm lato; bracteis exterioribus rotundis vel spathulatis, ad 12 mm longis. Flores cupula calycis d iue p 3 mm longis; corolla tubo 10-14 mm longo, extus glabro, lobis ad 4.5 mm longis; antheris 2.5-3.0 mm longis. Fructus elliptico- oblongi, ad 1.2 cm longi, glabri. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) FiGURE 15. C. Fruit (x3). [A Cephaelis dressleri Dwyer.—A. Leaf and inflorescence (x'2).—B fter Dressler 4677.] . Flower (x3).— 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Orchidaceae. It is distinguished by its large sessile or subsessile leaves, round at the apex, with many lateral veins, and subauriculate at the base. The glands within the larger bracts and within the calycine cup are diagnostic. OLÓN: Near Agua inta at rain E Santa a lumber road, Dressler 4677 (MO). DARIEN: Trail from Tacarcuna villag o Tacarcuna to Cerro Mali, 800-1300 m, Gentry 13598 (MO). SAN BLAS: Puerto Obaldia, aan 16801 (MO). COLOMBIA: CHOCO: Rio Tigre W of Unguia, base of Serrania del Darién, 300 m, A. Gentry & L. Aguirre 15191 (MO). 10. Cephaelis elata Swartz, Prodr. 45. 1788. TYPE: Jamaica, Swartz, not seen. Cephaelis punicea Vahl, Eclog. Amer. 1: 19. 1796. TYPE: Jamaica, von dae not seen. elata Aubl. ex Lam., Encycl. Méth. 7: 585. 1806. TYPE: n °ophaelis costaricensis Schlecht., Linnaea 28: 546. 1856. TYPE: not s ae punicea (Vahl) Schum. in Engl. & Prantl, Nat. Pfanzenfam. IV: 120. 1891. Evea elata (Sw.) Standley, Contr. U.S. Natl. Herb. 18: 123. Cephaelis chiriquiensis Standley, Ann. Missouri Bot. Gard. id 2 1941. TYPE: Panama, Woodson & Schery 610 (MO, holotype). Shrubs or trees to 8 m tall, the branches few to many, smooth, terete or angular, glabrate, occasionally constricted below the nodes, the nodes often swol- len (persistent stipules). Leaves narrowly oblong or obovate oblong, 10-25 cm long, 1.5-7.0 cm wide, obtuse, widely deltoid or rounded at the apex, acuminate, the acumen short, acute, basally cuneate to obtuse, the costa prominent, prom- inulous to subplane above, prominent beneath, the lateral veins (12—)16—22, widely arcuate or at first widely divergent and then arcuate marginally, the margin often slightly revolute, stiffly papyraceous to subcoriaceous, not herbaceous (Panama), lustrous and glabrous above, glabrous or pubescent on the costa and the veins beneath; stipules persistent, connate, turgid, cupshaped, to 1.2 cm long, each part of the sheath bilobed, the lobes usually hemispherical, as long as or longer than the sheath, obtuse, coriaceous, glabrous, the margins scarious, often widely so. Inflorescences terminal, the peduncle solitary or rarely 2-4 at the tip of the twiglet, 0.5-3.5 cm long; heads including 2 outermost bracts to 6.5 cm wide; outermost bracts ovate rotund, to 7 mm long, deep red or rusty orange. CANAL ZONE: Camp Pina, Duke 4400 (MO). Pena Road, Tyson & Chu 1700 (MO). Fort Sherman, Tyson & Dwyer 1215 (MO). CHIRIQUI: Bajo Chorro, Boqueté District, Davidson 103 (MO). Cerro Horqueta, Kirkbride 131, 132 í MO). Between Pinola and divide on Chiriquicito-Caldera Trail, Kirkbride & Duke 845 (MO). Bajo Chorro, Woodson & Schery 610 (MO). COcLE: Cerro Pajita, N o " e de Antón, Allen & gus 4168 (MO). Cerro Caracoral, Duke & Dwyer 15103 (MO). EI Valle, er & Correa 7998 (GH, US); Lewis et al. 1713 (GH, MO, US). corów: Santa Rita Ridge, PM 14146 (MO). DARIÉN: es Pirre, Bristan 462 (MO); Duke 6570 (MO). Summit camp between Sasardí and Mortí, Duke 10034 (MO). Cerro Pirre, Duke & Elias 13752 (GH, MO). Top of Cerro Mali. old helipad, Gentry & Mori 1366 (MO). Summit camp, Darién-San Blas border, Oliver et al. 3692 (COL, MO, UC, VEN). Cana-Cuasi Trail, Terry & Terry 1480 (MO). PANAMA: Cerro Jefe, Blum & Duke 2197 (MO). Cerro Campana, Blum et al. 2371 (MO). El Llano-Carti Road, 330 m, Croat 33742 (MO). Cerro Jefe, Duke 8018 (MO); Dwyer 444 (MO). Cerro Azul, Dwyer 1369 (MO). Cerro y. Folsom 2551 (MO). El Llano-Cartí road, 13.7 km N of Panamerican Highway, Folsom UA "3505 (both MO). Top of Cerro Jefe, tears 3614 (MO). Cerro Campana, vie 118 (MO Cerro Jefe, DEA & c 33 (MO). Altos del Río Pacora, Lewis et al. 2262 (COL, DUKE, K, MO), 2356 (COL, MO, UC, VEN). Cerro A Tyson 3189, 3205, 3231, 3254, 3285, 3288. 3373, 3389, 3425, 3546, 3577, hes (all Mo. a et al. 8282, 8503 both MO). SAN BLAS: SE of Puerto Obaldia, Croat 16750 (MO). E of Cangandi-Mandinga airport road, 2-5 mi S of Mandinga airport, Duke 14771 (MO). Between Rio Diabolo ar Rio Acuati near aa Duke 14865 (MO). VERAGUAS: 3.9-5.0 mi N of Santa Fé, Gentry 2951 (MO). Mouth of Río Concepción, Lewis et al. 2806 (COL, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 71 DUKE, MO, UC, VEN). Cerro Hornitos, 2238 m, Mori & Bolten 7496 (MO). 10 km NW of Santa Fé, road to Calovébora, Mori & Bolten 7639 (MO). N of Santa Fé, Escuela Agricola Alto de Piedra, Mori & Kallunki 2507 (MO) 11. Cephalis gaugeri Dwyer.!! Type: Panama, Mori et al. 3779 (MO, holotype). Subshrubs to V5 m tall, the stem rhizomatous at the base, the roots fibrous, the stem unbranched, ca. 0.3 cm wide, glabrous but with minute black puncta- tions, the nodes well spaced. Leaves succulent, oblong, 5.0-7.5 cm long, 1.8-3. cm wide, widely deltoid toward the apex, acuminate, the acumen to 0.3 cm long, ultimately obtuse, toward the base cuneate, obtuse or rounded, slightly inequi- lateral, the costa prominulous above, subplane beneath, red on drying, to 0.2 cm wide, the lateral veins ca. 7, arcuate, somewhat inconspicuous, the fine veinlets obscure, the margin lightly revolute, coriaceous, concolorous, smooth, glabrous; petioles to 1 cm long, rigid, glabrous; stipules free, often persistent, compressed- rotund, to 1 cm long, to 1.3 cm wide, with 2 triangular or oblong lobes, ca. 0.5 cm long, the lobes erose or with subulate teeth, the body of the stipule rigidly scarious, glabrous, the raphides dense. /nflorescences axillary, the peduncle ca. | em long, ca. 0.3 cm wide, the fruits disposed in a globose head, to 2.5 cm in diam. Fruits sessile, oblong, to 1.5 cm long, obtuse or truncate toward the apex, carnose, smooth, glabrous, red, purple toward the apex, the bracts few, large, imbricate, compressed rotund, to 0.8 cm long, perhaps much wider than long, subcoriaceous, glabrous except ciliolate toward the margin, the margin erose, the raphides dense. Cephaelis gaugeri is known only from Panama. It is named in honor of Mr. George Gauger, a microbiologist in the Canal Zone who has assisted this author in collecting plants in Panama. e new species is distinguished by its stem being rhizomatous, its leaves being succulent with few, inconspicuous lateral veins, and its stipules being large and lobed. PANAMA: Roads to Cerro Jefe and Altos de Pacora, Mori et al. 3779 (MO). 12. Cephaelis glomerulata Donnell Smith, Bot. Gaz. 16: 12. 1891. TYPE: Guate- mala, J. D. Smith 1637, not seen.—Fic. 16 Psychotria glomerulata (Donnell Smith) Steyermark, Mem. N.Y. Bot. Garden 23: 670. 1972. Subshrubs to 2 m tall, the stem simple above or occasionally with 2 terminal branches, the twiglets terete, glabrous, the nodes well spaced. Leaves lanceolate, ovate lanceolate or oblong, occasionally inequilateral, occasionally salicoid, 5— 13 cm long, 1.6-5.5 cm wide, deltoid or acute at the apex, acuminate, the acumen 14 Cephaelis ino ri Dwyer, spec. nov. Suffrutices ad 1⁄4 m alti rhizomati caule simplici. Folia succulentia, oblonga, 5.0-7.5 cm longa, 1.8-3.0 cm lata, versus apicem lato-deltoidea venis lateralibus . 7 in sicco ee en ine coriacea laevia glabra; petiolis ad 1 cm longis; stipulis comp o- rotun dis, ad 1c i 1.3 cm latis, prope medium lobis 2 ad 0.5 cm longis, in margine erosis gis, vel dentibus subulatis praeditis. /nflorescentiae axillares, pedunculo ca. 1 cm longo, ca. 0.3 cm lato fructibus in caespitem globosum ad 2.5 cm dispositis. Fructus sessiles oblongi, ad 1.5 cm longi, carnosi, laeves, glabri, rubri. 72 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 16. Cephalis glomerulata Donnell Smith.—A. Inflorescence (x3/s).—B. Flower (x2.4).—C. Fruit (x 1.2). [After Hammel 1819. | to 1.5 cm long, ultimately acute or obtuse, cuneate or obtuse at the base, the lateral veins 12-16(—20), widely arcuate, with several smaller veins between ad- jacent laterals, these soon branching, the intervenal areas spreading reticulate, chartaceous to papyraceous, glabrous, usually lustrous; petioles 0.3-1.5 cm long, glabrous; stipules connate, erect, somewhat inflated, cylindrical, to 3 mm long, truncate, glabrous. Inflorescences terminal, sessile or subsessile (the peduncle to 0.5 cm long), the bracts and flowers aggregated into an oblong or compressed rotund head, 1.5-2.5 cm long, often somewhat wider than long, the outermost bracts ovate rotund, to 1.5 cm long, to 1 cm wide, obtuse, rounded or bilobed at the apex, subcoriaceous, glabrous, venose, eglandular within, white or green, turning purple in fruit, with progressively smaller, narrower, and more concave bracts within, each flower enclosed by 2 tightly enveloping bracteoles. Flowers 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 73 sessile; hypanthium ca. | mm long, glabrous, the calycine cup about equal to the hypanthium, delicately petaloid, glabrous, eglandular within, the teeth 5, irregular but mostly subulate or triangular, to 1.2 mm long; corolla white, the tube cylin- drical, ca. 1 cm long, petaloid, glabrous outside, puberulent within in the middle of the tube for ca. 4 mm; stamens 5, the anthers subsessile, oblong, 2.2 mm long, obtuse, minutely apiculate, the filaments attached near the middle of the tube; style slender, about as long as the tube, the stigmas 2, dilated, oblong, 1.3 mm long, subplane. Fruits fleshy, oblong or subrotund, to 1 cm long, china blue at maturity, glabrous. Cephaelis glomerulata extends from Guatemala to Panama. *‘Uasika”’ (Duke 14378) BOCAS DEL TORO: Punta Pena, near Chiriquicito, ca. 1000 ft, Lewis et al. 2175 (COL, MO, UC, VEN). Old Bank Island, Wedel 1930 (MO, US), 1941 (US). Fish Creek, Chiriquí Lagoon, Wedel 2199 (MO, US). Fish Creek Mountains, Wedel 2309 (MO, US). Bastimentos, Chiriqui Lagoon, Wedel 2907 (MO). Isla Colón, Wedel 2968 (MO, US); Woodson et al. 1936 (F, MO). CANAL ZONE: Telephone cable trail between splice S16 and S49, bs Indio, Steyermark & Allen 17428 (MO). Fort Sherman near road to Gatün along Road S-5, 1 & Dwar 1215, 1243 (both MO). cocrÉ: 16.7 km N of turnoff to Coclesito ES Llano G NE yon ees (MO). coLGn: Santa Rita Ridge, Croat 14143 (MO); Dwyer 8404, 8 d. (both MO): Dwyer et al. 8995 (MO). Highway to 8 mi E of Santa Rita Ridge, 800 ft, Dwyer et al. 8986 (MO). Santa Rita Ridge road, pp 1887 (MO). Santa Rita Ridge, Gentry 6822 (MO); Lewis et 2 5272 (MO). Santa e Ridge, 2.7 mi by gravel road NE of Transisthmian Highway, Nee & Mori 3675 (MO). PANAMÁ: Road to DR 13 km N of El Llano, Busey 384 (MO). Cerro Jefe, 5 mi S of Summit, Busey & Croat 2 (MO). El Llano-Cartí Road, 10 mi from Panamerican Highway, 330 m, Croat 33756 (MO). Cerro Jefe, Croat 35869 (MO). Río Sancanti, ca. 2 mi upstream from Piria, ca. 120 m, Duke 14378 (MO). Goofy Lake, to 8 mi S of Goofy Lake toward Cerro Jefe, Dwyer 7103 (MO). Be ii Cerro Jefe and La Eneida, 2100-2900 ft, Dwyer 8241 (MO). El Llano- Cartí Road, Folsom 2616 (MO). El Llano-Cartí Road, 12 km N of Panamerican Highway, Folsom 3597 (MO). Cerro Jefe, ca. 1000 m, Gentry 6147 (MO). Cerro Jefe, ca. 3100 ft, Loue et al. 3459 (MO). Cerro Jefe to La Eneida, Hayden 1016 (MO, UC). El Llano to Cartí-Tupile, 16 km above Panamerican Highway, gps: et al. 2429 (MO). Altos del Río Pacora, 2500 ft, cen et al. 2262 (COL, DUKE, K, MO, UC, VEN). El Llano to Cartí-Tupile, 300-500 m, Liesner 1276 (MO). Gorgas Memorial Labs, 5-10 km NE a Altos de Pacora, ca. 600 m, Mori & Kallanks 3325 (MO). Cerro Jefe, 2700 ft, Tyson 3373 (MO). VERAGUAS: Mouth of Rio Concepcion, Hayden 2806 (COL, DUKE, MO, UC, VEN). 3-4 km W of Santa Fé, 2500 ft, Nee 11308 (MO). SAN BLAS: Inland from airport at Mandinga, Duke 8917 (MO) 13. Cephaelis insueta Dwyer." type: Panamá, Mori & Kallunki 3108 (MO, ho- lotype). Shrubs (?), the stems rhizophorous to the bases of the uppermost leaves, square, dark red when dry, glabrous, the nodes crowded, the petiole scars sub- rotund, to 0.8 cm long, the pith perhaps porate. Leaves sessile, oblong, 17—43 cm long, 8-16 cm wide, widely deltoid at the apex, acuminate, the acumen tri- angular, ca. 1 cm long, the blade spathulate toward the base, finally subauriculate, the costa subplane above or prominulous, prominent beneath, to 0.3 cm wide, the lateral veins ca. 42, scarcely arcuate, the submarginal veins 2, these somewhat 15 Cephaelis insueta Dwyer, spec. nov. sarune s (?) caule rhizophoro etiam basibus foliorum superiorium. Folia sessilia, oblonga, 17—43 cm l pus -16 cm lata, lamina basim versus spathulata ultime subauriculata venis lateralibus ca. 42 vix arcuatis E. submarginalibus 2 margini subparal- lelibus tenui-chartacea discoloria supra spy aiat puberula. Inflorescentiae E à 16 cm longae, ad 6 cm latae, pedunculo recto, ad 10 cm longo; Miel i minutis ovato-subulat . 1mm longis. Flores non visi. Fructus elliptico- lu. ad 0.9 cm longi, ad 0.4 cm lati, laeves glabri opaco- rubri calyce ca. 1 mm longo. 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 parallel to the margin, the one nearer the margin inconspicuous, slender, 0.1 mm from the margin, the other prominulous, | mm from the margin, the intervenal areas spreading reticulate, thin chartaceous, discolorous, glabrous above, mi- nutely puberulent beneath on the costa and the veins. Inflorescences terminal (only one seen in fruit) to 16 cm long, to 0.6 cm wide, puberulent, the peduncle erect, to 10 cm long, to 0.5 cm wide, flattened, the branches few, ascending, the lowermost perhaps opposite, to 4 cm long; bracteoles minute, ovate subulate, ca. 1 mm long. Flowers not seen. Fruits on short pedicels; elliptic oblong, to 0.9 cm long, to 0.4 cm wide, smooth, glabrous, dark red, the calyx persistent, ca. 1 mm long, each pyrene 4-sulcate. Cephaelis insueta is known only from Panama. The Latin adjective means *unusual" which seems appropriate as the stem of the new species is rhizopho- rous even to the apex; the basally spatulate leaves are striking. The 2 very reduced submarginal veins which parallel the margin suggest the situation in e.g. Faramea talamancarum Standley. The collectors of the new species note: "'detritus col- lected in the leaf axils penetrated by adventitious roots from the stem.” VERAGUAS: 11 km from Escuela Agricola Alto de Piedra forest along Río Dos Boca, tropical wet forest, Atlantic slope, Mori & Kallunki 3108 (MO). 14. Cephaelis ipecacuanha (Brot.) A. Rich., Bull. Fac. Med. 4: 92. 1818. Callicocca ipecacuanha Brot., Trans. Linn. Soc. London 6: 137. 1802. Type: Brazil, Gomez, not seen Ipecac sanha officinalis Arruda, Diss. Pl. Brazil 44. 1810. Not seen, reprinted in part as Appendix to Kos Travels in Brazil 2: 374—375. 1817. type: Described from PM on. Evea ipecacuanha (Brot.) Standley, Contr. U.S. Natl. Herb. 2 I 19 Ouragoga ipecacuanha (Brot.) Farw., Drugg. Circ. 61: 175. 1 Subshrubs to 0.25 m tall, the stems simple, arising from a rhizome, to 0.8 cm in diam., the nodes often crowded toward the apex of the stem, and often rough with stipular scars. Leaves oblong, obovate oblong, elliptic oblong, oblong sub- rotund, 8-14 cm long, 3.2-9.0 cm wide, obtuse or rounded at the apex, occa- sionally widely cuneate, acuminate, the acumen often vague, usually widely tri- angular, to 1 cm long, basally obtuse, rounded, attenuate acute or cuneate (and the lamina abruptly contracted), occasionally vaguely auriculate, the costa slen- der, prominulous above and beneath, the lateral veins ca. 6, arcuate, uniting well below the margin to form a submarginal undulate vein, with 1—2 smaller lateral veins between a pair of lateral veins, stiffly papyraceous or chartaceous, glabrous or occasionally sparsely puberulent above and beneath; petioles either absent or to 0.5 cm long, glabrous; stipules persistent, appressed to the stem, thin, oblong or ovate rotund, to 11 mm long, the body 3-5 mm long, with ca. 10 linear subulate awns to 7 mm long, these spreading fanlike or curled, glabrous. /nflorescences axillary and/or terminal, glabrous, the peduncles erect or deflexed, 1.2-4.0 cm long, ca. 0.15 cm wide, terminated by a bracteate cluster of flowers, the heads compressed rotund, 1.5 cm in diam., to 3.5 cm wide when bracts spread, the bracts elliptic oblong to subrotund, 8-10 mm long, 3.5-8.0 mm wide, glabrous, stiffly petaloid, with a few marginal, weak, subulate projections at the base, to 1.5 mm long, to 0.2 mm wide basally. Flowers sessile, distylous; hypanthium 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 75 0.1—0.2 cm long, glabrous, the calycine cup 0.1—0.2 mm long, eglandular within, glabrous, the teeth 5, unequal, irregular, scarious, tending to be triangular, to 0.5 mm long; corolla tube cylindrical, to 4 mm long, petaloid, glabrous outside, pu- berulent within from filament attachment to the mouth, the lobes 5, oblong or ovate oblong, ca. 1.5 mm long, acute; stamens 5, the anthers exserted or included, narrowly oblong, ca. 1.7 mm long, the filaments short or elongate, attached near the middle of the tube; ovarian disc rotund, ca. 0.3 mm long, the style filamentous, 0.2—0.6 cm long, the stigmas minute. Fruits fleshy, red, oblong or rotund, to 1 cm long, the pyrenes smooth, glabrous or with a small ringlike scar. Cephaelis ipecacuanha is well distributed throughout tropical America. The roots and rhizome contain an alkaloid emetin, an emetic and expectorant. This drug is known in English as ‘‘ipecac’’ or in Spanish as ‘‘Raicilla.”’ The United States Pharmacopeia lists Rio or Brazilian ipecac as Cephaelis acuminata Kar- sten, "known in commerce as ‘Cartagena, Nicaragua, or Panama ipecac’’’ which is supposed to have different pharmacological properties from Rio or Brazil ipe- cac. The name Cephaelis acuminata is an unpublished name which may refer to plants which do not occur in Panama. Panamanian material seen to date is quite uniform taxonomically and agrees with other material from South America which has been going under the name Cephaelis (or Psychotria) ipecacuanha. BARRO COLORADO ISLAND: Bartlett & Lasser 16723 (MO); Busey & Croat 237 (MO); Croat 4096, 4104, 4269, 6609, 12427, 15117 (all MO); Hayden 122, 127, 155, 1037 (all MO); Starry 55 (MO) DARIEN: Punta Alegre, Ambulo (MO). Rio Tuira and Rio Paca, Duke 5027 (MO). Line CC, Duke 5237 (MO). Río Chucunaque, 2-10 mi above Cuna-Darién border, Duke 8566 (MO). Without other locality, Sexton 146A, 146B (both MO). SAN BLAS: Headwaters of Rio Mulatupo, Elias 1770 (MO). 15. Cephaelis kennedyi Dwyer.'^ TYPE: Panama, Kennedy 1779 (MO, holotype). Subshrubs to 2 m tall, the branchlets terete, smooth, glabrous, the nodes well spaced. Leaves narrowly oblong or obovate oblong, 4—9 cm long, 0.8-3.0 cm wide, obtuse toward the apex, often abruptly acuminate, the acumen to 1 cm long, often falcate, ultimately obtuse or acute, basally acute or cuneate, slightly inequilateral, the costa prominulous above, prominulous beneath, ca. 0.35 mm wide, the lateral veins ca. 20, widely arcuate, subprominent, chartaceous, gla- brous; petioles to 1 cm long, 0.8 mm wide, glabrous; stipules connate, the sheath cupshaped, to 2 mm long, truncate, with narrowly subulate bodies, to 1 mm long, yellow when dry, with 2 terminal subulate bristles arising immediately below the margin of the sheath, these bifid above the middle, often longer than the sheath. Inflorescences terminal, sessile or subsessile, capitate, oblong, to 0.6 cm long, 16 Cephaelis kennedyi Dwyer, spec. nov. Suffrutices ad 2 m alti, glabri. Folia angusto-oblonga, 4— 9 cm longa, 0.8-3.0 cm lata, abrupto- acumin nata acu mine ad 1 cm longo, venis lateralibus ca. 20, i 2 mm lon l e bus, ad 0.3 mm vel dente uno; sí. tubo ad 3.2 mm longo; antheris oblongis, ca. 1 mm longis. Fructus a 76 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 to 0.5 cm wide, outer bracts 2, oblong, 5.5-6.0 mm long, ca. 4 mm wide, obtuse, subcoriaceous, glabrous, subplane, with numerous fimbriate glands within at the base; interior bracts 2, somewhat stiff, concave, with a few basal bristles; flowers 8-10, each flower enclosed in 3 bracteoles. Flowers with the hypanthium to 0.5 mm long, the calycine cup 0.5 mm long, somewhat swollen, stiffly petaloid, gla- brous, rigidly petaloid, the teeth variable, sometimes absent, sometimes disposed as 5 minute points, sometimes with 2 unequal teeth, to 0.3 mm long, sometimes with one tooth and sometimes the margin with one oblong petaloid obtuse lobe to 2.5 mm long, and to 1.5 mm wide; corolla with the tube cylindrical and inflated, to 3.2 mm long, rigidly petaloid, glabrous without, barbate within near the fila- ment attachment region, the lobes 5, the anthers oblong, ca. 1 mm long, the filaments linear, attached above the middle of the tube; style to 0.1 mm long, the stigmas 2, 0.2 mm long, the ovarian disc compressed rotund, 0.6 mm long, con- spicuously 2-lobed. Fruits not seen. Cephaelis kennedyi is named in honor of Dr. Helen Kennedy, a student of the flora of Panama. The new species is known only from Panama. It is clearly distinguished from its closest relative, C. mucosa (Jacq.) Sw., by its much re- duced heads and its calycine cup either lacking teeth or exhibiting a great range of variability in the shape and in the size of the teeth. A bladelike lobe on the calyx cup of one flower may be an anomaly. NAMÁ: 6 km from above the Panamerican Highway on the road from El Llano to Cartí-Tupile, Kennedy 1779 (MO). SAN BLAS: trail to Darién from Puerto Obaldía, 10-15 km WSW of Puerto Obaldía, Mori et al. 6859 (MO). 16. Cephaelis latistipulata Standley, J. Wash. Acad. Sci. 18: 281. 1928. TYPE: Costa Rica, Standley 39695 (US, holotype; F, photo). Subshrubs to 1.5 m tall, the stems unbranched, somewhat angular, glabrous, the internodes well spaced. Leaves obovate, elliptic or ovate, 14-21 cm long, (0.5—)7-9 cm wide, rounded or obtuse at the apex, the acumen short, acute and somewhat decurrent on the petiole at the base, the costa prominent, the lateral veins 11-12, coriaceous, smooth, glabrous; petioles elongate, to 7 cm long; stip- ules persistent, kidney shaped or round, 0.8-2.2 cm long, to 2 cm wide, briefly bilobed at the apex, the margin undulate and serrate, the lobes to 10 mm long, acute. /nflorescences in axillary heads, to 1.5 cm long, to 2 cm wide, the peduncle obsolete; involucral bracts 5, ovate or oblong, to 1.5 cm long, to 1.2 cm wide. Flowers sessile, the calycine cup with 5 teeth, these lanceolate, 2-5 mm long, pilose, denticulate; corolla funnel shaped, 4-6 mm long, glabrous or elongate pilose outside, white pilose within near the mouth, the lobes triangular, 1-2 mm long, pilose marginally; stamens subsessile, the anthers oblong, ca. 1 mm long; ovarian disc deeply bilobed, the style 3-5 mm long, glabrous or pilose, the stigmas | mm long. Fruits not seen. Cephaelis latistipulata is known from Costa Rica and Panama. I have drawn freely on Molina's (1951) description of the flower in the above diagnosis and his description of the bracts and bracteoles reads: ‘‘bracteas involucrales 5, cori- áceas, enteras o bipartidas, corto-pilosas en ambas caras, obtusas o acuminadas, ovadas o anchamente oblongas, de 12-15 mm de largo, de 10-12 mm de ancho, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 77 brácteas internas numerosas, pilosas por ambas caras, agudas o acuminadas, obovadas, márgenes sencillo-pilosos, de 10-12 mm de largo, 6-10 de ancho, brac- téolas lineares o oblanceoladas, subcoriáceas, pilosas por ambas caras, de 6-7 mm de largo, de 1-3 cm de largo." cHiRIQUÍ: Caná-Cuasí Trail, Terry 1481 (F). 17. Cephaelis nana (Standley) Standley, J. Wash. Acad. Sci. 17: 171. 1927. Evea nana Standley, J. Wash. Acad. Sci. 15: 105. 1925. type: Panama, Standley 27550 (US, holotype; F, isotype). Subshrubs less than 1 m tall, unbranched, glabrous, the internodes relatively short. Leaves elliptic to oblong, 9-22 cm long, 5-10 cm wide, obtuse to acuminate at the apex and the base, the lateral veins prominent, ca. 15, subcoriaceous, drying green; petioles to 4 cm long; stipules persistent, deeply bilobed, the narrow lobes 4-6 mm long, usually obtuse. /nflorescences solitary, terminal, subsessile or the peduncles to 4 cm, subglobose, to 1.5 cm long, to 3 cm wide, containing 3-5 heads; outermost bracts involucral, obovate or spatulate, acute or rounded at the apex, to 1.5 cm long, to 1.5 cm wide, free, purple, membranous, the internal bracts oblanceolate or spatulate, rarely narrowly oblong, to 1.2 cm long, to 0.6 cm wide, frequently with minute teeth at the base. Flowers not seen. Fruits subglobose, to 1 cm long, to 1 cm wide, fleshy, blue, the pyrenes ovoid or elliptic oblong, ca. 0.5 cm long. Cephaelis nana is known only from Nicaragua and Panama. CANAL ZONE: Hills N of Frijoles, Standley 27550 (F, US). 18. Cephaelis panamensis Dwyer." Type: Panamá, Mori & Kallunki 3714 (MO, holotype). Shrubs, the branchlets woody, smooth, glabrous, the nodes swollen, well spaced except crowded at the apex of the branchlet. Leaves usually sessile, verticillately disposed at the apex of the branchlet, elliptic or obovate oblong, 15- 28 cm long, 7.5-15.0 cm wide, acute or deltoid at the apex, the base 1-2 cm wide, conspicuously truncate, often somewhat auriculate, the costa prominulous above, prominent beneath, to 0.3 cm wide, the lateral veins 15-17, strongly prominulous beneath, the lowermost often strict, the upper arcuate, the undulate submarginal vein inconspicuous, the intervenal areas patulous reticulate, the smaller veins pinnatiform, chartaceous, concolorous, green when dry, minutely puberulent be- neath; petioles absent or to 1.3 cm long, lignose, to 0.3 cm wide; stipules decid- uous, the scars conspicuous, to 6 mm long, to 2.5 mm wide. Flowers not seen. Inflorescences terminal, the fruits massed into a rounded head, to 3 cm in diam., the peduncle absent, the outermost bracts apparently few, triangular oblong, to ephaelis panamensis Dwyer, spec. nov. ?Frutices s glabris. Folia apice ramuli verti- SET d spositis elliptica vel obovato- elliptica 15-28 cm longa, 7.5-15.0 cm lata, basi 1-2 cm lata conspicue truncata saepe auriculata venis lateralibus 15-17 ae eee infra minuto-puberula; petiolis nullis vel ad 1.3 cm longis. /nflorescentiae terminales fructibus oda caespite rotundo, ad 3 cm diam. pedunculo nullo. Fructus sessiles oblongi, ad 1.2 cm longi, glabri 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1.7 cm long, rigid, glabrous except ciliolate at the apex. Fruits sessile, oblong, rotund or truncate at the apex, to 1.2 cm long, smooth, glabrous, blackish green when dry, the calycine cup bowl shaped, ca. 1.5 mm long, to 2.5 mm wide, truncate, not lobed, rigid, minutely ciliolate, each pyrene drying tan brown with a slender median groove. Cephaelis panamensis is known only from Panama. Unfortunately flowers were not collected and I am not certain that the species belongs in Cephaelis. The conglomerate inflorescences appear to be subtended by several bracts. The fact that some leaves are sessile and are basally truncate or vaguely auriculate is diagnostic. The relatively large fruit is more suggestive of Cephaelis than o Psychotria. The calycine cup is toothless and somewhat bowl-shaped. : Rio Guanche, ca. 2.5 km upriver from bridge on road to Portobelo, tropical wet forest ine puis creek to the S of the river, 10-100 m, Mori & Kallunki 3714 (MO). 19. Cephaelis panchocoensis Dwyer.!'? TYPE: Panamá, Gentry & Mori 13895 (MO, holotype). Succulent subshrubs, to 0.75 m tall, the stems somewhat smooth, glabrous, drying red, the superior nodes well spaced. Leaves elliptic or obovate elliptic, 10-35 cm long, 3.5-14.0 cm wide, rounded or obtuse toward the apex, acuminate, the acumen short and wide, to 1 cm long, acute or attenuate acute basally, the costa immersed above, prominent beneath, the lateral veins 20-38, plane or prom- inulous above, leaving the costa at almost right angles and arcuate only near the margin or strict and sharply ascending, prominulous and conspicuous beneath, arcuate, the undulate submarginal vein 2-8 mm from the margin, not conspicu- ously undulate on proximal portion of the blade, the smaller veins inconspicuous, the intervenal areas smooth, chartaceous to subpapyraceous, discolorous, drying red to brown beneath, glabrous and minutely farinose above, minutely puberulent especially on the veins beneath; petioles absent or to 1 cm long; stipules free, ovate lanceolate, to 1.6 cm long, ca. 0.6 cm wide, obtuse?, drying red, subcori- aceous, apparently glabrous. Inflorescences terminal, capitate, 2-4 cm long, 2- 6 cm wide, the head appearing solitary or of 2-6(or more?) smaller heads aggre- gated tightly into a unit head on suppressed branches or several heads on 3 branches terminating a peduncle to 2.5 cm long, stout, to 4 mm wide, glabrous, drying red; bracts numerous, involucral, the outermost oblong, to 2 cm long, about '2 as wide as long, obtuse, stiff, venose, glabrous, the margin irregular, callose, drying tan, with a few glands within. Flowers sessile, the hypanthium ca. | mm long, glabrous, the calycine cup cylindrical, to 3.5 mm long, the teeth 18 Cephaelis panchocoensis Dwyer, spec. nov. Suffrutices succulenti. Folia elliptica, 10-35 cm longa, 3.5-14.0 cm lata, versus apicem rotundata vel obtusa venis PURA bus 20-38 chartacea ad iub rubra ad brunnea supra glabra sed farinosa us in venis minuto-puberula petiolis nullis 0-4.5 cm So stipulis ovato-lanceolatis ad j 6 cm longis, ca. 0.6 cm latis RAF due terminales RM , ad 6 cm i. glabrae pedunculo nullo vel ad 2.5 cm longo e ramis tribus ad 1.5 c cnc te rmina oribus in unum caespitem ad 2.2 cm in diam. vel in oie se: dispositis. Fore subsess les; 'hypanthio glabro cupula calycis cylindrica, ad 3.5 m longa; corolla alba tubo gra ca. 10 mm longo extus glabro; antheris ca. 2 mm longis. Fructus partim bracteolis haar, ia ad 8 mm longi, ue costati glabri, calyce ad 3 mm longo 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 79 5, short, obscured by the ciliate margin, the cup glabrous with a few glands within, these polymorphic, narrow oblong to ovate oblong, to 1.2 mm long; co- rolla white, narrowly cylindrical, to 10 mm long, ca. 0.6 mm wide in the middle, glabrous outside, villosulose within near the mouth, the lobes 5, oblong, to 3 mm long, attenuate above the middle with a minute adaxial projection ca. | mm below the apex; stamens 5, the anthers narrowly oblong, to 2 mm long, subsessile, attached just below the mouth; ovarian disc oblong, to 0.8 mm long, drying black; style linear, ca. 14 mm long, the stigmatic area scarcely wider than the style, to 2 mm long. Fruits oblong, to 8 mm long, ca. 4 mm wide, glabrous, red, lightly costate, the calyx cup persistent or deciduous. Cephaelis panchocoénsis is known only from Panama and Colombia. It is recognized by its many-veined leaves and terminal capitate inflorescences. The leaves dry a pink red beneath, have the lateral veins either leaving the costa at right angles or strict and sharply ascending. The undulate submarginal vein of the blade becomes subparallel to the margin on the lower 4 of the blade. Another diagnostic character is the callose margin of the bracts. The flowers appear to be few per head. My description of the flower is based on Gentry et al. 16756. The calycine cup is strikingly elongate with several glands within. On dissecting the calycine cup of the fruit of Gentry 4767, I observed several triangular glands regularly arranged at the base of the cup; these I was unable to find in dissecting the calycine cup of Gentry 16756. The style is noteworthy in being linear and extending beyond the elongate corolla tube. The new species is related to Cepha- elis dressleri Dwyer, described as new in this paper. Cephaelis panchocoénsis is an upland species, while C. dressleri is a lowland species. ANAMA: DARIEN: La Boca de Pirre, Bristan 1277 (MO). Cerro AE base = on Rio Perre- du. Gentry 4767 (MO). S slope of Cerro Tacarcuna above Río o base camp, 700-1000 m Gentry & Mori bon (MO). E of Tres Bocas, shortest headwater P Río Cuasí, Kirkbride & Duke 1198 (MO). coLoMBiA: cHOCO: Upper Rio Tigre near base of Serrania del Darién, E of Unguia, 250- 300 m, Gentry 2 al. 16756 (MO). 20. Cephaelis tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19. 1796. Tapogomea tomentosa Aubl., ij Pl. Viene 160. 1775. TvPE: not s Psychotria poeppigiana Muell.- Arg. in , Fl. Bras. 6(5): 370. TYPE: Rum seen. Evea tomentosa (Aubl.) Standley, EN a S. Natl. Herb. 18: 123. 1916. Shrubs to 2 m tall, the branches terete, ultimately angular, golden hirsute or villose, often constricted below the nodes. Leaves lanceolate or ovate lanceolate, 8—23 cm long, 3-9 cm wide, acute at the apex, attenuate acuminate, the acumen to 2 cm long, acute, rounded or obtuse at the base, the costa prominulous above and beneath, the lateral veins 9-12, arcuate, the intervenal areas spreading retic- ulate, membranous to chartaceous, densely golden villose above and beneath; petioles to 2 cm long, densely hairy; stipules connate, usually persistent, cylin- drical, the sheath 2-6 mm long, truncate or tapering toward the apex, each part usually with 2 erect narrowly subulate awns to 11 mm long, usually twice the length of the sheath, densely hirsute. Inflorescences terminal, 5-6 cm wide, the flowers conglomerate, the head subtended by numerous bracts, the outer pair large, flaring, deep red; peduncles 2.5-7.0 cm long, densely villose. Flowers with the hypanthium short, pubescent, the calycine cup scarcely measurable, the lobes 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 5, narrowly oblong or narrowly triangular subulate, to 2 mm long, petaloid, with a linear oblong gland at the sinuses to 0.5 mm long; corolla yellow, the tube narrowly cylindrical to 13 mm long, somewhat expanded near the mouth, petal- oid, densely elongate ciliate outside, barbate inside ca. 3 mm below the filament attachment, the lobes 5, triangular, ca. 3 mm long; stamens 5, the anthers sub- sessile, narrowly oblong, 2.5-3.5 mm long, attached at the mouth, usually slightly exserted; style slender, included or barely exserted, the stigmas digitiform, scarcely wider than the style, ca. 2.5 mm long. Fruits berrylike, bright blue, rotund to broadly oblong, less than 1 cm in diam. Cephaelis tomentosa is well distributed throughout tropical America. Standley (1938) remarked that **. . . in most herbaria (it) is represented by more specimens than perhaps any other tropical member of the family. Oddly enough I have not seen any herbarium material from the Province of Chiriqui, Panama; undoubtedly it occurs there." ‘‘Uasika’’ (Cuna, fide Standley). BOCAS DEL TORO: Santa Catalina, Blackwell et al. 2805 (COL, DUKE, MO, UC, VEN). Almi- rante, near Nigra Creek, McDaniel 5050 (MO). Water Valley, Wedel 811 (MO). Chiriquí Lagoon, Wedel 1081, 1279 (both MO). Water Valley, Chiriquí Lagoon, Wedel 1744 (MO). Old Bank Island, Chiriquí Lagoon, Wedel 2118 (MO). Little Bocas, Chiriquí Lagoon, Wedel 2534 (MO). BARRO COL- ORADO ISLAND: Aviles 31 (MO); Croat 4364 (MO); Ebinger 612 (MO); Shattuck 53, 531, 1045 (all MO); Starry 155 (MO); Woodson & Schery 987 (MO). CANAL ZONE: Crossing of Cruces Trail and Madden Dam Road, Bartlett & Lasser 16344 (MO). Near Juan Mina, Bartlett & Lasser 16556 (MO). Cerro Viejo on K 16 C, Blum 1224 (MO). Chagres, 4 mi S of river, Busey 389 (MO). Madden Forest Reserve, Candido F. 1 (MO). Pipeline Road, 0.2 mi N of ATTC Gate, Croat 4697 (MO). Pipeline Road near Gatun Lake, iol 4704 (MO). Radar Station Road 1 mi N of Summit Garden, Croat 10276 (MO). Madden Forest SE of Río Pedro Miguel, D'Arcy & D'Arcy 6099 (MO). Río Azote Caballo, Sabanas, Dodge s r 16844 (MO). Hill C-6, Ft. Sherman, Duke 4401 (MO). George Greene Memorial, Las Cruces Trail, Dwyer 6755 (MO). Chagres, Fendler 177 (MO). Gatún Station, Hayes s.n. (MO). Road between Gamboa and Summit Gardens, Kennedy 2257 (MO). Pipeline Road, Ken- nedy & Andrews 1896 (MO). Navy Reservation N of Gamboa, Robyns 65-54 (MO). Las Cruces, ieu 577 (MO). Pipeline Road, Stern & Chambers 18 (MO). 6 mi N of Gamboa, Tvson 1510 (MO). N of Gamboa, Tyson 6635 (MO). Fort Sherman, Tyson & Blum 1679 (MO). 1 mi N of Summit sud en, Tyson & Blum 1968 (MO). cocLe: El Valle, Aguilar CH3 (MO). Cerro Pilon, 700-900 m Duka 12184 (MO). La Mesa, El Valle, Folsom & Butcher 2807 (MO). Ft. Cerro Pilón, ca. 2000 m, Porter et id 4584 (MO, SCZ). Cerro Valle Chiquito, 700—800 m, Seibert 505 (MO). El Valle de Anton, , Woodson & Schery 171 (MO). Las Margaritas and El Valle, Woodson et al. 1297 (MO). ons n Mina gra Rio pcre Allen 4109 (MO). Santa Rita Ridge, East Ridge, Dwyer " "Waaa $405 (MO). Santa Rita of Agua Clara Rain Gauge, Kennedy 2764 (MO). Santa Rita idge, Lew i et al. 5263 ( Eo M SCZ, UC, eae 0.7 mi NE of Rio Piedras Bridge, Pilon to Portobelo e & Mori 3654 (MO). 8 km NE of Santa Rita, ca. Es : Wilbi ir & Weaver 10840 (MO). DARIÉN: E arde of Río Chico, 500—750 ft, yn 4598 (MO o Chico, across from Boca de Tesca, Duke 5214 (MO). Pico Piriaque, Duke 8138 (MO). Río E 2 mi upstream from Piria, ca. 120 m, Duke 14383 (MO). Puerto St. Dorothea, Dwyer 2228, 2255 (both MO). Quebrada **Camachi- muricate’’ near Casa de Bartolo, Kennedy 2861 (MO). Manené to Mouth of Río Cuasi, Kirkbride & ipt 1543 (MO). Caná, 1750 ft, mde et al. 654 (MO). Los SANTOS: Loma Prieta, Cerro Grande, 2400-2800 ft, Lewis et al. 2240 (COL, DUKE, UC, VEN). AMÁ: dum to Nuevo Emperador, Austin "s (MO). Cerro Azul, Pisis 198 (MO); Castrejon 39 (MO). Cerro Campana, Croat 12139 (MO). Nuevo Emperador, De Garcia 30 (MO). Near Jenené, Du 3882 (MO). Panamá Viejo, Duke 5730 (MO). Cerro Jefe, Duke 9459 (MO). Cerro nena ca. 2000 ft, Dwyer 2124, 3074 (both MO). mpana, ca. 3000 ft, Dwyer & Kirkbride 7854 (GH, MO); Ebinger 337 (MO). Cerro Azul, Ebinger 986 (MO). Nuevo Emperador, Fernandez 116 (MO). Cerro Jefe, Ferris 160 (MO). Cerro Campana, Gentry 1829 (MO). Río Maestro, 2 mi from coast, Gentry 2205 (MO). Cerro ae Gomez- Pompa et al. 3394 (MO); Guerra 6 ~ Nuevo Pian (Arraijan), Guerra 9 (MO). Las Guacas, on 41 (MO). 16 km above Panam n Highway, road from El Llano to Cartí-Tupile, ca. 350—400 , Kennedy 2531 (MO). Peluca eae Station, s & Dressler 2976 (MO). 12-16 km WR Panamerican Highway, El Llano to Cartí-Tupile Road, Kennedy et al. 3128 (MO). 2-3 mi S of Goofy Lake, Lewis et al. 261 (F, GH, K, MO, NY, PA, UC, US). Cerro Campana, Lewis et al. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 81 1920 (GH, MO, UC, US). Altos del Rio Pacora, 2500 ft, Lewis et al. 2302 (MO, UC). Cerro Campana, McDaniel 6887 (MO). Cerro Jefe, Mori 7108 (MO). Cerro Campana, 45 km SW of Panama City, Mori & Bolten 7687 (MO). Road past Cerro Azul, Mori & Kallunki 2182A (MO). 4.5 km N of Lago c. Azul, 675 m, Nee 7037 (MO). Cerro Azul, Porter et al. 4110 (MO, SCZ, UC, VEN). Cerro Campan Porter et al. 4160, nig (both MO). Capira, Saldana 6 (MO). Cerro Jefe, Tyson 3441, 3590 ‘bath MO). Cerro Campana, Webster & Breckon 16505 : MO). Cerro Jefe, Wilbur & Weaver 11096 (MO). Arraijan, Woodson et e 777, 1400 (both MO). SAN BLAS: Between Río Diablo and Río Acuatí, Duke 14893 (MO). VERAGUAS: Cerro Tuti, Folsom & Edw ends 3343 (MO) 21. Cephaelis vultusmimi Dwyer.'® TYPE: Panamá, Gentry & Dwyer 4811 (MO, holotype).—Fic. 17. Subshrubs to 0.5 m tall, the stem smooth, black or dark brown when dry, pilose, the hairs golden, slender, to 2 mm long, the nodes well spaced. Leaves oblong or oblong lanceolate, 9-15 cm long, 3-9 cm wide, deltoid or acute at the apex, acuminate, the acumen to 2.5 cm long, obtuse or truncate at the base, rarely subacute, often conspicuously inequilateral, the costa prominulous above, prominent beneath, the lateral veins ca. 11, strongly arcuate, subprominent be- neath, not forming an undulate subarea vein, rigidly chartaceous, bullate but not drying bullate, concolorous, green with weak curled hairs above, often with withered stiffer and more elongate hairs beneath, those along the costa to 2 mm long, perpendicular; petioles 0.5—1.8 cm long, to 0.3 cm wide; stipules persistent, erect or spreading, free, the body rotund, the lobes 2, erect, widely subulate, 1.5-2.0 cm long, to 0.4 mm wide at the base, caudate tipped, scarious, densely golden hirsute, the hairs to 2 mm long. Inflorescences terminal, solitary, the peduncle to 8 cm long, to 0.5 cm wide including the hairs, densely golden-hirsute or barbate, the hairs to 2 mm long, the head ca. 5 cm wide; the large exterior bracts connate toward the base, triangular to 3.5 cm long, to 4.5 cm wide, bright red toward the base, purple to yellow in the middle, green toward the apex, venose, ciliate on both sides, the hairs similar to those of the leaves. Flowers with the calycine teeth triangular, unequal, to 1 mm long, hirsute marginally with a gland between adjacent teeth, these to ! the length of the tooth Cephaelis vultusmimi is known only from Panama. It is the only new species in this paper without either a complete floral description or a fruit description. The inflorescence as found within the outermost bracts is very immature. I located one bud in which only the hypanthium and the calycine teeth were obvious. The corolla was very immature and little was discernible except the fact that the hairs were very dense. The bracts are strikingly unique and are suggested by the spe- cific name vultusmimi (clown's face). The leaves when seen in the field were bullate; this feature was lost in drying. A 2 x 2 Ektachrome of the species has been deposited in the herbarium of the Missouri Botanical Garden. COLÓN: Santa Rita Ridge Road, E. ridge, Gentry & Dwyer 4811 (MO). 19 Cephaelis uei den Pi spec. nov. Suffrutices ad 0.5 cm alti, caule dense piloso. Folia ded vel oblongo-lanceolata, 9-15 cm longa, 3-9 cm lata, apice deltoidea vel acuta, acumine ad m longo, s Ciel ca. 11, gr eres dense de: Spar 0.5-1.8 cm longis; SUL: liberis lobis 2 erectis lato-subulatis, 1.5-2.0 cm longis, apice audatis, dense hirsutis. Inflorescentiae terminales solitariae forma aes tomentosa simili haste Reno conspicue variegatis rubris purpureis luteis viridibusqu 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 17. Cephaelis vultismimi Dwyer.—A. Habit (x'2).—B. Vegetative node (x1). [After Gentry & Dwyer 4811.] 16. CHIMARRHIS Chimarrhis Jacq., Sel. Stirp. Amer. 61. 1763. TYPE: C. cymosa Jacq. Pseudochimarrhis Ducke, Arch. Jard. Bot. Rio de Janerio 3: 255. 1922. TYPE: P. turbinata (A. P. DeCandolle) Ducke (= Chimarrhis nnd A. P. DeCandolle). Trees. Leaves crowded at the apex of the branchlets; stipules caducous. Zn- florescences axillary, corymbose cymose, bracteate. Flowers small, odoriferous; 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 83 calyx cup campanulate, reduced, truncate or dentate; corolla white, the tube short and broad, villous within, the lobes 5, valvate in bud; stamens 5, inserted in the throat, the anthers dorsifixed, the filaments often exserted, villous at the base; style short, the stigmas 2, obtuse, ovules numerous on an axile peltate placenta. Fruits capsular, small, oblong, dehiscent, septicidally into 2 valves, each valve ultimately divided; seeds numerous, compressed, reticulate, the en- dosperm fleshy. Chimarrhis, a genus of about 14 species, is found in the West Indies, Costa Rica, Panama, and in South America. a. Leaves 10-20 cm long, 6-11 cm wide; stipules 20-30 mm long; fruits ca. 5 mm long ...... l. d latifolia aa. Leaves 3.5-9.5 cm long, 2-5 cm wide; stipules to 11 mm long; fruits ca. 2 mm long |... 2. C. parviflora 1. Chimarrhis latifolia Standley, Publ. Field pou Mus., Bot. Ser. 4: 265. 1929. TYPE: Panama; Cooper & Slater 260 (F, Trees to 25 m tall, the wood yellow, the buttresses high, the ultimate branch- lets angular, smooth, glabrous; nodes well spaced, the stipular scars ringlike, often drying black, the leaf scars +0.5 cm in diam. Leaves widely elliptic to elliptic oblong, 10—20 cm long, 6-11 cm wide, rounded at the apex, short acu- minate, the acumen widely triangular, ca. 0.5 cm long, obtuse, basally cuneate to abruptly attenuate acute, the costa prominulous above, prominent beneath, ca. 0.2 cm wide, rubescent, the lateral veins ca. 10, arcuate, conspicuous above and beneath, uniting to form an undulate submarginal vein, to 0.3 cm from the margin, stiffly papyraceous to thin coriaceous, discolorous, drying dark chocolate above, liver colored beneath, somewhat lustrous, glabrescent; petioles to 4 cm long, angular, twisted, glabrous; stipules deciduous, ovate, 2-3 cm long, acute, thin coriaceous, rubescent. /nflorescences axillary, paniculate, to 12 cm long, to 11 cm wide, the peduncle to 6 cm long, the branches disposed in a fan, to 3 cm long, the branchlets in a fan, the corymbs densely flowered. Flowers short ped- icellate to subsessile; hypanthium turbinate, ca. 2 mm long, glabrous, rubescent, the calycine cup to 0.8 mm long, somewhat expanded, stiffly petaloid, glabrous, teeth absent; corolla white, the tube compressed cylindrical, ca. 2 mm long, stiffly petaloid, rubescent, glabrous outside, white pubescent within above attachment of the filaments, the lobes 5, oblong, longer than the corolla tube, obtuse, pu- berulent within; stamens 5, the anthers ovate oblong, ca. 1 mm long, versatile, the filaments to 4 mm long, tapering to apex, ca. 0.35 mm wide in middle, bearded adaxially along lower 25, attached near the corolla mouth; style to 3.5 mm long, basally narrow, expanded above, the stigmatic lobes oblong rotund, ca. 1 mm in diam., crassate. Fruits capsular, bivalvular, to 5 mm long, the outer part forming a skirt-like structure enclosing the valves, striate, glabrescent. Chimarrhis latifolia is known from Costa Rica and Panama. The flowers were described from Allen 6302 (MO), Jalaca Station, Province of Puntarenas, Costa Rica, which is close to the Province of Chiriqui in Panama. Standley did not describe the flowers in his original diagnosis. 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 The wood was described by the collectors as ‘‘yellow, heavy and hard to split, the grain very crooked and crossed.” ‘‘Jagua amarilla.” CHIRIQUi: Progreso, Cooper & Slater 260 (F, holotype; US, isotype). N . Chimarrhis parviflora Standley, Trop. Woods 11: 26. 1927. TYPE: Panamá, Cooper & Slater 120 (US, holotype; F, NY, isotypes).—Fic. 18. Trees to 18 m tall, the branches smooth, terete, glabrous, ultimately angular, the nodes well spaced. Leaves oblong, 3.5-9.5 cm long, 2.5 cm wide, acute and short acuminate at apex, acute or cuneate at the base, scarcely inequilateral, the costa subplane to prominulous above, subprominent beneath, the lateral veins 9- 11, at first strict, then arcuate toward the margin, the intervenal areas smooth, the veinlets inconspicuous, papyraceous, scarcely discolorous, drying chocolate brown above, glabrous above, appressed villose especially on the costa and the veins beneath; petioles 0.5-1.6 cm long, scarcely pubescent; stipules free, nar- rowly triangular, to 11 mm long, densely appressed pubescent and farinose out- side, glabrescent within. Inflorescences axillary or pseudoterminal, to 4 per axil, minutely puberulent, cymose paniculate, to 15 cm long, to 9 cm wide, the pe- duncle when present 4—9 cm long, arcuate or strict, with 3 or 4 radiate branches terminating the peduncle 1—3 cm long, the cymules hemispherical, densely flow- ered. Flowers on short pedicels to 2 mm long; calyx and hypanthium very short, glabrous truncate the 4-5 teeth obtuse, ciliate; corolla funneliform, ca. 3 mm long, the lobes 4, rounded, glabrous outside, villose within; stamens 4, exserted, the anthers narrowly ovoid, to 0.7 mm long, the ovarian disc annular ca. 0.7 mm long, 0.65 mm wide, 0.8 mm tall; style ca. 2.5 mm long, the stigmas broader than long. Fruits capsular, obovoid or turbinate, ca. 2 mm long, 1 mm wide or almost as wide as long, the valves separating almost to the base, the inner face subplane, scarious, longitudinally grooved, the outer face weakly ribbed; seeds usually 4, plane, subrotund. 0.5-1 mm in diam. the wing enveloping the central glandular shiny fruit body. The species is known only from Panama and Costa Rica. Croat 11244, taken from a fallen tree, has the inflorescences with galls measuring 5 x 4 mm in di- ameter. ‘‘Yema de Huevo." BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 3, 120 (F). Daytonia Farm, Cooper 401 (F, US). PANAMA: Barro Colorado Island, Croat 11244 (MO). VERAGUAS: Valley of Rio Dos Bocas, road between Escuela Agricola Alto Piedra & Calovébora, 15.6 m NW of Santa Fé, Croat 27755 (MO). 17. CHIOCOCCA Chiococca P. Br., Civ. Nat. Hist. Jam. 164. 1756. TYPE: Lonicera alba L. = C. alba (L.) Hitchc. Chiococca L., Syst. Nat., ed. 10. 917. 1789. TYPE: C. racemosa L. Shrubs, vines, or trees, often scandent. Leaves membranous to coriaceous, petiolate; stipules usually cuspidate, persistent. Inflorescences axillary, racemose or paniculate, the racemes often secund. Flowers pedicellate; hypanthium com- pressed; calycine cup 5-lobed, persistent; corolla funnelform or campanulate, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) Od ANO? n FiGuRE 18. Chimarrhis parviflora Standley. Habit (x1). [After Croat 11244.] glabrous throughout, the lobes valvate; stamens with the filaments connate at the base; ovary 2-celled, each cell 1-ovulate. Fruits drupaceous, orbicular, usually laterally compressed, white at maturity. aa. Chiococca occurs in southern Florida, tropical America, and the West Indies. Vines or sprawling shrubs; stamen filaments included in the corolla tube. b. Corolla 3-4 mm long; leaves stiffly papyraceous to subcoriaceous, the lateral veins —5 DENN AEE AA ee a AO = ea ea et ET ERE ba bb. Corolla 6-10 mm long: leaves coriaceous, the lateral veins 5-8 ------------ 2C. durifplia Upright shrubs or trees; stamen filaments exserted. Leaves tapering at apex; corolla with tube 3.6-6.0 mm long -------- 4. C. phanaestemon cc. Leaves obtuse or subrotund at apex; corolla with tube ca. 2 mm long...... 3. C. jefensis 85 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1. Chiococca alba (L.) Hitchc., Ann. Rep. Missouri Bot. Gard. 4: 94. 1893. Lonicera alba L., Sp. Pl. 175. 1753. TYPE: not see Chioccoca racemosa L., Syst. Nat. ed. 10. 917. 1759. + TYPE: not s C. macrocarpa Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles THN 231. 1844. TYPE: not seen. C. petensis Lundell, Wrightia 5: 7. 1972. type: Guatemala, Contreras icine (LL). C. vestita Lundell, Wrightia 5: 8. 1972. rype: Belize, Lond 16425 C. vestita var. glaberrima Lundell, Wrightia 5: 9. 1972. TYPE: not seen. Vines or shrubs (trees?), the twiglets smooth, glabrous, usually ultimately wiry, the branchlets perpendicular, opposite. Leaves ovate oblong, oblong, rarely rotund, 2.2-13.5 cm long, 1.3-6.0 cm wide, tapering acutely, obtuse or somewhat rounded toward the apex, often acuminate, the acumen often arising from a con- tracted apex, 1.0-1.5 cm long, acute or obtuse, often falcate, basally acute, at- tenuate acute, obtuse or rounded, the costa slender, prominulous above and beneath, the lateral veins 4—5, arcuate, uniting to form a submarginal vein 4—5 mm from the margin in the larger blades, the intervenal areas smooth, the fine endings of the venules regularly evanescent, stiffly papyraceous to subcoriaceous, usually glossy above, glabrous; petioles to 0.8 cm long, glabrous; stipules often persistent, widely triangular to oblong, 1.5-5.0 mm long, acute or contracted at the apex into a median subulate awn, to 4 times the length of the body. /nflores- cences axillary, solitary, or often 2 per node, 2.5-8.0 cm long, racemiform or paniculate, the flowers occasionally solitary or in sessile cymules; peduncles slen- der, 0.7-7.0 cm long, ca. 0.35 mm wide, glabrous, the lowermost branches to 4.5 cm long. Flowers pedicellate, the pedicels 2-5 mm long; hypanthium subrotund, somewhat compressed, glabrous, the calycine cup ca. 1 mm long, the teeth acute, 5, about as long as the carnose, glabrous, eglandular cup; corolla white or yellow, urn shaped, 3-4 mm long, glabrous inside and out, petaloid, the lobes 5, oblong, slightly shorter than the tube, tapering obtusely; stamens 5, the anthers at first linear, then narrowly oblong, ca. 4 mm long at anthesis, the filaments slender, 1.2 mm long, pubescent, at the very base connate into a ring adnate to the corolla tube; style linear, 5-6 mm long, the stigmatic area turgid, to 1.3 mm long, the 2 lobes obtuse. Fruits white, fleshy, compressed, 4-8 mm in diam., the persistent calyx short. Chiococca alba ranges from southern Texas through Central America south to Argentina. It also occurs in the West Indies. *‘Lagrimas de Maria BOCAS DEL TORO: Bocas, Maccaw Hills, Wedel 553 (MO). Water Valley, Wedel 1408, 1732, 1787 (all MO). CANAL ZONE: Fort Sherman, Croat 14154, 15425 (both MO). Madden Forest, Croat 15266 (MO). Fort Kobbe, Duke 4207, 4716 (both MO). Albrook, Dwyer 6707 (MO). Madden Dam, Dwyer & Elias 7492 (GH, MO, UC, US); Ebinger 1960 (MO); Kirkbride 55 (MO). Farfan Beach Road, Kirkbride & Elias 65 (MO). Río Petitpie, road to Ft. Sherman from Gatün Locks (S2 or 82), Mori & d 2693 (MO). Cocoli, Tyson 1627 (MO). Fort Sherman, Tyson & Blum 3750 (MO). Curundu, son & Dwyer € (MO). Fort Kobbe, Woodson et al. 1415 (MO). Salamanca Hydrographic Sta- ion. Río Peque . as et al. 1594 (MO). cHiRIQUí: Boquete, Davidson 782 (MO). cocLé: El Valle, Woodson « et al. , 1253, 1760 (all MO). COLON: Portobelo, D'Arcy i (MO). 4 km E of Buena Vista, Nee 6796 o. Achiote, Tyson et al. 4538 (MO). DARIEN: Pucro to Río Pucro, Duke 5369 (MO). Pinas, Duke 10586, 10600, 10639 (all MO). Chepigana, Duke & 1 8 [#278 ?] Pew HERRERA: Las Minas and Pesé, Burch et al. 1322 (GH, K, MO, NY, UC, US). 12.5miSo ú, Lewis et al. 1635 D MO, UC, US). Los sANTOS: Loma Prieta, Duke 11886 (MO). Los Wei Wendehake 35 (MO). PANAMÁ: Nuevo Emperador, ded d (MO). Cerro Campana, Bartlett & Lasser 16908 (MO). es a Larga, Croat 12392 (MO). amé, Dodge 16731 (MO). Rio Canita, Duke 3841 (MO). Rio Mamoni, Duke 5681 (MO). Cerro Prisca Duke 8668 (MO). Isla del Rey, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 87 Duke 10422 (MO). Coronado Beach, Duke 11814 (MO). San José Island, Duke 12517 (MO). Cerro Azul, Dwyer 1509 (MO). Cerro Campana, Dwyer & Kirkbride 7823 (GH, MO). Cerro Azul, Ebinger 1960 (MO). Cerro Campana, Ebinger 1960 (MO). Chagres, Fendler 138 (MO). Panama Vieja, Maurice 797 (MO). Playa de Venado, Mejia 3 (MO). Old Fort Lorenzo, Tyson 1585 (MO). El Llano, Tyson 1742-A (MO). Cerro Azul, Tyson 2093 (MO). Playa Grande, San José Island, Tyson & Loftin 5088 (MO). Arraíjan, Woodson et al. 1404 (MO). vERAGUAS: Río Los Chorros, San Francisco, Dwyer 1354 (MO). Near Ponuga, Dwyer & Kirkbride 7439 (GH, MO, UC, pes Soná, Woodson et al. 545 (MO). SAN BLAS: Soskatupu, Duke 8967 (MO); Elias 1693 (GH, MO, US). 2. Chiococca durifolia Dwyer.2° TYPE: Panamá, Dwyer 11865 (MO, holotype). Vines or climbing shrubs, the terminal branchlets opposite, perpendicular, nigrescent, smooth, glabrous, the nodes well spaced, turgid (persistent leaf bas- es). Leaves oblong, ovate oblong, lanceolate, 3-13 cm long, 1.3-6.0 cm wide, deltoid obtuse or rounded at the apex, often with an acumen to 1.2 cm long, basally rounded, obtuse or cuneate, the costa plane or prominulous, often distally evanescent above, scarcely evident beneath, the lateral veins 4—5(-8), at first strict, then strongly arcuate, the submarginal vein well set in from the margin, coriaceous, smooth, glabrous, usually drying chocolate brown above; petioles to cm long; stipules connate, compressed hemispherical, annular in appearance, ca. 2.5 mm long, much wider than long, apiculate, subcoriaceous, shiny, glabrous. Inflorescences axillary, to 8 cm long, often secund, glabrous or glabrate, racemi- form or scarcely paniculate; peduncle absent or much reduced, to 1 cm long, the branches opposite, few; flowers solitary or in pairs; bracteoles minute, ovate subulate; pedicels to 0.6 cm long or obsolete. Flowers with the hypanthium rotund to subrotund, to 1.5 mm long, glabrate, drying black, the calycine cup scarcely developed, the teeth 5, erect, subulate, ca. 1.2 mm long, acute, puberulent; co- rolla white, the tube subcampanulate, 6-10 mm long, thickly petaloid, glabrous outside, puberulent ca. 2 mm above the base, the lobes 5, deltoid, scarcely shorter than the tube, acute; stamens 5, the anthers linear, 4-5 mm long, obtuse, the filaments slender, ca. 2 mm long, pubescent, connate at the base for ca. 0.5 mm; style slender for almost half its length, expanding above, the stigmas 2, somewhat unequal, oblong, ca. 1 mm long, the ovules oblong, ca. 1 mm long, obtuse. Fruits pedicellate, the pedicels shorter than the berry, white, rotund to oblong, strongly compressed laterally, to 0.8(-10) cm in diam., rounded or obtuse at the base, smooth, glabrous, delicately ribbed and venose, the persistent calyx coroniform, ca. 0.5 mm long. Chiococca durifolia is known only from Panama and Colombia. The relatively abundant material cited has been misidentified as C. pachyphylla Wernham. The well-defined calycine teeth and the flattened fruit distinguish it from this species, whose calycine teeth are almost obsolete and whose fruit is nearly globose in cross section. 20 Chiococca durifolia Dwyer, spec. nov. Vites vel frutices ei di oblonga ovato- oblonga vel lanceolata, 3-13 cm longa, 1.3-6 cm lata, venis lateralibus 5-8 supra evanescentibus, coriacea laevia glabra; petiolis ad 2 cm longis: sills = compre piene a , ad 2.5 mm longis. /nflo- rescentiae pedunculo nullo vel multo reducto. Flores dentibus calycis subulatis, ca. 1.2 mm longis; corolla alba tubo M LES 6-10 mm longo. Fructus pedicellis valde compressis, rotundi, ad 0.8(—10) cm diam., plan 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN (Vol. 67 : La Mesa above El Valle, Dwyer 11865 (MO). El Petrosa, 7 km N of El Copé, Alto Calvari ario, “900 m, Folsom 1330 (MO). COLÓN: es Rita ias Correa & Dressler 906 (MO); Dwyer et al. 9008 (MO); Gentry 1422 (MO). PANAMÁ: Cerro Jefe, Correa & Dressler 467 (MO); Croat 15235 (MO). Cerro Azul, Dwyer 1886 (MO). Cerro Jefe, Elias & Md 1799 (GH, MO). Panamerican Highway to Cerro Jefe summit; 21.7 km from Panamerican Highway, Folsom vira (MO). i Jefe, Kirkbride & Crebbs 9 (MO). Cerro Trinidad, SE slope, Kirkbride & Duke 1649 (MO). Cerro Azul, Tyson 2088 (MO). Cerro Jefe, Tyson et al. 3207 (MO). COLOMBIA: CHOCO: Bin Solano, PD & Fallen 17164 (MO) 3. Chiococca jefensis Dwyer.?! TYPE: Panamá, Dressler 3296 (MO, holotype).— Fic. 19 Trees (?), the branchlets terete, glabrous, the bark cracking, the nodes mod- erately well spaced. Leaves widely oblong, 5.5-13.5 cm long, 2.5-7.0 cm wide, obtuse or subrotund at the apex, acuminate, the acumen short, widely triangular, to 0.8 cm long, acute at the apex and the base, the costa plane or prominulous above, prominent beneath, to 0.12 cm wide, the lateral veins 7-8, arcuate, the marginal vein undulate, 1-2 mm distant from the margin, the venules inconspic- uous above, reticulate beneath, thin coriaceous, scarcely discolorous, farinose punctate above and beneath; petioles rigid, to 1.5 cm long, to 2.5 mm wide; stipules (one seen and this juvenile) connate, the sheath cylindrical, turgid, ca. 5 mm long, coriaceous, glabrous, with each part compressed rotund, ca. 1.5 mm long. Inflorescences terminal, paniculate, to 8 cm long, to 7 cm wide, glabrous or glabrate, shiny; peduncle absent or to 3 cm long, the branches few, spreading; bracts and bracteoles not seen; pedicels to 1 cm long, stiff, smooth, to 0.8 mm wide toward the apex. Flowers glabrous except for the corolla tube within; hy- panthium short, the calycine cup cupuliform, to 2 mm long, to 3 mm wide, con- spicuously truncate, subcoriaceous, the lobes absent; corolla white, the tube compressed cylindrical, ca. 2 mm long, rigidly carnose, the lobes 5, oblong, to 7 mm long, obtuse, conspicuously villose at the base within, otherwise glabrous; stamens 5, exserted, the anthers oblong, ca. 2 mm long, to 1.5 mm wide before anthesis, contorted after anthesis, subbasifixed, the filaments thick, 2-4 mm long, geniculate near the middle, conspicuously villose except near the bend, attached at the mouth; style thick, to 5 mm long, ca. 0.6 mm wide, the stigmas 2, oblong, 2.2-3.0 mm long, obtuse, each stigma longitudinally bilobate. Fruits rotund, to 1 cm in diam., plano compressed, smooth, glabrous, white, the persistent calyx to 1 mm long. Chiococca jefensis is known only from Panama. AMÁ: La Eneida, region of Cerro Jefe, Dressler 3296 (MO). Cerro Jefe, on top near Antenna, ca. 3000 ft, Folsom & Harp 1373 (MO). ?! Chiococca jeu Dwyer, spec. nov. Arbores (?) ramulis glabris. Folia lato-oblonga, 5.5-13.5 cm longa, 2.5-7.0 lata, apice obtusa vel subrotunda acumine brevi lato-triangulari, ad 0.8 cm longo, venis einn: 7-8 tenui-coriacea supra farinoso-punctata; petiolis rigidis, ad 1.5 cm longis, ad 2.5 mm latis; stipulis connatis vagina turgido-cylindrica, ca. 5 mm longa, ca. 5 mm lata. depen centiae terminales paniculatae, ad 8 cm longae, ad 7 cm latae, glabrae pedunculo nullo vel ad ongo. Flores pedicellis ad 1 cm longis; l, brevi cupula calycis ad 2 mm longa, dentibus nullis: corolla tubo compresso-cylindrico, ca. 2 mm longo, lobis 5, ad 7 mm longis; manaa 5, exsertis antheris oblongis, ca. 2 mm longis, filamentis crassis, 2-4 mm ouis “Wie crasso, ad 5 mm longo stigmatibus 2, 2.2-3.0 mm longis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 89 3296.) FIGURE 19. Chiococca jefensis Dwyer.—A. Habit (x!2).—B. Flower (2.5). [After Dressler 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 4. Chiococca phaenostemon Schlecht., Linnaea 9: 594. 1834. TYPE: México, Schlechtendal; not seen.—Fic. 20 C. staminea Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 11: 231. 1844. TYPE: not seen. Shrubs or trees, to 30 m tall, the twigs terete, smooth, glabrous, often nodose (persistent stipule bases). Leaves oblong or falcate lanceolate, 2.5-10.0 cm long, 0.7-3.3 cm wide, tapering to a narrow cusp, to 1.2 cm long, attenuate acute to cuneate at the base, usually slightly inequilateral, the costa plane or prominulous above, prominent beneath, the lateral veins ca. 8, often evanescent, delicately prominulous, at first strict, then arcuate and forking, the intervenal areas indis- tinct, subcoriaceous, lustrous, glabrous, delicately marginate; petioles to 2.5 cm long, ca. 0.12 cm wide; stipules connate, compressed hemispherical or widely deltoid, to 4 mm long, including the median subulate projection. Inflorescences terminal or axillary, when terminal about as long as the uppermost leaves and disposed as a spreading raceme or a narrow panicle, the peduncle 1.5-3.0 cm long, terminated by 3 or 4 primary branches, 2 or 3 of these longer, bearing short branches along their length or unbranched and with cymules. Flowers with the hypanthium oblong rotund, ca. 2.2 mm long, ca. 2.3 mm wide, smooth, glabrous, the calycine cup arising from a constricted base, ca. 0.8 mm long, glabrous, the lobes 5, oblong, ca. 1 mm long, convex in cross section, crassate, glabrous but puberulent marginally; corolla white or yellow, the tube campanulate, 3.6—6.0 mm long, constricted basally, thin carnose, glabrous, the lobes 5, subequal, as- cending, deltoid, ca. 3 mm long, ca. 1.8 mm wide at the base, glabrous; stamens 5, the anthers linear, ca. 4.2 mm long, ca. 0.15 mm wide, the filaments slender, ca. 0.5 mm long, barbate, connate for ca. 0.35 mm, not attached to the corolla tube; ovarian disc doughnut shaped, 0.9-1.0 mm long, 0.5 mm wide, 0.6—0.75 mm high, the style linear subulate, 6.5-9.0 mm long, ca. 0.25 mm wide, narrower toward the base, the ovary bilocular, the ovules 2, oblong, ca. 1 mm long, pendent from the apex of the locule. Fruits on pedicels to 0.3 cm long, oblong rotund to rotund, to 0.5 cm in diam., the calyx persistent, smooth, glabrous, occasionally delicately ribbed or sulcate. Chiococca phaenostemon occurs at elevations between 700 and 1800 m in Mexico, Guatemala, Honduras, Costa Rica, and Panama. 'HiRIQUÉ: Boquete, Dwyer 6966 (MO). Bambito, 1300 m, Maasola 31 (MO). Boquete, Stern et al. 2016 (MO). Volcán de Chiriqui, Terry 1359 (MO). Slopes below Cerro Horqueta, along Quebrada Horqueta, 1600 m, Webster 16704 (MO). Valley of Upper Rio Chiriqui Viejo, White & White 37, 38 (both MO). Finca Lerida, Woodson & Schery 224 (MO). 18. CHIONE Chione DC., Prodr. 4: 461. 1830. TYPE: C. glabra DC. Crusea A. Rich., Mém. Mus. Natl. . Nat., Ser. B. Bot. 5: 204. 1834, non Schlecht. & Cham. 0. TYPE: C. suh d ) A. Rich. Sacconia Endl., Gen. Pl. 451. Trees or shrubs. Leaves often coriaceous, petiolate; stipules often briefly connate, small, caducous. /nflorescences terminal, cymose or corymbose, pe- dunculate; pedicels bracteolate. Flowers with hypanthium turbinate, the calycine w 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 91 FIGURE 20. Chiococca phaenostemon Schlecht.-—Habit (x!2).—B. Flower, partly dissected (x5).—C. Ovary dissected (x5). [After White & White 37.] 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cup with 5 lobes, or the margin simply undulate; corolla reduced, funnel shaped, lacking hairs within, the lobes 5, imbricate, 2 of these exterior; stamens 5, the anthers large, exserted, dorsifixed, the filaments stout, inserted above the base of the tube; ovarian disc swollen, the style stout, the stigmas oblong, exserted, the ovary 2-celled, the ovules solitary, pendulous. Fruits drupaceous, 2-celled; seeds elongate, the testa membranous. Chione occurs in Central America and in the West Indies. There are about 15 species in the genus. It is the most poorly collected among the genera of the Rubiaceae of Panama containing several species. a. Leaves with the lateral veins 4-9; petioles 0.5-1.5 cm long. b. Leaves with the lateral veins ca. 4, thickly coriaceous; fruits rotund .. . C. buxifolia bb. Leaves with the lateral veins 5-9, papyraceous to coriaceous; fruits oblong Or donee oblong. Leaves with the lateral veins 5-6, the blades neither farinose punctate beneath nor barbate in the axils Leaves with the largest blades to 3 cm wide; inflorescences with 2 bran es; fruits ca. 2 cm long ------------------------------------------ . ` campanensis dd. Leaves with the s blades ca. 7 cm wide; inflorescences with branches; fruits ca. 1.4 cm long -------------------------------- 5. C. panamensis ce, d with the lateral veins cn 9, the blades either farinose punctate bene r barbate in the axils .. . C. costaricensis aa. Leaves with the lateral veins 13— 15; ‘petioles 1.8-3.5 cm long... 4. C. darienensis 1. Chione buxifolia Dwyer & Hayden, Phytologia 15: 55. 1967. rvPE: Panamá, Tyson et al. 3291 (MO, holotype; NY, isotype). Shrubs 3-5 m tall, glabrous, the branchlets terete, smooth, rimose. Leaves elliptic or oval elliptic, 2.0-6.5 cm long, 1.5-3.5 cm wide, deltoid or round obtuse at the apex, vaguely acuminate, the acumen obtuse, the costa plane above, prom- inulous beneath, the lateral veins ca. 4, strongly ascending, evanescent beneath when dry, depressed above, the marginal undulate vein well set in from the margin, the margin revolute, coriaceous, concolorous, drying deep brown, lus- trous. Flowers not seen but presumably solitary (the fruits are solitary at the tips of the twigs). Fruits sessile or on pedicels to 0.5 cm long, rotund, to 1.5 cm long, 0.75 cm wide, black; the persistent lobes of the calyx minute, to 1 mm long; seeds solitary, large, hard, the endosperm biporous in cross section. Chione buxifolia is known only from Panama. ANAMA: Cerro Jefe, 6.5 km by road N of Goofy Lake, Cerro Azul, forest remnant on slope, Nee 9308 (MO). La Chorrera, Rodriguez 36 (MO). Cerro Jefe, 2700-3000 ft, Tyson et al. 3291 (MO NY). 2. Chione campanensis Dwyer.? TYPE: Panamá, Wilbur et al. 1020 (MO, holo- type). Trees with the branches crowded, smooth, glabrous, the nodes well spaced. Leaves elliptic oblong, oblong, occasionally ovate oblong, 2.0-6.5 cm long, 1-3 22 Chione campanensis Dwyer, spec. nov. Arbores ?. Folia inar oblonga, oblonga vel ovato- oblonga, 2.6-5.0 cm longa, 1-3 cm lata em versus lato-de ea, basi obtusa vel lato- aes venis lateralibus ca. 6, coriacea ih eus ad 1c Pers stipulis elliptico- ion ad 4m longis, minuto-puberulis. “qan entiae Lae peduncu -]. ongo, ca. 0.1 cm lat Sabes ramis 2 angulari-ascendentibus fructibus paucis. Fructus ra i 0. 8 cm longis, angusto- ios. ad 2 cm longi, ca. 0.7 cm lati, glabri. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 93 cm wide, widely deltoid toward the apex, acuminate, the acumen widely trian- gular, to 3 mm long, obtuse, basally obtuse or widely cuneate, often inequilateral, the costa plane above or scarcely prominulous, prominulous beneath, the lateral veins ca. 6, prominulous beneath and conspicuous, strongly ascending, often branching dichotomously, the intervenal areas spreading reticulate, thin coria- ceous, discolorous, dark red, glabrous; petioles to 1 cm long, ca. 0.1 cm wide; stipules elliptic oblong, to 4 mm long, acute, erect, stiff, minutely puberulent. Inflorescences spreading, the peduncle 1.0-1.2 cm long, ca. 0.1 cm wide, gla- brous, the branches 2, angular ascending, the fruits few (flowers not seen). Fruits with pedicels ca. 0.8 cm long, carnose, narrowly oblong, to 2 cm long, ca. 0.7 cm wide, obtuse at the base, glabrous, shiny, the calycine cup persistent, to 1 mm long, the lobes absent, the margin undulate or with 5 inconspicuous, widely tri- angular lobes, to 0.2 mm long; disc of the ovary persistent, oblong or ovate oblong, to 5 mm long, truncate to obtuse at the apex. Chione campanensis is known only from Panama. It is closely related to C. buxifolia Dwyer & Hayden whose leaves have few lateral veins and are more obtuse at the apex. The fruits of C. campanensis are narrowly oblong, while those of C. buxifolia are rotund. PANAMÁ: Cerro Campana, trail to Campana Ridge, 2300 ft, Wilbur et al. 1020 (MO). 3. Chione costaricensis Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 111. 1940. rype: Costa Rica, Austin Smith 1778 (F, holotype). Chione allenii L. O. Williams, Phytologia 25: 462. 1973. rype: Costa Rica, Allen 5321 (F, holotype). Trees 6—7 m tall, the branchlets terete, smooth, glabrous, ultimately angular, the bark often flaking. Leaves elliptic ovate or lance oblong to oblong rotund, 6.5-14.5(-18) cm long, 3.5-7.0(-11.0) cm wide, acute, obtuse or rounded at the apex, acuminate, the acumen absent or when present widely deltoid, to 0.5 cm long, basally acute, rounded or obtuse, the costa slender, immersed or plane above, prominent beneath, the lateral veins 7-9, arcuate, prominulous beneath, the intervenal areas smooth, spreading reticulate, papyraceous, discolorous, gla- brous or puberulent, barbate in the axils; petioles to 1.5 cm long, ca. 0.1 cm wide in the middle, glabrous; stipules deciduous (one seen), connate, the sheath ca. 1.5 mm long, with each part with an obtuse lobe, to 3 mm long, glabrous. Inflo- rescences terminal, solitary, glabrous, shorter than the terminal leaves, the pe- duncle 2.0—5.5 cm long, the branches opposite, the pairs few, angular ascending, the lowermost to 5 cm long; bracteoles minute, subulate. Flowers (fide Standley) with the corolla tube 5-6 mm long, ca. 0.4 cm wide, somewhat dilated near the mouth, the lobes 4, round elliptic, almost equal to the tube, glabrous within; stamens exserted, the anthers ca. 3.5 mm long, narrowly oblong. Fruits pedicel- late for 1-4 mm, usually with 2 (or more?) often persistent bracteoles at the base, fusiform oblong, often falcately so, to 2.2 cm long, to 0.8 cm wide, delicately longitudinally striate, glabrous, capped by a calycine cup, to 1 mm long, the persistent annular disc protruding for 1 mm in penislike fashion. Chione costaricensis is known from Costa Rica and Panama. COLÓN: Salud, Lao & Holdridge 233 (MO). 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 4. Chione darienensis Dwyer.” Type: Panama, Duke 13571 (MO, holotype). Trees, the uppermost nodes crowded, the branchlets terete, ashen gray. Leaves oblong lanceolate or obovate lanceolate, 13-24 cm long, 4.5-7.8 cm wide, acute toward the apex, acuminate, the acumen to 1.3 cm long, subacute or cu- neate toward the base, scarcely inequilateral at the base, the costa subplane above, prominent beneath, the lateral veins 13-15, prominulous beneath, the intervenal areas evanescent above, spreading reticulate beneath, papyraceous, shiny, glabrous above except for the costa and the veins, the hairs small, white, beneath sparsely appressed villose, barbate in the axils; petioles rigid, 1.8-3.5 cm long, often twisted; stipules not seen. Inflorescences (only one seen) axillary, to 13 cm long, ca. 11 cm wide, the peduncle slender, 4.5-6.0 cm long, arcuately disposed, terminated by 2 branches, these short and slender, to 1.5 cm long, appressed ciliate, with the scars of the pedicels ca. 6 per branch; bracts ovate, to 1.5 mm long, acute, concave, the margins ciliolate. Fruits falcate oblong or obovate oblong, to 2.2 cm long, attenuate toward the base, often minutely pu- berulent, the calycine cup to | mm long. Chione darienensis is known only from Panama. DARIEN: Río Balsa, between Manené and Tusijuanda, Duke 13571 (MO). UA . Chione panamensis Steyermark, Ceiba 3: 19. 1952. TYPE: Panamá, Hagen & Hagen 2137 (NY, holotype; F, isotype). Trees or shrubs, to 25 m tall, the branchlets numerous, terete, smooth, gla- brous, the nodes well spaced, somewhat turgid. Leaves oblong or obovate-ob- long, 3.5-9.5(-10.5) cm long, 2-7 cm wide, widely deltoid to obtuse at the apex, cuneate, obtuse, rounded or truncate at the base, the costa prominulous and grooved above, scarcely prominent beneath, the lateral veins 5-6, opposite, strongly ascending, often forking well below the margin, coriaceous, lustrous, mostly drying dark brown, the margin often revolute on drying; petioles to 1.5 cm long, rigid; stipules not seen. /nflorescences terminal, solitary, glabrous, cy- mose corymbose, 4.0-8.5 cm long, 3-7 cm wide, the peduncle 2.5-4.0 cm long, angular, usually terminated by 3-5 flabellately disposed branches or the branches opposite, few and well spaced, the lowermost to 3.5 cm long; bracts deciduous. Flowers with the hypanthium and calycine cup turbinate, ca. 3 mm long, coria- ceous, glabrous, the cup eglandular within, the teeth not evident; corolla white, broadly cylindrical, ca. 4 mm long (in bud), wider than long, carnose, glabrous, the lobes 5, oblong rotund to compressed triangular, 3.3-4.0 mm long, often wider than long, obtuse or rounded at the apex, carnose, glabrous; stamens 5, the anthers widely oblong, ca. 3.5 mm long, the filaments ca. 3 mm long, thick; style subclavate, ca. 6 mm long, ca. | mm wide, the stigmas 2, oblong, ca. 1.5 mm ? Chione d rds nsis ipi bi spec. nov. sei ee Folia oblongo- lanceolatae vel obovato-lanceolata, 13-24 cm longa, 4.5-7.8 cm lata, acumine ad 1.3 cm longo, venis lateralibus 13-15, papyracea nitida supra glabra exe Mee venasque; petiolis Higidis, 1.8-3.5 cm longis. /nflorescentiae axillares, ad 13 cm longae, ca. 11 cm latae, pedunculo gracili, 4.5—6.0 cm longo; bracteis ovatis, ad 1.5 mm longis. Fructus isto. onic vel obovato-oblongi, ad 2.2 cm longi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 95 long, truncate. Fruits red, oblong, glabrous, to 1.4 cm long, marcescent, glandular shiny, glabrous. Chione panamensis is known only from Panama. HIRIQUi: Cerro Horqueta, Blum & Dwyer 2651 (MO); Hagen & Hagen 2137 (F); Kirkbride 158 (MO). PANAMÁ: El Llano-Cartí Road, 18 km from the Interamerican Highway, Mori & Kallunki 5125 (MO). SAN BLAS: El Llano-Cartí Road, 24.5-25 km from the Interamerican Highway, Mori & Kallunki 5533, 5557 (both MO). 19. CHOMELIA Chomelia Jacq., Enum. Pl. Carib. 12. 1760. TYPE: C. spinosa Jacq. Anisomeris Presl., Symb. Bot. Upsal. 2: 5. 1833. TYPE: A. spinosa C. B. Presl. Shrubs or trees, occasionally armed, the branchlets occasionally reduced in length. Leaves with intervenal areas occasionally lineolate; stipules usually per- sistent. Inflorescences cymose, axillary, the peduncle occasionally reduced in length; bracts and bracteoles small, the latter wasan a connate. Flowers tu- bular, the teeth persistent; corolla white or cream colored, the corolla salverform or funnel shaped, usually sericeous outside, the lobes valvate or imbricate; sta- mens generally not exserted, the anthers dorsifixed, sessile, attached near the mouth: ovary 2-celled, each cell with a pendulous ovule. Fruits fleshy, the seeds cylindrical, pendulous. Chomelia has over 400 species, about 50 of which occur in Central and South America. The genus has the majority of species in tropical Africa and tropical Asia. It also occurs in the Seychelles Islands. There are only 4 genera among the Panamanian Rubiaceae which are armed. Apparently only one of these, Uncaria, is consistently armed. Most species of Randia have spines. In the case of Machaonia and Chomelia only rarely are the species with spines. Literature: Steyermark, J. S. 1967. Chomelia in Botany of the Guayana Highland. Mem. New York Bot. Gard. 17: 333-334 a. s armed. . Leaves with intervenal areas not lineolate _ usss. 9. C. spino “a Leaves with intervenal areas lineolate ------------------------------------——- i C. llanto aa. Pu unarmed. Leaves with intervenal areas lineolate __--_-__________-------------------------- 1. C. atlantica cc. Leaves with intervenal areas not lineolate d. Sprawling shrubs; leaves with lateral veins ca. 6 _____.----------_--- 7. C. psilocarpa dd. Trees or upright shrubs; leaves with lateral v veins ruo: 18). Leaves ashen white beneath; blades 7.0—18.5 cm long ------ 4. C. leucophylla ee. Leaves not ashen white beneath; vali 4.5— 12.0 0 cm long. f. Flowers and fruits not secundly disposed. g. Inflorescence with peduncle scarcely measurable. |... 8. C. recordii gg. Inflorescence dm peduncle 1-3 cm long h. Leaves coriaceous; lateral veins 7-9. i. Leaves with petioles 0.5 cm long; stipules to 0.35 m long |... 3. C. endo arpa 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 H. Leaves with petioles to 1.5 cm long: Li: to | em Bis " coclensis s chartaceous; lateral veins 6 ` 6. — peninsularis ff. Flowers us fruits secundly disposed .. - _.. 5. C. panamensis 1. Chomelia atlantica Dwyer.** rvPE: Panama, Kennedy & Gra 2242 (MO, ho- lotype). Trees or shrubs to 6 m tall, the branchlets short or long, to 13 cm long, the bark gray, densely appressed pubescent or glabrescent, the nodes well spaced, at times armed, the spines solitary, to 0.4 cm long, rigid. Leaves oblong, 3-14 cm long, 1.5-7.5 cm wide, deltoid toward the apex, acute, basally attenuate acute, rarely subcordate, sometimes inequilateral, the lateral veins 8-10, strongly ar- cuate, the intervenal areas conspicuously lineolate, chartaceous, discolorous, pubescent above on the costa and veins, with elongate appressed hairs on the costa and the veins beneath, the hairs of the intervenal areas small, strigose; petioles slender, to 2.2 cm long; stipules free, often deciduous, narrowly trian- gular, to 10 mm long, acute with a median keel, densely pubescent especially toward the base. Inflorescences axillary, perhaps pedunculate; flowers ca. 3; bracteoles connate at the base of the hypanthium, each part provided with a conspicuous bristle to 2.5 mm long. Flowers green or cream colored, the hypan- thium oblong rotund, ca. 2 mm long, the hairs long, appressed, white, the calycine cup to 1.5 mm long, eglandular within, truncate, the lobes 4-5, subulate, to 6 mm long or longer, often curled; corolla with the tube 1.7-2.0 cm long, ca. 0.1 cm wide in the middle, with appressed, white and dense hairs outside, glabrous within, the lobes 4, narrowly ovate oblong, ca. 8 mm long, ca. 2 mm wide at the base, with a median keel, the latter with elongate, appressed hairs; stamens 4, somewhat included, the anthers narrowly oblong, 3.3 mm long, obtuse, the fila- ments ca. 0.5 mm long, attached near the mouth, the style linear, 4-11 mm long, the stigmas 2, ca. 1 mm long. Fruits not seen. Chomelia atlantica is known only from Panama. It is immediately distin- guished by the intervenal areas of the leaves being lineolate and the inflorescence being epedunculate. It is probably related to C. tenuiflora Bentham from Surinam, British Guiana, and Venezuela, which also has the bracteoles united below the hypanthium and a suppressed peduncle. : Swamp forest and ~ tierra firma, ca. 0.5 km from Río Buena-Ventura, near Portable. Ke nnedy & Gra 2242 (MO). vs EUM Pirre, Bristan 567 (MO). Caná-Cuasí trail (Camp 2), Chepigana, Terry & Terry 1448 (FP. M . ere atlantica ies spec. nov. Arbores vel frutices ad 6 m alti, interdum armati. Folia PM nga, m longa, 1.5-7.5 cm ma ad apicem deltoidea venis lateralibus 8-10 interveniis con- spicue a. subtus ciliis elongatis appressis in costa venisque si Menai petiolis ad 2.2 cm longis; stipulis ad 1 cm longis. idee eee fortasse epedunculatae iro us ca. 3 bracteolis asa cu- as freer basi hypanthii ad 1 mm longis, seta ad 2.5 mm lon "Flores cupula calycis ad 1.5 mm longa, truncata, dentibus 4-5 subulatis, ad 6 mm longis; Sci viridi v el cremea tubo ad 20 mm longo, lobis ca. 8 mm longis; antheris ca. 3.3 mm longis. Fructus non visi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 97 2. Chomelia coclensis Dwyer.” TYPE: Panamá, Croat 25396 (MO, holotype). Shrubs to 3 m tall, the branchlets terete, somewhat smooth, lightly puberulent, ascending at a sharp angle, the nodes well spaced, the lenticels many but incon- spicuous. Leaves ovate-oblong or oblong, 4.5-10.5 cm long, 2-5 cm wide, deltoid or subobtuse at the apex, acuminate, the acumen triangular to 0.8 cm long, ul- timately obtuse, obtuse or rounded at the base, the costa prominulous above, somewhat prominent beneath, the lateral veins ca. 7, arcuate, conspicuous be- neath, the smaller veins not conspicuous, coriaceous, discolorous, dark brown and glabrous above, beneath with a few white hairs especially on the costa and the veins; petioles rigid, woody, to 1.5 cm long, to 1.5 mm wide; stipules decid- uous, only seen at the apex of the twigs, rhombic oblong, to 1 cm long, 0.6 cm wide, acute, subcoriaceous, puberulent, the veins crowded, fanlike. /nflores- cences subterminal, axillary, cymose paniculate, to 7 cm long, to 3 cm wide, the peduncle to 3 cm long. Flowers with the hypanthium oblong, glabrous, ca. 1 mm long, the calycine cup expanded, 0.1 mm long, the lobes 4, triangular, 0.1 mm long; corolla white, the tube cylindric funneliform, ca. 1.5 mm long, petaloid, glabrous without, densely white villose toward the mouth, the lobes 4, ovate, to 2.5 mm long, to 1.8 mm wide, ultimately obtuse; stamens 4, the anthers oblong, to 0.7 mm long, dorsifixed, the filaments scarcely longer than the anthers, at- tached near the mouth; stigmas 2, oblong, to 0.3 mm long, obtuse, the style ca. 2 mm long, the ovules one per locule, pendent. Fruits not seen Chomelia coclensis is known only from Panama. It is recognized by its cori- aceous leaves and rigid, elongate peduncles. The flowers are much reduced in length. COCLÉ: La Mesa above El Valle, 800 m, Croat 25396 (MO). 3. Chomelia grandicarpa Dwyer.” TYPE: Panamá, Kennedy & Foster 2174 (MO, holotype) Trees to 6 m tall, the branchlets terete, glabrescent, smooth, moss covered. Leaves oblong, ovate-oblong or ovate, 4.5—11.0 cm long, 2.5-5.5 cm wide, deltoid to subacute at the apex, acuminate, the acumen to 1 cm long, ultimately acute, 25 Chomelia coclensis Dwyer, spec. nov. Frutices ad 3 m alti. Folia ovato-oblonga vel oblonga 4.5-10.5 cm longa, 2-5 cm lata, versus apicem deltoidea vel subobtusa basi obtusa venis lateralibus ca. 7 coriacea; petiolis ad 1.5 cm longis; stipulis see MM ad 1 cm longis. /nflorescentiae ad 7 cm longae, by 3 cm latae, pedunculo rigido, ad 3 cm longo. Flores hypanthio glabro cupula calycis ca. 0.1 mm longa; d alba tubo ca. 1.5 mm longo, lobis 4 ad 2.5 mm longis; antheris ad 0.7 mm longis. mee tus non v 36 Mri Yee Mia Bes r, spec. nov. Arbores ad 6 m altae. Folia oblonga, ovato-oblonga 1.0 c vel ovat onga, 2.5-5.5 cm lata, venis lateralibus 6-8 venis venulisque minoribus non visis iso papyracea e tenui-coriacea, in costa subtus villosa; stipulis po triangularibus; pe- tiolis ad 0.5 cm longis. /nflorescentiae axillares pedunculo forte gracile, ad 4.5 cm longo, floribus 2- 5 in ramis 2 pue reductis dispositis. Flores sessiles cupula calycis carnosa extus appresso-villosa lobis 4 inconspicuis. Fructus oblongi, ad 3.2 cm longi, glabri rubri carnei, seminibus 2 oblongis, ad 2 cm longis, proxime apicem truncatis ad basin attenuatis conspicue rostratis in sicco purpureo-nigris laevibus inconspicue sulcatis. 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 rounded or obtuse at the base, the costa slender, prominulous above and beneath, drying red, the lateral veins 6-8, arcuate, forming an undulate, submarginal vein well in from the margin, the intervenal areas smooth, the smaller veins not visible, rigidly papyraceous to thin coriaceous, drying deep red brown above, lighter beneath, glabrous above and beneath but moderately villose on the costa below; petioles stiff, terete, to 0.5 cm long, glabrate; stipules persistent only at the tips of the twiglets, triangular, to 3.5 mm long. Inflorescences axillary, the peduncle slender, 1.2-4.5 cm long, ca. 0.15 mm wide, often curved, pubescent, terminated by 2 flowers or by 2 short, spreading branches, to 8 mm long, each branch with 2-5 flowers; bracts and bracteoles not seen. Flowers (in young bud) sessile, the hypanthium scarcely measurable, the calyx urn shaped, ca. 1 mm long, carnose, appressed pilose outside, the margin subtruncate with 4 compressed-rotund lobes, the latter scarcely measurable; corolla tube cream colored or pink and streaked, carnose, pubescent outside, the lobes valvate; ovules pendent, one in each of the 2 cells, the style cylindrical. Fruits presumably rotund when fresh, fleshy, red, when dried oblong, to 3.2 cm long, to 2.5 cm wide, the seeds 2, oblong, to 2 cm long, wider at the apex than at the base, truncate toward the apex, conspicuously rostrate, the beak triangular, to 0.4 cm long, obviously attenuate toward the base, drying purple black, smooth, glabrous, vaguely sulcate. Chomelia grandicarpa is known only from Panama. The fruit is of extraor- dinary size for a Chomelia, and in the pressed condition appears like a young fruit of Pterocarpus (Leguminosae). This character, coupled with the few flowers per inflorescence and the subcoriaceous texture of the leaves, readily segregates the species from the remaining species in Panama. The fact that Folsom 3696 has pink flowers that are streaked is noteworthy. COLÓN: Santa Rita Lumber Road, 8.7 km from Transisthmian Highway, Folsom 3696 (MO). 2 mi up Río Guanche, lowland rain forest, 10-20 m, Kennedy & Foster 2174 (MO). 4. Chomelia leucophylla Dwyer.” TYPE: Panamá, Gentry 4262 (MO, holotype).— Fic. 21 Small trees, 4—10 m tall with the trunk branching above, the twigs terete, rigid, pubescent, the nodes well spaced. Leaves oblong elliptic, oblong or sub- rotund, sometimes falcate, 7.0—18.5 cm long, 2.2-11.0 cm wide, acute or deltoid at the apex, the acumen widely triangular, to 1 cm long, often falcate, at the base rotund or widely cuneate, the costa slender, prominulous above and beneath, the lateral veins 7—9, arcuate, thin chartaceous, discolorous, brown above, ashen white beneath, glabrous above except for the costa and veins, with the hairs beneath crowded, minute, white; petioles 0.5—2.3 cm long, to 0.13 cm wide; stipules free, narrowly lanceolate, 5-7 mm long, acute, the hairs curled. Inflo- 27 Chomelia leucophylla Dwyer, spec. nov. Arbores 4—10 m altae. Folia viue elliptica, ob- longa vel subrotunda, 7-18.5 cm longa, 2.2-11 cm lata, venis lateralibus 7-9 t -chartacea supra brunnea infra albo-cinerea; deu ad 2.3 cm longis; stipulis 5-7 mm longis, rd Inflorescentiae ad 3 cm longae, pedunculis 1-1.5 cm longis. Flores sessiles vel pedicellis ad 1.5 cm longis; cupula calycis brevi lobis 4 triangularibus subulatis vel foliaceis, 0.7-3 mm longis: corolla tubo ad 13 mm longo, lobis 2.5-3 mm longo; antheris ca. 2.2 mm longis. Fructus oblongi, ad 7 mm longi, puberuli. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 99 FiGuRE 21. Chomelia leucophylla Dwyer.—A. Habit (x4.5).—B. Flower, opened to show in- terior (x2.8).—C. Ovary cross section (7.5). [After Holdridge 6452. ] rescences axillary, to 3 cm long, the peduncles 1.0-1.5 cm long, slender, white puberulent, the flowers congested on short branches, the bracts and bracteoles small. Flowers sessile or subsessile, to 1.5 cm long, the hypanthium rotund, ca. 1.5 mm long, densely pubescent, the cup of the calyx short, provided with a few minute red glands within, the lobes 4, triangular, subulate or foliaceous, 0.7—3.0 mm long, acute, glabrous within, often reflexed; corolla with the tube narrowly 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cylindrical, 10-13 mm long, 0.8 mm wide in the middle, puberulent outside, pu- bescent within near the base, the lobes 4, subrotund, 2.5-3.0 mm long, with the margins crisp, glabrous within; stamens 4, included within the tube, oblong, 2.2 mm long, obtuse, the filaments short, ca. 0.5 mm long, attached slightly below the mouth; style to 5.5 mm long, ca. 0.25 mm wide, the stigmas narrowly oblong, ca. 3 mm long. Fruits immature, sessile, oblong, to 7 mm long, to 3.5 mm wide, truncate above, tapering at the base, puberulent, drying brown, with 6 ribs. Chomelia leucophylla is known only from Panama. It is readily recognized by the white undersurface of the leaf blades as suggested by the specific epithet. COLON: Lower Rio Guanche, Dressler 4346 (MO). Rio Trapiche, Holdridge 6452 (MO). Rio Guanche, ca. 3 km upriver from bridge, road to Portobelo, Mori & Kallunki 5181 (MO). DARIEN: Cerro Chucula, Gentry 4262 (MO). 5. Chomelia panamensis (Standley) Dwyer, comb. nov. Antirhea panamensis Standley, N. Amer. Fl. 32: 264. 1934. type: Panamá, Cooper 238 (F, holotype; Chione Ferne Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 138. 1967. Type: Panamá, Stern & Chambers 144 (MO, holotype). Trees of medium height, to 12 m tall, the bole slender, the crown small, the twigs smooth, the branchlets smooth, rimose, glabrescent, occasionally lenticel- late, the leaves often persistent and verticillate at tips. Leaves ovate, oblong, elliptic oblong, occasionally oblong rotund, 6-13(-18) cm long, 3.0-6.5 cm wide, apically acute and acuminate or obtuse and abruptly acuminate, basally cuneate or obtuse, the lateral veins 7-8, usually immersed above, prominulous beneath, the intervenal areas often striate lineolate, stiffly membranous to rarely stiffly papyraceous, concolorous or not, glabrate or short pilose on the costa and the veins and often with short strigose appressed hairs in the intervenal areas; petioles slender, 1.0-2.5(—5.0) cm long; stipules deciduous, ovate elliptic, 4.0-5.5 mm long, attenuate at apex, glabrous to puberulent, ciliate marginally. Inflorescences terminal and/or axillary, cymose, shorter than or slightly longer than the upper- most leaves, the peduncles to 6 cm long, 0.4-0.8 mm wide, strigillose, twisted, the first 2 branches from apex of peduncle short and soon bifurcating to form branches to 3.5 cm long, often conspicuously spreading, the flowers several, secundly disposed, ca. 1 mm apart. Flowers sessile; hypanthium oblong, 1 mm long, glabrous to puberulent, the calycine cup and the 4—5 oblong-rotund lobes ca. 0.5 mm long, glabrous within, the lobes minutely ciliolate, carnose; corolla greenish yellow, the tube ca. 7.5 mm long, 0.6 mm wide at the middle, puberulent outside, glabrous within except hirsute at very base of the tube, carnose, the lobes 4, oblong rotund, 1.5 mm long, or 3 of 4 equal in width, compressed rotund to oblong, 1 mm long and the 4th oblong, 1.8 mm long, carnose, glabrous within; stamens 4, the anthers linear oblong, ca. 3 mm long, conspicuously sagittate at the base, subbasifixed, the filaments ca. 0.5 mm long, attached to the tube 2-3 mm below the mouth; ovarian disc hemispherical, ca. 0.3 mm long, the style linear, 6-8 mm long, the 2 stigmas narrowly oblong, slightly wider than the style, 0.7 mm long, the ovary thick walled, the ovules 1 per cell, pendulous, narrow oblong, ca. 0.7 mm long. Fruits sessile, fleshy, oblong, 1.0-1.3 cm long, rounded, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 101 obtuse or truncate at the apex, smooth, glabrous, lightly costate, the ribs irregular and inconspicuous, black, and drying black; the calyx not persistent, the scar evanescent; seeds 2, enclosed by a very thick, hard endocarp. Chomelia panamensis is known only from Panama and Colombia. The wood, bright orange in color, is used in interior construction, for axe handles, paddles, etc. The fruit, while juicy, is not eaten (fide Duke). ‘‘Tablis’’; *‘Tapaliso’’; **Ta- palizo.”’ CHIRIQUÍ: Puerto Armuelles, Stern & Chambers 144 (MO). DARIEN: Río Ucurganti, Bristan 1167 (MO). Rio Yapé, Bristan 1372 (MO). Without other locality, Bristan & Duke 179 (MO). Santa Fé, Rio Sabana, Duke 4119 (MO). Rio Chico across from Boca de Resca, Duke 5220 (MO). Santa Fé, 9 (M 6290 (MO). Foothills, N slope of Cerro Pirre, Mori & Kallunki 5412 (MO). Campamento Buena Vista, Río Chucunaque above Río Tuquesa, Stern et al. 825 (MO). 3 mi N of Santa Fé, Tyson et al. 4639 (MO). 3 mi E of Santa Fé, Tyson et al. 4704 (MO). 2 mi E of Santa Fé. Tyson et al. 4842 (MO). Los SANTOS: Loma Prieta, 800-900 m, Duke 11889 (MO). PANAMÁ: Between Canasas and Sabalo, Duke 14472 (MO). SAN BLAS: Permé .G. P. Cooper 238 (F). COLOMBIA: CHOCO: | mi NE of Camp Curiche, 3 mi E of Curiche, Duke & Idrobo 11320 (MO, NY). 6. Chomelia peninsularis Dwyer.?* rype: Panamá, Croat 22440 (MO, holotype). Shrubs to 1.3 m tall, the branchlets numerous, terete, glabrous, the tips villose, densely lenticellate, the nodes well spaced. Leaves oblong to vaguely oblong rotund, occasionally ovate, 5.5-15.0 cm long, 2.8-8.0 cm wide, obtuse to deltoid at the apex, the acumen triangular, ca. 1 cm long, ultimately acute or obtuse, acute to cuneate at the base, the costa prominulous above and beneath, the lateral veins ca. 6, rigid and angular ascending for about 34 of their length then curving arcuately to form an undulate vein to 1 cm from the margin, the smaller veins between the laterals scarcely visible, stiffly chartaceous, somewhat discolorous, drying dark green above, smooth, glabrate with a few hairs on the costa, often barbate in the vein axils beneath, the hairs dense-ferrugineous; petioles stiff, 0.3— 0.8 cm long, ca. 1 mm wide in the middle, sparsely villose; stipules triangular, to 4 mm long, stiff, appressed pubescent. /nflorescences axillary or pseudo terminal, usually 2 per node, the peduncles wiry, 1.8-2.5 cm long, to 0.4 mm wide, glabrate, the peduncle ending as 2 spreading branches, the flowers 15 to 25 per inflores- cence, disposed as a few secund cymules or solitary, sessile or subsessile. Flow- ers with the hypanthium oblong or turbinate, ca. 1.5 mm long, obtuse or rounded basally, the hairs few, the calycine cup scarcely measurable, the teeth usually 5, triangular or compressed rotund, to 0.3 mm long or inconspicuous; corolla white (in bud) to 6 mm long, densely white puberulent outside, carnose, barbate basally 28 Chomelia peninsularis Dwyer, spec. nov. Frutices ad 1.3 m alti ramulis crebris teretibus. Folia oblonga vel oblongo-rotunda interdum ovata, ad 15 cm longa, ad 8 cm lata, basi acuta vel cuneata, venis lateralibus ca. @ vena submarginali undulata a margine bene distante, rigido-chartacea laevia solum in costa et venis sparse pubescentia axillis saepe barbatis; petiolis 0.3-0.8 cm longis, sparse villosis; stipulis see ad 4 mm longis, appresso- pubescentibus. Infloresc entiae axillares vel Flores sessiles vel subsessiles, hypanthio ca. 1.5 mm longis, dentibus inconspicuis vel 5 et ad 0.5 mm longis, obtusis vel acutis. Fructus non visi. 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 within, the lobes valvate, to 2 mm long; stamens 5, the anthers oblong, ca. 1.3 mm long, obtuse, usually exserted, the filaments shorter than the anthers; style slender, ca. 7 mm long, the stigmas thumblike, ca. 2 mm long; ovules 1 per cell, pendulous. Fruits not seen. Chomelia peninsularis is known only from lowland Chiriqui. It is readily dis- tinguished from C. panamensis (Standley) Dwyer, a large tree which is distributed primarily in Darien Province and in the neighboring Province of Choco, Colombia. The new species is a shrub with thicker leaves and with fewer lateral veins. CHIRIQUi: Burica Peninsula near Punta de Piedra, secondary forest, Croat 22440 (MO). 7. Chomelia psilocarpa Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 139. 1967. TYPE: Panamá, Hayden 143 (MO). Scandent shrubs, often in treetops, the branches glabrous, smooth, often drying white. Leaves elliptic, 6.5-13.0 cm long, 2.5-6.0 cm wide, acute at the apex, acuminate, the acumen to 1 cm long, apiculate, the apiculum ca. 1 mm long, attenuate acute at the base, the costa prominulous beneath, the lateral veins ca. 6, alternate, strongly arcuate, papyraceous, glabrate above, the hairs short, white, deciduous, beneath ciliolate, the veins appressed pilose, with the shorter hairs seriately disposed, densely barbate in the axils of the veins; petioles 0.7— 2.0 cm long, appressed subhirsute; stipules triangular or subulate, to 1 cm long, ca. 0.35 cm wide at the base, elongate acuminate, smooth, glabrate, diffusely pilose. Inflorescences axillary, cymose paniculate, to 9 cm long, ca. 6 cm wide, the peduncles 4-7 cm long, ca. 0.15 cm wide, stiff, ascending, 1-7 cm long, the cymules few flowered; bracts and bracteoles linear subulate, to 3 mm long, gla- brate, the bracteoles of the pedicels several, aggregated at the end of the pedicel. Flowers not seen. Fruits sessile or pedicellate, the pedicels to 2 mm long, elliptic, ca. 1 em long, to 0.6 cm wide, obtuse, the persistent lobes of the calyx 4, erect, linear subulate or linear ligulate, the 2 shorter 2-3 mm long, the 2 longer 5-6 mm long, the exocarp carnose, densely appressed pilose. Chomelia psilocarpa is known only from the Canal Zone. CANAL ZONE: BARRO COLORADO ISLAND: Croat 7931, 10307 (both MO); Hayden 143 (MO). 8. Chomelia recordii Standley, Trop. Woods 7: 9. 1926. TYPE: Guatemala, Record 8862 (Y, holotype; NY, isotype). C. englesingii Standley, Trop. Woods 16: 45. 1928. rype: Nicaragua, Englesing 49 (F, holotype). Anisomeris recordii (Standley) Standley. N. Am. Fl. 32: 227. 1934. A. englesingii (Standley) Standley, N. Am. Fl. 32: 227. 1934. a. Calyx with lobes lanceolate, villose outside _...... 8a. C. recordii var. recordii aa. Calyx with the lobes narrowly subulate, glabrous outside ` ordo cma ES E RR ENS 8b. C. recordii var. paucipube scens 8a. Chomelia recordii Standley var. recordii. Trees or shrubs, rarely armed, the spines lignose, to 1 cm long, the branchlets distichous, often short, terete, smooth, drying grey, glabrous or ultimately pu- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 103 bescent. Leaves oblong, ovate oblong, occasionally oblong rotund or subfalcate, usually 2 times as long as broad, 6-11(—12) cm long, 2.5-5.5 cm broad, tapering obtusely or rounded at the apex, occasionally deltoid, acuminate or only vaguely so, the acumen to 0.8 cm long, obtuse, basally cuneate to obtuse, occasionally rounded, the costa plane or prominulous above, the lateral veins 6-8, occasionally subimmersed above, arcuate, usually forking toward the margin, chartaceous, scattered villose above, occasionally densely white villose beneath, the hairs elongate, spreading, occasionally confined to the costa and the veins, the inter- venal areas glabrous or pubescent, the hairs often tufted in the axils, the margins often sparsely hirsute; petioles to 0.5 cm long, densely villose; stipules free, ovate lanceolate, to 8 mm long, acute, densely villose. Inflorescences axillary, of 2-3 flowers on a scarcely measurable and densely bracteate peduncle terminating a short or rarely elongate branchlet to 8 cm long; bracts foliose, lanceolate, to 5 mm long, acute, venose, villose. Flowers sessile; hypanthium oblong, ca. 1 mm long, densely appressed white villose, the calycine cup ca. 2.5 mm long, petaloid, villose outside, glabrous within, the lobes 4, unequal, lanceolate, to 3.5 mm long, acute, each with a distinct median vein; corolla white, the tube narrow cylindrical, ca. 17 mm long, 0.5-0.8 mm wide, petaloid, glabrate or densely appressed villose outside with white hairs, to 1.3 mm long, glabrous within, the lobes 4, oblong, ca. 5 mm long, obtuse, petaloid, glabrous adaxially, villose to glabrate within, the margins revolute, short fimbriate; stamens 4, the anthers sessile, narrowly oblong, ca. 2.8 mm long, attached ca. 2 mm below the mouth; style linear, ca. 17 mm long, the stigmas 2, narrowly oblong, ca. 1.5 mm long. Fruits oblong, ca. 8 mm long, ca. 3.5 mm wide, truncate, obtuse at the base, black or purple red, juicy, the hairs white, to 1.5 mm long, the calycine teeth ca. 5 mm long. Chomelia recordii is known from Guatemala, Nicaragua, Costa Rica, Panama, and Colombia. ‘‘Pata de Gallina" (fide Stern et al. 254) DARIÉN: Between Pinogana and Yaviza, Allen 244 (MO). Río Ucurganti, Bristan 1177 (MO). La Boca de Pirre, Bristan 1281 (MO). El Río Aruza, Bristan 1365 (MO). El Río Yapé, Bristan 1407 (MO). Río Uruceca, Bristan 1447 (MO). Without other locality, Bristan 1509 (MO). Río Pirre, Duke 4962A (MO). Río Chucunaque near Quebrada Bolcillo, Duke $580 (MO). 2 mi N of Santa Fé, Duke 10235, 10715 (both MO). 1-4 mi N of Pucro, Duke 13031 (MO). Quebrada Maskia off Río Pucro, above Pucro, Duke 13098 (F, MO). Between Canasas and Sabalo, Duke 14490 (MO). Trail from Paya to Payita, Stern et al. 254 (MO). Campamento Buena Vista, Río Chucunaque above Río Pie k Stern et al. 865 (MO). 2-3 mi N of Santa Fé, Tyson et al. 4626, 4833, 4843 (all MO). PANAM Cuipo forest, 28.5 km E of Bayano Bridge, Folsom 3538 (MO). COLOMBIA: CHOCO: "piel Casi- hada 6209 (COL). 8b. Chomelia recordii Standley var. paucipubescens Dwyer.?? TYPE: Panama, Gentry & Tyson 5038 (MO, holotype). Shrubs, the branches short, often with reduced spines at the tips of the short branchlets. Flowers with the calycine lobes narrowly subulate, glabrescent out- side. Chomelia recordii var. paucipubescens Dwyer is known only from Panama. 2m Chometia recordii Standley var. paucipubescens Dwyer, var. nov. Frutices ramulis brevibus saepe armati spinis se terminalibus. Flores lobis calycis linsati: subulatis glabrescentibus; co- rolla tubo extus glabrescent 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 CANAL ZONE: Near archeological site, edge of Madden Lake, Gentry & Tyson 5038 (MO). 9. Chomelia spinosa Jacq., Enum. Pl. Carib. 12. 1760. Type: Colombia, "'in syl- vaticis ad pedem montis de La Popa.” Jacquin, not seen.—Fic. 22 Ixora spinosa (Jacq.) Lam., Encycl. Méth. Bot. 3: 344. Guettarda armata Bartl. in DC., Prodr. 4: 457. 1830. TYPE EU Benth. in Oerst., Vidensk. Meddel. Dansk Niet Foren. Kjobenhavn 1852: 41. hold Hines (Benth.) Kuntze, Rev. Gen. i xd 1891. C. spinosa (Jacq.) Kuntze, Rev. Gen. Pl. 277. Guettarda costaricensis K. Schum. nomen d TYPE: not s B Brandegee, Univ. Calif. Publ. Bot. 6: 70. 1914. TYPE: México, Pringle 6940 (UC, holot Chomelia toil Standley, Contr. U.S. Natl. Herb. 18: 136. 1916. Type: Panamá, Pittier 4673 (US, holot C. purpusii (Bran degee) zs Bull. Torrey Bot. Club 52: 138. 1925. C. longicaudata Rusby, Bull. Torrey Bot. Club 52: 139. 1925. rype: Colombia, Smith 392 (US). Anisomeris nie ra inei Rusby, Bull. Torrey Bot. Club 52: 142. 1925 Trees to 20 ft tall or shrubs, the branchlets terete, smooth, the twiglets often rough with crowded nodes and withered stipular remnants, branched, the branch- es often uniformly as short as 3 cm, usually strict, the spines axillary, divergent, lignose, to 2 cm long, to 0.25 cm wide at base. Leaves often crowded at the end of twigs, oblong, ovate oblong, rarely ovate rotund, 3.7-14.0 cm long, 2.5-5.6 cm wide, acute, rarely obtuse, acuminate or cuspidate, acute at base, obtuse, often sharply contracted, occasionally truncate, often slightly inequilateral, the costa prominulous above, prominent beneath, densely or moderately villose be- neath, the lateral veins 8-10, prominulous above and beneath, arcuate, the in- tervenal areas not lineolate; petioles to 2 cm long, to 0.5 mm wide, densely pubes- cent; stipules free, narrowly triangular, 4-6 mm long, tapering into an awn, scari- ous, pilose, often persistent. /nflorescences axillary, 1 per axil, 2-5 cm long, the pe- duncle capillaceous, ca. 0.4 cm long, strict, arcuate or occasionally deflexed, the cymules terminal, few flowered, the flowers erect, sessile, disposed like a can- delabra. Flowers with the hypanthium rotund, 1.0-1.5 mm long, the calycine tube ca. 2 times the length of the hypanthium, the teeth 4, unequal, subulate, to 0.8 mm long, sericeous outside; corolla white, the tube cylindrical, to 18 mm long at maturity, to 0.65 mm wide at the middle, stiffly petaloid, the lobes 4, ovate lanceolate, to 6 mm long, conspicuously hooded at the narrow apex, sericeous outside, glabrous within, the margin scarious; stamens 4, the anthers linear ob- long, ca. 3.5 mm long, obtuse, sagittate at the base, barely exserted, the filaments slender, very short, attached just below the mouth; ovarian disc about 0.6 mm long, the stigmas 2, linear oblong, ca. 1 mm long, the style ca. 17 mm long, slightly exserted, glabrous. Fruits sessile, oblong or ovate oblong, 0.5-0.9 cm long, obtuse or truncate at the apex, the persistent calyx ca. 1 mm long, drying black, pubescent, delicately ribbed. This species extends from Mexico through Central America south to Colom- bia, Venezuela and northern Brazil. Lamarck's (1789) discussion of the species says that, "These flowers exhale during the night a very sweet odor.`` "Pata de Palome. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 105 FIGURE 22. Chomelia spinosa Jacq. Habit (<%). [After Allen 794. ] ZONE: 1 mi NW of Paraiso, near Gaillard Cut, Croat 12656 (MO). Summit Naval Radio Station, Croat 12852 (MO). Old Gamboa road, Between Summit Hills Golf Course E Summit Naval Radio, Croat 16668 (MO). Under Thatcher Bridge, D'Arcy & D'Arcy 6082 (MO). Fort Kobbe, Duke 3957 (MO). Albrook, U.S. Army Tropic Test Center, Dwyer 6731 (MO). Co colí, B 7212 (MO). Madden Dam, Boy Scout Road, Dwyer & Elias 7530 (GH, MO, US). Beach, Fort Kobbe, Gentry 6428 (MO). Mandingo Road, Hayden 60 (MO). Ancón Hill, Holdridge 6536 (MO). K-2 Road, Con tractors Hill, Lewis et al. 2882 (COL, MO, UC, VEN). Fort Kobbe, Luteyn 1088 (F). Gaillard Highway, Río Masambi Grande, Nee 7069 (MO). Madden Road, 1-2 km N of Gaillard Highway Junction, 80-110 m, Nee 76/4 (MO). 1 mi NE of Fort Clayton, Stimson 5192 (MO). Curundu, Survival School, Tyson 1059 (MO). Chiva- odis Trail, Miraflores Lake, Tyson 1362 (MO). Farfan Beach, Tyson 1810 (MO). Rodman Ammo Dump, Tyson 6658 (MO). Miraflores Lake, White 128 (F, MO). Miraflores, White 131 (F, MO). pese Hill, Woodson et al. 1329 (F, MO). CocLE: El Valle, Harvey 5188 (F). DARIEN: Chepigana, Duke & Bristan 265 (MO). HERRERA: Pesé, ca. 50 m, Allen 794 (F, MO). Las Minas and Pesé, ca. 600 ft, Duke 12337 (MO). 4 mi S of Los Pied Tyson 2652 (MO). Los SANTOS: Pocrí, Dwyer 1195 (MO). PANAMA: Madden Lake, Aguilera 16 (MO). Pacora, aeiiae is 190 (MO). Tocumen, Dressler 3898 (MO). E of Bejuco, Duke 4556 (MO). prin El Lla Río Mamoni, Duke 5530 (MO). Tocumen, Dwyer (MO); Dwyer & Hayden 16 (MO). 8 mi S of Goofy Lake, Dwyer 7125 (MO). 4 mi W of Chepo, Gentry 5555 (MO). Taboga Island, Gentry 5719 (M 9. Archeological Site, ps Lake, Gentry & Tyson 5038 (MO). Chepo, Pittier 4673 (US). VERAGUAS Roadside savanna, 2-4 mi E of Santiago, 30 m, Duke 12372 (MO). Santiago, 4 mi from Transisthmian Highway toward Atalaya, Dwyer & Kirkbride 7411 (GH, MO, UC, US). 5 mi E of Santiago, Tyson et al. 4274 (M 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 20. CINCHONA Cinchona L., Sp. Pl. 172. 1753; Gen. PI., ed. 5. 79. 1754. TYPE: C. officinalis L. Quinquina Boehmer in Ludwig, Def. Gen. Pl. 30. 1760. New name for Cinchona L. Kinkina Adans., Fam. 2: 147. 1763. Based on Cinchona L. Cascarilla R. ex Steud., B ed. 2: I: 302. 1840; not Adanson 1763. Nomen nudum. Trees or shrubs. Leaves petiolate; stipules interpetiolar, free, mostly decid- uous, glandular at the base. /nflorescences terminal, paniculate, the branches mostly opposite. Flowers 5-merous, the hypanthium pubescent, the calycine cup turbinate, the lobes 5, mostly reduced; corolla variable in color, hippocrateriform, pubescent outside, glabrous within, the lobes valvate; stamens partly exserted or included, the anthers slender, dorsifixed; ovary 2-celled, the ovules numerous, imbricate, attached peltately to a spongy placenta, the style slender, the stigmas 2, obtuse; ovarian disc pulviniform. Fruits capsular, mostly ovoid oblong, sep- ticidally dehiscent, opening from the base; seeds numerous, winged. Cinchona 1s found in Costa Rica, Panama and in the Andes of South America. There are about 40 species in the genus. 1. Cinchona pubescens Vahl, Skr. Naturhist. Selsk. 1: 119. 1790. TYPE: not seen.—FIG C. puede Mutis ex Humb., Ges. Naturfe Feeund. Berlin Mag. Neuesten Entdeck. Gesammten May 1: Naturk. 117. 1807. TYPE: not seen. C. pubescens a cordata DC., Prodr. 4: 353. 1830. TYPE: not seen. C. tucuyensis Karst., Fl. Columb. 1: 17. pl. 9. 1858. TYPE: not seen. Trees 5—8 m tall, the branchlets angular, glabrous, drying gray or reddish gray. Leaves oblong rotund to ovate rotund, to 30 cm long, to 20 cm wide, widely deltoid at the apex, obtuse to subrotund at the base, the costa slender, promi- nulous above and beneath, the lateral veins slender, 12-13, arcuate, the basal lateral veins often leaving the costa at right angles, the smaller veins pinnatiform, stiffly chartaceous, scarcely discolorous, drying red, glabrate above, appressed pilose on the costa and the lateral veins beneath; petioles to 8 cm long; stipules not seen. /nflorescences paniculate, to 35 cm long, the branches opposite, angular ascending, the lowermost to 15 cm long. Flowers not seen. Fruits pedicellate, the pedicels short, the capsules narrowly oblong or narrowly ovate oblong, to 3.5 cm long, to 0.7 cm wide, minutely golden puberulent, delicately longitudinally ribbed, the persistent calyx 1-2 mm long, wider than the apex of the capsule. Cinchona pubescens occurs in Costa Rica, Panama and in the Andes of South America. The bark is the source of quinine. *‘Quina Steyermark (1974) gives an elaborate dése dai of the flowers (from Span- ish): "Calyx and hypanthium obconic or obconic oblong, 2.5-3.0 mm long, 2.0- 2.5 mm wide, densely pale yellow sericeous. Calyx 1—2(—3) mm long, ca. 3 mm wide, moderately strigulose on the back, the teeth widely and slightly deltoid, acute to acuminate, much shorter than in C. officinalis, 0.2-1.0 mm long, strig- ulose on back. Corolla 15-16 mm long, the tube 10-13 mm long, minutely to- mentose on the back, glabrous within; lobes lanceolate or oblong, obtuse or rounded at the apex, 3-5 mm long, 1.5-2.0 mm wide, tomentulose on the back, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 107 FiGuRE 23. Cinchona pubescens Vahl.—A. Habit (x!4). [After Croat 15769.]—B. Dehisced fruit (x 1).—C. Seed (x5). [After Fosberg 20016, Colombia.] 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 densely villose along the margins within, otherwise glabrous. Anthers 2.5-3.0 mm long, the filaments ca. 2 mm long, attached approximately at the upper third; style 6-13 mm long, glabrous; disc scarcely puberulent to hispidulous. ` CHIRIQUi: Disturbed cloud forest, Monte Rey above Boquete, Croat 15769 (MO). Santa rolg region, 27 km NW of El Hato de Volcán, coffee finca Rattibor Hartmen called ''Ojo de Agua," 5000- 5300 ft, Mori & Bolten 7410 (MO). 21. COCCOCYPSELUM Coccocypselum P. Br., Civ. Nat. Hist. Jamaica. 144. 1756. '"Coccocipsilum" No- men and orthography cons. TYPE: C. repens Sw. Sicelium P. Br., Civ. Nat. Hist. e 144. 1756. Nomen Rejic. contra Coccocypselum P. Br. ased on ue i ard P Tontanea Aubl., Hist duci $ 108. 1775. TYPE: T. guianensis Aubl. Bellardia Schreb., p ne : . 1791, not Allioni 1785. Provisional name for Tontanea Aubl. Condalia - Mod Prodr. - Ten Nomen Rejic., not Condalia Cav. (1799). Rhamnaceae. Nomen $C. Mn .&P Lipostoma D. Don a Edinburgh Philos. J. 1: 168. 1829. LECTOTYPE: L. capitata (Graham) D. Don (Aeginetia ana a Graham). Herbs, often woody, prostrate or occasionally erect. Leaves petiolate; stipules free, persistent. Inflorescences terminal or axillary, the peduncle occasionally absent, the flowers disposed in a head; bracts and bracteoles small. Flowers 4-merous; calyx with the lobes narrow and elongate, persistent; corolla variable in color, infundibuliform, the mouth glabrous, the lobes valvate; stamens included or exserted, the anthers dorsifixed; ovary 2-celled, the ovules numerous in each locule, horizontal. Fruits baccate, ovoid or globose, shiny blue; seeds numerous, minute, somewhat angular. Coccocypselum ranges from Mexico through Central America, the West In- dies, and South America. There are about 20 species in the genus. Literature: d hag J. S. 1967. Coccocypselum in Botany of the Guayana Highland. . N.Y. Bot. Garden 17: 333-334. a Leaves with lateral veins 8-12, ovate lanceolate to oblong, to 7.5 cm long, often dryin olive green; pubescence not reddish . C. lanceolatum aa. Leaves with lateral veins 6-8, mostly ovate, a or ovate rotund, to 6.5 cm long, no d olive green, often with a reddish pubescen orescence with peduncles 1.0—3.8 cm one: leaves (2-)3-6.5 cm long, often with a reddish pubescence c. ue not cordate at the cule C. hirsutum ec. s cordate at the bas C. EUH bb. l. sessile; leaves 0. 84. 2 cm long, lacking a reddish E bene E 2. Li or eum 1. Coccocypselum cordifolium Nees & Mart., Nova Acta Acad. Cas Leop.-Carol. German Nat. Cur. 12: 14. 1824 S Ue pleuropoda Donnell Smith, Bot. Gaz. 52: 50. 1911. TYPE: Guatemala, ae i , not seen. Geocardia pleuropoda (Donnell Smith) Standley, Contr. U.S. Natl. Herb. 17. 445. one pleuropoda (Donnell Smith) Standley, N. Am. FI. 32: 148. 1921. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 109 Coccocypselum rothschuhii Loesner, Engler Bot. Jahrb. Syst. 60. 370. 1926. TYPE: not se Coccocypselum pleuropodum (Donnell Smith) Standley, Publ. Field Colombian Mus., Bot. Ser. 4: 81. 1929. Herbs, stems prostrate, the hairs dense, weak, spreading, the nodes well spaced. Leaves reniform, ovate or ovate rotund, 2.0-4.5 cm long, 1.5-3.2 cm wide, widely deltoid to obtuse at the apex, cordate at the base, the lateral veins 4—6, arcuate, usually drying purple beneath. /nflorescences with peduncle to 1.6 cm long. Coccocypselum cordifolium ranges from Mexico to Panama; it also occurs in eastern and southern Brazil. I have seen only one collection from Panama. Stand- ley & Williams’ (1975) description reads, "Inflorescence mostly 2—4-flowered; bracts linear or foliaceous; calyx and hypanthium long-hirsute; calyx lobes tinear or oblong linear, acute, 2-4 mm long; corolla 12 mm long, purplish white or sometimes blue; fruit densely hirsute, bright blue or sometimes greenish white." 2. Coccocypselum herbaceum P. Br., Civ. Nat. Hist. Jam. 144. 1756. TYPE: Ja- maica, not seen. C. repens Swartz, Prodr. Veg. Ind. Occ. 31. 1788. not C. repens H.B.K., 1819; not Condalia repens P. (1798). Tontanea herbacea (P. Br.) Standley, N. Am. Fl. 32: 147. 1921. Prostrate herbs, the stems slender, glabrescent or pilose. Leaves ovate or ovate subrotund, 0.8—4.2 cm long, 0.5-2.5 cm wide, widely deltoid at the apex, rounded or truncate at the base, often slightly contracted at the junction with the petiole, often inequilateral, the costa prominulous above and beneath, the lateral veins 6-8, arcuate, membranous, glabrescent to pilose above, the hairs short or long, the latter often scraggly, golden, bulbous at the base, often much longer above than beneath, pilose beneath but the hairs often restricted to costa and veins and then appressed and elongate; petioles 0.5-2.5 cm long, ca. 0.1 cm wide; stipules small, 2.5-3.0 mm long, the awn often 5 times longer than the body. Inflorescences sessile or subsessile, the heads few flowered. Flowers with the calycine lobes subulate, to 4 mm long, pilose; corolla light blue, the tube to 9 mm long, short pilose. Fruits ovoid at maturity, subrotund to ovate rotund when immature, to 1.5 cm long, pilose, the hairs gradually deciduous, china blue. Coccocypselum herbaceum is well distributed throughout tropical America and the West Indies. Steyermark (1967) discusses the synonomy of C. herbaceum at length. BOCAS DEL TORO: Santa Catalina, Blackwell et al. 2730 (MO, Bey eios Lagoon, Wedel 1134 (MO). CANAL ZONE: N of El Cope, 9.4 km above El Cope, 750-900 m, Croat 44632 (MO). Pipeline Road, D'Arcy 9273 (MO). Chagres, Fendler 179 (MO). Telephone cable trail between splice S16 and S49, Río a Steyermark & Allen Ae (MO). CHIRIQUi: Cerro Colorado, 20-28 mi d» San Félix, 1200-1500 m, Croat 33374 (MO). cocLE: La Mesa above El Valle, d 11904 (MO). L Mesa, Folsom & RR UMOR El Valle, feuis et al. 2512 (COL, DUKE, K, MO, UC, VEN). COLON: Miguel de la Borda, Croat 10028 (MO "n Río Guanche near Coastal rut D'Arcy 10192 (MO). Santa Rita Ridge, Dwyer & Gentry 9360 (MO). HERRERA: 12.5 mi S of Ocu, 1200 ft, Lewis et al. 1647 (MO). PANAMÁ: Alto de Pacora, Busey 804 (MO). Cerro Jefe, Croat 13059 (MO). Campo Tres beyond Cerro Jefe, Croat 27088 (MO). Pipeline Road near Gamboa, Clewell & Tyson 3218 (MO). Panamá Viejo, Duke 5734 (MO). Cerro Azul, Duke 9378 (MO). Cerro Campana, Gentry 1853 (M Cerro Jefe, summit, ca. 3000 ft, Hayden 1004 (MO). Cerro Azul, Mori & Kallunki 2167 (MO). Cerro 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Jefe, ca. 1000 m, Mori & Kallunki 3618 (MO). Cerro Jefe, Tyson et al. 3215 (MO), 4430 (SCZ). SAN BLAS: Road between Mandinga & Cangandi, Duke 14745 (MO). VERAGUAS: Rio Primero Braso, 2.5 km beyond Agriculture School near Santa Fé, 700-750 m, Croat 25458 (MO). To 7 km NW of Santa Fé, D'Arcy 10279 (MO). Mouth of Río Concepcion, Lewis et al. 2803 (MO, SCZ, UC). 3. Coccocypselum hirsutum Bartling ex DC., Prodr. 4: 396. 1830. TYPE: probably Trinidad, type collection not known. Tontanea hirsuta (Bartl.) Standley, N. Am. FI. 32: 147. 1921. Coccocypselum hirsutum Bartling ex DC. var. hirsutum. 3a. Coccocypselum hirsutum DC. var. hirsutum. Prostrate herbs, the nodes well spaced. Leaves ovate, ovate rhomboid, rarely oblong or reniform ovate, (2—)3.0-6.5 cm long, 1.5-5.0 cm wide, widely deltoid to obtuse at the apex, widely triangular to truncate at the base, usually slightly inequilateral, densely pubescent or with a few elongate hairs above, these golden and bulbous at the base, hirsute beneath, the hairs often elongate and stiff as well as short and soft, the hairs sometimes restricted to the margins of the blade; petioles to 2 cm long; stipules subulate, 3-6 mm long (Standley & Williams 1975). Inflorescences pedunculate, the peduncles 1—4 cm long, slender, glabrescent, often bearded. Flowers with the calycine lobes 4, linear lanceolate, 3-6 mm long, densely hirsute; corolla to 11 mm long. Fruits 2-4 per cluster, ovate, to 2 cm long, densely hairy or with a few hairs, often delicately lenticellate. Coccocypselum hirsutum ranges from Mexico to Panama. It also occurs in Trinidad, Colombia, and Bolivia. CANAL ZONE: Cerro Pelado, 1 km N of Gamboa, 200-220 m, Nee 11024 (MO). ee San Félix, Cerro Otoe, 3000 ft, Bort 69 (MO). Road from Boquete to David, oe 85 (MO). COLON: Rio Piedras, road to Portobelo, Blum et al. 2518 (MO, SCZ). Maria Chiquita, Dwyer & Kirkbride 7796 (MO). Santa Rita Ridge, Nee & Mori 3671 (MO). NE of Santa Rita ieee 650 m, Wilbur & Weaver 10837 (MO). PANAMA: Goofy Lake, 200 ft, Correa & Dressler 583, 584 (both MO). Cerro fe, D'Arcy 9751 (MO). Cerro Azul, D'Arcy & D'Arcy 6227 (MO). Hills NE Hacienda La Joya, p et al. 16896 (MO). Cerro Campana, summit, Duke 6009 (MO). Cerro Azul, Ebinger 421 (MO). o Jefe, 2900 ft, Elias & Hayden 1796 (MO). Cerro Azul, Finca Melo, Kant 47 (MO). Cerro Jefe, Kirkbride & Crebbs 28 (MO). Cerro Campana, Kirkbride 245 (MO); Lewis et al. 1927 (GH, MO, US), 1948 (MO), 3075 (COL, MO, UC, VEN); Porter et al. 4175 (MO). Cerro Jefe, 2900 ft, Porter et al. 5063 (COL, MO, UC). Cerro Campana, Witherspoon & Witherspoon $904 (MO) 3b. Coccocypselum hirsutum var. glabrum (DC.) L. O. Williams, Phytologia 25: 62. 1973.—Fic. 24. C. glabrum Bartl. ex DC., Prodr. 4: 397. 1830. TYPE: not s Tontanea glabra (DC.) Standley, J. Wash. Acad. Sci. 15: 104. 1925. Prostrate herbs. Leaves ovate to oblong, 3-6 cm long, to 5 cm wide, widely triangular to truncate at the base, glabrate above and below. /nflorescences with the peduncles to 4 cm long, glabrous. Fruits ovate, to 2 cm long. Coccocypselum hirsutum var. glabrum occurs in Mexico, Guatemala (Peten), Belize, Nicaragua, and Panama. CHIRIQUÍ: Boquete, ca. 4000 ft, Davidson 759 (MO). 4.1 mi from Boquete, road to David, Kirk- bride 85 (MO). COLON: Between Río Piedras and Puerto Pilón, Lewis et al. 3227 (MO). Santa Rita 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 111 E 24. um" hirsutum var. glabrum (DC.) L. O. Williams. Habit (<2). [After kula p Crebbs 28.] Ridge, 2 mi E of Transisthmian Highway, Lewis et al. 5242 (MO, SCZ, UC). ee Rita Ridge Road, ca. 1.9 km from Boyd-Roosevelt Highway, Mori & Kallunki 1808 (MO). AMA: Goofy Lake, Correa & l. 584 (MO). Cerro Campana, savanna near s Duke 6009 (MO). Goofy Lake, 2000 ft, Dwyer et al. 4137 (COL, DUKE, F, MEXU, MO, NY, MICH, PMA, VEN). Cerro Campana, 3⁄4 way to summit, Dwyer 4719 (MO). Cerro Azul, Antene apa 65 (MO). 4. Coccocypselum lanceolatum (Ruiz & Pavón), Pers. Syn. Pl. 1: 132. 1805. Condalia lanceolata Ruiz & Pavón, FI. Peru. Chil. a 1: 54. 1798. Type: Peru, ‘in memoribus uchero and Chinchao, Ruiz & Pavon, not see Pon ena canescens Willd. ex ne Sa io; 4: 139. 1929. rype: "Herb. Willd. n. 2844 (specimen Humboldtianum),` “yapaq canescens (Cham. & Siret] SB, N. Amer. FI. 32: 146. 1921. Herbs, prostrate or to 25 cm tall, the stems golden green, pilose, the nodes well spaced. Leaves ovate lanceolate to oblong, often somewhat falcate, 2.0—7.5 cm long, 1-3 cm wide, deltoid, obtuse to occasionally rounded at the apex, basally 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cuneate or obtuse, occasionally truncate, acute or obtuse, usually slightly in- equilateral, the costa plane or grooved above, plane to prominulous beneath, the lateral veins 8-12, arcuate, chartaceous, densely pilose beneath and above, vel- vety to the touch, often drying olive green, the veins often bearded beneath; petioles 0.3-1.3(—3.5) cm long; stipules mostly persistent, ovate triangular, 5-6 mm long, the awn longer than the body, densely hairy. Inflorescences subsessile or on peduncles 2—3(—5) cm long, 0.1—0.3 cm wide, densely pilose. Flowers sessile, numerous; hypanthium subrotund, ca. 2 mm long, densely hairy, the calycine lobes 4, erect, ovate or oblong, 3(—6) mm long; corolla white, rarely purple, the tube 4-5 mm long, petaloid, pilose outside, glabrous within, the lobes 4, oblong or triangular, ca. 2 mm long; stamens 4, the anthers sessile, attached near the middle of the tube, ovate, ca. 1 mm long; style ca. 2 mm long, the stigmas slender, about the same length as the style. Fruits bright blue, ovoid, to 7 mm in diam., pilose. Coccocypselum lanceolatum is well distributed throughout Middle America and extends into Brazil and Bolivia. The leaves of the Panamanian collections are smaller than those collected to the west of Panama. RIQUI: Boquete, Er ft, vi gig. 635, 756 (both F, MO). Bogues; Llanos Francia, Dwyer (F, G ), 2 (M O (MO). DARIEN: Gold mine, Cana, 500-600 m, Croat 37646 (MO). HERRERA: Between Las Minas and Pese, Burch et al. 1338 (MO). VERAGUAS: Trail between Canazas and foot of Cordillera Central, Rio Canazas, Allen 183 (MO). Near San José on Santiago-Santa Fé Road, 400 m, Nee 8179 (MO) 22. COFFEA Coffea L., Sp. Pl. 172. 1753; Gen. Pl. ed. 5: 80. 1754. TYPE: C. arabica L. Shrubs, rarely pubescent. Leaves occasionally ternate, usually petiolate, the stipules free, acuminate, persistent. Inflorescences with flowers conglomerate in the axils, the bracteoles connate into a calyxlike cup. Flowers sessile or pedi- cellate, the hypanthium variously shaped; calycine cup reduced, truncate, dentate or lobulate, often bearing small glands within: corolla salverform or funnelform, the lobes 4-8, contorted; stamens 4-8, sessile or on short filaments inserted in the throat, the anthers linear, basally dorsifixed, exserted or included, often twist- ed; ovarian disc swollen, the style slender or thick, the stigmas 2, narrow, the ovary 2-celled, the ovules solitary, subpeltately affixed to the middle of the sep- tum. Fruits baccate, dry or fleshy, the 2 carpels often coriaceous, convex but plane and sulcate on the inner face; seeds 2, convex, the testa membranous with a deep groove on the ventral side. This is a genus of about 40 species represented in the tropics of the Old World, especially in Africa. The 2 species, Coffea arabica and C. liberica, supply most of the coffee of commerce. Literature: Standley, P. C. & L. O. Williams. 1975. Coffea Linnaeus Coffee (part of Ru- biaceae of Guatemala). Fieldiana 24, part II: 44. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 113 a. Flowers 5-merous _______________---___------------------------------------------------------ 1. C. arabica aa. Flowers 6-8-merous ______________---..--------------------------------------------------- 2. C. liberica 1. Coffea arabica L., Sp. Pl. 172. 1753. TYPE: not seen.—FIG. 25. Shrubs or trees, to 20 m tall, the branchlets straight or flexuous, glabrous, scarcely or moderately swollen at the nodes, the latter well spaced. Leaves el- liptic oblong, 8-18 cm long, 2.5-7.5 cm wide, distinctly acuminate at the apex, the acumen to 1.5 cm long, often falcate or somewhat hooklike, usually ultimately obtuse, attenuate or widely cuneate at the base, the costa prominulous above, subprominent beneath, the lateral veins 9-11, uniting to form a submarginal un- dulate vein to 0.8 cm from the margin, thin coriaceous to stiffly papyraceous, usually drying green or grey green; petioles to 1.2 cm long; stipules linear lan- ceolate, to 1.2 cm long, acute. Flowers not seen. Fruits plump, oblong rotund, to 1.5 cm long, to 1 cm wide, rounded at the apex and the base, drying black brown or dull olive, glabrous, the calycine scar scarcely elevated, ringlike, to 2 mm in diam. Coffea arabica is native to Africa and is cultivated throughout the tropical areas of the world. Standley & Williams (1975) discuss the cultivation of Coffea arabica in Guatemala. The 5-parted flowers are described in their diagnosis: * . . 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 equalling or exceeding the tube; anthers exserted .. . CHIRIQUí: Cafetales above El Cantinero, D'Arcy 9846 (MO). Finca ‘‘Princesa Janca, Boquete, D'Arcy & D'Arcy 6453 (MO). Fred Collins Finca, Boquete, Ebinger 725 (MO). Quebrada Melliza, e, 700 6 mi S of Puerto Armuelles, 0-150 m, Liesner 457 (MO). CocLE: Forest behind Club Campestre, m, Duke 13252 (MO). El Valle, Ebinger 948 (MO). PANAMA: Rio Maje along river from waterfalls near Bayano Lake ca. 2 mi upstream, Croat 34553 (MO). Highway 79, Quebrada Sardinilla, 5 km NNE of cement factory, 55 m, Nee 6581 (MO). Cermeno, Ochoa 32 (MO). 2. Coffea liberica Bull ex Hiern., Trans. Linn. Soc. Ser. 2(1): 171, tab. 24. 1876. TYPE: Sierra Leon, Daniell (BM, not seen; photo, MO). Shrubs or trees to 20 ft tall, the twiglets subangular, glabrous, the nodes well spaced. Leaves oblong or obovate elliptic, 7.5-8.5 cm wide, short acuminate at the apex, rounded at the base, the lateral veins ca. 12, coriaceous, glabrous; petioles to 2 cm long, stiff. Flowers not seen. Fruits oblong subrotund, to 2.0— 2.5 cm long at maturity (fide Standley & Steyermark), lightly grooved. Coffea liberica is native to Liberia and to areas of West Africa. It is cultivated throughout tropical areas of the world. Standley & Williams (1975) describe the 6—8-parted flowers as `°. . . several in a cluster, subsessile, 2.5 cm long; bractlets shorter than the subtruncate calyx; corolla lobes about as long as the tube; anthers not exserted.’’ ‘‘Liberian Coffee." COLON: Achiote, Tyson et al. 4508 (MO). 23. CONDAMINEA Condaminea DC., Prodr. 4: 402, 1830. LECTOTYPE: C. corymbosa (R. & P.) DC. Shrubs or small trees. Leaves large, coriaceous; stipules large, bipartite. In- florescences terminal, spreading, cymose or corymbose; bracteoles absent. Flow- 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN \ FIGURE 25. Coffea arabica L. Habit (x%). [After Ochoa 32.] [Vol. 67 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 115 ers pedicellate; hypanthium turbinate campanulate, the calycine cup with 3—5 lobes or teeth; corolla funnelform or campanulate, the tube cylindrical, villous in the throat, the lobes 5, valvate, glabrous; stamens 5, the anthers exserted, narrow, sagittate, the filaments thick, inserted below the corolla mouth; ovarian disc de- pressed, the ovary 2-celled, the style stout, filiform, exserted, the stigma oblong, obtuse, the ovules numerous. Fruits capsular, turbinate, truncate, thick walled, dehiscing loculicidally into 2 valves; seeds minute, attached horizontally, the testa reticulate. Condaminea occurs sporadically throughout Middle America and occurs in tropical South America. There are about 3 species in the genus. a. Leaves oblong, to 50 cm long, the lateral veins ca. 20; petioles absent or to 1 cm lon EN . C. cory "ymbosa aa. Leaves lanceolate, to 15 cm long; the lateral veins ca. 10; petioles 0.7-1. 2c cm m long reer 2. C. petiolata 1. Condaminea corymbosa (Ruiz & Pavón) DC., Prodr. 4: 402. 1830.—Fic. 26. Macrocnemum corymbosum Ruiz & Pavón, Fl. Peruv. Chil. Prodr. 2: 48. 1799. TYPE: not seen. Shrubs or small trees to 7 m tall, the branches terete, stout, smooth, glabrous or puberulent. Leaves oblong, to 50 cm long, to 21 cm wide, deltoid to vaguely obtuse at the apex, basally truncate to subcordate, the costa plane or prominulous above, prominent beneath, to 3 mm wide, the lateral veins usually ca. 20, usually 1.5-2.5 cm apart, coriaceous, glabrous to golden puberulent beneath, the inter- venal areas often delicately wrinkled; petioles either absent or to 1 cm long, stout, rimose, glabrous; stipules interpetiolar, connate, to 4 or 5 cm long, to 0.6 cm wide at the base forming a sheath, each part with 2 subulate awns longer than the sheath, acute, coriaceous, puberulent, rubescent. /nflorescences terminal, spreading paniculate, to 40 cm long, the peduncle 15-20 cm long, with 3 primary branches at the tip, 10—15(—21) cm long, angular ascending or arcuate, each ter- minating in either 3 radially disposed secondary branches or in a few branches. Flowers with the hypanthium oblong, to 1 cm long, to 0.6 cm wide, tapering cuneately below, the calycine cup short, dilated, the margin with 4 or 5 broad, strongly compressed rotund or widely triangular lobes, coriaceous, glabrous; co- rolla white, the tube cylindrical, ca. 2x longer than wide, to 2 cm long, carnose, glabrous inside and out, the lobes 5, triangular or oblong, to 1 cm long, glabrous; stamens 5, the anthers narrowly oblong, ca. 7 mm long, ca. 1.2 mm wide, retuse at the apex, the filaments short, plane, attached at the upper 25 of the tube; style plane, attenuate at the base, to 1.5 cm long; stigmas 2, to 5 mm long, ca. 0.7 mm wide, truncate, the ovary thin walled, the locules 2-3, the ovules numerous on 2-3 intrusive, C-shaped placentas as seen in cross section. Fruits oblong or obovate oblong, 1-1.5 cm long, ca. 1 cm wide at the base, ligneous, glabrous, drying black, topped by a conspicuous and shallow ringlike scar, splitting longi- tudinally into 2 or 3 thin valves; seeds numerous. Condaminea corymbosa ranges from Costa Rica to Bolivia. CHIRIQUi: Boquete, Davidson 792 (F, MO, US). 12.4 mi N of David, Lewis et al. 692 (GH, K, MO, NY. PMA, UC, US). 17 km NE of San Félix, road to Cerro Colorado, 1000 m, Nee 10718 (MO). San Félix, Pittier 5252 (F, US). DARIÉN: Piñas, Duke 10621 (MO). 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 E 26. Condaminea corymbosa (Ruiz & Pavon) DC.—A. Leaves and inflorescence n. TH "Fruit (x3.5).—C. Seed (x21). [After Duke 10621.] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 117 2. Condaminea petiolata Dwyer.?" TYPE: Panama, Croat 34706 (MO, holotype). Shrubs to 2 m, the twigs slender, glabrous, obtusely angled, the nodes well spaced. Leaves lanceolate, to 15 cm long, to 4.5 cm wide, acute at the apex and the base, the costa prominulous above and beneath, the lateral veins ca. 10, strongly arcuate, plane above, prominent beneath, the intervenal areas open re- ticulate, chartaceous, discolorous, the hairs appressed, white, elongate, more dense beneath especially on the costa and the veins; petioles 0.7-1.2 cm long, ca. 0.2 cm wide, pubescent; stipules persistent at the apex of the twigs, connate basally, turgid, the sheath 1-2 mm long, each part with an erect triangular pro- cess, attenuate toward the apex, to 0.6 cm long, acute, vaguely thickly carinate, villose outside, densely villose at the base within and often with a few conspic- uous black glands on the margin within, the glands oblong, ca. 0.5 mm long. Inflorescences axillary, paniculate; peduncle slender, to 0.3 cm wide, the cor- ymbs disposed toward the apex of 2-3 pairs of short, arcuate, ascending, opposite branches, these 3-5 mm long; bracts not seen; bracteoles subulate. Flowers not seen. Fruits sessile, oblong, coriaceous, to 1 cm long, less than 2 times longer than wide, truncate at the apex, pubescent outside, delicately ribbed, the persis- tent calycine cup scarcely measurable or to 1 mm long, the 2 carpels loculicidally dehiscent, lignose, yawning apically, the endocarp smooth, thick, drying yellow, finally free from mesocarp; seeds numerous, plano compressed, fusiform, ca. 2 mm long, shiny. Condaminea petiolata is known only from Panama. It is distinguished by its leaves having few lateral veins and by its axillary inflorescence. The fruits are smaller than those of C. corymbosa. PANAMÁ: Río Maje, near new Bayano Lake, 100 m, Croat 34706 (MO). 24. COSMIBUENA Cosmibuena Ruiz & Pavón, Fl. Per. Chil. Prodr. 3: 2. 1802. Nomen conservan- dum. TYPE: C. obtusifolia Ruiz & Pavón not Cosmibuena Ruiz & Pavón (1794). Nomen rejiciendum (Chrysobalanaceae). Buena Pohl, Pl. Bras. Ic. 1: 8. 1827 Shrubs, glabrous, usually epiphytic. Leaves fleshy, petiolate; stipules large, deciduous. Inflorescences terminal, cymose or the flowers solitary; bracts and bracteoles present. Flowers large; hypanthium oblong or turbinate, the calycine cup tubular or campanulate, the teeth 5-6, usually unequal; corolla salverform or funnelform, the tube elongate, 5-6-lobed, the lobes contorted; stamens 5-6, the anthers linear, basifixed, included, the filaments short: ovarian disc cup —M Vp Dwyer, spec. nov. Frutices ad 2 m alti, Wei gracilibus glabris obtuse angulatis nodis ntibus. Folia lanceolata, ad 15 cm longa, cm lata, acuta, venis lateralibus ca. 10, pibe Au. chartacea discoloria; petiolis ad 1. 2 cm longis; stipulis connatis turgidis vagina 1-2 mm longa parte utraque processu triangulari erecto ad 0.6 cm longo praedita extus que. cm lato, ^e nini versus apicem jugorum 2-3 ramulorum brevium arcuate ascendentium opposito- rum -5 ongorum. Fructus sessiles oblongi, ad 1 cm longi, coriacei apice truncati extus pubescentes beya costati. 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 shaped, the style elongate, clavate at the apex, the stigmas 2. Fruits capsular, narrowly oblong or cylindrical, septicidally bivalvate from the apex, the epicarp separating from the endocarp; seeds numerous, oblong, the body small, the testa membranous, produced at each end into a narrow wing. a. Calyx cup ca. 1.1 cm long, the teeth only minute points; corolla tube to 7.5 cm long ______ Seip ays l. C. ovalis aa. Calyx cup 3.3-5 mm long, the teeth 1.5-3 mm long; corolla tube 6-10 cm long ------------ un s ey L a eaee eee 2. C. skinneri 1. Cosmibuena ovalis Standley, Contr. U.S. Natl. Herb. 18: 137. 1916. TYPE: Panama, Pittier 5074 (US, holotype: NY, isotype). Trees to 30 ft tall, 4 inches d.b.h., terete, drying dark, the nodes well spaced. Leaves rhomboid, oblong rotund or elliptic oblong, 12-17 cm long, 5.5-8.5 cm wide, deltoid, rounded or obtuse at the apex, acuminate, the acumen vague, short, basally cuneate or attenuate cuneate, the costa plane or prominulous above, prominulous beneath, to 0.2 cm wide, the lateral veins 5-6, at first strict and strongly ascending, arcuate toward the margin, the submarginal vein indis- tinct, stiffly chartaceous, discolorous, glabrous above except the puberulent cos- ta, glabrous beneath except puberulent along the lowermost part of the costa and along the lateral veins; petioles to 1 cm long, to 0.15 cm wide in the middle, puberulent, expanded at point of attachment; stipules not seen. Flowers 2 at the end of the twiglet, erect, the calyx collar shaped, to 1.1 cm long, to 0.6 cm wide, truncate, the teeth minute points to 0.5 mm long, acute, rigid, glabrous; corolla tube narrowly cylindrical, to 7.5 cm long, to 0.3 cm wide in the middle, glabrous, scarcely expanded at the apex, the lobes oblong, 2-2.5 cm long, 1-1.3 cm wide, obtuse, glabrous; style exserted, probably to 8.5 cm long, densely golden villose. Fruits not seen. Cosmibuena ovalis is known only from Costa Rica and Panama. The above description is sketchy, as only one collection was available except for the type. CHIRIQUI: Rio Macho de Monte, | mi E of Cuesta de Piedra, 2800 ft, Tyson 904 (MO). COCLE: 100—350 m, Ola, Pittier 5074 (NY, US). 2. Cosmibuena skinneri (Oerst.) Hemsl., Biol. Centr. Amer., Bot. 2: 12. 1881. n ded Oerst. Vidensk. Meddel. Dansk Naturhist. Foren. Kjobenhavn 26: 1852. TYPE: Nic- , Oersted 11134 (not seen, holotype, photo, Geen paludicola Standley, Contr. U.S. Natl. Herb. 18: 137. 1916. TYPE: Panamá, Pittier 4107 , holotype; NY, isotype). Trees to 25 ft tall, the branchlets terete, smooth, glabrous, often drying gray or tan, frequently angular, the nodes usually well spaced. Leaves obovate oblong, occasionally oblong subrotund, 6-16 cm long, 2.7-7.5 cm wide, obtuse or rounded at the apex, cuneate at the base, the costa prominulous above and beneath, often evanescent distally, 1-1.5 mm wide at the base, the lateral veins 5-7, prominu- lous, evanescent, strongly ascending, coriaceous, glabrous; petioles to 2 cm long, to 0.25 cm wide, lignose, glabrous; stipules deciduous, oblong rotund, ca. 1.5 cm long, 1 cm wide, obtuse, coriaceous, glabrous, often with a thickened keel below the middle. Inflorescences terminal, the peduncle short, to 1 cm stout, glabrous, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 119 ZEE Z — [JA — Z m 7 SS. NM Z f IN SS SK N FIGURE 27. Cosmibuena skinneri (Oerst.) Hemsl.—A. Flowering branch (x12). [After Croat 6326.|—B. Fruiting branch (x). [After Croat 5057,|—C. Flower (x 1).—D. Seed (x2). [After Gentry 5761.] drying much darker than the stem tip, the flowers 3-6, ascending. Flowers 1-2 cm long (pedicels and hypanthium), ca. 0.4 cm wide, glabrous, marcescent; ca- lycine cup 3.3-5 mm long, glabrous, coriaceous, the teeth 5, widely triangular, disposed as points 1.5-3 mm long, acute or obtuse; corolla white, the tube nar- 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 rowly cylindrical, 6-10 cm long, ca. 0.3 mm wide, drying brown, subcoriaceous, glabrous, the lobes 5, oblong, 1.9-3.5 cm long, 0.9-1.2 cm wide, obtuse, often glabrous, often drying yellow abaxially, brown adaxially; stamens 5, the anthers narrowly oblong, 1.0—1.2 cm long, ca. 0.15 cm wide, the filaments short, inserted near the middle of the corolla tube, anthers basifixed; stigmas 2, digitiform or oblong, to 1 cm long, 1.8 cm wide, obtuse, the style slender, as long as the corolla tube, ca. 1 mm wide, villose for ca. 6 mm below the stigmatic lobes, the latter oblong, ca. 5 mm long, obtuse. Fruits pedicellate for ca. 2 cm, narrowly oblong, often falcate, to 7.5 cm long, to 0.8 cm wide, terete in cross section, rounded at the apex with an annular scar, the calyx rarely persistent, to 0.3 cm long, the ovarian disc persistent as a cylindrical projection, to 2 mm long, ca. 1.3 mm wide, attenuate at the base, ligneous, glabrous, conspicuously lenticellate. Cosmibuena skinneri ranges from Mexico to Colombia. Croat has noted on labels that the flowers are pollinated by hawkmoths. CANAL ZONE: Gatun, Higgins 400 (MO). BARRO COLORADO ISLAND: Aviles 78 (MO); Bangham 473 (F): Croat 5057, 6326, 6554, 7833, 8519, 9029, 9853, 12285, 12444 (all MO); Foster 1912 (F); Knight 8 (F); Shattuck 863 (MO); Woodworth & Vestal 771 (MO). cocrÉ: El Valle, Allen 2798 (MO). 2 km from El Greco Hotel, El Valle, Correa & Dressler 997 (MO). Las Margaritas & El Valle, Woodson et al. 1738 (MO). COLON: Santa Rita Ridge, Correa & Dressler 695 (MO); D'Arcy & D'Arcy 6162 (MO). María Chiquita, E of Río Piedras, Dwyer & Kirkbride 7773 (GH, F, K, MO, NY, PMA, UC). Santa Rita Ridge, Dwyer & Gentry 9394 (MO). Vienta Frío, Pittier 4107 (US). DARIÉN: Río Pinas, Duke 10546 (MO). PANAMÁ: Cerro Campana, Correa & Dressler 1032 (MO); Duke 6008 (MO); Dwyer & Kirkbride 7829 (MO); Ebinger 917 (MO); Gentry 5761 (MO); McDaniel 6921 (MO). SAN BLAS: Puerto Obaldía, Gentry 1496 (MO). WITHOUT EXACT LOCALITY: Duke 6193 (MO). COLOMBIA cHoco: Between Camp Curiche and Quebrada Changame, 3.7 mi S of Camp Curiche, Duke & Idrobo 11574 (NY). 25. COUSSAREA Coussarea Aubl., Hist Pl. Guiane 1: 98. 1775. TYPE: C. violacea Aubl. Trees or shrubs. Leaves often coriaceous; stipules usually entire and connate, short, obtuse, often semiorbicular. /nflorescences terminal or rarely axillary, pa- niculate or umbellate, many flowered, the peduncle usually present, the branch- lets ultimately cymose; bracts slender or much reduced. Flowers usually 4-mer- ous, rarely sessile, the bud rounded or truncate; hypanthium occasionally with 4 lobes or teeth; corolla funnelform or salverform, the throat naked, the lobes 4, valvate; anthers subsessile; stigmas 2, the lobes filamentous, the ovary 2-celled, the ovules 2, longer than broad, erect, connate by means of a thin, evanescent septum. Fruits drupaceous, oval, I-seeded by abortion of the second seed, the seed vertical, ellipsoid, usually smooth, often one side convex. Coussarea occurs throughout tropical America. It includes about 100 species. One species of Viburnum, V. costaricanum (Oerst.) Hemsley, well collected in Chiriqui, may be easily mistaken in the fruiting stage for a Coussarea, but the fruit may be distinguished from Coussarea species by its persistent ovarian disc which is triangular but conspicuously capitate at the apex, measuring ca. 0.4 mm long. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 121 Literature: Dwyer, J. D. and M. Hayden. 1966. Notes on Coussarea (Rubiaceae) especially the Panamanian Species. Ann. Missouri Bot. Gard. 53: 368-374. a. Leaves coriaceous, inflorescence 3-5 cm wide. Leaves with lateral veins 16-25; petioles less than 1 cm long; stipules 0.25—0.5 lon g. c. Leaves petiolate, the lateral veins ca. 16; calycine cup ca. 3 mm long ---------- 3. C. durifolia cc. Leaves sessile, the lateral veins ca. 25; calycine cup ca. 9 mm long 5. C. garciae bb. mp ie lateral veins ca. 9; petioles 1-3 cm Sha stipule 1-1.5 cm long. d. ves 9.0-16.5 cm wide: fruits to 2 cm lon y C. latifolia . C. jefensis dd. qe 1-5 cm ide: fruits to 1 cm lon aa. Leaves chartaceous or if coriaceous (C. pantudato) the inflorescence to 6 cm md Leaves obviously petiolat Petioles to 0.5 cm long : Petioles 0.6-5.0 cm long. g Calycine cup ca. 16 mm long; corolla tube ca. 33 mm long 2. C. curvigemmia = 4. C. enneantha gg. "apk cup 0.2-7.0 mm long; corolla tube less than 20 mm long. Inflorescence patulous . open i. Leaves with lateral veins ; 11-15. J. Calycine cup ca. 0.5 mm long -------------- 14. C. veraguensis j Calycine cup ca. 2.5 mm long 8. C. loftonii il. Losses with lateral veins 5—10(—12). k. Leaves with lateral veins 5-6 |. 10. C. nebulosa kk. Leaves with lateral veins 7-10(—12). l. Calycine cup ca. 16mm long ------------ 4. C. enneantha ll. Calycine cup 0.5-7.0 mm ong. m. Leaves with lateral veins ca. 12; petioles 2-5 cm long . : — ^nsis mm. ipid with lateral veins 8— 10; petioles 0- cm lon Calyeiie cup ca. 5 mm long; flo nk. 13. C. roseo-cremea pin nn. Calycine cup 1-2 mm long; flowers white. o. Petioles (longest) 2.0-2.5 cm long; fruits to 2.7 cm long ___ 11. C. neei oo. Petioles o pd ca. 1.5 cm long; fruits 1.2-1.6 cm long |... A ee 12. C. ae ulata p. Leaves coriaceous; calyx p berulent, edentate. pp. Leaves thinly chartaceous; ca- lyx glabrous, often with 2 E jin N Inflorescence obviously contracted "e hh. ee. Leaves sessile- . C. garciae 1. Coussarea cerroazulensis Dwyer & Hayden, Ann. Missouri Bot. Gard. 53: 374. 1966. TYPE: Panamá, Dwyer 1368 (MO, holotype). Trees 5-10 m tall, the branchlets terete, densely villosulose, the nodes well spaced, often turgid with evident stipule bases. Leaves elliptic, 15-33 cm long, 5-14 cm wide, acute or conspicuously acuminate, the acumen slender, often narrowly caudate, to 2 cm long, basally cuneate, the costa prominulous above, 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 prominent beneath, the lateral veins ca. 12, widely arcuate, prominent beneath, densely golden pubescent, often velutinous to the touch, coriaceous; petioles 2— 5 cm long; stipules not seen. Inflorescences terminal, sessile, to 12 cm long in fruit, to 6 cm wide, with 3 pairs of opposite, well-spaced branches, the lowermost to 2 cm long, each terminated by a 3-flowered cymule. Flowers with the hypan- thium squarish in lateral view, coriaceous, densely golden villose, the calycine cup cylindrical, about as wide at the base as at the apex, to 7 mm long, ca. 3.5 mm wide, coriaceous, eglandular within, the hairs outside ferrugineous, long and spreading, the hairs within appressed, ferrugineous, the margin undulate, with 4 irregular triangular teeth, these often curved, to 1 mm long: corolla with the tube narrowly cylindrical, ca. 1.7 cm long, to 3.5 mm wide, somewhat falcate, ex- panded above the middle, coriaceous, densely appressed hairy outside, glabrous within, the lobes 4, narrowly oblong, ca. 15 mm long; stamens 4, the anthers narrowly oblong, ca. 7 mm long, sagittate at the base, the filaments plane, ca. 1 mm long, glabrous, attached near the middle of the tube; style slender, ca. 5 mm long, the stigmas subulate, somewhat plane, ca. 5 mm long. Fruits globose, 1—3 cm long, densely puberulent, the persistent calyx narrowly cylindrical, to 1 cm long, ca. 0.4 cm wide, the wall of the fruit 2-3 mm thick. Coussarea cerroazulensis is known only from Panama. Two striking features of the species are the elongate petioles, to 5 cm long, and the elongate filaments of the stamens which are almost as long as the elongate anthers. PANAMÁ: Cerro Jefe, Croat 13052 (MO), 15189 (MO, NY). Without other locality, Duke 8167 (MO). Goofy Lake and Cerro Jefe, Blum et al. 2270 (MO). Cerro Jefe, Duke & Dwyer 15064 (MO); Duke 8167 (MO). Cerro Azul, Dwyer 1368 (MO). Between Cerro Jefe and La Eneida, Dwyer 8199 (MO, NY). Cerro Azul, Gallegos et al. 5174 (MO). Cerro Jefe, 2900 ft, Gentry 2147 (MO). 2-8 mi S of Goofy Lake, Lewis et al. 2161 (DUKE, F, GH, K, MO, NY, PMA, UC, US). Gorgas Lab, ca. 25 km NE of Cerro Azul, 550 m, Mori & Kallunki 3267 (MO). NE of town of Cerro Azul, 20 km by road from Panamerican Highway, Mori & Kallunki 3654 (MO). 5-10 km NE of Altos de Pacora, ca. 750 m, Mori & Kallunki 4969 (MO). 2. Coussarea curvigemmia Dwyer, Phytologia 38: 215. 1978. TYPE: Panamá, Croat 14863 (MO, holotype).—Fic. 28. Shrubs or small trees, to 5 m tall, the branchlets terete, glabrous, ashen gray when dry, the nodes well spaced. Leaves oblong or lanceolate, 6.5—25.0 cm long, 2.0—13.5 cm wide, scarcely or strongly inequilateral, often falcate, acuminate, the acumen straight or curved, to 1.5 cm long, obtuse, cuneate at the base or some- times obtuse or lightly auriculate, the costa subplane to prominulous above, prominulous beneath, the lateral veins 6-9, widely arcuate, the venules 1—3 be- tween adjacent lateral veins, these branching and reticulate, the petioles lacking or to 0.5 cm long, glabrous; stipules triangular, to 4 mm long, rounded toward the apex, glabrous. /nflorescences solitary, terminal, glabrous to puberulent, thrysoid paniculate, to 6 cm long, to 4.5 cm wide; peduncle slender, 1-2 cm long, the primary branches alternate or opposite, to 2 cm long, ascending, the rachis often with 2 pairs of terminal branches, alternate or opposite, flabellate in ap- pearance, the flowers congested, sessile or subsessile. Flowers with the hypan- thium oblong or oblong rotund, often shorter than the calyx cup, the latter cylin- drical or collar shaped, 1.5-2.5 mm long, truncate, stiffly petaloid, glabrous or 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 123 FIGURE 28. Coussarea curvigemmia Dwyer.—A. Habit (x!2).—B. Fruiting branch (x%).—C. Flower, opened to show stamens and style (x3.5).—D. Ovary cross section (x5). [After Croat 7308. | 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 puberulent on the outside; the teeth 5 or absent, triangular, to 0.5 mm long, often with minute, glandular red spots on the margin; corolla white, the tube narrow cylindrical, often falcate, 6-16 mm long, ca. 1.5 mm wide, petaloid, glabrous or puberulent outside, glabrous within, the lobes 4, narrowly oblong, 7-8 mm long, glabrous within, not cucullate; stamens 4, the anthers linear, 4.5-5.7 mm long, acute with a minute apiculum, the filaments short, ca. | mm long, attached above the middle of the tube; ovarian disc prominent, to 0.65 mm high, ca. 1 mm wide, the style short or long, included or scarcely exserted, linear, 6-16 mm long, the stigmas 2, linear oblong, 1.5-3.0 mm long, ca. 0.1 mm wide. Fruits ellipsoid, often laterally compressed, 1.2-1.5 cm long, to 0.7 cm wide, glabrous, at first fleshy and spongy, the pulp glistening white, the surface often lenticellate when dry; seed solitary with a hard testa filling the cavity, the calyx persistent, to 1.5 mm long, to 1.2 mm wide. Coussarea curvigemmia is known only from Panama and Costa Rica. Most of the material cited below has previously been erroneously assigned to Cous- sarea impetiolaris Donnell Smith which unlike this species has subquadran- gular branches, a corolla tube up to 2 mm in width, much more elongate anthers which are up to 8 mm long, and a fruit which in the dried condition is about 19 mm.long. The type of Coussarea impetiolaris is Pittier 134008 (US, syntype) from Costa Rica. Most fruits on herbarium material have only the fibrous endocarp visible. One collection, Dwyer 2012 (MO) shows the fleshy outside of the fruit still present. These fruits measure to 1.5 cm in length; those with only the endocarp present measure to 1.2 cm in length. BOCAS DEL TORO: Almirante, eu 2770 (MO). cANAL ZONE: C-19 Road, Blum 1901 (MO). Nuevo Emperador, Blum 2393 (MO). K-6 Road, Dwyer 2853, 5021 d bi n ep Dam, Dwyer 9161 (MO); Hayden 74 (MO); Kant 24 WES Boy Scout Camp Road near Madden Dam, Mori 4040 (MO). Madden Dam, Porter et al. 4051 (MO, SCZ, UC, VEN). Viena Island. [ei 5475 (MO). BARRO COLORADO ISLAND: Croat 4892, 5465, 5873, 6201, 6282, 7308, 8600, 10295A, 10973, 14863 (all MO); Ebinger 171, 565, 610 (all MO): Foster 1779 (MO); Hayden 34 (MO); Killip 39987 (MO); WEE A loa 66-11-2-1259 (MO); Shattuck 135, 621 (both MO); Starry 50 (MO); Wetmore & Abbe O). COCLE: Pa Pilón, Dwyer & Lallathin 8630 (MO). COLON: Santa Rita Ridge, Dwyer et al. e (MO). ÉN: Santa Fé. Duke & Bristan 309 (MO). Yaviza, Duke 4875 (MO). El Real to Pinogana, Duke. 5012A (MO). Santa Fé. Duke pod 14091 (both og Tyson et al. 4665, rn (both MO). Río Chucunaque, Stern et al. 845 (MO). AMÁ: Juan Mina, Bartlett & Lasser 16572 (MO); Tyson 5459 (MO). Cerro Azul, Dwyer 7093 (MO. Pt Jefe, Ge Bn 6781 (MO). Villa Rosario, near Capira, Saldana 41 (MO). 3. Coussarea durifolia Dwyer.?!' TYPE: Panama, Liesner 1180 (MO, holotype). Small trees to 4 m tall, the twiglets thick, ultimately somewhat angular, smooth, glabrous, the nodes well spaced, somewhat turgid, the leaf scars trian- gular or subrotund, ca. 0.5 cm long. Leaves obovate or obovate oblong, to 22 cm long, to 12 cm wide, obtuse or rounded at the apex, acuminate, the acumen short, to 5 mm long, cuneate or obtuse at the base, the costa prominulous or subplane above, prominent beneath, convex, the lateral veins ca. 17, arcuate, the sub- ! Coussarea durifolia Dwyer, spec. nov. Arbores parvae vel frutices. Folia obovata vel obovata- Hn. apicem verus obtusa vel rotundata, brevi-acuminata venis lateralibus ca. 17, coriacea in sicco gadina glabra; petiolis nullis vel ad 1 cm longis; stipulis ovato- triangularibus. Inflorescentiae terminales, ramis paucis floribus patulis. Flores sessiles. Fructus non visi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 125 marginal vein undulate, 2-3 mm from the margin, the intervenal areas reticulate, coriaceous, emerald green beneath when dry, glabrous; petioles absent or to 1 cm long, to 0.35 cm wide, glabrous when dry; stipules free, ovate triangular, to 1 cm long, basally ca. 0.65 cm wide, obtuse, coriaceous, with minute dots outside, to 0.2 mm in diam. /nflorescences terminal, solitary, to 7 cm long, to 5 cm wide, glabrous, drying black; peduncle to 2 cm long, to 0.35 cm wide, the branches few, short, to 2 cm long, the flowers spreading. Flowers sessile, the perianth parts carnose; hypanthium oblong, 2.5-4.0 mm long, minutely puberulent or gla- brescent, the calycine cup turgid, collarlike, ca. 3 mm long, ca. 3.5 mm wide, truncate, the teeth 4 or scarcely visible, ovate oblong, 0.8-1.5 mm long, reflexed, obtuse; corolla cream colored, the tube narrowly hypocrateriform, ca. 1.4 cm long, minutely puberulent outside, glabrous within, the lobes 4, narrowly oblong, 7-9 mm long; stamens 4, the anthers narrowly oblong, ca. 6 mm long, ca. 0.7 mm wide, the filaments short, ca. 1.5 mm long, plane, attached near the middle of the tube; style slender, ca. 4 mm long, the stigmas 2, linear oblong, ca. 2 mm long. Fruits not seen. Coussarea durifolia is known only from Panama. It is readily recognized by its coriaceous leaves which dry an emerald green beneath. The lateral veins are unusually numerous for a Coussarea. The black punctations of the stipules are probably diagnostic. ANAMÁ: El Llano-Cartí highway, 20 km N of El Llano, Dressler 4352 (MO). El Llano-Cartí road, 12 mi above Panamerican Highway, Liesner 1180 (MO) ~ 4. Coussarea enneantha Standley, J. Wash. Acad. Sci. 18: 282. 1928. TYPE: Pan- amá, R. S. Williams 841 (US, holotype; isotype, NY).—FIG. 29. Shrubs 4(—7) m tall, the branchlets subterete, slender, the internodes well spaced. Leaves elliptic oblong, 9-16(—24) cm long, 3.2-8.0(-13.0) cm wide, acu- minate, the acumen to 1.5 cm long, usually falcate, obtuse, acute or cuneate at the base, somewhat inequilateral, the costa prominent, the lateral veins (7-)10— 12, joining to form a usually distinct submarginal vein, the venules reticulate, membranous to stiffly chartaceous, concolorous or discolorous, villosulose or villose on veins and costa above and beneath; petioles slender or stout, 1.2-3.5 (-4.2) cm long, 0.1—0.2 cm wide, pubescent; stipules connate, the sheath ca. 3 mm long, pubescent. /nflorescences terminal, usually sessile or the peduncle to 1 cm long, trichotomous, the branches spreading, curved, to 8 cm long, slender, ca. 1 mm wide, densely golden pubescent, each branch with 3 flowers at the apex. Flowers sessile; hypanthium including calycine cup narrow cylindrical, to 16 mm long, ca. 3.3 mm wide, the cup more than twice the length of the hypan- thium, villose outside, glabrous within, stiffly carnose; corolla white or cream colored, the tube to 33 mm long, ca. 4 mm wide, densely pubescent outside, sometimes glabrescent, the lobes 4, narrowly oblong, ca. 11 mm long, 2-3 mm wide basally, glabrous adaxially; stamens 4, the anthers narrowly oblong, ca. 7 mm long, ca. 0.7 mm wide, obtuse, the filaments ca. 6.5 mm long, attached submedially in the tube; stigmas 2, digitiform, ca. 3 mm long, the style ca. 13 mm long; ovary oblong in cross section, thin walled. Fruits sessile or pedicellate for 3 mm, oblong, to 3 cm long, to 1.5 cm wide, tapering at the apex and base, the calyx persistent, 1.0-1.7 cm long, densely pubescent. 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 29. Coussarea enneantha Standley.—A. Habit (x!2).—B. Flower (x1).—C. Flower, dissected showing stamens (x 1).—D. Fruit (x 1). [After Duke 15707.] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 127 Coussarea enneantha is known only from Panama. One collection, Dwyer & Correa 8892, has extraordinarily large leaves, up to 24 cm long and 13 cm wide; the indumentum is somewhat reduced on the lower surface of the blade and the fruits are only up to 2 cm long. “AS DEL TORO: Fish Creek Mountains, Wedel 2364 (MO). CocLE: Cerro Caracoral, 900 m Duke & Dwyer 15139 (MO). “aa yan Caracoral, Dwyer & Correa 8892 (MO). Cerro Pilón, Porter a al. 4402 (MO). DARIEN: Cerr . Bristan 536 (MO). Río Pirre, Bristan 1462 (MO). Cerr Campamiento, S of Cerro Pirre, Date 1570 7 (MO). Cerro Pirre, 3500-4500 ft, Duke & Elias 13781 (MO, UC, VEN). Cana-Cuasi trail on Cerro Campamiento, E of Tres Bocas, Kirkbride & Duke 1239 (MO, NY). Caná, Williams 841 (NY, US). PANAMÁ: 6.5 km on road N of Goofy Lake, Nee 9301 (MO). 5. Coussarea garciae Standley, Publ. Field Mus. Nat. Hist. Bot. Ser. 22: 180. 1940. TYPE: Colombia, Killip & Garcia 33524 (F, holotype). Shrubs or small trees 2.0-3.5 m tall, the branchlets stout, angular above, lenticellate, the nodes somewhat turgid, glabrous, the stipular scars thin, ringlike. Leaves sessile, obovate, to 50 cm long (or longer?), to 20 cm wide, obtuse or rounded at the apex, occasionally short acuminate, tapering narrowly and cu- neately toward the base and finally vaguely auriculate, the costa slender and prominulous above, prominent beneath, to 0.3 cm wide, lignose, the lateral veins ca. 25, to 3 cm apart, prominulous above, prominent beneath, the veinlets per- pendicular, thin coriaceous, occasionally papyraceous, concolorous, drying green, glabrous above and beneath; stipules free, deltoid, to 1.5 cm long, sub- coriaceous, marcescent, glabrous. /nflorescences terminal, to 5 cm long, the cen- tral axis suppressed or to 3 cm long, the branches few, short, the flowers densely aggregated. Flowers with the hypanthium ca. 2 mm long, the calycine tube cy- lindrical, ca. 7 mm long, rounded basally, somewhat turgid, glabrous, petaloid, with a few, well spaced, reddish glands within, the calycine teeth 4, narrowly oblong or narrowly ovate oblong, 10-12 mm long, ca. 2 mm wide at the base, acute; corolla blue, the tube narrowly cylindrical, somewhat curved like an S, ca. 15 mm long, wider above the middle, ca. 2 mm wide at the middle, glabrous, stiffly thin carnose, the lobes 4, narrowly oblong, ca. 6 mm long; stamens 4, the anthers oblong, 4.5-6.0 mm long, subsagittate basally, the filaments slender, 3 mm long, attached near the middle of the tube; style slender: 4 mm long, the stigmas digitiform, 2 mm long. Fruits on pedicels 0.3 cm long, elliptic oblong, 2.0-2.8 cm long, to 1.3 cm wide, obtuse at the apex and the base: light brown, smooth, glabrous, the raphides conspicuous under magnification, ribbed, the per- sistent calyx to 2 cm long, the lobes often withering, narrowly ovate lanceolate, to 12 mm long, glabrous. Coussarea garciae is known only from Darien and the Dept. of Choco, Co- lombia. Duke (11350) notes that the fruits are eaten in Colombia. ''Colorin.' NAMA: DARIEN: Between the ayaq Río Membrillo and Camp 7 on the road to San Blas, Duke /0878 (MO). Hydro Camp No. 14, Rio Salaqui, 6 days upstream from Rio Sucio, Duke 11350 (MO). Rio Tuquesa, lower Tuquesa Mining Co. camp, Mori 6982 (MO). Camp Summit, adjacent to Darién- San Blas border, Sea Level Canal, Route 17, 1000-2000 ft, Oliver et al. 3679 (MO). COLOMBIA: Between Rio Nercua-Quebrada Barrial and Quebrada Ambrosio near Camp Curiche, 10-294 m, Duke 11435 (MO). Bahia Solano, dense forest along Quebrada Jellita, 50-100 m, Killip & Garcia 33524 (F, US). 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 6. Coussarea jefensis Dwyer.22 TYPE: Panamá, Mori & Joly 7927 (MO, holotype). Trees to 6 m tall, the branchlets terete, smooth, glabrous, drying dark, the nodes well spaced, turgid, the leaf scars elevated, round, drying whitish with a central circular vascular bundle. Leaves oblong rotund to oblong, 3.0-8.5 cm long, 1-5 cm wide, rounded to obtuse at the apex, occasionally vaguely acumi- nate, obtuse or rounded at the base, the costa prominulous above and beneath, drying red, to 0.15 cm wide proximally, the lateral veins ca. 9, arcuate, prominent beneath, forming an undulate submarginal vein with usually 1 irregular interme- diate vein between adjacent lateral veins, the intervenal areas patulous reticulate, the margin slightly revolute, coriaceous, concolorous, drying dark, glabrous, glandular shiny; petioles 1.0-1.3 cm long, stiff, glabrous, to 1.5 mm wide; stipules connate, forming an expanded, coriaceous, glabrous and truncate cup, to 2.5 mm long. /nflorescences (in fruit) terminal, paniculate, glabrous, drying dark, to 8.3 cm long, to 4.5 cm wide; peduncle 3.5-4.0 cm long, plano compressed, the low- ermost branches opposite, paired, ascending, to 2 cm long, the upper branches to 6, alternate; bracts and bracteoles minute, ca. 1 mm long. Fruits subsessile, oblong, rarely vaguely curved, terete in cross section, to | cm long, drying black, glabrous, vaguely 2-lobed, not costate; seed solitary, oblong, to 5 mm long, to 4 mm wide, obtusely 3-angled in cross section, with a thin horizontal ridge on the adaxial surface. Coussarea jefensis is immediately distinguished from all other Coussarea in Panama by its thick-coriaceous leaves which are oblong rotund to rotund. The elevated large leaf scars are diagnostic. PANAMÁ: Cerro Jefe, ca. 2900 ft, Mori & Jolv 7927 (MO). N . Coussarea latifolia Standley, J. Wash. Acad. Sci. 18: 281. 1928. TYPE: Costa Rica, Tonduz 9574 (US, holotype). Trees to 15 m tall, glabrous throughout, the twiglets stout, 4-angled, lenticel- late. Leaves broadly elliptic obovate to broadly ovate elliptic, 16.5-25.0 cm long, 9.0-16.5 cm wide, acute or obtuse at the base, the costa and lateral veins stout, salient beneath, the lateral nerves 9, arcuate, papyraceous, concolorous, pale green when dry; petioles stout, 1.7-2.5 cm long; stipules persistent above, semi- orbicular, ca. 5 mm long, wider than long, intrapetiolar, shortly connate, broadly rounded at the apex. Inflorescences terminal, glabrous, to 6 cm long, to 5 cm wide, the peduncle to 2 cm long, to 0.35 cm wide. Flowers sessile or Se hypanthium short, glabrous, the calycine cup cylindrical, ca. 9 mm long, ca. 6 mm wide at the apex, the latter truncate, coriaceous, glabrous, dentate, olen: dular; corolla (seen only in part) narrow below, presumably falcate, coriaceous, ? Coussarea jefensis Dwyer, spec. nov. Arbores ad 6 m altae ramulis dp one nodis cicatricibus foliorum conspicue rotundis ornatis. Folia oblongo-rotunda ad oblonga, 3.0-8.5 cm longa 1-5 cm lata, apice rotundata vel obtusa venis rape ralibus ca. M coriacea, bes phun: 1.0 longis; stipulis connatis vagina puede , ad 2. m longa, truncata coriacea glabra. Infores entiae terminales paniculatae, ad 8.3 cm longae ad s cm d pce 3.5-4.0 cm longo, ramulis inferioribus ad 2 cm longis, superioribus ca. 6 alternatis; bracteis et bracteolis ae Fructus oblongi, ad 1 cm longi, aoh, vix 2-lobat 1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 129 glabrous outside, pubescent within; stamens 4, included in the tube, the anthers lunate or S-shaped, 5-6 mm long, tapering at the apex, cordate at the base, the filaments 2 to almost as long as the anthers, presumably attached at the middle of the tube; ovarian disc prominent. Fruits oblong, to 2 cm long, often somewhat falcate, obtuse or rounded at the apex, green to yellow, lenticellate or smooth, the calycine cup short, smooth, glabrous. Coussarea latifolia is known only from Costa Rica, Panama, and Colombia. BOCAS DEL TORO: Above Almirante, Gentry 2765 (MO). Fish Creek Hills, Chiriqui Lagoon, Wedel 2218, 2463 (both MO). COLON: 10 mi S of Portobelo, 2—4 mi from the coast, 10-200 m, Liesner 1071 (MO). PANAMÁ: Cerro Jefe, ca. 700 m, Correa & Dressler 1158 (MO). Between Cerro Azul and Cerro Jefe, Mori & Kallunki 2173 (MO). 8. Coussarea loftonii (Dwyer & Hayden) Dwyer, comb. nov. Faramea loftonii Dwyer & Hayden, Phytologia 15: 56. 1967. rvPE: Panamá, Tyson et al. 3323 (MO, holotype). Shrubs or trees, to 10 m tall, the branchlets terete, drying yellow grey, gla- brous, the nodes usually well spaced. Leaves salicoid, often falcate, 5-17 cm long, 1.8-5.0 cm wide, the larger blades regularly 3 times longer than wide, tapering acutely or deltoidly at the apex, acuminate, the acumen attenuate, usu- ally narrowly falcate, to 1.5 cm long, 0.1-0.3 cm wide in the middle, acute at the base, slightly inequilateral, the costa scarcely prominulous above, rimose, sub- plane beneath, the lateral veins 9-12, prominulous, at first strict and then arcuate, or strict to the margin, the intervenal areas markedly reticulate, papyraceous, concolorous, glabrous; petioles to 1 cm long when present, narrowly alate; sti- pules connate, often persistent, deltoid or compressed rotund, wider than long, 2.5-3.0 mm long, 3-4 mm wide, coriaceous, glabrous. Inflorescences terminal, to 5 cm long, to 5 cm wide, cymose paniculate, the peduncle to 2 cm long, ca. 1.5 mm wide, glabrous, the branches 3-4, the lowermost often paired, opposite or subopposite, divergent or ascending, to 1.5 cm long, the terminal cymules with 3-6 flowers: lowermost bracts rarely persisting, narrowly elliptic, to 15 mm long, ca. 1.5 mm wide, glabrous. Flowers with the pedicels 1.5-3.0 mm long, glabrous; hypanthium 0.5-1.2 mm long, smooth, glabrous, the calycine cup and teeth to 2.5 mm long, the cup with a few appressed glands, coriaceous, the teeth 4, sub- ulate, to 0.7 mm long; corolla white, the tube narrowly cylindrical, to 9 mm long, ca. 2 mm wide, glabrous, stiffly petaloid, the lobes 4, lanceolate, to 7.5 mm long, tapering narrowly toward the apex; stamens 4, the anthers narrowly oblong, 4- 5 mm long, ca. 0.5 mm wide, obtuse, caudate at the base, dorsifixed, the filaments ca. 1 mm long, attached near the middle or at the upper 4 of the tube; stigmas 2. subulate, to 2.2 mm long, the style 10-13 mm long. Fruits scarcely pedicellate, oblong, to 1.7 cm long, ca. 1 cm wide, smooth, glabrous, white, drying yellow, spongy within, the persistent calycine cup small, to 1.5 mm long, ca. 1 mm wide. Coussarea loftonii is known only from Panama. It was originally described from flowering material, but a recent collection, Lewis & Dressler 6567, has fruits which are obviously those of a Coussarea and not of Faramea. Nee 9297 de- scribes the flowers as very fragrant. 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 : Santa Rita Ridge, Correa & Dressler 667 (MO); ut sa 3852 (MO). Santa Rita Ridge oad, Ge "ntry et al. 8864 (MO). El Llano-Carti road 9.6-11 km from Interamerican Highway, Mori & "Kallunki 3521 (MO). Santa Rita Ridge Road, 14 km from the Pen Roosevelt Highway, Mori & Kallunki 4904 (MO). PANAMA: La Eneida, Lopez House, Correa & Dressler 818 (MO). Cerro Jefe, Lewis & Dressler 7567 (MO). NE of town of Cerro Azul, Mori & Kallunki 3641, 3649 (both MO); NEE 9297 Cerro Jefe, Tyson et al. 3323 (MO); Wilbur & Weaver 11373 (MO). 9. Coussarea morii Dwyer.? TYPE: Panama, Mori 7058 (MO, holotype). Trees to 12 ft tall, ca. 20 cm d.b.h., the twiglets slender, smooth, glabrous, the nodes well spaced. Leaves oblong, 10.0-24.5 cm long, 3.5-12.0 cm wide, obtuse to widely cuneate at the apex, acuminate, the acumen to 1.5 cm long, falcate, acute to obtuse at the base, the costa subplane above, subprominent beneath, the lateral veins 7-10, arcuate, prominulous beneath, the submarginal undulate vein vague, the intervenal areas patulous reticulate, thin chartaceous, scarcely discolorous, drying green, glabrous except for tufts of hairs in the axils of most veins; petioles 1.5-2.0 cm long, rigid, strict, glabrous, drying dark; stip- ules not seen; glands at stipular scar position numerous, narrow oblong, ca. | mm long, drying red. /nflorescences terminal and axillary, cymose paniculate, to 19 cm long, to 6 cm wide; peduncle ca. 3 cm long, ca. 0.3 cm wide, angular and compressed, drying black, glabrous; primary branches disposed as 3-4 pairs, opposite or subopposite, well spaced, often at right angles to the rachis, the lowermost to 2.5 cm long; cymes few, clustered at the tips of reduced secondary branches; bracts and bracteoles apparently absent except for 2 mammiform swell- ings in the axils of the uppermost leaves, 1 per axil, ca. 1 mm long, glabrous, drying black. Flowers sessile or subsessile, externally glabrous; hypanthium to 2 mm long, the calycine cup and hypanthium chalicelike, the cup ca. 2 mm long, turgid, petaloid, the teeth as vague points or distinctly 5, oblong, to 1 mm long, obtuse, the cup with a few appressed glandular spots within; corolla white, the tube narrow cylindrical to 5 mm long, to 1.3 mm wide in the middle, flaring at the very base; stamens 4, the lobes 4, oblong, slightly longer than the tube, to 8 mm long, the anthers narrow oblong, ca. 4 mm long, exserted, the filaments to 2 mm long, attached below the mouth; style slender, to 6.5 mm long, glabrous, the stigmas digitiform, to 1 mm long, as wide as the style; ovules 2, connate. Fruits not known. Coussarea morii is known only from Panama. It is named in honor of Scott Alan Mori, distinguished student of tropical American plants. The new species is closely allied to C. paniculata (Vahl) Standley, but differs in having both 33 Coussarea morii Dwyer, spec. Arbores pou rU. E oblonga, 10.0—24.5 cm longa, 3.5-12.0 cm lata, apice obtusa er we cuneata, acumine ad 1.5 cm longo, venis lateralibus 7— 10, tenui-coriacea alabea. petiolis 1.5-2.0 cm longis. Iflor: scentiae me: et axillares, cymoso- paniculatae, a cm longae 6c tae, pedunculo go glabro ramis primariis in jugis 3—4 dispositis. Flores sessiles vel subsessiles hypanthi ; longo glabro cupula calycis ca. 2 mm longa, turgida “see dentibus saepe inconspicuis vel aliter ad 1 mm longis, obtusis glabris; corolla tubo cylindrico, ca. 5 mm longo, leviter curvato extus glabro intus villosuloso supra medium; antheris 4 angusto- on is, ca. 4 mm longis exsertis Lun ad 2 mm longis infra faucem affixis: stigmatibus digitiformibus, ad 1 cm longis. Fructus not v 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 131 terminal and axillary inflorescences which are often twice the length of those of C. paniculata; usually a single small mammiform swelling in the axils of the uppermost leaves subtending the peduncle; thin chartaceous rather than coria- ceous leaves; glabrous hypanthium with the calycine cup glabrous and occasion- ally with teeth to 1 mm long; corolla with oblong lobes which measure about 2 mm in diameter. Dr. Mori records that the tree is 12 meters in height. Apparently C. paniculata does not exceed 8 meters in height (fide Steyermark, 1974). D : Río Tuquesa, at middle Tuqesa Mining Company Camp called Charco Peje, tropical wet forest, 2501 m, Mori 7058 (MO). 10. Coussarea nebulosa Dwyer.?* TYPE: Panamá, Croat 15868 (MO, holotype). Shrubs 3.0—3.5 m tall, the twigs smooth, dark grey when dry, often rimose, the nodes swollen, well spaced. Leaves oblong, 6.0-10.5 cm long, 2.5-3.5 cm wide, acute toward the apex, acuminate, the acumen narrowly deltoid, to 1 cm long, the costa plane or prominulous above, prominulous beneath, the lateral veins 5-6, arcuate, extending to the margin, subchartaceous, somewhat discol- orous, glabrous, the raphides minute, white, dense; petioles slender, to 1.2 cm long, ca. 0.13 cm wide; stipules connate, the sheath slightly turgid, ca. 2 mm long, each part of the sheath with 2 well-spaced awns shorter than the sheath. Inflorescences terminal, solitary, glabrous, to 5 cm long, to 3 cm wide, the pe- duncle to 2.5 cm long, the lowermost branches opposite, to 1.5 cm long, the upper branches few; lowermost bracts subulate, to 3 mm long. Flowers sessile; hypanthium subrotund, ca. 0.6 mm long, the calycine cup glabrous, 0.5 mm long, lacking glands within, the teeth 5, triangular, 0.5 mm long; corolla white, ca. 25 mm long, the tube ca. 18 mm long, carnose, glabrous outside, villose inside, the lobes 5-6 mm long; stamens 5, the anthers linear oblong, 4 mm long, 0.3 wide, the filaments ca. 0.5 mm long, the style 19 mm long, ca. 0.3 mm wide, the stigmas 2, ca. 2 mm long. Fruits not seen. Coussarea nebulosa is known only from Panama. The new species has the fewest lateral veins (5—6) among the Panamanian Coussarea ; its connate stipules, each part with 2 short awns, are unique among the Panama Coussarea. The type was collected near Boquete at about 4,000 ft elevation. This is probably the highest elevation at which a Panamanian Coussarea has been collected. The specific name “‘nebulosa’’ was selected as it suggests a cloud forest. CHIRIQUÍ: Uncut cloud forest at Monte Rey near Boquete, Croat 15868 (MO). 34 Coussarea nebulosa Dwyer, spec. nov. Frutices 3.0-3.5 m alti. Folia oblonga, 6.0—10.5 longa, 2.5-3.5 cm lata, versus apicem acuta, acumine ad 1 cm longo venis lateralibus 5—6 i unas glabra; petiolis ad 1.2 cm longis; stipulis connatis vagina subturgida, ca. 2 mm longis, parte utraque setis duabus quam vaginam brevioribus. /nflorescentiae terminales solitarieae glabrae, 5 cm longae, d 3 cm latae, pedunculo ad 2.5 cm longo. Flores cupula calycis 0.5 mm longa, glabra eglandulosa dentibus 5, 0.5 mm longis; corolla alba tubo ca. 18 mm longo, intus villoso, lobis 5-6 mm longis; staminibus 5, antheris lineari-oblongis, 4 mm longis, filamentis ca. 0.5 mm longis. Fructus non visi. men o N ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 11. Coussarea neei Dwyer.’ TYPE: Panamá, Nee 9820 (MO, holotype). Trees or shrubs, 4-7 m tall, the twigs leafless except at tips, gray, cracking, often flexuous, the nodes turgid. Leaves lanceolate or oblong-lanceolate, often inequilateral and falcate, 5-16 cm long, 1.8-5.0 cm wide, acute, obtuse or rounded at the apex, the acumen absent or conspicuous, to 1 cm long, acute to obtuse at the base, the costa immersed above, prominulous or subprominent beneath, the lateral veins 9-10, often widely arcuate, the intervenal areas spreading reticulate, thin papyraceous, discolorous, often drying brown, arachnoid pubescent only on the costa above, pubescent on the costa beneath, occasionally on the veins; petioles to 2.5 cm long, 1.3 cm wide; stipules free, compressed rotund, to 0.4 cm long, rotund at the apex, smooth, glabrous, ciliolate marginally. Inflorescences terminal, solitary, ferrugineous villose or hirsute, 2.5-8.0 cm long .Ü cm wide, the peduncle reduced, 0.5-0.8 cm long, ca. 1 mm vide. the branches 2, opposite, the cymules terminal, the flowers few. Flowers (in bud), with the hy- panthium oblong, to 3 mm long, ferrugineous hirsute, the calycine cup ca. 2 mm long, subcarnose, villosulose within, the lobes 4-5, subulate, often scarcely in- equilateral, to 3 mm long, erect, densely golden villose. Fruits sessile, narrowly oblong, to 2.7 cm long, to 0.8 cm wide, drying black, densely golden puberulent. Coussarea neei is known only from Panama. It is named in honor of Michael Nee, who collected extensively in Panama in 1973-1974. It is closely related to Coussarea enneantha Standley, as is evident from the structure of the inflores- cence. The new species has a much shorter calycine cup with conspicuous ca- lycine teeth. ARIÉN: Serrania de du ascent of Serrania de Pirre . Cana gold mine, 600—1000 m, Croat 37737 (MO). VERAGUAS: Valley Río oe Bocas, between Escuela Agricola Alto Piedra and Calove bora, 15.6 km N f Santa Fé, 450—550 m, Croat 27649 (MO). 5 mi NW of Sant Fé. above Río Primero Brazo, ns sner 823 (MO). 8. 8 km from Escuela Agricola Alto de Piedra, y rri & Kallunki 3216 (MO). 6-7 km W of Santa Fé, new road past agricultural school, 2900 ft, Nee 9708 (MO), 9820 (MO, holotype). 12. Coussarea paniculata (Vahl) Standley, J. Wash. Acad. Sci. 18: 282. 1928. Froelichia paniculata Vahl, Ecol. Amer. 3. 1796. not Moench. 1794. TvPE: French Guiana, not seen. Billardieria paniculata Vahl, Ecol. Amer. 10. 1796. rype: French Guiana, "In sylvis prope amnem alibiensem,`` not seen. Faramea martini DC. ex Griseb., Fl. Brit. W. I. 339. 1861, not DC. 1830. rype: French Guiana, not seen; photo in Delessert Herb. Coussarea froelichia A. Rich., Mém. Soc. Hist. Nat. Paris 5: 177. 1834. TYPE: not s C. darienensis Steyermark, Ceiba 3: 20. 1952. type: Panama, Allen 4576 (F, beoe. iem NY). Small trees. Leaves widely elliptic, 8.5-14.0 cm long, 3-6 cm wide, caudate acuminate at the apex, the acumen to 1.8 cm long, cuneate, acute, often attenuate 35 Coussarea neei Dwyer, spec. nov. APT a frutices, 4-7 m altae. d rna vel s lanceolata, saepe inaequilateralia et falcata, 5-16 cm longa, 1.8-5.0 cm lat s lateralibus 9-10, tenui-papyracea supra in costa solum one. pubescentia oe in ens et saepe in venis pubescentia; petiolis ad 2.5 cm longis; stipulis compresso-rotundis, ad 0.4 cm longis, nae rotundis. t de scentiae terminales solitariae can villosae vel hirsutae, 2.5-8.0 cm longae, 2.5-9.0 cm , pedunculo 0.5-0.8 cm longo, ca. 1 mm lato, ramis 2 oppositis dispositis mule terminalibus 2 flores ferentibus. Flores cupula calycis ca. 2 mm longa, lobis 4-5, ad 3 m MS Fructus sessiles angusto-oblongi, aa 2.7 cm longi, ca. 0.8 cm lati, in sicco nigri dense aureo-puberuli 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 133 at the base, the lateral veins ca. 8, stiffly chartaceous, discolorous or concolorous, glabrous except for a few hairs in the axils of the veins below; petioles to 1.5 cm long; stipules often deciduous. Inflorescences pyramidal paniculate, 6.5-9.0 cm long, to 6 cm wide, the peduncle to 1.5 cm long, the branches few, paired, perpendicular to the rachis, the lowermost to 2.5 cm long. Flowers subsessile or on pedicels to 0.5 cm long; calycine cup glabrous, turgid, to 1 mm long, smooth, the teeth scarcely measurable; corolla white, the tube to 3 mm long, the lobes ca. 6.5 mm long; anthers linear, to 5 mm long, the filaments to 1 mm long attached near the mouth. Fruits on pedicels to 0.5 cm long, ovoid, 1.2-1.6 cm long, 1.0- 1.5 cm wide, yellow, white or green, glabrous, drying black. Coussarea paniculata is known from Panama, Trinidad and northern South America. The isotype of C. darienensis is deposited in NY, not in the Missouri Botanical Garden herbarium, contrary to Steyermark's indication in the original description. DARIÉN: Río Chico, Yaviza, ca. 100 ft, Allen 4576 (F, NY). Puerto St. Dorothea, near Colombia- Panamá border on Pacific side, Dwyer 2222 (MO). 13. Coussarea roseo-cremea Dwyer.?* TYPE: Panamá, Dressler 4334 (MO). Shrubs to 3 m tall, the branchlets ultimately obtusely angular, smooth, gla- brous, the lenticels minute, few. Leaves oblong, 6.5-13.0 cm long, 3.5-7.0 cm wide, obtuse, rotund or widely deltoid at the apex, rounded or obtuse at the base, acuminate, the acumen short, to 0.5 cm long, acute or obtuse, the costa promi- nulous above and beneath, the lateral veins 7-8, widely arcuate, forming an undulate submarginal vein, the intermediate veins 1—4, soon branching; petioles moderately slender, stiff, glabrous, 1.0-2.5 cm long, to 0.12 cm wide; stipules often short connate, persistent, hemispherical, turgid, to 0.4 cm long, each part with 2 awns. Inflorescences solitary, terminal, cymose paniculate, to 6.5 cm long, to 5.5 cm wide, the peduncle to 2.5 cm long, the lowermost branches few, 1—3, opposite or verticillate, to 3 cm long, the flowers terminal, few. Flowers sessile; hypanthium subrotund, ca. 1 mm long, shiny, glabrous, the calycine cup to 0.5 mm long, subturgid, petaloid, glabrous, eglandular within; corolla pink, the tube narrow cylindrical, ca. 15 mm long, wider above the middle, puberulent inside and out, petaloid, the lobes 5, linear subulate, ca. 10 mm long, 1 mm wide at base, the margins contiguous above the middle, flat, venose near the mouth, petaloid; stamens 5, the anthers narrowly oblong, 3.3 mm long, subacute, not exserted, the filaments short, 1.1 mm long, attached above the middle of the tube; ovarian disc 0.4 mm long, 0.9 mm wide, the style slender, 15 mm long, the stigmas 2, subulate, 1.5 mm long. Fruits not seen. 36 Coussarea roseo-cremea Dwyer, spec. nov. Frutices ad 3 m alti. ps dunes 6.5-13 cm longa, 3.5-7 cm lata apice obtusa, rotundata vel lato-deltoidea, acumine m longo, basi rotundata vel ena. venis lateralibus 7-8; petiolis 1-2.5 cm longis: woods uad bir eee persistentibus hemisphaericis turgidis ad 0.4 cm longis de utraque aristis duabus praedita. Inflo- rescentiae solitariae terminales cymoso-paniculatae, ad m longae, ad 5.5 cm latae, pedunculo ad cm longo, floribus terminalibus paucis. Flores cupula c Rie s ad 0.5 mm longa, dentibus nullis; corolla roseo-cremea, tubo ca. 15 mm longo, lobis 5 lineari-subulatis ca. 10 mm longis: antheris angusto-oblongis, 3.3 mm longis, filamentis 1.1 mm longis. Fructus non visi. 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Coussarea roseo-cremea is known only from the single collection in Panama. It is closely related to Coussarea nebulosa Dwyer which also has connate stipules with awns, few lateral veins in the leaf blades and a very reduced calycine cup. Coussarea roseo-cremea, however, is a lowland species with pink flowers, small- er anthers and corolla lobes almost equal to the corolla tube in length. The Latin name roseo-cremea refers to the flower color. PANAMA: El Llano-Cartí Road, 6-10 km N of El Llano, Dressler 4334 (MO). 14. Coussarea veraguensis Dwyer.” TYPE: Panamá, Mori & Kallunki 4800 (MO, lotype) Trees to 4 m tall, the branchlets rigid, angular, glabrous, sharply ascending, the nodes well spaced, somewhat turgid. Leaves narrowly or widely elliptic, often slightly falcate, rarely inequilateral, 14-20 cm long, 3.5-10.0 cm wide, deltoid toward the apex, acuminate, the acumen to 2 cm long, often ultimately acute, cuneate or obtuse at the base, the lateral veins 11—14, arcuate, disappearing toward the margin, the intervenal areas reticulate, the raphides minute, white, rigidly membranous, concolorous, glabrous or puberulent below on the costa and veins; petioles thick, 0.5-1.0 cm long, to 0.25 cm wide, puberulent; stipules de- ciduous, not seen. /nflorescences terminal, racemelike, to 5 cm long, to 3 cm wide, the peduncle slender, 1-2 cm long, subtended by 2 large leaflike bracts 2.0— 5.5 cm long, 0.8-1.5 cm wide, each bract with an erect linear-oblong process in the axil, to 0.4 mm long, obtuse, black, glabrous, the flowers 15-20, either solitary on the rachis or in 2-3 sessile clusters. Flowers with the pedicels short, 1-3 mm long, glabrous; hypanthium oblong, ca. 2 mm long, glabrate, the calycine cup cupuliform, somewhat turgid, to 0.6 mm long, minutely puberulent only on the margin, inside with 10 rotund glands of various size, dark red: corolla white, the tube narrowly cylindrical, ca. 7 mm long, 1.5 mm wide, subfalcate in the bud, narrow toward the base, carnose, glabrous, the lobes 4, carnose, nar- rowly oblong, to 5 mm long, stamens 4, the anthers narrowly rectangular, ca. 4 mm long, obtuse, sagittate at the base, the filaments arising near the middle of the tube; disc of the ovary compressed rotund, to 0.8 mm long, the style plane compressed, ca. 5 mm long, the stigmas 2, fingerlike, to 2 mm long. Fruit not seen. Coussarea veraguensis is known only from Panama. It is distinguished by its racemelike inflorescence in which the flowers arise either in solitary fashion or are disposed as clusters on 2 or 3 reduced branches of the rachis. The 2 large of MA hy veraguensis Dwyer, spec. nov. Arbores ad 4 m altae. Folia angusto- et lato-elliptica, 14-20 cm longa, 3.5-10 cm c versus apicem deltoidea acumine ad 2 cm longo, plerumque ultime acuto, basi oae. vel obtusa venis lateralibus 11—14, rigido-membranacea, glabra vel puberula; petiolis 0.5-1 cm longis. ie n terminales racemo similes, ad 5 cm longae, ad 3 cm latae, pedunculo gracili, 1-2 cm longo, bracteis basi pedunculi 2 magnis foliis similibus, 2-5.5 cm longis, floribus 15-20 solitariis in rhachidi vel 2-3 fasciculis seesilibis dispo sitis. Flores pedicellis 1-3 mm 5 mm longis; staminibus 4, antheris angusto-rectangularibus ca. 4 mm longis. Fructus non v 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 135 leaflike bracts have a distinct oblong process at the base; the corolla tube appears to be shortest (only 7 mm long) among the Panamanian Coussarea except for C. villosula. Coussarea veraguensis appears to be closely related to C. carolina Standley, a Costa Rican species, and to C. paniculata (Vahl) Standley, which occurs in Panama and in Costa Rica. VERAGUAS: NW of Santa Fé, | km from Escuela Agricola Alto de Piedra, on the slopes of Cerro Tute, Mori & Kallunki 4800 (MO). 15. Coussarea villosula Dwyer, Ann. Missouri Bot. Gard. 53: 105. 1966. TYPE: Panama, Duke 5338 (MO, holotype). Small trees, the branchlets terete, nodose, minutely puberulent. Leaves lan- ceolate, to 20 cm long, to 8 cm wide, acuminate at the apex, acute basally, the lateral veins ca. 15, ca. 1 cm apart, the costa prominulous above, prominent proximally beneath, the margin delicately revolute, rigidly papyraceous, glabrous above, sparsely villosulose and conspicuously reticulate beneath; petioles 1-4 cm long, somewhat thick, puberulent; stipules triangular, to 8 mm long, hirsute. Inflorescences terminal, 2-4 cm long, narrow, to 2.5 cm wide, densely sericeous villose; peduncle slender, 2-3 cm long, the cymules 3—4-flowered; bracts subulate, to 2 mm long. Flowers with the buds fusiform, to 0.5 cm long; calycine cup bell shaped, to 2.5 mm long, ca. 1.6 mm wide, carnose, densely appressed villose outside, the hairs to 0.8 mm long, the teeth 5, scarcely measurable or to 0.5 mm long; corolla white, the tube about 3.2 mm long, densely appressed villose out- side, glabrous within, carnose, the lobes 5, imbricate, ca. 1 mm long, falcate and obtuse at the apex; anthers subsessile, somewhat wedge shaped, to 1.3 mm long; style slender, to 1 mm long; stigmas small; ovules 2, brown when dry, separated by a thin septum. Fruits not seen. Coussarea villosula is known only from Panama. Its spindle shaped buds and greatly contracted hairy inflorescence readily distinguish it. cocLÉ: El Valle from village to La Mesa, ca. 2000 ft, Dwyer & Correa 7925 (MO). DARIEN: Cerro Pirre, from Rio Pirre S to El Real, 750-1030, Duke 5338 (MO). 26. COUTAREA Coutarea Aubl., Hist. Pl. Guiane 314. 1775. TYPE: C. speciosa Aubl. Shrubs or small trees. Leaves membranous, glabrate, petiolate; stipules short, acute. /nflorescences terminal or axillary, cymose, the flowers occasionally sol- itary (not Panama). Flowers with the calyx 5—8-lobed, deciduous in fruit; corolla funnel shaped or campanulate, the tube asymmetric, curved, gibbous ventricose, the lobes 5-8, short, plicate imbricate; anthers exserted, the filaments free, not attached to the corolla tube; ovules numerous, the ovary 2-celled. Fruits capsular, compressed oval, usually coriaceous and conspicuously lenticellate, dehiscing longitudinally; seeds broadly winge Coutarea ranges from Mexico to Argentina, with about 7 species in the genus. I have some doubts as to whether this genus belongs in the Rubiaceae. The unique 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 corolla shape, the staminal filaments being free of the corolla tube, and the unsusal shape of the fruit are the principal characters which render the genus suspect. 1. Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl. Bras. 6(6): 196. 1889. Portlandia hexandra Jacq., Enum. PI. Carib. 16. 1760. rYPE: apne Cartagena, not seen. Coutarea flavescens Moc. & Ses., DC., Prodr. 4: 350. 1830. TYPE: C. hexandra pubescens K. an in Mart., Fl. Bras. 6(6): 198. 1889. TYPE: : not designated. Small trees or shrubs, the twiglets terete, glabrous, relatively smooth, the lenticels often abundant and conspicuous, the branchlets rigid or subarcuate, the nodes usually well spaced. Leaves usually ovate rotund to ovate, 4-12 cm long, 2-6 cm wide, round or occasionally deltoid at the apex, acuminate, the acumen short, subfalcate, acute or more often acuminate, basally rounded, obtuse or truncate, often vaguely inequilateral, the venation immersed or prominulous above, prominulous beneath, the costa slender, the lateral veins 6-12, scarcely arcuate, well spaced, stiffly papyraceous, glabrous above, glabrous to golden pubescent beneath, the hairs curled, scattered along the veins, often tufted in the axils; petioles short, to 1.5 cm long, glabrous or often golden villulose; stipules soon deciduous, widely triangular or compressed hemispherical, 2-4 mm long, acute. Inflorescences terminal and axillary, glabrate or minutely puberulent, the peduncle absent or short, the branches 3-5, often lenticellate, terminated usually by 3-flowered cymules or to single flowers; bracteoles linear subulate or ovate subulate, several located at the base of the pedicel; bracts lanceolate or subulate; pedicels short. Flowers large, showy, hypanthium oblong rotund or oblong, ca. 6 mm long, smooth, puberulent, the calycine cup ca. 1.5 mm long, glabrous within, with patches of oblong glands at the junction of the calycine lobes, these ca. 0.75 mm long, obtuse, the lobes 4-6, subulate, ca. 5.5 mm long, obtuse, marginally pubescent; corolla obovate oblong, slightly falcate and thus zygo- morphic, to 8 cm long, 1.8-3.0 cm wide, obtuse at the apex, attenuate acute at the base, petaloid, scattered pubescent outside, puberulent within, the lobes 2.0— 2.5 cm long, obtuse or acute; stamens 6, the anthers linear oblong, 1.7-2.2 cm long, ca. 1.5 mm wide, sagittate at the base, basifixed, the filaments free of the corolla tube, ca. 7.5 cm long; style linear, 7.5-9.5 cm long, ca. 1.5 mm wide, the stigmas 1.2-1.7 cm long, scarcely wider than the style, the ovary wall oblong in cross section, with conspicuous ducts. Fruits capsular, oblong obovoid or ob- ovoid rotund, 2.5-5.0(—7.0) cm long, 2.0-2.8 cm wide, strongly compressed, lig- nose, bisculcate, dark brown, lustrous, glabrous, with numerous white, punctate lenticels; seeds round or oblong, 2-15 mm long, brown, the body 2-5 mm long, alate, the wing membranous, entire or undulate. Coutarea hexandra is a common tree or shrub in the American tropics ex- tending from Mexico to Argentina. The fruits of Mori et al. 4097A are noteworthy being ca. 3 times longer than wide, the longest to 7 cm long. Dr. Thomas Croat (pers. comm.) notes that the flowers are bat pollinated. ‘‘Nino Muerto"; “Quina.” ` ARRO COLORADO ISLAND: Aviles X-5 (MO); Croat 5165, 7354, 8700 (all MO); Foster 1074 (MO); Gentry 1988 (MO); Zetek 3904 (MO). CANAL ZONE: Pipeline Road, Croat 16683 (MO). Fort Amador, D'Arcy 9264 (MO). Island beyond Fort Amador, D'Arcy & D'Arcy 6119 (MO). Ft. Kobbe, Duke 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 137 4257 (MO). Farfan Beach, Dwyer 3067 (MO); Dwyer et al. 457 (MO); Dwyer et al. 4693 (MO, SCZ). Madden Dam, Dwyer & Hayden 10 (MO). Farfan Beach, Dwyer & Hayden 7524 (COL, DUKE, K, MO, UC). Pipeline Road, Gentry 1447 (MO). Farfan Beach, Gentry 5501 (MO). Ancón Hill, Holdridge 6534b (MO). Farfan Beach, Lewis et al. (GH, K, MO, NY, UC M Tyson 1814 (MO). CHIRIQUÍ: San Félix to Cerro Flor, Allen 1953 (MO). CocLE: Nueva Gorgona, D'Arcy & D'Arcy 6283 (MO). 5 mi S of Antón, Tyson & Blum 2572 (MO). La Vinta, Woodson et al 1706 (MO). COLÓN: 4 km NW of Salamanca, 13 km NE of Buenos Aires, Nee 9067 (MO). DARIEN: Río Tuqueza below Quebrada Venado, Bristan 1078 (MO). Isla Saboga, Duke 10357 (MO). Punta Garachine, Duke 10490 (MO). 1- 4 mi N of Pucro, Duke 13028 gee Trail, Rio Pucro to Quebrada Maskia, Duke 13071 (MO). HERRERA: Pese, Allen 796 (MO). AMA: Burunga, Austin 28 (MO). Morro Island, D'Arcy & D'Arcy 6785, 6786 (both MO). Nueva Gorgona, Duke 4532 (MO). E of Bejuco, Duke 4548 (MO). El Llano, Duke 5864 (MO). Taboga Island, Dwyer 2807, 3067 (both MO). Cerro Azul, Gentry & Dwyer 5508 (MO). Juan Diaz, Hunter 252 (MO). Playa de Venado, Mejia 2 (MO). 6 mi SW of Arraijan, road to Bique, Nee 6898 (MO). Las Sabanas, Pittier 6757 (MO). Nuevo Emperador, vicinity of Arraijan, Rivas de Hoyos 6 apud N ú Chorrea, Tyson 5936 (MO). S of La Capitana, Tyson 6765 (MO). Chepo. Tyson 6776 (MO). A . Woodson et al. 1386 (MO). Isla Taboga, Woodson et al. 1495 (MO). Pacora and Chepo, Woodson el al. 1668 27. CRUSEA Crusea Cham. & Schlecht., Linnaea 5: 165. 1830. TYPE: C. rubra (Jacq.) Cham. & Schlecht. Herbs, often perennial, the stems usually 4-angled. Leaves mostly lanceolate, the lateral nerves conspicuous; petioles adnate to the stipular sheath, the latter setiferous. Inflorescences capitate, surrounded by foliaceous bracts. Flowers with the 4 calycine lobes alternating with minute glands, the calyx persistent in fruit; corolla funnelform, glabrous within, the 4 lobes valvate; ovary 2-celled, the ovules 1 per locule. Fruits capsular, didymous, the mericarps or cocci indehis- cent, thin, the endocarp transparent, smooth, ecostate, the ventral face grooved, narrow, the flanks of endosperm folded against the deep hilar ridge, the seeds 1 per locule, the testa cells contorted. The genus Crusea, according to Anderson (1972), extends from "Southern Arizona and New Mexico through Mexico and Central America to northern Pan- ama." Literature: Anderson, W. R. 1972. A monograph of the genus Crusea (Rubiaceae). Mem. N.Y. Bot. Garden 22(4): 128 a. Corolla red, rose or violet; diea cup 3-4 mm long; anthers ca. 3 mm lon MP mE EE E l. C. coccinea "a r, chiriquiensis aa. Corolla white; calycine cup p less than I mm long: anthers ca. | mm lo rae i ú ayer d qm longiflora 1. Crusea coccinea DC., Prodr. 4: 567. 1830. TYPE: Pavón, not seen.—Fig. 30. la. Crusea coccinea DC. var. chiriquiensis W. R. Anderson, Mem. N.Y. Bot. Garden 22: 45. 1972. TYPE: Panama, Woodson & Schery 358 (F, holotype; GH, MO, isotypes). Upright or creeping herbs, stems angular, ribbed, glabrous, drying purple. Leaves oblong or ovate oblong, 1.5-6.0 cm long, 0.5-2.0 cm wide, acute at the 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 30. Crusea coccinea var. chiriquiensis W. R. Anders.—A. Habit (x%).—B. Flower (x 1.55).—C. Calyx (x3.5).—D. Ovary in longisection (3.5). [After D'Arcy & D'Arcy 6519. ] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 139 apex, occasionally distinctly acuminate, the acumen to 1.2 cm long, basally cu- neate to acute, often slightly decurrent on the petioles, the costa slender, often indistinguishable from the lateral veins, plane or immersed above, prominent beneath, to 0.7 mm wide, the lateral veins usually 5, strongly arcuate ascending, arising at a sharp angle, immersed or plane above, prominent beneath, charta- ceous, not strongly discolorous, the upper surface striate, white hirtellous be- tween the veins, white hirtellous beneath; petioles 0.5-0.8 cm long, to 0.1 cm wide, these adnate to the lip of the stipular cup; stipules cylindriform, to 1.5 cm long, to 0.5 cm wide, membranous, glabrous or occasionally villosulose, the sheath extending beyond the next superior node, with a terminal fringe of 1-12 iege tipped linear setae, often hirtellous, to 1.4 mm long, ca. 0.2(-0.5) mm wide. Inflorescences terminal, sessile, the flowers several, ascending, the hypan- thiums hidden by the terminal stipules and the leaf bases. Flowers with the hy- panthium turbinate, 3-5 mm long, ca. 2 mm wide at the truncate apex, the ca- lycine cup cupuliform or campanulate, 3-4 mm long, ca. 3 mm wide at the apex, glabrous, petaloid, the lobes 4, unequal, lanceolate, to 8 mm long, ca. 1.2 mm wide at the base, acute, glabrous except hirsute marginally, usually with 2-3 small glands at the junction of 2 adjacent lobes, oblong, to 1 mm long, obtuse; corolla red, rose or violet, to 35 mm long, the tube narrowly funnel shaped, sharply attenuate below middle, to 0.3 cm wide near the mouth, glabrous within and outside, the lobes 4, elliptic or ovate oblong, 8-11 mm long, to 5 mm wide, with a few white hairs abaxially and terminally; stamens 4, exserted, the anthers oblong, ca. 3 mm long, twisted after shedding pollen, versatile, the filaments slender, plane, equal to the corolla lobes, to 7 mm long, attached near the mouth; style filamentous, as long as the corolla tube, minutely puberulent on the upper 14, the stigmas 2, slender, ca. 1.8 mm long. Fruits not seen. Crusea coccinea var. chiriquiensis is known from Costa Rica and Panama. Croat 15956, with only flower buds, is described on the label as ‘‘white. CHIRIQUí: Río Chiriquí Viéjo Valley, Allen 1369 (MO). Cerro Punta, Blum et al. 2426 (MO). NW side of Cerro Pando cloud forest, Croat 15956 (MO). Cerro Respinga above Cerro Punta, D'Arcy 10709 (MO): D'Arcy & D'Arcy 6519 (MO). Cerro Punta, Ridgway & Solis 2391 (MO, UC). Rio Chiriqui Viéjo, Monte Lirio, Seibert 220 (MO). Casita Alta to Cerro Copete, Woodson & Sc hery 358 O). 2. Crusea longiflora (R. & S.) Anderson, Mem. New York Bot. Gard. 22: 89. 1972. Spermacoce longiflora Willd. ex R. & S., Syst. Veg. 3: 531. 1818. Type: México, Humboldt & Bonpland (B-Willd., not seen). Crusea brachyphylla Schlecht. & Cham., Linnaea 5: 165. 1830. TYPE: Schiede & Deppe 532b (F, isotype). Spermacoce tenuiflora Ses. & Moc., Fl. Mex 1893. TYPE: not see Crusea chiapensis Brandegee, Univ. Cal. E a 6: 374. 1917. rvyPE: México, Purpus 7526, not seen RUNS brachyphylla fo. boi dca Loes., Verh. Bot. Vereins Prov. Brandenburg 65: 115. 1923. : Seler & Seler 12 Herbs to subshrubs, 0.5—50.0 cm tall, branched or unbranched, the stems slender, the hairs appressed, deflexed, the nodes well spaced. Leaves sessile, lanceolate, occasionally ovate oblong, 0.5-3.0 cm long, 0.2-0.6 cm wide, acute or deltoid at the apex, acute at the base, the costa slender, plane or immersed 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 above, prominulous beneath, the lateral veins (1—)2—5, strongly arcuate ascend- ing, without a submarginal undulate vein, the intervenal areas smooth, the vein- lets not evident, chartaceous to thinly succulent, concolorous, the hairs above strigose, appressed, golden colored or white, subulate, 0.2-0.7 mm long, the hairs beneath as above but mostly on costa and veins, the margin somewhat thick, minutely ciliolate or with short, thick, ascending hairs; stipules to 4 mm long, truncate, the setae 3-5, erect, the hairs stiff, to 1.7 mm long. /nflorescences terminal, 0.5—4.0 cm in diam., the uppermost leaves involucrate, the stem im- mediately below the involucre of leaves with numerous, weak, reflexed hairs. Flowers subsessile, the hypanthium oblong, shorter than the calyx, minutely puberulent; calycine cup 0.2-0.5 mm long, the lobes 4, subulate, erect, slightly spreading, 1.6-3.2 mm long, usually slightly unequal, the hairs slender, stiff, to 0.8 mm long, with a minute gland in the sinus of 2 adjacent lobes; corolla white, the tube slender, 0.6-0.7 cm long, ca. 0.2 mm wide, minutely puberulent to glabrous outside, glabrous within, the lobes 4, oblong, 1.0-2.2 mm long, obtuse; stamens 4, exserted, the anthers oblong, ca. 1 mm long, the filaments threadlike, to 1.8 mm long, attached at the mouth; style filamentous, about as long as the tube, the stigmas 2, 0.3-1.0 mm long, slightly wider than the style. Fruits not seen. Crusea longiflora extends from Mexico to Panama. Superficially the two Pan- amanian collections seem to belong to different species but floral dissections indicate that they are conspecific. On the dwarf plants of Davidse & D'Arcy 10348, the leaf blades show a reduction of lateral veins to 1-2 pairs. cHiRIQUÍ: Large old lava flow, 3 km NE of El Hato del Volcán, base of Volcán de Chiriquí, 1— 3 km E of highway, 1500-1800 m, Davidse & D'Arcy 10348 (MO). Volcán Chiriquí above Boquete, roadside, D'Arcy 9793 (MO). 28. DECLIEUXIA Declieuxia H.B.K.,?" Nov. Gen. Sp. Pl. folio 3: 275, tab. 281. 1819; quarto 3: 352, tab. 281. 1820. TvPE: D. chiococcoides H.B.K. Herbs or subshrubs, sometimes ericoid in habit. Leaves occasionally verti- cillate, often coriaceous, frequently sessile; stipules interpetiolar, often subulate. Inflorescences terminal or axillary, cymose, usually branching dichotomously, or spikelike and racemiform; bracts often absent. Flowers with the calycine cup reduced, the lobes usually 4, equal or unequal; corolla variable in color, infun- dibuliform or cylindrical, the lobes 4, valvate; ovary 2-celled, the ovules 1 per cell, attached basally. Fruits a schizocarp, at first fleshy, didymous, indehiscent, compressed, the thin endocarp usually hard; seeds compressed. Declieuxia ranges from Mexico, through Central America, the West Indies, and South America. There are about 40 species in the genus. 38 For synonymy see Kirkbride (1976). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 141 Literature: Kirkbride, J. H., Jr. 1976. A revision of the genus Declieuxia (Rubiaceae). Mem. New York Bot. Gard. 28(4): 1-87. 1. Declieuxia fruticosa (R. & S.) Willd. ex Kuntze, Rev. Gen. Pl. 1: 279. 1891.— Fic. 31 Houstonia fruticosa Willd. ex Roem. & Schult.,?? Syst. Veg. 3: 527. 1818. TYPE: Venezuela, Humboldt & Bonpland 223 (B-Willd., holotype; P, isotype, neither seen). Herbs, woody, to 3.5 m tall, the branchlets arising from the base or above, rarely restricted terminally, smooth, glabrous, delicately alate, the nodes well spaced. Leaves sessile, occasionally 3 per node, lanceolate, occasionally slightly falcate, to 5 cm long, to 1.2 cm wide, acute at the apex, obtuse to acute at the base, the costa prominulous above and beneath, the lateral veins ca. 5, arcuate, prominulous above and beneath, the intervenal areas reticulate, usually concol- orous, lustrous above, marginate; stipules deciduous. Inflorescences terminal, cymose paniculate, a peduncle like axis (really a primary cyme) often present, to 2 cm long, or the stem tip with 3 primary cymes, each with an axis terminated by a sessile flower and 2 lateral branches, these branching into small cymules, the more profusely branched inflorescences often flat topped; bracteoles tending to persist, subulate. Flowers with the hypanthium subrotund, truncate, ca. 0.8 mm long, glabrous, the calycine teeth 4, erect, linear oblong, ca. 0.8 mm long, obtuse, glabrous; corolla white, the tube cylindrical, ca. 6 mm long, petaloid, densely villose below the middle within, the lobes oblong, to 2 mm long, glabrous, not cucullate; stamens 4, the anthers exserted, oblong, to 1 mm long, dorsifixed, the filaments to 1.5 mm long, attached in the middle of the tube or below; ovarian disc bilobed, pulvinate, the stigmas 2, slender, to 0.3 mm long, the style slender, to 2.5 mm long, attenuate basally, the ovary irregularly ribbed, 2-carpellate. Fruits sessile, rotund and ca. 2 mm in diam. or bilobed (2 carpels developing) and then wider than long, drying black, marcescent, often reticulate, glabrous, usually marginate; bracteoles persistent, subulate. Declieuxia fruticosa ranges from Mexico south to Brazil. cHiRIQUÍ: San Félix to Cerro Flor, 100-800 m, nn 1930 (MO). Savanna below Boquete, ca. 800 m, D'Arcy 9773 (MO). 2 mi S of Boquete, 3-4 , D'Arcy & D'Arcy 6309 (MO). nde 4000 ft, Davidson 701 (MO); Dw yer Hayden 7593 a Parte h 69-80, 69-128 (both MO). Boquete, Llanos Francia, Stern et al. 1152, (both MO). Boquete, 1200-1500 m, Woodson & Schery 791 (MO). cocLé: Nata, Allen 827 fe. Santa Clara Beach, Croat 9588 (MO). Penonomé, Ebinger Hacienda La Joya, Dodge et al. 16897 (MO). Savanna near Rio Pacora and Chepo Highway, Duke 5898 (MO). Savannalike areas, road to Cerro Campana, Duke 6029 (MO). Forest n to grasslands, Cerro Campana, 2400-2700 ft, Duke 8701 (MO). Tocumen Airport, Dwyer 1251, 4428 (both MO). Cerr us rh 4847 (MO, SCZ, UC); Dwyer E Kirkbride 7824 (MO). a Arraijan, Guerra 24 (MO). Cerro Campana, Lewis et al. 3100 (COL, DUKE, K, MO, UC, VEN). Taboga Island, je. 489 (MO). Cerro Campana, 2600 ft, Mc Di 6824 (MO); Nee 6872 (MO): With- erspoon & Witherspoon 8365 (MO). Taboga Island, Woodson et al. 1467 (MO). VERAGUAS: Yeguada, 39 For synonymy see Kirkbride (1976) p. 17. 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 x Y j E T lA VW, wy f Mae | y A s Yy FIG ia fruticosa (R. & S.) Kuntze.—A. Habit (x.6)—B. Flower, opened to show i spend en. ee Aller s 1271.1 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 143 Hladik 383 (MO). San José on Santiago-Santa Fé Road, 18 km S of Santa Fé, 400 m, Nee 8/80 (MO). Canazas, Tyson 3740 (MO). 29. DEPPEA Deppea Schlecht. & Cham., Linnaea 5: 167. 1830. TYPE: D. erythrorhiza Cham. & Schlecht. Choristes Benth., Pl. Hartw. 63. 1840. rvPE: C. obtusifolia Benth. Shrubs. Leaves membranous, petiolate; stipules minute. Inflorescences ter- minal or axillary, the flowers rarely solitary, disposed in cymes or umbels, ebrac- teolate. Flowers pedicellate; calyx 4-lobed, the lobes persistent; corolla rotate, the tube often shorter than the lobes, glabrous within, the lobes 4, contorted; stamens 4, exserted, the filaments short, attached near the corolla mouth; style slender, the stigmas capitate or the 2 lobes evident, the ovary 2-celled, the ovules numerous, imbricate, the placentas peltately adnate to the septum. Fruits cap- sular, small, turbinate or obovoid, scarious to coriaceous, costate, loculicidally dehiscent from the apex; seeds numerous, subglobose, minute, obtusely angular, the testa hard, granulate. Deppea ranges from Mexico to Venezuela. The genus includes about 20 species. a. Flowers with hypanthium sparsely villose; fruits turbinate, 0.4—0.5 cm long, 8-costate; high 500 m l. mountain species, 1500—4 D. scene aa. Flowers with hypanthium glabrous; fruits rotund, to 0.2 cm long, 12-costate; low elevation species 2. D. panamensis 1. Deppea grandiflora Schlecht., Linnaea 19: 748. 1847. syNTYPES: México, Lei- bold 235, 253, 263, none seen.—Fic. 32 D. costaricensis Polak., Linnaea 41: 566. 1877. rype: Costa Rica, Polakowski 148 (not seen; photo, F) D. floribunda Hemsl., Diag. Pl. Nov. 31. 1879. TYPE: Botteri 937 (not seen; P, holotype; photo, MO). D. longipes Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. TYPE: Panama, Pittier 3070 (US. ho- otype). Trees 3-8 m tall, the branchlets terete, smooth, glabrous to puberulent. Leaves oblong, oblong rotund or oval, 6-17 cm long, 2.0-6.5 cm wide, deltoid at the apex, acuminate, the acumen to 1.5 cm long, acute, cuneate to acute at the base, the costa plane above, prominulous beneath, the lateral veins 8-12, arcuate, prominulous above and beneath, chartaceous, scarcely bicolorous, glabrous above except on veins, appressed villose on the veins beneath, the hairs often tufted in the vein axils; petioles to 4.5 cm long, to 0.2 cm wide, wiry, often arcuate and deflexed, densely pubescent: stipules deciduous, minute, broadly deltoid and glanduliferous (fide Standley). Inflorescences terminal or pseudo ter- minal, occasionally longer than the uppermost leaves, paniculate, the peduncles 3-4 cm long, villosulose, terminated by 3 primary branches, these to 2.5 cm long, strongly ascending, in turn branching, ultimately producing cymes. Flowers with the hypanthium oblong or turbinate, 1.6-2.0 mm long, ribbed, sparsely villose, the calyx cup scarcely measurable, the teeth reduced to 5 points, 0.2-0.6 mm 144 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 A Faramea altipetens Dwyer, spec. nov. Arbores ad 4 m altae, ramulis glabris ultime tortis. Folia oblongo-la ata, venis mE 14 interveniis conspicue reticulatis m minimis inulis; I tipulis ca. 0.4 mm lon ahr ed A. terminales solitariae, pedunculis 4-5 r radii is dispositis terminatis ramo ends e 1-2 jugis ramulorum oppositorum cymu V us Flores glabri; corolla alba tubo ca. 15 mm longo, lobis 4 ad 7 mm longis; antheris subsessilibus, ca. 4 mm longis. Fructus non visi 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 163 rolla white, the tube narrowly cylindrical, ca. 15 mm long, the lobes 4, crassate, triangular subulate, to 7 mm long; stamens included, 4, the anthers subsessile, linear ovate, ca. 4 mm long, minutely apiculate, inserted near the middle of the corolla tube; style ca. 9 mm long, the stigmas 2, slender, ca. 2 mm long. Fruits not seen. Faramea altipetens is known only from Panama but is to be expected in nearby Colombia. It is marked by the combination of short stipules and anthers and a relatively elongate corolla tube. The smallest veinlets are prominulous with the intervenal areas having a striking reticulate appearance. It is closely related to F. pauciflora Dwyer in which the flowers are white and few, ca. 3 per cyme. F. pauciflora, however, has much larger stipules, corolla tube, and anthers. DARIEN: Cerro Tacarcuna, 1500 m, Gentry & Mori 14013 (MO). 2. Faramea bocataurensis Dwyer.* TYPE: Panama, Kirkbride & Duke 806 (MO, holotype). Small trees, to 5 m tall, the branches smooth, glabrous, the nodes well spaced. Leaves lanceolate, 6-11.5 cm long, 2-2.7 cm wide, acute at the apex, acuminate, the acumen to 2 cm long, ca. 0.15 cm wide in the middle, acute or subobtuse at the base, the costa prominulous, the lateral veins 6-9, arcuate, prominulous, with 1-4 smaller veins between a pair of adjacent veins, these subparallel but soon branching, the intervenal areas spreading reticulate, chartaceous, bicolorous, gla- brous; petioles slender but stiff, ca. 0.5 cm long; stipules connate, the sheath cylindrical, ca. 0.7 cm long, stiffly petaloid, glabrous, each part with an awn 0.8 mm long. /nflorescences terminal, solitary, glabrous, to 3.5 cm long, epedun- culate, cymose racemose, the branches to 5 mm long or absent, the cymules few flowered, bracteoles deciduous, subulate, 0.8-2 mm long; pedicels ca. 4 mm long. Flowers purple, glabrous; hypanthium subrotund, ca. 0.5 mm long, the calycine cup turgid, to 0.35 mm wide, smooth, petaloid, eglandular within, with 4 subulate lobes, to 0.8 mm long; corolla (only one seen in bud) to 7.5 mm long, the tube cylindrical, rigidly carnose, to 6 mm long, ca. 1 mm wide toward the base, the lobes 4, narrowly ovate lanceolate, ca. 2 mm long; stamens 4, the anthers nar- rowly oblong, ca. 4 mm long, obtuse and subsagittate at the base, the filaments slender, ca. 1 mm long, affixed near the middle of the tube, the disc of the ovary to 1.3 mm long, 3 times longer than wide, the style linear, ca. 6 mm long, the stigmas 2, narrowly oblong, ca. 1 mm long. Fruits not seen. Faramea bocataurensis is known only from Panama. While the flowers are known only from the bud stage, the calycine cup is remarkably short as are the Ba duy bocataurensis Dwyer, spec. nov. Arbores parvae ad 5 m altae. Folia lanceolata, 6— 11.5 cm long .7 em lata, apice acuta caudato-acuminata, acumine ad 2 cm longo, basi acuta vel subobtusa, ea s lateralibus 6-9; petiolis ca. 0.5 cm longis; stipulis connatis vagina cylindrica, ca. cm n onde parte utraque seta ad oon mm longa. a terminales solitariae glabrae, ad 3.5 e aes praeditis: sessi d. saepe decidit, isle 0. 8-2 mm longis. Flores glabri; pedicellis ca. mm longis, cupula calycis turgida, ad 0.35 mm lata, ca. 0.5 mm longa, dentibus 4, ad 0.8 mm longis; corolla purpurea, ad 7.5 mm longa; antheris lineari-oblongis, ca. 4 mm longis. Fructus non visi. 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 calycine teeth. The leaves are as narrow as any species of Panamanian Faramea and have the fewest lateral veins. The reduced racemelike inflorescence is note- worthy. BOCAS DEL TORO: E slope of La Zorra to divide, Kirkbride & Duke 806 (MO). 3. Faramea bullata Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 294. 1929. TYPE: Panama, Cooper 507 (F, holotype; NY, isotype). Trees or shrubs, 2—4.5 m tall, glabrous, the branchlets smooth, the nodes well spaced. Leaves oblong, 15-21.5 cm long, 5.5-7 cm wide, acuminate, the acumen ca. 2 cm long, basally cordate or occasionally rounded, the costa prominent, the lateral veins ca. 15, immersed above, with 1-2 somewhat irregular intermediate veins between pairs of lateral veins, the submarginal veins 2, the outermost ca. 1.5 mm from the margin, the other ca. 5 mm from the margin, subcoriaceous, shiny green above, bullate, glaucescent and pruinose beneath; petioles ca. 0.5 cm long, glabrous; stipules presumably connate basally, each part oblong rotund, ca. 8 mm long, tapering acutely toward the apex, the awn subulate, to 14 the length of the sheath, coriaceous, glabrous. Inflorescences cymose paniculate, to 7.5 cm long, often almost as wide as long; peduncle thick or slender, to 3 cm long, the branches few, angular ascending; pedicels 2-6 mm long. Flowers with hypan- thium and calycine cup short, the teeth triangular or subulate, to 1 mm long; corolla blue or bluish white, the tube ca. 9 mm long, ca. 1.2 mm wide in bud. Fruits compressed rotund, to 1.5 cm wide, smooth, ca. twice as long as wide, drying black, smooth, glabrous; "seed gray, deeply and narrowly exsculpate at the base, smooth, ca. 1 cm long” (fide Standley). Faramea bullata is known only from Panama. In preparing the above descrip- tion I have drawn freely from Standley's original description. BOCAS DEL TORO: Kankintoe, 10 mi above Holstein near Almirante, Cooper 507 (F, NY, US). Punta Pena, Chiriquicito, Lewis et al. 2145 (F, MO). vERAGUAs: Caribbean slope above Río Primero Brazo, 5 mi NW of Santa Fé, Liesner 965 (MO) 4. Faramea caput-anguis Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 39. 1968. TYPE: Panamá, Dwyer & Gauger 7375 (MO, holotype). Trees to 8 m tall, the branchlets smooth, drying black and glabrous, the in- ternodes to 7 cm long. Leaves oblong, to 26 cm long, to 12 cm wide, acute or acuminate at the apex, the acumen to 1.5 cm long, acute at the base, the costa subplane above, prominent beneath, the lateral veins ca. 20, arcuate, prominent, 1-2 cm apart, the venules pinnatiform, thin coriaceous, often yellow green be- neath, minutely yellow puberulent; petioles 2-4 cm long, smooth; stipules per- sistent, triangular, to 3 cm long, to 0.8 cm wide, acute, lignose, glabrate. In- florescences usually 1 per axil; peduncles divergent, stiff, smooth, glabrous, expanding terminally, to 9 cm long; heads ovoid, apically blunt, solitary, cymose, at first gemmiform and fusiform oblong, to 2 cm long, ca. 1 cm wide, the outer bracts ca. 5, widely fusiform oblong, glabrous, foliose; flowers 7-9 per head, each 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 165 pair with 2 to several bracteoles. Flowers glabrous, the calyx narrowly campan- ulate, to 8 mm long, the lobes 4, unequal, subulate, the shorter lobes ca. 4 mm long, the longer lobes ca. 8 mm long; corolla white, tube ca. 25 mm long, narrow medially, dilated above, the lobes 4, lanceolate, to 7 mm long, acute; stamens 4, the anthers linear oblong, ca. 5.5 mm long, the filaments affixed near the middle of the tube: ovules orbicular. Fruits oblong, to 1.5 cm long, to 1 cm wide including persistent calyx, smooth, glabrous, glandular shiny, drying black, the calyx 2-5 mm long. Faramea caput-anguis is known only from Panama. PANAMA: Cerro Jefe, Busey 791 (MO). La Eneida, Cerro Jefe, Correa & Dressler 954, 1120 (both MO). Cerro Jefe, Dwyer et al. 7248 (MO); Dwyer & Gauger 7375, 7388A (both MO): Dwyer et al. 8236 (MO). 9.4 km N of Goofy Lake, 900 m, Folsom et al. 1963A (MO). Cerro Jefe, Luteyn & Foster 1140 (MO). 5. Faramea eurycarpa Donnell Smith, Bot. Gaz. 44: 113. 1907. TYPE: Costa Rica, Tonduz 12880, not seen. F. platycarpa Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 140. 1967. TYPE: Ecuador, Owenby 2711 (MO, holotype). Trees to 4 m tall, the ultimate branchlets becoming angular, smooth, glabrous, the nodes well spaced. Leaves oblong or oblanceolate, 5-16 cm long, 1.5-5.5 cm wide, rounded or obtuse at the apex, rarely cuneate, contracted into a slender acumen, the acumen to 1.3 cm long, ultimately obtuse, basally cuneate to obtuse, rarely truncate or auriculate, often inequilateral, the costa prominulous, the lat- eral veins ca. 13, at first nearly perpendicular, then widely arcuate, united to form a distinct undulate submarginal vein, to 0.3 cm from the margin with 1—2 smaller veins between 2 lateral veins, the intervenal areas spreading reticulate, charta- ceous to papyraceous, concolorous, glabrous, the margins delicately revolute; petioles slender to stout, to 0.5 cm long, glabrous; stipules connate, the 1 at the twig tip dome shaped, to 6 mm long, the awn shorter than the body, the inferior stipules more cylindrical, to 7 mm long, the sheath separating above into 2 tri- angular parts, each with an awn ca. 2 mm long, glabrous. /nflorescences terminal, solitary, oblong rotund to subcuneate, to 9 cm long, to 9 cm wide, paniculate, the peduncle 2-5 cm long, the lowermost branches opposite, to 3 cm long; the upper branches disposed as 2—3 pairs, usually opposite, the branchlets terminating in a fan of few-flowered cymules. Flowers with pedicels ca. 1 cm long, stiff; hypanthium ca. 0.75 mm long, glabrous, the calycine cup compressed cup shaped, shorter than the hypanthium, scarious and eglandular within, the teeth 4-5, ir- regular, unequal, triangular or subulate, to 0.8 mm long, acute; corolla purplish or white with a purplish apex, narrowly cylindrical, to 1 cm long, glabrous, car- nose: corolla lobes 4, ovate, to 3 mm long, occasionally slightly acuminate; sta- mens subsessile, the anthers 4, narrowly oblong, 3-4 mm long, minutely apicu- late, attached near the middle of the tube; style slender, ca. 8 mm long, the stigmas fingerlike, slightly unequal, to 1 mm long, scarcely wider than the style. Fruits drying black, bright blue, compressed rotund, wider than long, to 1.3 cm wide, smooth, glabrous, the calycine scar ringlike, scarcely elevated. 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Faramea eurycarpa is known from Costa Rica and Panama. It is also known from Colombia and Ecuador. Leon 537 (MO) from the Dept. Choco, Colombia, seems to be F. eurycarpa, although it reaches 10 m in height, has a strongly contracted inflorescence, and stipules to 1 cm. Forero et al. 3073 (MO), also from Dept. Choco, superficially resembles F. eurycarpa but has corolla lobes to 15 mm in length. COCLÉ: La Mesa, Dwyer 8318 ee m Santa Rita Ridge, Dwyer 8554 (MO). DARIEN: 1-3 mi N of de Duke 14005 (MO). PAN : Cerro Campana, Allen 1877 (F, MO, US); mud x Dressler 291 (MO); Croat 14217, 17240. 22779 (all MO); Duke 10737 (MO, OS); Dw yer et al. 4 (MO); Gentry 1831 (MO, NY), 4904, 4952 (both MO); Kirkbride & Hayden 277 (MO, NY); a a 337 (MO, NY); Lewis et al. 3049 (MO, SCZ, UC, VEN). El Llano-Cartí Road, 21.1 km from Pana- merican Highway, Mori & Kallunki 5112 (MO). El Llano-Cartí Road, 11-12 km from Panamerican igh (MO) i 6 16 km N of Panameric ea Highway, Nee & Dressler 9356 (MO). El Llano-Carti Road, Nee et al. 8769 ul Cerro Campana, 825- pen m, Nee & Stockwell 11581 (MO); Porter et al. 4162 (COL, DUKE, MICH, MO, NY , SCZ, ae N). 6. Faramea jefensis Dwyer & Hayden, Phytologia 15: 56. 1968. TYPE: Panama, Tyson et al. 3396 (MO, holotype; NY, isotype). Trees or shrubs, the branchlets strongly arcuate, ultimately angular, black when dry, the nodes somewhat turgid, the internodes to 5 cm long, minutely ciliolate. Leaves narrowly elliptic, 9-15 cm long, 2-5 cm wide, long cuspidate, the cusp to 1.5 cm long, ca. 1.5 mm wide, basally cuneate, often inequilateral, the costa prominulous above and beneath, the lateral veins ca. 20, scarcely ar- cuate, delicately prominulous beneath, the submarginal vein irregular undulate, ca. 3 mm from the margin, thin coriaceous, concolorous or discolorous, drying a dull yellow beneath, glabrous, the margin scarcely thickened; petioles to 0.8 cm long; stipules connate, cylindrical, ca. 2.5 cm long, the sheath separating into 2 triangular parts ca. 1⁄2 the length of the sheath, minutely puberulent. /nflores- cences rarely reduced to a single flower, usually cymose thrysoid, to 2.5 cm long, to 3 cm wide, the peduncle scarcely developed, crassate, ca. 3 mm wide, the branches few, to 1.5 cm long, ascending; bracts foliaceous, enclosing the juvenile inflorescence, ovate, 2-4 cm long, 1-2 cm wide, cuspidate, glabrous, variegated medially, the margins green; pedicels to 3 mm long. Flowers with the hypanthium and calyx to 6 mm long, glabrous, the calycine cup cupular, to 1.5 mm long, ca. 2 mm wide, the teeth 4, minute; corolla dark blue, the tube cylindrical, 9-12 mm long, the lobes 4, crassate, triangular in cross section, to 8 mm long; stamens 4, the anthers 3.5-4 mm long, inserted below the middle of the corolla tube; style ca. 8 mm long, the stigmas linear, ca. 2 mm long, erect, minutely papillate. Fruits sessile or subsessile, compressed rotund, 0.3-0.5 cm long, 1-1.6 cm wide, smooth, glabrous, drying black or yellow, the calycine cup scar annular, ca. 2 mm wide Faramea jefensis is known only from Panama. AMA: 6 miles above Goofy Lake, road to Cerro Jefe, Croat 15222 (MO). Cerro Jefe, 2900 ft, «on. et al. 7279 (MO); Dwyer & Gauger 7328 (GH, MO, NY), 7338A (MO): Foster & Kennedy 1875 (MO); Tyson et al. 3396 (MO). 1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 167 7. Faramea liesneri Dwyer.“ TvPE: Panama, Busey 9/3 (MO). Shrubs to 3% m tall, the branchlets terete, smooth, glabrous, ultimately an- gular, the nodes well spaced. Leaves oblong, 4.5-9.5 cm long, 1.3-4 cm wide, obtuse or widely deltoid at the apex, acuminate, the acumen often barely dis- cernible, ultimately obtuse, basally acute, the costa prominulous above, the lat- eral veins ca. 13, perpendicular to the costa, forming an undulate submarginal vein ca. 1.5 mm from the margin, the margin lightly revolute, subcoriaceous, glabrous, concolorous, drying emerald green or whitish green; petioles to 0.8 cm long, rigid, glabrous; stipules connate, glabrous, with each part oblong rotund, to 4 mm long, with a median keel continuing into a terminal rigid bristle shorter or longer than the body, coriaceous. Inflorescences terminal or axillary, umbel- liform, to 6 cm long, 1.5—4 cm wide, glabrous, the peduncle 1-2.5 cm long, the cymes radiately disposed; pedicels slender, 0.5-1 cm long. Flowers with the hypanthium oblong rotund, to 1.2 mm in diam., the calycine cup much reduced, truncate, erose and minutely puberulent marginally, the lobes inconspicuous, to 0.3 mm long, truncate or obtuse; corolla violet blue, the tube narrowly cylindrical, 5-6.5 mm long, ca. 0.1 cm wide, thin carnose, glabrous inside and out, the lobes 4, unequal, narrowly oblong, to 6 mm long, to 2 mm wide, obtuse, glabrous; stamens 4, the anthers oblong, ca. 3 mm long, obtuse, attached above the middle of the tube; style linear, to 2.5 mm long, the stigmas 2, to 0.8 mm long, ovarian disc to 0.6 mm long, rounded. Fruits on pedicels to 1.5 cm long, equal in length, slender, radially disposed, the pedicel scars obvious, ringlike, the bracteoles few, stipulelike, the body of the fruit round to compressed rotund, ca. 0.5-1.2 cm in diam., smooth, glabrous, often emerald green when dry, the calycine cup small. Faramea liesneri is known only from Panama. It is named in honor of Ronald Lee Liesner, Missouri Botanical Garden, who has contributed much to our knowledge of the flora of Panama. It is closely related to F. woodsonii Standley, a small-leafed species, which has only stiffly chartaceous leaves and has a much reduced inflorescence in which the flowers number about four. Faramea liesneri has stiffly coriaceous leaves which dry a striking emerald green. This, coupled with a much reduced, marginally pubescent calycine cup, as well as much reduced calycine Bex serves to distinguish the new species. : Road to Cartí, 8-19 km N of El Llano, 350-500 m, Busey 913 (MO); Folsom et al. 1495 (Oy pue 7733 (MO): Mori & Kallunki 2265, 2889 (both MO). 8. Faramea luteovirens Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. TYPE: Panamá, Pittier 4038 (US, holotype). Trees to 10 m tall, the twiglets smooth, shiny, terete, finally angular, glabrous, 7 Faramea lie 'sneri Dwyer, spec. nov. Frutices ad 2 m alti. Folia oblonga, 4.5-9.5 cm longa, 1.3-4 lata, a brevi, venis lateralibus ca. 15, subcoriacea glabra in sicco smaragdina vel albido-smaragdina; Mee ad 0.8 cm longis: stipulis connatis glabris parte yen oblongo- rotunda, mm longis, in medio carinata seta rigida quam vagina breviori vel vix longiori. Inflorescentiae terminales vel axillares, ad 6 cm longae, umbelliformes cymis radiate dispositis. Flores pedicellis 0.5— 1 em longis: cupula calycis brevissima lobis inconspicuis, ad 0.3 mm longis: corolla tubo angusto- eylindries, 5-6.5 mm longo, lobis 4, ad 6 mm longis; antheris ca. 3 mm longis. Fructus ca. 0.5 cm iam., smaragdini. 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 the nodes well spaced. Leaves oblong, obovate oblong, oblong rotund, to 15 cm long, to 8 cm wide, obtuse, rounded or truncate at the apex, cuspidate, the cusp to 1.5 cm long, often falcate, cuneate to acute, rarely truncate at the base, equi- lateral, the costa plane to prominulous above, prominent beneath, the lateral veins 10-13, widely arcuate, prominulous above and beneath, the intervenal areas reticulate, the veinlets often disappearing, the margin conspicuously thickened, coriaceous, glabrous, concolorous or discolorous, usually drying lemon yellow; petioles stiff, to 1 cm long, glabrous; stipules appressed, deltoid to compressed hemispherical, 2-4 mm long, the awn subulate, 3.5-5.5 mm long, coriaceous, glabrous. /nflorescences axillary or occasionally terminal, the peduncle usually suppressed (or if present to 2 cm long), branches few or absent, the flowers 1—2 per axil; pedicels strongly ascending, to 5 cm long, stiff, lignose, glabrous, ex- panded at the apex. Flowers with hypanthium oblong, ca. 9 mm long, coriaceous, glabrous, the calycine cup ca. 4 mm long with patches of minute appressed glands within, these covered with a membranous "skin," the calycine teeth absent, the margin undulate; corolla white, the tube narrowly cylindrical, falcate, ca. 2.2 cm long, ca. 4.5 mm wide medially, contracted below the middle, stiffly petaloid or subcoriaceous, glabrous, the lobes 3-4, lanceolate, 2.2-3.0 cm long, ca. 4.5 mm wide at the base, obtuse, coriaceous, glabrous, the margins deeply infolded; sta- mens 4, narrowly oblong, ca. 10 mm long, ca. 1.3 mm wide, obtuse, the filaments slender, dorsifixed, ca. 4 mm long, inserted on the lower ! of the corolla tube; stigmas 2, erect, linear, to 2.2 mm long, the style slender, ca. 22 mm long, ca. 0.35 mm wide, the ovary thick walled. Fruits subglobose, 8-1.5 cm in diam., crowned with a calycine cup, smooth, glabrous, drying black. Faramea luteovirens is known from Panama and Colombia. The label on Duke 10055 indicates that the flowers turn maroon. ''Huesito" Duke 14241. CANAL ZONE: Old Fort Lorenzo, Croat 15411 (MO). Fort Sherman, Dwyer 7180 (MO). Pipeline Road, vicinity of Río Mendoca, Gentry 7469 (MO). Fort Lorenzo, Johnston 1514 (MO). Road to Battery VII, Johnston 1662 (MO). Río Petitpie, Fort Sherman to Gatün Locks, Mori & Kallunki 2701 (MO). Headwaters Río Providencia, Nee & Gentry 8656 (MO). Gatün, Standley 27177 (MO). Fort Sherman, edid pee (US, S); Tyson & Dwyer 1187, 1189 (both MO). Fort San Lorenzo, Tyson 2230 (MO). OLORADO ISLAND: Croat 5217, 6725 (both MO). CHIRIQUÍ: 4.1 mi from Boquete, Road to David. pares 102A (MO). Cano Quebrado, Pittier 6664 (US). COLON: Near home of P. H. Zimmerman, Achiote, ts et al. 3326 (MO). Loma de la Gloria, Fato, Pittier 4038, 4098 (both US). DARIEN: Rio Tuira and Rio Paca, Duke 5024 (MO). Asnati, Duke 10055 (MO). Pinas, Duke ded (MO). Near Estero Grande off Río Maresa, Duke 10954 (MO). Río Perrecénega and Rancho , Duke & Elias 13883 (MO). Mortí Hydro, Río Morti, Duke 14212, 14241 (both MO). La Palma, Pittier 6697 (US). SAN BLAS: Isla Sokatupu, Duke 8956 (MO). Playon Chico, Stier 187 (MO). Í Pino, near Mulatupo, Elias 1701 (GH, MO). Río Mulatupo, headwaters, Elias 1764 (GH, MO). LOMBIA: CHO Municipio de Rio Sucio, Parque Nal. Los Katvos. Camino Tilupo Via Alto aia Leon 536 (MO e 9. Faramea occidentalis (L.) A. Rich., Mém. Soc. Hist. Nat. Paris 5: 176. 1834.— Fig. 38 Ixora occidentalis L., Syst. Nat. x | he TYPE: not seen Tetramerium V ai DC., .4:4 1830. TYPE: not seen. Faramea zeteki Standley, Contr. ir ped 5: 147. 1933. TYPE: Woodworth & Vestal 641 (F, holotype). Trees to 10 m tall, the branchlets terete, smooth, glabrous, ultimately angular. Leaves oblong or elliptic, occasionally ovate oblong, 8-20 cm long, 2.5-11 cm 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 169 38. Faramea occidentalis (L.) A. Rich.—A. Flowering branch (x4). [ Stern et al. FIGURE 24.]—B. Fruit (x4). [After Stimson 5385.] wide, rounded or obtuse at the apex, acuminate, basally cuneate or truncate, the costa subplane to prominulous above, prominulous to prominent beneath, the lateral veins 8-10, widely arcuate, usually with 1—5 irregular and soon evanescent veinlets between adjacent lateral veins, the intervenal areas spreading reticulate, slightly marginate, papyraceous to thin coriaceous, glabrous above, sometimes sparsely pubescent on the veins beneath; petioles stiff, often twisted, to 1.5 cm long, to 0.2 cm wide, glabrate: stipules widely deltoid, 1-2 cm long, 0.5-0.8 cm wide at the base, the terminal awn linear, about the same length or twice the length of the body, glabrous. /nflorescences terminal, occasionally axillary, pa- niculate, spreading, shorter than the terminal leaves, glabrous, the peduncle oc- casionally absent, mostly 1-2.5 cm long, the primary branches usually 2(4), either strongly ascending or the branches strict arcuate, divergent or deflexed, the cy- 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 mules few, sparsely flowered; pedicels 1-2 cm long. Flowers with the hypanthium oblong or rounded, the calycine cup cylindrical, ca. 2.2 mm long, stiffly carnose, glabrous with scattered glands within, these oblong, ca. 0.1 mm in diam., the teeth absent or evanescent; corolla white, the tube narrowly cylindrical, 1.7-2.2 cm long, carnose, glabrous, the lobes 4, lanceolate or ovate lanceolate, 1.1—1.8 cm long, ca. 0.45 cm wide; stamens 4, the anthers barely exserted, narrowly oblong, ca. 8 mm long, 0.8 mm wide, minutely apiculate, versatile, the filaments slender, ca. 2 mm long, inserted ca. 5 mm below the corolla mouth; ovarian disc prominent, ca. 0.5 mm long, the style slender, as long as the corolla tube or less than 4 the length, the stigmas 2, slender, erect, 2.5-5.5 mm long, obtuse, the ovarian wall very thick. Fruits rotund or occasionally compressed rotund, often wider than long, to 1.3 cm in diam., crowned with a cylindrical calycine cup, to 2 mm long, glabrous, drying black. Faramea occidentalis ranges from southern Mexico to Panama; it also occurs in the West Indies and throughout tropical South America. *‘Huesito,’’ Benja- min," “Palo Escrito” (Howell 119) L ZONE: Pina, near Waterways Exp. Sta., Duke 9252 (MO). Gatün, Dwyer 1781 (MO). George Greene Memorial, Las Cruces Trail, Dwyer & Elias 7522 (MO, UC). Boy Scout Road, Madden Dam, Dwyer 11974 (MO). Obispo, Hayes 66 (BM). Pipeline Road, 4 km NW of Ga pes Nee 7076 (MO). Road S-8, Quebrada La Furnia, 1.5 km SW of mouth of Chagres River, Nee 8636 (MO). Río Indio de Gatun, Pittier 2783 (BM, US). jede Forest, Stimson s (MO). Gamboa, Tyson et al. 4571 (MO). Madden Lake, Boy Scout Camp, Tyson 5446 (MO). RO COLORADO ISLAND: Croat 4024A, 4131A, 6310, 6597, 9302, 14570, m 14961, 14992, 17055. (all MO); Dwyer 1407 (MO); Knight 69-36 (MO); i. iip 66-11-2-0905, 66-10-20-0911, 66-8-28-1528, 67-1-17-1004 (all MO); Foster 827 (MO); Robyr -32 (M O); Woodworth & Mes 641 (F); Zetek 3992 (MO). COLON: Río Culebra, Pittier 4164 (U °. COCLÉ: Lower Río Anton, near El Valle, pum 1000 m, Allen 113 (MO). El Valle, Alston 1849 (US). DARIEN: Rio Chico, Allen 464 0 (BM). Rio Bayano above confluence with is Chepo, Duke 3979 (MO). 0-4 mi up Río Sabano, near Santa Fé, Duke 4153 (MO). Río Tuira, 2 upstream from Boca de Cupe, Duke Vide (MO). El Llano, Duke 5519 (MO). Without other locality, Duis 8347 (MO). Río Chucunaque, between Río Membrillo and Río Subcutí, Duke 8610 (MO). Near o Grande off Río Marea, Duke 10958 (MO). Río Corso headwaters off Río Pacora, ca. 500 m, Duke ee (MO). Quebrada Maskia at Rio Pucro, Duke 13099 (MO). Cativo Swamp near Tusi- a, 1 day up Rio Balsa from Manené, Duke 13534 (MO). Tumaganti, ca. 300 m, Duke 14155 (MO). Without De locality, Duke 15787 (MO). 1-3 mi N of Paya, Kirkbride & Duke 14013 (MO, US). Payita, ids rn et al. 248 (MO). Between Río Escucha Ruido & Río Tuira, Stern et al. 576 (MO). 2mi E of Santa Fé, Tyson et fas 4673, 4807 (both MO). Los SANTOS: Loma Prieta, 800-900 m, Duke 11843 (MO). PANAMA: Bayano Cuipo Forest, 34.6 km E of Bayano Bridge, Folsom 3533 (MO). Rio Pasiga, eg sharp bend, Du 2263A (MO). 6 km SW of Arraíjan, old road to Bique, Nee 6901 (MO). El Llano- Carti R oad, 2.3 mi N of Panamerican Highway, ca. 250, Nee & Dwyer 9242 (MO). Ríos ae & Corso a eel apa of Rio Corso, Oliver 2374 (MO, UC, VEN). Juan Diaz, Standley 30581 (MO). 1 Island, Madden Lake, Tyson 5468 (MO) 10. Faramea ovalis Standley, Contr. U.S. Natl. Herb. 18: 138. 1916. TYPE: Pan- ama, Pittier 3159 (US, holotype; C, BM, isotypes). Trees 3-13 m tall, the twiglets slender, the nodes well spaced, glabrous. Leaves small, oval, oblong rotund, or lanceolate, 4-9 cm long, 1.2-4.8 cm wide, rounded, obtuse, or cuneate at the apex, abruptly short acuminate, the acumen 0.3-1.0 cm long, cuneate to obtuse at the base, the lateral veins 6-12, widely arcuate, thin coriaceous, concolorous, glabrous, callose marginate; petioles to 1 cm long; stipules hemispherical, 3-9 mm long, the terminal awn longer than the body. Inflorescences to 5 cm long, to 4 cm wide, few flowered, the pedicels 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 171 umbelliform from the apex of a shorter peduncle. Flowers with the pedicels ca. 1 cm long; calycine cup undulate; corolla purplish or white, tinted pink, the tube ca. 12 mm long, the lobes 4-6 mm long. Fruits not seen. Faramea ovalis is known only from Panama. The small, often oval leaves are unique among the Faramea of Central America. Only one other species of Far- amea in Panama has strongly marginate leaf blades, F. luteovirens Standley. The notes on the type collection indicate ‘‘trunk straight, ca. 6 m, the crown pyram- idal.” Davidson 530, unlike the type, has lanceolate leaves. In addition the notes on the label indicate that it is 30—40 ft in height and has a corolla tube which is white and tinted pink. While these 3 differences from the type suggest that Da- vidson 530 is a second species, I prefer to regard it as F. ovalis. CHIRIQUI: Yo Boquete District, 5500 ft, Davidson 530 (MO). Cerro Horqueta, 1200-1700 m, Pittier 3159 (BM, C, US). ll. Faramea papulata Dwyer & Hayden, Phytologia 15: 57. 1967. TYPE: Panama, Tyson et al. 2218 (MO, holotype).—Fic. 39. Shrubs to 4 m tall, glabrous, the branchlets somewhat plane, the nodes well spaced. Leaves elliptic, 12-14.5 cm long, 5.5-7.5 cm wide, rounded at the apex, acuminate, the acumen 0.3-0.8 cm long, ultimately obtuse, basally rounded or widely cuneate, the costa prominent beneath, the lateral veins 10—12, arcuate, the submarginal vein conspicuous, ca. 0.5 cm distant from the margin, the inter- venal areas prominent, coriaceous, shiny, bullate or papillate, bicolorous, occa- sionally drying yellow; petioles 1—1.5 cm long; stipules connate at the base, di- morphic, some small and compressed rotund, others elliptic, to 0.7 cm long, elongate cuspidate, the cusp to 0.25 cm long, the stipular scars to 0.4 cm long. Inflorescences terminal, umbelliform paniculate, 5.5-9 cm long, to 7 cm wide, the peduncle smooth, to 4.5 cm long, ca. 0.3 cm wide, the primary branches 4, equal, rigid, ascending sharply, the terminal branches ternately disposed, equal, the flowers often ternately disposed on each branchlet; pedicels to 4 mm long. Flowers with pedicels ca. 2 mm long, the hypanthium urceolate, ca. 1.2 mm long, lack when dry; calycine cup crownlike, ca. 2.2 mm long (on immature fruit to 4 mm long), brown, the margin undulate, the teeth evanescent or to 0.1 mm long; corolla white and purple white, drying black, the tube narrowly cylindrical, to 13 mm long, scarcely expanded toward the apex, glabrous, thickly petaloid, the lobes 4, narrowly ovate triangular, to 13 mm long, carnose, glabrous; stamens 4, included, the anthers 4, oblong, to 5.3 mm long, the connective apically produced ] mm, the filaments short, inserted below the middle of the corolla tube; ovules 2, hemispherical, the septum evanescent; style ca. 3 mm long, the stigmas fin- gerlike, ca. 1.8 mm long, scarcely wider than the style; ovarian disc ca. 1 mm long, drying black. Fruits rotund, to 0.5 cm in diam., smooth, glabrous, black when dry. Faramea papulata is known only from the vicinity of Cerro Jefe, Province of Panama. ANAMA: Cerro Jefe, Dwyer & Hayden 4392 (MO). 9.4 km N of Goofy Lake, Folsom et al. 1977 (MO). Cerro Jefe, Mori 6521, 7991, 7998 (all MO); Mori & Kallunki 3630 (MO); Tyson et al. 2218, 3284, 3381 (all MO). 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 12. Faramea pauciflora Dwyer.** TYPE: Panama, Croat 27575 (MO). Trees to 4 m tall, the branchlets angular, twisted, somewhat smooth, the nodes well spaced. Leaves elliptic oblong, 15-19 cm long, 6-7.5 cm wide, rounded or obtuse at the apex, the acumen slender, to 1.5 cm long, ultimately obtuse, cuneate or acute at the base, the costa somewhat prominulous above, subprominent be- neath, the lateral veins 14-17, arcuate, prominulous, forming an undulate mar- ginal vein, rigidly papyraceous, somewhat discolorous, minutely puberulent on the veins and the costa; petioles to 1.5 cm long, ca. 0.13 cm wide, black when dry, minutely puberulent; stipules free, ovate triangular, to 1.5 cm long, 0.4—0.6 cm wide at the base, subcoriaceous, glabrescent, minutely rugulose, the apex drawn into a short bristle. Inflorescences axillary, the flowers 3 per cyme, or the flowers solitary; peduncle to 4 cm long, to 0.15 cm wide, black, glabrous; bracts and bracteoles not seen, perhaps regularly absent; pedicels perpendicular to the axis of the inflorescence or sharply ascending, to 4 cm long, ca. 0.1 cm wide, glabrous. Flowers with hypanthium similar to the pedicel but slightly expanded, glabrous, to 2.5 mm wide, the calycine cup cylindrical, to 1.2 cm long, to 0.5 cm wide, conspicuously truncate, the lobes 4, subulate, to 8 mm long, with 2 longer than the others, acute; corolla white, aromatic, the tube to 3 cm long, ca. 0.25 cm wide in the middle, expanded toward the apex, stiffly petaloid, glabrous on both sides, the lobes 4, these erect, narrowly subulate, to 1.3 cm long, to 0.35 cm wide, glabrous; stamens 5, sessile, narrowly oblong, ca. 8 mm long, obtuse at the apex, sagittate at base, inserted above the middle of the corolla tube; style not seen. Fruits not seen. Faramea pauciflora is known only from Panama. It is readily distinguished by its few-flowered inflorescences, its elongate calycine lobes and corolla tube, and its elongate anthers. VERAGUAS: Valley Rio Dos Bocas, Road toward Escuela Agricola Alto Piedra and Calovébora, 450-550 m, Croat 27575 (MO). 13. Faramea scalaris Standley, Contr. U.S. Natl. Herb. 13: 139. 1916. TYPE: Panama, Pittier 3265 (US, holotype; isotypes F, NY). Shrubs 4-6 ft tall, the branchlets smooth, glabrous, the nodes well spaced. Leaves sessile or subsessile, glabrous, narrowly elliptic, 5.5—15 cm long, 1.3-3.5 cm wide, tapering at the apex, acuminate, the acumen falcate, to 1.5 cm long, usually ultimately obtuse, basally cuneate to obtuse, the costa prominulous above and beneath, the lateral veins 8-13, perpendicular, scarcely arcuate, the inner submarginal vein more or less parallel to the margin, to 4 (-7) mm from the 48 Faramea pauc Vene Dwyer, spec. nov. Arbores ad 4 m altae. Folia elliptico-oblonga, 15—19 cm , 6-7.5 cm lata, apice rotunda vel obtusa, acumine ad 1.5 cm longo, basi cuneata, venis lateralibus 14-17 rigido- papyracea in venis costaque minuto-puberula; petiolis ad 1.5 cm longis; sti- floribus 3 in cymam unam dispositis, pedunculo ad 4 cm longo. Flores pedicellis ad 4 cm longis; cupula calycis ad 1.2 cm longa, truncata dentibus 4 subulatis, ad 8 mm longis, inaequalibus; corolla alba, tubo ad 3 cm longo, lobis 4 ad 1.3 cm longis; antheris e angusto-oblongis, ca. 8 mm longis. Fructus non visi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 173 FiGURE 39. Faramea papulata Dwyer & Hayden.—A. Habit (<'2).—B. Flower (x1.5).—C. Flower opened to show interior (x 1.5).—D. Fruit ( x2). [After Mori 7998.] margin, prominulous beneath, the outer submarginal vein close to the margin, less prominulous and conspicuous than the inner submarginal vein, with 1—3 smaller veins paralleling the adjacent lateral veins, these soon branching, stiffly chartaceous, concolorous, the margin revolute; stipules persistent, free, rectan- gular or deltoid, to 0.5 cm long with a short median awn arising from a median keel, stiffly chartaceous, glabrous. /nflorescences terminal, the peduncle 3-4 cm long, ca. 0.15 cm wide, flowers ca. 15, aggregated into a head subtended by 2 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 ovate rotund bracts, to 1.5 cm long, the terminal awn about as long as the body. Flowers subsessile; hypanthium and calycine cup ca. 2 mm long, the teeth tri- angular, 0.2-0.4 mm long; corolla bluish white, the tube narrowly cylindrical, 1— 1.2 em long, ca. 0.1 cm wide, the lobes 4, 0.3-0.5 mm long. Fruit not known. Faramea scalaris is known only from high elevations in the Province of Chi- riquí. It may be conspecific with the well-collected F. suerrensis (Donnell Smith) Donnell Smith. Unfortunately the fruit of F. scalaris is not known. HiRIQUÍ: Bajo Chorro, near Boquete, 6000 ft, eie d 94 (US), 430 (MO), 436 (US). Between Alto de las Palmas and Cerro Horqueta, Pittier 3265 (F, NY, US). 14. Faramea suerrensis (Donnell Smith) Donnell Smith, Bot. Gaz. 44: 112. 1907. F. iie K. Schum. & Donnell Smith var. suerrensis Donnell Smith, Bot. Gaz. 31: 115. 1905. : Costa Rica, Donnell Smith 6589, not n. Trees to 15 ft tall, the branches stout terete, ultimately angular, glabrous, the nodes well spaced. Leaves oblong, ovate oblong or elliptic, to 14(30—) cm long, to 7(10.5—) cm wide, usually 244 times longer than wide, acute to obtuse at the apex, cuspidate, the cusp often falcate, acute, obtuse or truncate, to 2 cm long, occasionally vaguely auriculate at the base, the costa prominulous above, prom- inent beneath, the lateral veins 13-15, usually with a smaller vein intermediate between 2 laterals paralleling these, at first strict, then arcuate and joining a prominent marginal vein, 0.5—0.7(1.0—) cm within the margin, usually with a sec- ond less prominent vein ca. 2 mm from margin the intervenal areas spreading reticulate, occasionally bullate, chartaceous to subcoriaceous, concolorous, gla- brous; petioles absent or to 2 cm long, glabrous; stipules connate, cylindrical, oblong rotund, oblong rectangular to hemispherical, 6-7 mm long, 5-7 mm wide, minutely mucronate or retuse, glabrous. Inflorescences terminal, cymose cor- ymbose, glabrous, to 9 cm long, sometimes epedunculate with 2 branches arising from the tip of the twiglet, sometimes long pedunculate, the peduncle to 5 cm long and the secondary branches ascending and flabellate or penicillate. Flowers with the pedicels slender, to 2.5 mm long; hypanthium oblong rotund, ca. 1.2 mm long, glabrous, the calycine cup rimlike ca. 0.3 mm long, glabrous but with a few glandular spots within, the teeth minute, to 0.3 mm long; corolla white (in bud), blue or purple, the tube narrow, to 6 mm long, ca. 1.2 mm wide, petaloid, gla- brous; stamens 4, the anthers oblong, ca. 2.2 mm long, ca. 0.4 mm wide, minutely apiculate, the filaments attached near the middle of the tube: ovarian disc oblong rotund, ca. 1 mm long, the style short, the ovules ca. 0:3 mm in diam. Fruits reniform or compressed oblong, to 0.7 cm long, to 1.5 cm wide, about twice as wide as long, apiculate, smooth, often delicately ribbed, farinose or minutely rugulose. Faramea suerrensis is known from Costa Rica, Panama, and Colombia (Cho- co). The fruit of Duke & Lallathin 14977 (MO) is reported as blue. OCAS DEL TORO: Punta Pena, vicinity of Chiriquicito, Lewis et al. 2145 (MO). Chiriquí Lagoon, Wedel 1430 (F). Water Valley, Wedel 1511 (MO, US), 1610 (MO). CHIRIQUÍ: d in road to Cerro Colorado and Escopeta above Río San Félix near town of San Félix, ca. 13 mi N of Río San Félix 800-1200 m, Croat 33428 (MO). cocLé: Cerro Pilón, Duke & Lallathin 14977 aes El Cope, El 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 175 Potroso, Atlantic slope of Alto Calvario, Folsom & Lantz 1887 (MO). Cerro Pilón, Lallathin 5093 MO). coLON: Camp Bejita, District Donoso, Holdridge 6251 (MO). Santa Rita Ridge, Mori & Kal- lunki 4919 (MO); Mori & Crosby 6425 (MO). VERAGUAS: 6.4 km outside Santa Fé, road to Agricultural School, Folsom 2965 (MO). Cerro Tute, Folsom & Edwards 3336 (MO). COLOMBIA: CHOCO: Alto de Buey, 500-1200 m, Gentry & Forero 7287 (MO). 15. Faramea talamancarum Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 332. 1929. TYPE: Costa Rica. Carleton 135 (F, holotype, US). Trees to 15 ft tall, the branches slender, terete, smooth, glabrous, the nodes well spaced. Leaves oblong to obovate oblong, to 17 cm long, to 6 cm wide, cuspidate, the cusp to 1.5 cm long, erect or falcate, to 0.25 cm wide, obtuse, cuneate or acute at the base, scarcely inequilateral, the costa slender, prominu- lous above and beneath, the secondary veins 10-15, almost perpendicular, the submarginal vein not elevated or impressed, membranous to chartaceous, con- colorous; petioles to 1 cm long, glabrous; stipules persistent, connate, the sheath cylindrical, 2-5 mm long, the awn 4-5 mm long. Inflorescences terminal, solitary, paniculate, cymose corymbose, 8-10 cm long, glabrous, sometimes the peduncle absent and usually 3 branches arising from the stem tip, sometimes 2-3 cm long, ca. 0.15 cm wide, the branches spreading or strongly ascending, in 2 or more pairs, opposite, 2-4 cm long. Flowers with the pedicels slender, 8-10 mm long; hypanthium compressed rotund, to 0.7 mm long, carnose, glabrous, the calycine cup inflated, glabrous, ca. 0.6 mm long, with several rubescent glandular spots within, the teeth 4, deltoid, to 0.3 mm long, acute; corolla blue, the tube narrow cylindrical, expanding apically, 10-17 mm long, the lobes oblong, to 6.5 mm long; stamens 4, the anthers narrowly oblong, 2.8—4 mm long, 0.6-0.8 mm wide, obtuse, cordate at the base, the filaments dorsifixed, to 1 mm long, attached above the middle of the tube; ovarian disc coroniform, to 0.35 mm long, the stigmas 2, upright, scarcely thicker than the style, 1-1.5 mm long, the style 1.8-3 mm long, .2 mm wide, the ovules about as long as broad. Fruits not seen. Faramea talamancarum is known from Costa Rica and Panama. OCAS DEL TORO: Río Teribe, between Quebrada Huron & Quebrada Schlunyik, Kirkbride & Duke 496 (MO). Vo ge d Peak, N Río Teribe, lg n & Duke 584 (MO). Fish Creek Mts, Wedel 2191 (F, MO), 2243 (MO, US). CoLON: Santa Rita Ridge, Dwyer 8555 (MO). DARIEN: Airstrip, Caná gold mine, 480 m, Croat 37968 (MO). Paya, coe et al. 230 (MO, US). Cana, Stern et al. 656 O). 16. Faramea terreyeae Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 189. 1940. TYPE: Panamá, Terry & Terry 1479 (F, holotype, isotype MO). Shrubs to 8 ft tall, the branchlets terete, smooth, glabrous, the nodes well spaced. Leaves lanceolate or elliptic to 13 cm long, to 3.6 cm wide, at least 3 times longer than wide, often falcate, acute at the apex, cuspidate or caudate acuminate for ca. 1.5 cm, ca. 1.5 mm wide in the middle, prominulous above, subplane and grooved beneath, the lateral veins ca. 20, strict, joining a delicate and vaguely undulate vein ca. 0.2 cm from the margin, but without a distinctly elevated submarginal vein, papyraceous, discolorous; petioles to 0.5 cm long, slender, ca. 0.8 mm wide, glabrous; stipules connate, cylindrical, to 3 cm long, ca. 0.5 cm wide, the awn ca. 1 cm long. Inflorescences terminal, to 2.5 cm long, 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 the peduncle ca. 1 cm long, the lowermost bracts folded, ovate rotund or com- pressed rotund, to 2 cm long, with a slender awn arising from a slender keel-like thickening, papyraceous, glabrous. Flowers not clearly seen (cf. text below). Fruits not known. Faramea terreyeae is known only from the type collection. It is closely related to F. jefensis Dwyer which has a very reduced inflorescence subtended by 2 large leaf-like bracts. F. terreyeae has papyraceous leaves whereas those of F. jefensis are thin coriaceous; the hypanthium and the calyx of F. terreyeae are about 2 mm long, while those of F. jefensis are about 6 mm long. Unfortunately the flowers of F. terreyeae are poorly known. Standley described the inflorescence as umbellate and capituliform; he noted that in addition to the 2 outermost large bracts there are 2 much shorter inner bracts. His description of the floral parts is meagre (translated from Latin): `“ Flowers short-pedicellate; hypanthium widely obovoid, 1.5 mm long, the calyx ring shaped, ca. 0.6 mm long, truncate, remotely and minutely denticulate.”’ DARIEN: Cana-Cuasi Trail, Chepigana, 3000 ft, Terry & Terry 1479 (F, MO). 17. Faramea woodsonii Standley. Ann. Missouri Bot. Gard. 28: 470. 1921. TYPE: Panama, Woodson & Schery 164 (MO, holotype). Shrubs, the branchlets slender, terete, smooth, glabrous, the nodes well spaced. Leaves oblong, 3-7 cm long, 1-2.6 cm wide, tapering obtusely or oc- casionally acutely toward the apex, acuminate, the acumen 0.4-1 cm long, obtuse, cuneate to occasionally obtuse at the base, the costa slender, prominulous above and beneath, the lateral veins ca. 9, at first strict, then arcuate, forming an un- dulate vein 1-1.5 mm from the margin, 1—2 smaller, irregular, and soon branching veins arising between the lateral veins, the other conspicuous submarginal vein delicate, ca. 0.35 mm from the margin, the margin delicately revolute, the inter- venal areas spreading reticulate, stiffly chartaceous, concolorous, drying gray or green, glabrous; petioles to 0.4 cm long; stipules connate, the sheath not deeply incised laterally, ovate rotund, to 4 mm long, glabrous, the median awn of each part almost as long as the sheath, extending along the body as a median keel. Inflorescences terminal, the flowers ca. 4, spread radially, the peduncle obsolete. Flowers to 2 cm long; hypanthium oblong, ca. 1 mm long, glabrous, the calycine cup ca. 0.5 mm long, with minute glands within, the teeth minute, disposed as 5 points; corolla blue, the tube narrow cylindrical, to 11.5 mm long, ca. 1.3 mm wide below, ca. 3 mm wide near the mouth, petaloid, glabrous, the lobes 4, ovate oblong, to 7 mm long; stamens 4, the anthers subsessile, oblong, 3-4 mm long, attached at the middle of the tube; style subulate, 1.5-3 mm long, the stigmas 2, minute, ca. 0.5 mm long, the hypanthium angular in cross section. Fruits on slender pedicels longer than the pericarp, the latter compressed rotund, to 0.8 cm long, smooth, glabrous, the persistent calyx ca. 0.5 mm long. Faramea woodsonii is known only from Panama. CocLÉ: El Valle, Allen 3544 (MO); Duke 12152 (MO); Duke & Dwyer 13914 (MO); Dwyer 11868 MO); Woodson & Schery 164 (MO). PANAMA: El Llano-Cartí Road, 16-1872 km N of Panamerican Highway, 400—450, Nee & Tvson 10940 (MO). 177 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae ~ FIGURE 40. Ferdinandusa panamensis Standley & Williams. Habit (x14). [After Wedel 2250. ] 38. FERDINANDUSA Ferdinandusa Pohl, Pl. Bras. Icon. Decr. 2: 8. 1828-1829. Substitute name for Ferdinandea Pohl, 1827. rype: F. speciosa Pohl. Ferdinandea Pohl, Flora 10: 153. 1827, not Ferdinanda Lagasca, 1816. TYPE: No type indicated. Aspidanthera Benth. in Hook., J. Bot. 3: 217. 1841. TYPE: A A. rudgeoides Benth. Gomphosia Wedd., Ann. Sci. Nat. Bot. 3: 10: 14. 1848. rvPE: No type indicated. Trees or shrubs. Leaves with stipules interpetiolar. /nflorescences terminal, cymose paniculate or corymbose paniculate. Flowers 4—-5-merous, the calycine cup cupuliform, the teeth reduced; corolla variable in color, infundibuliform or hypocrateriform, the lobes contorted in the bud, obliquely 2-lobed, emarginate: stamens ca. 4, the anthers exserted or included, often variable in size, the fila- ments often variable in length; ovary 2-celled, the ovules several to numerous in 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 each locule. Fruits capsular, cylindrical to rotund, loculicidally dehiscent, the seeds few to numerous, peltately affixed to an axile placenta, winged, the wing occasionally laciniate. Ferdinandusa, a genus of about 20 species, is found in Panama and in South America. 1. Ferdinandusa panamensis Standley & Williams, Ceiba 3: 34. 1952. TYPE: Pan- ama, Wedel 2232 (F, holotype; isotype, MO).—Fic. 40. Trees to 60 ft tall, the branchlets terete, smooth, villose, minutely resinous pustulate (bases of deciduous hairs), the nodes well spaced. Leaves oblong, oc- casionally falcate, 10-17 cm long, 6.0—9.5 cm wide, obtuse and short acuminate, the acumen to | cm long, obtuse, occasionally truncate, the costa slender, prom- inulous above, prominent beneath, the lateral veins 12-15, prominulous, arcuate, stiffly papyraceous, glabrous above and beneath, the intervenal areas reticulate; petioles to 1 cm long, stout, villose, the lamina decurrent on the upper side; stipules deciduous, deltoid lanceolate, ca. 1 cm long, acuminate, glabrous. /nflo- rescences terminal, longer than the uppermost leaves, the peduncle shorter than the uppermost leaves, glabrous with 2 branches arising at the tip, the flowers several, ascending. Flowers with the hypanthium short, subrotund, ca. 2.5 mm in diam., glabrous, the calycine teeth reduced to 4—5 fleshy points; corolla white, the tube cylindrical, expanded at the apex and at the base, 1.5-2.3 cm long, 0.14 cm wide in the middle, fleshy to subcoriaceous, glabrous, the lobes 5, imbricate, oblong, ca. 0.8 mm long, not cucculate; stamens 5, the anthers dimorphic, usually 2 long and 2 short, dorsifixed, the thecae of some anthers dimorphic, oblong to rotund, the filaments linear, to 7 times the length of the anthers, attached below the middle of the tube; ovarian disc prominent, the style subulate to 11 mm long, the stigmas 2, ca. 1.5 mm long, wider than the style, oblong rotund. Fruits not seen. Ferdinandusa panamensis is known only from Panama. BOCAS DEL TORO: Fish Creek, Chiriquí Lagoon, Wedel 2232 (F, MO), 2250 (MO). 39. GALIUM Galium L., Sp. Pl. 105. 1753. Gen. PI., ed. 5. 46. 1754. TYPE: Galium mollu- goL Erect or decumbent /ierbs or vines, the stems often 4-angled, slender. Leaves 3-4 or more at a node, verticillate; stipules foliaceous. Inflorescences terminal or axillary, in dichasial panicles, the flowers not subtended by an involucre. Flowers bisexual, unisexual or polygamous dioecious, pedicellate; hypanthium articulate with the pedicel; calycine cup ovoid or globose, the lobes reduced or absent; corolla white, yellow or green, the lobes 3-4, valvate in the bud; stamens 2-4, exserted, the styles 2, the stigmas capitate; ovary 2-celled, the ovules soli- tary. Fruits didymous, fleshy or dry, separating finally into closed segments or only one carpel maturing; seeds concave. Galium is a genus of about 400 species. It is well distributed throughout the world, especially in the temperate climates. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 179 Literature: Dempster, L. T. 1973. The Polygamous Species of the Genus Galium (Rubi- aceae) Section Lophogalium of Mexico and Southwestern United States. U. Calif. Publ. Bot. 64: 1-36. . 1978. The e bc (Rubiaceae) in Mexico and Central America. U. Calif. Publ. Bot. 7 Greenman, J. M. 1898. | of the Mexican and Central American Species of Galium and Relbunium. Contr. Gray Herb. n.s. 14: 455-457. a. Leaves 6-8(-10) at a node ---------------------------------------------------------- 2. G. mexicanum aa. Leaves 4 at a node. . Leaves with lateral veins evident; fruit with n uncinate hairs ____ à G. uncinulatum bb. Leaves with lateral veins not evident; fruit glabro . G. aschenbornii 1. Galium aschenbornii Schauer, Linnaea 20: 701. 1847. TYPE: México, Aschen- borne 92 (JE, not seen). i acuminata Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 11: 127. 1844. Type: México, Galeotti 362 (K, holotype). Pus gemniflorum Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 11: 126. 1844. Type: México, Galeotti 2604 (BR, holotype; isotype, K, not seen). G. galeottianum Walp., Repert. Bot. Syst. 6: 17. a TYPE: not s G. uropetalum Hemsl., Biol. Centr. Amer., Bot. 2: 66. 1881. TYPE: gd» seen. Vines, the stems angular, obtusely winged, glandular shiny, glabrescent, often with a few regularly arranged, retrorse, stumpy hairs. Leaves sessile, 4 per node, ovate oblong, to 5 mm long, acute, the lateral veins not evident or scarcely visible, subcoriaceous, glandular shiny above and presumably beneath, with sev- eral golden brown glands beneath. Inflorescences terminal, 1-2-flowered, to 1 cm long. Flowers not seen. Fruits subrotund, to 4 mm in diam., glabrous, drying black, obviously pedicellate. Galium aschenbornii occurs in Mexico and Central America. Dempster pro- vides an excellent description of the flowers and fruits (1978) based on a study throughout the range of the species: "flowers few, or sometimes many, | to several in imperfect, perfect, or compound dichasia borne on short leafy deter- minate branches; flowers staminate, pistillate, or perfect, on the same plant, or sometimes on different plants; corollas rotate, glabrous, usually red or pink, but sometimes yellow, white, or greenish, the lobes much prolonged at the apex; fruits glabrous, fleshy, black and wrinkled when mature, 3-7 mm across when CHIRIQUí: Volcán de Chiriquí, ca. 7000 ft, Davidson 882 (MO, left-hand side of sheet). Chiriquí Volcano, Killip 3582 (US). 2. Galium mexicanum H.B.K., Nov. Gen. Sp. Pl. 3: 337. 1818. NEoTYPE (Demps- ter, 1978): México, Solbrig & Ornduff 4540 (UC). Original Humboldt & Bon- pland type apparently lost.—Fic. 41. g . mexicanum var. glabratum Greenm., Contr. Gray Herb., 14: 458. 1898. TYPE: México, Nelson 83 (US e . mexicanum var. platyphyllum Greenm., Contr. Gray Herb. 14: 458. 1898. TYPE: México. Nelson 5 (US). 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 41. Galium mexicanum H.B.K. Habit (x24). [After White 206. | Vines, the stems weak, stramineous, angular, branching profusely, the hairs stiff, minute, reflexed, uncinate, occasionally weak and spreading, the nodes well spaced. Leaves sessile or subsessile, usually 6—8(-10) at a node, occasionally 4, linear oblong or narrowly obovate oblong, 0.4-2.0 cm long, 1.3-2.0 mm wide, rounded or acute at the apex, tapering narrowly at the base, the costa prominulous above and beneath, the hairs well spaced, uncinate, rigid, the margin often re- volute and with stiff, uncinate hairs, the blade glandular shiny, pubescent be- neath, the hairs with bulbous bases, often deciduous, the blade then granulose. Inflorescences cymose paniculate, to 2 cm long (? longer). Flowers minute, 0.6— 1.5 mm long; hypanthium subrotund, ca. 0.35 in diam., pubescent; calycine cup and lobes absent; corolla white, greenish white, occasionally purple or red, the tube scarcely measurable or to 0.2 mm long, the lobes 4, (2) ovate oblong, 0.5— 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 181 0.7 mm long, the tip acuminate, often deflexed adaxially, the acumen to % the length of the body of the lobe, petaloid, dorsally pubescent; stamens 4, the anthers oblong, ca. 0.2 mm long, the filaments 0.2-0.5 mm long, attached at the junction of the corolla lobes; style short, the stigmas terminal, capitate. Fruits compressed rotund, less than 1 mm long, wider than long at maturity, the hairs dense, thick, falcate, ca. 0.1 mm long. Galium mexicanum ranges from Mexico to Panama. CHIRIQUÍ: Volcan Barú, E slope, 1 mi from summit, ca. 3000 m, Croat 34905 (MO). El Baru, above Boquete, 1200-1800 m, D'Arcy 99/9 (MO). Near Boquete, D'Arcy & D'Arcy 6463 (MO). E slope of Volcan de Chiriquí, WNW of Boquete, Davidse & D'Arcy 10187 (MO). 11 km WNW of Boquete, Davidse & D'Arcy 10224 (MO). Bajo Mono, Boquete, Davidson 595 (MO). Volcán de Chiriquí, 8000 ft, Davidson 935 (MO). W side of Volcán Barú, oak forest, 2000-2500 m, Mori & Bolten 7355 (MO). Valley of upper Río Chiriquí Viejo, Monte Lirio, 1300-1900 m, Seibert 276 (MO). mi N el El Volcán, ca. 5000 ft, Tyson 5721 (MO). Volcán de Chiriqui, White 206 (MO, US). Finca Lerida to Peña Blanca, 1750-2000 m, Woodson & Schery 334 (MO). Potrero Muleto to summit, Volcán de Chiriquí, 3500-4000 m, Woodson & Schery 443 (MO). 3. Galium uncinulatum DC., Prodr. 4: 600. 1830. TYPE: not seen. G. obstipum Schlecht., Linnaea 9: 592. 1834. TYPE: Mexico, Schiede 248 (HAL, not seen, isotypes MO, NY; P, W, neither seen). G. nelsonii Greenm., Proc. Amer. Acad. Arts 33: 460. 1898. Type: México, Nelson 628 (US). Herbs, the stems slender, weak, 4-angled, much branched, hispid, Leaves 4-nate, subsessile, elliptic oblong, or ovate oblong, 1.5-2.3 cm long, 0.5-1.3 cm wide, rounded, obtuse or deltoid at the apex, cuneate or rounded at the base, the apiculum minute but obvious, the lateral veins 2, basal and arcuately ascending, usually with 2 other more delicate veins arising along the costa, membranous, hispid on both sides; stipules not seen. Inflorescences terminal or axillary, di- chotomously corymbose, to 7 cm long; peduncle to 4 cm long, with 2 or more frequently foliose bracts to 1⁄2 the length of the regular leaves, subtending a pair of branches, the flowers few. Flowers pedicellate, the pedicels 0.1-1.0 cm long; hypanthium rotund, ca. 0.7 mm in diam., the hairs abundant, strict, white, stiff, terminally uncinate, to 0.7 mm long, the calycine cup and teeth not visible; corolla greenish yellow or yellow, the tube short, the lobes 4, ovate or oblong, ca. 1 mm long, acute, pubescent inside and out. Fruits rotund or compressed rotund, didy- mous, the body 1-2 mm wide, the hairs dense, often obscuring the body, radiate, tawny, usually hooked at the apex. Galium uncinulatum ranges from Arizona, Texas, and Mexico to Central America. 1300-1900 m, Seibert 205 (MO). Rio Chiriqui Viejo Valley, near El Volcan, White 184 (MO). Finca Lerida, 1750 m, Woodson & Schery 219 (MO). 40. GENIPA Genipa L., Gen. Pl. ed. 5. 87. 1754. LECTOTYPE: G. americana E. Trees. Leaves opposite, coriaceous, petiolate; stipules interpetiolar, often per- sistent, triangular, connate basally. Inflorescence of a single axillary or terminal 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cyme, the flowers several, occasionally solitary. Flowers large, bisexual, 5—6- merous; calyx campanulate or turbinate, truncate, the teeth absent or of 5-6 short lobes; corolla short cylindrical, the lobes 5—6, contorted in bud, often longer than the tube; stamens 5-6, the anthers usually sessile, inserted at the mouth, dorsi- xed, exserted; ovary 2-celled, the ovules numerous, usually disposed horizon- tally. Fruits baccate, usually globose, smooth, leathery, the wall thick, the seeds numerous, large, usually trapeziform, disposed in several rows in a fleshy pulp. Genipa is a genus of about 7 species, occurring in tropical Florida, Mexico, Central America and in South America. 1. G. americana a. Leaves drying black; corolla pubescent outside aa. Leaves drying brown; corolla glabrous outside. b. eaves lanceolate, 3. 7-5.2 cm wide, deltoid at the apex; corolla lobes ca. 4.5 m l G. curv nm wide; fruits ca. 3 cm in diam... bb. Leaves obovate rotund, obovate or oblong, 3.5-10.0 cm wide, rounded or obtuse at the apex; corolla lobes ca. 1 cm wide; fruits 3.5—5.0(-7.0) cm in diam. _____- . G. williamsii l. Genipa americana L., Syst. Nat., ed. 10. 2: 931. 1759. TYPE: not seen— Fic. 42 Gardenia genipa Swartz, Prodr. Veg. Ind. Occ. 51. 1788. rYPE: not s Genipa oblongifolia Ruiz & des Fl. Peruv. Chil. 2: she 1798. TYPE: oe seen. G. grandifolia Pers., Syn. Pl. 1: 198. 1805. TYPE: not s G. caruto H.B.K., Nov. Gen, Sp. Pl. 3: 407. 1820. TYPE: : Venezuela and Colombia, Humboldt & Bonpland (P, not s G. humilis Vell., Fl. Flum. 94. Ic. 2: pl. 142. 1825. TYPE: not G. pubescens DC., Prodr. 4: 379. 1830. TYPE: 2 de la Osa pen seen. G. barbata Presl., Symb. Bot. Upsal. 2: 13. 1 TYPE: Brazil, Blanchet, not see Genipa americana var. caruto (H.B.K.) ee nti in Mart., FI. in . 6(6): 352. 1889. G. excelsa Krause, Bot. Jahrb. Syst. 40: 327. 1908. TYPE: not seen. G. americana var. caruto fo. grandifolia Chod. & Hassl., Bull. "Herb. Boissier ser. 2, 4: 171. 1904 TYPE: Paraguay, Hassler 7851 (G. isotype). G. excelsa Krause, Bot. Jahrb. Syst. 40: 327. 1908. TYPE: not seen. Trees 20 m tall, the twigs terete, ultimately angular, the bark smooth, glabrous, the nodes well spaced or crowded terminally. Leaves usually ~ ascending, elliptic, elliptic oblong or obovate oblong, 11-28 cm long, 6-13 cm wide, deltoid or somewhat rounded toward the apex, acuminate, the acumen e. attenuate toward the base, the costa prominulous above, prominent beneath, the lateral veins 11-20, arcuate ascending, thin coriaceous to coriaceous, shiny and glabrous above, often drying black, glabrous to velutinous pubescent beneath, the hairs dense but short; petioles stout, to 1 cm long; stipules free, ovate, to 1.5 cm long, acute, coriaceous to thickly scarious, glabrous to puberulent toward the scarious margins, the margins thickened, pubescent, drawn into a subulate process. Inflo- rescences terminal, cymose, 4-10 cm long or much longer; peduncle short, the bracts ovate, 3-4 mm long, acuminate, the branches to 9, few or many flowered. Flowers to 3.5 cm long, bisexual on pedicels to 1.2 cm long; hypanthium narrowly ae calycine cup turgid, ca. 0.6 cm long, stiffly carnose, the margin truncate, teeth as mere points, glabrous except often with a few hairs marginally; 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 183 FIGURE 42. Genipa americana L.—A. Flowering branch (x .45). [After Blum & Dwyer 2477 .\— B. Flower (x2.7).—C. Ovary in cross section (x2.7). [After Bartlett & Lasser 16465.]—D. Fruit (x.45). [After Croat 7913. ] corolla cream white, the tube short cylindrical, ca. 1.5 cm long, 0.8-1.0 cm wide, densely velutinous hairy or villose outside and in, the hairs to 2 mm long, the lobes 5-6, oblong, 1.5-2.5(-3.0) cm long, longer than the tube, widely deltoid at the apex, reflexed at anthesis; anthers 5, subsessile, narrow oblong, 8.0-9.5 mm long, the connective produced for ca. 1.5 mm at the apex, ultimately acute or obtuse, the filaments attached near the mouth; stigmatic lobes 2, narrowly lan- ceolate, erect, ca. 7 mm long, partially exserted. Fruits oblong or subrotund, to 9 cm in diam., glabrous, silvery gray outside, the persistent calyx short, giving a craterlike effect, ca. 1 cm in diam. 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Genipa americana is found in tropical Florida, Mexico, Central America, West Indies, and tropical South America, extending to Paraguay. Duke (1968) states that according to the Cuna: ‘‘Pulp of the fruits is edible raw; it is used to make sweet and fermented beverages; sometimes pickled with vinegar and onions. The flowers are attractive to bees and the fruits to cattle. Indians paint their bodies black with the pulp; some OICS personnel claimed it does improve the complexion and state that it is insecticidal. It is the ‘Tree of Life’ to the Cuna. ‘Apples of Immortality’ arise from the Earthmother’s placenta and umbilical cord; the juice turns the skin black, thus hiding the individual from the devils that cause disease. It appears to be bacteriocidal, germicidal, and insect repellent. It is said to be the only medicine capable of flushing the parasitic catfish which attacks man south of the study areas. The Cuna have a special song of the jagua (Genipa americana) known as ‘quanekigala.’’’ In Panama the most common name is *'Jagua"'; other common names are ‘‘Guaytil Blanco"; "Guaitil Blanco"; Jagua Amarilla"; ‘Jagua Blanca"; ''Jagua Colorado"; and ‘‘Jagua de Mon- tana." It is used throughout its range for construction of buildings, furniture and carts. Julian A. Steyermark (The Botany of the Guayana Highland, Mem. N.Y. Bot. Garden 23: 349. 1972) discusses in detail the typification of Genipa americana. CANAL ZONE: Cerro Viejo, K 16 C, Blum 1265 (MO). Fort Kobbe, Duke 3943, 4189, 4259 (all MO). Farfan Beach, Duke 11734 (MO). Río Mar, Duke 12427 (MO). K-9 Road, Ebinger 516 (MO). Mandingo River, Hayden 57 (MO). Arraijan, Hayden 112 (MO). Albrook, U.S. Army Tropic Test site, Hayden 134A (MO). Survival School, Fort Clayton, Kirkbride & Elias 268 (MO), Howard Air MO, US), É i H ; I M Seibert 409 (MO). Cerro Luisa, vicinity of Pedro Miguel Locks, Stern & Chambers 8 (MO). Survival School, Fort Clayton, Tyson 1051 (MO). BARRO COLORADO ISLAND, Croat 7913 (MO). Cardenal Creek, Fort Clayton, Tyson 1279 (MO). Miraflores, Mee 126 (MO). BARRO COLORADO ISLAND: Robyns 65- 33 (MO); Shattuck 415 (MO). CHIRIQUÍ: 12.4 mi N of David, us et al. 728 (GH, MO). cocrÉ: Río Hato Airstrip, Blum & Dwyer 2477 (MO). PROCU. Dwyer 2002 ier DARIEN: Rio Ucurganti, Bristan 1185 (MO). Isla Casaya, Duke 10390 (MO). HERRERA: Guararé, Rodriguez 26 (MO). Los SANTOS: 4 mi S of Limón, Correra & Galegos 66 (MO). Punta ey Dum 9755 (MO). Candelaria, Duke 12453 (MO). Monagre Beach, Tyson et al. 3021 (MO). PANAMÁ: Río Pacora, Bartlett & Lasser 16465 (MO). Cerro Campana, Castillo 2 (MO). Río Hi Duke 4763 (MO). Farfan Beach, Duke 11734 (MO). Río Mar, Duke 12427 (MO). La Bayoneta, Perlas Archipelago, Dwyer 1733 (MO). Tocumen, Dwyer 2894, 4347 (both MO). San José Island, Johnston 1358 (MO). Palo Seco, Mejia 20 (MO). Isla del Espiritu Santo, Tyson 5558 (MO). Isla Taboga, Woodson et al. 1480 (MO). vERAGUAS: Santiago, Dwyer & Kirkbride 7427 (COL, DUKE, MO, UC), 7429 (GH, MO) 2. Genipa curviflora Dwyer.*® Type: Panamá, Nee & Warmbrodt 10347 (MO, holotype). Shrubs or small trees to 4 m, the branchlets angular, smooth, minutely pu- berulent, the nodes turgid, often well spaced. Leaves lanceolate, 11.5—20.0 cm long, 3.7-6.8 cm wide, deltoid toward the apex, the acumen short, to 1 cm long, ultimately obtuse, cuneate toward the base, the costa prominulous and often 49 Genipa curv idet: Dwyer, spec. nov. Frutices vel be ramulis puberulis. Folia lanceolata, 5-20.0 cm lon 7—6.8 cm lata, venis lateralibus 7-12, tenui-coriacea supra glabra subtus = ie petiolis a 3. " cm longis; stipulis ovato- Meu ie aA Inflorescentiae terminales ad 4 c ongae, floribus ca. 7. Flores pedicellati i calycis ad 0.5 mm longa, coriacea glabra remisi nullis; corolla tubo angusto-cylindrico, ca. 1.5 cm longo, falcato, glabro lobis 4 carnosis basi unila- Pea auriculatis; antheris sessilibus lineari- Ie ca. 8.5 mm longis. Fructus subsessiles rotun- di, ca. 3 cm in diam., laeves glabri 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 185 sulcate above, subprominent beneath, the lateral veins 7-12, at first stiff, then arcuate toward the margins, uniting to form an undulate submarginal vein ca. 5 mm distant from the margin, prominulous beneath, the intervenal areas spreading, conspicuously thin coriaceous, concolorous, glabrous above, minutely puberulent beneath; petioles to 2.5 cm long, ca. 0.2 cm wide, lignose; stipules free, often deciduous, ovate trapeziform, to 1.7 mm long, acute, at first scarious then some- what ligneous, puberulent. /nflorescences terminal, to 4 cm long; peduncle absent or to 0.5 cm long, the flowers ca. 7, perhaps flabellately disposed. Flowers (in bud) on pedicels to 1 cm long, rigid, shiny, the calycine cup acetabuliform, ca. 0.5 mm long, coriaceous, glabrous, the margin undulate with lobes absent or 4, widely triangular, ca. 0.2 mm long; corolla white, the tube narrow cylindrical, ca. 1.5 cm long, falcate, glabrous, carnose, the lobes 4, the buds conspicuously ob- tuse at the apex, to 1 cm long, ca. 0.45 cm wide, carnose, concave, auriculate unilaterally basally; stamens 4, the anthers sessile, linear oblong, ca. 8.5 mm long, ca. 0.8 mm wide, obtuse toward the apex, attached above the middle of the tube: ovarian disc coroniform, ca. 1 mm long, black when dry; style slender, to 1.2 cm long, plane, the stigmas 2, narrowly triangular, ca. 1 mm long, acute, scarcely wider than the style, pubescent, the ovary sometimes lacking, the ovar- ian wall thin, the locule ample, the ovules numerous. Fruits subsessile, rotund, ca. 3 cm in diam., smooth, glabrous, purple black when dry, the calycine scar ringlike, ca. 0.6 cm in diam. Genipa curviflora is known only from Panama. Few flowers were available for dissection and these are in the bud stage. On dissecting two buds of Duke 10678 I found that the locule of the ovary is located at the proximal end of what appears to be the pedicel. Two buds of Nee 10347 showed no ovary in the presumed pseudopedicel, although the style and stigmas were evident. Thus, it seems advisable to call the structure in the latter a true pedicel and its counterpart (Duke 10678) a pseudopedicel. Genipa curviflora is closely related to G. williamsii Standley, known only from Panama, being distinguished by its narrower leaf blades which taper some- what at the apex and which are only thin coriaceous. The fruits are smaller than those of G. williamsii. DARIÉN: Camp Summit between Mortí and Sasardí, 1800 ft, Duke 10678 (MO). PANAMÁ: El Llano to Cartí Road, ca. 14 km N of Panamerican Highway, 350-500 m, Folsom & Kauke 1403 (MO): Folsom et al. 1475 (MO). El Llano-Cartí Road, 8 km N of Panamerican Highway, 450 m, Nee & Warmbrodt 10347 (MO). 3. Genipa williamsii Standley, J. Wash. Acad. Sci. 8: 642. 1918. TYPE: Panamá, R. S. Williams (US, holotype). Trees 4-20 m tall, the twiglets stout, ultimately 4-angled, the bark rough, smooth, glabrous, the nodes moderately well spaced, the leaf scars triangular rotund, ca. 5 mm in diam. Leaves oblong rotund to obovate oblong, to 33 cm long, to 16 cm wide, rounded or obtuse at the apex, not acuminate, cuneate, rounded or obtuse at the base, the costa slender, prominulous above, prominent beneath proximally, the lateral veins 9-12, arcuate, with an undulate, submarginal vein well set in from the margin, coriaceous, scarcely discolorous, drying a deep brown, glabrous; petioles 0.5-6.0 cm long, stout, to 0.4 cm wide, rigid, lignose, 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 often curved basally; stipules deciduous, presumably rhomboid oblong, to 3.2 cm long, to 1.8 cm wide, striate, coriaceous, glabrous except ciliolate, thickened basally. Inflorescences terminal, the flowers 3—4(—7) on a suppressed peduncle, to 1 cm long, the bracteoles triangular, to 3 mm long. Flowers perhaps monoe- cious, subsessile or jpeg the pedicels to 1 cm long, ca. 0.2 cm wide, drying black, glabrous; buds ca. 5 cm long, clavate, the mass of corolla lobes oblong, to 2 cm long, ca. 1.3 cm wide; hypanthium often blending in with pedicel, tur- binate, ca. 5 mm long, drying black, glabrous, the calycine cup scarcely measur- able, platelike, with 5 vague lobes, compressed rotund, ca. 1 mm long, wider than long, glabrous, coriaceous; corolla white, the tube 1.8—3 cm long, ca. 0.5 cm wide in the middle in bud, to 1 cm wide in open flowers, coriaceous, glabrous outside, the lobes 5, imbricate, triangular oblong, to 2.1 cm long, to 1.5 cm wide, glabrous outside, often with a few scattered white hairs within; male flowers with anthers sessile, 1.8-2.0 cm long, ca. 1 mm wide, attached below the mouth; rudimentary style linear, ca. 2.5 cm long, ca. 1 mm wide; stigmas 4, unequal (at least one of 4), oblong, to 4 mm long, acute with a few hairs adaxially. Fruits rotund, to 7 cm in diam., smooth, glabrous, drying black, the wall to 1 cm thick, the seeds numerous, plano compressed, oblong or subrotund, to 1 cm long, yel- low, embedded in a white pulp. Genipa williamsii is known only from Panama and Colombia. This species is easily recognized by a combination of characters: leaves rounded and obtuse at the apex, drying brown; buds elongate and clavate at the apex; fruits very large and drying black. I have not seen female flowers. One of the striking floral char- acters is the fact that the flower buds are elongate and clavate while the open flowers appear much shorter than the buds. According to Duke (8026) the ‘‘fruit (is) very bitter." NAMA: COCLÉ: Furlong's Finca, Cerro Pilón, 880 m, P is 37571 (MO). 3 km NE of El Valle, Mori & Kallunki 2959 (MO). La Mesa, 4 km N of El Valle, , Nee & Dwyer 9152 (MO). DARIEN: , Williams 814 (US). PANAMA: on ro uda Duke n Dol Cerro Azul T Cerro Jefe, Dwyer 5041A (MO). 9.4 km N of Goofy Lake, 900 m, Folsom et al. 1987 (MO). 4.8 km N of highway, W of * ne Cerro Jefe, Gentry 5517, 6138 (both. MO). El Llano-Cartí Road, 12.7 km from Panamerican , Mori et al. 4688 (MO). El Llano-Cartí Road, 8-11 km from Panamerican Highway, Mori dm TA (both MO). E slope of Cerro Jefe, Tyson 3397 (MO). Cerro Jefe, Tyson et al. 4441 (MO). COLOMBIA: CHOCO: Municipio de Novita. Vereda Llanadas. Ladera norte del Cerro Torra. Filo al Oeste del Río Surama. Camino al Alto del Oso, 600—900 m, Forero et al. 3082 (MO) 41. GEOPHILA Geophila Don, Prodr. Fl. Nepal. 136. 1825. Nomen conserv. TYPE: G. reniformis D. Don, nom. illeg. — G. herbacea (Jacq.) Schumann. TE E. F. Wight, Contr. U.S. I. Herb. 9: 216. 1905. rYPE: Psychotria herbacea Jacq. = Geophila herbacea (Jacq.) Schuman Gan Standley, Contr. U.S. Natl. Herb. 17: 444. 1914. TYPE: not indicated. Creeping herbs. Leaves often cordate, membranous; petioles elongate. /nflo- rescences axillary, usually pedunculate, the flowers few in a terminal cluster. Flowers with the calycine tube dentate or lobate; corolla funnelform, pilose in the throat, the lobes valvate; stamens 4-7, the filaments slender, inserted in the throat; ovary 2-celled. Fruits fleshy, drupaceous, the 2 nutlets plano convex, compressed dorsally, usually costate. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 187 Geophila, a genus of about 30 species, occurs throughout the tropical areas of the world. Literature: Steyermark, J. A. 1977. A new member of the Rubiaceae from Panama. Phy- tologia 35: 401—402. £e Leaf blades with auricles not overlapping at the base; bracts lanceolate or lance oblong. b Leaves with abundant cystoliths beneath, the een 2.5-6.0 cm long, 2.0-4.5 cm wide; peduncles 0.1—0.2 cm long at anthesis, glabro l. G. croatii bb. Leaves icr cystoliths beneath, the blades 2— Í cm ipe 1.0-8.5 cm wide; pedun- cles 0.8—5.0(—8.0) cm long at anthesis, puberulou 2. G. macropoda aa. Leaf blades with auricles overlapping at the base; a subulate |... À... . G. repens 1. Geophila croatii Steyermark, Phytologia 35(6): 401—402. 1977. TYPE: Panama, Croat 6647 (MO, holotype). Creeping herbs, glabrous, conspicuously rhizophorus at the nodes, the roots fibrous. Leaves widely ovate or lance oblong, 2.5-6.0 cm long, 1.5—4.5 cm wide, acute or acuminate at the apex, cordate at the base, the lateral nerves 4-5, prom- inulous beneath, uniting to form a vein 1-5 mm from the margin, membranous, glabrescent or the costa and the veins with a few hairs with abundant cystoliths beneath; petioles 0.5-9.0 cm long, glabrous to puberulent; stipules ovate, ca. 3 mm long, obtuse, glabrescent. /nflorescences capitate umbellate, the peduncle at anthesis 1-2 mm long, elongating in fruit, to 20 mm long, the flowers usually 5— 7 in a head; bracts lance oblong or lanceolate, 3.0-4.5 mm long, 1.0-1.5 mm wide, glabrous; pedicels less than 1 mm long at anthesis, to 3 mm long in fruit, glabrous. Flowers with hypanthium and calyx 2.0-4.5 mm long, the calycine lobes 5, narrowly oblanceolate or lanceolate, 2.5-3.5 mm long, much longer in fruit, glabrous; corolla white, ca. 5 mm long, with hairs clustered below filament attachment, the lobes slender; stamens with filaments connate below. Fruits ovoid or globose, 3-4 mm in diam., red, often in a cluster of 3; seeds oblong, to 4 mm long, plane on one side, somewhat twisted, the ribs 3, faint. Geophila croatii Is known only from Panama. CANAL ZONE: Barro Colorado Island, Croat 6647, 10989 (both MO). Gaillard Highway between Summit Garden and Gamboa, Sullivan 27 (MO). 2. Geophila macropoda (Ruiz & Pavón) DC., Prodr. 4: 537. 1830. usss macropoda Ruiz & Pavón, Fl. Peruv. Chil. 2: 63. 1799. Type: Peru, Ruiz & Pavón, not see Coccoc sium mac dp (Ruiz & A pee Bull. Torrey Bot. Club 18: 107. 1891. PO s, ia H.B.K., Nov. Gen. Sp. Pl. 3: 379. 1818. Type: Colombia, between Barthomome and Nares, Humboldt & Meus. um seen. Geophila vitas olia t H.B.K.) DC., Prodr. 4: 537. 1830. Geophila herbacea violaefolia (H.B.K.) Chod. & Hassl., Bull. Herb. Boissier II. 4: 180. 1904. Geocardia violaefolia (H.B.K.) Standley, Contr. U.S. Natl. Herb. 17: 445. 1914. Creeping herbs, the stems weak, glabrous, often farinose. Leaves 1-4 at a node, cordate or ovate cordate, 2-9 cm long, 1.0-8.5 cm wide, occasionally wider than long, acute at the apex, cordate at the base, the auricles not overlapping, occasionally inequilateral, the costa plane above, prominulous beneath, the lateral 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 veins 4—6, the intervenal areas reticulate, papyraceous, glabrous above, glabres- cent beneath, the cystoliths not evident; petioles weak, flexuous, 0.8-14.0 cm long, rubescent, pubescent; stipules not seen. Inflorescences with peduncles 0.3— 3.5 cm long, much longer in fruit, puberulous, the flowers few, aggregated ter- minally in clusters to 0.8 cm wide; bracts lanceolate, to 3 mm long. Flowers with the hypanthium rectangular, ca. 1.5 mm long, | mm wide, the calycine cup ca. 1 mm long, stiffly petaloid, glabrous outside and inside, with a few glands proxi- mally within, these separate or clustered in 2’s or 3’s, oblong, ca. 0.1 mm long, mostly obtuse, the teeth 5, linear lanceolate, slightly unequal, 2.2-3 mm long, to 0.5 mm wide; corolla white, not seen; stamens not seen; ovarian disc hemispher- ical, to 0.8 mm long, smooth and glandular shiny, the style slender, to 3 mm long, glabrous, the stigmas 2, digitiform, ca. 0.8 mm long. Fruits (usually 2 maturing), ovoid, to 1 cm long, glabrous, the calyx persistent, 1-2 mm long. Geophila macropoda occurs in Costa Rica, Panama and Colombia. ‘‘Nomas Tuwat'' (Cuna, fide Duke 14317). BOCAS DEL TORO: Purser farm, Armuelles, Dickson 2292 (MO). Vicinity of Nievecita, Msn et al. 1851 (F, MO). DARIEN: Río Pirre, trail from El Real to Tucutí, Duke 5201 (MO). PANAMA: Piria- Canasas Trail near Piria, Duke 14317 (MO). sAN BLAs: Headwaters, Río Mulatupo, Elias 1739 (GH, O). 3-4 hours up Río Mulatupo, Kirkbride 217 (MO). 3. Geophila repens (L.) Johnston, Sargentia 8: 281. 1949.—FiG. 43. Rondeletia repens L., Syst. Nat. ed. 10. 928. 1759. Type: Jamaica, not seen. Psychotria repens (L.) L., Amoen. Acad. 5: 377. 1759. P. herbacea Jacq., Enum. Pl. Carib. 16. 1760. a Bae aet not seen. Cephaelis reniformis H.B.K., Nov. Gen. Sp. PI. 3: . 1819. TYPE: not seen. Geophila cordata Miq., Linnaea 17: 72. 1843. TYPE: mum. Focke 471. Mapouria cordata (Miq.) Müll.-Arg., Mart. Fl. Bras. 6(5): 426. 1881. Geophila herbacea (Jacq.) Schum., Engl. & Prantl. Nat. Pflanze nfam. 4. 4: 119. 1891. Geocardia herbacea (Jacq. ) Standley, Contr. U.S. Natl. Herb. 17: 444. 1914. Carinta repens (L.) Smith & Downs, Sellowia 7: 88. 1956. C. repens (L.) Brem., Verh. Kon. Nederl. Akad. Wetens 54: 118. 1963. Redundant combination. Carinta repens var. americana Brem., Verh. Kon. Nederl. Akad. Wetens, 119. 1963. TYPE: not seen. Prostrate herbs, rooted at the nodes, the stems weak, the nodes well spaced. Leaves cordate, occasionally ovate rotund, (1.2-)1.7—4.5(—5.0) cm long, 1.3-4.2(- 5.0) cm wide, obtuse, widely deltoid or rounded at the apex, basally cordate, auriculate, the auricles overlapping, the lateral veins 5, the 3 proximal veins arising from a common point, prominulous beneath, the veinlets inconspicuous, glabrous to scabridulous above, glabrous beneath except on the costa, usually with cystoliths on the lower surface; petioles 0.6—4.5(—6.0) cm long; stipules not seen. Inflorescences capitate with the peduncle 0.8—-5.0(-8.0) cm long, glabrous to papillate at anthesis, the flowers few; bracts subulate, to 5 mm long, pilose, the outermost often with 2 short lateral lobes at the base. Flowers (in bud) with the hypanthium oblong or rectangular, ca. 1 mm long, glabrous, the calycine cup funnel shaped, ca. 0.8 mm long. glabrous outside and inside, eglandular within, the calycine lobes 5, subulate or lanceolate, ca. 3 mm long, slightly unequal, to 1 mm wide at the base, pubescent marginally; corolla white or orange, clavate in bud, to 7 mm long (10 mm long, fide Johnston), fleshy, petaloid, pubescent es- pecially at the apex outside, glabrous and with numerous raphides within; stamens 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 189 FIGURE 43. Geophila repens (L.) Johnston. Habit (x l). [After Tyson & Blum 1658. | 5, the anthers oblong, ca. 1 mm long, obtuse; style slender, the stigmatic lobes 2, short. Fruits red, ovate oblong, to 0.5 cm long; calyx cup and lobes persistent, to 3.5 mm long; seeds twisted and rugulose. Geophila repens extends from Mexico throughout Central America, and in the Guianas, Venezuela, Colombia, south to Argentina. It also occurs in the West 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Indies. Duke 14345 notes that the herb is used medicinally by the Bayano Cuna Indians, who call it, “Nomas Puruiwat.”’ OCAS DEL TORO: Changuinola, Dunlap 565 (F). Chiriquicito to 5 mi S along Rio Guarumo, Lewis et al. 2097 (MO). Pumpkin River, Wedel 2577 (MO). CANAL ZONE: 2 mi from Gatun Locks, Blum 1491 (MO). Summit Garden, Mori & Kallunki 1885 (MO). Balboa, Standley 25503 (MO). Old Fort Lorenzo, Tyson 2207 (MO). Gamboa, Tyson 4570 (MO). Fort Sherman, Tyson & Blum 1658 (MO). Miraflores Lake, White 252 (F). Salamanca Hydrographic Station, Río Pequeni, AE et ~~ Hee (F, MO). Vicinity of Summit Garden, Witherspoon & apt sala 870 00 (MO). ADO ISLAND: Bailey & Bailey 3 (F); Rancham 558 (F); Bartlett & Las 16738 (MO); Cas 4347, 5834, 6464, 7976, 10171, eed 15121 (all MO); Ebinger 230 (F, MO); aay "rg 48163 (MO); Hayden 153, 1030 (both MO, UC, VEN); Salvoza 967 (F); Shattuck 130 (F, MO); Starry 286 (F, MO); Woodson & Schery 986 (MO). ME San Bartolomé, Burica Peninsula. Woodson & Schery 919 (F, MO). DARIÉN: Río Chico, from junction with Río Chucunaque to ca. 1 hour by outboard, Burch et al. 1080 (MO). Pinogana ue El Real, Croat & Porter 15477 (MO). 10 mi S of El Real, S bh) Mais 5447 (MO). Río Pucro to Quebra Maskia, Duke 13055 (MO). Quebrada ‘‘Comachim ` Correa Casa de Bartolo, Kennedy 2854 = Rio Tuquesa, middle of Tuquesa Mining Co.. “camp called cn Peje, Mori 7054 (MO). Paya, Rio Paya, es et al. 431A od LOS SANTOS: Headwaters Río Ped- regal, 25 mi SW of Tonosí, ges et al. 2903 (MO). PAN : El Llano and Río Maou Duke 5553 (MO). Piria Canasas Trail near Piria, Duke 14345 (MO). Ana. Woodson et al. 769 (MO). 42. GONZALAGUNIA mR deus Ruiz & Pavón, Fl. Peruv. Prodr. 12. 1794. TYPE: G. dependens Pavón. Dugge na West, Bidar. Ste. Croix 269. 1793. Nomen nudum. type: D. richardii, based on Barleria Buena Cav., Anales Hist. Nat. 2: 278. 1800, not Pohl 1827. TYPE: B. eagle oe Gonzalea Pers., Syn. Pl. 2: 132. 1805. New name for Gonzalagunia Ruiz & Pav Shrubs or small trees, the branchlets slender. Leaves petiolate or subsessile; stipules free, usually triangular subulate. Inflorescences terminal, elongate, slen- der, spiciform or thrysiform. Flowers with the hypanthium rotund, the calyx with 4-5 small, persistent lobes; corolla funnelform or salverform, the tube villous, the lobes 4-5, valvate or imbricate; stamens 4-5, the filaments slender, attached to the tube; anthers slender, included; ovarian disc crenulate, the style slender, the stigmas small, 2-4-lobed, the ovules numerous, the placentas small. Fruits baccate, globose, 2-4-sulcate, the pyrenes 2 or 4, chartaceous to bony; seeds numerous, minute, subglobose, deeply foveolate. Gonzalagunia, a genus of about 25 species, is common throughout the Amer- ican tropics and the West Indies. It is among the easiest genera of the Rubiaceae to recognize because of its pendent spiciform inflorescences. Literature: Donnell-Smith, J. 1899. Undescribed plants from Guatemala and other Central American Republics. Bot. Gaz. 27: 336. Standley, P. 1921. New Rubiaceae from Colombia and Costa Rica. Contr. U.S. Natl. Herb. 17: 446. ——N. 1929. Studies of American plants II: Rubiaceae. Field Mus. Nat. Hist., Bot. Ser. 4: 278-281 a. Leaves 10-13 cm long; lateral veins of larger leaves 6-12. Largest bracts 5-8(-12) mm long: flowers blue; fruits dicoccus __ 1. G. bracteosa 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 191 bb. Maie bracts 3-5 mm long; flowers white or red; fruits 4-coccus. Leaves subsessile or with petioles to 0.3 cm lon Stipules subulate, 8-9 mm long: calycine lobes ca. 2 mm long, obtuse 3. G. ovatifolia dd. Stipules ovate or deltoid, ca. 6 mm long; calycine lobes ca. 0.7 mm long, acute 6. G. rudis ec. "dgio kd petioles 0.5-2.0 cm lon s 7-22 cm long; stipules 8- 10 mm long, corolla pink, the tube ca. 8 mm bu densely appressed villose within -------------------------- . G. rosea ee. Leaves 5-13 cm long; stipules ca. 3 mm long; corolla white, x tube 10.0- .5 mm long, glabrous within . G. panamensis aa. Leaves 15-24 cm long; lateral veins of larger leaves 12-16 (except G. si Lateral veins of larger leaves 12-16; flowers white Flowers pedicellate 2. G. kallunkii Flowers sessile or subsessile 7. G. veraguensis ff. Lateral veins of larger leaves 6-12; flowers pink |... 5. G. rosea 1. Gonzalagunia bracteosa (Donnell Smith) Robinson, Proc. Amer. Acad. Arts 45: 405. 1910 io bracteosa Donnell Smith, Bot. Gaz. 33: 252. 1902. type: Guatemala, Donnell Smith 6583 (F, hee bracteosa (Donnell Smith) Standley, Contr. U.S. Natl. Herb. 18: 125. 1916. Shrubs 1.5-2.0 m tall, the lowermost branches occasionally prostrate, the branches smooth, glabrous to villose, wiry, to 0.15 cm in diam., the nodes often swollen, well spaced. Leaves oblong, obovate oblong or narrowly elliptic, 8— 18(—22) cm long, 2.5-8.5 cm wide, widely deltoid or scarcely obtuse at the apex, short acuminate, the acumen to 1.5 cm long, acute, subcuneate or obtuse at the base, the costa prominulous above, prominent beneath, the secondary veins 6- 9(—11), strongly arcuate ascending, the intervenal areas spreading reticulate, char- taceous, pubescent, the hairs 1.0-1.5 mm long, appressed, white, more dense beneath; petioles short or absent, to 0.5 cm long; stipules connate, cylindrical, the sheath closely appressed to stem, thin, densely golden villose or hirsute, occasionally carinate, the hairs more dense along the keel; each division tapering into an acute awn to 5 mm long. Inflorescences axillary, pendent, (5-)15-25 cm long, of ca. 20 flower clusters well spaced along an often flexuous rachis; pedun- cles 4-9 cm long; bracts several subtending the mass of flowers, deflexed, nar- rowly ovate lanceolate, 5-8(-12) mm long. Flowers sessile, the hypanthium and calycine cup subrotund, ca. 1.2 mm long, carnose, densely villose, the calycine lobes 5, to 0.5 mm long, densely villose outside, glabrous within, with a minute gland at the base of adjacent lobes; corolla blue (in bud), to 4 mm long, the tube apparently glabrous outside, villose within, the lobes 5, villose outside and within; stamens 5, the anthers oblong, to 1 mm long, minutely acuminate, affixed near the orifice; style and stigmas to 2.8 mm long, plano compressed, the style slender along most of its length, expanding above to 0.7 mm wide, the style and 2 stig- matic lobes sagittate, the lobes to 0.4 mm long; ovarian disc compressed cylin- drical, to 0.2 mm long. COLON: Río Guanche, ca. 3-5 km above bridge on Portobelo Road, Croat 37012 (MO). Río G ca. 2.5 km upriver from bridge, road to Portobelo, Mori et al. 6470 (MO). DARIÉN: Cocalito, Dwyer 4360 (MO). Caná- Cuasí Trail (Camp 1). Chepigana, Terry & Terry 1621 (F. MO). PANAMÁ: Tres. 3 mi NE of Altos de Pacora, 500-800 m, Croat 22701 (MO, US). El Llano- Cartí Road, Croat 33763 (MO); D'Arcy 10613 (MO); Liesner 1201 (MO, US); Mori et al. 4588 (MO); Mori & Kallunki 5587 (MO). 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 2. Gonzalagunia kallunkii Dwyer.^ rype: Panamá, Mori & Kallunki 3141 (MO, holotype). Shrubs to 2 m tall, the twiglets terete, smooth, the hairs short, dense, golden, the nodes well spaced, the pith occasionally porate. Leaves elliptic oblong, 17- 24 cm long, 5.0-11.5 cm wide, acute, short acuminate, the acumen short, widely triangular, to 1.5 cm long, often slightly falcate, basally acute, cuneate to atten- uate acute, the blade slightly decurrent on the petioles, the costa prominulous above, prominent beneath, the lateral veins 12-14, arcuate, the undulate sub- marginal vein to 3 mm from the margin, membranous, discolorous, dark re brown above when dry, dull gray green beneath, the hairs numerous, often ap- pressed strigose, sometimes glabrescent; petioles to 2 cm long, rigid, lignose; stipules free, triangular, to 8 mm long, acute, with a median and densely bearded keel, the hairs golden. Inflorescences densely pubescent, to 35 cm long, to 2 cm wide: peduncle to 6 cm long, to 0.12 cm wide at the base. Flowers with pedicels to 0.5 cm long, the hypanthium and calycine cup compressed rotund, to 3 mm long, truncate, densely hairy, the calycine lobes 4, triangular, 1-1.5 mm long, hirsute outside, glabrous within; corolla white (in bud), the tube slender, 5.5-7.0 mm long, densely villose outside, glabrous within; stamens 4, subsessile, the anthers to 1.3 mm long, attached at base of tube; style 3 mm long, the stigmas ca. 0.7 mm long. Fruits white, rotund, ca. 0.5 cm in diam., densely appressed villose. Gonzalagunia kallunkii is known only from Panama. It is named in honor of Jacquelyn Kallunki, who has contributed much to the collection of valuable her- barium material in Panama. It is closely related to G. rosea Standley which has pink flowers, fewer lateral veins, and glabrate fruit. ERAGUAS: Valley of Dos Bocas, between Alto Piedra and Calovébora, 350-400 m, Croat 27462, 27733 (both MO). NW of Santa Fé, 11 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 3141 (MO) 3. Gonzalagunia ovatifolia (Donnell Smith) Robinson, Proc. Am. Acad. Arts 45: 405. 1910 Gonz ju : ovatifolia Donnell Smith, Bot. Gaz. m 336. 1899. svyNTYPES: Costa Rica: Donnell Smith 6600 (both F); Pittier & Tonduz 9171; Panamá: Hart 113. hig na aie f (Donnell Smith) Standley, C U.S. Natl. Herb. 18: 126. 1916. Shrubs to 3 m tall, the branchlets pubescent. Leaves subsessile, ovate oblong, to 10 cm long, to 5 cm wide, acuminate, the lateral veins (6—)8—12, discolorous, dark chocolate color above, lighter beneath, glabrescent above, sericeous be- neath; stipules subulate, 8-9 mm long, triangular at the base, 2-3 mm wide. 50 Gonzalagunia kallunkii Dwyer, spec. nov. Frutices ad 2 m alti, ramulis laevibus aureo-pu- erulis. Folia elliptico-oblonga, 17-24 cm longa, 5.0-11.5 cm lata, acuta, acumine ad 1.5 cm longo, venis lateralibus 12-14, membranacea glabrescentia vel ciliis crebris appresso-strigosis; petiolis ad 2 cm longi ipuli l S stipulis ad longis carina media dense aureo-barbata ornatis. Inflorescentiae ad 35 cm longae floribus rare solitariae pleru in fasciculis florium paucorum sessiles vel subsessiles bene distantes separatis. Flores hypanthio dense puberulo, rotundi, ca. 3 mm diam.: corolla alba tubo ad 7 mm longo extus a appressis M Re albis praedito, lobis ad 2.5 mm longis intus glabris. Fructus rotundi ad 0.5 cm diam., dense hirsuti 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 193 Inflorescences terminal, (8-)17—27 cm long with peduncles 2-3 cm long or seem- ingly sessile and subtended by regular-sized leaves, the cymules 1-5-flowered, sessile, the bracts subulate, 3-5 mm long. Flowers with the hypanthium com- pressed rotund, ca. 1.5 mm long, the calycine lobes 4, oblong, ca. 2 mm long, usually curved, ciliate, with 1 to several rotund glands at the junction of the lobes; corolla white, the tube to 2 cm long, strigose, the lobes 4, obtuse, often as long as the tube; stamens 4, not exserted; ovarian disc pulvinate and bearing a few hairs, the style with the 4 stigmas included. Fruits tetracoccus, to 1 cm in diam. with white flesh persisting, hirsute, drying white ca. 3 mm in diam. ; seeds angular, honeycomb reticulate. This species is known only from Costa Rica, Colombia and Panama. Donnell Smith (1899) cited Hart 113, collected in Laguna de Chiriqui (Bocas del Toro), Panama in 1895. AS DEL TORO: Changuinola Valley, Dunlap 102 (F), 496 (F, BM). Without other locality, Wedel. 341 (MO). cociÉ: La Mesa above El Valle, denda 3653 (MO). SAN BLAS: Mainland opposite Achituppu, Lewis et al. 100 (F. GH, MO). COLOMBIA: CHOCO: Trail between Alto Coriche and Camp Curiche, E of Boca Curiche, 10—400 ft, Duke & Idrobo 11306 (NY). 4. Gonzalagunia panamensis (Cav.) Schum. in Mart., Fl. Bras. 6(6): 292. 1889.— Fic. 44 Buena panamensis Cav., Anales Hist. Nat. 2: 279. 1800. TYPE: dedeg Qe (?MA). Gonzalea cornifolia H.B.K., Nov. Gen. Sp. a 3: 416. 1820. TYP G. panamensis Spreng., Syst. Veg. 1: 417. 1825. TYPE: Panamá, pap iii (BM, not seen; photo, i leptantha A. Rich. in Sagra, Hist. Cuba 11: 16. 1850. Type: Cuba, Ramon de la Sagra, not Guarani id iier Maza, Anales Soc. Esp. Hist. Nat. 23: 289. 1894. TYPE: not seen. Gonzalea hayesii Wernham, J. Bot. 51: 219. 1913. Type: Panama, Hayes (B Duggena hayesii (Wernham) Standley, Contr. U.S. Natl. Herb. 18: c 1916. D. leptantha (A. Rich.) Standley, Contr. U.S. Natl. Herb. 18: 125. Shrubs to 10 ft tall, the branchlets terete, smooth, villosulose, the nodes well spaced, mostly 3-6 cm apart. Leaves elliptic or ovate elliptic, 5-13 cm long, 1- 4(—6) cm wide, acute at the apex, usually acuminate, the acumen to 2 cm long, cuneate to subobtuse at the base, often markedly inequilateral, the costa prom- inulous above and beneath; petioles slender, 1.0—1.5(-2.0) cm long, ca. 0.8 mm wide, pubescent; stipules ovate oblong, widely triangular to subulate, to 3 mm long, the awn usually evident, weak, as long as or longer than the body. Inflo- rescences terminal, solitary, to 31 cm long, the peduncles flagelliform, 1.5—4.0 cm long, to 0.2 cm wide, pubescent, the cymules sessile or subsessile, 0.5-1.0 cm apart; bracts linear subulate or triangular subulate, shorter than the hypan- thium or much longer, to 7 mm long, pubescent; pedicels short or absent. Flowers with the hypanthium subrotund, ca. 0.8 mm long, puberulent, the calyx ca. | mm long, the teeth 4, triangular, ca. 0.4 mm long, the cup stiffly petaloid, glabrous and eglandular within; corolla white, the tube narrowly cylindrical, 10.0-13.5 mm long (or longer), ca. 1 mm wide, petaloid, sparsely hairy outside, sometimes glabrescent, glabrous within, the lobes 4, oblong rotund, 3.2 mm long: stamens 4, the anthers narrowly oblong, 2.0-2.2 mm long, the filaments short, ca. 0.7 mm long, attached 3-4 mm below the mouth: style slender, 7-10 mm long, the stigmas 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 pe » < Ake //, LANDE NI" D» A s LA \ = n FIGURE 44. Gonzalagunia panamensis (Cav.) Schum.—A. Habit (x'2).—B. Flower, opened to show stamens (x 51⁄).—C. Ovary, cross section (x 51⁄2). [After Tyson 1422.] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 195 4, ca. 0.7 mm long. Fruits 4-lobed, rotund, 3-4 mm in diam., glabrous to pubes- cent, white or purple black at maturity. Gonzalagunia panamensis ranges from Mexico to Colombia, and it also occurs in the West Indies. A common name in Panama is ‘‘Niguita.” Stier 200 reports the common name "Ina Kapuriwiwa" by the San Blas Cuna Indians. Steyermark (1972) pointed out that Gonzalagunia cornifolia (H.B.K.) Standley (1929) is the "Gonzalagunia panamensis of Standley and other authors, not G. panamensis (Cav.) Schum," but he was referring to Standley's use of his name for plants of Venezuela, not his use of the name for plants of Panama. Literature: Steyermark, J. A. 1972. The Botany of the Guayana Highlands—Part IX. Mem. New York Bot. Gard. 23: 312. BOCAS DEL TORO: Changuinola Valley, usd nie (BM, US). CANAL ZONE: Ancon, Heriberto (US). Cerro Luisa, Road C- i: Regis 10773 (MO). Ft. Sherman, Duke & Mussell 6639 (MO). Quarry Heights, Dwyer 2608 (MO). Fort Sherman, Dwyer 2M MO). Coco Solo, Dwyer & Duke 7883 (GH, MO, UC, US). K-9 Military a Ebinger 543 (MO). Balboa Heights, Greenman & Greenman 5032 (MO). Ancon, ae 12052 (US). Gamboa, Pittier 3711, 3785 (both US). Ancón, Rowlee & Rowlee 396 (US). Ancón Hill, Seibert 404 (MO). Balboa, "ped 26977 (US). Corozal, Standley 27358 (US). Fort Sherman, mouth of Chagres, Tyson 1536 (M Fort San Lorenzo, Tyson 2214 (MO). Miraflores, White & White 57 (MO). CHIRIQUI: El Hato mine nia. 816 (MO). Frances Arriba School, 14 mi N of David, Lewis et al. 675 (GH, K, MO, NY). c : El Valle, Blum et al. 2381 (MO); Duke 13271 (MO). Penonomé, Dwyer 20/0 (MO). El Valle, Dwyer 2381 (MO). Near El Copé, Garner 38 (MO). El Valle, Seibert 428 (MO). COLON: Portobelo, Croat 11412 (MO); Elias & Kirkbride MO, UC, VEN); Folsom 3753 (MO); Gentry 1738 (MO). N side Río Guanche, 12 km upstream from Puerto nd Highway, Nee 7088 (MO). HERRERA: 10 mi S of Ocu, Tyson 2844 (MO). PANAMA: Campana, Allen & Alston 1865 (MO). Pacora and Chepo, Dodge et al. 16667 (BM, MO). San José Eins) Evanson 544 (US). Chagres, Fendler 288 (MO). San José Island, oe 76 (MO). Las Cumbres, Rivera 18 (MO). Chepo, Tyson 1422 (MO). Veracruz, Wendehake 9 (MO). SAN BLAS: Between Río Diablo ae Rio ibis near Narganá, Duke 14859 (MO). Mainland opposite Achituppu, Lewis et al. 100 (GH, MO). Niatuppu, ye 200 (MO). VERAGUAS: 5 mi NE of La Mesa, Blum & Tyson 668 (MO). Santiago, . 1253 (MO). Ponuga, Dwyer & Kirkbride 7441 (GH, MO, US). Santiago, Dwyer et al. 7557 (COL, MO, VEN UC). — ~ 5. Gonzalagunia rosea Standley, Ann. Missouri Bot. Gard. 25: 836. 1938. TYPE: Panamá, White & White 7 (F, holotype; isotypes, MO, NY). Trees, small, to 5 m tall, or shrubs, the branchlets terete, stiff or flexuous, densely pubescent. Leaves oblong, ovate oblong or elliptic, 7-22 cm long, 2-8 cm wide, deltoid at the apex, often with an acumen to 1.5 cm long, basally cuneate or rounded, the costa prominulous above and beneath, the lateral veins 8-13, prominulous, arcuate, papyraceous, the hairs moderately dense, strigose or spreading, often farinose above (bases of deciduous hairs), usually densely golden villose beneath; petioles 0.5-2.0 cm long, 0.1-0.2 cm wide, pubescent; stipules oblong triangular or triangular, 8-10 mm long, acuminate, the acumen often 4 times the length of the body, keeled, golden pubescent. Inflorescences terminal, spiciform, 15-35 cm long, often nodding, to 5 cm wide, pubescent; peduncles to 4 cm long, to 0.18 cm wide, the cymules well spaced; bracts small. Flowers with the hypanthium obovoid, ca. 3 mm long, ca. 1.4 mm wide, villosulose, the ca- lycine cup ca. 1 mm long, petaloid, the teeth 4-5, triangular, ca. 0.6 mm long, with 2—4 subrotund glands at the junction of the lobes; corolla pink, the tube slender, ca. 8 mm long, ca. 1.6 mm wide in the middle, densely pubescent outside, densely appressed and white villose within, carnose, the lobes 5, 2 mm long, 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 thick, the margins infolded, densely villose adaxially; stamens 4, the anthers oblong, ca. 2.5 mm long, ca. 0.25 mm wide, the filaments short, inserted 24 way up the tube; stigmas 4, unequal, subfalcately digitiform, 0.6-1.0 mm long, the style filamentous, ca. 9 mm long, the ovary 4-celled, thick-walled, the ovules numerous, rubescent. Fruits 4-coccous, 2-3 mm in diam., glabrate. Gonzalagunia rosea 1s known from Costa Rica and Panama. Mori & Dressler 7875 from Cerro Colorado, Chiriqui, is noteworthy in having glabrous leaves. BOCAS DEL TORO: Between Quebrada Gutierrez and E slope La Zorra, Kirkbride & Duke 737 (MO). CHIRIQUI: Cerro Horqueta, Blum & Dwyer 2638 (MO). Bambito and Cerro Punta, Croat 10676 (MO). Top of peak between Barü and Respinga, ca. 3000 m, D'Arcy 10125 (MO). La Popa, above Boquete, D'Arcy 10859, 10946 (both MO). Bajo Grande, Di Ariy & D'Arcy 6561 (MO). Bajo AM Davidson 152 (MO, US), 259 (MO). Cerro Horqueta, Duke et al. 13615 (MO); Dwyer et al. 490 (M / . Boquete, Dwyer 6992, 6995 (both MO). Cerro Horqueta, Dwyer & Hayden 2 (MO): Dod & Lallathin 8774 (MO). 2 Volcán on road to Costa Rica, Ebinger 816 (MO, US). Quebrada Hondo and divide on Caldera-Chiriquicito Trail, Kirkbride & Duke 927 (MO). 27 km NW of El Hato del Volcán, 5000-5300 ft, Mori & Bolten 7199 (MO). Cerro Colorado, 50 km N of San Félix, 1200-1500 m, Mori & Dressler 7875 (MO). Cerro Punta, Ridgway & Solis 2409 (MO, UC). Boquete, Stern et al. 1028 (MO, US). 1 m SW of Cerro Punta, Tyson 5646 (MO). Bambito, Tyson 5861 (MO). Río Chiriqui Viejo Valley, White & White 7 (F, MO), 80 (MO). Quebrada Im Herr dae & Redi 262 (MO). DARIÉN: Upper Río Membrillo and Camp 7, Duke 10908 (MO). HERRERA: 10 mi ' Ocú, Tyson et al. 2844 (MO). vERAGUAS: NW of Santa Fé, 11 km from SEE pres Alto de Piedra, Valley Rio Dos Bocas, Mori et al. 3841 (MO). 6. Gonzalagunia rudis (Standley) Standley, J. Wash. Acad. Sci. 17: 170. 1927. Duggenia rudis Standley, Contr. U.S. Natl. Herb. 18: 125. 1916. TYPE: Panama, Pittier 2378 (US). Gonzalagunia acutifolia Rusby, Descr. S. Amer. Pl. 131. 1920. Small trees, to 3 m or shrubs, the branchlets terete, slender, 0.1—0.5 cm wide, puberulent, the nodes well spaced. Leaves ovate oblong, trapeziform oblong, often falcate and inequilateral, 3-11 cm long, 1-4 cm wide, deltoid to acute at the apex, acuminate, the acumen vague or to 1.5 cm long, acute, rounded, obtuse or occasionally truncate at the base, the costa prominulous above and beneath, the lateral veins 8-10, strongly arcuate, papyraceous, glabrescent or pubescent above, the hairs white, strigose, villose beneath, especially on the veins; petioles to 0.3 cm long, the blades sometimes sessile; stipules ovate or deltoid, ca. 6 mm long, the awn subulate, as long as the body, the body sometimes much reduced and the awn then to 5 times longer. /nflorescences terminal, occasionally ap- pearing axillary as on reduced side branches, solitary, exceeding the terminal leaves, to 20 cm long, flagelliform, the peduncles 0.5-3.0 cm long, slender, to 0.1 cm wide, the flowers occasionally solitary or 3—4 in sessile or subsessile cymules, these 0.5-1.0 cm apart; bracteoles linear subulate, to 5 mm long. Flowers with the hypanthium rotund, ca. | mm long, truncate, villose, the calycine tube ca. 0.3 mm long, the teeth 4, unequal, triangular, ca. 0.7 mm long, obtuse to acute, with a minute gland in the sinuses; corolla white, the tube cylindrical, ca. 3 mm long, expanded at the apex, white villose on the outside and within, the lobes 4, oblong rotund, ca. 1.5 mm long, with papillae and hairs on upper surface; stamens 4, the anthers narrowly oblong, 1.0-1.3 mm long, the filaments slender, ca. 0.5 mm long, inserted near base of tube; style linear, 2.5-3.0 mm long, the stigmas 3-4, ca. 0.3 mm long. Fruits white, coccoid, rotund, to 0.5 cm in diam., fleshy, pubescent, the calycine lobes persistent, often radiating in starlike fashion. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 197 Gonzalagunia rudis has a limited range, being known only from Panama and Colombia. In Panama it is known as "'Niguita." Among the Bayano Cuna of Panama it is called ""Usiburnu," ‘‘Puruiwat’’ and ‘‘Inna Ulu” (Duke, 14460). Duke 14548 notes that it is used by the Choco Indians for snakebite. Stier 177 lists the common names "'Ipar kae Tummat” (San Blas) and ‘‘Ipa Kae” (San Blas) and notes that it is used in "'Ibia ina" (apparently as a medicinal). CHIRIQUÍ: 21 km WNW of Puerto Armuelles, Busey 459 (MO). 4 mi S of Puerto Armuelles Liesner 379 (MO). cANAL ZONE: Mohinga Swamp, Bartlett & Lasser 16856 (MO). Gamboa, Clew ell & Tyson 3217 (MO). Pipeline Road, Croat 9383, 12341 (both MO). Pipeline Road, within 5 mi of Gamboa Gate, D'Arcy 9285 (MO). Río Indio, Dodge & Allen 17298 (MO). Ga Pica. Ebinger 904 MO). Pipeline Road, Gentry & Dressler 1967 (MO). 25 km NE of Cerro Azul on Río Piedras, Mori & Kallunki 3367 (MO). Chagres at El Vigia, Pittier 2378 (US). Gatün, Standley 27180 (US). Frijoles, Moe 27560 (US). Gamboa, Tyson 1521 (MO). COLON: Tumba Vieja, Dodge et al. 16935 (MO). : Río Tuquesa below Quebrada Venado, Bristan 1033 (MO). Río Pirre, Bristan 1314 (MO). ws uceca, Bristan 1441 (MO). Río Chico, Duke & Bristan 450 (MO). Río Tuira and Río Paca, Puts Ha (MO). Río Pirre, 10 mi S of El Real, Duke 5450 (MO). Río Balsa, Duke 8715 (MO). Río Sambú, Duke 9306 (MO). Cerro Pirre, Duke & Elias 13813 (MO, UC, VEN). Río Morti, Duke 14134 (MO). Trail p dbi Duke 14328 (MO). Canasas and Sabalo, Duke 14460 (MO). Río Tuira, Duke 14548 (MO). rto Sta. Dorothea, Dwyer 2275 (MO). Río Cuasí, Kirkbride & Bristan 1439 (MO). Yaviza, cuni er al 147 (MO). Río Paya, Stern et al. 365 (MO, US). Los sANTOS: Río Pedregal, 25 mi SW of Tonosi, Lewis et al. 2922 (COL, MO, UC, VEN). PANAMA: Río La Maestra, Allen 22 MO). Near Jenené, Duke 3851, 3889 (both MO). El Llano to Río Mamoní, Duke 5596 (MO). Piria- Cañasas trail near Piria, Duke 14328 (MO). Fort Lorenzo, Ebinger 458 (MO). Río Chagres, 5 mi SW of Cerro Brewster, Lewis et al. 3443 (MO). 25 km NE of Cerro Azul on Río Piedras, 550 m, Mori 16713 (MO). Río Cuadí, Duke et al. 3658 (MO, UC, N). Cangandé Mandinga Airport Road, Duke 14789 (MO). Rio Diablo and Rio Acuatí, Duke M Isla Pino, near Mulatupo, Elias 1702 (MO, UC). Playon Chico, Mainland, Gentry 6413 (MO). pia opposite Achituppu, Lewis et al. 132 (GH, K, MO, NY, UC, US). Alligandi, Lewis et al. 143 (GH, K, MO, UC, US). Trail to inland village of Armila, 3-8 km SW of Puerto Obaldía, Mori et al. 6800 (MO). Playon Chico, Stier 157, 177 (both MO). 7. Gonzalagunia veraguensis Dwyer.?! TYPE: Panamá, Mori 6673 (MO, holotype). Shrubs to 2 m tall, the twigs terete, golden velutinous, the nodes well spaced. Leaves elliptic or elliptic falcate, 5-25 cm long, 3-9 cm wide, deltoid or acute at the apex, acuminate, the acumen triangular, ultimately acute, cuneate to acute at the base, the costa prominulous above, prominent beneath, the lateral veins ca. 14, arcuate, not forming a distinct marginal vein, prominulous, usually with 1 short, soon branching or evanescent smaller vein between a pair of lateral veins, the numerous smaller veins arising from the veins often in a subperpendicular fashion, the intervenal areas spreading reticulate, rigidly papyraceous or subcori- aceous, discolorous, somewhat lustrous above, with a few scattered hairs on the costa and the veins above or in young leaves scattered over the upper surface, densely shaggy villose beneath, the hairs slender, appressed, to 2 mm long, more 51 Gonzalagunia veraguensis Dwyer, spec. nov. Frutices ad 2 m p ramulis angulatis em appresso-ferrugineis. Folia lanceolata, 15-28 cm longa, 4.5-8.5 cm lata, enis pie bus 10-16 e EE ad T m latis, 1.02. 5 cm distantibus; bracteis ad 0.5 cm longis. Flores sessiles; calyce lobis subulatis, 1.5-2.0 mm longis: corolla tubo cylindrico, ad 7.5 mm longo, extus ciliis Yum appressis, lobis ^ ca. 3 mm longis; antheris ca. 1.8 mm longis; stigmatibus 4 inaequalibus, ad 0.5 m longis. Fructus non visi. 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 abundant on the costa and the veins; petioles 0.3-3.2 cm long, lignose, stout, pubescent; stipules often persistent, ovate or triangular, to 1.5 cm long, to 0.6 cm wide, acute, glabrous outside except for a densely pubescent median keel which arises from a densely pubescent broad basal band ca. 1.5 mm deep. /nflo- rescences terminal, solitary, spikelike, 15-60 cm long, to 2 cm wide, the flowers in well spaced aggregates; bracts linear, usually as long as the pedicels. Flowers sessile or subsessile; hypanthium rotund, ca. 2 mm long, sericeous, the calycine cup to 0.7 mm long, glabrous, often with a few irregular glands in the sinuses, the latter triangular or subulate, to 1.8 mm long, acute; corolla white, the tube narrowly cylindrical, to 7 mm long, thickly petaloid, the hairs outside dense, white, appressed, puberulent to sparsely ciliate within, the lobes 4, oblong, ca. 3 mm long, obtuse; stamens 4, the anthers ca. 1.8 mm long, obtuse, the filaments slender, almost equaling the anthers, inserted 1.5-3.0 mm above the base of the tube; ovarian disc annular, ca. 0.2 mm deep, the style ca. 3 mm long, the stigmas 4, unequal, digitiform, to 0.5 mm long. Fruits not seen. Gonzalagunia veraguensis is known only from Panama. It is readily recog- nized by its many veined shaggy villose (beneath) leaf blades and its very short corolla. VERAGUAS: Valley of Río Dos Bocas between Alto Piedra above Santa Fé and Calovébora, 350- 400 m, Croat 27462, 27733 (both MO). Near Escuela Agricola Alto de Piedra, Mori 6673 (MO). 43. GUETTARDA Guettarda L., Sp. Pl. 991. 1753; Gen. Pl., ed. 5: 428. 1754. TYPE: G. speciosa L. Matthiola L., Sp. Pl. 1192. 1753, not R. Brown, (1812. Nomen conserv., Cruciferae). TYPE: M. SC Hale ae P. Br., Civ. Nat. Hist. Jam. 205. 1756, nomen rejic. not J. Ellis ex L. 1759 (Nomen conserv., Styracaceae). Laugeria e. Enum. Pl. Carib. 2. 1760. type: L. odorata Jacq. Trees or shrubs. Leaves occasionally ternate, usually petiolate; stipules de- ciduous. Inflorescences axillary, cymose, the flowers occasionally solitary, usu- ally secundly disposed or congested; bracteoles present or absent. Flowers with the hypanthium ovoid or globose, the calycine cup tubular, cupular or campan- ulate, truncate, edentate or 2—9-toothed: corolla funnelform or salverform, the tube elongate, the lobes 4-9, imbricate, 2 of them exterior; stamens 4-9, inserted in the tube, the anthers slender, dorsifixed, included, the filaments frequently wanting; style slender, the stigmas capitate or 2-lobed, the ovary 2-9-celled, the ovules | per cell, pendulous. Fruits drupaceous, globose, ovoid or oblong, thinly carnose, usually with a fine velvety indumentum, often acutely angular, the pu- tamen ligneous or bony, the testa membranous. Guettarda, a genus of about 80 species, occurs throughout tropical America. It also occurs in New Caledonia (fide Willis 1966). a. Inflorescences with cymes 3-flowered; peduncles s - 5. G. ramuliflora aa. i odd with cymes more than 3-flowered: ds slender. b s with lateral veins 6-10 C. Fruits obovate we. less than 2.2 cm long |... Loss 6. G. sanblasensis cc. Fruits rotund, 7.0-1.5 cm in diam 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 199 Leaves 10—16 cm long; petioles to 2 cm long __ 2. G. foliacea Leaves 2.0-5.5 cm long; petioles to 0.7 cm long ------------------ 4. G. odorata bb. "di with lateral veins 10— y ves glabrate above; fruits compressed rotund, ca. 2 cm long, not sharply . macrosperma ee. D. villose or villosulose above; fruits oblong, to 0.8 cm long, sharply angle G. chiriquiensis 1. Guettarda chiriquiensis Standley, Ann. Missouri Bot. Gard. 25: 838. 1938. TYPE: Panama, White & White 22 (F, holotype; MO, isotype).—Fic. 45. Trees to 30 ft tall, the branches terete or angular, the twiglets angular, stout, densely puberulent, the nodes usually well spaced. Leaves ovate rotund to lan- ceolate, 6.5-18.0 cm long, 2.5-12.5 cm wide, acute or somewhat rounded apically, frequently cuspidate, cuneate to occasionally obtuse or rounded at the base, rarely truncate, the costa prominulous above, prominent beneath, the lateral veins 10-11, prominulous, especially beneath, those in the lower 4 of the blade strict but marginally arcuate, those of upper ^ markedly arcuate, stiffly papyraceous to subcoriaceous, usually discolorous, villose or villosulose or both; petioles to 4.5(-10.0) cm long, usually rigid, densely pubescent; stipules often persistent at the ends of the branchlets, ovate or oblong, 12-13 mm long. Inflorescences ax- illary, 1 per axil, about as long as the petiole of the adjacent leaf, occasionally longer, the peduncles 1-2 cm long, densely golden pubescent, bifurcating, the branches 2.5-4.0 cm long, the flowers unilaterally disposed on the branch, usually 8-12 per branch. Flowers sessile; hypanthium oblong, ca. 2.5 mm long, the ca- lycine cup ca. 1 mm long, the 4 teeth deltoid, ca. 0.5 mm long, densely pubescent outside, corolla white or purplish white, the tube narrowly cylindrical, ca. 12 mm long, ca. 1.5 mm wide in the middle, densely golden hirsute outside, the hairs deflexed, glabrous within, carnose, the lobes 4, oblong, the margin conspicuously lobed and scalloped, the whole ca. 6 mm long, and 5 mm wide at the base, glabrous within, puberulent outside; stamens 4, the anthers oblong, ca. 2.8 mm long, ca. 0.7 mm wide, obtuse, dorsifixed, the filaments plane, ca. 0.35 mm wide, attached just below the orifice; stigmas 2, oblong, 1 mm long, subconcave adax- ially, the style 15-18 mm long. Fruits sessile, oblong, to 0.8 cm long, ca. 0.5 cm wide, the remnants of calycine cup persistent as an apiculum, sharply 4-angled and 4-sulcate, drying grey-green, the pubescence dense. Guettarda chiriquiensis is known from Panama and Colombia but may be expected in adjacent Costa Rica. The solitary Colombian collection cited below occurs at a much lower elevation than the Panama collections. CHIRIQUÍ: Between Cerro Punta and Las Nubes, 1800-2100 m, Croat 26394 (MO). Valley of Río Chiriquí Viéjo, N of Volcán City, 5200-5600 ft, Duke 9037 (MO). El Hato, 5000-6000 ft, Ebinger 822 (MO). Boquete, 1400 m. Maasola 28 (MO). 27 km NW of El Hato del Volcán, 5000-5300 ft, Mori & Bolten 7189 (MO). Cerro Punta, 6000 ft, Tyson 5820 (MO). Above Jaramillo Arriba, N slopes of Cerro Palo Alto, 4700 ft, Webster 16684 (MO). Volcán de Chiriquí, 1700 m, Woodson & Schery 508 MO). coci É: El Valle, Correa 316 (MO). Cerro Pilon, Duke 12111 (MO); Dover 10566 (MO); Dwyer & Correa 7946, 7975 (both MO): Dwver & Lallathin 8618 (MO). La Mesa, Dwyer & Nee 11936 (MO); Gentry 5637, 6837 (both MO). El Potroso, Folsom & Collins 1554 (MO). Cerro Pilón, Mori 6555 (MO). La Mesa, Nee & Dwyer 9180 (MO). Cerro Pilón, Porter et al. 4416 (COL, DUKE, MICH MO. NY. SCZ, UC, VEN). PANAMA: Cerro Campana, Correa & Dressler 368 (MO); Croat 12158 (MO); Dwyer & Kirkbride 7835 (MO); Gentry 4917 (MO): McDaniel 6857 (MO); Mori & Kallunki 3572 (MO). COLOMBIA: CHOCO: Rio San Juan just above Istmina, 100 m, Gentry & Fallen 17650 (MO). 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 aes 45. Guettarda chiriquiensis Standley.—A. Flowering branch (x1⁄).—B. Flower —C. Ovary sections (x3). [After Dwyer & Neë 11936.].—D. Fruiting branch "em [After D s 5637.] 2. Guettarda foliacea Standley, Contr. U.S. Natl. Herb. 18: 139. 1916. TYPE: Panamá, Pittier 4031 (US, holotype; isotype, F). Trees, to 15 m tall, or shrubs, the twiglets terete, smooth, glabrous, ultimately pubescent, often with spreading lateral branches. Leaves often crowded at the apex of the twiglets, obovate oblong to elliptic, 3-17 cm long, 1.2-6.5 cm wide, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 201 widely deltoid or rounded at the apex, the acumen short or absent, less than 1 cm long, basally obtuse or truncate, rarely acute or subcordate, usually inequi- lateral, the costa prominulous beneath, the lateral veins 8—10, arcuate, the veinlets pinnatiform, stiffly membranous or chartaceous, concolorous or discolorous, the hairs above white or scattered, the hairs beneath short or long, appressed or spreading, often strigose, appearing shaggy, the axils often barbate; petioles 0.8— 2.0 cm long, slender, puberulent; stipules not seen. /nflorescences axillary, usu- ally spreading, the peduncle slender, to 3 cm long, ca. 0.5 mm wide, pubescent, the few branches short and often secundly disposed; bracts deciduous, oblong, to 6 mm long, ca. | mm wide, sericeous, the calycine cup cupuliform, scarious, usually slightly longer than the hypanthium, sericeous, truncate, the teeth absent; corolla white, the tube narrowly cylindrical, to 1.5 cm long, ca. 1 mm wide in the middle, sericeous outside, glabrous within, the lobes 4-5, oblong, ca. 3 mm long, obtuse, thinly carnose, sericeous outside, the anthers subsessile, narrowly ob- long, 3 mm long, attached ca. 2 mm below the mouth; style slender, the stigmas compressed, capitate rotund, ca. 0.25 mm long. Fruits sessile, rotund, 0.8— 1.5(—2.0) cm in diam., finely velutinous, red at maturity. Guettarda foliacea occurs in Panama and Colombia. ‘‘Guyababa’’; Espino." BARRO COLORADO ISLAND: Bailey & Bailey 202, 611 (both F); Croat 6029, 6061, 6114, 6240, 6540, 7405, 7422, 7670 (all MO), 1145 (F), 11764 (MO), 15094 (F), 15160 (MO): Dwyer 1416 (MO); Ebinger 267 (MO), 390 (MO, US); Foster 951, 1002 (both F); Knight 69-1, 69-40 (both MO); Shattuck 8 (F); Starry 222 (F); Zetek 3580 (MO). CANAL ZONE: Road S-11, NW of Escobal, Croat 12471 (MO). Pipeline Road, Croat 15094 (MO). Coco Solo, Gentry 6191 (MO). Barbacoas Railroad Station, Hayes 167 (F). Pipeline Road, Mori & Kallunki 2051 (MO). Trinidad River, Pittier 4031 (F , US). CHIRIQUI: Progreso, Cooper & Slater 227 (US), 236 (F, US). DARIEN: Río Ucurgantí, Bristan 1175, 1178 (both MO). S of El Real, Duke 5036 (MO). Río Chucunaque between Río Membrillo and Río ee iE cuti, Duke MO). Río Sabana, above Santa Fé, Duke 14085 (MO). Piria-Cañasas uw il nea Duke 14310 (MO). Between Canasas and Sábalo, Duke 14478 (MO). El Real Qu adig "Trapiche; Duke & Bristan 322 (MO). Campamento Buena Vista, Stern et al. 957 (US). 2 mi E of Santa Fé, Tyson et al. 4845 (MO). PANAMÁ: Río Chepo, Río Bayano, Duke 3987 (MO). San José Island between Bodega Bay and E Mata Puerco, Duke 12552 (MO). Llanos of Río Mar, Dwyer 1799 (MO). Río Pasiga, Gentry 2287 (MO). Between Canita and dam site, Gentry & Tyson 1704 (MO). El Llano, road to aae María, ati 6873 (MO). vERAGUAS: Rio Trinidad, below Rio de Jesús, Dwyer 1321 (MO). go, Stern et al. 996 (MO, US). coLoMBiA: CHOCO: Behind beach, El Valle, sea level, Gentry & Falle n 17231 (MO). 3. Guettarda macrosperma Donnell Smith, Bot. Gaz. 18: 204. 1893. TYPE: Gua- temala, Heyde & Lux 3160, not seen. Trees to 6 m tall, much branched, the twiglets terete, glabrous to pubescent, the nodes moderately well spaced, the lenticels minute, often inconspicuous. Leaves often crowded toward the apex of the twiglets, elliptic oblong, oblong rotund or occasionally pyriform, 12-20 cm long, 6-13 cm wide, deltoid to obtuse at the apex, acuminate, the acumen short, the apex of the blade often eroded, basally obtuse to widely cuneate, occasionally acute at the base, the costa prom- inent beneath, the lateral veins ca. 12, prominent, rigid, often light in color and contrasting with the rest of the blade, at first angular ascending then arcuate toward the margin, the ultimate veinlets vaguely lineolate, stiffly chartaceous, often brittle when dry, concolorous, glabrous or pubescent on the costa and the veins above, beneath with the hairs short, white, strigose, the vein axils often shortly barbate; petioles to 4 cm long, 0.10—0.25 cm wide, stiff, often curved, 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 glabrous to pubescent; stipules deciduous, not seen. Inflorescences axillary, the peduncles to 1.5 cm long, to 1.5 mm wide, stiff, the pubescence minute, green, velutinous. Flowers not seen. Fruits sessile, compressed rotund, ca. 2 cm wide, the pubescence minute, velutinous, yellow green or green when dry, delicately ribbed, the ribs disposed as faint and scarcely elevated lines, fruit wall thin; seeds 5—7, in a hard, white, pinkish or molasses-colored pulp. Guettarda macrosperma extends from Guatemala to Panama. The Panama- nian collections, while relatively abundant, lack flowers, making critical identi- fication difficult. Donnell Smith in his original diagnosis describes the mature leaves of G. mac rosperma as "294 x 11⁄ in." with ''6—-7 costal nerves." The fruits are described as ‘‘obovoid oblong (6-7 x 3-4 inches)." The leaves of Pan- amanian material are considerably larger, the lateral nerves are about 12, about double the number listed for the type which was collected at about 3,000 ft at Santa Rosa, Guatemala. The fruits of the Panamanian material do not exceed an inch in diameter while Donnell Smith describes the fruits as 4 inches in diameter. Unfortunately, I did not see the type to verify this incredibly large fruit. Paul Standley identified Wedel 266 as G. macrosperma. It and Carrasquilla et al. 387 have much smaller leaves even though the fruit is like that of the exsiccatae below. About 1⁄2 of the Panamanian material was collected near sea level in the limestone outcrops at Madden Dam. CANAL ZONE: Madden Lake, Croat 38326 (MO). Fort Sherman, Duke & Mussell 6638 (MO). Madden Dam, Boy AN Road, Dwyer & Elias 7504 (GH, MO); Dwyer & ving ieh 20 (MO); Ebinger (MO). Area W of Limón Bay, Gatún Locks, Johnston 1704 (MO). Madden Dam, Boy Scout Road, Kirkbride 42 (MO): Kirkbride & Elias 260 (MO); Lewis et al. 15 (GH, K. MO. NY, UC, US). Pipeline Road, 4 km NW of Gamboa, Nee 7225 (MO). Sotto Caballo, Smith et al. 3318 (F. MO). COLON: Río Boquerón, 3 km from La Pelona, Carrasquilla et al. 387 (MO). Between Garotte and La Guaira, D'Arcy 9349 (MO). Portobelo, Dwyer 5149 (MO). Maria Chiquita, Dwyer E dye: 7772 (GH, C, US). DARIEN: Santa Fé, Td 14085 (F). SAN BLAS: Ustupo, D'Arcy 9534 (MO). Ailigandí River, Duke & Bristan 349 (MO). Río Guadí, Camp Diablo, drill site 22. Duke et al. 3646 (MO). Isla Pino near Mulatupo, 200 ft, Elias 1705 (MO, UC). 4. Guettarda odorata (Jacq.) Lam., Tab. Encycl. Méth. 2: 219. 1819. Laugeria odorata Jacq., Enum. Pl. Carib. 16. 1760. type: Cuba. Jacquin, not seen. Shrubs or trees, to 3.5 m tall, the branchlets terete, distichous, well spaced, the bark drying dull grey, often mottled, occasionally conspicuously lenticellate. Leaves oblong, 2.0-5.5 cm long, 1.0-2.5 cm wide, cuneate to rounded at the apex, acuminate, the acumen vague, basally cuneate, obtuse or vaguely rounded, the costa plane, immersed or prominulous above, prominent beneath, the lateral veins (6—)8-9, arcuate, the veinlets lineolate, irregularly perpendicular to lateral veins, the intervenal areas often mosaic-like, papyraceous, glabrate above, villose or merely puberulent on the costa and the veins as well as the intervenal areas, below the hairs of the veinlets occasionally shaggy: petioles 0.2-0.7 cm long, slender, puberulent or villose, often curved; stipules ovate deltoid, ca. 2 mm long, acute, pubescent. Inflorescences axillary, 1-2 per axil, 3-5 cm long, the pedun- cles 1.5-3.0 cm long, capillaceous or wiry, to 0.5 mm wide, pubescent, producing I cymule at the apex, this usually 5-flowered, or bifurcating and producing a pair of 5-8-flowered cymules. Flowers sessile or subsessile; hypanthium subrotund, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 203 ca. 1 mm long, carnose, the calycine cup ca. 2.5 mm long, carnose, eglandular within, the teeth absent; corolla white or reddish, the tube narrowly cylindrical, to 16.5 mm long, ca. 1 mm wide, puberulent outside, glabrous within, the lobes (4-)5(-6), oblong, 2.5-4.2 mm long, thinly carnose; stamens (4-)5, the anthers linear oblong, 1.5-3.0 mm long, ca. 0.2 mm wide, dorsifixed, subsessile or sessile, occasionally the filaments as long as the anthers, attached immediately below the mouth or at the upper 2⁄4 of the tube, the ovarian disc annular, the style linear, 8-14 mm long with appressed ascending hairs, the stigmas 2(-3?), capitate, ca. 0.2 mm long, slightly wider than the style. Fruits rotund, to 0.7 cm in diam., the persistent calycine cup to 1.3 mm long, the pericarp smooth, wrinkled when dry, minutely grey green, puberulent, the hairs eventually deciduous, Guettarda odorata occurs in Panama and in northern South America. It has also been reported from the West Indies. "Espino Blanco"; ‘‘Huesio Negro." Steyermark (1972) discusses in detail his designation of a Jacquin collection from Cuba as the lectotype. L ZONE: Fort Kobbe, Allen 1407 (F): Duke 3957, 4241 (both MO). Farfan Beach, Dwyer & Hayden n 7532 (GH, MO, US). Barro Colorado Island, Ebinger 267 (MO). Survival School, Curundú, Tyson 1059 (MO). Fort Clayton, Tyson 6779 (MO). Miraflores Lake (?), White 139 (MO). Ancón Hill, Woodson et al. 1331 (F, MO). Fort Kobbe, Woodson et al. 1407 (MO). CHIRIQUI: 6 mi S of Puerto uelles, 0-150 m, Liesner 428 (MO). cocrÉ: Penonomé, Dwyer 2033 (MO). DARIEN: Mouth of Río Yapé Allen 371 (F, MO). Río Pirre, Duke 4962 (MO). Río Cucunatí at Puente Quemado, Duke 8806 (MO). HERRERA: Oct, Ebinger 1062 (MO). La Cabuya, Las Minas, Lao 139 (MO). PANAMA: Pedro Gonzales, Allen 2597 (F). Calzada Larga, Croat 12385 (MO). Rio Mar, Savanna, Duke 12426 (MO). Isla Taboga, Hayes 637 (F). Las Sabanas, Pittier 6702 (F). San José Island, Stimson 5326 (MO). Isla Taboga, Woodson et al. 1482 (F, MO). E of Pacora, Woodson et al. 723 (F, MO). VER- AGUAS: 5 mi N of gea t Río Santa María, Blum & Tyson 616 (MO). 5 mi NE of La Mesa, Blum & Tyson 667 (MO). 2 mi W of Santiago, Transisthmian Highway, Dwyer et al. 7544 (GH, MO). Quebrada S of Santa Fé, 450 m, Nee 8040 (MO). San José, Santiago-Santa Fé Road, 400 m, Nee 8170 (MO). 5. Guettarda ramuliflora Beurl., Kongl. Vetensk. Acad. Handl. 132. 1854. TYPE: Panama, Billberg (S). Trees, to 8 m tall or shrubs, the twigs terete, puberulent, ultimately finely velutinous, the nodes well spaced. Leaves ovate oblong or sagittate, 6-9 cm long, 3-5 cm wide, acute to widely deltoid at the apex, acuminate, the acumen short, somewhat vague, obtuse to widely deltoid at the apex, acuminate, the acumen short, somewhat vague, obtuse to widely cuneate at the base, the costa promi- nulous above, prominulous beneath, the lateral veins 6-8, arcuate, with short, white, appressed, striate, densely appressed hairs on the costa and the veins beneath, the intervenal areas lineolate but not conspicuously so, concolorous, membranous; petioles slender, 1-3 cm long, ca. 0.1 cm wide, appressed villosu- lose; stipules not seen. /nflorescences axillary, to 3.5 cm long, the peduncle stout, to 1 cm long, velutinous, the flowers few, the cymes 3-flowered (fide Standley 1928) crowded at the end of the peduncle. Flowers sessile, villose; hypanthium broader than long, 1 mm long, the calycine cup turgid, cupuliform, ca. 3 mm long, truncate, edentate; corolla white, the tube narrowly funnelform, to 2.2 cm long, to 0.2 cm wide, carnose, villose outside, glabrous within, the lobes 5, ob- long, to 5 mm long, to 3.5 mm wide, obtuse, the margin with occasional shallow 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 notches; stamens 5, sessile, the anthers narrowly oblong, obtuse, acutely sagittate at base, attached ca. 3 mm below the mouth, the ovarian disc cylindrical, 1.3 mm long, villosulose at the base, truncate, the style linear, ca. 7.5 mm long. Fruits not seen Guettarda ramuliflora is known only from Panama. PROVINCE UNKNOWN: Billberg (S). CANAL ZONE: Devil's Beach, woods back of beach, NW part of Canal Zone, Johnston 1717 (MO). 6. Guettarda sanblasensis Dwyer.? Type: Panama, Duke et al. 3646 (MO, holo- type). Trees 15-20 m tall, the branchlets terete, ultimately puberulent, the nodes moderately well spaced. Leaves elliptic or elliptic rotund, 5-14 cm long, 4-10 cm wide, toward the apex, rounded, deltoid, subacute or acuminate, the acumen inconspicuous or to 1.5 cm long, ultimately acute, at the base cuneate or widely cuneate, the costa plane above, prominulous beneath, the lateral veins 8-10, arcuate, the venules pinnatiform, rigidly chartaceous, concolorous, green, shin- ing, the hairs on both sides minutely strigose, those on the veins appressed, short, velvety; petioles to 3 cm long, ca. 0.12 cm wide, stiff; stipules not seen. /nflo- rescences axillary, the peduncle to 3 cm long, pubescent, the branches to 3 cm long. Flowers not seen. Fruits obovate oblong, to 2.2 cm long, to 1.2 cm wide, subtruncate at the apex, minutely velutinous, yellow green when dry, attenuate toward the apex, the calycine cup ca. 2 mm long, ca. 1.5 mm wide. Guettarda sanblasensis is known only from Panama. It is distinguished by its apically subtruncate fruit. : Headwaters of Río Cuadí, Camp Diablo, drill site 22, N 82.2, E 87.8, 273 ft, Duke et al. 3646 (MO. holotype). 44. HAMELIA Hamelia Jacq., Enum. Pl. Carib. 16. 1760. LECTOTYPE: H. erecta Jacq. = H. patens Jacq. Hamellia L., Sp. Pl., ed. 2, 246. 1762, orth. mut. d Jacq. Tangaraca Adans., Fam. E t 147. 1763. TYPE: not i Lonicera Adans., Fam. Pl. 11: 157. 1763. Not Pee x 1753) TYPE: not indicated. Duhamelia Pers., Syn. Pl. 2 Ps 1805. TYPE: not indicated. Tepesia Gaertn., Fruct. Sem. Pl. 3: 72. 1805. type: T. dubia Gaertner. Shrubs or small trees. Leaves opposite or in whorls of 3-5, petiolate; stipules free, entire or 3-lobed, small. /nflorescences terminal, of thryses, dichasia or cymes, the floral axes often spreading, the flowers usually secund; bracts decid- uous. Flowers with the hypanthium 5-lobed, the lobes often reduced, persistent 5 Guettarda sanblasensis Dwyer, spec. nov. Arbores 15-20 m altae. Folia elliptica vel elliptico- rotunda, 5-14 cm longa, 4-10 cm lata, venis lateralibus 8-10 arcuatis, rigido-chartacea; petiolis ad 3 cm longis. at scentiae in gui axillares pedunculo ad 3 cm longo. Flores haud suppetunt. Fructus obovato-oblongi, ad 2.2 cm longi, ad 1.2 cm lati, minute velutini in sicco luteo-virides cupula calycis 2 mm longa, 1.5 mm lata Porn. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 205 in fruit; corolla tubular or inflated, orange, red or yellow, the lobes 5, imbricate; stamens 5, the connectives produced apically, the filaments attached at the base of the corolla; ovary (4—)5-celled, the style slender, the stigmas 1-5. Fruits fleshy, the ovarian disc often persistent and forming a truncated cone; seeds numerous, plano compressed, foveolate. Hamelia ranges from southern Florida through Mexico, Central America, the West Indies and South America. There are about 16 species in the genus. Literature: Elias, T. 1976. A monograph of the genus Hamelia (Rubiaceae), Mem. New York Bot. Gard. 26(4): 81-144. Wernham, H. F. 1911. A Revision of the Genus Hamelia. Jour. Bot. London 49: 206-216. a. Corolla funnel shaped, yellow (sect. Amphituba lla tube 1.0-1.3 cm long; fruits 0.5—0. 7 em long nul l. H. axillaris bb. Corolla tube 1.8-5.0 cm long; fruits 7-16 | Calycine lobes 3-5 mm long: stipules subulate ------------------------ 2. H. calycosa cc. Calycine lobes 0.2-2.5 mm long; stipules triangular to ovate d. Corolla tube 4-7 mm broad at the apex: gredi > 10-25- flowered . FRNETCILURNUES . H. xeroc "carpa dd. Corolla 10-20 mm broad at the apex: inflorescence 4_15-flowered TT 3. H. macrantha aa. E rae pe yellow, orange or red (Sect. Hamelia). Lateral v airs eins 3-5 Inflorescence a much reduced dichasium, flowers usually 2-5; leaves 0.8-2 r .8) cm broad... 2 aa == . H. m ff. Inflorescence a sciences dichasium « or cyme, flowers 3-8; leaves 2-7 2m road PEPPER cM EE . H. rovirosae ee. Lateral veins 5-11 pairs Floral parts and leaves densely villous. Leaves with lateral veins 3-5 pairs; seeds 1.0-1.2 mm long ---------------- See AS LS SERS ee ne Oe 6. H. rovirosae hh. Leaves with lateral veins 7-10 pairs; seeds 0.6— s. 9mm long .. 5. H. patens gg. i m ge leaf undersides glabrous to puberulen Flow 12-15 mm long; leaves with 11-16 a “of lateral veins; corolla . H — alae, SOS ey NEHME RT rs . magnifolia ii. Flowers 15- 25; mm n long; leaves with 7- 10 pairs of lateral veins; >: olla yellow to red - I - - . H. patens 1. Hamelia axillaris? Swartz, Nov. Gen. Sp. Pl. Prodr. 46. 1788. LECTOTYPE: (Elias, 1976): Jamaica, Swartz (S). Shrubs to 4 m tall, the branchlets terete to angular, glabrous to puberulent. Leaves opposite or rarely ternate, elliptic, narrowly elliptic or obovate, 5-17 cm long, 2-7 cm wide, acuminate at the apex, often conspicuously so, attenuate at the base, glabrous above, often puberulent beneath and barbate in the vein axils, the lateral veins 8-11, arcuate, chartaceous to membranous, glabrous or sparsely puberulent; petioles 1—4(—6.5) cm long, occasionally densely puberulent; stipules triangular, 2-6 mm long. /nflorescences axillary and terminal, of few- to many- flowered compound dichasia, often disposed as scorpioid cymes; peduncles to 53 For a list of synonyms see Elias (1976). Only this name has been used for Panamanian material. 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1.5 cm long, the branches 1.5-4.5 cm long, spreading; bracts ovate, ca. 0.5 mm long, puberulent. Flowers sessile or on pedicels to 1 mm long, sparsely puberu- lent; calycine lobes to 1.5 mm long, puberulent, often densely so; corolla yellow, the tube infundibular, expanding at the apex, to 11.5 mm long, the lobes ovate, to 2 mm long, puberulent to glabrous; anthers 5-7 mm long, the apiculum ca. 0.5 mm long, the filaments to 5 mm long, the style slightly shorter than the tube, the stigmas ca. 1.2 mm long. Fruits globose or subglobose, 5-7 mm long, ultimately black; seeds ca. 1 mm long. Hamelia axillaris is a weedy shrub extending from Mexico through Central America to northern South America. It also occurs in the West Indies. CANAL ZONE: Road S-10, N of Escobal near junction of Road S-1, Croat 12484 (MO). Río ndosa, Navy Reservation, 8 km N of Gamboa, Dressler 4087 (MO). Camp Pina, Fort Sherman, Duke 4398 (MO). Madden Dam, Boy Scout Road, Dwyer & Elias 7491 (GH, MO, UC, US). N base of Mindi Hills, Johnston 1637 (MO). BARRO COLORADO ISLAND: Croat 6299, 10940, 11862, 11901 (all MO); Hayden 150 (MO); Starry 284 (MO). COLON: Portobelo, Ebinger 118 (MO). DARIEN: Rio Tu- queza, below Quebrada Venado, Bristan 1034 (MO). Rio Aruza, Bristan 1367 (MO). Rio Balsa be- tween Quebrada Chusomocatre and Rio Areti, Duke 8718 (MO). Quebrada Madkia off Rio Pucro pay Pucro, Duke 13013 (MO). Tumaganti, Duke 14160 (MO). Rio Tuira between Rio Punusa and o Mangle, Duke 14635 (MO). Between Rio Perrecénega and Rancho Frio, Duke & Elias 13860, 13867 (both GH, MO, US). Between Paya and Palo de Los Letras, Duke & Kirkbride 14034 (MO). Rio Tuquesa, Middle Tuquesa Mining Co. Camp called Charco Peje, Mori 7090 (MO). Between Rio Escucha Ruido and Río Tuira, Stern et al. 578 (MO). Cana, 1750 ft, Stern et al. 670 (MO). Campa- mento Buena Vista, Rio Chucunaque, Stern et al. 828 (MO). 3 mi E of Santa Fé, Tyson et al, 4715 (MO). Los SANTOS: Los Toretos, Dwyer 242 ). PANAMÁ: Piria-Cañasas Trail near Piria, c m, Duke 14331 (MO). Cerro Azul, ca. 2000 ft, Dwyer 2072 (MO). Within 2 mi of Achiote, Folsom 3871 (MO). Dam site near Río dee DR d Chepo, Gentry & Tyson 1676 (MO). Cerro Cam Mass et al. 1731 (MO); Mori & Kallunki 1937 (MO): Webster & Breckon 16510 (MO). Co Memorial Laboratory, ca. 25 km NE of Cerro Azul, Río Piedras, 550 m, Mori & Kallunki 3459 (MO). 2. Hamelia calycosa J. D. Smith, Bot. Gaz. 12: 132. 1887. TYPE: Guatemala, Tuerckheim 454 (US, holotype; NY, isotype). H. chiapensis Brandegee, Univ. Cal. Publ. Bot. 6: 71. 1914. rype: México, Purpus 7030 (F, MO, N H. boyacana Standley, Field Mus. Nat. Hist., Bot. Ser. series 11(5): 213. 1936. TYPE: Colombia, Lawrance 503 (NY, holotype, M : H. d a Field Mus. Nat. Hist... Bot. Ser. 13. No. 364: 96. 1932. rype: Perú, Klug 3664 , holotype) Shrubs or trees to 10 m tall, the twiglets terete, smooth, glabrous, ultimately puberulent, branching freely, the nodes well spaced. Leaves oblong, often 3—4- nate at the ends of twigs, lanceolate or trapeziform lanceolate, 1.5—11.0 cm long, 0.9-3.5 cm wide, tapering acutely at the apex, acuminate, the acumen ca. 0.7 cm long, acute to cuneate at the base, the costa grooved, plane or prominulous above, the lateral veins 4-7(-9), arcuate, membranous or papyraceous, glabrous above and beneath except occasionally pubescent on the costa beneath, often barbate in the vein axils beneath; petioles slender, 0.8—2.0 cm long, wiry, glabrate; stip- ules subulate, to 2.5 mm long, puberulent. Inflorescences terminal, rarely axil- lary, the scorpioid cymes lax and spreading, the flowers many in 3-4 fascicles; peduncles to 2 cm long, often minutely puberulent, the floral axes with up to 4 flowers, 0.6-4.0 cm long; bracts triangular to subulate, to 2 mm long, glabrous. Flowers secund, the pedicels to 5 mm long; hypanthium oblong, puberulent, the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 207 calycine cup scarcely measurable, the lobes oblong, to 1.5 mm long, fleshy, pu- berulent; corolla pale orange or yellow, striped maroon, the tube somewhat in- fundibuliform, 30 mm long, constricted narrowly at the base, the base ca. 1.5 mm wide, ca. 7.5 mm wide in the middle of the tube, ca. 1 cm wide at the mouth, puberulent outside, glabrous within, the lobes ovate, to 6 mm long; stamens 5, the anthers linear oblong, to 2 cm long, ca. 1 mm wide, the connective extended apically as a rotund petaloid appendage, ca. 0.75 mm long, sagittate at the base, the filaments plane, ca. 4 mm long, attached ca. 6 mm from the base of the tube; style to 15 mm long, the stigmas 5, connate into a mass, to 1.5 mm long. Fruits cylindrical, 7-15 mm long, 4-6 mm wide, the ovarian disc conical and truncate; seeds to 1.2 mm long, minutely foveolate. Hamelia calycosa extends from Mexico to Peru. : N rim of El Valle, Allen 1785 (GH. MO, NY). Cerro Pilón, 700-900 m, Duke 12151 (MO). "Eon Pilón, Duke & Dwyer 13898 (COL, MO, E UC). N of El Valle, Lewis et al. 1797 (COL, DUKE, K, MO, UC). La Mesa, 850 m, Mori 6599 (MO). PANAMÁ: Río Cascada, 300—400 m, Correa & Dressler 1652 (MO). vERAGUAS: Alto Piedra, Santa Fé, 1200 m, Lao 515 (MO). 3. Hamelia macrantha Little, Caribbean Forest. 9: 274. 1948. Type: Ecuador, Spruce 6193 (K, holotype, not seen). H. grandiflora Spruce, in Wernham, J. 49: 209. 1911. LECTOTYPE: Brazil, Spruce 6193 (K, holotype; photo, MO); SYNTYPE: eun pM Triana 1759 (F, frag.). Not L’Heritier, 1788. Shrubs or trees to 5 m tall, the branchlets terete, ultimately angular, minutely puberulent, the nodes well spaced. Leaves ternate toward the apex, broadly oblong to ovate, 13-23 cm long, 4-15 cm wide, rounded, obtuse or widely deltoid at the apex, rounded, subtruncate to cuneate basally, the costa plane to promi- nulous above and beneath, the lateral veins 11-15, membranous to chartaceous, glabrous above, glabrous to rufous puberulent beneath; petioles 6.0-9.5 cm long, glabrous or puberulent; stipules ovate, 3.1-3.5 mm long, acute, glabrous or rarely pubescent. /nflorescences terminal or axillary, the dichasia many flowered; pe- duncle terete, to 8 cm long, the floral axes to 2 cm long; bracts ovate, to 0.7 mm long, acute, glabrate to villosulose. Flowers sessile or on pedicels to 2 mm long; hypanthium 3-5 mm long, the calycine cup scarcely measurable, the lobes tri- angular to ovate, to 1.3 mm long, acute, glabrous to tomentose; corolla yellow, the tube infundibular, 2-3 cm long, constricted basally, glabrous, the lobes ovate, to 4 mm long, glabrous; stamens included, the anthers to 20 mm long, the con- nective produced at the apex, ovate, ca. 1 mm long, sagittate basally; style to 20 mm long, the stigmatic lobes linear, 5-8 mm long. Fruits ellipsoid, to 1.5 cm long, red to finally lustrous black at maturity; persistent ovarian disc conical and truncate; seeds to 1 mm long, delicately foveolate. Hamelia macrantha ranges from Costa Rica south to Ecuador. CANAL ZONE: Pipeline Road, Agua Salud, Kennedy & A dine s 1894 (MO). COLON: Río Gua Correa et al. 1898 (MO). cociÉ: N of El Valle, road to La Mesa, 1000 m, Allen 3724 (MO). El d near La Mesa, Correa & Dressler 1000 (MO). Between Cerro “pilón and El Valle, Duke & Dwyer 13939 (MO). El Valle, ca. 2200 ft, Dwyer 1913 (MO). Cerro Pilón, 900 m, Dwyer & Lallathin 8625, stipe (both MO). El Valle, ca. 1000 m, Gentry 5694 (MO). Cerro Pilon, Lallathin 10-4, 28-C (both O). PANAMA: El Llano Carti- Tupile Road, 12 mi above Panamerican Highway, Liesner 1236 (MO). 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 4. Hamelia magnifolia Wernham, J. Bot. 49: 210. 1911. TYPE: Guatemala, Stand- ley 64569 (F, holotype; NY, US, isotypes). Shrubs or small trees, to 6 m tall, the branchlets angular to tetragonal, gla- brous, the nodes well spaced. Leaves rarely ternate at the tips of the branchlets, ovate to ovate elliptic, occasionally puberulent; petioles 1.0-5.5 cm long; stipules deltoid, 4-7 mm long, to 2 mm wide at the base, glabrate. Inflorescences terminal, of compound dichasia, often appearing cymose, the floral clusters numerous and many flowered; peduncle to 3 cm long. Flowers with the calycine cup to 2.5 mm long, the lobes deltoid, ca. 0.6 mm long or shorter, puberulent marginally; corolla yellow, the tube 7.5-12.0 mm long, glabrous except often minutely puberulent at the base, the lobes to 1.5 mm long, about as broad at the base; anthers narrowly oblong, 6-8 mm long, the connective produced apically as an ovate or deltoid appendage, ca. 1 mm long, falcate and ultimately acuminate, basally sagittate. Fruits oblong, to 1 cm long or longer; persistent ovarian disc ca. 1 mm long. Hamelia magnifolia is known only from Costa. Rica and Panama. Kirkbride & Duke 757 list the common name "Ojo movado.’ OCAS DEL TORO: Gualaco, Cerro Galera Chorca, Allen 5055 (F). CHIRIQUí: San Bartolo Arriba, W of Puerto Armuelles, Croat 26705 (MO). Chiriquicito-Calderas Trail between Quebrada Higueron and Gutierrez, Kirkbride & Duke 757 (MO). Puerto Armuelles, Woodson & Sche ery 845 (MO). San Bartolomé Peninsula, Woodson & Schery 868 (MO). DARIEN: Tres Bocas on Río Cuasí. Kirkbride 1361 (MO). VERAGUAS: Isla de Coiba, pem 1565 (MO). 5. Hamelia patens? Jacq., Enum. Pl. Carib. 16. 1760. rype: Dominican Republic, Jacquin, not seen. Shrubs or small trees, the branchlets glabrous to sparsely villous. Leaves opposite or usually 3-nate or 4-nate (rarely 5-nate), elliptic, ovately elliptic or obovate elliptic, 4-18 cm long, 2.2-7.8 cm wide, deltoid to acuminate at the apex, rounded to acute at the base, the costa slender, the lateral veins 7-10, usually prominulous beneath, chartaceous to membranous, glabrous to densely villous beneath; petioles 0.5-4.0 cm long; stipules triangular to awl shaped, 2-6 mm long, villous. /nflorescences terminal, solitary or of a few fascicles, the flowers in dichasia, often helicoid, the flowers few to numerous, peduncle glabrate to vil- lous, 0.4—4.0 cm long, the floral axes to 4.5 cm long; bracts triangular, less than | mm long, villous, deciduous. Flowers secund, to 2.5 cm long, sessile or sub- sessile, the hypanthium and calycine tube bell shaped, to 4 mm long, glabrate to villous, the lobes ovate, ca. 1 mm long, glabrate or villous, persistent; corolla orange red to red, 12-20 mm long, 2-3 mm wide, glabrous to villous outside, the lobes ovate, to 2 mm long, glabrous to villous; stamens often exserted, the anthers 8-12 mm long, the terminal appendage ca. 0.5 mm long, the filaments shorter than the anthers, attached in the upper !^ of the tube; style slender, about equaling the corolla tube, the stigmas oblong, about '/s the length of the style; ovary 5-celled. Fruits ellipsoid or rotund, to 1 cm long, black at maturity, the ovarian disc often slightly rostrate; seeds 0.5-1.0 mm long. * Fora list of synonyms see Elias (1976). Only this name has been used for Panamanian material. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 209 Hamelia patens occurs in tropical Florida, throughout the West Indies, Mex- ico, Central America, and in South America south to Bolivia. Key to varieties (adapted from Elias, 1976): a. Leaves villous; hypanthium and calyx villous -------------------------------------- 3 Sa. var. pater aa. Leaves glabrous above; hypanthium and calyx usually glabrous ---------------- Sb. var. Pase 5a. Hamelia patens Jacq. var. patens. `ANAL ZONE: Gaillard Highway, near Summit Garden, Croat 9777 (MO). Road 2C2, Cerro Luisa, Croat 10774 (MO). Albrook, Dwyer 6744 (MO); Dwyer & Hayden 4 (MO); Dwyer & Robyns 11 MO): Ebinger 847 (MO). Road K2 off road to Cocoli, Kirkbride 80 (US). Farfan Beach, Kirkbride & Elias 72 (MO). Balboa, Mori & Kallunki 1711 (MO). Cocoli, Riley 133 (MO). Road K-9, Stern et al. 308, 347 (both MO). Albrook, Stimson 5146 (MO). Contractors Hill, Stimson 5242 (MO). Madden m. Stimson et al. 5085 (MO). Curundt, Tyson 1073 (MO). Chiva-Chiva Trail near Miraflores Lake, Tyson 1404 (MO). Cocoli, Tyson 1635 (MO). Barro Colorado Island, White 122 (MO). Miraflores Lake, White & White 188 (MO). CHIRIQUi: Cerro Horqueta, ca. 7000 ft, Blum & Dwyer 2608 (MO). i M quete, Cerro Horque Pate & Hayden 7686 (GH, MO, NY, UC, US). N of San Felix, 36.2 a by road from compu Highway, Mori > d ae 6012 (MO). Finca Lerida to Boquete, c 1300-1700 m, Woodson et di 1106 (MO). cocLÉ: Forest behind Club Campestre, ca. 700 m Duke 13263 (MO). Las iil coco and El Valle, TE et al. 1726 (MO). DARIEN: Without other locality, Bristan 96 (MO). RERA: Road between Las Minas and Pesé, ca. 600 ft, Duke 12335 (MO). Ocu, Ebinger 2002 (MO. "m 5 mi S of Ocu, 1200 ft, Lewis et al. 1668 (GH, MO, UC, US). Los SANTOS: Upper Río Pedregal, 350-400 m, Croat 34490 (MO). Los Toretos, Dwyer 2424 (MO). Pedasi, Dwyer 2483 (MO). Guaybo, Stern et al. 1903 (MO). Tonosi, Tyson et al. 3108 (MO). PANAMÁ: Isla Pedro Gonzales, Allen 2594 (MO). Río Congo, Carrasquilla et al. 208 (MO). 2 mi from U. of Panama, Transisthmian Highway, D'Arcy & D'Arcy 6108 (MO). Isla Chepillo, Duke 10317 (MO). Tocumen, Dwyer 1850 (MO). Lidice, Jaramillo I (MO). Cerro Silvestre, Reece 29 (MO). 5 mi N of Panama City, Stimson 5045 (MO). Arraijan, Woodson et al. 1367 (MO). Isla Taboga, ta et al. 1436 (MO). VERAGUAS: 5 mi N of San M Blum & Tyson 607 (MO). 2-4 mi E of Santiago, Duke 12357 de ESL Dwyer & Epor 7419 (G, MO, US). Montijo, 8 mi S of Santiago, Tyson 6034 (M 5b. Hamelia patens var. glabra? Oerst., Vidensk. Meddel. Dansk Naturhist. For- en. Kjøbenhavn 1852: 42. 1853. rvPE: Nicaragua, Oersted s.n. (C, holotype, not seen).—Fic. 46 OCAS DEL TORO: RR track near Milla 5, Croat & Porter 16462 (MO). Isla Bocas del Toro, Durkee 71-150 (MO). Río Changuinola, Dwyer 42, 2758 (both MO). Quebrada Huron, 300—400 ft, Kirkbride & Duke 445 ere poo das to 5 mi S at junction d Ríos Changuinola and Teribe, Lewis et al. 821 (COL, Changuinola, Lewis et al. 974 (MO, SCZ, UC). Shepherd Island, McDaniel 5160 (MO). E un Wedel 96, 2483, 2930 (all MO). Water Valley, Wedel 664, 2752 (both MO). Chiriquí Lagoon, Wedel 1236, 2598, 2901 (all MO). Nievecita, Woodson et al. 1850 (MO). CANAL ZONE: Cruces Trail and Madden Dam Road, Bartlett & Lasser 16346 (MO). Juan Mina, Bartlett & Lasser 16514, 16531 (both MO). Fort Sherman, Blum & Dwyer 2131 (MO). Pipeline Road near Gamboa, Clewell & Tyson 3252 (MO). Road S-10 N of Escobal, Croat 12436, 12461 (both MO). Farfan Beach, D'Arcy & D'Arcy 6072 (MO). Coco Solo Weather Station, Duke 4282 (MO). Camp Pina, Duke 4418A (MO). Madden Dam, Boy Scout Camp Road, Dwyer 6928, 9/63 (both MO); Dwyer & Elias 7476 (GH. MO, US), 7483 (GH, MO), 7498 (COL, MO, UC), 7499 (MO). Madden Dam, Dwyer & Lallathin 8835 (MO). a San Lorenzo, a 459 (MO). Frijoles, Greenman & Green- man 5156 (MO). Mandingo Road, Hayden 61 (MO). Gatun, Hladik 525, 538 (both MO). Belo Madden Dam pee station. Mesi s.n. (MO). a oy Scout Camp Road, Madden Dam, Kirkbride 40. 41. 51 (all MO). Farfan Beach Road, Kirkbride & Elias 72 (MO). Rio B Bridge, SE of Gamboa, d 4268 (MO). Pipeline Road, 16 mi N of Gamboa, Lewis et al. 5461 (MO, SCZ). 55 For a list of synonyms see Elias (1976). Only this name has been applied to Panamanian material. 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 RE 46. Hamelia patens var. glabra Oerst.—A. Habit (x4). [After Elias 1699.|—B. Fruit- ing branch (x1). [After Elias 7498. | RR 32 mi W of Gamboa, Nee & Mori 3613 (MO). Rodman Marine Base, Nowicke et al. 3579 (COL, MO, UC). Chiva-Chiva Trail, 2 mi N of Miraflores Locks, Tyson 1382, 1400 (both MO). Old Fort enzo, Tyson 1539, 1551 (both MO). 5 mi NW of Cocoli, Tyson 1622 (MO). Gatün RR Station, Tyson 2280 (MO). 1 mi N of Summit Garden, Tyson & Blum 1941 (MO). Farfan Beach, Tyson et al. 3177 (MO). Miraflores, White 115 (MO). Pipeline Road, Witherspoon & Witherspoon 8293 (MO) Salamanca Hydrographic Station, Rio Pequeni, Woodson et al. 1613 (MO). BARRO COLORADO ISLAND: Aviles 11 A (MO): Bartlett & gus 16746 (MO); Croat 4053, 4273, 4392, 6362, 6370, 6814, 6962 (all ); Dwyer et al. 8471 (MO); Ebinger 36 (MO); Hayden 39, 1039, 1040, 1041 (all MO); Shattuck 167, 924 (MO); Starry 76 (MO); Tyson 4211 (MO): Wetmore & Abbe 41, 173 (both MO): Woodworth & Vestal 622 (MO); Zetek 4270 (MO). CHIRIQUI: Puerto Armuelles, D'Arcy 10071 (MO). Chiriquí- V i 200 is et al. 642 (GH, MO). cocLe: El Valle, road to La Mesa, Croat 13312 (MO). Club Campestre, ca. 700 m, Duke 13263 (MO). Cerro Pilón and El Valle, 700-900 m, Duke & Dwyer 13898 A (MO). El Valle, Dwyer 1833, p^ (both MO). El Valle, vicinity of Cerro Pilon, Dwyer & Correa 7969 (MO, NY, . La Mesa, Dwyer & Duke 8294 (MO). Cerro Pilón, Kirkbride 1084 (MO). El Valle, Lao 280 (MO); Lewis 4 al. 1788 (COL, MO, UC, VEN). Río Grande, Rosario 5 (MO). El Valle, Seibert 425 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 211 (MO). Llano Bonito, S of Las See yop 529 (MO). EI Valle, Tyson et al. 3967 (MO). Las Margaritas and El Valle, Woodson et al. MO). COLON: Río Gatuncillo, vicinity of Santa Rosa, Allen 4160 (MO). Sardinilla, Blum & D po 525 (both MO). Sabanita, Croat 11068 (MO). Por- tobelo, D'Arcy & D'Arcy 6684 (MO); Dwyer 5008 (MO). Chagres, Gutierrez 23 (MO). 2.4 mi SW of Pii ña, Lewis et al. 1852 (COL, MO, UC). I mi S of Gatuncillo, Stimson 5260 (MO). Achiote, Tyson et al. 4510 (MO). DARIEN: Río Tuqueza below Quebrada Venado, Bristan 1073 (MO). Río PE Croat & Porter 15522, 15453 (both MO). 2-3 mi S of El Real, Duke 4841 (MO). Río Pirre, 2-5 m above El Real, Duke 5082 (MO). Cerro Pirre, S of El Real, 100-600 m, Duke 5292 (MO). 10 mi S of El Real, Río Pirre, Duke 5449 (MO). Between Río Membrillo and Río Yaviza, Duke 8620 (MO). Río Pinas, Duke 10564 (MO). Cerro Chucula, drainage Río Pavarando, Gentry 4253 (MO). Upper Río Tuqueza, Le Clezio 132 (MO). Río Tuquesa, Charco Chiva, Mori 6972 (MO). Without locality, Sexton 218 (MO). Yaviza, along Río Chucunaque, Quebrada Barbua, Stern et al. 85 (MO). Trail El Real to Pinogana, Stern et al. 274 (MO). Trail between Paya and Payita, Stern et al. 354 (MO). Tucuti, Chepigana, Terry & Terry 1406 (MO). 3 mi N of Santa Fé, Tyson et al. 4622 (MO). As SANTOS: Las Tablas, Dwyer 1157A (MO). 17.8 mi S of Macaracas, 1100 ft, Lewis et al. 1607 (MO, UC). P PANAMÁ: Panama City, Lefevre Park, Bartlett & Lasser 16335 (MO). Road to Cerro Cam M Croat 12047 (MO). Río Canita, Jenené, Duke 3818 (MO). Río Bayano above confluence with Río Chepo, Duke 3985 (MO). 6 mi E of Chepo, Duke 4063, 4082 (both MO). Río Mamonica, 4 mi beyond Chepo, Duke juela, Dwyer 1042, 1045 (both MO). Tocumen, Dwyer 4065 (MO). 1 km E of El Llano, Gentry 6467 (MO). San Se Island, Johnston 147 (MO). Transisthmian Highway and Madden Dam, Lewis et al. 5303 (MO, SCZ, UC). P oa pal 12 (MO). 25 km NE of Cerro Azul, Rio Piedras, 550 m, Mori & Kallunki 3286 (MO). La C era, Rodriguez 14 (MO). El Llano, Tyson 1739 (MO). SAN BLAS: Puerto Obaldia, Croat 16959 pes Sasardí, Duke 10135 (MO). Río Diablo and Río Acuati, Duke Tasa Ailigandí, Dwyer 6848 (MO). Soskatupu Island, = 1662 (COL, MO, UC), 1699 (MO). waters of Río Mulatupo, Elias 1755 (GH, MO, UC, US). Mainland poa Playon Chico, Ganis 6371 (MO). 3-4 hrs up Río Mulutupu on foot, Kirkbride 219 (MO). VERAGUAS: Isla de Coiba, Dwyer 1565 (MO). Tute, Dwyer 4283 (MO). 3.9-5.0 mi N of Santa Fé, 5001000 m, Gentry 2952, 2999 (both MO). 6. Hamelia rovirosae Wernham, J. Bot. 49: 211. 1911. Type: México, Rovirosa 499 (NY, US, isotypes). H. patens var. coronata J. D. Smith, Bot. Gaz. 4: 4. 1905. Type: Guatemala, Tuerckheim 8532 (US, holotype; F, NY, isotypes). H. r Blake, Contr. Gray Herb. 52: 105. 1917. Type: British Honduras, Peck 104 (G, type). Shrubs, the branchlets villous, the hairs septate. Leaves ternate, occasionally opposite, elliptic to ovate, 5-16 cm long, 2-7 cm wide, acuminate, attenuate at the base, the lateral veins 3-5, arcuate, glabrous to sparsely villose above, villous beneath; petioles 0.3—2.0 cm long; stipules linear, often widely so, 5-8 mm long. Inflorescences terminal, dichasial or often helicoid; peduncles to 3 cm long, vil- lous, the floral axes 1.5-3.5 cm long, each axis 3-8-flowered, the flowers secund, sessile or on pedicels 0.1—0.2 cm long. Flowers with the hypanthium to 4.5 mm long, villous, the calycine lobes oblong or triangular, 2.0-5.4 mm long, villous, glabrous within: corolla reddish orange to scarlet, the tube 1.8-2.2 cm long, 2-3 mm wide, villous, the lobes ovate, 1-2 mm long, acute, villous outside; anthers 10-12 mm long, the terminal appendage ovate, ca. 0.5 mm long, the filaments 7— 9 mm long; ovary 5-celled. Fruits cylindrical, to 1.2 cm long, 0.5-0.8 cm wide, red, finally black; seeds ca. 1 mm long Hamelia rovirosae is found in Mexico and throughout Central America. BOCAS DEL TORO: Changuinola Valley, Dunlap 192 (F). 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 7. Hamelia sanguinea Elias, Mem. New York Bot. Gard. 26(4): 116. 1976. New name for H. pauciflora Standley. H. pauciflora Standley, Contr. U.S. Natl. Herb. 18: 139. 1916. rype: Panamá, Pittier 3469 (US, holotype), not H. pauciflora Willd. (1825). Shrubs or small trees, 4-7 m tall, the branchlets terete to angular, glabrous or puberulent, these usually forming numerous short branchlets. Leaves mostly ternately disposed, ovate, trapeziform oblong or ovate rotund, 2.0-8.5 cm long, 1.0-3.5 cm wide, attenuate at the apex, acuminate, the acumen triangular, to 1 cm long, acute, attenuate acute to cuneate at the base, the costa prominulous or grooved above, the lateral veins (3—)4-5, arcuate, membranous to subcharta- ceous, glabrate above, usually minutely papillate and with numerous raphides, sparsely short ciliate beneath, often only on the costa and the veins, ciliolate; petioles slender, wiry, scarcely measurable or to 3 cm long, pubescent; stipules deciduous. Inflorescences with dichasia 2—5(—9)-flowered, terminal on short lat- eral branches; peduncles 1.0-2.5 cm long, the floral axes to 5 cm long, each with up to 4 flowers, the flowers secund, those in the axils of dichasia sessile or subsessile, the others on pedicels. Flowers with the hypanthium oblong, to 4.5 mm long, glabrous, the raphides numerous; calycine cup obsolete, the lobes ovate, to 1.2 mm long, acute: corolla orange red to red, glabrous, the tube cylin- drical to 2 cm long, ca. 0.3 cm wide, the lobes ovate, ca. 1.5 mm long or longer, pubescent outside; anthers linear, 8-10 mm long, the terminal appendage ca. 0.5 mm long. Fruits suborbicular, about twice as long as wide at maturity, the per- sistent ovarian disc rostrate; seeds to 1.2 mm long. Hamelia sanguinea is known only from Panama. CANAL ZONE: | mi above Madden Lake, Duke 4479 (MO). Madden Dam, Boy Scout Camp Road, Dwyer & Elias 7488 (GH, MO, UC, US), 7494 (GH, MO, US), 7514 (MO, UC), 7515 (COL, MO, UC): Dwyer & Hayden 21 (MO); Dwyer & Lallathin 8834 (MO). Madden Dam, Ebinger 875 (MO). Boy Scout Road, Madden Dam, Hayden 77 (MO). Forest of Gatün, Hladik 512 (MO). Madden Dam Road, Boy Scout Camp Road, Kirkbride 50 (MO). Madden Dam, Lewis et al. 22 (MO, SCZ, UC). Trail, Río Petitpie, road Ft. Sherman from Gatün Locks (S2 or 82), Mori & Kallunki 2679 (MO). Boy Scout Road, Madden Dam, Porter et al. 4017 (MO, SCZ). Boy Scout Camp, Madden Lake, Tyson 5438 (MO). Boy Scout Camp R oad, Madden Dam, Tyson & Dwyer 1201 (MO). Salamanca Hydro- graphic Station, Río Pequeni, TONO 1595 (MO). DARIEN: Quebrada Venado and Peje Swamp, headwaters of Río Tuquesa, Bristan 1021 (MO). Near Refugio, 15-21 mi N of Santa Fé, Duke 10265 (MO). Morti Hydro, 260 m, Duke 14236 (MO). 8. Hamelia xerocarpa Kuntze, Rev. Gen. Pl. 1: 284. 1891. TYPE: Costa Rica, Kuntze 20281 (NY, holotype). x Nes iue Wernham, J. Bot. 49: 209. 1911. Type: Nicaragua, Tate 200 (K, holotype, not seen). wleei Standley, J. Wash. Acad. Sci. 15: 7. 1925. type: Costa Rica, Rowlee & Stork 75 (US, olotype). H. ueni) a Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 116. 1940. rvPr: Colombia, Haught 2176 (F Shrubs or small trees, the branchlets ultimately angular, scarcely lenticellate, villous. Leaves ovate to elliptic, 8-37 cm long, 4-14 cm wide, acuminate, atten- uate to rounded at the base, the lateral veins 11-18, chartaceous, pilose above, sericeous along the veins, especially in the axils; petioles 0.8—7.0(—8.5) cm long, villous; stipules triangular, 7-17 mm long, cuspidate, often with 2 reduced lateral 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 213 teeth, glabrate. Inflorescences terminal, of dichasia or scorpioid cymes; pedun- cles 1.5-5.5 cm long, villous, the floral axes to 15 cm long, ascending, often ferrugineous pilose, the flowers secund, ca. 25 on each axis, pedicels to 0.2 cm long. Flowers with the hypanthium 3-4 mm long, glabrous to spreading pilose; calycine lobes ovate, 1-2 mm long, glabrous to pilose; corolla yellow salverform, to 3.5 cm long, the tube 0.4-0.7 cm wide, glabrous to densely puberulent, the lobes ovate, 4-8 mm long, acute; anthers 10-16 mm long, acute, the connective extended for ca. 0.1 mm at the apex, acute, sagittate at the base, the filaments 8-15 mm long; ovary 5-loculate, the style ca. 20 mm long, the stigmas 5, to 5mm long. Hamelia xerocarpa is known from Costa Rica, Panama, and Colombia. 2381 (F. MO). DARIEN: Río Tuquesa, Middle Tuquesa Mining Co. Camp called Charco Peje, Mori VERAGUAS: NW of Santa Fé, 11 km from Escuela Agricola Alto de Piedra, 450-550 m, Mori et al. 3828 (MO). 45. HEMIDIODIA Hemidiodia Schum. in Mart., Fl. Bras. 6(6): 29. 1888. TYPE: Spermacoce ocim- ifolia Willd. — Hemidiodia ocimifolia (Willd.) Schum. Perennial herbs, often woody and profusely branching from the base. Leaves narrow, the veins conspicuous; stipules adnate to the petioles, forming a setif- erous sheath. Inflorescences axillary, epedunculate, the flowers fasciculate. Flowers with the calycine lobes 4; corolla funnelform, the lobes 4, valvate; sta- mens attached near the mouth, exserted. Fruits of 2 mericarps separating at maturity from a median septum, opening basally. Hemidiodia consists of a single species which extends from Mexico to South America. It also occurs in the West Indies and in Malaya. Literature: Bremekamp, C. 1934. Diodia (Gronov.) L. in Notes on the Rubiaceae of Suri- nam. Rec. Bot. Neerl. 31: 305. 1. Hemidiodia ocimifolia ( Willd.) Schum. in Mart., Fl. Bras. 6(6): 29. 1888.—Fi1G. Spermacoce ocimifolia Willd. ex R. & S. Syst. Veg. 3: 530. 1818. TYPE: not seen. Subshrubs to 1 m tall, often branching from the base, the stems usually an- gular, smooth, puberulent or glabrous, the bark thin, often exfoliating, the nodes well spaced. Leaves narrowly lanceolate, ovate lanceolate to trapeziform lan- ceolate, to 6.5(—7.5) cm long, to 1.3 cm wide, acute at the apex, usually attenuate acute at the base, the costa prominulous to plane, the lateral veins ca. 7, strongly arcuate, chartaceous, puberulent or farinose puberulent above and beneath, the petioles almost absent or 0.3-1.0 cm long, ca. 0.25 mm wide; stipules adnate to 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 TW BECK be FIGURE 47. Hemidiodia ocimifolia (Willd.) Schum.—A. Habit (xV2).—B. Flower (x6).—C. Fruits (x 5). [After McDaniel 6875.] the petioles, the body fan shaped, 2-5 mm long, membranous, turgid, puberulent, with 8-12 erect or spreading setas. Inflorescences disposed as sessile heads in the leaf axils, to 0.6 cm long. Flowers with the hypanthium oblong, ca. 1.5 mm long, petaloid, glabrous, the calycine teeth 4, scarcely visible or as triangular 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 215 points, to 0.15 mm long, glabrescent; corolla with the tube short cylindrical, ca. 1.3 mm long, petaloid, glabrous, the lobes 3-4, ca. equal to the tube or much shorter, oblong, obtuse, glabrous outside; stamens 3-4, the anthers oblong ro- tund, ca. 0.2 mm long, the filaments shorter than or as long as the corolla lobes, attached at the mouth; stigmas compressed rotund, ca. 0.15 mm wide, the style ca. 1.3 mm long, the septum of the ovary thin. Fruits oblong, 3-4 mm long, bicarpellate, each carpel 1-seeded, separating from a median septum, dehiscing at the base, densely pubescent, the calycine lobes persistent, to 0.2 mm long. The species ranges from Mexico to Brasil. Stern et al. 241 list the common name "Yerba Meona"' and indicate that the leaves are ‘‘used for kidney and urinary disorders, boiled as a tea.” CAS DEL TORO: Station Milla 7.5 on p pr Almirante Railroad, Croat 38095 (MO). Chan- es River, sss mi inland, Lewis et al. 887 (G, H, K, MO, NY, UC, US). Chiriquicito to 5 mi S along Río Guarumo, Lewis et al. 2075 (MO, SCZ, UO). CANAL ZONE: S-5 Road, Caribbean side, Blum 1505 (MO). pone Colorado Island, Croat 12549, 12840 (both MO). Pipeline Road within 5 mi of Gamboa Gate, D’Arcy 9274 (MO). Hill C6, Fort Sherman, Duke 4354, 4374 (both MO). Pipeline Road, 16 mi N of Gamboa, Lewis et al. 5464 (MO). Navy reservation, N of Gamboa, Robyns 69-50 MO). cocLÉ: El Copé, Burch et al. 1374 (COL, MO, UC). Cerro Pilon, Duke 10275 (MO): Dwyer et al. 4538 (MO, SCZ, UC), 4563 (MO). PANAMA: Cerro Azul, Croat 11568 (MO). Cerro Campana, Croat 12048 (MO). Cerro Jefe, D'Arcy 3967 (MO). Cerro Campana, Duke 5996 (MO). Cerro Azul, Duke 9352 (MO). Río Pacora, Duke 12016 (MO). Cerro Campana, Dwyer & Kirkbride 7822 (MO). Chagres, Fendler 145 (MO). Cerro Jefe, Folsom a al. 2533 (MO). Rio Pasiga, Gentry 2362 (MO). Cerro Jefe, Gentry 6755 (MO); Kirkbride & Crebbs 8 (MO). Cerro Azul, Lewis et al. 228 (F, GH, K, MO, NY, UC, US). Cerro Campana, McDaniel 6875 (MO). Road to Altos de Pacora, Mori & Kallunki 2351 (MO). TV tower, Cerro Peñón, 3 km S of Alcalde Diaz, Nee 8591 (MO). Cerro Campana, Porter et al. 4183, 4289 (both MO), 4890 (MO, SCZ). Cerro Azul, Tyson 2082 (MO). Cerro Jefe, Tyson et al. 4309 (MO). Macapale Island, Madden Lake, Tyson 5484 (MO). W of El Llano, Tyson 6876 (MO). bat I et al. 1396 (MO). SAN BLAS: Ailigandi, mainland, Lewis et al. 67, 83 (both MO). : Ustupo, D'Arcy 9518 (MO). Road from Santa Fé past Agricultural School to base o pin Tute, Folsom 1576 (MO). 3.9-5.0 mi N of Santa Fé, Gentry 2975A (MO). 46. HILLIA Hillia Jacq., Enum. Pl. Carib. 3, 18. 1760. Type: H. parasitica Jacq. eira Vell. ex Vand., Fl. Lusit. Bras. Sp. 21. 1788. TYPE: F. vellozoana J. A. & J. H. Schultes. videt. Vell., Fl. Flum. 141. 1825, Not Bureau 1868. LECTOTYPE: S. illustris Vell. Shrubs, mostly epiphytic, glabrous. Leaves succulent, drying coriaceous, pet- iolate; stipules thin, caducous. Flowers often large, solitary, terminal; hypan- thium obovoid or cylindrical, the calycine cup absent, the lobes 2-5 or absent, caducous or persistent in fruit; corolla salverform or funnel shaped, the tube elongate, expanded toward the mouth, glabrous, the lobes 3-7, contorted in bud; stamens included; ovary 2-celled, the ovules numerous. Fruits capsular, narrowly oblong, or cylindrical, the 2 valves dehiscing septicidally; seeds imbricate, basally appendaged, bearing a distal tuft of hairs. Hillia, a genus of about 20 species, occurs in the West Indies, Mexico, Central America, and South America. a. Leaves small, 0.6-1.3 cm lon b Leaves ca. 0.7 cm long, Qe at the apex; corolla tube ca. 1.5 cm long .. og hh a te a a ed Se gsc a UM I H. chiapensis bb. Leaves 0.6-1.3 cm long, tapering toward the apex; corolla tube to 2.8 cm long ------ 5. H. panamensis 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 aa. ii: iai 2-8 cm lon es 2-3 times longer than wide: calycine lobes ca. 1.7 cm long . 3. H. UNUM ec. ems less than twice as long as wide; Mais, lobes less than 1 cm long or abse d. Corolla tube 2-3 cm long __- MERECE . H. s m" dd. dd tube 4-7 cm long. Trees to 5 m tall; leaves 0.6-2.8 cm wide, rounded at the apex, e. tan beneath; corolla tube 6-7 cm n ng EEE A 2. chiriquiensis ee. Epiphytic shrubs to 2 m tall; o4 cm wide, usually Rue at the apex, drying black bos Velas MEA tube 4-5 cm long. |... Loose 6. H. tetrandra 1. Hillia chiapensis Standley, J. Wash. Acad. Sci. 16: 16. 1926. TYPE: México, Purpus 262 (US, holotype). la. Hillia chiapensis Standley var. chiapensis. Epiphytic shrubs, much branched, heathlike in habit, the branches rough, 4-angled, the angles delicately alate, glabrous, the nodes constricted, the point of petiole insertion depressed and with an annular orifice. Leaves ericoid, elliptic, oblong elliptic or oval, to 0.7(-0.9) cm long, rounded at the apex, acute to obtuse at the base, the costa and lateral veins not visible in dried material, coriaceous, glabrous, marcescent; petioles to 0.2 cm long; stipules deciduous, occasionally persistent at tip of branchlets, obovate to oblong, 3-4 mm long, rounded, gla- brous, venose. Flowers solitary, sessile; corolla tube ca. 15 mm long, the lobes ca. 7 mm long. Fruits capsular, to 3 cm long. Hillia chiapensis ranges from Mexico to Panama. NAMA: Finca El Indio, Cerro Jefe, Correa et al. 1770 (MO). Summit of Cerro Jefe and forests dione pu beyond summit, to 2700 ft, Hayden 1021 (COL, DS, MO, UC). Ib. Hillia chiapensis subsp. grandifolia Dwyer,?* Type: Panamá, Nee 9698 (MO, holotype) Leaves to 3(-4.5) cm long, to 1.2(-1.8) cm wide; fruits to 4 cm long. The new subspecies is known only from Panama. It differs from H. chiapensis subsp. chiapensis in having much wider leaves and more elongate fruit. =RAGUAS: Lower montane wet forest 6-7 km W of Santa Fé on new road past agricultural iud 2900 ft, Nee 9698 (MO). 2. Hillia chiriquiensis Dwyer.? TYPE: Panamá, Mori & Kallunki 5810 (MO, ho- lotype).—Fic. 48 Trees to 5 m tall, branching freely, ericoid in appearance, ultimately black when dry, the branchlets thick, the nodes well spaced or crowded, ultimately 56 Hillia chiapensis ee sida Dwyer, subsp. nov. Folia ad 3(—4.5) cm longa, ad 1.2(-1.8) cm lata; fructus ad 4 cm lon ; ET plerumque invisibilibus. GU zer petiolis ad 0.8 cm cage sipol CU ad L5 em longis. Flores terminales solitarii, lobis calycis 4 oblongis, ca. 0.4 cm longis, ca. 0.8 cm latis; corolla viridi, tubo infundibuliformi, ad 7 cm longo, 0.3 cm lato, lobis 4—5, antheris subsessilibus, ca. 1.2 cin longis, apice connectivo ad 1 mm producto. Fructus non visi. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 217 I] U \ j Äl LA \; ' W C. 2 i IN IB SS. Z Da SS DN Figure 48. Hillia. A-C. H. chiriquiensis Dwyer.—A. Habit (<'2).—B. Flower, opened to show interior ( x 3).—C. Ovary cross section (x 11.5). [After Mori & Kallunki 5810.]—D. H. tetrandra Swartz. Fruit (x.6). [After Dwyer & Gauger 7377.] angular, nodose in appearance. Leaves crowded at the apex of the twigs, nar- rowly obovate oblong, 2.0-6.5 cm long, 0.6-2.8 cm wide, rounded at the apex, acute or cuneate toward the base, the costa scarcely prominulous above, only visible on the proximal side of the blade, the lateral veins ca. 4, inconspicuous, at times invisible above, arcuate, the margin lightly revolute, coriaceous, discol- orous, smooth, drying tan beneath, glabrous; petioles to 0.8 cm long, thick, to 0.25 cm wide; stipules deciduous, persistent only at the tips of the twigs, oblong, 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 to 1.5 cm long, obtuse, coriaceous, glabrous. Flowers terminal, solitary, calycine lobes 4, oblong, 4-6 mm long, 2-4 mm wide, obtuse, coriaceous, glabrous; corolla green, the tube infundibuliform, 6-7 cm long, to 0.3 cm wide, dilated toward the apex, the lobes 4—5, white with a pink tinge, ovate oblong or oblong, to 2.5 cm long, to 1.3 cm wide, obtuse, glabrous; stamens 4—5, the anthers sub- sessile, oblong, ca. 1.2 cm long, the acumen to 1 mm long, sagittate at the base, attached about 1.5 cm below the mouth; style linear, to 5.5 cm long, ca. 0.1 cm wide, lignose when dry, glabrous. Fruits not seen. Hillia chiriquiensis is known only from Panama. It is remarkable in being a fair-sized tree and in having the longest corolla among the Panamanian Hillia. CHIRIQUI: N of San Felix, Chiriqui-Bocas del Toro border, on Cerro Colorado Copper Mine Road, lower montane rain forest, 5000-5500 ft, Mori & Kallunki 5810 (MO). 3. Hillia ligulifolia Dwyer.” Type: Panamá, Nee 9812 (MO, holotype). Epiphytic trees, 3.5-5 m tall, the branchlets numerous, glabrous, drying dark, tending to be ƏpñƏsiie and arcuate, ultimately quadrangular, the nodes well spaced below, crowded at the tips of the twigs. Leaves narrowly oblong or vague- ly obovate oblong, to 8 cm long, 1.5-2.5 cm long, the blades usually 21⁄2 times longer than wide, conspicuously rounded or truncate at the apex, basally cuneate to attenuate acute, the costa thick and prominulous proximally, evanescent dis- tally, the lateral veins 4-5, inconspicuous, scarcely prominulous, strongly as- cending, coriaceous, concolorous, drying dark brown, glabrous; petioles variable in length and thickness, 0.1—1.5 cm long, the thicker petioles short, the blades decurrent; stipules deciduous, often persisting at the tips of the twigs, of the same shape and texture as the leaves, to 2 cm long. Flowers solitary, terminal, sessile, the hypanthium enclosed by 2 bracts (stipules?) and by 4 bracteoles of the same texture as the leaf blades, to 2 cm long; bracts connate, each part with 2 subrotund lobes, to 0.6 cm long; bracteoles 4; calycine lobes 4, narrowly oblong, 1.0—1.4 cm long, 0.2-0.3 cm wide, unequal, coriaceous, glabrous; corolla tube ca. 4 cm long, the lobes 1 cm long, white, the lateral margins tinged with red; stamens 5, the anthers narrowly oblong, ca. 12 mm long, the connective produced at the apex; style slender, to 5.5 cm long, the stigmas oblong, ca. 5.5 mm long, ca. 2.5 mm wide. Fruits sessile, narrowly oblong, cigar shaped, often somewhat falcate, (3.5-)5-6 cm long, to 1 cm wide before dehiscence, rounded at the apex, obtuse at the base, drying deep red, glabrous, the calyx often persistent, the lobes to 1.8 cm long, to 0.6 cm wide, rounded at the apex, venose Hillia ligulifolia is known only from Panama. It is distinguished from other Hillia of Panama by its very large calycine lobes and its leaves, many of which * Hillia ligulifolia Dwyer, spec. nov. Arbores. Folia angusto-oblonga vel vix obovato-oblonga, saepe liguliformia, ad 8 cm longa, 1.5-2.5 cm lata, apice rotunda vel truncata, basim versus cuneata angusto-oblongi, saepe falcati, (3.S—)5—6 cm longi, ad 1 cm lati, obtusi basi, apice rotundati:; lobis calycis persistentibus ad 1.8 cm longis, ad 0.6 cm latis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 219 are truncate at the apex. Many of the leaf blades have a tonguelike or straplike shape. Noteworthy is the fact that the petioles are quite variable in length and thickness. The persistent calycine lobes give the large fruits a striking appearance. Only one other Panamanian large-leafed Hillia is known in the fruiting stage: H. tetrandra. CHIRIQUi: Cerro Colorado, 34.1 km beyond bridge over Rio San Félix, 1390 m, Croat 37199, 37256 (both MO). Cerro Colorado, 35.6 km from Rio San Félix bridge, 1390 m, Sullivan 388 (MO). PANAMA: Cerro Jefe, ca. 1000 m, Mori & Joly 7926 (MO). VERAGUAS: Lower montane wet forest 6— 7 km W of Santa Fé on new road past agriculture school, 2900 ft, Nee 98/2 (MO). 4. Hillia palmana Standley, J. Wash. Acad. Sci. 18(6): 164. 1928. TYPE: Costa Rica, Maxon & Harvey 8045 (US, holotype). Epiphytic shrubs, the branchlets terete, finally compressed, black when dry and covered with gray spots, glabrous. Leaves persisting toward the apex of the branchlets, obovate or obovate rotund, 1.5—4.5 cm long, 1.0-2.2 cm wide, round or truncate at the apex, cuneate basally, the costa scarcely distinguishable, the lateral veins not visible, thickly coriaceous, concolorous, glabrous, the margin slenderly callose; petioles thick, to 1 cm long, to 0.3 cm wide; stipules deciduous, persistent toward the apex, obovate rotund, to 1.5 cm long, rounded, coriaceous, glabrous, the veins conspicuous or evanescent. Flowers terminal, solitary, to 4 cm long, enclosed partially by 2 large bracteoles, these compressed rotund, to 2 cm long, scarious, glabrous, venose; hypanthium rectangular, to 1.8 mm long, glabrous, the calycine cup none, the lobes obviously absent; corolla white, the tube infundibuliform, 2-3 cm long, ca. 0.3 cm wide, rigid, glabrous, the lobes 4, compressed rotund, to | cm long, wider than long, glabrous; stamens subsessile, the anthers linear oblong, ca. 3 mm long, affixed near the mouth; style ca. 6 mm long, the stigmas 2, subulate, ca. 3 mm long. Fruits not seen. Hillia palmana is known only from Costa Rica and Panama. The leaves are strikingly ligulate. As few flowers are available for dissection I elected to dissect one whose corolla measured only 2 cm in length. No ovules were located; this coupled with the absence of fruit in the two known collections is cause for spec- ulating that the flowers are unisexual. In this dissection the style and stigmas measured only about !^ the length of the corolla tube. VERAGUAS: NW of Santa Fé, | km from Escuela Agricola Alto de Piedras, Mori & Kallunki 4873 O). 5. Hillia panamensis Standley, N. Am. Fl. 32(2): 117. 1921. TYPE: Panamá, Pittier 3190 (US, holotype). H. hathewayi Fosberg, Sida 2: 387. 1966. rype: Costa Rica, Hatheway 1371 (US, holotype). Epiphytic trees, to 4 m tall, the branchlets ericoid in habit, ultimately bearing very short and numerous branchlets, the nodes proximate, the leaves tending to persist only at the tips of the twiglets. Leaves elliptic oblong, 0.6-1.2 cm long, 0.4-0.5 cm wide, tapering slightly toward the apex, subacute to obtuse at the base, frequently cuneate, the costa and lateral veins invisible above, scarcely visible beneath, the lateral veins ca. 2, strongly arcuate ascending, subcoriaceous, 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 concolorous, drying dark, glabrous; petioles absent or to 0.2 cm long; stipules (Standley) narrowly oblong, to 4 mm long, obtuse, brownish. Flowers terminal, solitary; hypanthium narrowly oblong or turbinate, ca. 2 mm long, glabrous, the calycine lobes 4, linear oblong, to 9 mm long, ca. 0.8 mm wide, obtuse, gla- brous, drying red, venose; corolla white, the tube narrowly cylindrical, to 2.8 cm long, ca. 0.8 cm wide in the middle, glabrous inside and out, the lobes 4, ovate triangular, to 1.2 cm long, obtuse, glabrous, petaloid, venose; stamens 4, the anthers subsessile, affixed immediately below the mouth, oblong, ca. 1.8 mm long, obtuse, the style linear, ca. 12 mm long, the stigmas 2, ca. 3 mm long, scarcely wider than the style. Fruits linear oblong, 1.0—2.5 cm long; seeds oblong with terminal hairs to 6 mm long CHIRIQUÍ: Cerro Horqueta, 1650 m, Croat 26957 (MO): Pittier 3190 (US). 6. Hillia tetrandra Swartz, Prodr. 58. 1788. TYPE: Jamaica, Swartz (not seen). H. maxonii Standley, J. Wash. Acad. Sci. 18: 163. 1928. rype: Nicaragua, Maxon 7501 (US, holo- type). Small shrubs, to 2 m tall, often epiphytic, the twiglets terete or angular, smooth, glabrous, the nodes well spaced. Leaves oblong rotund or ovate oblong, mostly equilateral, to 7 cm long, to 4 cm wide, usually deltoid, occasionally obtuse at the apex, obtuse, rounded or cuneate at the base, the costa plane proximally above, subprominulous proximally beneath, evanescent distally, the lateral veins ca. 5, arcuate, evanescent above and beneath, coriaceous, drying dark brown; petioles to 1 cm long, to 0.25 cm wide, shiny, puberulent to gla- brescent, the lamina decurrent on the upper side; stipules oblong, to 1.5 cm long, tapering, obtuse, coriaceous, glabrous. Flowers terminal, solitary, subsessile; hypanthium oblong, ca. 10 mm long, ca. 3.5 mm wide, glabrous, the lobes often deciduous, 0.5-1.0 cm long or perhaps scarcely measurable; corolla white or yellow with the tube narrowly cylindrical, 4—5 cm long, 0.2-0.4 cm wide, often subfalcate, glabrous inside and out, the lobes 5, oblong, ca. 1.5 cm long, ca. 1.2 cm wide, glabrous; stamens 5, the anthers oblong, ca. 11 mm long, 2 mm wide, obtuse, the filaments short, ca. 1 mm long; style linear, ca. 5.5 cm long, ca. 0.9 mm wide, the stigmas about as wide as the style, to 12 mm long, villose. Fruits sessile, narrowly oblong, 5(-7) cm long, 0.4-0.9 cm wide, truncate or rounded, smooth, glabrous, lignose; seed ca. 2 mm long with a tuft of tawny hairs to 12 mm long Hillia tetrandra occurs in the West Indies and throughout most of Central America. One collection, Maxon & Harvey 8092 (US), has calycine lobes to 4 mm long; these are obtuse at the apex. In other respects it resembles the Panamanian material of Hillia tetrandra. Mori & Kallunki 5815 notes. "flower not opened when collected at 5:00 p.m. but opened in bag by 7:00 p.m. Moth pollinated?" HiRIQUÍ: San Félix, Maxon & Harvey 8092 (US). N of San Félix at Chiriqui- des Ps Toro bodie Cerro Colorado Copper Mine Road, 5000-5500 ft, Mori & Kallunki 5815 (MO). c : Cerro Valle Chiquito, Seibert 495 (MO). COLON: Santa Rita Ridge, Dwyer & Ge ntry 9394 (MO). Pond Cerro Campana, Allen 1872 (F); Croat 14801 (MO). Goofy iri. road to Cerro Jefe, Correa & Dressler 941 (MO). Cerro Campana, Dressler 3522 (MO). Cerro Jefe, Dw ver & Gauger 7377 (MO); Mori & Kallunki 6095 (MO). vERAGUAS: 5 mi W of Santa Fé, Cost 23079 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 221 47. HIPPOTIS Hippotis Ruiz & Pavón, Prodr. Fl. Peruv. 33. 1794. TYPE: H. trifolia Ruiz & Pavón. Shrubs or trees. Leaves with the blades membranous or thinly chartaceous, the intervenal areas lineolate, petiolate, the stipules large, caducous. /nflores- cences cymose (or flowers occasionally solitary). Flowers large, 5-merous, tend- ing to become zygomorphic, the calyx large, spathaceous or bilabiate; corolla tubular or funnelform, usually curved, the lobes short, plicate valvate; stamens often unequal in length, inserted near the middle of the tube; style slender, elon- gate, the ovary 2-celled, the ovules numerous, the placentas adnate peltately to the septum. Fruits baccate, round; seeds numerous, foveolate. Hippotis, a genus of about a dozen species, is known from Panama, Colombia, Venezuela, Ecuador, and Peru. 1. Hippotis albiflora Karst., Fl. Colomb. 1: 33. 1858. TYPE: not seen.—Fic. 49. Duroia panamensis Dwyer, Ann. Missouri Bot. Gard. 55: 38. 1968. Type: Panama, Kirkbride & Duke 571 (MO, holotype). Trees to 8 m tall, the branchlets subterete, drying lightly rimose, densely pilose terminally, nodose. Leaves obovate elliptic, to 26 cm long, to 15 cm wide, widely deltoid at the apex, acuminate, the acumen to 1.5 cm long, the costa prominulous above, prominent and convex beneath, the lateral veins ca. 12, at first strict, then slightly arcuate toward the margin, 2.0-2.5 cm apart in larger blades, prominent beneath, the secondary veins irregularly pinnatiform, the ul- timate veins enclosing minute, parallel striae, the hairs appressed, white, ca. 1- 2 mm apart, somewhat scabrous, pilose beneath, with some hairs more rigid and longer, ciliolate; petioles 1.0-2.5 cm long, lignose, pilose; stipules free, decidu- ous, ovate elliptic, here the younger ones to 1.5 cm long, densely pilose. Flowers 1(?-3), terminal or axillary; hypanthium oblong, 8-10 mm long, densely appressed pilose, the hairs yellowish white, ascending, 4-5 mm long, the calycine cup tu- bular for 10 mm then spathaceous for ca. 15 mm, thickly petaloid, densely villose outside, the hairs stiff, white, to 2.5 mm long, glabrous and eglandular within; corolla white or greenish white, curved, trumpet shaped, to 45 mm long, ca. 6 mm wide in the middle, densely white pilose outside, glabrous within, stiffly petaloid or subcoriaceous, the lobes 5, elliptic or ovate, ca. ] cm long, ca. 0.8 cm wide, densely white pilose outside; stamens 5, included, the filaments slightly unequal, to 11.5 mm long and to 0.8 mm wide, plane, densely pilose, attached below or near middle of the tube, the anthers oblong, ca. 3 mm long; ovarian disc oblong rotund, to 3 mm long, the style elongate, subplane, ca. | mm wide, the stigmas not seen. Fruits nutlike, elliptic, to 3 cm long, to 1.5 cm wide, often 5- 7-ribbed, densely pilose, the hairs yellowish white, the calyx persistent, about as long as the fruit; seeds elliptic oblong, ca. 2 mm long, 3-angled, foveolate under magnification. Hippotis albiflora occurs in Costa Rica, Panama and Colombia. Kirkbride & Duke 571 notes the: "corolla tube greenish white, anthers yellow." McDaniel 5079 notes that the plant is a "vine" but this is highly unlikely. 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 49. Hippotis albiflora Karst. Habit (x14). [After Kirkbride & Duke 571.] COSTA RICA: HEREDIA: Puerto Viejo, Sarapiqui, 10°26’N, 84*01'W, Finca La Selva, ed 1044 (MO). PANAMÁ: BOCAS DEL TORO: Duw € E Peak, N of Río Teribe, across from West ie Kirkbride & Duke 571 (MO). 4 mi N of Alm . McDaniel 5079 (MO). COLOMBIA: CHOCC in forest NW of Alto Curiche, ca. 300 m, Duke ? pus 11258 (MO). SANTANDER: Magdalena ia Campo Capote, 30 km E of Carare, 300 m, Gentry et al. 20083 (MO). 48. HOFFMANNIA Hoffmannia Swartz, Prodr. Fl. Ind. Occ. 30. 1788. TYPE: H. pedunculata Swartz. O'Higginsia Ruiz & Pavón, Fl. Peru 1: 55. 1798. TvPE: O. obovata R. & P sa Pers., Syn. Pl. 1: 133. 1805. LECTOTYPE: H. obovata Pers., “Higginsia” orth. mut. Steudel Nom. ed. 2, 1: 767. 1 an a DUE H. & B B., PI. A Aeq. 2: 165. 1817. TYPE: E. caripensis H. & B. Campylobotrys Lem., Fl. Serres Jard. Eur. 3: 269. 1847. rype: C. discolor Lem. Shrubs, subshrubs or small trees, rarely herbs. Leaves membranous to thickly chartaceous, usually replete with cystoliths, petiolate or sessile, the stipules de- ciduous. Inflorescences axillary, occasionally subtended by roots (rhizomatous), cymose, usually pedunculate, the flowers spreading or conglomerate. Flowers usually pedicellate, the hypanthium with numerous cystoliths, the calycine tube short, the teeth often 4, usually shorter than the hypanthium; corolla tube usually reduced, glabrous within, the lobes imbricate; stamens 3-5, the anther connective occasionally produced apically; style elongate, slender, glabrous, the stigmas 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 223 usually 2, if connate then clavate, linear or lanceolate. Fruits baccate, usually 2-celled; seeds numerous, small, foveolate. Hoffmannia ranges from Mexico to Argentina. There are more than 150 species in the genus, 1⁄2 of which may be reduced to synonomy when a monograph of the genus is written. This is the most challenging among the larger genera of Rubiaceae in Middle America. The fact that the stipules drop off when the leaf buds emerge contributes to the taxonomic difficulty of the genus. The flowers and fruits exhibit little variability for the size of the genus. Literature: Dwyer, J. D. 1969. The Genus Hoffmannia (Rubiaceae) in Panama. Annals of the Missouri Botanical Garden 56: 269-286. Standley, P. C. 1934a. North American Flora. Rubiales—(Pars) 32: 190—208. Williams, L. 1973. Hoffmannias from Mexico and Central America. Fieldiana (Botany) 36(6): 51—60. Leaves bullate . Leaves with lateral veins 22-25 ___________---------------------------------------- ¢ 22. Hie ind d Leaves with lateral veins 8-14 ______---_------------------------ RENE S aA H. bullata l'. Leaves smooth, not bullate Petioles with conspicuous lateral inflations (vescicles). Inflorescences axillary and occasionally t oe inflations of petioles, if present, poorly developed; lateral veins of leaves ca. 10 ------------------------------ . H. morii 4'. Pe diia. strictly axillary; inflations y^ petioles well defined; lateral veins of leaves 11- 5. Leaf blades to 25 cm long; petioles to 4 cm long; fruits red -~--~ 26. H. vesciculifera 5'. Leaf blades to 40 cm long; petioles to 13 cm long; fruits white -—— 12. H. kirkbridei a . Petioles without conspicuous lateral inflations. Leaves petiolate. 7. Hypanthium with hairs to 1.7 mm long; inflorescence usually contracted and pe- duncle short. 8. Inflorescences densely capitate and fasciculate, the a? much reduced or absent, flowers not unilaterally or helicoidally dispos 9. Small trees; petioles 0.5-0.8 cm long; stigmas eee cease I; - aeruginosa 9'. Herbs; petioles 1—4 cm long; stigmas linear oblong ---------- . H. areolata 8'. Inflorescences patulous. the peduncle to 1.5 cm long, flowers a and helicoidally disposed ---------------------------------------------------- 10. H. gentryi - '. Hypanthium with hairs less than 1 mm long or absent; inflorescences usually pedunculate and spreadin 10. Fruits (immature) at least 2 times longer than wide; stigmas linear oblong, spreading, ca. !⁄4 the length ani the style. ll. Peduncles absent or to 1 c a plants terrestrial, 1.5-2.5 m tall. 12. Leaves chartaceous, the lateral veins ca. 4; ae aha to 10 m TIRE ——— — ———MO8Á 23. H. rexmontis . Leaves membranou us; lateral veins 10-11; peduncles padlike an scarcely measurable, occasionally to 8 mm long -_----------------- m . H. RE 11’. Peduncles to 4. 5 cm long: plants probably epiphytic, ca. 0. mt . H. po very 10'. Fruits less than 2 times longer than wide; stigmas if linear dn short and spreading. 13. C Mara Tohes 2 or more times longer than the tube; leaf sang often with patches of raphides on upper surface forming a thin crust ....... - 4, eee s 13’. Corolla lobes less than 2 ‘times ‘the d of the tube; leaf blade lacking patches of raphides forming ac 14. Calycine s to 12 mm in fruit; edi visible poa the w of the fruit... eo Ssh eiocnAl ne sce . H. longicalycina " ord abe 1-3 mm long; seeds not visible as the w of the R ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 5, piger subtended by roots; stamens 3-4 16. Leaves oblong rotund; flowers yellow S yellow brown or ve cw red; corolla lobes ca. 3 times the length of l corolla tube; anthers 4; stigmas clavate ain d 7. H. davidsoniae ll: — E Lepves oblong or m elliptic; flowers red; corc the length of the corolla pus anthers usually 3; s narrowly oblong : A elas ord aere not subtended by roots; stamens 4(—6), often n H. ostaur Ta Leaves in bunc pairs MA =e flowers per axil = = witheringioides 17’. Leaves in pairs similar ir in size; flower xil. her connective not Sac ace or a a 19. Flowers ca. 2 cm long at e dra rud s 3-4 loc lobes 4—5(—6); stamens 3-4; ova ulate |... "OR s s TREE H. lewisiana 19’. Flower s to l. 5c em n long at vui calycine lobes al arly 4; stamens (3—)4(—5): stigmas 2-3: ovaries 2-3 loculate. 20. Leaves membranous; end 3(-4) .. PONES . H. ostaurea 20’. Leaves thin chartaceous * subcoria ceous; stamens 4(— 21. Stigmas lanceolate, va tipped n . H. ane ', Stigmas capitate pur to linea oblong, blunt tipped. 22. Leaf blade. rubescent, the pigment extractable « on boil ing; flowers subse 24. N "— H. rubripiementa 22'. Leaf blades not rubes not yielding pigment on oil ing; Wee Obviously pedi- 23. [TR Sea c s subcoriaceous, not m branous and done io on drying. 24. Leaves widely ob- long to ovate sub- tund, to 11 cm coriaceous, MR 6. Henna . Leaves elliptic obovate elliptic, 9- N AR ou . H. pilin 23’. Leave: es OEE N and ee on drying. 25. Lea to 25 cm ionis podes 0.5 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 225 mm wide; calycine lobes to 1.1 mm long; corolla tube pubescent; sta- mens 3(-4) |... -— 19. H. ostaurea 25’. Leaves 10-16 long: rusos ca. 0.25 mm wide; ca- lycine lobes 0.4- 0.8 mm long; corol- la tube glabrous; stamens4 |. . . EROS 5. H. capillacea '. Anther connective produced apically or basally. 26. Leav 9.5 cm wide; anther connective produced basally; flowers mu CDS PR ETE . H. . fimbrianthera 26'. Leaves usually 3 em wide; a connective produced REPAS flowers = oo a '. Leaves sessile. 27 Piae piden with peduncles 3-6 cm long 28. Leaves rounded toward the apex: cal eine lobes 3-4 mm long |... Bere eee seat eee eee l. E denii 28'. Leaves acute or -acuminate toward the apex; calycine lobes less pns mm lon 29, Inflorescences solitary in axils; plants prostrate 25. H. subauriculata 29'. Inflorescences several per axil or cauliflorous; plants Dpp - a SER _ EE 2. x araneopedaria 21". Inflorescences with peduncles ca. 30 cm n long RE . H. manussatani 1. Hoffmannia aeruginosa Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1313. 1938. rype: Costa Rica, Austin Smith A86 (F). Shrubs to 4 m tall, the branchlets slender, terete, the bark pale brown, fer- rugineous villosulose, the hairs deciduous leaving a minutely tuberculate surface. Leaves oblong, to 20 cm long, to 8 cm wide, short acuminate at the apex, widely to narrowly cuneate at the base, the costa subplane and slender above, promi- nulous beneath, the lateral veins ca. 15, arcuate, prominulous, the intervenal areas often appearing lineolate under magnification, stiffly chartaceous, scarcely discolorous, drying green or brown, glandular shiny, glabrescent, minutely foveo- late, the raphides moderately abundant, white, divergent, to 0.5 mm long, more conspicuous beneath, the costa ferrugineous pubescent, sometimes glabrescent: petioles 0.5-0.8 cm long, curved or reflexed, densely ferrugineous villosulose, to 3 mm wide; stipules not seen. /nflorescences in the uppermost axils, subsessile glomerules of 3-10 flowers, the reduced peduncles ferrugineous pubescent or purple villose. Flowers sessile, with the hypanthium to 1.7 mm long, villose with elongate, soft and ferrugineous hairs; calycine teeth subulate, to 1.5 mm long: corolla densely villose outside, the lobes acute: anthers ca. 3 mm long, acute at the apex, the connective not produced at the apex or base: stigmas 2, rotund or clavate, the ovary 2-loculate. Fruit not seen. The species occurs in Costa Rica and Panama. The collection of Austin Smith (H659) from Guadalupe de Zarcero, Costa Rica, has the notes on the label: "Growing in deep forest shade; gregarious but very local. Base of the stem dull. 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 dark brown. Buds deep chokeberry red. Petioles and some of the veins of the under surface of the leaves tinged with pinkish red." BOCAS DEL TORO: Chiriqui Lagoon, Wedel 2289 (MO). 2. Hoffmannia araneopedaria Dwyer.? TYPE: Panamá, Gentry & Mori 14166 (MO, holotype). Shrubs to 2 m tall, stem simple, lignose, terete, to 1.8 cm in diam.; bark with inconspicuous longitudinal lines, glabrous; nodal scars ringlike, the nodes well spaced. Leaves sessile, oblong, 12.5-32 cm long, 6-9 cm wide, short acuminate, obtuse or widely deltoid toward the apex, the acumen 1-1.2 cm long, basally attenuate and elongate cuneate below the middle, ultimately 8-10 mm wide and truncate or scarcely auriculate, the costa scarcely prominulous to immersed above, prominulous beneath, the costa slightly prominulous above, prominulous on the lower side only near the apex, prominent at the base, 2.5 mm wide, the lateral veins arcuate, the undulate submarginal veins inconspicuous, either absent or to 0.5 cm from the margin, the intervenal areas reticulate, stiffly papyraceous, concolorous, drying brown, glabrous above, minutely puberulent on the costa and veins beneath. /nflorescences cauliflorous or axillary; peduncles 8-12, 0.8- 3 cm long, wiry, unequal, radially disposed from a single axil, arising from a compressed hemispherical swollen pad, ca. 1 cm wide, 0.5 cm long, the flowers to 6 on each axis, secundly disposed. Flowers (in bud) with pedicels red when living, 5-7 mm long, the hypanthium turbinate, | mm long, glabrous, ca. 8-ribbed, the calycine cup ca. 0.2 mm long, subturgid, the 4 lobes widely triangular, ca. 0.3 mm long, glabrous; corolla yellow, the tube carnose, glabrous, the 4 lobes triangular oblong, 2.8 mm long; anthers subbasifixed, the filaments short; ovarian disc cushionlike, narrower toward the apex, ca. 0.6 mm long, the style slender, the 2 ovate stigmas to 0.8 mm long. Fruits (perhaps immature), turbinate or oblong rotund, ca. 0.6 cm long, truncate, glabrous, not ribbed, white in living state. Hoffmannia araneopedaria is known only from Panama. It is most closely related to H. subauriculata Standley which has similarly shaped, sessile leaf ades. The specific name araneopedaria means "resembling a spider's legs” a reference to the several wiry peduncles suggesting the legs of a "Daddy long- legs." Some inflorescences are cauliflorous, an uncommon condition in Hoff- mannmia. DARIÉN: Cerro Mali on Colombian border, S slope just below summit, lower montane wet forest- premontane wet forest transition, 1400 m, Gentry & Mori 14166 (MO). 2 ont d araneopedaria Dwyer, spec. nov. Frutices ad 2 m alti, caule simplici glabro. Folia sessilia oblonga, 12.5-32 cm longa, 6-9 c m lata, versus apicem B vel lato-deltoidea, venis late- ralibus ca. 20, Tigido- papyracea supra labia subtus minuto- idus e scentiae pedunculis 8— 12 ab una axilla radiatis et ab area tumida compresso-haemosphaeric . | em lata, ca. 0.5 cm alta, floribus in axe utroque ad 6 secundatim dispositis. Flores mecs llis, a pals calycis ca. 0.1 mm longa, dentibus 4 lato-triangularibus, ca. 0.3 mm longis, glabris: corolla lutea tubo glabro lobis 4 oblongo- triangularibus, ca. 2.8 mm longis. Fructus his fortasse juveniles turbinati vel oblongo-rotundi, ca. 0.6 cm longi, truncati glabri non costati, in vivo albi. 1980] DWYER— FLORA OF PANAMA (Family 179. Rubiaceae) 229 3. Hoffmannia areolata Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 52. 1940. TYPE: Panama, Davidson 40 (F, holotype; isotype, MO). Herbs to 2 m tall, the stems smooth, glabrous or villose. Leaves elliptic, to 25 cm long, to 8 cm wide, attenuate acuminate at the apex, the acumen to 3 cm long, cuneate at the base, the costa immersed above, plano compressed beneath, the lateral veins ca. 25, arcuate, glabrescent above, appressed villosulose on the costa beneath, chartaceous, concolorous, the raphides numerous, scattered, white, linear, to 0.3 mm long; petioles 1-4 cm long; stipules not seen. /nflores- cences sessile, glomerules in the uppermost axils, to 2 cm long or longer, often wider than long. Flowers yellow or greenish, the hypanthium oblong, covered with elongate ferrugineous hairs to 1.6 mm long, the calycine lobes 4, ca. 3.5 mm long, obtuse; corolla ca. 4.5 mm long, the tube cylindrical, ca. 1 mm wide, the lobes shorter than the tube; stamens 4, ovate, ca. 1.5 mm long, the connective not produced at the apex or the base; stigmas 2, oblong, spreading, ca. 1.5 mm long, the style ca. 2.5 mm long. Fruits scarlet (fide Davidson). Hoffmannia areolata is known only from Panama. CHIRIQUÍ: Boquete, near Bajo Chorro, Davidson 40 (F, GH, MO); Woodson & Schery 661 (F, GH, MO). 4. Hoffmannia bullata L. O. Williams, Fieldiana: Botany 36: 52. 1973. TYPE: Guatemala, Steyermark 38719 (F, holotype). Subshrubs to 0.3 m tall, the stems simple, the lowermost nodes rhizophorous, the hairs dense, ferrugineous, the lenticels many, the nodes moderately well spaced. Leaves narrowly obovate oblong, 5-15 cm long, 1.5-5.0 cm wide, often slightly falcate, widely deltoid or rounded at the apex, the acumen inconspicuous, obtuse or cuneate toward the base, often abruptly contracted toward the base, the costa plane above or immersed, prominulous beneath, the lateral veins 8-14, arcuate, conspicuous and prominulous beneath, chartaceous, usually bullate at least on drying, discolorous, drying tan or silvery, glabrous above, the costa and veins below with golden, dense hairs, the raphides minute, white; petioles ca. 1.3 cm long, pubescent; stipules not seen. Inflorescences axillary; peduncles slender, to 2.5 cm long, ca. 0.7 mm wide. Flowers with hypanthium oblong or turbinate, glabrous, lightly costate, often seemingly alate on pressing, the teeth 4, to 1.5 mm long: (Williams) "dull brick red; calyx and hypanthium 3-4 mm long, the hypanthium turbinate, glabrous, the calyx lobes triangular or lance triangular, acute, sparsely villosulose: corolla about 1 cm long, glabrous, the lobes oblong, obtuse or acute, 2-3 times as long as the tube; stamens with short filaments, the anthers exserted." Fruits red; pedicels to 2 mm long, oblong or turbinate, 6—7 mm long, truncate at the apex, 4-angled, smooth, glabrous, lightly costate, the raphides abundant, the persistent calyx lobes obtuse. Hoffmannia bullata is known from Mexico to Panama. Unfortunately the single Panamanian collection cited below lacks flowers so the floral description above is largely from Louis Williams’ original diagnosis. Steyermark 37241 (F), from the slopes of Volcan Tajumulco, Guatemala, 1,300-1,500 m, notes that the flowers have: "calyx pale green; corolla lobes spreading, salmon-brick color: anthers pale yellowish white, as in stigma." 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Hoffmannia bullata is closely related to H. pusulata Dwyer, which has more numerous lateral veins and has been collected at a much lower elevation, less than 200 m. PANAMA: Cerro Campana, 800 m, Croat 12137 (MO). Hoffmannia capillacea Dwyer, Ann. Missouri Bot. Gard. 56: 274. 1969. TYPE: Panama, Elias 1783 (MO, holotype). Subshrubs, the stems lignose, smooth, glabrous, the petiole scars somewhat conspicuous, the nodes ca. 1 cm apart. Leaves oblong, 10-16 cm long, 5-7 cm wide, subacute and short acuminate at the apex, the acumen to | cm long, acute at the base and scarcely symmetrical, the costa subplane on both sides, the lateral veins ca. 10, subprominulous, the intervenal areas prominulous and spreading, membranous, discolorous, glabrous; petioles to 5 cm long, ca. 0.2 cm wide, lightly alate, strict; stipules not seen. /nflorescences cymes in the uppermost axils, gla- brous; peduncles 1-15 mm long, ca. 0.3 mm wide, capillaceous; flowers 2-4 on spreading branches, often somewhat radially disposed. Flowers (in bud) with the pedicels 6-8 mm apart, capillaceous, to 0.4 mm wide, glabrous: hypanthium ca. 3 mm long, squarish, glabrous, carnose, the lobes 4, acute, glabrous; stamens 4, the anthers 2.5-3 mm long, the connective not produced at the apex or base, the filaments short, ca. 0.7 mm wide; stigmas 2, oblong rotund, ca. 1.5 mm long, wider than the style. Fruits not seen. uA Hoffmannia capillacea is known only from Panama. : Alto Calvario, 7 km N of El Copé, Folsom 3314 (MO). SAN BLAS: Headwaters of Rio Kalana, Elias 1783 (MO). 6. Hoffmannia cercidifolia Dwyer, Ann. Missouri Bot. Gard. 56: 275. 1969. TYPE: Panamá, Duke 15597 (MO). Shrubs to 1% m, the branchlets slender, smooth, glabrous. Leaves ovate subrotund or widely oblong, 7-11 cm long, 5.5-8 cm wide, short acuminate and deltoid apically, the acumen falcate or straight, to 1 cm long, ultimately subacute, basally obtuse or truncate, the costa subplane above with a thin median ridge, subplane beneath, the secondary veins 7-9, arcuate, prominulous beneath, dis- colorous, dark red brown and glabrescent above, beneath glabrous or golden puberulent on the costa and veins, glistening, above honeycomb-like under mag- nification; petioles slender, to 4.5 cm long, ca. 0.15 cm wide; stipules not seen. Inflorescences to 1 cm long, the flowers few, nodding, the peduncle reduced. Flowers (buds only) with the pedicels short, the hypanthium short, campanulate, ribbed, glabrous, the calycine cup short and scarcely measurable, the 4 triangular teeth ca. 1 mm long, carnose, glabrous except puberulent marginally; corolla white with the tube short, ca. 2.5 mm long, subcarnose, pubescent outside, gla- brous within, the lobes 4, oblong, ca. 4 mm long; stamens 4, the anthers ca. 3.5 mm long, ca. 0.8 mm diam., the connective not produced at the apex or base, the filaments compressed; style ca. 5.5 mm long, the 2 stigmas clavate capitate. Fruits not seen. The species is known only from Panama. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 229 DARIEN: Cloud Forest, Cerro Campamento, S of Cerro Pirre, Duke 15597 (MO). VERAGUAS: 16 km NW of Santa Fe, road to Calovébora, 300-500 m, Mori & Kallunki 6119 (MO). 7. Hoffmannia davidsoniae Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 53. 1946. Type: Panama, Davidson 216 (F, holotype; isotype, MO). Herbs to 1.5 m tall, the stems terete, smooth, glabrous, usually drying tan, the internodes to 9 cm long. Leaves oblong rotund, 8-17 cm long, 4-12 cm wide, rounded and acuminate at the apex, the acumen 0.5-1.8 cm long, basally rounded or tapering widely, abruptly cuneate, the costa prominulous to subplane above, prominent beneath, the lateral veins 12-15, widely arcuate, prominulous, 0.5-1.2 cm apart, thinly and stiffly succulent, lustrous, brown, white puberulent to gla- brous above, minutely puberulent on veins beneath; petioles 2-7 cm long; stipules not seen. /nflorescences cymose, mostly subtended by roots, the peduncles elon- gate, strongly ascending, 5-10 cm long, 0.1—0.2 cm wide, rubescent, glabrous, terminated by 3-4, usually radially disposed and spreading branches, the cymules ascending, each branch 7-17-flowered. Flowers with the pedicels stiff, glabrous, to 5 mm long, ca. 0.3 mm wide; buds obviously acute at the apex; hypanthium strongly 6—8-ribbed, emerald green to red in living state, to 3 mm long, the ca- lycine lobes flaring, deltoid, 0.3-0.5 mm long; corolla to 6 mm long, the lobes 4, much longer than the tube, glabrous; anthers 4, oblong, to 3.8 mm long, the connective not produced at apex or base, the filaments to 0.6 mm long; ovary 2-loculate, the style 4 mm long, the stigmas 2, clavate oblong, ca. 1.1 mm wide, each somewhat uncinate. Fruits oblong to 3.5 mm long, truncate, the persistent calycine teeth scarcely measurable, glabrous, delicately costate. Hoffmannia davidsoniae is known only from Panama. My own collection notes on this species (Dwyer 8789) are more elaborate than usual: "To 1 m tall; leaves smooth, lustrous, brown, stiffly succulent; inflores- cence 17-flowered, glabrous, the peduncle thick, red green; hypanthium strongly angular, green; ovarian disc cushionlike; corolla rose, the tube glabrous, in bud flushed with red or pink, the anthers 4, the stigmas 2, clawlike, angular.” RIQUI: Cerro Horqueta, 7000 ft, Blum & Dwyer 2623 (MO). Río Chiquero, D'Arcy 70983 (MO). es Chorro, 6000 ft, Davidson 216 (F, MO). Cerro Horqueta, Dwyer & Lallathin 8789 (MO). j uebrada Chiquero, 1500 m, Woodson & Schery 595 (MO). COcLE: Cero Pilon, 2900 ft, Lallathin 133, 135 (both MO). 8. Hoffmannia eliasii Dwyer, Ann. Missouri Bot. Gard. 56: 276. 1969. TYPE: Pan- amá, Kirkbride & Duke 1129 (MO). Herbs to 10 cm high, succulent, the stems simple, horizontal at the base, lignose, 15-20 cm long, to 1.2 cm wide; roots fibrous, lignose. Leaves oblong or obovate elliptic, to 25 cm long, 9.5 cm wide, deltoid or subobtuse at the apex, scarcely acuminate, cuneate to narrowly cuneate at base, the costa prominulous above to subplane beneath, often lightly alate, the secondary veins 12-16, slightly arcuate, prominulous, to 1.8 cm apart, the intermediate veins irregular and soon branching, then evanescent, the ultimate venules spreading pinnatiform, the mar- ginal vein conspicuous, ca. 7 mm from the margin, stiffly chartaceous, scarcely discolorous, glabrescent, the hairs on the costa and the veins few; subsessile or petiolate, the petioles to 12 cm long, often lightly alate; stipules not seen. /nflo- rescences 1-3 cm long, to 2 cm wide, subtended by roots, the peduncles 2 per 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 axil, often stiff, to 4.5 cm long, often marcescent, canaliculate, pilose, the hairs weak and spreading. Flowers sessile or on pedicels to 4 mm long; hypanthium campanulate, ca. 2 mm long, glabrous, black when dry, ribbed, the calycine teeth 4, somewhat unequal, erect, subulate, 1-1.8 mm long, minutely puberulent; co- rolla glabrous, the tube cylindrical to 2.5 mm long, the lobes 4, lanceolate, longer than the tube; stamens 3, the anthers 3-4 mm long, subsessile, the filaments ca. 0.2 mm long, the style narrow cylindrical, the connectives not produced at the base or the apex; stigmas 2, unequal, oblong, to 0.7 mm long, scarcely wider than the style. Fruit (immature) with pedicels strongly ascending, to 4 mm long, dif- fusely pilose, oblong, to 5 mm long, to 2.5 mm wide, the calyx persistent, the lobes 1-2.5 mm long, red when dry, moderately ribbed, minutely puberulent. The species is known only from Panama and nearby Colombia. resin Caná-Cuasí Trail, Chepigana, 4000 ft, Terry & Terry 1508 (F, GH, MO). Rio Cuasi 2.5 mi S of Tres Bocas, Kirkbride & Duke 1129 (MO). SAN BLAS: Headwaters Rio Malatia, Elias 1779 (MO). Mountains above Puerto Obaldia, Gentry 1504 (MO). COLOMBIA: CHOCO: Rio Nercua, Quebrada Barrial and Quebrada Ambrosio near Camp Curiche, Duke 11438 (MO). 9. Hoffmannia fimbrianthera Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969. TYPE: Panama, Kirkbride & Duke 556 (MO, holotype). Subshrubs, the stems smooth, reed-like, glabrous, terete, the internodes to 15 cm long. Leaves oblong rotund, often subfalcate, to 19 cm long, to 9.5 cm wide, deltoid at the apex, short acuminate, the acumen to 1.5 cm long, basally acute or narrowly cuneate at the base, the costa plane above, subprominent beneath, the lateral veins ca. 15, divergent-arcuate, conspicuously prominulous beneath, the veinlets spreading pinnatiform, the submarginal vein ca. 8 mm from the mar- gin, papyraceous, scarcely discolorous, drying grey, conspicuously honeycomb- like under magnification, glabrescent above, granulose puberulent only on the veins beneath; petioles slender, lax, to 7 cm long, to 1.2 mm wide, minutely bby puberulent; stipules not seen. Inflorescences sessile, the flowers few, (3-)5(-10). Flowers subsessile or on pedicels to 1 mm long; hypanthium rotund oblong, ca. 3 mm long, somewhat smooth, stiffly carnose, ferrugineous puberu- lent, the calycine lobes 4, spreading uncinate subulate, to 1 mm long, puberulent; corolla carnose, ferrugineous puberulent outside, minutely farinose within, the lobes 4 (?); stamens 4, the anthers oblong, to 3.2 mm long, minutely apiculate, the connective petaloid fimbriate at the base for 0.3 mm long; style to 3.5 mm long, 0.2 mm wide, wider toward the apex, the stigmas 2, clavate, ca. 0.35 mm wide. Fruits not seen. Hoffmannia fimbrianthera is known only from Panama. BOCAS DEL TORO: Río Teribe, just below Puerto Palenque, 350 m, Kirkbride & Duke 556 (MO). 10. Hoffmannia gentryi Dwyer.® type: Panamá, Gentry & Mori 14061 (MO, holotype). Subshrubs to 1.5 m tall, the branchlets weak, angular when dry, villose, the hairs persistent or deciduous, curled, swollen at the base, the nodes moderately "" Hoffmannia gentryi Dwyer, spec. nov. Suffrutices ad 1.5 m alti, ramulis debilibus. Folia oblonga vel obovato-oblonga, 12-20 cm longa, 5-10 cm lata, plerumque versus apicem deltoidea, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 231 well spaced. Leaves oblong or ovate oblong, 12-20 cm long, 5-10 cm wide, deltoid toward the apex, occasionally acute, acute or obtuse at the base, rarely subtruncate, the costa prominulous on both sides, the lateral veins ca. 10, arcuate and scarcely prominulous, the intervenal areas lightly reticulate, the undulate submarginal vein to 5 mm from the margin, membranous, scarcely discolorous, hairy on both sides, the hairs on upper surface few, subulate, spreading or ap- pressed, mostly confined to the costa and the veins, below more dense, often curled and giving a webby effect; petioles weak, slender, often curved, to 4.5 cm long, densely hairy; stipules deciduous. Inflorescences axillary, to 2.5 cm long; peduncle slender, to 1.5 cm long, ca. 0.3 mm wide, the cymes 4—-5-flowered, the flowers unilaterally and helicoidally disposed. Flowers subsessile, densely villose, the hypanthium turbinate, to 2.5 mm long, the lobes 4, narrowly oblong; corolla light green, subcampanulate, the tube about as long as the calycine teeth, perhaps glabrous outside, raphides abundant, the lobes 4, longer than the tube; stamens 4, erect, the anthers oblong, ca. 4 mm long, subsessile, attached near the mouth; style slender, exceeding the anthers, the stigmas connate, clavate, ca. 1.5 mm long. Fruits not seen. Hoffmannia gentryi is named in honor of one of the co-collectors of the type, a distinguished student of the flora of Panama. It is known only from Panama, although it no doubt occurs in adjacent Colombia. It is distinguished by its mem- branous leaves adorned with adpressed and often curled hairs; the hypanthium is replete with elongate hairs and the calycine cup has teeth as long as the corolla tube. The flowers being unilaterally and helicoidally disposed distinguishes it from H. aerolata to which it seems closely related. pa Cerro Tacarcuna, summit camp, along stream N of camp, 1550-1650 m, Gentry & Mori 14061 (MO 11. Hoffmannia haydenii Dwyer, Ann. Missouri Bot. Gard. 56: 277. 1969. TYPE: Panamá, Hayden 177 (MO, holotype). Herbs, the stems drying quadrangular. Leaves sessile, obovate elliptic or oblong, to 35 cm long, to 14 cm wide, rounded toward the apex, short acuminate, the acumen to 1.5 cm long, basally narrow cuneate and ultimately auriculate, the costa prominulous, slender and ridgelike above, beneath subplane, to 2.5 mm wide proximally, puberulent, the lateral veins ca. 20, somewhat plane above, prominulous beneath, coriaceous, discolorous on drying, glabrescent and brown above, puberulent on costa and veins beneath; stipules not seen. Inflorescences 1.5-7 cm long, the peduncles 1.2-6 cm long, subtended by roots, capillaceous, ca. 0.3 mm wide, the branches strict, puberulent, red in living state, to 2.5 mm long, puberulent. Flowers with buds cuneate and acute at the apex; hypanthium puberulent, scarcely ribbed, the calycine lobes 4, erect lanceolate, 3-4 mm long, purple red when dry, puberulent; corolla lobes 4, ca. 5 mm long, puberulent outside, longer than the tube; stamens 4, the anthers ca. 4 mm long, the connec- venis lateralibus ca. 10, gajene utrinque ciliata praesertim subtus; petiolis PrE ad 4.5 longis. Inflorescentiae ad 2.5 cm longae, pedunculo gracilissimo, cymis floribus 4-5 DEDE W: dE Flores hypanthio dense elongato-ciliato dentibus 4 ad 3 mm longis; corolla tubo brevi lobis m longis: staminibus subsessilibus, antheris 4 oblongis, ca. 4 mm longis, exsertis. Fructus non visi. 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 tive not produced at the apex or the base, the filaments short; stigmas 2, aggre- gated. Fruits not seen. Hoffmannia haydenii is known only from Panama. CHIRIQUI: Finca Collins, ca. 5000 ft, Havden 117 (MO). 12. Hoffmannia kirkbridei Dwyer, Ann. Missouri Bot. Gard. 56: 278. 1969. TYPE: Panamá, Kirkbride & Duke 429 (MO, holotype). Subshrubs to 2 m tall, the stems glabrous, drying minutely rugulose. Leaves elliptic oblong to 40 cm long, to 20 cm wide when dry, the costa prominulous and canaliculate above, subprominent beneath, distinctly corky alate toward the base, the lateral veins 15-25, ca. 3 cm apart, arcuate, scarcely prominulous above, conspicuously prominulous beneath, the intermediate veins more divergent, soon evanescent, the veinlets spreading, irregularly pinnatiform with patches of simple appressed hairs on the costa and veins beneath, the hairs few, stellate, stiffly chartaceous, discolorous when dry: petioles to 13 cm long, ca. 3 mm wide in the middle, inconspicuously swollen below the base of the blade, the vescicles 0.6— 10 cm long, ca. 1 cm wide when dry. Inflorescences often 3 per axil, the peduncles reflexed, 1-1.5 cm long, ca. 0.13 cm wide, with minute patches of crystalline hairs and raphides, the hairs few, to 0.7 mm long, rarely stellate. Flowers (juvenile fruit) subsessile; hypanthium short, moderately villose, the raphides thick, white: calycine cup somewhat turgid, villose, the lobes 4, thickly carnose to 4.5 mm long, to 3.5 cm wide, ciliate, the veins few, ascending, the wall of the hypanthium thick. Fruits (immature) rhomboid, ca. 6 mm in diam.: seeds thickly reticulate. Hoffmannia kirkbridei is known only from Panama. ‘AS DEL TORO: Buena Vista, Almirante, Cooper 225 (F). Quebrada Huron, ca. 300—400 ft. Kirkbride & Duke 429 (MO). 13. Hoffmannia lancistigma Dwyer, Ann. Missouri Bot. Gard. 56: 279. 1969. TYPE: Panamá, Dwyer & Hayden 7735 (MO, holotype). Shrubs, the stems canelike, terete, smooth, the hairs minute, dense, ferru- gineous, moniliform, deciduous. the nodes 2.5-3.5 cm apart. Leaves strongly ascending, well spaced, oblong, to 19 cm long, to 9 cm wide, obtuse at the apex, scarcely short acuminate, ultimately obtuse, cuneate at the base, often asym- metrical, the costa prominulous above and beneath to 0.15 cm wide, the lateral veins ca. 15, arcuate, prominulous beneath, the intermediate veins spreading pinnatiform, papyraceous or stiffly chartaceous, minutely granulose and honey- comb-like under magnification, the hairs few beneath, except on the densely pubescent costa; petioles to 7.5 cm long, ferrugineous; stipules not seen. Inflo- rescences usually 1 per axil, often arising from a leafless part of the twig, the peduncle ascending, to 10 cm long, slender, ca. 2 mm wide, strict or arcuate, glabrescent or with few crystalline hairs. Flowers glabrous (?); hypanthium ob- long, to 3.8 mm long, ribbed, the calycine teeth 4, erect, triangular, ca. | mm long: corolla tube thin carnose, the lobes 4(—6), ca. 4.5 mm long, carnose; stamens 4, the anthers ca. 3.8 mm long, to 1.2 mm wide, the connective not produced at 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 233 apex or base, the filaments ca. 1 mm long, flat, to 1 mm wide; style ca. 4 mm long, the stigmas 2, lanceolate, ca. 2.8 mm long, to 0.8 mm wide, thick, wider than the style, constricted medially. Fruits not seen. Hoffmannia lancistigma is known only from Panama. CHIRIQUÍ: Cerro Horqueta, ca. 6000 ft, Dwyer & Hayden 7735 (MO, SCZ, UC). 14. Hoffmannia lenticellata Hemsl., Diag. Pl. Nov. 30. 1879. sYyNTYPES: México, Botteri 974; Bourgeau 2992; Linden I, none seen. Shrubs to 1.5 m, the branchlets smooth, minutely puberulent to glabrescent, the nodes well spaced, the pith porate. Leaves elliptic to oblong elliptic, 8-15 cm long, 2.5-5 cm wide, widely deltoid or rounded at the apex but distinctly short acuminate, the acumen to 1.5 cm long, slender, often falcate, attenuate acute toward the base, slightly decurrent along the petiole, the lateral veins 14—20, arcuate, plane and somewhat inconspicuous above, prominulous beneath, not forming a distinct marginal vein, the veinlets inconspicuous, rigidly papyraceous, scarcely discolorous, presumably glabrous above, glabrescent to minutely pu- berulent on the midvein and the lateral veins, often the upper surface with small patches of white round crystals; petioles 0.5-2.5 cm long; stipules not seen. Inflorescences axillary, sessile or on reduced peduncles, the cymes few flowered. Flowers not seen. Fruits somewhat congested, puberulent, to 4 cm long, the peduncle absent or to 0.5 cm long; pedicels to 2 mm long, the pericarp oblong, to 4 mm long, truncate, the lobes 4, erect triangular or oblong, to 1 mm long, acute to obtuse, puberulent, concave within, rigid, the persistent ovarian disc compressed annular, the fruit body with 8 narrow but prominent ribs. Hoffmannia lenticellata extends from Mexico to Panama. This species demands extensive study. Many collections labelled H. lenticel- lata from Middle America have been misidentified. The value of the patches of crystals on the leaf blades as a distinguishing character needs assessment. Stand- ley in his key to species of Hoffmannia in North American Flora used the char- acter that the corolla lobes are 2 times as long as the tube; no flowers were available on the single collection from Panama seen. CHIRIQUÍ: Road Between Cerro Punta and Las Nubes, 1800-2100 m, Croat 26399 (MO). 15. Hoffmannia lewisiana Dwyer, Ann. Missouri Bot. Gard. 56: 279. 1969. TYPE: Panamá, Ridgway & Solis 2413 (MO). Shrubs, the branchlets smooth, glabrous, canaliculate. Leaves oblong, 17-19 cm long, 7-8 cm wide, somewhat attenuate acute and acuminate at the apex, the acumen to 1.5 cm long, acute and almost symmetrical at the base, the costa subplane above, canaliculate, prominulous beneath, to 0.15 cm wide, the lateral veins ca. 12, plane above, prominulous beneath, arcuate, ultimately spreading reticulate, papyraceous, concolorous, glabrous above, pubescent beneath only on the costa and veins, petioles to 4.5 cm long, glabrous; stipules not seen. Inflorescences to 6.5 cm long, spreading, the peduncles short, the branches nu- merous, 2.4 cm long, 0.8-1.8 mm wide, the cymes few, alternate or often sub- 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 radiate. Flowers with the buds clavate; pedicels 1-7 mm long, ca. 0.4 mm wide, densely white puberulent; hypanthium oblong, to 7 mm long, carnose, puberulent and conspicuously ribbed, the teeth 4—6, erect, deltoid, unequal, often one con- spicuously reduced, to 2.5 mm long, puberulent; corolla with the tube to 11 mm long, carnose, the lobes 5, to 5.5 mm long, acute, puberulent outside; stamens 4-5, the anthers ca. 3.5 mm long, ca. 1.3 mm wide, the connective not produced at the apex or base, the filaments compressed, ca. | mm wide; style 12-14 mm long, the stigmas (2—)3-4, capitate. Fruits not seen. Hoffmannia lewisiana is known only from Panama. CHIRIQUI: Cerro Punta, 6800 ft, cloud forest and distrubed edge, Ridgway & Solis 2413 (MO). 16. Hoffmannia longicalycina Dwyer, Ann. Missouri Bot. Gard. 56: 280. 1969. TYPE: Panamá, Kirkbride & Duke 1278 (MO, holotype). Subshrubs, the branchlets smooth, glabrous, the nodes ca. 1 cm apart. Leaves oblong, tapering cuneately toward the apex, acuminate, the acumen short, to 1 cm long, ultimately acute, narrowly cuneate at the base, the costa prominulous and ridged beneath, to 2 mm wide, the lateral veins ca. 16, spreading arcuate, prominulous, rigidly chartaceous, slightly discolorous, green when dry, glabres- cent above, appressed ciliate on the costa and the veins beneath, the hairs often rigid and appressed falcate; petioles to 10 cm long, angular to compressed, ca. 0.2 cm wide, the hairs white, thickly subulate, forming a thin crust; stipules not seen. Flowers not seen. Fruits oblong, terete in cross section, to 12 mm long, ca. 7 mm wide, delicately costate, pubescent with weak ferrugineous hairs to 2 mm long: persistent lobes of calyx 5, erect, linear lanceolate, to 12 mm long, ca. 2 mm wide, petaloid and pubescent; seeds numerous, thickly reticulate, ca. 0.5 mm in diam., visible through the thin wall. Hoffmannia longicalycina is known only from Panama. : Cana-Cuasi Trail between Cerro Campamento and La Escalera to "paramo," E of Tres Boza. Kirkbride & Duke 1278 (MO). 17. Hoffmannia manussatani Dwyer." Type: Panamá, Allen 4978 (MO, holo- type).—Fic. 50 Herb to 3 ft tall, the stem perhaps unbranched, 4-angled, smooth, glabrous, the nodes well spaced. Leaves sessile, obovate oblong or spatulate, ca. 20 cm long, 6.0-6.5 cm wide, narrowly acuminate at the apex, the acumen to 1.5 cm long, acute, the blade obviously contracted below the middle, at base vaguely auriculate, to 2.5 cm wide basally, the costa prominulous above with a central 91 p ipe manussatani Dwyer, spec. nov. Herbae caulibus 4- aei uis glabris. Folia Vides sans g onga vel spathulata, ca. 20 cm longa, 6.0-6.5 cm lata, acuminata, basi infra me- dium a uato- rA proximaliter modice auriculata venis Tee ca. 18 chartacea glabra. oxi scentiae axillares pedunculo elongato ad 30 cm longo, floribus terminalibus in caespitibus pauci- floris congestis ad 2.5 cm latis, jugis tribus bractearum bene distantibus in pedunculo, bracteis magnis plerumque ovato-oblongis, ad 9 mm longis. Flores hypanthio ferrugineo- d lobis calycis 4, ca. 3 mm er corolla lobis 4, ad 4 mm longis; staminibus sessilibus antheris 4, ad 4 mm longis. Fructus non vis 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 235 NINA NN AE Y MI "t Y UN E £e SS ¿a Ficure 50. Hoffmannia manussatani Dwyer.—A. Habit ( x15).—B. Flower bud (x5).—C. Sta- mens (x 5). —D. Style (x 5).—E. Ovary cross section (x5). [After Allen 4978. | depression, prominulous beneath, ca. 0.2 mm wide proximally, the lateral veins ca. 18, arcuate, the submarginal vein inconspicuous and proximate to the margin, the veinlets pinnatiform, chartaceous, concolorous, drying reddish, glabrous; stipules widely triangular, to 0.6 cm long, wider than long, thickly chartaceous, glabrous. Inflorescences axillary, elongate pedunculate, the peduncles flagelli- 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 form, to 30 cm long, twisted, angular, puberulent, 1.6-2.3 mm wide basally, with 4 pairs of bracts, the basal bracts much reduced, rotund, to 3.5 mm long, the other 2 pair well spaced, one pair at the apex of the lower 1⁄4 of the peduncle, the other at the base of the upper 1⁄4, rotund or ovate rotund, 1.6-2.5 cm long, obtuse or rounded at the apex, stiffly chartaceous, ascending, the costa and veins evi- dent, disposed as in the leaves, the flowers few, loosely aggregated in a terminal rotund or compressed rotund head, to 2.5 cm in diameter, presumably of few cymules; bracts oblong or ovate oblong, 7-9 mm long, stiff, ciliolate marginally and on the keel, usually narrowing toward the apex. Flowers sessile (in bud), the hypanthium turbinate, ca. 2 mm long, densely ferrugineous villosulose, the ca- lycine cup short, the calycine lobes 4, narrowly triangular, ca. 3 mm long, ca. 1 mm wide, obtuse, thickly carnose especially dorsally, ferrugineous ciliolate at the base outside and marginally, glabrous within; corolla lobes 4, oblong, 74 mm long, carnose; stamens sessile, the anthers 4, rectangular oblong, to 4 mm long, to 1.2 mm wide; style slender, to 5.5 mm long, ca. 0.4 mm wide, obtuse at the apex, the stigmas not distinct, the ovarian disc compressed, to 1.8 mm wide, the ovules numerous, subrotund, ca. 0.05 mm wide, compressed, reticulate. Hoffmannia manussatani is known only from Panama. A combination of ses- sile leaves and a peduncle of extraordinary length bearing 3 pairs of bracts along its length readily distinguish it from all other species of Hoffmannia. BOCAS DEL TORO: Róbalo Trail, slopes of Cerro Horqueta, 6000-7000 ft, Allen 4978 (MO). 18. Hoffmannia morii Dwyer. rype: Panamá, Mori & Kallunki 5915 (MO, ho- lotype). Subshrubs to 1 m tall, the stem simple, obtusely angular, rimose, glabrous or glabrescent, covered with mosses, the nodes to 4 cm apart. Leaves widely oblong, 13-25 cm long, 7-12 cm wide, deltoid at the apex, obtuse or widely cuneate toward the base, the costa subplane above, prominent beneath, the lateral veins ca. 10, arcuate, not joining to form a conspicuous submarginal vein, thin coria- ceous when dry, slightly discolorous, glabrous above, puberulent beneath on the veins and costa, often moss covered; petioles to 3 cm long, to 1 mm wide, not vesciculiferous or with poorly developed vescicles. Inflorescences axillary and terminal, the flowers numerous in globose heads, 2 cm in diam. Flowers with pedicels ca. 2.5 cm long; hypanthium subrotund or turbinate, ca. 2 mm long, the hairs elongate, to 3 mm long, the calycine cup ca. 2 mm long, the lobes 4, narrowly oblong, ca. 4.5 mm long, thickly petaloid or subcoriaceous with purple hairs to 2 mm long outside; corolla tube narrowly cylindrical, glabrous, 5 mm long, 2.5 mm wide, thickly petaloid, the lobes 4, oblong, triangular, ca. 3 mm long, cu- cullate; stamens 4, the anthers oblong ca. 3 mm long, the filaments ca. 1 mm . eee morii Dwyer, spec. nov. Suffrutices ad | m alti. Folia lato- -oblonga, 13-25 cm longa, 7-12 cm lata, apice deltoidea, venis lateralibus circa 10 arcuatis . tenui-coriacea supra glabra subtus in venis ES costa puberula saepe. /nflorescentiae axillares - terminales floribus dense confertis in capitula globosa circa 2 cm diam. Flores pedicellis 2.5 mm longis; cupula a calycis 2 mm longa, lobis 4 angusto-oblongis 4.5 mm longis, extus ciliis purpureis elongatis a 2 mm longis ornatis, intus glabris; corolla tubo 5 mm longo, lobis 4 oblongo-triangularibus, 3 mm longis; staminibus 4 antheris oblongis 3 mm longis filamentis | mm longis; stylo 6.5 mm longo stigmatibus ovatis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 237 long, attached above the middle of the tube: style slender, ca. 6.5 mm long, slightly expanded toward the apex, the stigmas 2, ovate, ca. 2-8 mm long, ca. 1.5 mm wide, conspicuously thickened. Fruits not seen. Hoffmannia morii is known only from Panama. It is named in honor of Dr. Scott Mori whose collections of Panamanian Rubiaceae have been of great as- sistance. The new species is closely related to H. vesciculifera Standley. The latter has conspicuous inflations or vescicles on the petioles which are not present or at least not well developed in the new species, although some smaller leaves of the new species have conspicuously dilated petioles. The leaf blades average 10 lat- eral veins while H. vesciculifera has 12-25 lateral veins. The hypanthium of the new species is hairy in contrast to H. vesciculifera and the calycine lobes are much shorter than in Standley’s species. This species is known only from the type collection. CHIRIQUÍ: N of San Félix at Chiriqui-Bocas del Toro border, road to Cerro Colorado mine, 5000- 5500 ft, Mori & Kallunki 5915 (MO). 19. Hoffmannia ostaurea Dwyer, Ann. Missouri Bot. Gard. 56: 280. 1969. TYPE: Panamá, Kirkbride & Duke 501 (MO, holotype). Succulent subshrubs, the branchlets canaliculate when dry, glabrescent, with crustose patches of hairs (under magnification), the juvenile branchlets with many raphides, the internodes 1.5-3 cm long. Leaves elliptic, to 25 cm long, to 7.5 cm wide, deltoid and short acuminate apically, the acumen to | cm long, the costa plane on both sides, the lateral veins 12-15, spreading and arcuate, slender, the intermediate veins irregular pinnatiform with intervenal areas conspicuously spreading, thinly membranaceous, concolorous, glabrous but with vermiform ra- phides to 0.3 mm long; petioles arcuate ascending, to 8 cm long, 0.2-0.15 cm wide, glabrous, the raphides to 0.3 mm long; stipules not seen. /nflorescences to 1.5 cm long, the peduncle capiliaceous, the flowers few. Flowers (buds) with the hypanthium oblong rotund, ca. 2 mm long, smooth, the calycine lobes 4, unequal, subulate, to 1.2 mm long, the raphides thick, white; corolla subcarnose, the lobes presumably 4: filaments short; stigmas 2, capitate, ca. | mm long. Fruits on pedicels to 5 mm long, rotund, to 5 mm in diam., red and red purple when dry. Hoffmannia ostaurea is known only from Panama. BOCAS DEL TORO: Rio Teribe between Quebrada Huron and Quebrada Schlunjik, Kirkbride & Duke 501 (MO). 20. Hoffmannia pittieri Standley, Contr. U.S. Natl. Herb. 18: 140. 1916. TYPE: Panama, Pittier 3247 (US). H. panamensis Standley, Contr. U.S. Natl. Herb. 20: 204. 1919. TYPE: Panama, Pittier 3056 (US). Shrubs to 8 m tall, the stouter branches smooth, drying tan, terete, glabrous, the pith porate. Leaves elliptic or obovate elliptic, 9-35 cm long, 8-12 cm wide, 2.5-3 times longer than wide, acute to cuneate, rarely subrotund at the apex, short acuminate, the acumen to 2 cm long, basally acute to attenuate acute, the 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 costa prominulous above, prominulous to prominent beneath, often ligneous, to 2.5 mm wide, the lateral veins (11—)15—20, widely arcuate, 1(—2.2) cm apart, with 1-2 conspicuous irregular veins between each pair of lateral veins, chartaceous, usually slightly discolorous, usually drying tan or brown, glabrous; petioles 5.5— 7 cm long, 1.5-3 mm wide; stipules not seen. /nflorescences in the uppermost axils, spreading, 3-9 cm long, the primary peduncle compressed as a nodal pad, or elongate, the fasciculate branches 1—4, the flowers usually spreading. Flowers with the pedicels to 15 mm long, stiff, glabrous; hypanthium carnose, ribbed, puberulent to glabrous, the calycine teeth 4, triangular, 0.6-2 mm long; corolla yellow to greenish white, to 13.5 mm long, subcarnose, glabrous to sparsely puberulent, the lobes 4(—5), 2.3-4.3 mm long, 1.7-3 mm wide; stamens 4, the anthers 2.3-4.5 mm long, the connective not produced at apex or base, the fila- ments to 1.8 mm long; ovary 2-4-loculate, the style to 11 mm long, the stigmas usually ball shaped, 1.2-1.8 mm long, wider than the style, each stigma ovate oblong to cordate. Fruits subrotund; calycine lobes persistent, to 6.5 mm long, glabrous, drying red brown, scarcely ribbed; seeds brown, foveolate. Hoffmannia pittieri is known only from Panama. Folsom 2211 and 2212 have exceptionally large fruits, to 1.4 cm long, and drying tan. Apparently the fleshy portion surrounding the seeds did not shrink as usual on drying. The fruits of the 2 collections are described as "white and pink.” CHIRIQUÍ: Róbalo Trail, N slope of Cerro Horqueta, 6000-7000 ft, Allen 4962 ee Cerro Hor- queta, 7000 ft, Blum & Dwyer 2599 (MO). Cerro Hong hrs Blum & Dwyer 2643 (MO). Cerro Hor- queta, 7000 ft, Blum & Dw yd 2647 (MO). Las Nubes, 5.5 km NW of Río Chiriquí Ta W of Cerro Punta, 2200 m, Busey 679 A (MO). Cerro Horqueta, Cor hrane et al. 6244 (MO). Monte Rey above Boquete, Croat & Porter 15655 (MO). Between Boquete & Monte Rey, Croat 15823 (MO). Alto Reone a and above, 2800 m, D'Arcy 9967 (MO). Above Boquete on La Popa, 5400 ft, D'Arcy & a 6371 (MO). Above Boquete, road to La Popa, D'Arcy & D'Arcy 6372 (MO). Bajo Chorro, Boquete, 6000 ft, Davidson 47 (MO). Bajo Chorro, bas e e, . h 51 (MO). Cerro Horqueta, Duke et al. 13651 (M r Dwyer 8765, 8790 (both MO). Boquete, Finca Collins, ca. 5000 ft, Dwyer & Hayden 7654 (GH, K, MO, UC, US). NW of B pd Hoa. y 5800 ft, Dwyer et al. 525 (MO). B Finca Collins, Dwyer & Hayden 7665 ,U Boquete, Cerro Horqueta, Dwyer & Hayden 7745 (GH, MO). Cerro TRAE Dwyer & Lallathin 8783 (MO). Boquete, Folsom 2211, 2212 (both MO). Boquete, Finca Collins, Hayden 163 (MO). Boquete, Finca Collins, Kirkbride 117 (MO). Cerro Horqueta, 5000-7000 ft, Kirkbride 147 (MO). Between Pinola & Quebrado Hondo, toward summit on Chiriqui Trail, Kirkbride & Duke 894 (MO). Cerro Punta, 6800 ft, Ridgway & Solis 2393 (MO). Upper Río Chiriqui Viejo, near Monte Lirio, 1300— 1900 m, Seibert 165 (MO). Palo Alto, E of dn Stern et al. 1025 (MO). Boquete, Finca Collins, 5500 ft, Stern et al. 1127 (MO). 6 mi above Cer jupe Boquete Trail, ca. 750 ft, Tvson 7168 (MO). Above Jamarillo Arriba, N slopes of Cerro Palo Alto, 4900 ft, Webster 11674 (MO). Valley of upper Rio Chiriquí Viejo, White & White 2, 26 (both MO). Río Chiriquí Viejo Valley, uis 74 (MO). Bajo Mona & Quebrada Chiquero, 1500 m, Woodson & Schery 538 (MO). Bajo Mona & Quebrada Chi- quero, 1500 m. Woodson & Schery 561 (MO). Casita Alta, Volcán de Chiriquí, Gss erai. 839 (MO). COCLE: Cerro Pilón, 2900 ft, Lallathin 434 (MO). 21. Hoffmannia psychotriifolia (Benth.) Griseb., Fl. Brit. W. Ind. 321. 1861.— Fic. 51 Higginsia ee Benth. in Oerst. Vidensk. Meddel. oe Naturhist. Foren. Kjøbenhavn 1852: : Costa Rica, Oersted 11507, not see Shrubs to 2.5 m tall, the stems ultimately slender, terete, glabrous, rubescent. Leaves oblong, to 20 cm long, to 8 cm wide, usually 3-4 times longer than wide, acute and finally attenuate acuminate at the apex, the acumen to 2.5 cm long, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) WZ TS, - We Ws ` QNS, IGURE 51. Hoffmannia psychotriifolia (Benth.) Griseb.—A. Habit (x .45).—B. Flower, opened to show interior (x 4.95).—C. Ovary cross section (x4.95). [After Croat 15976. | acute and oblique at the base, the lateral veins 10-12, widely arcuate, stiffly chartaceous. concolorous or discolorous, glabrescent, usually with minute, scat- tered white raphides on both the surfaces; petioles to 9 cm long, and 2 mm wide, stipules not seen. Inflorescences in the axils of the uppermost leaves, the pedun- 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 cles reduced to a pad or to 8 mm long, the flowers radially disposed, ca. 7 per inflorescence. Flowers on slender pedicels to 4 mm long, white, to 12 mm long; hypanthium 2-4 mm long, scarcely ribbed, often with abundant red raphides, the calycine teeth 4, deltoid, 0.8—2 mm long, acute; corolla 8-11 mm long, the lobes 4, thickly petaloid, often longer than the tube; stamens 4, the anthers 2—2.8(—4) mm long, the connective not produced at the apex or the base, the filaments 0.5— l mm long; ovary 2-3-loculate, the style to 5.5 mm long, the stigmas 2, linear, spreading at maturity, 2-3.3 mm long. Fruits on radiate, slender pedicels, shorter than the fruit; fruit oblong to ovate oblong, obtuse or truncate at apex, to 8 mm long, at least 2 times longer than wide, at first delicately ribbed, smooth, glabrous, often drying yellow, finally fleshy, in vivo china red, pink or purple. The species ranges from Guatemala to Panama, and it also occurs in the West Indies. HIRIQUI: Central valley of Río Chiriquí Viejo, Allen 1329 (MO). Cerro Horqueta, 7000 ft, Blum & Dwyer 2626 (MO). Monte Rey above Boquete, Croat & Porter 15702 (MO). NW side of Cerro Pando, Croat 15976 (MO). Between Concepción and El Hato de Volcán, D'Arcy 4254A (MO). Bajo Chorro, Davidson 198 (MO). NW of Boquete, Cerro Horqueta, 5000-5800 ft, Dw M et al. 518 (GH, f ; US). Boquete, Finca Collins. dd & Hayden 7653, 7666, 7681 (GH, MO, US). Boquete, Finca Collins, Dwyer & Hayden 7664 (GH, MO, UC, US). Boquete, Ebinger 671 (MO). Finca Collins, Boquete, Hayden 162 (MO). Finca ae Boquete, 6000 ft, Kirkbride Ae O). 27 km NW of El Hato del Volcan, 500-5300 ft, n "s Bolten di (MO). Palo Alto of Boquete, 5000 ft, Stern et al. 1015 (MO). Quebrada Velo, 0 m, Woodson & 3e ine 258 TS ). Bajo Mona & Quebrada Chiquero, Woodson & Schery 564 (MON Río Chiriquí V Valley, White 53 (MO). Casita Alta, Volcán de Chiriqui, 1500-2000 m, Woodson et al. 984 4 (MO). COCLÉ: Between Cerro Pilón and EI Valle, Duke & Dwver 13972 (MO). 22. Hoffmannia pustulata Dwyer. ryPE: Panamá, Dressler May-June 1972 (MO- 217890, holotype). Subshrubs to 0.3 m tall, the stems subprostrate and rooting at the nodes, the bark somewhat rough, the lenticels minute. Leaves deciduous, but crowded to- ward the apex of the twiglets, narrowly obovate oblong, 4.5-8 cm long, 1.6—-4.5 cm wide, deltoid to obtuse toward the apex, acute at the base, the costa prominent above and beneath, the lateral veins 22-25, plane above, prominulous beneath, the intervenal areas conspicuously bullate above, stiffly chartaceous, concolor- ous, ferrugineous villosulose beneath; petioles 0.4-1 cm long, the hairs many, ferrugineous; stipules oblong, ca. 3 mm long, leaf-like. /nflorescences axillary, the peduncles 0.5-5 cm long, slender, ca. 1.2 mm wide, the hairs few, the cymules on reduced branches, the flowers few. Flowers scarcely pedicellate; calycine cup much reduced, the lobes 4, thickly subulate, to 1.5 mm long, the hairs few, to 0.8 55 Hoffmannia de AE Dwyer, spec. nov. Suffrutices ad 0.3 m alti, verve in parte inferiori subprostratis et in nodis radicibus. Folia pr obovato-oblonga, 4.5-8 cm long t versus apicem eio s. BRA lateralibus 22-25 interveniis supra conspicue bullatis rigido- charta acea s 0.4-1 cm Inflorescentiae pedunculis 0.5-5 cm longis, gracilibus, cymulis in brevissimis ramis dispositis floribus paucis. Flores breviter pedicellati, cupula calycis brevissima dentibus 4 crasso-subulatis, 1.5 mm longis, ciliis paucis, ad 0.8 mm longis, glandulis 2-3 minutis inter dentes dispositis; corolla tubo brevissimo, 1 mm longo, glabro, lobis 4 oblongis, ad 2 mm longis; staminibus 4, antheris oblongis, ca. 2mm longis, filamentis brevibus, ca. | mm longis; stigmatibus, ca. 1.2 mm longis, stylo ad 2 mm longo. Fructus immaturi oblongi-rotundi, ca. 5 mm in diam., glabri, raphidibus multis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 241 mm long, weak, with 2-3 minute glands between adjacent lobes, these 0.1 mm long, red; corolla pinkish green, the tube much reduced, to 1 mm long, glabrous, the lobes 4, petaloid, oblong, to 2 mm long, obtuse, slightly cucullate and carinate, the raphides abundant; stamens 4, the anthers oblong, ca. 2 mm long, obtuse, the filaments short, ca. 1 mm long; stigmas 2, thickly subulate, ca. 1.2 mm long, the style to 2 mm long, thick, the ovules numerous, the ovarian disc pulvinate, to 0.4 mm long, 1 mm wide. Fruits (immature), oblong rotund, ca. 5 mm in diam., glabrous, the raphides many. The new species is known only from Panama. Dressler has supplied important notes concerning the plant: "Shady bank in forest; leaves pink beneath, dark red green above; sepals red, petals pinkish green, anther and stigma cream.” The dominance of red and pink in several plant parts is striking. Dressler pressed the plant after cultivating it. Hoffmannia pustulata is closely related to the recently described H. bullata illiams which also has bullate leaves. The wide-ranging H. bullata (Mex- ico to Costa Rica) has the leaves petiolate and the lateral veins 8—15 pairs, while H. pustulata has leaves with petioles to 1 cm long and with 22-25 pairs of lateral veins. The corolla of H. bullata is "dull brick red measuring about 10 mm long," while H. pustulata has a pinkish green corolla about 3 mm long. COLÓN: Río Guanche, Dressler May-Jun. 1972 (MO). 23. Hoffmannia rexmontis Dwyer.** TYPE: Panamá, Croat 15934 (MO, holotype). Subshrubs 1—1.5 m tall, the stems succulent, olive colored when dry, con- spicuously rimose, minutely puberulent, the nodes well spaced; stipular scars conspicuously annular. Leaves widely oblong lanceolate, 13-18 cm long, 6-8 cm wide, deltoid and short acuminate at the apex, the acumen to 1.5 cm long, 0.15- 0.3 cm wide, the costa plane or ridged above, prominulous beneath, deltoid to obtuse at the base, the lateral veins ca. 14, widely arcuate, the submarginal vein undulate, 0.5 mm from the margin, the intervenal areas spreading reticulate, membranous, concolorous, minutely white farinose on both sides; petioles 6-7 cm long, slender, to 0.15 cm wide; stipules not seen. Inflorescences axillary, the peduncle to 1 cm long, 0.1 cm wide, the flowers few, the bracts subulate, to 1.5 mm long. Flowers not seen. Fruits pedicellate, the pedicels 0.3-10 mm long, the scars of the pedicels rotund, to 2 mm in diam.; fruits elliptic oblong or obovate oblong, to 0.9 cm long, to 0.38 cm wide, often truncate at the apex, attenuate toward the base, more than twice as long as wide. yellow green when dry, gla- brous, the ribs 8, slender; persistent calycine cup to 1 mm long, the lobes trian- gular or subulate, to 1.5 mm long, glabrous. es d re um Dwyer, spec. nov. Suffrutices 1-1.5 m alti, caulibus succulentis. Folia lato- E lan -18 cm longa, 6-8 cm lata apice deltoidea, venis lateralibus ca. 14 lato- membranacea Prem. minute-albido-farinosa:; petiolis ad 6 cm longa. /nflorescentiae pe- ce ad | cm ies. floribus paucis; bracteis subulatis ad 1.5 mm longis. Fructus pedicellis 0.3- 2 mm longis elliptico-oblongi obovato-oblongi, ad 0.9 cm longi, glabri costis gracilibus 8; calycis dentibus, ad 1.5 mm longis. TE 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Hoffmannia rex-montis is known only from Panama. It is distinguished by the fruit drying a dull yellow green and measuring about 24 times longer than wide. The fruits in the living state are red, pink or white (fide Croat). The only other species of Hoffmannia in Panama with similar fruit measurements is H. psycho- triifolia (Benth.) Griseb. The pedicels leave a buttonlike scar which is a distin- guishing character. The leaves tend to be concolorous on drying, but a few of the dried leaves suggest that they are darker above in the living state. CHIRIQUÍ: NW side of Cerro Pando, cloud forest, Croat 15920, 15934 (both MO). 24. Hoffmannia rubripigmenta Dwyer, Ann. Missouri Bot. Gard. 56: 283. 1960. TYPE: Panamá, Duke 10870 (MO, holotype). Herbs, woody, low growing, the branchlets glabrous, the raphides minute, oblong. Leaves obovate elliptic, to 23 cm long, to 9.5 cm wide, widely deltoid at the apex, ultimately obtuse, narrowly cuneate at the base, scarcely decurrent on petiole, the costa prominulous above and beneath, the lateral veins ca. 8, arcuate, the intermediate veins more divergent, ramose and evanescent; petioles to 1 cm long, delicately alate; stipules not seen. /nflorescences a little longer than the petioles, peduncle slender erect, glabrous, to 2 cm long, ca. 0.5 mm wide, red when dry, the flowers ca. 8. Flowers (in bud) subsessile, narrowly oblong, ca. 1 mm wide; hypanthium oblong, to 2.4 mm long, carnose, the wall thick, to 3.5 mm wide, glabrous, smooth, the teeth erect, triangular, to 1.5 mm long; corolla with the tube presumably short, the lobes 4, glabrous; stamens 4, the anthers ca. 4 mm long, the connective not produced at the apex or base; style cylindrical, the stigmas erect, oblong, ca. 1.3 mm long, scarcely thicker than the style. Fruits not seen. Hoffmannia rubripigmenta 1s known only from Panama. DARIÉN: Between upper Río Membrillo and Camp 7 on construction road to San Blas, 100—800 m, Duke 10870 (MO). 25. Hoffmannia subauriculata Standley, J. Wash. Acad. Sci. 18: 179. 1928. TYPE: Costa Rica, Standley & Rojas 50596 (US, holotype). Herbs, short, woody, the stems strongly quadrangular, smooth, glabrous, the roots fibrous. Leaves sessile, elliptic to obovate elliptic or obovate spatulate, 9— 30 cm long, 5-10 cm wide, acute and short acuminate at the apex, basally cuneate, often widely so, to 2.5 cm wide at the base, ultimately obtuse or vaguely auricu- late, the costa porcate above, prominent beneath, the lateral veins 10-25, widely arcuate, prominent beneath, usually 1.5-2.5 cm apart, with usually a conspicuous intermediate vein between a pair of lateral veins, chartaceous, glabrous, the raphides numerous, linear; stipules not seen. Inflorescences with peduncles wiry but not stiffly erect, arising at or near the ground level, to 4 cm long, the cymules few flowered. Flowers (in bud) with the hypanthium glabrous, the calycine teeth 4, subulate, less than 1 mm long, puberulent; stamens and petals not seen; ovary 2-locular. Fruits oblong, to 6 mm long, the ribs 4; seeds ca. 1 mm long, coarsely foveolate. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 243 Hoffmannia subauriculata is known from Costa Rica and Panama. BOCAS DEL TORO: Without other locality, Wedel 36] (MO). DARIEN: Cerro Pirre, Bristan 627 ) 26. Hoffmannia vesciculifera Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 285. 1929. TYPE: Panama, Cooper 231 (F).—Fic. 52. Shrubs to 1.5 m tall, probably unbranched, the hairs densely ferrugineous and arachnoid flocculose. Leaves oblong rotund to obovate oblong, to 25 cm long, to 15 cm wide, briefly acuminate at the apex, cuneate to rounded at the base, the costa immersed above, prominulous to plano compressed beneath, the lateral veins 12-25, widely arcuate to substrict, chartaceous, discolorous, drying brown, usually foveolate above, the hairs diffuse, to 5 mm long, often calcified at the base, confined to veins and venules; petioles to 4.5 cm long, conspicuously in- flated for most of their length, the inflations to 1 cm wide, located nearer the blade than the node, densely ferrugineous flocculose; stipules not seen. /nflores- cences to 5 cm long, the peduncles to 2 cm long or absent, the flowers numerous in densely bracteate axillary heads, these rounded or compressed rotund, the bracts linear lanceolate, to 1.5 cm long, 0.3-1.5 mm wide, with 1—2 irregular ascending submarginal veins, the margins glandular pectinate ciliate, the hairs to 4 mm long. Flowers yellow; pedicels to 4 mm long, glabrous; hypanthium oblong, truncate, glabrous, delicately 4-ribbed, the calvcine lobes 4-5, unequal in length nd width, lanceolate, 7.5-10 mm long, tapering above the middle, 1.5-2 mm wide, the hairs ferrugineous, marginal, to 2 mm long, usually densely aggregated above the middle, with a minute gland at junction of adjacent calycine lobes; corolla with the tube cylindrical, to 20 mm long, glabrous, the lobes 4, shorter than the tube, hirsute outside; stamens 4-5, the anthers oblong, ca. 4.5 mm long, the filament ca. 0.3 mm long, the connective not produced at the apex or base, the style slender, the stigmas clavate, spreading at maturity, wider than and about 14 the length of the style. Fruits oblong rotund, to 5 mm long, ca. 4.5 mm wide, delicately ribbed, the ribs ca. 10, the pericarp thin walled, the raphides oe the hairs deciduous; the persistent calycine lobes 4, linear lanceolate, to long, the hairs deciduous in patches; seeds rectangular to cuneate, piu re- ticulate. Hoffmannia vesciculifera Standley is known from Costa Rica and Panama. AS DEI ias pone e above Quebrada Huron, Kirkbride & Duke 611 (MO). Chiriquí- aa. Trail, n Quebrada Higueron and Gutierrez, Kirkbride & Duke 761 (MO). Water Valley, Chiriquí ype We del p (MO). Fish Creek Mts., Chiriquí Lagoon, Wedel 2284 (MO). CHIRIQUÍ: Cerro Colorado, - a N of San Félix, 1200-1500 m. Mori & Dressler 7784 (MO). Buena Vista gen Chiriqui Trail, 750 m, G. P. Cooper 231 (F, holotype). Cerro Colorado, upper mining mi from San E Croat 33386 (MO). Cerro Colorado, 34.1 km beyond bridge over Río San Fé dix. pe 37307 (MO). cocr É: El Valle, Dressler 4071, 4386 (both MO). Cloud forest of Cerro Caracoral, Duke & Dwyer 15150 (NY). La Mesa, Dwyer & Nee 11952 (MO). Cerro Colorado, Bocas Road, 1500 m, ud & Collins 1821 (MO). Cerro Caracoral Gaital, Kirkbride 1117 (MO). COLÓN: Río Iguanita, 300 D'Arcy 11256 (MO). vERAGUAS: NW of Santa Fé, 11 km from Escuela iran Alto de Piedra. Valley Río Dos Bocas, 450-550 m, Mori et al. 3810 (MO). 16 km from Santa ipa slope near Río Calovéborita, Mori et al. 6693 (MO). NW of Santa Fe, 11 km from a Agricola, Alto de Piedra, Mori et al. 3847 (MO). Guabal, 16 km NW of Santa Fé, 500 m, i» ord 4796 (MO). 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 27. Hoffmannia witheringioides Dwyer.” type: Panamá, Croat 25551 (MO. ho- lotype). Subshrubs to 1.5 m tall, the branchlets angular, black brown when dry, pu- bescent, the hairs dense, curled, the nodes well spaced. Leaves ovate, oblong, rotund or oblong rotund, at first seemingly anisophyllous but actually pseudo anisophyllous, the smaller leaves on short axillary branchlets, the blades then to 4 cm long, to 2.3 cm wide, acuminate, the larger leaves to 13 cm long, to 8 cm wide, rounded or obtuse toward the apex, attenuate cuneate toward the base, the costa slender and prominulous above, subprominent beneath, the lateral veins ca. 10, the venules irregular pinnatiform, arcuate, chartaceous, discolorous, dark brown above, liver colored beneath, the hairs on both sides short, vermiform subulate, turgid toward the base, rough to the touch; petioles 3.5-4 cm long, to 1.5 mm wide, with dense, curled hairs; stipules not seen. Flowers 1-2 per axil, the peduncle absent, the pedicels to 0.7 cm long, slender with curled hairs; hy- panthium turbinate, to 4 mm long, truncate, the hairs curled, lightly costate, the calycine cup 0.5 mm long, the lobes 4, deltoid, to 1 mm long; corolla dark yellow green, the tube cylindrical, to 4 mm long, scarcely expanded near the mouth, petaloid, villose outside, glabrous within, the lobes 4, ovate oblong, to 4 mm long, obtuse; stamens 4, the anthers subsessile, oblong, to 2.2 mm long, the connective to 0.5 mm long at the apex, rounded; style to 5 mm long, the stigmatic lobes oblong, to 1.2 mm long, to 0.5 mm wide, obtuse. Fruits not seen. Hoffmannia witheringioides is known only from the solitary collection in Pan- ama. It bears a striking resemblance to the solanaceous genus Witheringia. The pseudo-anisophylly of the species is striking. The reduction of the flowers to one or two, coupled with the absence of a peduncle seems unique for the Hoffmannia of Middle America. AGUAS: Río Segundo Braso, 8 km beyond Escuela Agricola Alto Piedra beyond Santa Fé, 750 m m, - radi 25551 (MO). 28. Hoffmannia woodsonii Standley, Ann. Missouri Bot. Gard. 28: 471. 1941. TYPE: Panamá, Woodson & Schery 503 (MO, holotype). Shrubs 2—4 m tall, the twigs glabrous, smooth, the raphides dot-like white. Leaves oblong, 5-12 cm long, 1-5(—6) cm wide, attenuate acuminate at the apex, the acumen to 1 cm long, basally attenuate cuneate or acute, the costa plano compressed beneath, the lateral veins 4-8, prominulous above, subprominent beneath, membranous to thinly chartaceous, glabrous or puberulent, the raphids scattered, white, subgranular above; petioles 2-6 cm long, to 0.8 mm wide; stip- cm bue foliis onus ad 13 cm ne ad 8 cm latis, venis s ja circa a 10. chartacea discoloria ciliis utrinque brevibus vermiformi-subulatis; petiolis 3.5-4 cm longis. Flores 1—2 per axillam, pe- a alle. pedicellis ad 0.7 cm longis ciliis crispis ornatis, cupula E circa 0.5 mm longa, idi 4 deltoide. ad 1 mm longis; corolla tubo ad 4 mm longo, extus villoso intus glabro, lobis 4, ad m longis; antheris 4, subsessilibus oblongis, ad 2.? mm longis, apice connectivo ad 0.5 mm longo: ds ad 5 mm longo, lobis stigmatis oblongis 1.2 mm m longis. a 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 245 on nye a = (Z: us ASIN FiGuRE 52. Hoffmannia vesciculifera Standley.—A. Habit (x!2).—B. Flower (x 1).—C. Fruit (x2.5). [After Dressler 4386. | ules not seen. Inflorescences 1—2 per axil, cymes to 1.5 cm long, the peduncles reduced to a pad or to 6 mm long, ca. 0.5 mm wide, glabrous to ferrugineous puberulent, the flowers usually 2-6 per cymule. Flowers with the pedicels 5-7 mm long, often radially disposed or deflexed candelabraform capillaceous, gla- brous to ferrugineous puberulent; hypanthium square in cross section; subcar- nose, glabrous (puberulent?), the calycine teeth 4(—5), cuneate subulate, 0.5-1.3 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 mm long; corolla white, cream, yellow, to occasionally whitish pink or orange, 7-8 mm long, the lobes 4, elongate, delicately carnose, glabrous, often delicately ribbed; stamens 4, the anthers partly exserted, 3-4.7 mm long, the connective produced at the apex, the acumen ovate deltoid, ca. 1 mm long; ovary 2-locellate, the style 4-5.5 mm long, the stigmas 2, oblong capitate, 1-1.2 mm long, wider than the style. Fruits (not seen in fleshy mature stage) oblong subrotund, truncate, to 4.5 mm long, red; persistent calycine lobes glabrous, delicately ribbed, the ribs well spaced, varying in prominence. Hoffmannia woodsonii is known only from Panama. Most collections are from above 600 m but a few collections have been made near sea level. BOCAS DEL TORO: Between Guttierrez Quebrada and E slope of La Zorra, Río Mali, Chiriqui Trail, Kirkbride & Duke 715 (MO). CANAL ZONE: Farfan Beach, Dwyer & Lallathin 8791 (MO). Barro Colorado Island, Croat 10125, 10213, 14567 (all MO). CHIRIQUI: Methodist camp. near Nueva Swissa, Croat 13511 (MO). Between Boquete & Monte Rey, Croat 15634 (MO). 10-11 mi W of Puerto rmuelles, Croat 22003 (MO). 6 mi beyond Escuela Agricola Alto Piedra, 730 m, Croat & Folsom 34024, 34025 (both MO). Road between Alto Piedra School and Río Calovébora, 1.8 mi beyond school, 500 m, Croat & Folsom 34112 (MO). Cerro Pads Duke et al. 13634 (MO). 12 mi W of Puerto Armuelles, qe 202 (MO). Dos Lagunas, 4 km W of El Hato del Volcán, 1300 m, hay & Bolten 7396 (MO). Quebrada Velo, Woodson & Schery 260 (MO). Callejon Seco, Volca Chiriquí, Woodson 7 Schery 503 (MO). cocLe: Cerro Pilón, Dwyer & Lallathin ee 8681 (all MO). Cerro Gaital Caracoral, Dwyer & Correa 8910 (MO). Club Campestre to La Mesa, Dwyer 10526 (MO). EI Valle, Lewis et al. 1732 (GH, MO, US). Cerro Pilón, 5 km NE of EI Valle, Mori 6560 (MO). Río Guanche, ca. 3-5 km aboye Portobelo road bridge, Croat 36998 (MO): COLON: Mori & Kallunki 5189 (MO). Río Guanche, 2.5 km upriver from bridge, road to Portobelo, Mori et al. 6447 (MO). Los SANTOS: Loma Prieta, Dae 11849 (MO). PANAMA: Cerro Campana, Croat 14723, 14783 (both MO). El Llano-Carti Road, 9.6 km from Panamerican Highway, Mori & Kallunki 1855 (MO). Cerro Cam- pana, 870-1000 m, Mori & Kallunki 3564 (MO). 5-10 km NE of Altos de Pacora, 750 m, Mori & Kallunki 4987 (MO). d and of Panamerican Highway, El Llano-Cartí Road, 400-450 m, Nee & Dressler 9341 (MO). AGUAS: Between Santa Fé and Río Calovébora, 1.8 mi beyond Escuela Agricola, 735 m, bent d Folsom 34268 (MO). 49. ISERTIA Isertia Schreb., Gen. Pl. 1: 342. n. 602. 1789. TYPE: J. coccinea (Aubl.) Vahl. Creatantha Standley, Field Mus. Nat. Hist., Bot. Ser. 8: 344. 1931. rype: C. peruviana Standley. Trees or shrubs. Leaves petiolate, the stipules free, interpetiolar, erect, per- sistent. /nflorescences large, terminal, cymose corymbose or cymose paniculate. Flowers large, pedicellate, the hypanthium usually subovoid, the calycine cup short, the teeth usually 4—6; corolla elongate, tubular funnelform or salverform, rigidly petaloid, hairy within, the lobes 4—6, valvate; stamens exserted, the an- thers linear; ovary 2-6-celled. Fruits baccate, globose, the seeds small, numer- ous, foveolate. Isertia occurs throughout Middle America and tropical South America. There are about 25 species in the genus. a. Leaf-blades usually dull grey green beneath; lateral veins 15-22; corolla tube to 20 m long: fruits to 0.7 cm in diam. Dp S ean Leaf-blades bright gray green, grey o or r tan beneath when dry, usually described as whitish beneath; lateral veins ca. 22: corolla tube to 50 mm long; fruits 1.5 cm in dia m > 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 247 1. Isertia haenkeana DC., Prodr. 4: 437. 1830. Type: México, Haenke, not seen. I. spicaeformis Hemsl., Biol. Centr. Amer. Bot. 2: 31. 1881, not DC. 1830. Trees to 6m tall or shrubs, the branchlets angular, usually densely villosulose or villose, the nodes well spaced, often constricted. Leaves oblong, widely lan- ceolate, occasionally obovate oblong, to 46 cm long, to 28 cm wide, cuspidate, the cusp to 2.5 cm long, deltoid to acute at the apex, subobtuse to cuneate at the base, often attenuate, occasionally slightly inequilateral, the costa slender, sub- immersed or slightly prominulous above, prominent beneath, especially proxi- mally, the secondary veins 15-22, rigid, ascending, the intervenal areas with perpendicular veinlets, membranous, papyraceous, rarely subcoriaceous, usually dull grey green beneath when dry, glabrous above, appressed villosulose beneath; petioles 0.5-6 cm long, the lamina decurrent adaxially, pubescent; stipules tra- peziform or ovate elliptic, to 1.6 cm long, acute, carinate, villose. Inflorescences terminal, solitary, puberulent, thrysiform, pyramidal, oblong or subrotund, rarely wider than long, shorter than the terminal leaves, to 21 cm long, to 10 cm wide, the peduncles absent or to 5 cm long, often stout, to 0.6 cm wide, angular, the basal branches usually opposite, the lower branches of epedunculate inflores- cences more remote from the upper branches, these appearing whorled, densely flowered. Flowers with the hypanthium and calycine cup subrotund, ca. 4 mm diam., to 2 mm long, sparsely pubescent outside, but with patches of glands at or near the apex of the cup within, the teeth 4-5, scarcely evident; corolla yellow, the tube narrowly cylindrical, ca. 20 mm long, glabrous outside, pubescent above the middle within, the lobes ca. 6 mm long, slightly cucullate, about % the length of the tube or less, golden villose within; stamens 5, the anthers oblong, ca. 3.5 mm long, acute at the apex, the filaments crassate, about as long as the anther, attached near the corolla mouth; ovarian disc prominent, padlike, the style linear, ca. 25 mm long, the stigmas 5, linear oblong, ca. 5 mm long. Fruits depressed rotund, to 7 mm in diam., glabrous to puberulent; seeds brown, to 1 mm long, favose. Isertia haenkeana occurs in Central America, extending into Colombia and Venezuela. It also occurs in Cuba. *'Canelito," "Fruta de Mono," "Huevo de ono," "Murcielago," Pansus Tumatti’’ (San Blas). OCAS DEL TORO: Almirante, Blum 1388 (MO); McDaniel 5049 (MO). Chiriquí Lagoon, Seibert 1546 (US). ded Bocas, Wedel 2488 (MO, US), 2539 (MO). Santa Catalina, Blackwell et al. 2703 NAL ZONE: Madden Dam Road, Bartlett & Lasser 16442 (MO). Summit Gardens, Croat 14980 (MO). "Pipeli ine Road within 5 mi of Gamboa Gate, D'Arcy 9275 (MO). Drowned Río Azote Caballo, Dodge et al. 16842 (MO). Fort ipu fi eid (MO). Between Puentos de Las Americas & Fort Kobbe, Duke & Mussell 6552 (MO). George Greene Memorial, Dwyer 6756 (MO). Miraflores Locks, Dwyer & Robyns 2 (MO). Albrook, U. s. p rmy Tropic Test Site, Dwyer & Robyns 52 (MO). Chagres, Fendler 43 (MO). Road to Corozal, Gervais 129 (F). Ancón, Gervais 149 (US). Mount Hope, Johansen 57 (MO). K-10 & K-15 Road, Luteyn 1584 (MO). Headwater Río Providencia, 7.5 km SW of Gatün Locks, Nee & Gentry 8678 (MO). Miraflores & Corozal, Pittier 2/99 (US). Gamboa, Pittier 3786 (US). Agua Clara, Pittier 5931 (US). Ancón Hill, Seibert 389 (F, MO). Road K-15, Smith et al. 3285 (F). Pipeline Road, Smith & Smith 3303 (F). Coroza ] Road, Standley 26832 (US). Summit, Standley 26940 (MO). Balboa, Standley 27011 (US). Rio Tapia, Standley 28111 (US). Fort Randolph, Standley 28605 (US). Balboa, Standley 29570 (US). Summit, Standley 29640 (US). France Field, Standley 30340 (US). France Field, Standley 30426 (US). Cerro Luisa, above Pedro Miguel Locks, Stern & Chambers 2 (MO). Fort Sherman, Stimson 5100 (MO, US). Albrook, poca 5062 (MO). ~ 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Miraflores Locks area, Tyson 1144 (MO). Gamboa Pipeline Road, Tyson 1488 (MO). Beacon Hill, Albrook Air Force Base, Tyson 2040 (MO, US). Farfan Beach, White 141 (F, MO). Summit Garden Witherspoon & Withe UE 8692 (MO). Ancón Hill, Woodson et al. 1333 (F, MO). BARRO COLORADO ISLAND: Aviles 29 (F, MO); Bangham 370 (US); Brown 73 (F); Croat 4055, 4063, 5076, 6107, 6331, 7050, 11611 (all MO): Dodge 3466 (US); Dwyer 8475 (MO); Hayden 23 (MO); Kenover 548 (US); Luteyn 781 (F); Shattuck 625 (F, MO), 765 (F); Standley 41056 (US); Starry 133 (F, MO); Woodson & Schery 964 (F, MO): Wondwanih & Vestal 352 (MO). coci É: Cerro Pilon, vp 2 (MO). El Valle, Wilbur 11140 (F). COLON: Camp Pina, Allen 3589 (F). Nuevo Limón, Allen 5141 (F). Guasimo on Río Miguel de la Borda, Croat 9997 (MO). Sabanita, Croat 11072 (MO). Santa ns Ridge, Croat 13175 we Ocean Trail between Rio Indio and Miguel de la Borda, Croat 36923 (MO). Rio Guanche near Coastal Highway, D'Arcy 10207 (MO). Road to Portobelo between Rio Piedras & Portobelo, Elias & Kirkbride 1638 (GH, MO, UC, US). Pina, Kirkbride & Hayden 316 (MO). 6 mi SW Portobelo, Luteyn 1420 (MO). Santa Rita Ridge, Lewis et al. 5254 (MO, MOCZ, US). 7 km NE of Puerto Pilon, ad to Portobelo, Nee 6680 (MO). 3-4 km NW of Salamanca, 13 km NE of Buenos Aires, Nee 9097 (MO). Eon Rita Hills, Smith & Smith 3441 (F). DARIÉN: Pinogana to Yaviza, Duke 5166 (MO). Río Tuira between Pucro & Río Ponusa, Duke 5388 (MO). S of El Real on Río Pirre, Duke 5418 (MO). Caná, Stern et al. 653 (MO). Tucutí, Terry & Terry 1404 (F, MO). Caná, Williams 754 (US). PANAMÁ: Pacora, Allen 1121 (F, US). Without exact locality, Andersson (F). Burunga, Austin 14 "ande Cam- pana, Castillo 27 say San José Island, Correa 89 (MO). El Llano-Carti Road, ca. 10 km N of Panamerican Highway, D'Arcy 10660 (MO). 5-6 mi E of Chepo, Duke 4050 (MO). edd El Llano and Río Mamoní, Duke 5606 (MO). San José Island, Duke 12544 (MO): Erlanson 264, 306 (both US). Arraijan, Guerra 8 (MO). Capira, Jaramillo 36 (MO). San José Island, Johnston 77 (US). Chimán, Lewis et al. 3241 (MO). Capira, Maurice 803 (MO). 6 km SW Arraijan, road to Bique, Nee 6903 MO). Capira, Paul 135 (US). Las Sabanas, Pittier 6705 (US). Juan Franco Race Track, Standley 27741 (US). Tumba Muerta Road, Srandley 29753 (US). San José Island, ipis (MO). Arraijan, Woodson et al. 1366 (F, MO). sAN BLAs: Permé, Cooper 653 (F, US). Ustupo, D'Arcy 9500 (MO). Mulatupu (Río Ibedi), Duke 8485 (MO). Isla de Pinas, Kirkbride 214 (MO). pu cud pu, Lewis et al. 118 (MO). Ailigandi, Lewis et al. 138 (GH, MO, UC, US). Niatuppu, Stier 201 (MO). VERAGUAS: Río Concepción, Lewis et al. 2788 (COL, DUKE, MO, UC, VEN). 2. Isertia hypoleuca Benth., Hook. L. Jour. Bot. 3: 220. 1841. TYPE: British uiana, Schomburghk 281 (K, not seen). I. coccinea var. hypoleuca (Benth.) Schum. in Mart., Fl. Fa 6(6): 286. 1889 Cassupa panamensis Standley, Contr. U.S. Natl. Herb. 18: 135. 1914. TYPE: Panamá, Pittier 3889 US). Isertia panamensis (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser. 8: 346. 1931. Trees to 20 m tall, the branches stout, grooved, densely puberulent. Leaves oblong, to 40 cm long, to 27 cm wide, deltoid at the apex, shortly cuspidate, the cusp slender, to 2.5 cm long, often falcate, obtuse, deltoid to cuneate at the base, the costa subplane to prominulous above, slender, prominent, especially proxi- mally beneath, the secondary veins ca. 22, strict, subcoriaceous, discolorous, usually chocolate brown above, grey green, drying grey or tan brown beneath, glabrate above except puberulent on the costa and the veins, densely puberulent beneath; petioles to 0.9 cm long, to 0.4 cm wide, stout, often curved, puberulent, delicately winged on the adaxial side; stipules oblong lanceolate, 1-2 cm long, acute, coriaceous, pubescent marginally. Inflorescences terminal, solitary, pu- berulent, shorter than or as long as the terminal leaves, to 35 cm long, to 15 cm wide, oblong, ovate or subrotund in outline, thyrsiform; peduncles stout, to 7(—8?) cm long, to 0.7 cm wide, lignose, angular, the branches alternate or subverticillate except the lowermost which are often disposed as 2 opposite pairs, stiff, well spaced; bracts and bracteoles minute, widely subulate. Flowers sessile or con- spicuously pedicellate; hypanthium oblong, truncate, 2.0-3.5 mm long, coria- ceous, glabrous, marcescent, the teeth scarcely evident or absent, with small patches of obtuse glands within the cup, corolla white, turning brown to red, the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 249 FiGURE 53. /sertia hypoleuca Benth.—A. Habit (xV2).—B. Flower opened to show interior (x34). x E cross section (x3!2). [After Stimson & Gallegos 5254. ]—D. Fruit (x'5). [After Croat 16756.] 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 tube narrow cylindrical, to 50 mm long, 2.0-3.5 mm wide at the middle, puber- ulent outside, densely villose within above the middle, coriaceous, the lobes 5, oblong, 9-15 mm long, cucullate, subcoriaceous, villose within below the middle; stamens 6, the anthers oblong, ca. 7.5 mm long, acute, minutely apiculate, dor- sifixed, the surface honeycomb-like, the filaments short, attached a few mm below the mouth; style linear, 25-27 mm long, ca. 0.7 mm wide, ciliolate at the apex; the stigmas 2, erect, oblong, ca. 5.5 mm long, 1.8-2.5 mm wide, the ovary oblong rotund in cross section, the wall thick, replete with reddish glands, the septum thick, 2-loculate, the placentas T-shaped, crassate, the ovules minute. Fruits rotund or oblong, to 1.5 cm in diam., smooth, glabrescent, drying black. The species is known from Panama, Colombia, Surinam, French Guiana, Brit- ish Guiana, Venezuela, and Brazil. BOCAS DEL TORO: Chiriquicito, Lewis et al. 2054 (GH, MO, US). Nances Cay Island, Wedel 580 (MO). Old Bank Island, Wedel 1937 (MO, US). CANAL ZONE: Pipeline Road, Folsom 2848 (MO). Rio Pina-Río Media Divide, Johnston 1603 (MO). CocLE: La Mesa, Gentry 6845 (MO). Above El Valle, Road to La Mesa, 2500 ft, Tyson 6930 (MO). CoLGN: Rio Piedras, road to Puerto Pilon, Correa 143 O). Guasimo, Croat 9929 (MO). 1 mi N of Maria Chiquita, Croat 11368 (MO). Santa Rita Ridge, Croat 13175 (MO). María Chiquita, Dwyer & Kirkbride 7801A (MO), 7812 (GH, MO, US). Santa Rita Ridge, Gentry 1856 (MO). Gatun, Lao 127 (MO). Between Pina and Gatun, Lutevn 1309 (MO). Santa Rita Ridge, Stimson & Gallegos 5254 (MO). Achiote, Tyson et al. 4551 (MO). DARIEN: Quebrada Venado and Peje Swamp, Bristan 1056 (MO). Cerro Pirre, Bristan 1228 (MO, US). Rio Paya, Duke & Kirkbride 14069 (MO). Puerto St. Dorothea, Dwyer 2201 (MO). PANAMA: Cerro Jefe, Duke 9393 MO). Cerro Jefe and La Eneida, Dwyer 82/3 (MO). Capira, Bro. Maurice 803 (US). Cerro Azul, Tyson 2196, 4051 (both MO). Cerro Jefe, Tyson 3330 (MO). SAN BLAS: Puerto Obaldia, Croat 16756 (MO). VERAGUAS: Cerro Tute, Gentry 6218 (MO). COLOMBIA: CHOCO: Rio Mutata, Rio El Valle, base of Alto De Buey, 100-150 m, Gentry & Fallen 17303 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 251 INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms: numbers with daggers (+) refer to names incidentally mentioned. Aeginetia 55, 55* palustris 43 capitata 108+ parviflora 43+ Alibertia 47, 19 i edulis 191, 19, 21* Billardieria garapatica 21 paniculata 132 bn 19 anamensis | Allenanthus 4+, 22 erythrocarpus dh 22 patinoi 447+, 467, 47 universitatis 467 Borrera 49 Borreria 27, 48 articularis 49* Amaioua 4*, 2 capitata cormbosa 27, 301 — "var. suaveolens 54 fagifolia 2 densiflora guianensis 27*, 27 laevis 5 magnicarpa 29 latifolia 52 eblinae 33* pringlei 53 spathulata 33 pumilio 54 Anisomeris 95 radicosa 49 brachyloba 104 spinosa spinosa 957 — var. suaveolens 54 Antirhea Bouvardia 27, 55 panamensis 100 glabra Anti 34 flor: + Peeves iria St, 38 blaerioides 387 lavarum 38 Aspidanthera 177 rudgeoides 177+ Bertiera 3+, 5+, 4 aequaliramosa 43 panamensis 248 angustifolia 43 gs coma 27, 47, 59, 671, 787 nensis 427, ac oF 43 —var. leiophylla 44 —var. gracilis 55 — var. obtusa 55 triphylla 55* Buena 190 panamensis 1907, 193 8 candidissimum 57*, 57 Cascarilla 106 Cassupa S be r^ 61* 252 bristanii 62 camponutans 63 chiriquiensis 70 correae 64 costaricensis 70 cro atij ia dichro dimorphandrioides 66 discolor dressleri p 791 elata 70 gaugeri 71 glomerulata 71 guapilensis 67 insueta 73 Visier dd 21, 74 kenne iun 76 cosa $97, 76* nana 77 Buc 67 panamens panchocoënsis 78 Chimarrhis 47, 82 decurrens 22 latifolia 83 alba durifolia 87 jefensis 88 s. 86 ee 87+ petens a 90 racemos: 1 847, 86 var. glaberrima 86 Chione 3t, 4t, 90 allen buxifolia 92, 93+ englesingii 102 filipes 104 grandicarpa 97 leucophylla 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mb deni" a — Var. une officinalis 1067 p tucuyensis 106 Coccocipsilum 108+ Coccocypselum 108, 108* canescens | cordifolium 108 anceolatum 11 macropodum 187 pleuropodum 109 repens 108+, 109 Parr 109 Coffe Minn Tes 113 — Condaminea 4+, 57, 113 corymbosa 113+, 115 unie 19% O ‘obtusifolia l me neri Tt Coussarea 4t, St, 120, 1257, loftonii 129 mibuena 4*, 5*, 117, 1291, 131* [ Vol. 67 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) morii 130 nebulosa 131, 1347 132 s M 1307, 1317. 132, s —var. ane 137 glabra longiflora 139 rubra 137+ ieclienzis 4*, 140 chiococcoides 140* fruticosa ye Deppea 47, ee 143 46 —var. alsinoides 146+ —var. australis 146, 1467 —var. rts 146 mexica TUS Da 147. 149* connellii virginiana 150* Dressleriopsis 4*, 153 54 panamensis 221 Elaeagnia 156 auriculata 156 nitidifolia 157 ruiz-teranii 1587 vdd 38 a 59 iets 61 di ehroa 65 elata 70 E lends 67 guianensis 591 ipecacuanha 74 , 1297, 161, altipetens m bocataurensis 163 bullata 164 caput-anguis 164 corymbosa 161* loftonii 129 luteovirens 167, 171* martini 132 BEGIdenpos 4*, 168 ) — var. suerrensis 174 woodsonii 167+, 17 —var. glabratum 179 —var. platyphyllum 179 1647, 171+ 233 254 mollugo 178+ nelsonii 181 obstipum 181 uncinulatum 181 nropetalum 179 americana 1817, 182 —var. caruto —fo ee nol 182 barbata 18 caruto 182 curviflora 184 ; 19 erandifolia 182 humilis 182 oblongifolia 182 pubescens 182 herbacea 1867, 188 —violaefolia 187 violaefolia 187 Gomphos 177 Gonna "de 190+ acutifol ependens 190+ kallunkii 192 l ntha 193 Gonzalea 190 chiriquiensis 199 costaricensis 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN foliacea 200 EKRIR 204 Hamelia 27, 3t, 4t, 204, 2047 klugii 206 macrantha 207 magnifolia 208 magniloba 212 patens 208 var. coronata 211 2 = ° ct e3 2 = £ N Hillia 5+, 215, 2187, 219+ chiapensis 216 —subsp. chiapensis 216+ —subsp. grandifolia 216 var. chiapensis 216 chiriquiensis 216 tetrandra 219%, 220 Sra 22+ Hippot albifiora 221 davidsoniae 229 eliasii 229 fimbrianthera 230 [Vol. 67 236%, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) gentryi 230 haydenii 231 alsinoides 146 longicalycina 234 ma anussatani 234 E cns 238, 2427 pustulata 2 rexmontis 241 rubripigmenta 242 subauriculata 2267, 242 vesciculifera 237+, 243 Sho 244 woodson Soi. fruticosa 141 Ipecacuanha 5 par dn 74 Isertia 37, coccinea gr p 248 spicaeformis 247 Ixora 2*, 4* occidentalis 168 Lonicera "204, 204* alba 8 M St, 95+ dida 57 candissima 57 Posoqueria 3+, 47, 5+ Pseudochimarrhis 82 turbinata 827 Pseudorhachicallis 38 Psychotria 2+, 3+, 44, 607, 627, 781 Richardia 2+, 5+ Ronabea ecta 627 Rondeletia 2+, 37, 5+ s 188 ubia acuminata 179 Rubiaceae 1, 27, 3+, 4t, S+, 22+, 27%, 951, 1357, 1545, 1907, 223+, 237+ —tribe E 5t Rustia 57, Sabicea ambigua 30*, 31 edulis 19 Sacconia 90 Solenandria 159% 255 ixoroides 159+ longiflora 139 ocimifolia 213+, 213 ocimoides 53 spinosa 49 suffruticosum 49* tenuiflora 139 Steudelago 159 Sturmia 34 lucida 34+ Styracaceae 198+ Tangaraca 204 violacea 59% ANNALS OF THE MISSOURI BOTANICAL GARDEN Tardavel 49 laevis 50 Tetramerium odoratissimum 168 Tocoyena 3+, 47, S+ costaricanum 1207 Witheringia 244+ [Vol. 67 THE BOTANY AND NATURAL HISTORY OF PANAMA A symposium to signal completion of the Flora of Panama will be held in Panama 14-17 April 1980. Sessions will include the physical setting, archaeology. vegetation, flora, fauna and demography of Panama and nearby countries as well as other natural history related topics. Conservation of endangered landscapes will be a theme. Both invited and contributed papers will be given, and either Spanish or English may be used. Limited funds may be available to assist costs of attending this meeting. The symposium is sponsored jointly by the Missouri Botanical Garden. the University of Panama, and the Smithsonian Tropical Research Institute. Contact: Prof. Mireya D. Correa A. Escuela de Ciencias Naturales y Farmácia Estafeta Universitaria University of Panama Republic of Panama The previous issue of the ANNALS of the Missouri Botanical Garden, Vol. 66, No. 4, pp. 591—904 + i-v was published on 22 February 1980. ANNALS OF THE MISSOURI BOTANICAL GARDEN FLORA OF PANAMA BY ROBERT E. WOODSON, JR., AND ROBERT W. SCHERY and Collaborators Part IX FAMILY 179. RUBIACEAE-—PART Ilt JOHN D. DWYER 49. IXORA Ixora L., Sp. Pl. 110. 1753; Gen. PI., ed. 5. 48. 1754. TYPE: I. coccinea L. Sideroxyloides Jacq., Sel. Stirp. Amer. 19. 1763. TYPE: S. ferreum Jacq. Siderodendron Schreb., Gen. Pl. 1: 71. 1789. TYPE: S. floribundum A. Rich. Trees or shrubs. Leaves occasionally ternate, usually thick; petioles occa- sionally wanting; stipules free, usually acuminate. Inflorescences terminal or ax- illary, corymbose, the pedicels of the flowers bracteolate. Flowers variously col- ored, the hypanthium ovoid, the calycine cup 4(-5)-lobed; corolla salverform, slender, the lobes 4—5, contorted; stamens 4(—-5), the anthers linear, dorsifixed, exserted; ovarian disc swollen, the style slender, the stigmas 2, exserted, the ovary 2-celled, the ovules solitary, peltately attached to the middle of the septum. Fruits baccate, leathery or fleshy; seeds concave convex, the testa membranous. Ixora, a genus of about 400 species, is widely distributed throughout the trop- ics of the world. a. Inflorescences capitate; corolla tube 30-45 mm long. . Leaves sessile or subsessile; corolla red or orange 1. I. coccinea bb. Leaves obviously Seien corolla wh ite 2. I. findlaysoniana + Part I appeared i in Volume 67(1) of the Annals of the Missouri Botanical Garden Missouri Botanical Garden, Box 299, St. Louis, Missouri 63166, and St. Pan University, 221 North Grand, St. Louis, Missouri 63103. ANN. Missouni Bor. GARD. 67: 257—522. 1980. 0026-6493/80/0257-0522/$26. 65/0 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 aa. bal jdn cymose paniculate, spreading; corolla tube 3-5 mm Leaves 16-24 cm long; peduncles ca. 2.2 cm long; flowers oe or addu fruits ca. t cm in diam I. flo ue cc. Leaves 7-16 cm long; ee 5-9 cm long; flowers with pedicels to 5 mm lon fruits ca. 0.4 cm in dia 4. I. Mei — . Ixora coccinea L., Sp. Pl. 110. 1753. TYPE: ?Ceylon, herb. Linn. (LINN 131.1, not seen, microfiche MO). Shrubs 0.5—2.0 m tall, much branching, the twiglets glabrous, the nodes well spaced. Leaves sessile or subsessile, oblong, ovate oblong, elliptic or occasion- ally obovate oblong, 2.0-8.5(-—16.0) cm long, 1.2-6.2 cm wide, rounded, deltoid to subacute at the apex, often with a terminal bristle to 1 mm long, cordate, obtuse, basally truncate or subobtuse, the lateral veins 10-13, joining to form an irregular, undulate submarginal vein, the intervenal areas spreading reticulate, coriaceous, glabrous; stipules oblong, to 6.5 cm long, usually shorter, the median awn as long as the body. Inflorescences terminal, the flowers in dense corymbs, the aggregate to 7 cm long, occasionally wider than long, the peduncle usually suppressed or to 1.2 cm long, occasionally with 1—2 pairs of opposite branches, the lowermost to 2 cm long; bracts and bracteoles ovate, to 0.8 cm long, acute. Flowers sessile; hypanthium subrotund to oblong rotund or turbinate, glabres- cent, the calycine cup reduced, the lobes 5, ascending, triangular, ca. 1 mm long, acute; corolla red or orange, the tube linear infundibuliform, 3.5—4.5 cm long, 0.1 cm wide, glabrous or minutely puberulent, the lobes ovate oblong or oblong, 1.2- 1.5 cm long, 0.8 mm wide, acute to widely deltoid, glabrous; stamens 4, the anthers soon deciduous, narrowly ovate oblong, 3.5—4.2 mm long, caudate acu- minate for ca. 1 mm, sagittate basally, the filaments shorter than the anthers, attached at the mouth; style scarcely exserted or exserted for 5 mm, the stigmas | mm long. Fruits not seen. Ixora coccinea is native to Asia. It is commonly cultivated in Central and South America, in the West Indies and in Madagascar. The inflorescence is usu- ally spreading and the peduncle is obvious. Croat 15624 notes: "'fruits red to dark violet, mesocarp sweet, pulpy; seeds 2." One collection, Correa 1142, has large leaves reaching to 16 cm in length and distinctly petiolate. "Buquet de Novia"; ‘*Cacho de Toro"; “Jazmin de Coral." S DEL TORO: Bocas, airport runway, Lazor et al. 2430 (MO). CANAL ZONE: Diablo Heights, EE a "10. 1142 (both MO). Frijoles, Ebinger 311 (MO). Coco Solo, Gentry 6063 (MO). Gaillard Highw: etween Summit Garden and Gamboa, Sullivan 21 (MO). Miraflores Locks, Tyson 1138 (MO). Fort Sherman, Tyson & Blum 3788 MO). BARRO COLORADO ISLAND: Croat 4061 A, 7051, 15264 (all MO); Hayden 15264 (MO); Lourteig et da 3027 (MO). COCLE: Penonomé, Quiros 5 (MO). Turnoff from main highway to Cerro Azul, Reece 1 (MO). DARIEN: El Real, Lazor & Correa 3368 (MO). PANAMA: Juan Diaz, Mendez s.n. (MO 2. Ixora findlaysoniana Wall. ex G. Don, Gen. Syst. 3: 572, 1834. TYPE: not seen. Shrubs or trees, to 4.5 m tall, the twigs terete, smooth, glabrous, ultimately angular, the pith porate, the nodes 1.5-5.0 cm apart. Leaves narrowly obovate to oblong, occasionally subfalcate, 7-18 cm long, 2.5-6.5 cm wide, obtuse to cuneate at the apex, occasionally rounded, basally cuneate, often attenuate acute, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 259 " NM a ey = T WSs Z NY mS ATP, NR WAR = SS FIGURE 54. Ixora floribunda (A. Rich.) Griseb. Habit (x!4). [After Dwyer & Robyns 35.] the costa prominulous or immersed above, subprominent beneath, especially proximally, the lateral veins ca. 9, arcuate, thinly coriaceous, concolorous, gla- brous; petioles to 2 cm long, 0.10-0.22 cm wide; stipules free or occasionally connate, oblong or hemispherical, 0.4-0.7 mm long, with 1-2 subulate awns long- er than the body, arising from 1-3 scarcely elevated longitudinal ridges on the body. Inflorescences terminal, capitate, to 4.5 cm long, ca. 9 cm wide, rounded. Flowers glabrous; hypanthium short, the calycine cup ca. 0.5 mm long, eglan- 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 dular, the calycine lobes foliaceous, widely elliptic, to 3.5 mm long, to 2 mm wide, acute, thickly petaloid; corolla white, the tube narrowly cylindrical, to 32 mm long, | mm wide, petaloid, the lobes 4, spreading or reflexed, oblong, 8 mm long, to 3 mm wide; stamens 5, exserted, the anthers linear ovate, 1.8 mm long, acuminate at the apex, sagittate at the base, the filaments short, attached at the mouth: ovarian disc doughnut shaped, compressed, the style linear, as long as the corolla tube, the stigmas narrowly oblong, to 2 mm long. Fruits not seen. Ixora findlaysoniana is widely cultivated in the tropics; it is probably native to Southeast Asia. ''Lila CANAL ZONE: Summit Gardens, Nee 10859 (MO). PANAMÁ: Arraijan, Carrasquilla & Rincon 295 (MO). Paitilla, Correa 40 (MO). SAN BLAS: Trail to inland village of Armila, 3-8 km SW of Puerto Obaldia, Mori et al. 6805 (MO). Niatuppu, Stier 222 (MO). VERAGUAS: San Francisco, Dwyer 1299 (MO). 3. Ixora floribunda (A. Rich.) Griseb., Cat. Pl. Cub. 134. 1866.—FiG. 54. Siderodendron floribundum A. Rich. in Sagra, Hist. Cuba 11: 24. 1850. Type: Cuba, Richard (?P, not seen). Small trees, to 5 m tall, the twigs terete, smooth, glabrous, the nodes well spaced. Leaves oblong or obovate oblong, 16-24 cm long, 7-10 cm wide, widely cuneate to subobtuse at the apex, acuminate, the acumen to 1.5 cm long, ulti- mately acute or obtuse, attenuate acute to somewhat rounded at the base, the costa slender, immersed, plane or somewhat prominulous above, proximally prominent beneath, the lateral veins 9-13, arcuate, thinly coriaceous, concolor- ous, glabrous; petioles stout, often curved, to 2.5 cm long, to 0.25 cm wide in the middle with a delicate adaxial wing, glabrous; stipules deltoid, to 5 mm long, ca. 1 mm wide at the base, acute, stiffly scarious, rugulose, densely villose on the margin and within, otherwise glabrous. Inflorescences terminal, pyramidal pan- iculate, shorter than the terminal leaves, epedunculate or with peduncles to 2.2 cm long, angular, pubescent, the branches disposed as 2-3 pairs, to 2 cm long, the flowers aggregated into subrotund cymules, ca. 1 cm in diam. Flowers sessile or shortly pedicellate; hypanthium subrotund, ca. 1 mm long, villosulose, with numerous oblong glands within, ca. 0.1 mm long, the lobes 5, oblong, obtuse, ca. 0.5 mm long; corolla white, the tube narrowly cylindrical, ca. 4 mm long, ca. 0.7 mm wide, glabrous outside, densely white villose within, the lobes 4, oblong, ca. 4 mm long, concave, ca. 1.5 mm wide, petaloid, glabrous; stamens 5, the anthers narrowly oblong, ca. 3.3 mm long, dorsifixed, the filaments plane, 0.6-1.5 mm long; stigmas 2, unequal, oblong, connate, ca. 1.5 mm long, the ovary oblong in cross section, the wall thick, the septum thick, the ovules several. Fruits sessile, rotund, ca. 1 cm in diam., the persistent calycine tube ca. 0.5 mm long, drying red, smooth, sparsely villose. Ixora floribunda occurs in Salvador, Nicaragua, Costa Rica, Panama, and Colombia, and the Antilles. The stipules of the type of Siderodendron floribun- dum A. Rich. are described as '"'obtussimis," which does not agree with my description of the stipules as "acute." “Cacho de Toro” (Holdridge 6219). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 261 CANAL ZONE: Albrook Site, U.S. Army Tropic Test Center Site, Dwyer & Robyns 35 (MO). DARIEN: Between Pinogana and Yaviza, Allen 298 (MO). Los SANTos: Coabal, Progreso, W slope above Rio Cobachén, Holdridge 6219 (MO). PANAMA: Pacora, Paul 785 (MO). 4. Ixora nicaraguensis Standley, Wernham, J. Bot. 50: 243. 1912. TYPE: Nicara- gua, Seemann 117 (?BM, not seen). I. rauwolfioides Standley, Trop. Woods 11: 25. 1927. type: Panama, Cooper & Slater 93 (US, ho- Shrubs or trees, to 7.5 m tall, the twiglets drying grey, smooth, glabrous, somewhat twisted. Leaves elliptic, oblong or oblanceolate, 7-16 cm long, 2-7 cm wide, short to long acuminate at the apex, acute at the base, the lateral veins 10— 15, slender, arcuate, prominulous, the intervenal areas spreading reticulate, char- taceous, discolorous, glabrous; petioles 0.4—1.0 cm long; stipules free, triangular, 0.5-0.8 cm long, acuminate to caudate acuminate at the apex. Inflorescences cymose paniculate, spreading, sessile or pedunculate, the peduncles 5-9 cm long, the branches paired, few, well spaced, many flowered; pedicels to 5 mm long. Flowers with the hypanthium turbinate, ca. 1 mm long, puberulent, the calycine cup 0.5 mm long, stiffly carnose, rubescent, glabrous, eglandular within, the teeth absent or scarcely developed; corolla white, the tube cylindrical, 3-5 mm long, glabrous outside, glabrous within except villose at the mouth, the lobes 4, oblong, 3-4 mm long, rounded, rubescent, subcarnose; anthers 4, narrowly ovate oblong, 3 mm long, acuminate, the acumen petaloid, the filaments 1 mm long; ovarian disc ca. 0.3 mm long, the style linear, 3 mm long, the stigmas oblong, ca. 1.5 mm long. Fruits pink, rotund to oblong rotund, to 4 mm in diam., smooth, glabrous. Ixora nicaraguensis extends from Belize to Panama. CAS DEL TORO: Changuinola Valley, Cooper & Slater 93 (US). cHiRIQUÍ: Puerto Armuelles, Woodson & Schery 851 (MO). San Bartolomé, Burica Peninsula, Woodson & Schery 851 (MO). San Bartolomé, Burica Peninsula, Woodson & Schery 880 (MO). DARIÉN: Río Urucea, ibis 1451 (MO). 50. JOOSIA Joosia Karst., Fl. Colomb. 1: 9. Tab. 5. 1858. TYPE: Colombia, J. umbellifera Karst. Trees or shrubs. Leaves usually elliptic or oblong, acute to long acuminate at the apex, tapering basally; petioles short; stipules deciduous except usually per- sistent at the tips of the twigs, elongate, free. Inflorescences terminal, cymose umbellate, the flowers secundly disposed. Flowers with the hypanthium short, the calycine cup and teeth short; corolla white, the lobes valvate, ovate, bifid at the apex, the margins conspicuously scalloped; stamens with anthers often ses- sile, attached below the middle of tube; ovary 2-celled, the ovules numerous, the placenta adnate to the middle of the septum. Fruits capsular, linear or narrowly oblong, splitting into 4 spirally coiled valves; seeds imbricate, membranous, winged, the wing usually bifid at the base, entire or dentate at the apex. Joosia, a genus of 4 species, is known from Panama, Colombia, and Peru. 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1. Joosia panamensis Dwyer.% Type: Panamá, Croat 25932 (MO).—FIG. 55. Shrubs or small trees, the branchlets glabrescent, often covered with mosses. Leaves elliptic or rarely obovate trapeziform, 16-22 cm long, 6-9.5 cm wide, acute or widely deltoid toward the apex, cuneate or acute at the base, slightly inequilateral, the costa prominulous above, prominent beneath, the lateral veins 10-15, arcuate, the veinlets dense, arising at right angles from the costa, stiffly chartaceous, glabrescent above, beneath with white, appressed hairs; petioles to 1.5 cm long, villose; stipules free, ovate oblong, ca. 2 cm long, ca. 0.5 cm wide at the base, obtuse, red when dry, the hairs few, white. Inflorescences terminal, perhaps solitary, cymose umbellate, the peduncle, rigid, slender, to 8 cm long, ca. 0.15 cm wide, often twisted, densely pubescent, the flowers secundly disposed and crowded on short branches arising radiately from the apex of the peduncle; pedicels ca. 2 mm long. Flowers with the calyx cup ca. 2.5 mm long, ca. 3 mm wide, cup shaped, expanded, densely ciliolate outside, glabrous within, the lobes 4, oblong, ca. 2 mm long, to 1.5 mm wide, rounded at the apex, glabrous within; corolla white, the tube narrowly cylindrical, ca. 9 mm long, ca. 1.8 mm wide, carnose, the hairs outside appressed, minute, glabrous within, the lobes 5, ovate oblong, to 3 mm long, with 2 ovate elliptic appendages within, to 2.8 mm long, petaloid, glabrous, scalloped; anthers 5, sessile or subsessile or filaments to 2.5 mm long, narrow oblong, ca. 3.5 mm long, attached to the tube at or below the middle, often near the base; style (2-)4-10 mm long, the stigmas 2, unequal, scarcely wider than the style, to 2 mm long. Fruits linear oblong, to 2 cm long, to 0.35 cm wide, terete, scarcely angular, the ribs few, prominulous, minutely glandular punctate; seeds spindle shaped ca. 6 mm long, the wing bifid or ero- sulose at the base, acute, entire or fimbriate at the apex. Joosia panamensis is known only from Panama. It is related to J. umbellifera Karst., a well-known Colombian species. It differs in having much shorter fruits, which measure only about a 4 as long as those of J. umbellifera, much shorter seeds, and presumably shorter stipules. BOCAS DEL TORO: Chiriqui Trail between Buena Vista Coffee Finca & Cerro Pilon, Kirkbride & Duke 680 (MO). COCLE: cae Calvario around Rivera sawmill, 7 km N of El Copé, 700-900 m, ries 3240 (MO). COLON: 2-3 mi up Rio Guanche, Kennedy & Foster 2136 (MO). PANAMA: 16-20 e Panamerican Highway er from El Llano ar Tupile, 400 m. Kennedy 2694 (MO). El. J ano Carti Road, 18 km from Panamerican Highway, Mori et al. 4592 (MO). VERAGUAS: Road between Escuela Agricola Alto de Piedra bene Santa Fé and Río Dos age 730-770 m, Croat 25932 (MO). Valley of Rio Dos Bocas between Alto Piedra and Calo ihe 350-400 m, Croat 27471 (MO). Caribbean slope above Rio Primero Brazo, 5 mi NW of San a Fé. Liesner 1014 (MO). 8.8 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 3220, 2 (both MO). 16 km from Santa Fé, near Rio 'Caloveborita, Mori 6688 (MO). 66 Joosia panamensis Dwyer, spec. nov. Frutices vel arbores parvae. Folia elliptica vel interdum obovato-trapeziformia, 16-22 cm longa, 6-9.5 cm lata, venis lateralibus 10-15, arcuatis; petiolis ad 1.5 cm longis; stipulis ovato-oblongis, ad 2 cm longis. /nflorescentiae terminales cymoso-umbellatae, pedunculo ad 8 cm longo, floribus in ramis brevibus secundatim dispositis, ramis radiate dispositis. jin pedicellati, pedicellis ad 2 mm longis, cupula calycis ca. 2.5 mm longa, extus dense ciliolata, lobis 4, ca. 2 mm longis; corolla alba, tubo ca. 9 mm longo, extus pubescente, lobis 5, ovato-oblongis, M mm longis, ad marginem fimbriatis; stamina 5, antheris sessilibus vel subsessilibus, angusto- oblongis, ca. 3.5 mm longis, infra medium tubi affixis. Fructus lineari-oblongi, ad 2 cm longi, teretes, costis paucis prominulis; seminibus ellipticis, ad 6 mm longis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 263 Wert SM DOES PORA SY WS AA A (o S V S SS Si NAN BY FIGURE 55. Joosia panamensis Dwyer.—A. Habit (x!2).—B. Flower opened to show interior (x2).—C. Ovary longitudinal section (x5). [After Kennedy & Foster 2136.] 51. LADENBERGIA Ladenbergia Klotzsch in Hayne, Arzn. Gew. 14: pl. 15. 1846. LECTOTYPE: L. dichotoma Klotzsch. Cascarilla Wedd., Ann. Sci. Nat. Bot. Sér. 3. 10: 10. 1848, not Cascarella Adanson, 1763, not Cascarilla Raf. 1838. LECTOTYPE: not selected. 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Trees or shrubs. Leaves opposite, occasionally verticillate, coriaceous, peti- olate; stipules large, often deciduous, glandular within at the base. /nflorescences terminal, paniculate. Flowers with the hypanthium ovoid or turbinate, the caly- cine cup cupuliform, corolla with the tube somewhat funnelform or salverform, replete with fibers, glabrous within, the lobes 5, valvate, papillose within and on the margins; stamens $5, the anthers linear, included, dorsifixed; ovarian disc pulvinate, the ovary 2-celled, the style slender, the stigmatic lobes 2, included, slender, the placentas adnate to the septum, the ovules numerous. Fruits cap- sular, narrowly clavate or cylindrical, septicidally bivalvate from the apex to the base; seeds numerous, imbricate, peltate, the body oblong, surrounded by a mem- branous, dentate or laciniate wing. Ladenbergia is known from Costa Rica, Panama, and tropical South America. It is a genus of about 30 species. Mori & Dressler 7850 (MO) is probably a new species but has only immature leaves. It is the only material seen from the Province of Chiriqui although La- denbergia has been collected several times in Costa Rica near the Panamanian border. It was collected in the Province of Chiriqui on Cerro Colorado, 50 km N of San Felix on the continental divide in cloud forest, alt. 1,200-1,500 m. a. Leaf blades bullate l. L. bullata aa. Leaf blades not bullate. b Leaves 8-15 cm long, 3.0-4.3 cm wide; lateral veins ca ---————-—— 2. L. laurifolia bb. Leaves (10-)20-45 cm long, more than 6 cm wide; eie veins 10-12... 22... -. 3. L. macrocarpa l. Ladenbergia bullata (Wedd.) Standley, Trop. Woods 34: 41. 1933. Cascarilla bullata Wedd., Ann. Sci. Nat. Bot. Sér. 3. 10: 11. 1848. TYPE: not seen. Trees or shrubs to 5 m tall, the twigs pubescent or glabrescent, ultimately angular, the nodes well spaced below, somewhat crowded terminally. Leaves oblong rotund, 11.5—38.0 cm long, 8-26 cm wide, rounded or obtuse at the apex, not acuminate, basally rounded, obtuse or truncate, the costa plane or immersed above, prominent beneath, to 0.35 cm wide, the lateral veins 8-11, widely arcuate, strict, prominent beneath, the intervenal areas spreading reticulate, coriaceous, drying dark brown above, bullate, glabrescent to golden villosulose above, golden appressed pilose beneath; petioles 2.0-4.5 cm long, rigid, lignose, golden villo- sulose; stipules not seen. /nflorescences terminal, solitary, golden villosulose, cymose paniculate, to 14 cm long, to 10 cm wide, the peduncles 0.5-4.5 cm long, the lowermost branches opposite, 1-2 cm long, angular, usually with 2 pairs of opposite, strongly ascending, superior branches. Flowers sessile or subsessile; hypanthium subturbinate, ca. 1 cm long, densely golden villosulose, the calycine cup cupuliform, ca. 3 mm long, coriaceous, pubescent outside, glabrous within, the lobes 5, 1-2 mm long, oblong to rotund, obtuse or rounded, coriaceous, golden villosulose on margins and outside, glabrous within, drying much darker than the hypanthium; corolla white, the tube narrowly cylindrical, 3.5-5.0 cm long, 0.35-0.50 cm wide, densely villosulose outside, the hairs occasionally tuft- ed, glabrous but rugulose within, coriaceous, the lobes narrowly oblong, 2.0-2.5 cm long, crassate, carnose and rugulose within; stamens 5, the anthers with tips 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 265 barely exserted from tube, narrowly oblong, ca. 7 mm long, the filaments ca. 2.5 mm long, glabrous, somewhat geniculate at point of attachment, attached ca. 1 cm below the mouth; style ca. 25 mm long, plano compressed, villose, the stig- matic lobes 2, ca. 1.8 mm long. Fruits with pedicels to 1.5 cm long, narrowly oblong, to 9 cm long, the valves at dehiscence to 2 cm wide, obtuse or rounded at the apex, cuneate at the base, smooth, rugulose, glabrescent at maturity, with a few slender longitudinal prominulous lines outside, the calycine cup persistent in unopened fruit, finally deciduous in open fruit. PANAMA: 8 mi S of Goofy Lake toward Cerro Jefe, Dwyer 7080 (MO). Cerro Campana, Gentry 1840 (MO). Cerro Jefe, Gentry et al. 3407 (MO); Gentry & Dwyer 5542 (MO). Cerro Azul, just E of Goofy Lake, Gentry & Mori 13403 (MO). El Llano-Carti Road, Liesner 1260 (MO); Mori et al. 4681 (MO). 2. Ladenbergia laurifolia Dwyer,% spec. nov. TYPE: Panama, Croat 25586 (MO). Trees to 5 m tall, the twigs terete, the bark grey when dry, lenticellate, the nodes well spaced, the scars of the pedicels oblong rotund, ca. 4 mm in diam. Leaves narrowly oblong or narrowly obovate oblong, 8-15 cm long, 3.0-4.3 cm wide, deltoid toward the apex, acuminate, the acumen triangular, to 0.6 cm long, obtuse, widely cuneate or obtuse at base, the costa prominulous above, sub- prominent beneath, dark red, the lateral veins ca. 10, subplane on both sides, arcuate, the submarginal vein undulate and inconspicuous, the margin lightly revolute; petioles lignose, rigid, glabrous, to 2 cm long; stipules deciduous. /n- florescences terminal, to 11 cm long, golden puberulent, dark red when dry, the peduncle to 0.2 cm wide, the branches 3, terminating a small branch, the lateral branches shorter and with 3 secondary branches at the apex, the cymes few flowered, the middle branch similar to the lateral branches but with the cymes spreading and often on branches to 1.5 cm long; bracts oblong ovate, to 2 mm long, puberulent, the bracteoles deciduous, to 1 mm long. Flowers sessile; hy- panthium turbinate, ca. 4 mm long, densely sericeous, the calycine cup somewhat swollen, ca. 0.5 mm long, the lobes 5, widely triangular or ovate, 0.3-1.2 mm long, obtuse, carnose, glabrescent outside except at the base and the margins, red when dry, minutely punctate; corolla white, the tube narrowly hypocrateri- form, to 1 cm long, to 0.5 cm wide, densely sericeous outside, stiffly carnose, the lobes 5, oblong, to 1.2 cm long, to 0.3 mm wide, glabrous within; stamens 5, the anthers linear oblong, ca. 5 mm long, ca. 0.6 mm wide, obtuse, the filaments ca. | mm long, attached near the middle of the tube; ovarian disc cushion like, ca. 0.4 mm long, the style slender, ca. 1.5 cm long, the stigmas 2, linear oblong, ca. 2.2 mm long, puberulent(?), the ovules membranous, winged, plano com- pressed, oblong, ca. 0.3 mm long. Fruits not seen. Ladenbergia laurifolia is known only from Panama. The general aspect of the 87 Ladenbergia pe Dwyer, spec. nov. Arbores ad 5 m altae. Folia angusto-oblonga vel sierra -obovato-oblonga, 8-15 cm longa, 3.0-4.3 cm lata, versus T deltoidea venis lateralibus ca. 10; petiolis ad 2 cm pits Inflorescentiae ee ad 11 cm longae, aureo-puberulae, cymis pauciforis bracteis oblongo-ovatis, ad 2 mm longis. Flores sessiles; cupula calycis ca. 0.5 mm longa, lobis 5 1.2 mm longis; corolla alba tubo ad 1 cm longo, lobis 5, ad 1.2 cm longis; antheris ca. 5 mm Ses subsessilibus. Fructus non visi. 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 L. nour gam FıGURE 56. Ladenbergia macrocarpa (Vahl) Klotzch. Habit (<3/10). [After Mori & Kallunki 5501.] leaf resembles that of a Lauraceae, the color suggesting the dried blades of a Myristicaceae. L. laurifolia has the smallest leaves among the Central American species of the genus. The flowers are markedly small. VERAGUAS: Beyond Río Tres Brazos, 9 km from Santa Fé, Croat 25586 (MO). 3. Ladenbergia macrocarpa (Vahl) Klotzch in Hayne, Arzn. Gew. 14: 15. 1846.— Fic. 56. Cinchona macrocarpa Vahl, Skiv. Naturh. Selsk. 1: 120. 1790. TYPE: not seen. Cascarilla macrocarpa (Vahl) Wedd. Ann. Sci. Nat. Bot. Sér. 3. 10: 13. 1848. Trees to 40 ft tall, the trunks often buttressed, the branches smooth, glabrous, usually angular. Leaves oblong or obovate oblong, 10-45 (or more) cm long, 6- 24 cm wide, obtuse at the apex, acuminate, the acumen short or evanescent, the costa plane or prominulous, to 0.5 cm wide, expanded toward the petiole, prom- inent beneath, the lateral veins 10-12, prominent beneath, coriaceous, glabrous to densely golden pubescent above, glabrous to densely golden villosulose or 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 267 appressed pilose beneath, often the axils barbate, shiny, dark brown when dry; petioles to 4.5 cm long; stipules crowded toward the apex of the twigs, persistent or deciduous, oblong rotund, 1.5-3.5 cm long, to 2.5 cm wide, rounded at the apex, coriaceous, glabrescent, venose, shiny. Inflorescences terminal, solitary, minutely pubescent or glabrescent, to 20 cm long, to 22 cm wide, presumably epedunculate, the several branches sharply ascending and terminating the twig- lets, 4-9 cm long, angular. Flowers scarcely pedicellate; hypanthium narrowly turbinate, to 0.8 cm long, tapering gradually into the pedicel, drying blackish, smooth, glabrous or puberulent, the calycine cup bowl shaped, to 0.7 cm long, the teeth triangular to rotund, 1.5-2.8 mm long, often obtuse; corolla with the tube narrowly cylindrical, 3.5-5.0 cm long, apically 3-4 mm wide, the lobes narrowly oblong, ca. 3.5 cm long, ca. 0.35 cm wide, densely golden pubescent; anthers 5, subsessile, ca. 6 mm long, ca. | mm wide, attached near the mouth. Fruits scarcely pedicellate, narrowly oblong, often slightly falcate, 7.0-13.5 cm long, 0.8-1.2(-1.8) cm wide, obtuse at the apex, attenuate toward the base, lig- nose, glabrescent to densely golden pilose, drying black brown or golden tan, delicately longitudinally striate ribbed, the persistent calycine cup 6-8 mm long, the calycine lobes ca. | mm long, the valves at dehiscence to 1.8 cm wide; seeds narrowly oblong or subfusiform, 1.0-2.5 cm long, the body 1-5 mm long, the wings often split or erosulose at the distal pole, acute at the proximal pole. A collection of this species, Mutis 6219 (F), from Colombia has a manuscript name L. colombiana which has not been validly published. Ladenbergia macrocarpa is known from Panama, Colombia, and south to Boliva. *‘Fruta Mono," ‘‘Michurajo.”’ ARIEN: Cloud Forest on Cerro Pirre, 3700 ft, Duke 6571 (MO). W slope of Cerro Pirre, Duke 6590 (MO). Cana-Cuasi Trail between Cerro Campamento and La Escalera, E of Tres Bocas, Kirk- bride & Duke 1313 (MO). N slope of Cerro Pirre, lower montane rain forest, 700-950 m, Mori & Kallunki 5501 (MO) 52. LIMNOSIPANEA Limnosipanea Hook. f. in Hook., Icon. Pl. Ser. 3. 1(11): 38, pl. 1050. 1868. TYPE: Brazil, Limnosipanea spruceana Hook. f Sipania Seemann, Bot. Voy. Herald 136. 1853. TYPE: S. palustris Seem., not Sipanea Aubl. 1775, (Rubiaceae). Herbs, slender, erect, branching above, the branches dichotomous. Leaves 3 to numerous at a node, verticillate, the uppermost occasionally opposite; stip- ules obsolete, often reduced to a subulate projection. Inflorescences cymose with flowers solitary or in few-flowered terminal clusters in the angles or along the length of usually dichotomous branches. Flowers with the calycine lobes some- what foliaceous, persistent; corolla variously colored, hypocrateriform, the tube narrow, glabrous within, the mouth glabrous or variously pubescent, the lobes 5, oblong, contorted; stamens 5, inserted at the mouth, the anthers oblong, dor- sifixed, the filaments slender; ovary 2-celled, the ovules numerous, disposed on a median septum. Capsule ovoid, scarious, loculicidally dehiscent, the seeds numerous. 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Limnosipanea is a genus of about 7 species, one of which is in Panama, the others in tropical South America. Literature: Standley, P. C. 1928. Flora of Panama Canal Zone, Contr. U.S. Natl. Herb. 27: 355. Steyermark, J. 1974. Limnosipanea Hook. f. Flora de Venezuela 9(1): 384. Wernham, H. F. 1917. The Genus Sipanea. J. Bot. 50: 171-175. 1. Limnosipanea palustris (Seem.) Hook. f. in Hook., Icon. 1050 (p. 38). 1868. die palustris Seem., Bot. Voy. Herald 136. 1853. rvPE: Panama, Seemann 347 (BM, excl. spec- n Microlicia, Melastomalaceae, not seen; MO, photo of holotype). Perennial? herbs, the stem slender, erect, branching only in the upper !^, pubescent, the nodes well spaced. Leaves ternate, the uppermost opposite, ses- sile, erect, except the basal leaves often deflexed, oblong, ca. 8 mm long, ca. 2 mm wide; stipules not seen. Flowers small, solitary or in few-flowered cymules, terminating a pair of dichotomously disposed, angular, ascending branches, pre- sumably a solitary flower in the angle of the dichotomy. Limnosipanea palustris is known only from the Seemann collection (Panama) and another from Colombia (Goudot?), neither seen by the author except for a photograph of the first. As the species is known to me only by a photograph, floral details have been omitted in the diagnosis. The type collection (Seemann 347, BM) has two different elements, one a Melastomataceae which superficially looks like Limnosipanea palustris but is probably a species of Microlicia. The specimen, on close examination, seems to have been first mounted on a separate piece of paper, this then pasted to the overall sheet containing the type material of Limnosipanea palustris. Standley (1928) said about Limnosipanea palustris: ‘The only North Amer- ican species, L. palustris (Seem.) Hook. f. was collected once in a swamp near Panama City, but has not been found again. It is a perennial herb with hirsute stems, ternate, sessile ovate-lanceolate leaves, and terminal cymes of small pink flowers." He may have drawn freely on Seemann's original description of Sipania palustris: "caule hirsuto basi repente, folis ternis sessilibus ovato-lanceolatis acutis glabris, floribus in dichotomia solitariis sessilibus, laciniis calycinis lineari- lanceolates acutis bi-aristatis, corolla glabra vel intus subpuberula, capsula subro- tunda, setoso-pilosa. In swamps, near Panama, growing together with Noterophila inundata Mart.” (Melastomataceae). Noterophila has been reduced to synonymy under Microlicia. On the other hand Standley may have seen the type sheet under discussion in the British Museum. In his diagnosis (repeated in North American Flora) he described the leaves as ''ovate-lanceolate" when in actuality they are oblong. The leaves of Microlicia are strikingly ovate-lanceolate, as is easily seen on the photograph of the sheet under discussion. Thus, it is possible that both Seemann and Standley used the Microlicia for parts of their diagnosis of Limnosipanea palustris, and they may have failed to 269 DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) Seis eg — OS —— FIGURE 57. Lindenia rivalis Benth. Habit (x*/;). [After Stern et al. 7.] 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 note that the sheet has been annotated with the name Noterophila (Melastoma- taceae). The enlargement on the photograph taken by Nowicke (MO) is of the Melastomataceae rather than the Rubiaceae element. 53. LINDENIA Lindenia Benth., Pl. Hartw. 84, 351. 1841. TYPE: L. rivalis Benth. Siphonia Benth., Pl. Hartw. 84. 1841, not Schreber, 1791, nomen nudum. Shrubs, glabrous or pubescent, the branchlets terete. Leaves short petiolate; stipules connate, short. Inflorescences terminal, cymose, the flowers few; bracts subulate. Flowers pedicellate, large, showy; hypanthium turbinate, 5-ribbed, the calyx with 5 elongate lobes, equal or unequal, persistent in fruit; corolla salver- form, elongate, the lobes 5, oblong, contorted in bud; stamens 5, inserted at the top of the corolla tube, the anthers sessile, linear, exserted; style slender, the stigmatic lobes 2, pilose, erect. Fruits clavate or pyriform, 2-celled, septicidally bivalvate, the valves splitting apically, often elastically coiled; seeds numerous, angulate. Lindenia is a genus of 2 species, one in the Old World and one in the New World. Literature: Darwin, S. P. 1976. The genus Lindenia (Rubiaceae). J. Arnold Arb. 57: 426- 1. Lindenia rivalis Benth., Pl. Hartw. 84. 1841. TYPE: México, Hartweg 581 (K, not seen; photo, MO).—Fia. 57. L. acutiflora Hook., Icon. Pl. 475. 1842. TYPE: México, Linden 358 (not seen). Subshrubs to 1.5 m tall, branching freely above, the leaves terminally aggre- gated, the branchlets stout, terete, smooth, drying black, often with an ericoid appearance, glabrous or minutely puberulent. Leaves salicoid or narrow elliptic, occasionally subfalcate, 4-12 cm long, 0.8-2.5 cm wide, acute at the apex and the base, occasionally vaguely acuminate, the costa subplane to prominulous above, prominulous beneath and rubescent, the lateral veins 7-8, strongly ar- cuate, reaching the margin or anastomosing to form a submarginal undulate vein, smaller intermediate veins soon evanescent, papyraceous to thinly coriaceous, glabrous above, glabrous or puberulent beneath especially on the costa and the veins, the margin often revolute; petioles absent or 0.2-1 cm long, the lamina slightly decurrent, dilated proximally. Flowers solitary, terminal, large, showy; hypanthium oblong, ca. 1 cm long, puberulent, the calycine cup scarcely mea- surable, the calycine lobes subulate, subequal, ca. 20 mm long, thinly carnose, puberulent with several rows of minute rubescent glands located basally within; corolla white, the tube slender, to 20 cm long, ca. 0.35 cm wide in middle, scarcely expanded at the apex and the base, pubescent outside, glabrous within, the lobes 5, elliptic oblong or rhombic oblong, 1.5-2.5 cm long, to 1.2 cm wide, subacute or obtuse at apex; stamens 5, the anthers linear ovate to oblong, ca. 1 cm long, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 271 FIGURE 58. Machaonia acuminata H. & B.—A. Habit (x'%).—A. Flower opened to show interior (x5). Fruit (x5). [After Tyson et al. 4273.] glabrescent with a few short hairs, the filaments attached near the mouth, the style slender, about as long as the tube, villose below middle, the stigmatic area ca. 1 cm long, barely exserted, divided above middle into 2 lobes, these oblong, ca. 0.2 em wide, obtuse. Fruits capsular, pyriform, to 4 cm long (including per- sistent calycine teeth), minutely pubescent, lignose, costate, drying dark, at first rupturing terminally at the ovarian disc position. 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Lindenia rivalis is common throughout Central America and Mexico; it has been erroneously reported from the island of Tobago. The only other species in the genus, L. vitiensis Seem. occurs in Fiji and New Caledonia. ‘‘Guabita de Quebrada.” L ZONE: Road K-6 E of Arraijan, Croat 15036 (MO). K-9 Road, Dwyer 3008 (MO). Pipeline Riad ^ " km from beginning of road, Mori & Kallunki 2043 (MO). Río Cocolí, Road K-9, Ster rn et .7 i z Tra waterfall on steep wooded slopes, Croat 34411 (MO). Río Maje 4-5 mi above waterfall near new Bayano Lake, 100 m, Croat 34710 (MO). Panamerican Highway, near Jenené, Río Canita, Duke 3842 (MO). Río Bayano, crossing Panamerican Highway, Duke 3984 (MO). Río Pasiga, to above waterfall on 2nd main fork, Gentry 2298 (MO). 54. MACHAONIA Machaonia H. & B., Pl. Aeq. 1: 101. 1806. TYPE: M. acuminata H. & B. Shrubs or trees, often armed. Inflorescences terminal, cymose paniculate, the flowers bracteate. Flowers with the calyx obovoid or turbinate, the calycine cup cupuliform, 4—5-lobed; corolla with the tube funnelform, hairy within, the lobes 4—5; anthers dorsifixed, included or exserted, the filaments short or long; ovarian disc fleshy, the style slender, the stigmas 2, the ovary usually 2-celled, the ovules solitary, pendulous. Fruits of 2 trigonous, indehiscent 1-seeded cocci separating from below upwards from the persistent axis; seeds elongated, cylindrical, the testa thin. Machaonia is found in Mexico, Central nr tropical South America, and the West Indies. It is a genus of about 30 specie Literature: Standley, P. 1934. 71. Machaonia H. & B. in North Am. Flora 32(3): 215-222. a. Leaves with 8-10 lateral veins; buds with the corolla 1.0-1.5 mm wide at the apex; anthers 0.3-0.5(—0.6) mm lon 1. M. acuminata aa. ar D with 6—7 lateral veins; buds with the corolla ca. 2.5 mm wide at the apex; anther a. 0.6 mm long namens 1. Machaonia acuminata H. & B., Pl. Aeq. 1: 101. 1806. TYPE: Ecuador, Hum- boldt & Bonpland, not seen.—Fic. 58 M. tysonii Dwyer, Ann. Missouri Bot. Gard., 53: 106. 1966. TvPE: Panama, Dwyer 5061 (MO, ho- lotype). Shrubs or small trees, occasionally armed with spines at nodes, to 1.2 cm long, lignose, occasionally with very small spinules, the branchlets terete, smooth, occasionally lenticellate, glabrous or puberulent, the nodes well spaced, frequently distichous. Leaves oblong, ovate oblong, ovate rotund, 4.5-10.0 cm long, 1.5-6.0 cm wide, deltoid to obtuse at the apex, often falcate cuspidate, the cusp usually short, the base acute, obtuse or rounded, occasionally slightly in- equilateral, the costa immersed or prominulous above, prominent beneath, the lateral veins 8-10, arcuate, immersed or prominulous above, prominulous be- neath, chartaceous, often stiff, usually concolorous, glabrate to villosulose above 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 273 and beneath, the intervenal areas reticulate, the veinlets immersed beneath; pet- ioles 0.5-2.0 cm long, slender, to 0.1 cm wide, puberulent, the hairs often dense only on the adaxial side; stipules free, small, compressed rotund, ovate oblong or triangular, 2.0-3.5 mm long, acute or constricted terminally into a subulate extension about as long as the body, pubescent. Inflorescences terminal, ex- ceeding the uppermost leaves, thrysoid paniculate, 4-15 cm long, 3—9(-15) cm wide, the peduncles 2-4 cm long, to 0.2 cm wide, pubescent, the lowermost 2- 4 pairs of branches usually opposite, usually strict and divergent, to 7 cm long, usually with 1 or 2 pairs of opposite secondary branches or with branches ter- nately disposed, the cymules terminal, densely flowered. Flower buds apically 1.0-1.5 mm wide, white, 4.2-5.0 mm long, the hypanthium turbinate or oblong, to 2 mm long, carnose, densely villosulose, the calycine cup scarcely measurable, glabrous and eglandular within, the lobes 5-6, erect, unequal, oblong, 0.6-1.0 mm long, villosulose outside and marginally; corolla tube compressed cylindrical, to 1.5 mm long, sparsely puberulent or occasionally glabrous outside, densely white villose within; stamens 5, the anthers oblong or subrotund, 0.3-0.6 mm long, dorsifixed, the filaments about as long as the anthers, attached near the apex of the tube; stigmas 2, erect, linear, 0.3-0.6 mm long, the style 1.0-3.5 mm long. Fruits turbinate, to 7 mm long, puberulent. This species extends from Mexico southward to Brazil. CANAL ZONE: Farfan Beach, Dwyer & Hayden 7533 (GH, K, MO, UC, US); Dwyer & Lallathin 8788 (MO). Coco Solo, Gentry 6055 (MO). cocLE: Río Hato Airstrip, Blum & Dwyer 2468 (MO). Ca. 14 mi W of Rio Hato, D’Arcy & D’Arcy 6277 (MO). Rio Hato Airstrip, Dwyer 5061 (MO). La Pintada, Escobar 11 (MO). í 1-1 5301 (MO); Woodson et al. 1205 (MO). COLON: Miguel de la Borda, Croat 9843 (MO). 2 S of Portobelo, Croat 11413 (MO). DARIEN: Puerto St. Dorothea, Dwyer 2293 (MO). HERRERA: " Chitre, Allen 1100 (MO). Ocü, Ebinger 1081 (MO); Lao 146 (MO). PANAMA: Bejuco, Woodson et al. 1680 (MO). SAN BLAS: Mulatuppu (Río Ibedi), Duke 8468 (MO). Ailigandí, Dwyer 6808 (MO). vERAGUAS: Santiago, Croat 10733 (MO); Dwyer 1270 (MO), 7429 (COL, MO, UC), 7553 (MO, UC). San Francisco, Río Los Chorros, Dwyer 1354 A (MO). Santiago, Stern et al. 993 (MO). Soná, Tyson et al. 4265 (MO). 5 mi E of Santiago, Tyson et al. 4273 (MO). 2. Machaonia martinicensis (DC.) Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 193. 19 Tertrea martinicensis DC., Prodr. 4. 481. 1830. TYPE: not see , seen. M. rotundata var. dodgei vere Field Mus. Nat. Hist., Bot. Ser. 18: 1326. 1938. TYPE: Costa Rica, Dodge 7752 (F, holotype). Shrubs, vines, or trees, to 10 m tall, the branchlets occasionally armed at the nodes with spines to 1.3 cm long, lignose, the twiglets terete, smooth, glabrate, the nodes 2-5 cm apart. Leaves opposite or rarely alternate, widely oblong, occasionally elliptic, 4-11 cm long, 1.8-4.5 cm wide, acute or rounded at the apex, usually contracted into a short acumen 0.3-1.0 cm long, basally widely cuneate to subobtuse, the costa slender, plane or prominulous above, prominu- lous to prominent beneath, the lateral veins 6—7, arcuate, the intervenal areas minutely mosaiclike, chartaceous, usually concolorous, glabrous above and be- neath except often minutely barbate in the axils; petioles to 1.2 cm long, 0.5-1.2 mm wide, glabrous or puberulent on the adaxial side; stipules minute, 2.5—3.0 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 i? FiGuRE 59. Macrocnemum glabrescens (Benth.) Wedd.—A. Habit (x'2).—B. Flower opened to show interior (x2). [After Busey 404. | mm long, the body compressed rotund to triangular, the subulate process longer than or shorter than the body. /nflorescences terminal, thrysoid paniculate, to 11 cm long, the branches plane or angular, pubescent, the lowermost 2-4 pairs of branches usually opposite, strict and divergent, mostly with 1 or 2 pairs of op- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 275 posite secondary branches; bracts usually linear, 5 mm long, sometimes larger and leaflike, the cymules terminal, densely flowered. Flowers 5.5-6.5 mm long, the buds usually 2.5 mm wide at the apex; hypanthium turbinate, the hypanthium and calyx ca. 2.5 mm long, glabrate, the hairs small and somewhat webby or larger and subulate, to 0.2 mm long, white, drying red, the calycine lobes 5, erect, unequal, to 1.2 mm long, obtuse, villosulose and often sparsely pubescent on the margins; corolla white or pale green, the tube turgid, compressed cylindrical, to 2.5 mm long, sparsely villose outside, densely villose at the mouth within: sta- mens 5, the anthers oblong, ca. 0.6 mm long, the stigmas 2, linear, to 1.2 mm long, the style ca. 3.5 mm long, frequently exserted. Fruits turbinate, to 6 mm long, glabrate. Machaonia martinicensis is found in Jamaica, Panama, Costa Rica, and Co- lombia. Duke 9719 notes that ''the flowers are very attractive to ants and bees." BOCAS DEL TORO: Macaw Hills, Wedel 548 (MO). Nances Cay Island, Wedel 585, 2862 (both MO). Water Valley, Wedel 834, 1751, 1788 (all MO). Fish Creek Hills, Wedel 2446 (MO). PANAMÁ: Coco Solo Weather Station, Duke 4287 (MO). COLOMBIA: ANTIOQUIA: Punta Las Barcas Turbo, sea level, Duke 9718 (MO) 55. MACROCNEMUM Macrocnemum P. Br., Hist. Jam. 165. 1756. TYPE: M. jamaicense L. Trees or shrubs, the branchlets terete. Leaves opposite, petiolate; stipules oblong or obovate. Inflorescences terminal or axillary, paniculate. Flowers with the calycine cup cup-shaped, the teeth 5, persistent; corolla funnelform or sal- verform, the lobes 5, in bud drawn in so as to give a maltese cross pattern in top view, spreading at anthesis, wider than long, pubescent within, valvate or redu- plicate; anthers oblong, included, the filaments partly villose; ovary 2-celled, the ovules numerous. Fruits capsular, bisulcate, dehiscing loculicidally into 2 valves; seeds winge Macrocnemum ranges from Central America to Colombia, and it also occurs in the West Indies. It is a genus of about 20 species, easily recognized by the configuration of the corolla lobes in the bud stage. a. Corolla tube glabrous outside; anthers ca. 1 mm long... l. M. glabrescens aa. Corolla tube golden puberulent outside; anthers ca. 0. 7mmlong |... . . 2. M. pastoense 1. Macrocnemum glabrescens (Benth.) Wedd., Ann. Sci. Nat. (Paris) 4. 1: 76. 1854.—Fic. 59. Lasionema glabrescens Benth., Bot. Voy. Sulphur 105. 1844. TYPE: not indicated. Trees to 8 m tall, the branchlets terete, ultimately angular, glabrous, the nodes usually well spaced. Leaves obovate oblong, spathulate, oblong, often vaguely falcate, widely deltoid to obtuse at the apex, acuminate or cuspidate at the apex, the cusps to 1 cm long, basally obtuse, acute or rounded, often inequilateral, the costa plane or immersed above, prominulous beneath, the lateral veins 7-9, ar- cuate, chartaceous to papyraceous, usually discolorous, glabrescent above and beneath, occasionally glabrous, often barbate on the costa and the veins beneath; 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 petioles 0.5-1.5 cm long, usually stiff; stipules deciduous, foliaceous, oblong or ovate oblong, 1.0-1.5 cm long, obtuse. Inflorescences terminal or axillary, thry- soid paniculate, to 30 cm long, either sessile and with usually 3 branches arising from the stem tip, or pedunculate and the peduncles to 11 cm long, glabrescent, the branches few, well spaced, usually opposite or ternately disposed, the low- ermost to 14 cm long, arcuate or strict, the cymules terminal, few flowered, spreading; bracteoles subulate, divergent. Flowers sessile or subsessile, dark- ening at maturity, the buds clavate capitate; hypanthium oblong, ca. 3.2 mm long, ca. 2.2 mm wide, villosulose, the calycine tube somewhat inflated, 0.5 mm long, petaloid, glabrous inside and out, venose, the calycine teeth 5, triangular, to 1 mm long, minutely puberulent on the margin; corolla pink, the tube cylindrical, 5 mm long or longer, ca. 2 mm wide, stiffly petaloid, glabrous outside and in except usually villose at filament attachment point, the lobes 5, oblong rotund, .5 mm long, glabrous, venose; stamens 5, the anthers oblong rotund, ca. 1 mm long, dorsifixed with a glandular patch at point of filament attachment, the filaments unequal, slender, subulate, 2.5—4.0 mm long, densely villose below the middle, attached at about middle of the tube and continuing downwards in riblike fashion; stigmas 2, oblong, ca. 0.5 mm long, convex in cross section, the style thickly subulate, 6-8 mm long, ca. 0.25 mm wide in middle. Fruits elliptic or fusiform, to 2.5 cm long, to 0.35 cm wide before opening, on opening the valves united at the apex and the base, the interior of valves smooth, yellow, the outside delicately ribbed, pubescent, glabrous, capped by a saucerlike calycine cup and ovarian disc, both to 1 mm long; seeds membranaceous, fusiform, to 3 mm long, often falcate, alate, the surface with crowded irregular lines. Macrocnemum glabrescens is known from Costa Rica, Panama, and Colom- bia. Duke (8041) indicates the fruits are eaten by loro. Busey and Mahler (351) note that the tree was covered with ants. ‘‘Palo cuadrado"; “Palo blanco’’; ‘*Madrono’’; ‘Guanabana Torete”; ‘‘Cedro Macho’’; ‘Cedro Chumaquino."' In the type photo collection at the Missouri Botanical Garden there is a collection of Wagner from Panama deposited in the Herbarium Regium Monacense (Vien- na). It is definitely M. glabrescens, but it is not the type. CAS DEL TORO: Daytonia Farm, Cooper 448 (MO). Between Buena Vista and Cerro Pilón, Kirkbride & Duke 680 (MO). CANAL ZONE: Chiva-Chiva Trail, Allen 2467 (F). Río Puente, between tunnel and natural bridge, Dodge & Allen 17317 (MO). Chiva-Chiva Trail, Fosberg 27392 (F). Summit, Harvey 5227 (MO). Mindi Hills, Johnston 1734 (MO). Near Gamboa, Río Mandinga, Kennedy et al. 2382 (MO). Without other locality, Lindsay 455 (MO). Gorgona, A 4741 (F, MO). Río Masambi Grande, 2 km N of Summit Garden, 80-100 m, Nee 9417 (MO). 1 mi N of Summit, Tyson et al. 2756 (MO). BARRO COLORADO ISLAND: PUR 50 (MO), 273 (F); HU & Bailey 233 (F, MO), 234 (F), 244 (MO); Bangham 172, 372 (both MO); Brown 26, 76, 142, 193 (all F); Chapman s.n. (F); Croat 4578, 4806, 4944, 4955, 5806, 5933, 7190, 7848, 8699, 11256A, 13169 (all MO); D'Arcy 3929 (MO); Dwyer et al. 8459 (MO); Ebinger 293 (MO); Foster 694, 698 (both F); Frost 26, 135 (both F): Shattuck 423, 659 (both F, MO), 8/1 (MO); Wetmore & Abbe 218 (F); Wilson 1 (F); Woodworth & Vestal 333 (F, MO). cuiRiQUÍÉ: Corotü, 6 km SW of airport, Puerto Armuelles, Busey 4 )9knS x SW of Puerto Armuelles, Busey 728 (MO). 12 mi W of Puerto Armuelles, Croat 22155 (MO). 2 mi SW of Puerto Armuelles, Croat 22454 (MO). CocLE: El Valle, Allen 114 (F, MO), dii d Mur MO); Hunter & Allen 302 (MO). COLON: Gatuncillo, Correa et al. 734 (MO). DARIEN: Rio , Bristan 56 (MO). Río Aruza, d 1346 (MO). Pidiaque, Duke 8041, 8042 (both MO). Río E Duke 8249 (MO). Río Sabana m above Santa Fé, Duke 10203 (MO). Between upper Río Membrillo and Ca on construction pv to San see Duke 10873 (MO). Manené near Cuasi, Kirkbride & Bristan 1482 (MO). La Palma, Pittier 6 (F). PANAMA: Upper Rio Pequeni between Salamanca Hydrographic Station and Rio Bn te ^n 17276 (MO). Las Cumbres, Duke 11825 (MO). Isla 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 277 Pedro Gonzales, Dwyer 1716 (MO). NW slope of Red Hill, San José Island, Johnston 1309, 1366 (both MO). Alhajuela, Pittier 2397 VA cra Santa Fé, Allen 4378 (MO). Santiago, Dwyer 1351 (MO). Isla de Coiba, Dwyer 2374 (M 2. Macrocnemum pastoense Karst., Linn. 30: 150. 1859. TYPE: Colombia, Karst., not seen. Trees to 8 m tall or shrubs, the branchlets terete, ultimately angular, smooth, minutely puberulent. Leaves oblong, oblong rotund or obovate oblong, 6-17 cm long, 4-10 cm wide, widely deltoid to rounded at apex, acuminate, the acumen short, widely deltoid, to 0.5 cm long, ultimately obtuse or acute, the base rounded or obtuse, the blade occasionally contracted at proximal 1⁄4, the costa plane or prominulous above, prominent beneath, the intervenal areas spreading reticulate, papyraceous, tending to dry dark at least on the upper side, glabrous above, puberulent beneath, barbate in the axils; petioles to 3 cm long, stiff, puberulent; stipules persistent only below inflorescence, oblong, oblong rotund to ovate ob- long, 3 cm long, obtuse or acute, rimose, puberulent. Inflorescences solitary, terminal, occasionally terminal and axillary, cymose paniculate, to 22 cm long, to 10 cm wide, the peduncle to 12 cm long, terminated by a pair of reduced leaves, the lowermost branches opposite, to 7 cm long, divergent or arcuate ascending, the branches divided into 2 branchlets. Flowers sessile, the hypan- thium densely golden puberulent, the calycine cup short, the 5 triangular teeth ca. | mm long, longer than the cup, golden puberulent; corolla purple or lilac, the tube narrow cylindrical 6-8 mm long, puberulent outside, glabrous within, thinly carnose, the lobes 5, in bud disposed as a hemispherical capitate mass to 0.3 cm long, 0.5 cm wide, when expanded the lobes hemispherical, wider than long, puberulent, the margin undulate; stamens 5, the anthers oblong, ca. 0.7 mm long, curved when mature, subversatile, the filaments slender, to 6 mm long, villose below the middle, attached below the middle of the tube; style slender, ca. 7 mm long, glabrous, the stigmas 2, when separated scarcely wider than the style, 0.3 mm long. Fruits not seen. Macrocnemum pastoense is known only from Panama and Colombia. DARIEN: Pinas, Duke 10610 (M(). Rio Tuira, near mouth of Rio Paya, Gentry 4426 (MO). Yaviza along Rio Chucunaque at El Punteadero, Stern et al. 173 (MO). COLOMBIA: CHOCO: Rio Mutata tributary Río El Valle, between base of Alto de Buey and mouth of river, 100-150 m, Gentry & Fallen 17459 (MO). 56. MANETTIA Manettia L., Mant. Pl. 553. 1771. Nomen conserv. TYPE: M. reclinata L. Lygistum P. Br., Civ. Nat. Hist. Jam. 142. 1756. Nomen rejic. TYPE: Petesia lygistum L. Nacibea Aubl. Hist. Pl. Guiane 1: 95. 1775. TYPE: no type indicated. Conotrichia A. Rich., Mém. Soc. Hist. Nat. (Paris) 5: 277. 1829; Richard, Mem. Fam. Rub. 830. TvPE: C. alba (Aubl.) A. Rich Herbs or vines, sometimes woody. Leaves petiolate, the stipules small, per- sistent. /nflorescences axillary, cymose, occasionally terminal, the flowers oc- casionally solitary. Flowers variable in color, the calycine lobes usually 4, short or long, usually with teeth or small glands between the lobes; corolla tubular or 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 funnel shaped, 4-lobed; anthers exserted or included; ovary 2-celled, the ovules numerous. Fruits capsular, obovoid or turbinate, septicidally bivalvate, splitting from the apex; seeds compressed, winged. Manettia occurs sporadically in Central America, abundantly in tropical South America, and the West Indies. It is a genus of about 130 species. Literature: Wernham, H. F. 1918. The genus Manettia. J. Bot. 57 (Suppl.): 1-44. a. Herbs, upright or scarcely upright. b. Plants terrestrial; leaves sessile, the blades linear ovate or linear elliptic; flowers M pedicels to 6 mm long l. boc dcc bb. Plants aquatic; TAA with petioles ca. 1 cm long, the blades ovate or ux ova flowers sessile . M. droit aa. Vines. c. Calyceine lobes (6—)8; flowers red or magenta rarely white __________________ 4. M. reclinata cc. Calycine lobes 4; flowers white 2. M. flexilis 1. Manettia bocataurensis Dwyer.°8 TYPE: Panama, Croat & Porter 16334 (MO, holotype). Herbs, the stems weak, scarcely erect, smooth, glabrous, the nodes well spaced. Leaves sessile, narrowly ovate or narrowly elliptic, 2.5-7.0 cm long, 0.3- 1.0 cm wide, acute at the apex and at the base, the costa plane above, subplane beneath, the lateral veins ca. 5, strongly arcuate, the veins of the intervenal areas not visible, thin membranous, glabrous except the minutely pubescent, slightly revolute margins, concolorous, shiny, with minute white cystoliths beneath; stip- ules compressed hemispherical, 2.0-3.7 mm long, the awns 3, subulate, equal or unequal, to 2.5 mm long. Flowers solitary, axillary; pedicels slender, to 6 mm long, puberulent; hypanthium subrotund, 1.5 mm long, glabrescent, the calycine lobes subulate, to 1.8 mm long, puberulent; corolla white, the tube cylindrical, to 1.2 mm long, glabrous, the lobes short. Fruits rotund, ca. 3.5 mm in diam., black when dry, glabrate, the calyx persistent. Manettia bocataurensis is known only from Panama. It is related to M. steno- phylla Donnell Smith, from Costa Rica, the type of which is Tonduz 12969 (F). The latter species has some of the flowers short-pedicellate, in cymes, and to 8 mm in length. BOCAS DEL TORO: Railroad near station at Milla 10, Croat & Porter 16334 (MO). 68 Manettia bocataurensis Dwyer, spec. nov. Planta vix erecta caulibus glabris. Folia sessilia lineari-ovata vel lineari-elliptica, 2.5-7.0 cm longa, 0.3-1.0 cm lata, venis lateralibus ca. 5 forte arcuatis, tenui-membranacea glabra praeter margines D pura pubescentes; stipulis compresso- mm longis; corolla alba tubo cylindrico, ad 1.2 mm longo, glabro, lobis brevibus. Fructus rotundi, ca. 3.5 mm diam., glabrescentes. (x 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 279 E 60. Manettia flexilis Brandegee.—Habit (x!⁄4).—B. Flower opened to show interior 5). — p nt cross section (x712). [After D'Arcy & D'Arcy 6616.] 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 2. Manettia flexilis Brandegee, Univ. Calif. Publ. Bot. 6: 196. 1915. TYPE: México, Purpus 7118 (UC, holotype).—Fic. 60. M. estrellae Standley, J. Wash. Acad. Sci. 15: 6. 1925. TYPE: Costa Rica, Standley 39224 (US, holotype). Vines, the stems flexuous, the hairs short, curled, white. Leaves oblong, ovate oblong or trapeziform, to 5(-7?) cm long, 1.5 cm wide or wider, acute or long attenuate (Standley) at the apex, acute to obtuse at the base, the costa subim- mersed above, prominulous to plane beneath, the lateral veins 7-8, arcuate, mem- branous, glabrous to villosulose above, short pilose or villosulose beneath; pet- ioles slender, to 1 cm long, to 0.5 mm wide, puberulent; stipules not seen. Inflorescences axillary, umbelliform, the peduncle reduced, the flowers conglom- erate. Flowers on short pedicels to 0.5 cm long; hypanthium to 0.5 mm long, densely villose, the calycine cup to 0.4 mm long, the calycine lobes 4, lanceolate, ca. 3 mm long, ca. 0.8 mm wide, venose, the hairs dense and curled outside, at least 1 erect oblong gland to 0.35 mm long at a sinus; corolla white, the tube (in bud) short, villose outside, glabrous inside, the lobes 4, to 3 mm long, ca. 0.7 mm wide, densely villose abaxially; stamens 4, the anthers ca. 0.7 mm long, the filaments short. Fruits turbinate, 5-7 mm long, sparsely villose. Manettia flexilis is known from Guatemala, Costa Rica, Panama and Colom- bia. CHIRIQUÍ: 5.4 km from Hato de Volcán on road to Las Lagunas, Correa & Lazor 1481 (MO). W of Cerro Pando (Río Chiriquí Viejo), D'Arcy & D'Arcy 6616 (MO). Bajo Mono, Boqueté, Davidson 468 (MO). 3. Manettia hydrophila Dwyer.5*? TYPE: Panama, Lewis et al. 2822 (MO). Herbs, often prostrate and procumbent, the stems spreading or erect and weak, the hairs dense, weak, wrinkled, the nodes well spaced. Leaves ovate or rhomboid ovate, 2-5 cm long, 1.2-2.8 cm wide, acute toward the apex, basally attenuate acute or subobtuse, often inequilateral, the costa subplane above, plane beneath or scarcely immersed, the lateral veins ca. 6, arcuate, prominulous above, sulcate beneath, membranous, shiny, with minute white cystoliths on both sides, the costa and veins often with appressed uncinate hairs, the margins pu- bescent; petioles ca. 1 cm long, puberulent; stipules short, widely triangular, ca. | mm long, black when dry. Inflorescences terminal or axillary and opposite, the peduncle to 1.5 cm long, the cymes contracted, the flowers alternate and well spaced on a flexuous rachis or on 2 branches terminating the peduncle, the branches to 4 cm long. Flowers sessile; hypanthium compressed roiund, ca. 1 69 Manettia hydrophila Dwyer, spec. nov. Herba aquatica nodis rhizophoris pie uda ciliis up crispis. Folia ovata vel rhomboideo-ovata, 2-5 cm longa, un cm lata, venis lateralibus arcuatis, membranacea cystolithis multis albis utrinque costa venisque pe saep indus ressis pas praeditis; petiolis ca. 1 cm longis; stipulis brevibus lato- triangularibus, c ps mm longis. /nflorescentiae terminales vel PAPE pedunculo ad 1.5 cm longo, cymis contractis vel dioe: pois nbn distantibus, ad 4 cm longa. Flores sessiles lobis calycis 5 ovato- e eae tis m lon ca. 0.4 mm latis; Cael rosea, tubo ad 5 mm longo lobis 4-5, ca. 5 mm longis; aha ad l. pa longis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 281 mm long, the hairs appressed, white, the calycine lobes 5, ascending, narrowly linear lanceolate, to 2.8 mm long, 0.4 mm wide, puberulent on both sides with a distinct, red, rotund gland, 0.05 mm long, in the sinuses; corolla rose colored, the tube cylindrical, to 5 mm long, 3 mm wide, glabrous, the lobes 4-5, ovate oblong, 5 mm long, 3 mm wide, obtuse; stamens 4, the anthers oblong, 1.3-1.5 mm long, obtuse at the apex and base, the filaments ca. 1 mm long, attached ca. 1.5 mm from the base, the ovarian disc ringlike, 0.1 mm long, the style much reduced, the stigmas lanceolate, acute. Fruits rotund, 3.5 mm in diam., the hairs ascending, uncinate, to 0.5 mm long, drying at first brown, ultimately the surface of the wall deciduous, the remainder scarious, white, smooth; seeds subrotund or oblong, ca. 0.5 mm long, flat, red, shiny. Manettia hydrophila is known only from Panama. It is unique in being a prostrate or erect aquatic. VERAGUAS: Mouth of Rio Concepcion, in swamp, Lewis et al. 2822 (MO). 4. Manettia reclinata L. Mant. Pl. 2: 553. 1771. TYPE: not seen. Vines, the stems slender, often with longitudinal ridges, glabrate or puberu- lent. Leaves ovate oblong, 3.0-5.5 cm long, 1.0-2.5 cm wide, acute at the apex, often acuminate, the acumen to | cm long, basally cuneate to obtuse, slightly inequilateral, the costa plane to prominulous above, prominulous to subplane beneath, the lateral veins 6—7, arcuate, the margin often villosulose, membranous; petioles to 2 cm long, slender, ca. 0.7-1.0 mm wide, puberulent; stipules small, widely triangular, to 1 mm long, wider than long, minutely puberulent and ru- gulose. Inflorescences axillary or terminal, glabrous, I-several flowers per axil, the peduncle, if present, short, terminated by a pair of bracteoles subtending the pedicels; pedicels capillaceous, to 3.5 cm long. Flowers with the hypanthium oblong, ca. 3 mm long, the calycine cup ca. 1 mm long, with minute oblong glands within to 0.15 mm long, the lobes (6—)8, subequal, subulate to narrowly oblong, 6-7 mm long, marginally ciliate; corolla red, scarlet, magenta, occasionally white, the tube ca. 7 mm long, basally turgid, glabrate, minutely puberulent to densely villose outside, densely villose within near the mouth and with a few hairs near the base, the lobes 4, oblong, 7-8 mm long, 3-4 mm wide, obtuse, petaloid, venose, marginally ciliate, glabrous adaxially; stamens 4, the anthers oblong, ca. 3 mm long, the filaments ca. 1 mm long, attached near the mouth; stigmas 2, linear oblong, ca. 2.5 mm long, the style ca. 11 mm long. Fruits capsular, com- pressed rotund, 0.6-0.8 cm in diam., smooth, glabrous or puberulent, each valve with 3 delicate ribs, the calycine lobes persistent, curled; seeds ca. 3 mm in diam., the wing round, the margin undulate. Manettia reclinata ranges from Mexico to northern South America. It also occurs in Cuba and Jamaica. Duke 14934 notes that the Indians chew the fruits which stain their teeth black. *‘Churco-guidave’’ (Choco). Sullivan 509 and Fol- som 3691 have an average of 6 calyx lobes and the flowers are described as "white." They may represent a new species. AS DEL TORO: Changuinola Valley, Dunlap 254 (MO). Chiriquí Lagoon, Wedel 1217, d 1669, 1714 (all MO). CANAL ZONE: Between Gatün and Fort Sherman, Burch et al. 1017 (F, GH, 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 MO, NY, UC, US). Farfan Beach, Burch et al. 1407 (F, GH, K, MO, NY, UO). Río Indio Hydro- graphic Station and Natural Bridge, Dodge & Allen 17485 (MO). Madden Dam, Dwyer 8389 (MO). M boa, Nee 8992 (MO). | km NW of Gamboa on E side of Ammunition Depot, Nee 9542 (MO). opal Rodman Marine Base and Chorrera, Nowicke et al. 3591 (MO, UC, VEN). BARRO COLORADO IS Croat 4375, 6705, 7467, 12740 (all MO); Shattuck 159 (F, MO), 486 (MO). 13 mi N of Rio San Félix bridge, 800-1200 m, Croat 33406 (MO). CHIRIQUI: Frances Arriba School, ca. 14 km N of David, Lewis et al. 684 (MO). 12 mi W of Puerto Armuelles, Liesner 205 (MO). CocLE: La Mesa, ge 2345, 2853 (both F); Croat 37483 (MO): oi io 3891 (MO); Sullivan 509 (MO). DARIÉN: Ridge of Yaviza, Duke 6538 (MO). Río Mortí Hydro Camp, Duke 15414 (MO). Río Balsa, Duke & Nic ud 14934 (MO). Los SANTOS: Headwaters, Rio Pedregal, Lewis et a es (MO, UC, VEN). Between Tonosi and Macaracas, Oliver et al. 3545 (MO). PANAMA: Between Capira and Potrero, Dodge & Hunter 8616 (MO). Cerro Campana, Dressler 3524 (MO). Cerro Jefe. Duke 8005 (MO). Cerro Cam- pana, Lewis et al. 1922 (MO), 3313 (MO, UC, VEN); Kirkbride & Hayden 282 (MO). Between isse and Río Yguana, McBride 2634 (F). El Llano-Cartí Road, 5 km N of Panamerican Highway, . 300 m, Nee 7880 (MO). Bella Vista, und 5338 (F). Cerro a Tyson 3423 (MO). SAN BLAS: Opposite Ag deed Lewis et al. 145 (F, GH, K, MO, NY, PA, UC, US). Ustupo, D'Arcy 9491 (MO). AGUAS: Soná, Allen 1024 (F). 0.2 mi pm fork in road at Escuela Alto Piedra, road to Río Calovabors. 750 m, Croat & Folsom 33899 (MO). Mouth of Rio Concepción, Lewis et al. 2841 A (MO). 57. MITRACARPUS Mitracarpus Zucc. ex Schultes in R. & S., Syst. Veg. 3. Mant. Pl. 210. 1827. TYPE: Mitracarpus scaber Zucc. Herbs, annual or perennial. Leaves usually narrow, often subsessile; petioles united with the stipules to form a setiferous sheath. /nflorescences of terminal or axillary heads. Flowers minute, 4-merous, the calycine cup short, the lobes unequal; corolla funnel shaped, the lobes 4(-3), valvate; stamens included or somewhat exserted, the anthers dorsifixed. Fruits capsular, 2-celled, each cell l-seeded, transversely or obliquely circumscissile, the calyx persistent; seeds ellipsoid or ovoid, ventrally 4-lobed and with a cruciform groove. Mitracarpus occurs in Mexico, Central America, South America, the West Indies, and in Africa. It is a genus of about 40 species. a. Stems with numerous hairs; leaves hairy above; nutlets with a single set of diagonally crossed grooves on ventral face M. hirtus aa. Stems with few hairs; leaves glabrous or ‘adi above; nutlets with a double 2 of diagonally crossed grooves on ventral face -2-2-2222 . M. breviflorus — . Mitracarpus breviflorus A. Gray, Pl. Wright. 2: 68. 1853. Originally ‘‘Mi- tracarpium breviflorum.” TYPE: México, Wright 1120, not seen. Herbs to 1.5 dm, branched, the stems slender, angular, sparsely hairy. Leaves sessile, usually opposite, occasionally 4-several per node, lanceolate, ovate lan- ceolate, to 2 cm long, to 0.4 mm wide, the lateral veins few, vague, the blades stiff, scabrid above, otherwise glabrous, glabrous beneath, the margin revolute. Inflorescences with heads to 1 cm wide. Flowers white. Fruits to 3 mm long, glabrous below the line of dehiscence, pubescent above, the horns to 2 mm long, unequal, pubescent, the seeds oblong, ca. 0.5 mm long, with 2 diagonally crossed grooves on the ventral surface. Mitracarpus breviflorus is known from Arizona, Mexico, Costa Rica, and 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) E UÜ ^ Í k ; er Gi z ) Y z "n A ge a im 32^ Ce RC ENT AE F . Mitracarpus hirtus (L.) DC.—A. Habit ( = =: EP. £2 4 LEA LEP y l Cy << ar pie ^ s c Ë g LA $0, ^ > E E h u EN yA L Ç 1 LM ay. y fj " (rg 25. 3 y KAN I 7, Z N | Y. j | 4634 FIGURE 72. Psychotria hammelii Dwyer.—A. Habit (x1!2).—B. Fruit (x312). [After Hammel 40. Psychotria haematocarpa Standley, J. Wash. Acad. Sci. 18: 274. 1928. TYPE: Costa Rica, Standley & Valerio 47194 (US, holotype). Shrubs to 2 m tall, branching freely, the branchlets terete, ultimately angular, glabrous. Leaves narrowly or widely lanceolate or oblong lanceolate, sometimes falcate, 5-15 cm long, 1-5 cm wide, acute at the apex or obtuse, often acuminate, the acumen to 2 cm long, basally acute or attenuate acute, the costa slender 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 383 above, prominulous beneath, the lateral veins 10-13, prominulous beneath, white when dry, arcuate, with 1-3 veins intermediate between a pair of lateral veins, these soon branching, the submarginal vein undulate, 0.2-0.7 cm from the margin, stiffly chartaceous, concolorous, glabrous; petioles 0.3-1.6 cm long, rigid, 0.1 cm wide, glabrous; stipules free, widely compressed rotund, the body to 2.5 mm long, coriaceous when dry, glabrous, the awns 2, subulate, longer than the body. Inflorescences terminal, 1.5 cm long, the peduncle short, to 1 cm long, ca. 0.5 mm wide, glabrous, the branches (0—)1—3, to 1.5 mm long, radially disposed, each branch (when fruits deciduous) terminated by a few scars; bracteoles deciduous from the base of the branches, few, subulate, ca. 1 mm long. Flowers not seen. Fruits sessile, orange when dry, rotund, 4—6(—8) mm in diam., each pyrene ul- timately 4-sulcate, smooth, glabrous, the calycine cup often persistent, 0.8—1.5 mm long, the teeth triangular, ca. 0.5 mm long. Psychotria haematocarpa is known from Costa Rica, Panama, and Colombia. The type from Costa Rica was collected at 900 m. The species is recognized by a combination of characters: the leaf blades with the lateral veins usually white when dry; the submarginal vein well set in from the margin; the inflorescence much reduced, and fruit drying orange. Several collections from Nicaragua may be Psychotria haematocarpa. These collections are labeled P. pervilis Standley & Williams, an unpublished name: Shank & Molina 4675, 4780 (both US), 4887, 4960 (both F). The first two of these have inflorescences with elongate reflexed bracts to 1.2 cm long, suggestive of P. officinalis. COCLÉ: Cerro Pilon, ca. 2700 ft, Duke & Lallathin 15000 (MO); Dwyer 8331 (MO); Gentry 3640 (MO). 3 mi NE of El Valle, Mori & Kallunki 2962 (MO). COLON: Rio Guanche, D'Arcy 9687 (MO). Santa Rita Ridge trail, 17-35 km from Boyd-Roosevelt Highway, 400-800 m, Mori & Crosby 6319 (MO). Santa Rita Road, ca. 6 km from Boyd-Roosevelt Highway, Mori & Kallunki 2154 (MO). DARIEN: Loma Cuasi, behind Manene, Duke 13612 (MO). 1 mi SE of Rio Tuira, Duke 14575 (MO). Between Río Diabolo and Río Acuati, Duke 14882 (MO). Puerto St. Dorothea, Dwyer 2267 (MO). VERAGUAS: Between Escuela Agricola Alto Piedra and Calovebora, 15.6 km NW of Santa Fe, 450- 550 m, Croat 27568 (MO). 8.8 km from Agricola Alto de Piedra, Mori & Kallunki 3212 (MO). co- LOMBIA: CHOCO: Municipio de Riosucio, Parque Natural Nal. Los Katyos, Subida al Alto Del Limon, 300—380 m, Leon 435 (MO). 4]. Psychotria hammelii Dwyer.?? TYPE: Panama, Hammel 4634 (MO, holotype; isotype PMA).—Fic. 72. Subshrubs, the ultimate branchlets terete, smooth, glabrescent, moderately slender, dark red when dry, the nodes well spaced, subturgid. Leaves narrowly elliptic to obovate elliptic, often scarcely falcate, 8.5-12.0 cm long, 2.5—4.0 cm wide, deltoid toward the apex, acuminate, the acumen to 2 cm long, 0.5-0.8 cm wide, cuneate or acute at the base, the costa plane or immersed above, promi- nulous beneath, the lateral veins ca. 15, immersed above, widely arcuate and 39 Psychotria hammelii Dwyer, spec. nov. Suffrutices. Folia angusto-elliptica vel obovato- -ellip- tica, 8.5—12.0 cm longa, 2.5-4.0 cm lata, acumine ad 2 cm longo, costa supra plana vel immersa venis lateralibus ca. 15 areis inter venas supra bullatis laminis rigido-membranaceis supra poene Sid puberulis; petiolis gracilibus; stipulis liberis oblongis, ad 2 cm longis, processibus duobus, longis. /nflorescentiae cymoso-paniculatae ad 12 cm longae, ad 4 cm latae, pedunculo ad 5 mm d Fructus sessiles oblongo-rotundi, ad 6 mm longi, ad 5 mm lati, glabri in sicco rubri. 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 forming an undulate submarginal vein, the intervenal areas patulous reticulate, distinctly bullate above, rigidly membranous, discolorous, dark red and glabrous above, puberulous beneath; petioles slender, stipules free, persistent only at the apex of twigs, oblong, to 2 cm long, with 2 slender erect processes 3 or more mm long, the body petaloid, apparently split lengthwise near the base or middle at maturity; venose, glabrous, red when dry; raphides numerous, minute. /nflores- cences cymose paniculate, to 12 cm long, to 4 cm wide, the peduncle slender, to 5 cm long, to 0.15 cm wide, delicately winged, usually terminated by 2-3 branch- es, 1.0-1.6 cm long, unequal, spreading, each with 2-3 secondary branches, the cymes 2-3-flowered, bracteoles narrowly oblong, to 4 mm long. Flowers not seen except for the persistent calyx. Fruits sessile, oblong rotund, to 6 mm long, to 5 mm wide, obtusely costate, glabrous, drying red, the persistent calyx cup 0.7 mm long, the lobes triangular oblong, to 1 mm long, glabrous. Psychotria hammelii is known only from Panama. It is recognized by the bullate character of the leaves. The stipules are diagnostic, being large, drying bright red, and having 2 erect antenna-like awns. Whether the stipules are nat- urally torn lengthwise as seen in the type material demands investigation. The collectors of the type note: **mature fruits red; leaves bullate, metallic dark above; leaf often similar to that of Urticaceae; 2 seeds, smooth on the bottom with a line; stoloniferous."' OCAS DEL TORO: E slope of La Zorra, forest to divide, mossy cloud forest, Kirkbride & Duke 812 (MO). vERAGUAS: W of Alto de Piedra W of Santa Fé, 3200-5600 ft, Hammel 4634 (MO, PMA). 42. Psychotria horizontalis Swartz, Nov. Gen. Sp. Pl. Prodr. 44, 1788. TYPE: Hispaniola, Swartz, not seen. ee H.B.K., Nov. Gen. Sp. Pl. 3: 358. 1819. TYPE: Venezuela, Humboldt & Bonpland, seen P. s a Riley, Kew Bull. 1927: 124. 1927. TYPE: Panama, ies Ps S holotype). P. l var. glaucescens (H.B.K.) Steyermark, Mem t. Garden 23: 472. 1972. Shrubs to 5 m tall, usually branching freely, the branchlets terete, slender, smooth, glabrous, the terminal nodes usually proximate. Leaves oblong, occa- sionally elliptic or obovate elliptic, 6-16 cm long, 2-8 cm wide, deltoid or sub- obtuse at the apex, rarely acute, acuminate, the acumen to 1.5 cm long, often falcate, ultimately obtuse, basally cuneate, obtuse or attenuate acute, occasion- ally cordate, rarely truncate, the costa prominulous above, often distinctly grooved, prominent beneath and usually with an evident pore in the axils of the secondary veins, the lateral veins 5-10, arcuate and forming an undulate, sub- marginal vein to 1 cm from the margin, stiffly chartaceous, concolorous, glabrous above, glabrous to puberulent beneath; petioles absent or to 0.5(—0.9) cm long, slender, to 0.13 cm wide, glabrous or minutely puberulent; stipules deciduous, ovate oblong, to 8 mm long, obtuse or acute, rubescent outside, glabrous, setose within. Inflorescences terminal, solitary, paniculate, to 8 cm long, usually 2 times longer than wide, puberulent or glabrous; peduncles 1.5-6.0 cm long, usually terminating in 5 radially disposed branches or the rachis extended and the branch- es in 2 well-spaced pairs, the cymules terminal. Flowers pedicellate, the pedicels to 2 mm long, the hypanthium short, the calycine cup ca. | mm long, pubescent, petaloid, eglandular or with 5 well-spaced glands within, glabrous within, the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 385 teeth 5, 1.0-2.5 mm long, sometimes unequal, acute; corolla white or yellow, the tube cylindrical, 2-4 mm long, puberulent to glabrate outside, with soft, white, elongate hairs within near the point of filament attachment, the lobes 5, oblong, petaloid, venose, about % the length of the tube, obtuse or acute; stamens 5, exserted, the anthers narrowly oblong, ca. 1 mm long, the filaments slender, to 2 mm long, attached near the mouth; ovarian disc pulvinate, ca. 0.3 mm long, the style 1.5—2.0(—7.0) mm long (fide Croat), the stigmas 2, digitiform, ca. 1 mm long, papillose. Fruits subglobose, 4-5 mm in diam., minutely puberulent, dis- tinctly furrowed, the calyx often persistent, to 2 mm long. Psychotria horizontalis ranges from Mexico south to Ecuador. It also occurs in the West Indies. It is usually recognized by a combination of characters: in- florescence with branches usually 5-radiate, ‘‘domatia’’ usually in the axils of the veins, the fruits with persistent elongate calyx. CANAL ZONE: Madden Dam, Allen 2563 (F). Fort Kobbe, Duke 4701, 4197 (both MO). Fort “masaq Duke 6910 (MO). K-9 Road, ie iid 2620, 2858 (both MO). Madden Dam, Dwyer & Hayden 23 (MO); Dwyer & Lallathin 8816 (MO). Mandingo Road, Hayden 59 (MO). Madden Dam, Kirkbride 39 (MO). Río Petitpie, road to Fort Sherman from Gatün Locks, Mori & Kallunki 2658 (MO). RR and Panama Canal, 6 km W of Gamboa, Nee 7207 (MO). 1 km NW of Gamboa, Nee 9543 (MO). 2: Standley 27181 (US). Las Cruces Trail, Standley 39178 (US). K-9 Road, Stern et al. 15 (MO). oad K-9, Río Cocoli, Stern et al. 325 (MO). Miraflores Lake, Chiva-Chiva Trail, Tyson 1345 (MO). Du Fort Lorenzo, Tyson 2233 (MO). Fort Sherman, Tyson & Dwyer 1186, 1197 (both MO). 12 mi S of Colón, Tyson et al. 4472 (MO). Miraflores, White 139 (MO). BARRO COLORADO ISLAND: Aviles 9 (MO); Bailey & Bailey 393, 566 (both F); Croat 4110, 4138 (both MO), 4383 (F, MO), 4752, 4985, 5083, 5137, 5498, 5578, 6011, 6274, 6300A, 6383, 6824, 9226, 10758 (all MO); Dwyer 1411 (MO); Ebinger 251 (MO), 556 (F, MO), 878 (MO); Hayden 15, 38, 138, 175 (all MO); Shattuck 1025 (F, O ip He : omé i (MO). Aue s 3859 (MO). Near home of P. H. Zimmerman, Folsom et al. 3327 (MO). DARIÉN: Río ues Duke 4953, 4954 (both MO). Río Mortí near Mortí Arriba, Duke 8443 (MO). Río Chucunaque, Cativo Swamp, Duke 11743 (MO). Río Mortí, Duke 14178 (F). Mortí Hydro, Duke 14238 (MO). Piria- 'Cañasas trail near Piria, Duke 14313 (MO). Cañasas and Sabalo, Duke 14470 (MO). Río Mortí, Mortí Hydro, Duke 15822 (MO). Without other locality, Duke & Bristan 102 (MO). El Real, Duke & Bristan 288 (MO). Without other locality, Sexton 115 (MO). Yaviza, Stern et al. 164 (MO, US). HERRERA: Ocú, Ebinger 1058 (F, pn 1071 (MO). Los sANTOS: Río Pedregal, 25 mi SW of Tonosí, Lewis et al. 2915 (COL, MO, UC, VEN). Guaniquito, near Tonosí, Luteyn & Foster 1382 F, A Á .P pelag Beach, Duke 11801 (MO). San Jose Island, Erlanson 213, 236, 477, 556 (MO). Bella Vista, Maxon & Valentine 6937 (US). Trapiche Island, Miller 1900 (US). Los Asientos, Wendehake 40 (MO). Isla Taboga, Woodson et al. 1459 (F, MO). 43. Psychotria impatiens Dwyer.!? TYPE: Panama, Mori & Kallunki 5446 (MO). Shrubs to 2 m tall, the twigs leafless except terminally, terete, finally angular, the bark grey when dry, glabrous, the nodes moderately turgid, crowded. Leaves (immature) ovate lanceolate, to 2.6 cm long, 0.7—1.0 cm wide, acute at the apex, cuneate at the base, the costa prominulous on both sides, the lateral veins ca. 7, 19 Psychotria impatiens Dwyer, spec. nov. Frutices ad 2 m alti foliis saepe persistentibus solum versus apicem ramulorum, glabri. Folia hic immatura ovato-lanceolata, ad 2.6 cm longa, 0.7-1.0 cm patulis glabris. Inflorescentiae terminales compresso-paniculatae, pedunculo nullo vel conspicue re- ducto floribus paucis. Flores subsessiles; dentibus calycis 0.3 mm longis; corolla tubo 5-6 mm longo extus glabro. Fructus non visi. 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 arcuate, the submarginal vein to 1 mm from the margin, membranous, concol- orous, glabrous; petioles absent or to | mm long; stipules persistent toward the tips of the twigs, connate basally, ovate elliptic, to 3 mm long, 1.5 mm wide, acute, glabrous. Inflorescences terminal, compressed paniculate, shorter than the leaves, to 1.2 cm long, to 1.5 cm wide, the peduncle obsolete, with few, ascending branches, the flowers ca. 12, the bracteoles minute, ovate. Flowers subsessile, the hypanthium to | mm long, glabrous, the calycine cup moderately turgid, 0.3 mm long, the lobes 5, triangular, 0.3 mm long, often spreading or somewhat reflexed; corolla white, the tube narrowly cylindrical but inflated toward the apex, 5-6 mm long, glabrous outside, villose within above the middle, the lobes 5, narrowly oblong, to 3 mm long, sparsely pubescent within; style linear, to 7 mm long, the stigmas oblong, to 1.7 mm long, obtuse. Fruits not seen. Psychotria impatiens is known only from Panama. Its precocious inflores- cence is unique among the Psychotria of Panama. The juvenile leaves are virtually sessile and crowded at the tips of the twigs. The nodose character of the twiglets is diagnostic as are the spreading subcoriaceous stipules which seem connate basally, drying a dull brown. The much reduced inflorescence is noteworthy. slope of Cerro Pirre, lower montane rain forest (cloud forest), ca. 700-950 m, Mori & Kallunki | (MO). 44. Psychotria indigotica Dwyer.'°! TYPE: Panama, Hladik 511 (MO, holotype). Trees to 7 m tall, the branchlets terete, smooth, glabrous, dull yellow when dry, the nodes well spaced. Leaves ovate oblong, 8.5—15.0 cm long, 4—9 cm wide, deltoid toward the apex, often acuminate, the acumen to 0.5 cm long, obtuse or rounded at the apex, the costa slender above, prominulous beneath, yellowish white when dry, the lateral veins 5-7, arcuate, prominent beneath, yellowish white, with 1—2 veins between a pair of lateral veins, these soon disappearing, the smaller veins pinnatiform, conspicuous and yellowish white, the undulate submarginal vein only ca. 3 mm from the margin, thinly coriaceous, glabrous, greenish yellow when dry; petioles rigid, to 3.5 cm long; stipules connate, crater shaped, to 0.2 cm long, coriaceous, glabrous, puberulent on both sides, without teeth or lobes. /nflorescences terminal, cymose paniculate, to 7 cm long, the peduncle 1.0-2.4 cm long, rigid, glabrous, the branches for the most part disposed as 2 pairs, the lowermost to 2.5 cm long, arcuate or angular ascending, the cymes few flowered, the branches often abnormal and swollen, to 0.4 cm wide. Flowers not seen. Fruits rotund, to 1.5 cm wide, blue, glabrous, the pyrenes oblong, to | cm long, truncate, conspicuously costate, often yellow green when dry, the calyx conspicuous; seeds blue black, with a T-shaped indentation as seen in cross section, often but one seed developing. 101 Psychotria indigotica Dwyer, spec. nov. Arbores ad 7 m altae ramulis glabris. Folia ovato- oblonga, 8.5-15.0 cm longa, 4-9 cm lata, venis rioris 5-7 in sicco bop luteis, tenui-coriacea glabra; petiolis ad 3.5 cm longis; stipulis connatis crateriformibus, ad 0.2 cm longis, coriaceis glabris extus intusque puberulis dentibus nullis. [s le terminales EURO ae ad 7 cm longae, pedunculo 1.0-2.3 cm longo, glabro ramulis pro maxime parte 2-jugis dispositis uem ad 2.5 cm longis cymis paucifloris in ramis reductis dispositis. Fructus rotundi, ad 1.5 cm lati, ubi carne praesente azurei glabri seminibus indigoticis 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 387 Psychotria indigotica is known only from Panama. It is so named because of its blue-black seeds. 45. Psychotria insignis Standley, Contr. U.S. Natl. Herb. 18: 130. 1916. TYPE: Panama, Pittier 4410 (US, holotype; F, isotype). Shrubs, the branchlets stout, angular, ferrugineous hirsute. Leaves ovate or oblong, 14-22 cm long, 7-9 cm wide, falcately acuminate, cordate at the base, the costa prominent beneath, the lateral veins 17-20, widely spreading and de- pressed arcuate, uniting submarginally to form a distinct undulate vein, stiffly chartaceous, concolorous, drying chocolate red; petioles to 5 cm long, ferrugin- eous hirsute; stipules deciduous, oblong, ca. 1.5 cm long, obtuse, ferrugineous hirsute. Inflorescences terminal, ferrugineous hirsute, 3 cm long, wider than long, the peduncle to 2 cm long, the branches 5, radiate, 1-2 cm long, the cymules densely flowered, usually 3 per branch, ca. 1 cm in diam., the bracts not seen. Flowers sessile, the calyx cup shaped, to | mm long, densely puberulent, the lobes oblong, 1.0-1.5 mm long. Fruits not seen. This species is known only from the type collection in Panama. SAN BLAS: Hills of Sperdi, near Puerto Obaldía, Pittier 4410 (F, US). 46. Psychotria liesneri Dwyer.'°? TvPE: Panama, Mori & Kallunki 3658 (MO, holotype). Shrubs to 2 m tall, the ultimate branchlets slender, smooth, glabrous, drying ashen gray, the nodes well spaced, scarcely swollen. Leaves elliptic, 3.0-7.5 cm long, 1.7-2.0 cm wide, acute at the apex, acuminate, the acumen to 0.5 cm long, ultimately obtuse, acute at the base, the costa plane above and beneath, ca. 0.8 mm wide proximally, drying white above, the lateral veins ca. 8, occasionally with domatia in the axils, arcuate, uniting well below the margin to form an undulate vein, the smaller veins not visible, the margin scarcely callose, often delicately involute, thinly coriaceous, lustrous, glabrous, scarcely discolorous; petioles slender, to 1 cm long; stipules deciduous, triangular, to 3.5 mm long, obtuse. Inflorescences (in fruit) terminal, shorter than the leaves, puberulent; peduncle stiff, 1.3 cm long, slender, wiry, ca. 0.7 mm wide, terminated by 3 divergent branches to 2.3 cm long, wiry, the flowers presumably very few, di- vergent. Flowers not seen. Fruits subsessile, rotund, to 4 mm in diam., glabrous, drying black, the persistent ovarian disc often umbonate, subplane, to 1.7 mm in iam. Psychotria liesneri occurs only in Panama. It is named in honor of Mr. Ronald Liesner, student of the flora of the Republic. It is closely allied to the small- 102 Psychotria liesneri Dwyer, spec. nov. inp s ad 2 m alti, ramulis ed ape Lond nodis bene i: Folia elliptica, 3.0-7.5 cm longa, 1.7-2.0 cm lata, acuta costa utrinque plana venis lateralibus venis minoribus inconspicuis, e coriacea glabra; stipulis irianguleribus obtusis. Infloresc entiae E pedunculo ca. 1.3 cm longo. Fructus rotundi, ad 4 mm lati, glabri, in sicco 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 73. Psychotria limonensis Krause —A. Habit (x'2).—B. Flower, exploded to show interior (x2). [After Hayden 26. | leaved P. chagrensis Standley which has bilobed stipules and a much more del- icate peduncle. PANAMA: 8 mi S of Goofy Lake toward Cerro Jefe, 2200 ft, Dwyer 7098 (MO). NE of Cerro Azul, premontane rain forest, Mori & Kallunki 3658 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 389 47. Psychotria limonensis Krause, Bot. Jahrb. Syst. 54: Beibl. 119: 43. 1916. TYPE: Costa Rica, Pittier 12681 (US).—Fic. 73. Subshrubs or shrubs to 3 m tall, the branchlets terete, ultimately angular, glabrous, smooth, the nodes well spaced. Leaves widely oblong, often oblong rotund, regularly ca. 2 times longer than wide, 11—30 cm long, 3.5-12.0 cm wide, widely deltoid at the apex, rarely acute and falcate, vaguely acuminate, the acu- men deltoid, to 1.5 cm long, obtuse, basally cuneate, rarely truncate, often slightly inequilateral, the costa subplane or prominent beneath, often rubescent beneath, subperpendicular to widely arcuate, often rubescent, chartaceous, discolorous, fleshy, glabrous; petioles to 5 cm long, to 0.3 cm wide, glabrous; stipules free, compressed rotund, ca. 0.8 cm long, ca. 0.9 cm wide basally, coriaceous, gla- brous. Inflorescences solitary, terminal, paniculate, subrotund, to 5.5 cm long, usually slightly wider than long; peduncles 1.5-2.0 cm long; bracts ovate or tri- angular, ca. | mm long, pubescent. Flowers ca. 3 mm long; hypanthium ca. 0.8 mm long, glabrous to puberulent, the calycine cup short with a few appressed, round glands within, the teeth minute, triangular, acute; corolla white or cream colored, the tube ca. 2 mm long, glabrous outside, villose within; stamens 5, the anthers oblong, ca. 0.9 mm long, the filaments short, ca. 1 mm long, villose, attached near the mouth; style slender, ca. 1 mm long; stigmas linear, ca. 1 mm long, the ovarian disc subrotund, ca. 0.4 mm long. Fruits subglobose or oblong, 6—7(-10) mm in diam., red at maturity, the pyrenes hemispherical, to 4 mm long, the ventral face plane, the dorsal face convex, the ribs 5 per pyrene. Psychotria limonensis occurs in the lowland rain forests of Central America and northern South America. . TORO: Almirante, Cooper 453 (F). Changuinola, Lewis et al. 822 (GH, MO, UC, US). Water. Valley, Wedel 1407, 1770 (both MO). Old Bank Island, Wedel 2137 (F, MO). CANAL ZONE: Fort Sherman, Blum & Dwyer 2108 (MO). Coco Solo, Duke 4264 (MO). Galena Point, Dwyer 6626 (MO). Fort — Pad & Robyns 136 (MO). Galena Point, Dwyer & Robyns 160 (MO). BARRO ; (M , & Vestal 310 (F). coLÓów: Portobelo, Ebinger 12 M US). DARIÉN: Quebrada and Peje Swam Bristan 1051 (MO). Without other locality, Sexton "os SAN BLAS: 3-4 hours by foot from Río Mulutupu, Kirkbride 226 (MO). Molina near river, Stier 39 (MO). 48. Psychotria longipedunculata Dwyer.!? TYPE: Panama, Croat 27621 (MO, ho- otype Shrubs to 1.2 m tall, smooth, angular, minutely puberulent when dry, the nodes well spaced. Leaves elliptic, to 18 cm long, to 9 cm wide, deltoid toward the apex, the costa prominulous above, prominent beneath, the lateral veins 12— 1? Psychotria longipedunculata Dwyer, spec. nov. Frutices ad 1.2 m alti ramulis minuto-pu- berulis. Folia Ee ca, ad 18 cm longa, ad 9 cm lata, venis lateralibus 12-15, rigido-membranacea Lene: ata; petiolis ad 3 c m longis, saepe reflexis. Inflorescentiae d cymoso-paniculatae pedun- o 14-16 cm ipee foribus i in caespitem terminalem oblon ad 3 cm longum dispositis bracteis Treni ankuan. ad 1 cm longis. Flores sessiles cupula salya 07 mm longa, lobis 5, 1.5 mm longis; corolla ad 2.5 mm longa, tubo extus glabro; antheris 0.8 mm longis. Fructus obovato-oblongi, ad 6.5 mm longi, costis linearibus aliquot glabri praediti. 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 15, arcuate, with 1-2 smaller and shorter veins between 2 lateral veins, the in- tervenal areas spreading reticulate, stiffly membranous, concolorous, shiny above, smooth, glabrate; petioles to 3 cm long, to 0.15 cm wide, often reflexed, glabrous; stipules not seen. Inflorescences axillary, cymose paniculate, the pe- duncle 14-16 cm long, to 0.18 cm wide, strongly ascending, rigid, wiry, the flow- ers disposed in an oblong terminal mass, to 3 cm long, to 3 cm wide; bracts linear subulate, to 1 cm long. Flowers (buds) sessile, the hypanthium short, the calycine cup 0.7 mm long, glabrous outside, eglandular and glabrous within, the lobes 5, oblong, 1.5 mm long, obtuse, petaloid; corolla to 2.5 mm long, the tube short, glabrous outside, villosulose near the filament attachment, the lobes 5, shorter than the tube, cucullate; stamens 5, the anthers oblong, 0.8 mm long, obtuse, the filaments short; ovary disc oblong hemispherical or compressed-rotund, 0.5—0.65 mm long, the style short, the stigmas oblong. Fruits obovate oblong, often slightly falcate, to 6.5 mm long, obtuse toward the apex and base, glabrous, the ribs linear, the raphides abundant. Psychotria longipedunculata is known only from Panama. Its extraordinary elongate peduncles distinguish it from all other Central American Psychotria. ERAGUAS: Valley of Río Dos Bocas, 15.6 km NW of Santa Fé, road between Escuela Agricola Alto Pein and Calovebora, Croat 27621 (MO). 49. Psychotria luxurians Rusby, Mem. Torrey Bot. Club 6: 50. 1896. TYPE: Bo- livia, Bang 1741 (US, holotype). P. berteriana subsp. luxurians (Rusby) Steyermark, Mem. New York Bot. Gard. 23: 534. 1972. Trees or shrubs to 25 m tall, the branches terete, smooth, glabrous, distinctly hollow, often constricted below the nodes, the latter well spaced, the branchlets ultimately subplane. Leaves oblong, often widely so, elliptic, 8.6-24.5 cm long, 3.2-15.0 cm wide, tapering obtusely or deltoidly at the apex, acuminate, occa- sionally inequilateral, the costa prominulous above, prominent or subplane be- neath, to 0.22 cm wide proximally, the lateral veins 10-18, arcuate, prominulous and often evanescent above, prominent beneath, the intervenal areas with smaller pinnatiform veins, papyraceous, usually discolorous, glabrous above, often pu- berulent on the veins and the costa beneath; petioles to 4 cm long, to 0.2 cm wide, usually glabrous, occasionally puberulent; stipules free, oblong or oblong rotund, often compressed and wider than long, 5-7 mm long, 3.7-8.0 mm wide, obtuse, rounded, often undulate or vaguely retuse, appressed setose within. In- florescences terminal, shorter than or exceeding the terminal leaves, to 22 cm long, paniculate, usually pyramidal paniculate; peduncle 1-6 cm long, 0.1—0.4 cm wide, subplane, angular, puberulent, the lowermost branches opposite, to 7 cm long, usually arcuate ascending or perpendicular, the axis with several paniculate branches, ultimately bearing densely flowered cymules; bracts of the primary branches linear oblong or subulate, to 5 mm long; hypanthium and calycine cup to 1.5 mm long, the calycine cup ca. 0.5 mm long, carnose, glabrous, the teeth widely deltoid, unequal, to 0.5 mm long, usually with a minute gland at the junction of 2 teeth; corolla tube cylindrical, to 3 mm long, glabrous outside, the lobes 5, ca. | mm long, villose within; stamens 5, the anthers ca. 1 mm long, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 391 obtuse, the filaments crassate, short; stigmas 2, subulate, ca. 1.2 mm long, villose, the style ca. 1.5 mm long. Fruits sessile, oblong, ca. 3 mm long, glabrous, the pyrenes with the ventral face of the endocarp flat and with a fissure, the dorsal face with 5-6 ribs. Psychotria luxurians ranges from Panama south to Bolivia. cHiRIQUÍ: Cerro Colorado, 50 km N of San Felix, 1200-1500 m, Mori & Dressler 7801 (MO). COCLE: El Valle, Croat 13396 (MO). Cerro Pilón near El Valle, 700-900 m, Duke 12125, 12126 (both 5641 (both MO). Cerro Pilón, ca. 2900 ft, Dwyer 8342 (MO). 8.5 mi from Club Campestre toward La Mesa, Dwyer 10559 (MO). El Valle. foot of Cerro Pilón, Dwyer & Correa 7989 (GH, MO, US). Cerro Gaital Caracoral, 3100 ft, Dwyer & Correa 8872 (MO). El Valle, Folsom 3117 (MO). Summit Alto Clavario, 900 m, Folsom & Robinson 2413 (MO). La Mesa above EI Valle, Gentry 7435 (MO). El Valle de Antón, $i 1215 (MO). El Valle to La Mesa, Spellman et al. 558 (MO). COLON: Forested near Guasimo, Croat 9949 (MO). Santa Rita Ridge, Croat 13838, 15311 (both MO); Croat & Porter 15598 (MO); Dwyer 8559 (MO); Dwyer et al. 9003 (MO). Near Agua Clara rainfall station, Foster 1750 (MO). Santa Rita Ridge, Gentry 1873 (MO). DARIÉN: Cerro Pirre, 750-1030 m, Duke 5332 (MO). Upper Río Membrillo and Camp 7, road to San Blas, 100-800 m, Duke 10892 (MO). Cerro Pirre, 2500-4500 ft, Duke & Elias 13787 (MO). Cuasi-Cana Trail, Cerro Campamiento, E of Tres Bocas, Kirkbride & Duke 1249 (MO). Cuasí-Caná Trail, Chepigana, Terry & Terry 1448, 1524 (both MO). PANAMÁ: Campo Tres, 5 km NE of Altos de Pacora, Busey 826 (MO). 3 mi N of Cerro Azul, Croat 11593 (MO). Cerro Jefe, Croat 13058 (MO). Altos de Pacora, 10 mi beyond Goofy Lake, Croat a Dwyer & Blum 3255 (MO). Cerro Jefe, Dwyer & Gauger 7354 (GH, MO, UC, US). Cerro Campana, Dwyer & Kirkbride 7843 (MO). Cerro Jefe, Elias & Hayden 1809 (GH, MO). El Llano Carti Road, 7.8 km N of Panamerican Highway, Folsom 3563 (MO); Same locality, 14 km N of Panamerican Highway, Folsom et al. 1484 (MO). Between Cerro Azul and Cerro Jefe, Foster p (MO). Cerro Jefe, Gentry 2126 (MO). Cerro Campana, Hayden 119 (MO). Cerro Jefe, Hayden 1001 (MO). Río Cerro Jefe, Mori & Kallunki 2768 (MO). El Llano-Cartí Road, 10.8 km from Interamerican Highway, Mori & Kallunki 4135 (MO). Cerro ie Tyson et al. 3222, 3255, 4435 (all MO). VERAGUAS: Cerro Tute, cloud forest, 1000 m, Mori 6258 (MO). Road to Calovebora, Mori & Kallunki 5356 (MO). 50. Psychotria macrophylla Ruiz & Pavon, Fl. Peruv. Chil. 2: 56. 1799. TYPE: ‘‘Nemoribus versus Pillao ad Iscutunam,’’ Huanuco, Peru, Ruiz & Pavon, not seen. P. anomothyrsa Schum. & Donnell Smith, Bot. Gaz. 35: 3. 1903. rype: Guatemala, Von Tuerckheim 7909, not seen. P. tolimensis Wernh., Kew Bull. Miss. Inform. 68. 1914. TYPE: ERN Sprague 237 (K). Hoffmania striata Rusby, Descr. S. Amer. Pl. 134. 1920. TYPE: not P. macrophylla Ruiz & Pavón var. angusha Standley, Publ. Field Mur Nat. Hist., Bot. Ser. 22: 1940. rvPE: Panama, Terry & Terry 1597 (F, holotype). Shrubs to 4 m tall, the branchlets subterete but ultimately angular, smooth, glabrate, the nodes 2-6 cm apart, the pith porate. Leaves strongly ascending, elliptic or oblong, often narrowly oblong, rarely oblong rotund, often somewhat falcate, 9—45 cm long, 3-10 cm wide, deltoid to acute at the apex, usually some- what contracted into a short acumen, the latter to 2 cm long, basally acute or cuneate, the costa subplane or immersed above, prominulous and occasionally grooved beneath, the lateral veins ca. 13, arcuate, often widely so, often with 1— 2 smaller veins between 2 adjacent lateral veins, scarcely prominulous, papyra- ceous, rarely membranous, glabrous or with small, well spaced, strigose, white hairs above and beneath; petioles 1-7 cm long, flexuous or stiff; stipules (fide 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Standley, only one seen), free, caducous, short, broad, apiculate. Inflorescences axillary, paniculate, to 18 cm long, to 9 cm wide at the base, the peduncles 1.5— 14.0 cm long, 1.2 mm wide, glabrate to pubescent, the hairs variable, the branches opposite, rarely verticillate, well spaced, usually 2-3 pairs in larger inflores- cences. Flowers with the hypanthium subrotund, ca. 0.5 mm long, villosulose outside, the calycine cup ca. 0.8 mm long, glabrous and eglandular within, the calycine lobes scarcely visible or as 5 triangular teeth, ca. 0.4 mm long; corolla white or yellow, short, to 4 mm long (including lobes), glabrous outside, villose within; stamens 5, the anthers oblong, ca. 1.2 mm long, the filaments short; style ca. 2.2 mm long, the stigmas 2, small, ca. 0.35 mm long. Fruits subrotund or subovate, drying white, 3.5-5.0 mm in diam., the pyrenes 2, strongly compressed, the dorsal face of each pyrene 4-5 costate, the ribs conspicuous. Psychotria macrophylla ranges from Mexico to Bolivia. Despite the fact that narrow-leafed collections can be easily segregated, I have found too much inter- gradation in leaf width to favor retention of var. angustissima Standley. A col- lection of particular interest is Mass & Dressler 1703 from El Llano-Carti Road, in which the inflorescences are almost sessile. BOCAS DEL TORO: Río Teribe, Kirkbride & Duke 512 (MO). Changuinola, Le n xi al. 969 (GH K, MO, UC, US). Chiriquicito to 5 mi S of Río Guarumo, Lewis et al. n (GH US). Water Valley, Chiriquí Lagoon, Wedel 1572 (MO). Nievecita, Woodson et al. 1840 (M D ae Baj Chorro, Boquete, Hi 229 (MO). Cerro Horqueta, Duke & Kirkbride 108 (MO); Duke et hi 13643 (MO). Boquete, Dwyer & Hayden de (GH, K), 7676 (MO). El Boquete, Maxon 4934 (US). Cerro Colorado, 50 km N j^ San Félix, 1200-1500 m, Mori & Dressler 7859 (MO). Las Signas Camp. Pittier 3271 (US). Upper Río Gariche, Seibert 360 (MO). Chiriquí Viejo Valley, White 87 (MO). Río Caldera, Wilbur et al. 11070 (MO). Bajo Mono, along Río Caldera, Woodson et i 1017 (MO). COCLE: Cerro Pilon, Croat 13468 (MO). La Mesa, Dwyer 8057 (MO). Cerro Pilón. Dwyer 8634 (MO). El Valle, Dwyer & Correa 7932 (GH, MO, US). Cerro Gaital Caracoral, Dwyer & Correa 8863 (MO). La Pintada, Hunter & Allen 542 (MO). Boca del Toabre, Lewis et al. 5533 (MO). Cerro Pilon, 5 km NE of El Valle, Mori et al. 6558 (MO); Porter et al. 4617 (COL, MO, SCZ, UC, VEN). COLON: Río n ca. 2.5 km upriver from bridge to Portobelo, Mori & Kallunki 3707 (MO). Río Guanche, m S of Portobelo, Nee & Gentry 8687 (MO). DARIEN: El Real, Bristan 108 (MO). Rio Yape, Pons 1370 (MO). Río Pirre, Bristan 1486 (MO). Cerro Pirre, Duke 5292, 6555 (both MO). Cerro Campamiento, Duke 15652, 15672, 15718 (all MO). Cocalito, Dwyer 2162, 2194 (both MO). S slope of Cerro Tacarcuna, 1250-1450 m, Gentry & Mori 13940 (MO). Top of W ridge, Cerro Tacarcuna, Elfin Forest, 1800-1850 m, Gentry & Mori 14021 (MO). 0-2 mi E of Tres Bocas. Kirkbride & Duke 1182 (MO). Cerro Campamiento, Kirkbride & Duke 1219, 1356 (both MO). Caná-Cuasi Trail, Che- pigana, Terry & Terry 1530 (MO). Caná Cuasi Trail, Terry & Terry 1597 (F). PANAMÁ: Cerro Ca ampana, Correa & Dressler 1016 (MO): Croat 14739 (MO). Cerro Jefe, Duke 15238 (MO). Cerro Campana, pe & Kirkbride 7860 (MO); oe 249 (MO); Kirkbride & Hayden 288, 301 (both MO). Cerro Campana, Lewis et al. 3090 (MO, UC); Luteyn 1014 (MO). El Llano- ple Road, 400 m, Maas & Dre ins 1703 (MO). Cerro ne Mori & Kallunki 3541 (MO). vER AS: NW of Santa Fé, 1 Santa Fe, 11 km from Escuela Agricola Alto de Piedra, Valley Rio dos Bocas, Mori et al, 3824 (MO). Road from Santa Fé to Calovebora, 16 km from Santa Fé, Atlantic slope near Rio Caloveborita, Mori et al. 6692 (MO). 51. Psychotria magna Standley, Contr. U.S. Natl. Herb. 18: 131. 1916. TYPE: Panama, Pittier 4092 (US, holotype; photo, F). P. compressicaulis Schum., Bot. Jahrb. Syst. 54: Beibl. 119. 44. 1916. Shrubs. Leaves elliptic, oval or elliptic oval, 18-25 cm long, 8.0-11.5 cm wide, tapering obtusely or acutely at the apex, the acumen well defined, narrowly tri- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 393 angular, to 2.5 cm long, cuneate at the base, the lateral veins 17-20, minutely puberulent beneath, the blade papyraceous; petioles 2-4 cm long, lignose; stipules not seen. Inflorescences terminal, cymose, paniculate, to 10 cm long, to 10 cm wide. Flowers with the calyx and hypanthium ca. 1.5 mm long, the cup undulate and ciliolate, the teeth absent; corolla yellowish white, 3-4 mm long, glabrous outside, densely villose within at the mouth, the lobes 1.5 mm long; anthers 5, minute, oblong, 0.7 mm long, obtuse. Fruits not seen. Psychotria magna is known only from Costa Rica and Panama. COLÓN: Loma de la Gloria, Fató, Pittier 4092 (F. US). 52. Psychotria marginata Swartz, Nov. Gen. Pl. Prodr. 43. 1788. TYPE: Jamaica, Swartz, not seen. Uragoga marginata (Swartz) Kuntze, Rev. Gen. Pl. 2: 960. 1891. Shrubs to 3 m tall, the branchlets terete, smooth, glabrous, the nodes well spaced. Leaves elliptic, oblong or obovate oblong, rarely oblong rotund, to 20 cm long, to 6.8 cm wide, rounded or deltoid at the apex, acute to attenuate acute at the base, rarely truncate, the costa plane and often grooved to prominulous above, prominulous beneath, often subplane, to 1.3 mm wide proximally, the lateral veins 9-15, arcuate, prominulous above and beneath, occasionally mi- nutely pubescent or farinose beneath, often with domatia in the axils, these often mammiform, | mm long, puberulent, the intervenal areas usually spreading re- ticulate below, evanescent above, papyraceous, concolorous, often drying me- tallic gray or brown; petioles to 2.3 cm long, to 0.17 cm wide, glabrous; stipules free, usually appressed, oblong, 1.2-1.6 cm long, acute or obtuse or tapering into an obtuse tip, scarious, glabrous. Inflorescences terminal, conspicuously open and spreading, solitary, paniculate, 6-15 cm long, the peduncle slender, to 8 cm long, the branches opposite, usually disposed as 3 well-spaced pairs, the lower- most to 8 cm long, strongly divergent; bracts triangularly ovate, to | mm long, glabrous or pubescent. Flowers with pedicels to 2 mm long, the calycine cup ca. 0.4 mm long, thickly petaloid, eglandular within, glabrous outside, the teeth al- most absent or widely triangular, unequal, ciliolate, shorter than the calycine cup; corolla yellow, yellow green, or white, the tube cylindrical, ca. 3 mm long, glabrous outside, pilose within, the lobes oblong, 0.4-1.8 mm long, glabrous or with tufts of hairs proximally; stamens 5, included, the anthers narrowly oblong, 1.0-1.2 mm long, obtuse: ovarian disc subglobose, to 0.5 mm long, the style shorter than the corolla tube, somewhat dilated above, the stigmas to 0.7 mm long. Fruits rotund, to 4 mm long, glabrous, the calycine cup usually persistent, minute, the pyrenes plane on the dorsal face, the convex face 4—S-ribbed, the albumen ruminate on the ventral face. Psychotria marginata is common throughout tropical America and is well collected, perhaps more so than any other species of Rubiaceae in Panama. In Darien the species is known as "'Gorojito"" (Duke 14271). AS DEL TORO: 1.5 m W of Almirante, Blum 1361 (MO). Island of Bocas del Toro, Durkee 71- 133 (MO). Rio Changuinola, Changuinola, Dwyer 4400 (MO). Cerro Bonyic, above Quebrada Huron, 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 | 0, Y | Pa (a NP Z by Ro X Y 2 Na. C =$: Pa - Nd c J) Pa Lor ZA (2 , XE < fy “ATL N / / QW. N $ j Š CX Zx LIN Ky Y JE \ / FIGURE 74. Psychotria maxonii Standley.—A. Habit (x1⁄4).—B. Flower (x 5).—Flower, opened to show interior (x 5).—Fruit (x5). [After Mori & Bolten 7589.] Kirkbride & Duke 591 (MO). Almirante, McDaniel 5129 (MO). Chiriquí Lagoon, Wedel 1079, 1191, 1347 (all bn Water Valley, Wedel 1722 (MO). Old Bank Island, Wedel 2011, 2160 (both MO). CANAL Z : Cruces Trail and Madden Dam Road, Bartlett & Lasser 16343 (MO). 2 mi from Gatün Locks, num 1494 (MO). Fort Sherman, m Hollow, Blum & Dwyer 2107 (MO). Galena Point, Blum & Dwyer 2148, 2152 (both MO). boa pne on TTC lands, Correa 483 (MO). E of Gamboa, Croat 4784 (MO). Las Cruces Pe Ma dai Forest, Dwyer 1959 (MO). Fort Sherman site, USATTC, Dwyer 8591 (MO). George Greene Memorial, Dwyer & Hayden 6 (MO). Fort Sherman, Skunk Hollow, Dwyer & Robyns 135 (MO). Chagres, " Fendler 64 (MO). Pipeline Road, Fosberg 55254 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 395 (US). Las Cruces Trail, Hunter & Allen 461 (MO). Pipeline Road, NE of Gamboa, Kennedy 2334 (MO). Pipeline Road, Agua Salud, Kennedy & Andrews 1903 (MO). Colon Highway, Road to Fort Sherman, Lazor & Blum 5364 (MO). Shore of Gatún Lake, Lewis et al. 1830 (GH, MO, US). Madden Forest Preserve, Las Cruces Trail, Lewis et al. 5320 (COL, MO, SCZ, UC, VEN), 5340 (MO, SCZ, UC). Road S-8 at Quebrada La Furnia, 142 km S of mouth of Chagres, Nee 8637 (MO). Pipeline Road, 9 km NW of Gamboa, Nee 9598 (MO). Pipeline Road, 19 km NW of Gamboa, Nee & Smith 11105 (MO). Fort Sherman to Fort San Lorenzo, Porter et al. 4993 (MO, SCZ, UC, VEN). Splice S-16 and S-49, Río Indio, Steyermark & Allen 17456 (MO). 6 mi N of Gamboa, Tyson 1440, 1517 (both MO). Fort Sherman, Tyson & Blum 1659 (MO). 1 mi N of Summit, Tyson & Blum 2762 (MO). 12 mi S of Colón on Rio Providencia, Tyson & Blum 3978 (MO). Fort Sherman, Pena Road, Tyson & Chu 1711 (MO). Pipeline Road, Witherspoon & Witherspoon 8630 (MO). BARRO COLORADO ISLAND: Croat 4033, 4099, 4105, 4303, 5083, 5678, 5681, 5775, 5838, 5843, 6394, 6519, 6548, 7134, 7325, 7718, 7838, 10315 (all MO); Ebinger 615 (MO); Hayden 14, 28, 129 (all MO); Knight 69-4 (MO); McDaniel 5012 (MO); Robyns 65-5, 65-10 (both MO), 65-27 (MO, US); Shattuck 116, 274, 932 (all MO); Starry El Valle, Allen 1187 (MO). COLON: Rio Guanche, D’Arcy 9686 (MO). Portobelo, Ebinger 108 (US). Vicinity of Achiote, Folsom 3858, 3860 (both MO). 6 mi SW of Portobelo, Luteyn 1416 (MO). 4 km NW of Salamanca, 13 km NE of Buenos Aires, Nee 9091 (MO). DARIEN: Río Yapé, Bristan 1386 (MO). Río Chico across from Boca de Tesca, Duke 5223 (MO). Summit Camp, between Sasardi and Morti, Duke 10042, 10048 (both MO). Camp Clarita, near Chucunaque-Bayano Divide, Duke 15538 (MO). Cerro Campamento, S of Cerro Pirre, Duke 15960 (MO). Río Ganglon, Duke & Bristan 362 (MO). Río San Jose, Duke & Bristan 396 (MO). Río Pirre, Duke & Bristan 8296 (MO). Paya and Palo de los Letras, Duke & Kirkbride 14306 A, 14070, 14271 (all MO). Tacarcuna Massif, above Rio Pucro, Gentry et al. 16958 (MO). Between Tres Bocas and Cerro Campamiento, Kirkbride & Duke 1357 (MO). Rio Tuquesa, Tuquesa Mining Company, Mori 7032, 7044 (both MO). Trail from Paya to Pucro, Stern et al. 416 (MO). Cana-Cuasi Trail, Chepigana, Terry & Terry 1422 (MO). 3 mi N of Santa Fé, Tyson et al. 4633, 4692 (both MO). Los sanTos: Loma Prieta, Cerro Grande, Lewis et al. 2201 (COL, MO, UC, VEN). 16 mi S of Macaracas, Tyson et al. 2902 (MO). PANAMÁ: Río la Maestra, Allen 11 (MO). Cerro Campana, Correa et al. 1015 (MO). Between Panama and Chepo, Dodge et al. 16673 (MO). Jenine, Duke 3886 (MO). Río Pita, Duke 4803 (MO). Piria-Canasas Trail, Duke 14342 (MO). Cerro Azul, Dwyer 2105, 2189 (both MO). Cerro Jefe, Dwyer 8526 (MO). Río Pasiga to above waterfall, Gentry 2294 (MO). Río Espave to Rio Agua Clara, Gentry 3764 (MO). Cerro Trinidad, Kirkbride & Duke 1648 (MO). Goofy Lake to Cerro Jefe, Lewis et al. 253 (GH, K, MO, UC, US). Chimán, Lewis et al. 3248 (MO, UC, VEN). Arraijan, Woodson et al. 772 (MO). SAN BLAS: Río Diabolo and Río Acuatí, Duke 14869 (MO). Headwaters of Río Cuadi, Camp Diabolo, Duke et al. 3644 (MO). VERAGUAS: Isla de Coiba, Collenette 469 (MO). 53. Psychotria maxonii Standley, Proc. Biol. Soc. Wash. 37: 53. 1924. TYPE: Costa Rica, Maxon & Harvey 8250 (US).—Fic. 74 Epiphytic subshrubs, ericoid in appearance, the branchlets slender, terete, smooth, glabrous, the nodes relatively well spaced. Leaves oblong to elliptic, occasionally trapeziform elliptic, 0.8—1.8 cm long, 0.25-0.55 cm wide, acute or obtuse, often acuminate, the acumen to 0.6 mm long, acute at the base, the costa evanescent above, plane and almost evanescent beneath, the lateral veins not visible, fleshy, drying papyraceous, dark brown above, buff colored beneath, smooth, glabrous, marginate; petioles to 0.2 cm long, glandular shiny; stipules intrapetiolar, connate, the sheath scarious, to 1.5 mm long, truncate, with fim- briate appendages outside. Inflorescences terminal and axillary, cymose, shorter than or slightly longer than uppermost leaves, glabrous, the peduncle 3-7 mm long, usually terminated by 3 primary radiate branches with 3-5 flowered cymes, the bracts and bracteoles subulate, minute. Flowers short pedicellate; hypan- thium rotund to oblong, 0.6-0.7 mm long, glabrous, the calycine cup almost indistinguishable, eglandular within, the teeth 4, deltoid, 0.35-0.60 mm long, thickly petaloid; corolla white, the tube narrowly cylindrical, ca. 2.5 mm long, ca. 0.65 mm wide, petaloid, glabrous except for a few hairs at the point of filament 396 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 attachment, the lobes oblong, ca. 1 mm long, ca. 0.6 mm wide at the base, acute; stamens in part exserted, the anthers oblong, ca. 0.3 mm long, obtuse, the fila- ments short, attached at the upper !⁄4 of the tube; stigmas 3-4, slender, to 0.5 mm long, the style slender, ca. 2.5 mm long, the ovary 3-4-celled, the ovules obovate oblong, plano convex. Fruits pedicellate, the pedicels to 3 mm long, oblong rotund, ca. 3 mm in diam., smooth, shiny, glabrous, waxy, pink or orange, the ribs few, not prominent. Psychotria maxonii is known only from Panama and Costa Rica. The fimbriate stipules readily separate it from the other epiphytic Psychotria in Middle Amer- ica. CAS DEL TORO: Cerro Horqueta, 6000 ft, Allen 4984 (NY); Hagen & Hagen 2152 (NY). Elfin Pu at divide on riu song Caldera Trail, Kirkbride & Duke 981 (MO). Cerro Hornitos, 40 km NW of Gualaca, 2238 Mori & Bolten 7503 (MO, US). N of San Félix at Chiriquí-Bocas del Toro border, road to Cerro Colorado Copper Mine, 5000-55 00 ft, Folsom et al. 4680 (MO); Mori & Kallunki 5923 (MO). VERAGUAS: Cerro Tute, 750-1000 m, Mori et al. 7589 (MO). 54. Psychotria micrantha H.B.K., Nov. Gen. Sp. Pl. 3: 363. 1819. TYPE: “Am. Austr., Kunth (B, not seen; photo, MO). Mapouria micrantha (H.B.K.) Wernh., Kew Bull., 1914: 69. 1914. Psychotria hindsii Standley, Publ. Field Columbian Mus. Bot. Ser. 7: 97. 1930. TYPE: Colombia, Hinds s.n. (K. holotype, not seen). P. rufescens of authors, not H.B.K. ex R. & S. Shrubs or trees to 5 m tall, the branchlets often with pith hollow, terete but ultimately compressed and angular, usually densely rufous villosulose. Leaves oblong lanceolate, 12-30 cm long, 5-12 cm wide, acute, rarely obtuse at the apex, often acuminate to 1.5 cm long, basally cuneate to acute, often attenuate at the base, the costa prominulous above, prominent beneath, to 0.3 cm wide proxi- mally, the lateral veins 18-26, laxly arcuate, papyraceous, concolorous, the hairs minute, white, strigose on the upper surface, densely rufous villosulose beneath, usually barbate along the costa and the lateral veins; petioles stout, ca. 2 cm long, to 0.3 cm wide, slightly winged (decurrent petiole), densely rufous puberulent; stipules free, usually persistent at the terminal nodes, appressed, deltoid to ovate, 1.0-1.5 cm long, ca. 0.7 cm wide at the base, rufous hirsute, the 2 lobes 1-5 mm long. Inflorescences either terminal or terminal and axillary, shorter or longer than the uppermost leaves; peduncle 6-7 cm long, rufous pubescent, terminated by ca. 6 primary radially disposed branches or occasionally with tertiary branches likewise disposed, the ultimate cymules conglomerate, 0.4—1.0 cm in diam.; bracts subtending the primary branches narrowly subulate, to 0.5 mm long, the brac- teoles of the lesser branches similar in shape. Flowers with the hypanthium ob- long, ca. 0.3 mm long, densely villose, the hairs white, to 0.4 mm long, the calycine cup ca. 0.5 mm long, glabrous within, lacking glands within, the teeth 5, compressed oblong, ca. 0.25 mm long; corolla white, the tube broadly cylin- drical, ca. 2.2 mm long, glabrate outside, villose on the upper % within, the lobes 5, oblong, ca. 1.8 mm long, villose, somewhat cucullate apically; stamens 5, the anthers oblong, ca. 1.5 mm long, the filaments slender, attached at the middle of the tube; ovarian disc hemispherical, ca. 0.5 mm long, shiny black, the stigmas 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 397 to 1.3 mm long. Fruits globose to ellipsoid, ca. 4 mm in diam., pubescent, red; pyrenes with ventral face flat, 5-ribbed on the dorsal surface. Psychotria micrantha occurs in Nicaragua, Panama, Colombia, Venezuela, Ecuador, and Peru. The fact that the pyrenes are pubescent is noteworthy. The common names used by the Cuna Indians of the San Blas (Duke 8466) are '* Uas- icuit’’ and *' Esuargit.' OCAS DEL TORO: Almirante, Blum 1385 (MO); McDaniel 5051 (MO). Chiriqui Lagoon, Wedel 1163 (MO). Water Valley, Wedel 1809 (F, MO). Old Bank Island, Wedel 1869 (MO). Little Boca, Chiriqui Lagoon, Wedel 2508 (F, MO). BARRO COLORADO ISLAND: Croat 6235, 16583 (both MO); Shattuck 1115 (F, MO). CANAL ZONE: Cerro Gordo, Road 15 B, Croat 15125 (F, MO). Fort Sherman, Croat 15361 (MO). Madden Dam, Dwyer & Lallathin 8822 (MO). Curundu, Tyson & Blum 4458 (MO). cocLE: Cerro Gaital Caracoral, Dwyer & Correa 8882 (MO). COLON: N side of Río Guanche, V5 mi upstream from Puerto Pilon and Portobelo Road, Nee 7/04 (MO). Palmas Bellas, Thoms 36 (MO). DARIEN: Rio Pirre, Croat & Porter 15532 (MO). Rio Chucunaque and Rio Canclones, Duke 5114 (MO). Rio Chico, Duke 5217 (MO). Paya, Stern et al. 176, 222 (both MO). El Real, Stern et al. 611 (MO). PANAMA: Camino de Cruces, Caceres 16 (MO). Cerro Campana, 2 mi above m Highway, Croat 12045 (MO). Tocumen, Dwyer s.n. (MO). Rio Tocumen, Standley 26604 (G). s BLAS: Mulatuppu (Rio Ibedi), Duke 8466 (MO). Mai nland opposite Ailigandi, Dwyer 6838 (MO). Río Mulatuppu, Elias 1729 (GH, MO, US). Mainland opposite Ailigandi, Lewis et al. 156 (F, GH, K, MO). 55. Psychotria microdon (DC.) Urban, Symb. Antil. 9: 539. 1928. Rondeletia microdon DC., neci = 408. 1830. TIRE: e De la Ossa, not se Psyc podes roo Sesse & Moc., Fl. Mex., ed. 1894. TYPE: Vicinity Put de la Aquadilla, . Sesse & Mocino, ah seen. Shrubs or trees to 5 m tall, the branchlets terete, rigid, smooth, often drying white or olive white, disposed at right angles to the twigs, the nodes often crowd- ed, the leaf-bearing nodes swollen. Leaves oblong or obovate, 5-9 cm long (or longer), 2-4 cm wide, obtuse to deltoid at the apex, acute to attenuate acute at the base, the costa prominent above and beneath, the lateral veins 5—6, arcuate, slender; petioles slender, to 1.5 cm long; stipules connate, persistent, triangular, to 2 mm long, mucronate or briefly bilobate, puberulent or glabrate outside, setiferous at the base within. Inflorescences terminal, cymose corymbose, not exceeding the uppermost leaves, the peduncle absent and then with 3-5 ascending branches to 7(-9) cm long, or pedunculate, the peduncle to 5 cm long, slender, ca. 0.12 cm wide, terminating in (2—)5 radially disposed branches, glabrous or puberulent, the flowers few; bracts ovate triangular, to 1.5 mm long, with ap- pressed hairs outside and a large gland at the base within. Flowers subsessile, distylous, the hypanthium short, the calycine cup urceolate, inflated, scarious to carnose, glabrous and eglandular inside, minutely puberulent outside, the teeth scarcely evident or the margin merely undulate; corolla cylindrical, to 8 mm long (in bud), somewhat constricted toward the base, petaloid, glabrous inside and out, the tube oblong subrotund or ovate oblong, 3-4 mm long, obtuse, venose, occasionally fleshy; stamens 4-5(-7), the anthers oblong, 1.8-2.2 mm long, the filaments short, attached above the middle of the tube; style slender, 1.0-2.5 mm long, the stigmas sagittate or subulate, longer than the style, the ovarian disc compressed rotund, ca. 0.3 mm long, ca. 1 mm in diam. Fruits subrotund, to 6 398 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 QSP Vo O EO Q © ( yx T 5 P 6759 7 M O og e Q `° Q Ó Q o o J FIGURE 75. Psychotria mima Standley.—A. Habit (x !4).—B. Flower (x3).—C. Flower opened to show interior (x3).—D. Fruit (x5). [After Dwyer & Gauger 7368.] mm long, to 5 mm wide, glabrous, the pyrenes with the ventral face (endocarp) plane, the dorsal face with a single rib. Psychotria microdon ranges from Mexico south to Ecuador. Its range includes the Guianas and the West Indies. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 399 CANAL ZONE: Madden Dam, Dwyer & Lallathin 8809 (MO); Ebinger 868 (MO); Hayden 72 (MO). Salamanca Hydrographic Station, Rio Pequeni, Woodson et al. 1565 (F, MO). cHiRIQUÍ: 2 mi SW Puerto Armuelles, Croat 22456 (MO). cocLÉ: 1-5 mi S of Anton, Tyson & Blum 2561 (MO). DARIEN: Río Tuquesa below Quebrada Venado, Bristan 1069 (MO). Río Pirre, Croat & Porter 15523 (MO). o Camp called Charco Peje, Mori 7103 (MO). Yaviza, Río Chucunaque, Stern et al. 92 (MO). El Real, Río Tuira, Stern et al. 799 (MO). Los SANTOS: 1-2 mi W of Candelaria, Duke 12436 (MO). Pedasi, Río Purio, Dwyer 2504 (MO). PANAMÁ: Trapiche, Allen 2623 (F). Panama City, Bartlett & Lasser 16332 (MO). Nueva Gorgona Beach, Duke 4531 (MO). Isla Tabogilla, Duke 5881 (MO). Taboga Island, Dwyer 2181 A (MO); Gentry 5718 (MO). Tabogilla Island, across from N shore of Punta Chame, Mori & Kallunki 5764 (MO). Taboga Island, Pittier 3528, 3531 (both MO). Chepo, Pittier 4561 (MO). Matias Hernandez, Pittier 6713 (MO). Old Panama, Riley 149 (MO, US). Bella Vista, Standley 25349 (US). Taboga Island, Standley 27067 (US). Matías Hernandez, Standley 31993 (MO). Saboga Island, Tyson 5584 (MO). Pacheca jog a son 5608 (MO). I mi E of El Llano, Tyson 6831 (MO). Taboga Island, Woodson et al. 1471 (F 56. Psychotria mima Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 204. 40. TYPE: Brazil, L. B. Smith 1878 (F, holotype).—Fic. 75 Trees to 5 m tall, the branchlets slender, terete, glabrate to puberulent. Leaves oblong to ovate oblong, 6.0-19.5 cm long, 2.0-8.5 cm wide, deltoid to obtuse at the apex, acuminate, the acumen to 1.7 cm long, widely cuneate to subobtuse at the base, occasionally inequilateral, the costa immersed or somewhat prominu- lous above, prominulous beneath, smooth, the lateral veins ca. 10, the middle veins to 2 cm apart, arcuate ascending, often with parallel intermediate evanes- cent veins, thinly coriaceous, usually drying gray or tan gray, glabrate but often rufous puberulent on the costa or on the lateral veins; petioles to 2.5 cm long, to 0.18 cm wide; stipules briefly or almost completely connate, cylindrical, to 0.8 cm long, each part 2-lobed at the apex, the lobes obtuse, each shorter than the sheath, stiffly scarious, glabrous or glabrescent. Inflorescences terminal, pyram- idal paniculate, to 45 cm long, the peduncles absent or elongate, to 0.2 cm wide at the base, sinuous, glabrous, the branches opposite, spreading or reflexed, well spaced, the lowermost to 10.5 cm long, the secondary branches floriferous, paired, opposite, well spaced, at right angles to the secondary branches, the bracts subulate, to 0.5 cm long, reflexed, persistent, the cymules few flowered, spreading. Flowers with the hypanthium rotund, to 1.3 mm long, puberulent, the calycine cup ca. 0.3 mm long, glabrous and with minute appressed rotund glands within, the calycine teeth 5, subequal, deltoid, to 0.3 mm long, acute, puberulent marginally and without; corolla white or cream maroon, the tube linear cylindri- cal, to 12 mm long, ca. 2 mm wide, glabrous outside and in except villosulose at the anther attachment position, the lobes 5, narrow oblong, cucullate, glabrous, petaloid; stamens 5, the anthers oblong, ca. 1.7 mm long, occasionally minutely apiculate, dorsifixed, the filaments short, attached near the base of the tube; stigmas 2, linear, to 2 mm long, the style compressed, 4-7 mm long. Fruits (fragmentary) subrotund, to 3 mm in diam., smooth, black, lightly ribbed. Psychotria mima occurs in Panama, Colombia and Brazil. The inflorescence in the living plant is blue or purple. One Colombian collection (Forero 797 [MO]) has excellent fruits which are compressed rotund. The inflorescence is epedun- culate or one may consider the tip of the twiglet as bearing 3 discrete peduncles, the median to 12.5 cm long 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 BOCAS DEL TORO: Between Buena Vista coffee finca and Cerro Pilón cloud forest, Kirkbride & Duke 678 (MO). coLów: Santa Rita Ridge, 19 km from paved highway, Dwyer 8574 (MO). DARIEN: Demartaganala, Camp Summit, Duke 15827 (MO). Camp Summit near Darien-San Blas border, Oliver et al. 3694 (COL, MO, UC). PANAMA: Cerro Jefe, Duke 9401 (MO); Dwyer & Gauger 7368 (MO); Dwyer & Gentry 9469 (MO). Gorgas Mem. Lab., 5-10 km NE of Altos de Pacora, Mori & Kallunki 3423 (MO). Cerro Jefe, Tyson et al. 3246 (MO). VERAGUAS: 5 mi W of Santa Fe, Escuela Agricola Alto Piedra, 800-1200 m, Croat 23077 (MO). 5 mi NW of Santa Fe, 700-1200 m, Croat 23156 (MO). Valley of Río Dos Bocas, 15.6 km NW of Santa Fe, Croat 27787 (MO). NW of Santa Fe, 1.5 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 5309 (MO). COLOMBIA: CHOCO: Region of Rio Pichma, Waunana, 100 m, Forero 797 (MO). 57. Psychotria morii Dwyer.1!% TYPE: Panama, Mori 3139 (MO, holotype). Shrub to 1.25 m tall, the stems smooth, glabrous, perhaps succulent, the pith porate, the nodes well spaced. Leaves large, elliptic oblong, to 31 cm long, to 14 cm wide, acuminate, the acumen triangular, to 1 cm long, ultimately acute, widely deltoid or subobtuse, basally attenuate acute or slightly inequilateral, the costa prominulous and grooved above, prominulous beneath, the lateral veins ca. 15, arcuate, prominulous on both sides, the marginal vein absent, the intervenal areas spreading reticulate, membranous, discolorous, dark green above when dry, the hairs curled, subulate, often with the basal cell red; petioles 8-11 cm long, to 0.25 cm wide, the hairs dense, webby; stipules connate, swollen and collar shaped, to 0.6 cm long, truncate, scarious, glabrescent, the hairs few. Inflorescences axillary, to 8 cm long, to 3 cm wide, the peduncle to 4 cm long, the branches 3, often radiate, these to 1.5 cm long, the cymes congested; bracts to 1 cm long, concave, the margins scarious, sparsely ciliolate, except the margins strongly ciliolate. Flowers with the hypanthium somewhat turbinate, 1.2 mm long, gla- brous, the calycine cup 0.5 mm long, petaloid, densely ciliate outside, glabrous within, the cystoliths dense, the teeth 5, triangular, 0.5 mm long, acute with a thin veil located between the teeth, the hairs of the sheath many, erect; corolla white ca. 2.5 mm long, carnose, ciliate outside, glabrous within, the lobes 5, somewhat longer than the tube, cucullate; stamens 5, the anthers oblong, 1.8 mm long, obtuse at the apex, sagittate at the base, the filaments short, attached below the middle; ovarian disc barrel shaped, 0.3 mm long, the style 0.7 mm long, the stigmas 2, linear oblong, scarcely wider than the style. Fruits rotund, ca. 6 mm in diam., red, the fleshy part, when dry, punctate, glabrous, the pyrenes glabrous, not costate. Psychotria morii is known only from Panama. It is named in honor of Dr. Scott Mori, distinguished collector of tropical American plants and authority on 1% Psychotria morii Dwyer, spec. nov. Folia magna elliptico-oblonga, ad 31 cm longa, ad 14 cm lata, ance lato-d deltoidea vel subobtus use, acumin nata, basi attenuato-acuta venis lateralibus ca. 15, Flores cupula calycis ca. 0.5 mm s denso-ciliata intus glabra dentibus ca. 0.5 mm longis, vagina diaphana inter dentes sein ciliis. n multis erectis; corolla ca. 2.5 mm longis, lobis cucullatis; antheris ca. 1.8 mm longis. Fructus rotundi, ca. 0.6 cm diam., pyrenis laevibus non- costatis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 401 the Lecythidaceae. The new species is distinguished by its relatively large collar- shaped stipules and by the thin interdental veil of the calyx. The smoothness of the pyrenes is also noteworthy. VERAGUAS: N of Santa Fe, 6.5 km from Agricultural School, Mori 3139 (MO). 58. Psychotria mortoniana Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 355. 1938. TYPE: Costa Rica, Skutch 2697 (F, holotype; NY, isotype). Shrubs to 6 m tall, the branchlets 4-angled, glabrescent or villosulose, the nodes well spaced. Leaves oblong or elliptic, 8.5-27.0 cm long, 3-11 cm wide, deltoid at the apex, acuminate, the acumen to 2.5 cm long, usually falcate, acute to attenuate acute at the base, the costa subplane above, prominent beneath, to .2 cm wide proximally, the lateral veins 22-25, arcuate, to 1 cm apart, stiffly chartaceous, often discolorous, darker green or brown above, yellow green be- neath, villosulose above, especially on the costa and veins, subvelutinous be- neath, the costa and veins barbate; petioles slender or stout, to 2 cm long, to 0.2 cm wide, densely villosulose; stipules free, oblong rotund, 6-12 mm long, with 2 proximate erect awns at the apex, to % the length of the body, the body appressed, scarious, velutinous. Inflorescences terminal, pyramidal paniculate, to 12.5 cm long, the peduncle ca. 4 cm long, the rachis with about 3 pairs of well spaced, opposite branches, the bracts spreading or deflexed, subulate, 0.7-1.5 cm long. Flowers sessile, the hypanthium short, the calycine cup short, eglan- dular within or with a minute erect gland at the junction of 2 lobes, the lobes 5, unequal, triangular to oblong, to 1 mm long, petaloid, carnose, venose, marginally pubescent; corolla white, the tube cylindrical, somewhat inflated, ca. 2.2 mm long, petaloid, glabrous, the lobes 5, erect, oblong, venose, somewhat cucullate, to 1.5 mm long, obtuse; stamens 5, the anthers included in the tube, oblong, ca. 1 mm long, the filaments short, attached just below the middle of the tube; style slender, 2.8—4.5 mm long, the stigmas 2, oblong, often plumose, 0.4—0.7 mm long. Fruits sessile, enclosed by 1 or 2 bracteoles, oblong, occasionally subrotund, ca. 3 mm long, white hirsute, obtusely ribbed. Psychotria mortoniana is known from Costa Rica and Panama. CHIRIQUÍ: W of Cerro Pando, ca. 6000 ft, D'Arcy & D'Arcy 6663 (MO). Los sANTOS: Loma Prieta, Cerro Grande, 2400-2800 ft, Lewis et al. 2218 (MO). 59. Psychotria nervosa Swartz, Prodr. 43. 1788. TYPE: Jamaica, Swartz, not seen. P. rufescens H.B.K., Nov. Gen. Sp. Pl.3: 364. 1818. LECTOTYPE: Venezuela, Humboldt & Bonpland, seen. P. hirta H.B.K., Nov. Gen. Sp. Pl. 3: 364. 1818. rype: Colombia, Humboldt & Bonpland, not seen. 2 822. P. lanceolata Nutt., Amer. Jour. Sci. Ser. 1, .] TYPE: East Florida, no collector given. P. oligotricha DC., Prodr. 4: 514. 1830. TYPE: Jamaica, Porto Rico, dr d no collectors given. P. portoricensis r 515. 1830. TYPE: Porto Rico, Bertero, not see DC., . 4: ; Myrstiphyllum undatum (Jacq.) Hitchc., Ann. Rep. Missouri Bot. Gard. 4: 95. Psychotria nervosa TE puedas B.K. ) op Mem. New York ih Gard. 23: 480. 1972. ot. Gard. 23: 481. 402 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Shrubs to 5 m tall, the branchlets terete, pubescent to glabrate, the nodes well spaced or occasionally crowded at the apex of the twiglets. Leaves lanceolate or oblanceolate, often narrowly so, occasionally falcate, 4-15 cm long, 1.8-7.0 cm wide, tapering acutely or obtusely into a distinct or indistinct acumen, the acumen to 2 cm long, ultimately acute or obtuse, basally acute, attenuate acute or cuneate, the costa plane or grooved above, subprominent beneath, the lateral veins 10— 15, arcuate, occasionally strict but arcuate toward the margin, the submarginal undulate vein conspicuous, chartaceous, often drying reddish, ferrugineous vil- losulose or villose above and beneath, occasionally glabrate above, often with bases of deciduous hairs giving a punctate appearance; petioles either absent or to 1.5 cm long; stipules connate, at least at the very base, often becoming free from the node as a movable cylinder, each part oblong rotund, 0.8-1.5 cm long, obtuse, venose, pilose. Inflorescences terminal, occasionally pseudoterminal, solitary or ternately disposed, ferrugineous pubescent, umbelliform paniculate or conglomerate capitate, 1-5 cm long, to 7 cm wide, either epedunculate with several radiate branches or pedunculate, the peduncles 0.5-1.3 cm long, occa- sionally the peduncle with one cymule or a rachis with 2-3 pairs of opposite branches terminated by cymules. Flowers with the hypanthium ca. 0.5 mm long, the calycine cup saucer shaped and expanded, glabrous to puberulent outside, glabrous and eglandular within, to 0.5 mm long, the margin of the cup undulate or with triangular teeth, to 0.35 mm long; corolla white, the tube narrow at the base, somewhat campanulate above, glabrous to puberulent outside, villosulose near the filament attachment position within, the lobes 5, ovate oblong, almost equal to the tube, villosulose at base; stamens 5, the anthers oblong, to 0.5 mm long, the filaments slender, to 2 mm long, attached at the mouth; style slender, ca. | mm long, the stigmas upright, ca. 0.4 mm long. Fruits sessile, yellow orange to red, ovoid oblong to oblong, to 0.7 cm long, about 2 times longer than wide, conspicuously sulcate, glabrous to rufous pubescent, the remnants of the calyx scarcely measurable. Psychotria nervosa occurs in subtropical Florida, the Bahamas, the West Indies, in Central America and northern South America. CANAL ZONE: Las Cruces Trail, Bartlett & Lasser 16444 (MO). Albrook, Blum 1213 (MO). Cerro Viejo on K 16 C, Blum 1221 (MO). C-15 Road, Blum & Dwyer 2459 (MO). Fort Sherman, Croat 12515 (MO). Fort Kobbe, Duke 4728 (MO). ids Doug emer Duke & Mussell 6640 (MO). Madden Dam, Dwyer 8394, 8813 (both MO). Fort Sher eeu & Robyns 152 (MO). Albrook, Dwyer 6638, 6706, 7315 (all MO); Hayden 66 (MO). Las C ail, Hunter Fi Allen 699 (MO). pos pM road to Fort Sherman from Gatün Locks, Mori & alui 2682 (MO). C-29 Road, Nee 8993 (MO). Madden Dam, Porter et al. 4052 io SCZ, UC). Curundu, Vd" 4176 (MO). Fort Agi Tyson & Dwyer 1183, 1193 (both MO). Pipeline Road, Witherspo oon & Witherspoon 8397 (MO). Salamanca drin c etd Station, Menu et al. 1588 (MO). BARRO COLORADO ISLAND: Bailey & Bailey 63 (F). MO) COC n, Dwyer 8 ( . Llano Bonito, Seibert 520 (MO). Las Margaritas and El Valle, mu et T 1745 (F, ip COLÓN: Garotte, D'Arcy 4002 (MO). Isla Grande, D'Arcy 4014 (MO). Río Guanche, ca. 3 km upriver from bridge, road to Portobelo, Mori & Kallunki 5213 (MO). DARIÉN: Yaviza, Río P Y28358 Stern et al. 145 (MO). PANAMA: Río Pacora, Bartlett & Lasser 16957 (MO). Calzada Larga, Croat 12395 (MO). San José Island, Duke 12540 (MO). Pedro Gonzales (Perlas Archipelago), Dwyer 1725 (MO). Santa Clara, Dwyer 4264 (MO). Cermeno, Dwyer 6630 (MO). Cerro Azul, Dwyer 7128 (MO). San Jose Island, Johnston 142 (MO). SAN BLAS: Ailigandi, Dwyer 6815 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 403 60. Psychotria nubiphila Dwyer.!° TYPE: Panama, Croat 15990 (MO, holotype). Trees to 4 m tall, the ultimate branchlets glabrous, the nodes well spaced, turgid with the bases of the persistent stipules. Leaves often inequilateral, elliptic, 10-15 cm long, 5-7 cm wide, deltoid toward the apex, basally acute to attenuate acute or cuneate, the costa prominulous above and beneath, the lateral veins 10— 12, arcuate, the intervenal areas smooth, the ultimate venules evanescent, sub- papyraceous, concolorous, glabrous above, puberulous on the costa and with the hairs crowded in the vein axils beneath; petioles to 3.5 cm long, to 0.25 mm wide, rigid but at times reflexed; stipules connate, the sheath cup shaped, to 1 mm long, glabrous except red glandular punctate, truncate, the margin erose. /nflores- cences terminal, solitary, cymose paniculate, to 13 cm long, to 9 cm wide, the peduncle 1.5-3.5 cm long, 0.3 cm wide, twisted, glabrous, the lowermost branch- es opposite, strongly ascending, to 6 cm long, the upper branches 3-4-jugate, opposite, well spaced. Flowers with the hypanthium turbinate, 1 mm long, red when dry, the calycine cup short, turgid, to 0.5 mm long, the teeth irregular, ciliolate; corolla green, the tube cylindrical, 3.5 mm long, rigidly petaloid, glabrate outside, densely barbate near the mouth within, the lobes 5, reflexed, triangular, 3.5 mm long; stamens (4—)5, the anthers oblong, | mm long, obtuse, the filaments unequal; style 3 mm long, the stigmas short. Fruits not seen. Psychotria nubiphila is known only from Panama. CHIRIQUÍ: NW side of Cerro Pando, Croat 15990 (MO). 61. Psychotria oerestediana Standley, Contr. U.S. Natl. Herb. 23: 1390. 1926. TYPE: not designated. Mapouria obovata Oerst., Biol. Centr. Amer., Bot. 17. 1863. TYPE: not s Psychotria quinqueradiata Polak., Linnaea 41: 570. 1877. TyPE: Costa "Rica, Polakowsky 94 (B, olotype; US, photo). Psychotria morae Polak., Linnaea 41: 470. 1877. rype: Costa Rica, Polakowsky 171, not seen. Psyc hotria obovata (Oerst.) ERR Biol. Centr. Amer., Bot. 2: 50. 1881. Not P. obovata Ruiz & Pavón (1799). TYPE: not s Shrubs 1.0—1.5 m tall, the branchlets smooth, glabrous, the nodes well spaced. Leaves elliptic, obovate elliptic or oblong rotund, 4.0-11.5 cm long, 1.8-6.0 cm wide, widely deltoid to rounded or obtuse at the apex, acuminate, the acumen vague or obvious, usually widely triangular, ultimately obtuse, basally cuneate, attenuate acuminate to acute, the costa prominulous above, prominent beneath, the lateral veins 8-12, at first strict, arcuate toward the margin, the submarginal vein near or far from the margin, occasionally with minute **domatia"' in the axils 105 Psychotria "n Dwyer, spec. nov. Arbores ad 4 m altae ramulis ultime glabris. Folia elliptica 10-15 cm longa, 5-7 cm lata, venis lateralibus 10-12, subpapyiaosa supra glabra in costa infra puberula ciliis in axillis confertis; petiolis ad 3.5 cm longis; stipulis connatis ad 1 mm longis, glabris praeter rubro-granuloso-punctatis truncatis Te pe Inflorescentiae terminales solitariae cymoso-paniculatae, ad 13 cm longae, ad 9 c e, pedunculo 1.5-3.5 cm longo. Pi cupula calycis ca. 0.5 mm SA dentibus brevissimis kp ton ciliolatis; corolla tubo ca. 3.5 m longo; antheris ca. | mm lon 404 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 of the lateral veins beneath, stiffly papyraceous, smooth, glabrous, drying gray red or gray; petioles 0.2-1.0 cm long (longer in extra Panama material); stipules deciduous, connate, the sheath ca. 2 mm long, each part with 2 lanceolate lobes longer than the sheath, glabrous. Inflorescences terminal, 1-3 cm long, usually | cm long, the flowers aggregated into subrotund heads, the peduncle absent or if present to 1.8 cm long, the flowers on scarcely measurable branches (in extra Panama material to 3.5 cm long and radially disposed); bracts and bracteoles deciduous. Flowers sessile, drying red or red black; hypanthium 0.5 mm long, glabrous, the calycine cup cupuliform, shorter than the hypanthium, truncate, the teeth scarcely evident, presumably 5, glabrous; corolla white, the tube cylin- drical, to 4 mm long, to 1 mm wide, glabrate outside, villose at and above the filament attachment position within, petaloid, the lobes 5, short or to 2 mm long (extra Panama material), glabrate; anthers oblong, 1.0-1.3 mm long, the filaments to 2.3 mm long, attached near the middle of the tube; ovarian disc black, usually longer than the calycine cup, the style to 2.3 mm long, the stigmas narrowly oblong, to 1.3 mm long. Fruits subsessile, oblong, to 6 mm long, to 4 mm wide, obtusely ribbed, glabrous, drying black. Psychotria oerestediana ranges from Mexico to Panama. Unfortunately I have seen only 4 collections from Panama. BOCAS DEL TORO: Milla 71⁄2, railroad 10 km NW of Almirante, Nee & Hansen 14085 (MO). CANAL ZONE: Chagres, Fendler 59 (MO). coct É: El Valle, N rim, Allen & Alston 1853 (MO). Forest behind Club Campestre, El Valle, Duke 13246 (MO). 62. Psychotria officinalis (Aubl.) Raeuschel ex Sandw., Kew Bull. 1931; 473. 1931. Nonatelia officinalis Aubl., Hist. Pl. Guiane 1: 182. 1775. type: Cayenne, French Guiana. Aublet, not seen. Psychotria involucrata Willd. Sp. Pl. 1. 966. 1798. Not Swartz 1788. Shrubs or trees to 4.5 m tall, glabrous, the nodes well spaced. Leaves elliptic, oblong elliptic or ovate elliptic, 4-22 cm long, 1.5-11.0 cm wide, deltoid, acute or occasionally tapering somewhat obtusely at the apex, acuminate, the acumen to 3 cm long, acute, cuneate or obtuse at the base, the lateral veins 6-11, arcuate, the submarginal vein close to the margin, stiffly membranous to papyraceous, 4— 22 cm long, 1.5-11.0 cm wide, concolorous, usually drying green, often lustrous above, glabrous above and usually so beneath; petioles 0.5-2.2 cm long, stiff, glabrous; stipules small, free, subrotund to cup shaped, 0.5-5.0 mm long, the 2 awns shorter than or to 4 times longer than the body, glabrate. Inflorescences terminal, occasionally in the uppermost axils, usually glabrous, usually purple, occasionally puberulent, often subrotund or oblong, paniculate, to 7.5 cm long, to 4.5 cm wide; peduncle 1.3-3.2(—5.0) cm long, the lowermost branches often 3-5, radiate and giving an umbellate appearance, the branches few, usually op- posite; bracts of the floral branchlets 1—3, usually blue or purple, the largest bract subulate, involute, 0.4-1.3 cm long, to 1 mm wide. Flowers sessile. soon going to fruit, the calyx and hypanthium 1.5 mm long, puberulent, the calycine cup short, with 5 ovate or subrotund lobes, 0.2-0.3 mm long, often wider than long, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 405 with a gland at each interdental position; corolla white, the tube and lobes 5 mm long, the tube ca. 1.0 mm long, partly glabrous and partly pilulose outside, hairy within, the lobes ovate, ca. 2 mm long, glabrous within except basally; stamens , the anthers 0.8-1.2 mm long, the filaments to 2.5 mm long; ovarian disc lobed, the style to 4 mm long. Fruits obovoid or subrotund, to 3.5 mm wide, glabrous or occasionally with a few hairs, costate. Psychotria officinalis occurs in Middle America and northern South America. AS DEL TORO: Between Quebrada Huron and Puerto Palenque, Kirkbride & Duke 504 (MO). baer anu Peak, N of Río Terebé, Kirkbride & Duke 580 (MO). Between Quebrada Menaco and Water Valley, Wedel 1608 (MO). Fish ede Wedel 2196 A (MO). CANAL ZONE: Hill C-6, Fort Sherman, Duke 4413 (MO). 1 mi N of Summit Garden, Tyson & Blum 1494 (MO). Fort Sherman, Tyson & Blum 1653 (MO). CocLE: La enti (oda 13328 (MO). Cerro Pilón, ca. 3000 ft, Duke & adipe 15001 (MO). La Mesa, Dwyer 8311 (MO). Cerro Pilón, Dwyer 8346 (MO). La Mesa, 8.5 mi m Club Campestre, El Valle, Dwyer 10520 (MO). El Valle, La Mesa, Dwyer 11907 (MO). Cerro Gaital and Caracoral, Dwyer & Correa 8895, 8904 (both MO). Cerro Pilón, Dwyer = or dee 8606, 8620 (both MO). El Cope, El Potroso, Folsom & Lantz 1882 (MO). Alto de Calva of E Pa Folsom & Robinson 2348 (MO). La Mesa, N of El Valle, ca. 1000 m, En 6852 PAN La . Kennedy et al. 2084 (MO). 4 km N of El Valle, Nee & Dwyer 9216 (MO). La Mesa, Nee & Hale 9619 (MO). Cerro Pilón, above El Valle, Porter et al. 4397, 4415, 4474 (all MO). DARIÉN: Between Río Balsa and Río Areti, Duke 8748 (MO). Cativo swamp near Tusijuanda, Duke 13523 (MO). Puerto St. Dorothea, Dwyer 2223, 2251 (both MO). Cana-Cuasi Trail between Cerro Campa- miento and La Escalera, Kirkbride & Duke 1338 (MO). Cerro Pirre, 300-700 m, Mori & Kallunki 5396 (MO). PANAMÁ: Cerro Jefe, road to Altos de Pacora, Croat 22650 (MO). Panama Viejo, Duke 5728 (MO). 2 mi upstream from Río Sancanti, near Piria, Duke 14377 (MO). Cerro Jefe, to 3100 ft, Dwyer 9431 (MO). 9.4 km N of Goofy ws m et al. 1983 (MO). Cerro Jefe, Gentry 3497 (MO); Gentry et al. 3428, 3429, 3461 (all MO). o Campana, Lewis et al. 1907 (GH, MO, UC, US). Chimán, Lewis et al. 3261 (MO). 6.5 km LE ud N of Cerro Azul, Nee 9306 (MO). Road past Cerro Azul, Mori & Kallunki 2189 (MO). Cerro Azul. Tyson 2139 (MO). Cerro Jefe, Tyson et al. 3320 (MO); Wilbur et A 11339 (MO). SAN BLAS: 3—4 hr by foot up Río Mulutupu, Kirkbride 218 (MO). VERAGUAS: 5 mi W of Santa Fé, road past Escuela Agricola Alto Piedra, 800-1200 m, Croat 23013 (MO). Road M ed Escuela Agricola and Alto Piedra and Río Dos Bocas, Croat 25946 (MO). Valley of Río Dos Bocas, between Alto Piedra and Calovebora, Croat 27457 (MO). Valley of Río Dos Bocas, 15.6 km NW of Santa Fé, Croat 27638, 27640 (both MO). Cerro Tuté, Dwyer s.n. (MO). 6-7 km W of Santa Fé, 2900 ft, Nee 9712 (MO). COLOMBIA: CHOCO: Curiche and Alto Curiche, Duke 9604 (MO). 63. innu olgae Dwyer & Hayden, Ann. Missouri Bot. Gard. 55: 42. 1968. PE: Panama, Dwyer et al. 8193 (MO, holotype). Trees or shrubs to 8 m tall, the branchlets terete, ultimately subplane, smooth, glabrous, usually the nodes turgid, well spaced below the tips, crowded termi- nally. Leaves strongly ascending, obovate elliptic, oblong rotund, subrotund or elliptic, 4.5-11.0 cm long, 1.5-5.0 cm wide, rounded, obtuse to deltoid at the apex, cuneate at the base, the costa prominulous above, prominulous to promi- nent beneath proximally, the lateral veins 5-8, arcuate, 0.8-1.5 cm apart, eva- nescent above, prominulous beneath, thickly coriaceous, lustrous, glabrous, oc- casionally drying yellow, the margins conspicuously revolute; petioles absent or to 1.5 cm long, lignose, glabrous; stipules often persistent, ?free, broadly cup shaped 0.2-1.5 cm long, much wider than long, usually with 2 mammiform bulges at each side, coriaceous, glabrous, fimbriate ciliate within. Inflorescences ter- minal, glabrous, to 11 cm long, to 5 cm wide, cymose paniculate, the peduncle 406 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 to 4.5 cm long, stiff, the branches disposed as 4-5 pairs, the cymules few flow- ered, terminal; bracts and bracteoles ovate subulate, ca. 2.5 mm long. Flowers sessile or short pedicellate; hypanthium 1.3 mm long, the calycine cup short with 1-2 minute glands at the junction of the lobes, the lobes 5, triangular, ca. 0.5 mm long, the calyx spread at anthesis; corolla white, to 5 mm long, the tube short, somewhat expanded basally, puberulent or glabrous outside, densely barbate within above the middle, the lobes 5, narrowly oblong, 2-4 mm long, cucullate; stamens 5, the anthers narrowly oblong, 2.5-3.0 mm long, the filaments short, 1 mm long; style thickly cylindrical to linear, 4-11 mm long, the stigmas small. Fruits obovate oblong, 1-2 cm long, | cm wide, obtuse at the apex, acute basally, carnose, red, 10—12-ribbed. Psychotria olgae is endemic to Panama. Its thick coriaceous leaves and large fruit distinguish it from all other Psychotria of Middle America. ANAMA: Cerro Jefe, Dwyer 7288 (MO). Cerro Jefe and La Eneida, Dwyer et al. 8193 (MO). Cerro Jefe, on top near antenna, Folsom & Harp 1368 (MO). Cerro Jefe, ca. 1000 m, Mori 7133 O). 64. Psychotria orchidearum Standley, J. Wash. Acad. Sci. 18: 276. 1928. TvPE: Costa Rica, Standley & Valerio 50863 (US, holotype). 64a. Psychotria orchidearum Standley ssp. orchidearum. Epiphytic subshrub, to 30 cm, the branchlets terete, smooth, glabrous, the nodes well spaced, occasionally swollen. Leaves oblong, obovate oblong, oblong rotund to elliptic, 1.8-8.0 cm long, 0.8-3.5(—4.5) cm wide, often inequilateral, tapering obtusely, deltoid or acute at the apex, usually conspicuously shortly acuminate, acute to obtuse occasionally rounded at the base, often inequilateral, the costa subplane above, prominulous beneath, the lateral veins 4—6(—7), eva- nescent or obvious, strongly arcuate, the intervenal areas indistinct, succulent, drying thinly to thickly coriaceous, concolorous or discolorous, smooth or wrin- kled on drying, glabrous; petioles to 1 cm long, slender to stout, 0.15-0.32 cm wide, glabrous; stipules annular, cylindrical, 3-4 mm long, usually slightly wider than long, stiffly scarious, smooth, glabrous. /nflorescences bright red, terminal or occasionally axillary, glabrous, to 6 cm long, occasionally slightly wider than long, paniculate when the branches are paired and arising along the rachis or subcorymbiform when the branches arise verticillately from the apex of the pe- duncle, the peduncles to 2.5 cm long, to 1.2 mm wide, the branches strongly ascending or reflexed, to 2 cm long, branching dichotomously or terminating in several verticillate branches, the flowers few. Flowers subsessile or with pedicels to 2.5 mm long; hypanthium red, pubescent and minutely granulose, the calycine cup short or absent, usually with 4 triangular teeth, to 0.4 mm long, acute; corolla white or magenta, the tube narrowly cylindrical, to 3 mm long, glabrous outside, villose within at the point of filament attachment, the lobes 4, oblong, ca. 0.35 mm long, obtuse; stamens 4, the anthers oblong, rotund, ca. 0.6 mm long, the filaments short; stigmas 4, the styles linear, the ovary 4-celled, the ovules 1 per 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 407 locule, attached basally. Fruits orange red, fleshy, subrotund to obovate-rotund, ca. 3-4 mm in diameter, truncate, glabrous, usually appearing 8-sulcate (4 py- renes) on drying. Psychotria orchidearum is known from Costa Rica and Panama. The use of the specific name orchidearum is a stop-gap measure, as the taxonomy of the epiphytic Psychotria is in disarray. The 4-carpellate nature of the pistil is note- worthy. ANAL ZONE: Between Fort San Lorenzo and Fort Sherman near Pavon Road junction with Road 82 (S 2), Mori & Kallunki 2722 (MO). PANAMÁ: 10 km N of El Llano on Carti Road, ca. 400 m, Busey 919 (MO). Cerro Jefe, ca. 950 m, Elias & Hayden 1804 (GH, MO). 16 km above Panamerican Highway, El Llano to Carti Tupile, Kennedy et al. 2431 (MO). La Eneida, Luteyn & Foster 1123 (MO). Cerro Jefe, ca. 950 m, Tyson et al. 3199, 3400, 4381 (all MO); Witherspoon et al. 8274 (MO). 64b. Psychotria orchidearum subsp. persedens Dwyer.1% TYPE: Panama, Tyson 5331 (MO, holotype). Similar to Psychotria orchidearum subsp. orchidearum except the leaves tend to be less than 2!4 times longer than wide, more distinctly acuminate at the apex, and if drying yellow, not strongly wrinkled. There is some overlapping between material identified as subsp. orchidearum and the new subsp. The leaves of subsp. persedens are much more variable in shape: oblong, obovate oblong, oblong rotund to subrotund. In general, they do not have the ligulate or straplike appearance of subsp. orchidearum. The leaf blades may dry as to appear smooth: Folsom 4476, 4235; Duke & Elias 13757. The leaves may be very wide: Gentry et al. 3497 and Witherspoon et al. 8274. In Maas & Dressler 727 the leaves are subrotund. Whether the veins are con- spicuous or inconspicuous in dried leaves of subsp. persedens seems of little importance. The subspecific name ‘‘persedens’’ refers to the epiphytic habit of the plant, meaning ^'residing over a long period of time.” In stained sections of the leaf, fixed in the field in Formalin, Acetic Acid and Alcohol (FAA) and embedded in paraffin after treatment in tertiary butyl alcohol, I noted a distinct band of cells located near the center of the mesophyll. This was not observed in sections of several non-epiphytic species of Panamanian Psycho- tria. CHIRIQUÍ: San Felix, 800-1200 m, Croat 33429 (MO). Cerro Horqueta, Hagen & Hagen 2093 . Cerro Pando, ca. 2000 m, Mori & Bolten 7321 (MO). cocLÉ: Cerro Pajita, N of El Valle, Allen & E. 4181 (MO). El Valle, Road from Interamerican Highway, Correa 332 (MO). La Mesa, ca. 800 m, Croat 25393, 37446 (both MO). Top of El Petroso, 800 m, D'Arcy 11350 (MO). Cerro Pilon, Duke 12148 (MO). El Valle, Duke 13186 (MO). Elfin Forest, Cerro Caracoral, ca. 1000 m, Duke & Dwyer 15082 (MO). La Mesa, Dwyer 8261, 10539 (both MO). Cerro Gaital Caracoral, Dwyer & Correa 106 Psychotria orc din of subsp. persedens Dwyer, subsp. nov. Differt a P. orchidearum subsp. orchidearum foliis saepe lato-oblongis oblongo-rotundis ad subrotundis vulgo apice conspicue brevi- cuminatis in sicco sae n e tenui-coriaceis et laevibus; laminis foliorum si in sicco luteis et valde rugosis minus quam 2!2-plo longioribus quam latioribus. 408 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 8874 (MO). Cerro Pilon, Dwyer & Lallathin 8969 (MO). El Valle to La Mesa, Dwyer & Nee 11928 (MO). 7 mi N of El Cope, Folsom 3217, 4144 (both MO). Summit, Alto Clavario, 900 m, Folsom & Robinson 2414 (MO). Cerro Gaital Caracoral, Kirkbride 1108 A (MO). El Valle, Nee & Hale 9637 O); Wilbur et al. 11117 (MO). DARIEN: Cerro Pirre, Bristan 544, 589 (both MO). Gold Mining Camp, Headwaters of Rio Tuquesa, Croat 27259 (MO). Cerro Pirre, Croat 37893 (MO); Duke 6118 (MO): Duke & Elias 13757 (COL, MO, UC), 13833 (MO); Folsom 4235 (MO). Summit of PE Pirre, hg & Clewell 7155 (MO). Top of ge Mali, Gentry & Mori 13647, 1369] (both — W peak of Cerro Tacarcuna massif, 1800-1850 m, Gentry & Mori 13994 (MO). Cerro Tacarcu 1720 m, ei et al. 16899 (MO). posed Trail Econ Cerro Campamiento and La Escalera, Kirkbride & Duke 1296 (MO). PANAMA: Cerro Campana, Allen 1879 (MO). 10 mi N of El Llano, Busey 360 A (MO). Cerro Campana, Croat 14782, 17196 (both MO). Cerro Jefe, ca. 2900 ft, Dwyer et al. 3497, 7277 (both MO). Cerro Campana, Gentry 4945, ate — MO). El Llano, Gentry 5801 (MO). Cerro Campana, Kennedy et al. 2072, 2085 (both MO); Maas & Dressler 727 (MO); did & Kallunki 2467, 2484, 3580, 3603 (all MO). El Llano, Nee & Ds "0347 (MO). Cerro Campana, Porter et al. 4210 (MO, SCZ, UC), 4960 (MO, UC, VEN). Cerro Jefe, ca. 2900 ft, Porter et 5 5073 (MO). Cerro Campana, Tyson 5331 (MO). Cerro Jefe, Tvson et al. 3292, 3426, 4346 (all MO). SAN BLAS: 2-5 mi S of Mandinga Airport, Duke 14790 (MO). VERAGUAS: Cerro Tute above Santa Fe, Gentry 6271 (MO). 5 mi W of Santa Fe, Liesner 967 (MO). Cerro Tute, NW of Santa Fe, 2 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 5276 (MO) 65. Psychotria oresbia Dwyer.!% Type: Panama, Gentry & Mori 14150 (MO, ho- lotype) Trees to 6 m, the twiglets slender, smooth, glabrous, the nodes well spaced, swollen. Leaves oblong, 3-9 cm long, 2-4 cm wide, rounded or obtuse at the apex, acuminate, the acumen to | cm long, obtuse or rounded at the base, the costa slender, prominulous above and beneath, the lateral veins 8—10, slender, widely arcuate, the undulate submarginal vein well set in from the margin, the margin delicately callose, the intervenal area reticulate, smooth, stiffly charta- ceous, discolorous, glabrous; petioles slender, stiff, 0.8-1.0 cm long, scarcely alate; stipules tending to persist at the end of the twiglets, connate, cup shaped, the sheath to 2 mm long, toothless, coriaceous, with 2 triangular lobes shorter than the sheath, glabrous, sometimes with minute crustlike excrescences. Inflo- rescences terminal, racemose in appearance, to 4 cm long, to 3 cm wide, the peduncle 5 mm long, the rachis stiff, the 5 pairs of opposite branches arranged distichously, the uppermost about equal to the lowermost, to 1.5 cm long, diver- gent, strict, glabrous but with minute excrescences, each branch with 1—2 3- flowered cymules; bracts at the base of branches ovate, 2.2 mm long, acute, somewhat clasping, spreading or somewhat ciliolate. Flowers sessile or subses- sile, hypanthium 1 mm long, glabrous, drying black, the calycine cup expanded, 0.5 mm long, the margin undulate, the teeth scarcely measurable or widely oblong triangular, to 0.8 mm long, acute, carnose, glabrous; corolla cream colored, the tube (almost mature) subcampanulate, ca. 2 mm long, somewhat constricted above the base, ca. 0.8 mm wide at the base, glabrous outside, villosulose only at the base within, the lobes 5, narrowly oblong, about as long as the tube, obtuse and somewhat constricted terminally with an obtuse beaklike projection, to 0.5 107 Psychotria oresbia Dwyer, spec. nov. Arbores ad 6 m altae. Folia oblonga, E 9 cm MR -4 cm lata, versum apicem rotundata venis iolis ad 1 cm longis; stipulis terminaliter persistentibus connatis vagina ad 2 mm longis, coriacea i o OUS bre- vibus praedita. /nflorescentiae terminales in aspectu racemosae ad 4 cm lon O 2 =, oO 3 E. oy z zz No S | © =. == a nd Q E > m = fm G ° > "3 [e] . =. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 409 mm long, located adaxially and perpendicular to the axis of the lobe, located ca. 0.4 mm below the apex of the lobe; stamens 5, the anthers narrowly oblong, ca. 2.2 mm long, obtuse, the filaments slender, ca. 1 mm long, attached near the middle of the tube; ovarian disc rotund, ca. 0.5 mm long, the style 1.5-2.0 mm long, the stigmas 2, ca. 1 mm long. Fruits not seen. Psychotria oresbia is known only from Panama but is to be expected in ad- jacent Colombia. It is distinguished by its oval-oblong leaves, its inflorescence with several pairs of branches distichously arranged and scarcely varying in length. The corolla lobes are striking, having thumblike projections located below the tips and extending in perpendicular fashion from the lobe. The petiole of one leaf has an oblong bladelike dilation distally measuring | cm long, ca. 0.5 cm wide. This resembles the wing of the leaf of Citrus. The leaf in question may be anomalous. DARIÉN: Cerro Tacarcuna W of ridge, trail toward Río Pucuro, just below summit camp, lower montane wet forest, 1500-1600 m, Gentry & Mori 14150 (MO). 66. Psychotria ostaurea Dwyer & Hayden, Ann. Missouri Bot. Garden 54: 143. 1967. TYPE: Panama, Wedel 1461 (MO, holotype). Trees to 8 m tall, the branchlets glabrous, the bark cracking, the pith often porate. Leaves elliptic rotund to ovate oblong, 15-28 cm long, 7.5—15.0 cm wide, deltoid at the apex, distinctly acuminate, basally obtuse to widely cuneate, the costa subprominent above and beneath, somewhat rubescent when dry, the lateral veins 9-14, arcuate, glabrous above, finely puberulent on the costa and the veins beneath, the intervenal areas spreading reticulate, rigidly papyraceous, concol- orous, shiny above; petioles to 3.5 cm long, glabrous, stiff; stipules free, persis- tent at the ends of the twiglets, oblong rotund or ovate oblong, 0.2—0.9 cm long, obtuse or rounded at the apex, scarcely retuse or with 2 obtuse lobes, to | mm long, stiff, glabrous, drying dark. Inflorescences terminal, spreading, cymose paniculate, to 12 cm long, to 8 cm wide, the peduncles 2.0-4.5 cm long, minutely puberulent, terminated by 4 branches, arcuate and umbelliform ascending or pa- niculate, the branches to 6 cm long, spreading, the flowers few; bracts and brac- teoles deciduous. Flowers sessile or on pedicels to 2 mm long; hypanthium | mm long, glabrous, the calycine cup saucer shaped, 1 mm long, undulate and truncate, the teeth absent, pale yellow when dry; corolla white, the tube cylindrical, to 1.3 cm long, scarcely turgid at the base, glabrous or minutely puberulent outside, villose basally within, the lobes 5—6, thick, to 5 mm long, obtuse; stamens 5, the anthers exserted or not, narrowly elliptic, 4-5 mm long, the filaments to 2 mm long, attached near the mouth; style to 8 mm long, the stigmas subulate or digi- tiform, to 2.5 mm long, sessile. Fruits sessile, oblong rotund, to 1 cm in diam., glabrous. Psychotria ostaurea is known only from Panama. CAS DEL TORO: Trail above Quebrada Menaco, in transition cloud forest, Chiriqui Trail, Kirk- bride & Duke 661 (MO). Water Valley, Wedel 1461 (MO). cocLé: Base of Cerro Pilon, ca. 2000 ft, Gentry 3669 (MO). 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 67. Psychotria panamensis Standley, Contr. U.S. Natl. Herb. 18: 132. 1916. TYPE: Panama, Pittier 3194 (US, holotype). Trees to 14 m tall, the branchlets smooth, drying gray, glabrous, the nodes well spaced. Leaves lanceolate or ovate lanceolate, rarely oblong rotund, occa- sionally narrow, 5-20 cm long, 1-9 cm wide, tapering acutely or obtusely toward the apex, occasionally with a cusp to 2 cm long, acute, cuneate or occasionally obtuse at the base, often slightly inequilateral, the lateral veins 7-15(—18), widely arcuate, the costa prominulous above, prominent and often angular beneath, oc- casionally with domatia in the axils of the lateral veins beneath, chartaceous to subcoriaceous, usually drying deep brown or chocolate brown above, reddish brown beneath, glabrate above and beneath; petioles 1.0—4.5 cm long, slender to somewhat stout, glabrous; stipules connate basally, ovate oblong, to 1 cm long, to 0.65 cm wide at the base, obtuse, appressed, stiff, ciliolate. /nflorescences terminal, occasionally axillary, often not solitary but 2 or 3 together, cymose paniculate, to 12 cm long to 10 cm wide, puberulent or glabrate; peduncles to 6 cm long, 0.1—0.2 cm wide, often stiff, lignose, 4-angled, the branches often stiff, lignose, 4-angled, opposite or occasionally ternate, the lowermost branches to 6.5 cm long, the upper branches disposed in 4—5 sets; bracts and bracteoles ovate subulate, to 1 mm long, ferrugineous villose especially adaxially, the pedicels bearing an apical bracteole ca. 0.5 mm long. Flowers (immature) usually sessile or subsessile; calycine cup to 1 mm long, glabrous, eglandular within, the teeth 5, irregular, subtriangular, to 0.5 mm long; corolla white or yellow green, to 4 mm long, thickly petaloid, glabrous outside, densely villose near the mouth, the lobes 5, oblong, 1.2-2.2 mm long, slightly cucullate, villose at the base; stamens 5, the anthers oblong, ca. 1.1 mm long, obtuse, the filaments short, attached near the mouth; stigmas 2, unequal, narrowly oblong, ca. | mm long, the style ca. 3.2 mm long. Fruits on pedicels shorter than the fruit; oblong to subrotund, to 1.2 cm long, drying dark, often rusty red, glabrous, fleshy, the 2 pyrenes often giving the persistent fleshy surface a ribbed effect or flesh deciduous, pyrenes glabrous, ribbed. Psychotria panamensis is known only from Panama. The leaves are usually large but occasional specimens have reduced leaves, e.g., White 248 (F, MO). CANAL ZONE: Miraflores Lake, White 248 (F, MO). CHIRIQUI: Bambito to Cerro Punta, Allen 308 (F. MO). Río Chiriquí Valley, Allen 1387 (F. MO). Boquete, ce 363 Ng , MO). Puerto Armuelles, Davidson 1120 (F). Boquete, Dwyer & Hayden 7963 (COL, DUKE, . MO. NY VEN). Las Siguas Camp, Cerro Horqueta, Pittier 3194 (US). Río ede Viejo, poen 10, 69, 159 (all F, MO), 187 (F). cocrÉ: El Valle, Duke 13151 (MO). COLON: Santa Rita Ridge Road, Dwyer & ene fary (MO). DARIÉN: Caná-Cuasi Trail (Camp 2), Chepigana, Terry & Terry 1436 (F, MO). : 9.4 km N of Goofy Lake, 900 m, Folsom et al. 1968 (MO). El Carti Road, 16-1814 b by pe N of Panamerican Highway, 400—450 m, Nee & Tyson 10939 (MO). vERAGUAS: Road from Santa Fe, past Agricultural School to base of Cerro Tute, 1000-1200 m, Folsom 1603 (MO). 68. Psychotria parvifolia Benth. ex Oerst., Vidensk. Meddel. Naturhist. Foren. Kjøbenhavn 1852: 36. 1853. rype: Costa Rica, Oersted 11637 (F, photo of type). Shrubs to 2 m tall, densely branched, the branches slender, the branchlets slender, puberulent or shortly pilose. Leaves ovate or oblong, 1.0-3.5 cm long, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 411 to 1.5 cm wide, acute or obtuse, tapering at the base, the blade decurrent, mem- branous, glabrous, drying black, usually with axillary domatia; petioles short, slender; stipules deciduous, ovate, ca. 4 mm long, ferrugineous. Inflorescences terminal, sessile, cymose paniculate, the cymes few flowered, the bracts decid- uous. Flowers usually sessile; calyx truncate; vaguely dentate; corolla white, 3— 4 mm long, glabrous outside, villous within, the lobes 4. Fruits small, rotund, red. Psychotria parvifolia is found in Guatemala, Costa Rica, and Panama. CHIRIQUI: Boquete, Maxon 4958 (US). 69. Psychotria perparva Dwyer.1% TYPE: Panama, Gentry et al. 16921 (MO, ho- lotype). Herbs to 0.5 m tall, perhaps unbranched, the stem succulent?, strongly com- pressed on drying, smooth, glabrous, the nodes well spaced. Leaves large, ob- long, to 30 cm long, to 20 cm wide, obtuse or widely triangular at the apex, acumen absent, basally cuneate, attenuate acute or rounded at the base, occa- sionally inequilateral, the costa ventrally convex, prominulous, prominent and convex beneath, to 3.5 mm wide below, the lateral veins 25-30, widely arcuate, joining to form a scarcely undulate submarginal vein to 3 mm from the margin, occasionally a pair arising from the costa as a unit but immediately diverging, the base of each lateral vein dilated, the intervenal areas patulous reticulate, scarcely visible above, invisible beneath, chartaceous, succulent, discolorous, minutely farinose punctate, especially below; petioles to 2.5 cm long, to 0.4 cm wide; stipules deciduous (one seen) connate?, 2-4 mm long, to 12 mm wide, ringlike or hemispherical, truncate or rounded, coriaceous, glabrous, with a small bulge or pair of bulges in a median or subterminal position, 2-3 mm long. Inflorescences axillary, the peduncles to 16 cm long, to 0.25 cm wide, sinuous, sparsely puber- ulent, the rachis slender, to 4 cm long, the branches disposed as 2 pairs, opposite, well spaced, the lowermost to 2 cm long, each branch in turn with a pair of reduced branchlets at or above its middle, these terminated by a much reduced glomerule or 3 compressed glomerules, each to 0.7 cm wide; bracts and bracteoles tending to persist, similar, the lowermost bracts persistent, triangular subulate, to 12 mm long, often as long as the flowerless portion of the primary branches, closely appressed to the latter; bracts subtending the glomerules several, involu- crate, the outermost oblong triangular, to 6 mm long, to 3 mm wide, thinly carnose, venose, puberulent (perhaps only marginally). Flowers sessile, few per glomerule; hypanthium oblong, apparently shorter than the calycine teeth in juvenile state, 108 psychotria UR Dwyer, spec. nov. Herbae to 0.5 m altae glabrae nodis bene distantibus. Folia oblonga, ad 30 cm longa, ad 20 cm lata, acumine nullo venis lateralibus 25-30 areis inter venas subtus vix bipes chartacea minuto-farinosa; petiolis ad 2.5 cm longis, ad 0.4 cm latis; stipulis connatis?, 2-4 mm longis. /nflorescentiae axillares, pedunculo ad 16 cm longo, rhachi ad 4 cm longa ramis 2-jugatis oppositis bene distantibus inferioribus ad 2 cm longis, ramo utroque jugo ramulorum reductorum praedito, ramulis 3 glomerulis terminalibus florum praeditis. Flores dentibus calycis 5, 1.2-1.7(-2.0) mm longis; corolla ad 2.2 mm longa; antheris ca. | mm longis. Fructus sessiles ad 0.65 cm longi, apice rotundi, glabri. 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 glabrous?, truncate at the apex, rounded at the base, the calycine teeth 5, oblong triangular, 1.2-1.7(-2.0) mm long, fleshy, apparently glabrous; corolla white, to 2.2 mm long in the mature bud, carnose, puberulent outside, within villose near the base of the anthers, the latter oblong, ca. 1 mm long, sessile?; style ca. 0.8 mm long, the stigmas 2, upright, ca. 0.35 mm long, scarcely wider than the style; ovarian disc prominent, compressed oblong, ca. 0.5 mm long, truncate. Fruits (one seen) sessile, to 0.65 cm long, rounded at the apex, the calycine lobes persistent, glabrous, the raphides abundant. Psychotria perparva is known only from Panama. It is named "perparva" because of its diminutive size. It has very large leaves for its height and the lateral veins are very numerous. The bulges on the surface of the stipules are diagnostic, although only one stipule was observed. The small glomerules of flowers asso- ciated with the structurally complex inflorescence are noteworthy. ANAMÁ: Cerro Tacarcuna Massif between summit camp on westernmost ridge and Pico Tacar- cuna, mostly elfin forest, 1650-1800 m, Gentry et al. 16921 (MO). 70. Psychotria philacra Dwyer.!'?? TvPE: Panama, Gentry & Mori 13916 (MO, holotype). Shrubs to 3 m tall, the branchlets terete, slender, smooth, glabrous, the nodes well spaced. Leaves persisting at the tips of the twigs, ascending, elliptic, often slightly falcate, 4.0—6.5 cm long, 1.5-3.0 cm wide, acute or subobtuse at the apex, acuminate, the acumen triangular, to 1 cm long, ultimately obtuse, basally acute or attenuate acute, the costa prominulous, the lateral veins ca. 7, arcuate, prom- inulous, the submarginal vein inconspicuous, chartaceous, concolorous, glabrous, minutely granulose beneath; petioles slender, to 1 cm long, to 0.6 cm wide; sti- pules deciduous. /nflorescences (in fruit) umbelliform, to 5 cm long, to 3 cm wide, the peduncle slender, to 3 cm long, ca. 0.5 mm wide, glabrous, terminated by 3 radially disposed branches, these to 1.8 cm long. Flowers deciduous, few, 5-8 on each branch, the pairs of flowers ca. 0.4 cm apart. Fruits subsessile, compressed-rotund, to 1 cm in diam., glabrous, obtusely 12-costate, the persistent disc of the ovary umbonate. DARIÉN: Cerro Tacarcuna, 1250-1450 m, Gentry & Mori 13916 (MO). 71. Psychotria pilosa Ruiz & Pavón, Fl. Peruv. Chil. Prodr. 2: 60. 1799. TYPE: eru, Ruiz (B, isotype labeled “Psychotria pilosa in Peruviae nemoribus," not seen; photo, MO). 199 Psychotria pn n Dwyer, i nov. Frutices ad 3 m alti, ramulis glabris. Folia elliptica, 4,0-6.5 cm longa, 1. m lata acuminata iue lateralibus ca. 7, chartacea glabra; petiolis ad 1 cm longis. a snie E, glabrae ad 5 cm longae, ad 3 cm pn n e culo gracili, ad 3 cm longo, ca. 0.5 mm lato, ramis 3 radiate dispositis terminato, eis ad 1.8 c floribus (hic pam 5-8 in utroque ramo, jugis florum a hi dad oo eae: compresso-rotundi, ad 1 cm diam., glabri obtuse 12-costati P ovarii umbon 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 413 Shrubs to 3 m tall, the branchlets terete, hirsute, the nodes occasionally prox- imate, usually well spaced, 2.0-5.5 cm apart. Leaves oblong, occasionally oblong rotund, 10-40 cm long, 4.5-17.0 cm wide, rounded or deltoid at the apex, the acumen to 1.5 cm long or longer, often falcate, the base obtuse, acute or widely cuneate, occasionally subrotund, the costa slender, prominulous above, promi- nent beneath, the lateral veins 15-20, widely arcuate, prominent beneath with 1— 2 subparallel veins between adjacent lateral veins, the minor veins pinnatiform, the intervenal areas patulous reticulate, stiffly chartaceous, concolorous, pilose above and beneath; petioles to 2.5 cm long, densely pubescent; stipules connate, the sheath to 2 cm long, each part oblong rectangular or ovate oblong with 2 widely subulate lobes, to 0.7 cm long, ca. 0.3 cm wide at the base, often somewhat divergent, widely scarious margined, villose outside. Inflorescences terminal, solitary, often nodding, cymose paniculate or disposed as a capitate mass of flowers, 6-12 cm long, 2.5-6.0 cm wide; peduncle elongate, 5.0-6.5 cm long, ca. 0.25 cm wide, villose, the lowermost branches 3-4, radiate, 0.5-2.0 cm long; bracts oblong or ovate lanceolate, 4.8—7.0 mm long, usually acute or deltoid at the apex, thickly petaloid, villose inside and out. Flowers sessile; hypanthium oblong, densely bearded, the calycine cup scarcely measurable, eglandular with- in, the lobes (4—)5(-6), unequal, triangular or subulate, 1-2 mm long, densely hirsute; corolla white, to 1.5 mm long, villosulose within at filament attachment, the lobes 5, oblong, 0.5 mm long, cucullate; stamens 5, the anthers oblong, 0.65 mm long, the filaments 0.5 mm long; style 1 mm long, the stigmas minute. Fruits with the pyrenes oblong, to 4 mm long, obtuse at the apex, costate, densely bearded, the hairs soft, white, usually more dense at each pole, dark blue or china blue. Psychotria pilosa is known from Costa Rica, Panama, and from northern South America to Peru. Among the Panamanian Psychotria it is readily distin- guished by its large scarious-margined stipules, its many-veined leaves which are hairy beneath, and its densely pilose inflorescence. Most collections note that it is 2-3 m tall. Croat 25777 records a height of 5.5 m. The single collection from San Blas, Liesner 1311, has leaves twice the size of the other collections from the Republic. The fruits are described on several collections as blue, china blue, dark blue purple, and black. The clusters of fruits are described by Mori et al. 3819 as ‘‘grapelike.’’ Mori & Kallunki 3152 note that the tree is "slender, few and laxly branched." AS DEL TORO: Without other locality, Wedel 174 (MO). Nievecita, MM ha b 1831 (MO). "eur | mi E of Cañas Gordas, near Costa Rica border, Croat 22325 (MO). c JH Pilón, Dwyer 8309, 8349 (both MO); Dwyer & Lallathin 8605, 8964 (both MO); Dwyer et E 4485 MO). El Valle, La Mesa, Webb a et al. 3029 (MO). Cerro de ee 2-C, 2D, 30-C (all MO). i Mesa, 4 km N of El Valle, 850 Nee & Dwyer 9192 (MO). RIÉN: Cerro Pirre, Bristan 469 (MO); Duke 5346 (MO); Duke & Elias | 13783 (GH, MO, US). PANAMÁ: Car Tupile road, 12 mi above Panamerican Highway, 200-500 m, Liesner 1199 (MO). Gorgas Memorial Lab, 5-10 km NE of Altos de Pacora, Mori & Kallunki 3418 (MO). SAN BLAS: Road from El Llano to Carti-Tupile, 300-500 m, Liesner 1311 MO). vERAGUAS: Valley Río Dos Bocas, 11-13 km beyond Agricultural School, Santa Fé, 350—500 m, Croat 25777 (MO). Valley of Rio Tercero Braso beyond Escuela Agricola Alto Piedra, Santa Fé, Croat 27340 (MO). 11 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 3152, 3180 (both MO); Mori et al. 3819 (MO). 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 72. Psychotria pithecobia Standley, Publ. Field Columbian Mus., Bot. Ser. 8: 187. 1930. TvPE: Costa Rica, Tonduz 12432 (F, holotype). Epiphytic subshrubs, to 3⁄4 m tall, the stems angular, smooth, glabrous, the nodes well spaced, 2-8 cm apart. Leaves elliptic to oblong, often ovate elliptic to obovate elliptic, 5.0-9.5 cm long, 1.2-4.5 cm wide, usually less than 21⁄2 times longer than wide, acute, obtuse or somewhat rounded at the apex, acuminate, the acumen to | cm long, usually finally acute, the costa prominulous above and below, slender, usually drying yellow, the lateral veins 10-13, arcuate, promi- nulous, conspicuous on drying, not forming an undulate submarginal vein, the veins between the laterals few, these either arising from the costa or arising solely from the laterals themselves, coriaceous on drying, concolorous, drying a dirty yellow or brown, glabrous; petioles 0.5—1.0 cm long, slender or crassate; stipules connate, compressed cylindrical, 2-4 mm long, truncate and scarious above. Inflorescences terminal, occasionally both terminal and axillary, cymose panic- ulate, 6-10 cm long, 2.5-9.5 cm wide, usually patulous, the peduncle 2.0—4.5 cm long, slender, stiff, with about 3 pairs of opposite primary branches, the ter- minal primary branches few and alternate, the secondary branches usually disposed as 1 obvious opposite pair, the terminal secondary branches much reduced and very few, the cymes usually 2-5, each with 2-6 flowers; bracts and bracteoles narrowly oblong, to 3.5 mm long, acute. Flowers pedicellate, the ped- icels 1-5 mm long; hypanthium ca. 1 mm long, glabrous, the calyx cup ca. 0.5 mm long, glabrous, the teeth 4, with perhaps 2 (or more?) very reduced accessory ones, triangular, to 0.5 mm long; corolla white, often tinged with pink, the tube in bud cylindrical, ca. 4 mm long, glabrous outside, villosulose within near fila- ment attachment, the lobes short; stamens 5, the anthers oblong, to 0.8 mm long, obtuse, the filaments very short, attached near the middle of the tube; style about as long as the corolla tube, linear, the stigmas 4, very slender, ca. 0.7 mm long. Fruits subrotund, 3.5-5.0 mm wide, red to orange red, turning black, glabrous, the calyx deciduous. Psychotria pithecobia is known only from Costa Rica and Panama. It is dis- tinguished from P. orchidearum Standley by having more lateral veins in the leaves and the flowers on more elongate pedicels. CHIRIQUÍ-BOCAS DEL TORO BORDER: Cerro Colorado, 11.2 km along ridge road from main road to Escopeta, 1700 m, Folsom 4880 (MO). CHIRIQUI: Bajo Chorro, Boquete, 6000 ft, Davidson 218 (MO). Top of Cerro Colorado, 1500 m, Folsom & Collins 1786 (MO). Bajo Mono, Folsom et al. 2254 (MO). Bajo Mono and Quebrada Chiquero, 1500 m, Woodson & Schery 571 (MO). 73. Psychotria pittieri Standley, Contr. U.S. Natl. Herb. 18: 132. 1916. TYPE: Panama, Pittier 2266 (US, holotype). P. dispersa Standley, J. Wash. Acad. Sci. 18: 184. 1928. rype: Costa Rica, Standley & Valerio 45548 (US, holotype). Key to ae a. Inflorescence with conspicuous white indumentum; leaves usually arias beneath _ . P. pittieri subsp. pittieri aa. Inflorescence without conspicuous white indumentum; leaves wes beneath 73b. P. pittieri subsp. oinochrophylla 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 415 73a. Psychotria pittieri Standley subsp. pittieri. Shrubs 1.0-1.5(—2.5) m tall, much branched above, the branches ascending, terete, smooth, slender, glabrous, the twiglets puberulent, the nodes well spaced. Leaves oblong or lanceolate, 5-15 cm long, 2.0—3.5 cm wide, tapering cuneately at the apex, often cuspidate, the cusp to 1.2 cm long, often falcate basally cuneate, acute to obtuse, often vaguely inequilateral, the costa prominulous above, prom- inent beneath, often with smaller parallel veins evanescing toward the margin, papyraceous, regularly drying dark green above, gray green beneath, glabrate above except on the costa, white pubescent beneath, especially on the costa and veins, or only minutely puberulent on the veins beneath; petioles to 1 cm long, slender, villosulose; stipules with at least the bases persistent, connate, to 6 mm long, each part with 2 awns, the awns subulate, longer than the sheath, the sheath pilose and appressed setose within, the setae conspicuous. Inflorescences ter- minal, paniculate, usually shorter than the terminal leaves, ovoid or oblong, the peduncles often nodding, slender, 1.0—4.5 cm long, the primary branches whorled, 3-4 per whorl, to 1 cm long, perpendicular to arcuate ascending, villose, the bracts linear subulate, to 6 mm long, the bracteoles shorter, usually irregularly 3-lobed, the secondary branches short, bearing a few flowers. Flowers sessile; hypanthium shorter than the calycine teeth, the teeth 5, subulate, 0.5-0.8 mm long, acute, puberulent, the calycine cup with 5 patches of minute oblong glands at the base within, these ca. 0.1 mm long; corolla white to yellow, 4—6 mm long, puberulent outside, the tube pubescent at the mouth within, the lobes 5, short, thinly carnose; stamens 5, the anthers oblong, ca. 0.9 mm long, included or exserted, the filaments almost lacking or elongate; style puberulent, the stigmas 0.5-1.0 mm long. Fruits subrotund, 3.5-6.0 mm wide, violet or turning to blue, the pyrenes ovoid, ca. 3 mm long, puberulent, 5-costate, concave in cross section. Psychotria pittieri Standley ranges from Belize to Panama. The white or gray indumentum of the inflorescence and often of the leaf undersides of subsp. pit- tieri permits ready identification. BOCAS DEL TORO: Nievecita, Bartlett 3 (MO). — n 1380 (MO). Railroad stop, milla 7.5, Croat & Porter 16269 (MO). Río Teribé between Que a Huron and Quebrada Schlunjik, Kirkbride & Duke 479 (MO). Chiriquicito to 5 m S along Río don umo, Lewis et al. 2095 (MO), 2112 (COL, DUKE, K, MO, UC). Without bain Wedel 217, 360 (both MO). Nievecita, Woodson et al. 1838 (MO). CANAL ZONE: Fort Sherman, U.S.A. Tropic Test Site, Dwyer & Robyns 134 (MO). Pipeline Road, Río Agua Salud, Kennedy 1855 (MO). Pipeline Road, Mori & Kallunki 1722 (MO). Pipeline Road 9.2 km from Gamboa, Mori & Kallunki 6487 (MO). Pipeline Road 19 km NW of spoon & Witherspoon 8627 (MO). BARRO COLORADO ISLAND: Croat 4107, 6469, 8732, 9547, 10100, 10797, 11243, 11599, 12205, 14581, 15104A, 15170, 15255 (all MO); vit d 56 (MO); Hayden 6, 9, 146, 156, 1028 (all MO); Kennedy 1847 (MO). cuiniQUÍ: Cerro Pando, NW side, Croat 15898 (MO). Boquete, Davidson 781 (MO). Upper Río Gariche, Seibert 359 (MO). pb. E Valle, Allen 214 (MO). El Valle, Cerro Pilón, Dwyer & Correa 7957 (COL. DUKE, K. MO, UC). Volcan to Rio Serano, 16 km from Rio Serano, Folsom 4049 (MO). Cerro Pilon, Lallathin "s I (MO). Boc sie Toabré, Lewis et al. 5520, 5543 (both MO, SCZ, UC, VEN). COLON: ` Croat & Porter 15609 (MO). Tumba Vieja, Dodge et al. 16932 (MO). Colon to Portobelo, 24.6 km of Transisthmian ighway, Folsom 3743 (MO). W of Portobelo, Gentry 1753, 5141 (both MO). 10 mi d of Portobelo, pis ee (MO). Rio Guanche, cutover woods along trail, Mori & Kallunki 3021 (MO). Rio Guanche, 2.5 km upriver from bridge, road to Portobelo, Mori x ier Iki 3713 (MO); Mori et al. ee (MO). DARIEN: Ascent of Cerro Pirre, Duke 5336 (MO). ecg ici 5138 (MO). Los TOS: 25 mi SW of Tonosí, Lewis et al. 2904 (COL, MO, UC, suyaq AMÁ: Cerro Campana, pa 17217 (MO). Cerro Azul, Dwyer 5133 (MO). Goofy Lake, Dwyer y pes 4130A (MO, SCZ). 416 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Cerro Jefe, Dwyer & Hayden 8095 (MO, UC); Gentry et al. 3362 (MO). Base of Serrania de Maje, ol , Ki del Río Pacora, Lewis et al. 2315A (MO). 5 mi SW of Cerro Brewster, Lewis et al. 3393 (COL, UKE, MO, UC, VEN). El Llano-Cartí Road, 8-11 km from Interamerican Highway, Mori 7734 (MO). Junction of Río Pacora and Río Corso, Oliver 2382 (GH, MO, US). Cerro Azul, Porter et al. 1486 (MO). Flood Plains and hills along Río Armula, ca. 10 m WSW of Puerto Obaldía, Mori 6820 (MO). 73b. Psychotria pittieri subsp. oinochrophylla Dwyer.!!° TvPE: Panama, Duke & Elias 13668 (MO). Shrubs with the leaves purple beneath. Inflorescences moderately puberulent. Psychotria pittieri subsp. oinochrophylla is known only from Panama. Among the Panamanian Rubiaceae the purplish undersurface of the leaf blades is rare. Occasionally P. marginata Ruiz & Pavón shows this character as does P. santa- rosae Standley. DARIÉN: Cerro Pirre, cloud forest and/or mossy forest, 2500-4000 ft, Duke & Elias 13668 (MO). 74. Psychotria polyphlebia Donnell Smith, Bot. Gaz. 33: 253. 1902. sYNTYPES: Guatemala, Donn. Sm. 6854 (US); Costa Rica, Tonduz 9577 (US). Subshrubs, stoloniferous, the stems unbranched, terete, often sinuous, smooth, glabrous, the pith partially hollow. Leaves oblong, obovate oblong, to 14 cm long, to 7.5 cm wide, obtuse or widely deltoid at the apex, occasionally vaguely acuminate, cuneate at the base, often inequilateral, the costa prominulous above, prominent beneath, ca. 1.1 mm wide, the secondary veins to 25, widely arcuate, salient above, prominulous to prominent beneath, joining near the margin, papy- raceous, sparsely or densely appressed hirsute beneath, occasionally diffuse gran- ulose above with the bases of deciduous hairs, often with white raphides; petioles slender, to 6 cm long, ca. 0.15 cm wide, often curved, puberulent; stipules ?con- nate, disposed as a bowl-like band between the petioles, 1-2 mm long, the central acumen blunt, shorter than or longer than the body, glabrous. Inflorescences terminal or subterminal, the peduncles 0.5-3.5 cm long, ca. 0.1 cm wide, pubes- cent, the flowers numerous in a subrotund head to 1.5 cm wide, the bracts oblong, narrowly oblong or subulate, the larger bracts to 5(-8) mm long, ciliate and con- cave. Flowers sessile; hypanthium short, ca. 1 mm long, the calycine cup not measurable, the margin undulate or ?denticulate; corolla white or yellow, the tube narrowly cylindrical, to 4.5 mm long, delicately petaloid, pilose outside, densely villose at filament attachment position within, the lobes 5, narrowly ob- long, shorter than the tube, vaguely cucullate; stamens 5, the anthers ca. 0.6 mm long, obtuse, the filaments slender, ca. 2.5 mm long, attached just below the mouth; ovarian disc compressed rotund, ca. 0.3 mm long, the style filamentous, ca. 2.5 mm long, the stigmas slender digitiform, ca. 0.5 mm long. Fruits sessile, subrotund, 6-8 mm in diam., glabrous, red, the strongly compressed pyrenes with the ventral face subconcave, the dorsal face 3-costate. 10 Psychotria pittieri subsp. oinoc SOLAR Dwyer, spec. nov. Frutices foliis subtus oinochro- phyllis. Inflorescentiae modice puberulae 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 417 Psychotria polyphlebia is known only from Costa Rica, Panama, and Colom- bia. BOCAS DEL TORO: Between Quebrada Lagoon and Cerro Bonyik, 90-270 m, Kirkbride & Duke 641 (MO). DARIEN: Cerro Pirre, Bristan 575 (MO). Cuasi-Cana Trail, Cerro Campamiento, Kirkbride & Duke 1243 (MO). VERAGUAS: Guabal, 16 km NW of Santa Fé, 500 m, Dressler 4781 (MO). NW of Santa Fé, 8.8 km from Escuela Agricola Alto de Piedra, Mori et al. 4004 (MO). Second branch of o š : Be- tween Curiche and Alto Curiche, Duke 9615 (MO). Rain forest N of Alto Curiche, 300 m, Duke & Idrobo 11220 (MO, NY). Novita, N of Cerro Torra, road to Alto del Oso, 600-900 m, Forero et al. 75. Psychotria psychotriaefolia (Seem.) Standley, Contr. U.S. Natl. Herb. 18: 133. 1916 deeds AM Seem., Bot. Voy. Herald 138. 1852. TYPE: Panama, Seemann 482 (BM, een; photo Shrubs or small trees, the branchlets terete, smooth, glabrous to ferrugineous tomentose, the nodes well spaced. Leaves lanceolate or obovate oblong, 4-18 cm long, 2.0-8.5 cm wide, usually acute, occasionally deltoid, often drawn into a triangular apical point, basally attenuate acute, rarely subobtuse, the costa prominulous to plane, sometimes grooved above, prominent beneath, to 0.15 cm wide proximally, the secondary veins 12-20, arcuate, uniting to form a submar- ginal vein ca. | mm from the margin, prominulous above and beneath, the inter- venal areas smooth, the veinlets evanescent, papyraceous to subcoriaceous, often gray or gray red above, often drying dull red beneath, glabrous above, glabrate or farinose puberulent beneath, especially on the veins; petioles absent or to 0.7 cm long, the lamina decurrent; stipules connate but at maturity often tearing and thus appearing free, closely appressed, thin, deciduous, each part to 2 cm long, the sheath cylindrical, subtruncate, the 2 awns long attenuate, shorter than the sheath, ciliate. /nflorescences axillary or terminal, the peduncles rarely absent, usually 3-5 per axil, spreading radially, to 2.5 cm long, often branched apically, the branches 2-3 times as long as the peduncle or shorter than the peduncle, rufous pubescent, angular, the flowers in conglomerate heads, to 1.5 cm wide. Flowers with the hypanthium short, the calycine cup much shorter than the calycine lobes, the latter narrowly oblong, to 1.5 mm long, acute or obtuse, glabrous inside and out; corolla white, the tube short, 1.2-2.0 mm long, petaloid, glabrous outside, villose within at filament attachment, the lobes 5, oblong, cu- cullate, slightly shorter than or as long as the tube; stamens 5, the anthers oblong rotund, ca. 0.5 mm long, the filaments ca. 0.8 mm long; stigmas 3, subulate, the style slender, the ovarian disc oblong, ca. 0.7 mm long. Fruits ovoid to oblong, ca. 7 mm long, usually 2 times longer than broad, drying red, the mesocarp thick, fleshy, sweet, the 2 pyrenes oblong elliptic, the ventral side flattened, the dorsal side with 10 grooves, glabrescent, the raphides numerous, the calycine cup per- sistent but withered. Psychotria psychotriaefolia extends from Costa Rica to northern South Amer- ica, being reported from Colombia, Venezuela, and Ecuador. 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 PANAMÁ: BOCAS DEL TORO: Almirante, G. P. s 557 (P). d Me, Dunlap 229 B (F, US). Susurba, Río Chu h. Duy 4399 (US). Río Teribe between Quebrada Treglo and Puerto Palenque, Kirkbride & Duke 525 (MO). Changuinola to 5 mi S at nct ai Rio Changuinola and Rio Teribe, Lewis et al. 963 n. K, MO, NY, UC, US). Chiriquicito to 5 mi S along Río Guarumo, Lewis et al. 2103 (GH, MO, UC, US, VEN). Without other locality, Wedel 364 (F, MO). CANAL V Without other locality, Duke 6130 (MO). Fort Sherman, Duke 9244 (MO); Dwyer 8590 (MO). Madden Dam, Boy Scout Road, Dwyer & Hayden 7 (MO). Frijoles, Ebinger 90 (MO, US). Chagres, Fendler 58 (MO, US). Fort Sherman, Hayden 104 (MO). Mindi Hills, Johnston 1636 (MO). Pipeline Road, Kennedy & Wilder 3103 (MO). 3 mi E of Coco Solo Hospital, Lazor & Blum 5396 (MO). Rio Masambi Grande 80-100 m, Nee 94/9 (MO). Rio Tapia, Standley 38129 (US). Rio Pedro sn Standley 29941 (US). France Field, Standley 30370 (US). Fort Sherman, Standley 31071 (US). 6 mi N of Gamboa, Tyson 1496 (MO). Madden Lake, Boy Scout Camp, Tyson 5447 (MO). Fort Witherspoon 8301 (MO). BARRO COLORADO ISLAND: Croat 12883 (MO). CHIRIQUI: Boquete, Davidson 709 (F, US), 849 (F). cocLE: El Valle, Allen & Alston 1853 (F, US). Cerro Gaital Caracoral, Dwyer & ae 8950 (MO). COLON: 4 m SW of Portobelo, Croat 14114 (MO). Río Guanche ca. 2.5 km upriver from bridge, road to Portobelo, Crosby & Denslow 6475 (MO). Río Guanche, 1—4 km upstream from Portobelo, Gentry 8813 (MO). 2-3 mi up Río Guanche, Kennedy & Foster 2135 (MO). Río Guanche, 2.5 km from bridge, road to Portobelo, Mori et al. 6475 (MO). Río Guanche, 6 km S of Portobelo, Nee & Gentry 8725 (MO). DARIÉN: Río . ys below Quebrada Venado, Bristan 1037 (MO). Río Aruza, Bristan 1368 (MO). Cativo Cam . 6 mi NW of El Real, Duke 4824 (F, MO). ío Tucuti, ca. 2 hrs by piragua above Tucuti, Duke 5263 (MO). 10 mi S of El Real on Rio Pirre, Duke 5412 (MO). Rio Sambu, 0-5 mi above Rio Venado, Duke 9273 (MO). Río Pucro to Quebrada Maskia, Duke 13081 (MO). Rio Morti, Drill Site 7, Duke 14185 (F, MO). Rio Pirre near town of Pirre, Gentry & m 6938 (MO). Manene to mouth of Río Cuasi, Kirkbride & Bristan 1449 (MO). Upper Rio „Tuque Le Clezio 52 (MO). Río Tuquesa, Lower Tuquesa Mining Camp, Mori 6933 — PANAMÁ: Rio Chagres, Duke 4474 (MO). Goofy Lake, Dwyer & Hayden 8042 (COL, DUK UC, VEN). Gorgas Memorial Labs, 25 km NE of Cerro Azul on Rio Piedras, Mori & Pu os (MO). Tumba Muerto Road, Standley 29816 (US). Near Arraijan, Woodson et al. 776 (MO). s BLAS: Río Cuadi, ca. 6 mi SW of Mulatupu, Duke 14210 (F, MO). Headwaters of Río Cuadi, Cam an Diabolo (Drill Site 22), Duke et al. 3616 (MO, SCZ), 3655 (MO). bp Ra Coiba Island, Collenette 466 (US). COLOMBIA: CHOCO: Madurex Logging Camp, Duke 9973 (MO). 76. Psychotria pubescens Swartz, Nov. Gen. Sp. Pl. Prodr. 44. 1788. TYPE: not seen P. hebeclada DC., Prodr. 4: 513. 1830. TYPE: not seen. Shrubs or small trees, to 4 m tall, the branchlets terete to angular, the indu- mentum variable, rarely glabrescent, the nodes well spaced, the stem often con- spicuously constricted below the node. Leaves lanceolate to oblong lanceolate, 8-17 cm long, 3.0-8.5 cm wide, acuminate at the apex, often abruptly so, acute or attenuate at the base, the lateral veins 9-18, often purple, membranous, pu- berulent to glabrate above, puberulent beneath; petioles slender, to 2 cm long, puberulent; stipules tending to persist, connate, the sheath to 2 mm long, the awns linear, to 5 mm long, conspicuously puberulent to pilose. /nflorescences greenish, terminal, usually solitary, paniculate, shorter than the terminal leaves, to 12 cm long, the lateral branches opposite or whorled, the rachis usually purple, the basal branches to 3 cm long, the uppermost gradually shorter; bracts linear, to 1.5 cm long, puberulent. Flowers on short pedicels; hypanthium short, sub- rotund, to 1.5 mm long, pubescent, the calycine cup scarcely measurable, gla- brous or puberulent, usually with ca. 3 minute glands at the junction of the lobes, the lobes 5, unequal, oblong to deltoid, 0.1—0.5(—1.5) mm long, ciliolate; corolla white, yellow or green, the tube cylindrical, ca. 5 mm long, puberulent outside, pilose within near the filament attachment and pubescent near the mouth, the lobes (4—)5, 2-3 mm long, glabrous within; stamens 5, the anthers oblong, 1.0— 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 419 1.8 mm long, obtuse, the filaments 0.5—1.0 mm long; stigmas 2, erect or deflexed, 0.3-0.65 mm long, the style 3.0—4.5 mm long, crassate, the ovarian disc mound shaped or oblong rotund, to 7 mm long. Fruits subrotund, 3.0—4.5 mm long, the calycine cup persistent, glabrescent, the pyrenes 2, 4—5-costate on the surface. Psychotria pubescens is common throughout Mexico and Central America. Seibert 435 (MO) from El Valle is different from other Panamanian material as the inflorescences are few-flowered. ‘‘Iskarmas Tumat’’; ‘‘Tibisci’’ (Bayano Cuna). CANAL ZONE: Cocoli, Riley 174 (MO). Madden Dam, Seibert 561 (MO). 1 mi below Chilibre, Seibert 1502 (MO). Curundu, Tyson 1046, 1047, 1050 (all MO). Fort Clayton, Tyson 1284 (MO) Curundu, Tyson 1322 (MO). 6 mi N of Gamboa, Tyson 1479 (MO). Farfan Beach, Tyson 1829 (MO). Curundu, Tyson 4173 (MO); Tyson & Dwyer 4454 (MO). Miraflores Lake, White 247 (MO). Ancón 883, 1070 (both MO). cuiRiQUÉ: Boquete, savannas, 4000 ft, Davidson 846 (MO). cocrÉ: El Valle, Seibert 435 (MO). COLON: Behind Portobelo, Croat & Porter 15599 (MO). Portobelo, Ebinger 126 MO). Colon to Portobelo, 0.6 km from Portobelo, Folsom 3758 (MO). DARIEN: Rio Urutí, Bristan 221 (MO). Río Tuqueza below Quebrada Venado, Bristan 1071 (MO). Río Pirre, Croat & Porter 15513 (MO). Río Bayano, near Panamerican Highway, Duke 4001 (MO). El Real to Pinogana, Duke 5154 (MO). Río Chucunaque near Pinogana, Duke 5178 (MO). Río Pucro to Quebrada Maskia, Duke 13066 (MO). Río Bayano, above Piria, Duke 14400 (MO). Río Tuquesa, lower Tuquesa mining camp called Charco Chiva, Mori 6936 (MO). 1 mi from Pinogana near Quebrada Chiuto, Stern et al. 122 (MO). Paya, Río Paya, Stern et al. 178 (MO). PANAMÁ: Juan Diaz, Castillo 27 (MO). Road to Cerro Campana, 3 mi from Interamerican Highway, Croat 12049 (MO). Río Bayano, near Panamerican Highway, Duke 4001 (MO). Tocumen, Dwyer 1853 (MO). El Llano, Gentry 5562 (MO). Chilibre, Mosley 28 (MO). 2 mi E of El Llano, Tyson 1738 (MO). 4 mi W of Chepo near Panamerican Highway, Tyson 6742 (MO). Tocumen, Villamil 24 (MO). Arraijan, Woodson et al. 1360 (MO). Isla Taboga, Ma da - al. 1525 (MO). SAN BLAS: N of Mandinga airport, Duke 14845 (MO). Río Diabolo and . Nargana, Duke 14864 (MO). Mainland opposite Achituppu, Lewis et al. 111 (MO). Mainland pets Ailigandí, Lewis et al. 177 (GH, K, MO, UC, US). VERAGUAS: Rio Trinidad, below Rio de Jesus, Dwyer 1315 (MO) 77. Psychotria pumiliocarpa Dwyer.''! TYPE: Panama, Maas & Dressler 1612 (MO, holotype). Shrubs 1.0—1.7 m tall, the branchlets smooth, wiry, glabrous or moderately appressed villose, the hairs white, the nodes well spaced. Leaves elliptic oblong or lanceolate, 6.5—14.5 cm long, 2-4 cm wide, acute or acuminate, the acumen triangular, to 1.5 cm long, occasionally falcate, acute or attenuate acute and usually slightly inequilateral at the base, the costa slender, often rubescent, prom- inulous above and beneath, the lateral veins 15-20, arcuate, joining to form an indistinct undulate submarginal vein, chartaceous, discolorous or concolorous, glabrous above except occasionally scattered villosulose on the costa above, the costa and lateral veins beneath with spreading, short, white or golden hairs, the intervenal areas with short, appressed, strigose hairs; petioles slender, to 1.8 cm long, puberulent; stipules connate, the sheath turgid, truncate, 2-4 mm long, each u1 Psychotria pumiliocarpa Dwyer, spec. nov. Suffrutices ad 1.7 m alti, ramulis laevibus gra- cilibus. Folia elliptico-oblonga vel lanceolata, 6.5—14.5 cm longa, 2-4 cm lata, acuta acuminataque venis lateralibus 15-20, membranacea vel tenui-chartacea in costa venisque subtus minute aureo- puberula interveniis ciliis brevibus appressis strigosis albis praeditis; petiolis ad 2.8 cm longis. Inflo- rescentiae terminales, ca. 6.5 cm longae, ca. 4 cm latae, ramis ca. 4 jugis dispositis. Fructus sessiles oblongi-rotundi pyrenis 2.5-3.0 mm longis laevibus glabris profunde costatis. 420 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 part with 2 well spaced awns as long as or slightly longer than the sheath, pu- berulent. Inflorescences terminal, cymose paniculate, often drying red, to 6.5 cm long, to 4 cm wide, the peduncle, if present, 2-4 cm long, slender, deflexed, if absent, then with 2 axes arising from the tip of the twiglet, the branches disposed as 2-3 well spaced pairs, the lowermost opposite, the others mostly alternate, spreading, the cymes few flowered; bracts subulate to narrowly oblong, 2-4 mm long. Flowers (only partly seen) sessile; hypanthium oblong, less than 1 mm long, pubescent, the calycine cup somewhat saucer shaped, ca. 0.2 mm long, scarious, the margin wavy, the teeth not evident; corolla and stamens not seen; ovarian disc about as long as the calycine cup, the style ca. 1 mm long. Fruits sessile, small, oblong-rotund, the pyrenes 2.5-3.0 mm long, smooth, glabrous, replete with cystoliths, deeply ribbed, the traces of the calyx scarcely measurable. Psychotria pumiliocarpa is known only from Panama. It is distinguished by a combination of characters: membranous or thin chartaceous leaves, connate stipules with 2 erect awns, a toothless saucerlike calyx, and small fruit. VERAGUAS: Alto de Piedra, forest, 800 m, Maas & Dressler 1612 (MO). 78. Psychotria quadrangulata Dwyer.'? type: Panama, Mori & Kallunki 5363 (MO, holotype). Trees, 10 m tall, the twiglets slender, 4-angled, the bark smooth, glabrous, the nodes widely separated, to 10 cm apart. Leaves 6-19 cm long, 2-8 cm wide, obtuse or deltoid at the apex, the acumen triangular, 0.5-1.0 cm long, ultimately acute or obtuse, cuneate to obtuse at the base, the costa subplane to prominulous above, scarcely prominent beneath, drying dull pink, to 0.15 cm wide proximally, the lateral veins 9-10, prominulous beneath, arcuate, not forming an obvious submarginal vein, drying yellow red, the margin delicately callose, chartaceous, mostly somewhat discolorous, glabrous above, beneath minutely puberulent on the costa and the veins and with a few scattered elongate hairs on the costa; petioles slender, stiff, to 2.5 cm long, to 0.2 cm wide, with a few scattered hairs: stipules persistent, perhaps briefly connate, the sheath compressed rotund, 3-5 mm long, each part with 2 erect horns, ca. 4 mm long, glabrous. Inflorescences terminal, cymose-paniculate, often pyramidal, 6-10 cm long, to 5 cm wide at the base; peduncle 4-5 cm long, slender, rigid, minutely puberulent; bracts and brac- teoles minute, ovate, deciduous, the basal branches usually opposite, occasion- ally 4-verticillate, the branches above usually 4-verticillate, well separated from the lowermost branches, the terminal branches few, reduced, the cymules few, few flowered, patulous. Flowers sessile or subsessile, the hypanthium 0.5-0.7 U Psychotria quadrangulata Dwyer, spec. nov. Arbores 10 m alti ramulis glabris quadrangulatis. Folia 6-19 cm longa, 2-8 cm lata, venis lateralibus 9-10, chartacea subtus in costa venisque minuto- puberula; petiolis irure ad 2.5 cm longis; stipulis persistentibus Boii breviter connatis vagina compresso-rotun m longa bicornuata. /nflorescentiae termina moso-paniculatae, 6—10 cm longae, pedun culo 4.5 cm longo, ramis inferioribus oppositis vel 4- uoa cymulis paucis paucifloris patulis. “Flores lobis calycis oblongo-rotundis, 0.5-0.7 mm longis. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 421 P a te ts Š EN NOR W — t M SW ius AW af 4 tL | EE >a FIGURE 76. Psychotria racemosa (Aubl.) Raeuschel.—4A. Habit (x3/;).—B. Flower (x6).—C. Flower opened to show interior (x6).—0D. Ovary, cross section (x6).—E. Fruit (x6). [After Hayden 128.] mm long, glabrous or with a few short hairs, the calycine cup short, the lobes oblong rotund, 0.5-0.7 mm long; corolla (immature) glabrous outside. Fruits not seen. Psychotria quadrangulata is known only from Panama. VERAGUAS: NW of Santa Fe, 2.7 km from Escuela Agricola Alto de Piedras, road to Calovéboro, Mori & Kallunki 5363 (MO). 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 79. Psychotria racemosa (Aubl.) Raeuschel, Nomen, Ed. 3: 56. 1797.—Fic. 76. P. racemosa (Aubl.) Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. Nonatelia racemosa Aubl., Hist. Pl. Guiane Fr. 186. 1775. TYPE: in sylvis territorii PRU French Guiana, Aublet, not seen. Shrubs to 1% m tall, the branchlets slender, terete, smooth, minutely puber- ulent, sometimes contracted below the nodes, the latter well spaced. Leaves oblong or lanceolate, occasionally subfalcate, to 18 cm long, to 8 cm wide, cus- pidate at the apex, the cusp to 1 cm long, acute to attenuate acute at the base, usually equilateral, the costa prominulous above, prominulous to subprominent beneath, to 0.13 cm wide at the base, the lateral veins 12-14, widely arcuate, prominulous, the veinlets prominulous, membranaceous to chartaceous, smooth, glabrous; petioles to 3 cm long, to 1.3 mm wide; stipules tending to persist, erect, free, the body scarcely measurable or to 1.5 mm long, the lobes 2, subulate, 8— 13 mm long, glabrous, fimbriate within. /nflorescences usually terminal, solitary, shortly pedunculate, 2-5 cm long, about as long as broad. Flowers with the hypanthium short, glabrate, the calycine cup scarcely measurable, lacking glands within, the teeth irregular and unequal, subulate to oblong rotund, to 0.8 mm long, acute and/or obtuse; corolla white, the tube cylindrical, 2.5 mm long, pet- aloid, glabrous or papillose puberulent outside, villose within near the filament attachment, the lobes oblong or ovate oblong, 0.8 mm long, cucullate, often papillate puberulent within; stamens 5, oblong, ca. 1 mm long, attached above the middle of the tube; stigmas 5, linear subulate, ca. 0.7 mm long, the style slender, 2 mm long, often papillate puberulent; the ovary 5-celled. Fruits usually somewhat congested, rotund, 5-6 mm in diam., 5-lobed, maturing from orange red to black; pyrenes ca. 3 mm long, lightly costate. This species is widespread throughout tropical America and is immediately recognized by the 5-carpellate ovary and fruit BOCAS DEL TORO: Halfway between Quebrada Huron and Puerto Palenque, Kirkbride & Duke 505 (MO). Punta Pena, near Der Lewis et al. 2144 (MO, UC, VEN), 2147, 2153 (both MO). Fish Creek Mountains, Wedel 2265 (MO). CANAL ZONE: Gamboa Pi line, Correa & Haines 482 (MO). Drowned Forest, Quebrada Bonita, Dodge & Allen 17068 (MO). Drowned Forest, Quebrada Ancha, Dodge & me OR 17044 (MO). George Greene Memorial, Madden Forest. Dwyer & Hayden 11 (MO). Coco Solo, Gentry 6422 (MO). Madden Dam Road, Hayden 75 (MO). Fort Clayton, Hayden 174 (MO). Pipeline Road, Río a Salud, Kennedy 1856 (MO). 8 km NW of Gamboa, Nee 6433 pd Splice S16 and S49, Río Indio, Steyermark & Allen 17454 (MO). Cocoli. Tyson 1617 (MO). I mi N of Summit Garden, Tyson & Blum 1949 (MO). George Greene Forest Preserve, Tyson et al. oe (MO). BARRO COLORADO ISLAND: Aviles 44, 88A (both MO); Croat 4031A, 5750A, 5971, 6182, 6189A, 6199A, 6763, 10338, 10842, 10929, 11246, 11475, 15116, 16533 (all MO): Hayden 128 (MO), 1027 (GH, MO, UC, US); Hladick s.n. (MO); Killip 39961 (MO); Shattuck 26 (MO). CHIRIQUI: San Bartolome, Woodson & Schery 881 (MO). COLÓN: 4 km NW of Salamanca, 3 km NE of Buenos Aires, 340-410 m, Nee 9090 (MO). DARIÉN: Río Menus Bristan 1390 (MO). Río Chico, from Yaviza to ca. | hr by outboard from junction with Rio Chucunaque, Burch et al. 1122 (F. GH. K. MO, US). Río Tucuti ca. 2 hrs by outboard above Tucutí, Duke 5268 (MO). 10 mi S of El Real, Duke 5416 (MO). Near Tusijuanda, | day up Rio Balsa from Manené, Duke 13529 (MO). Ca. 1 mi SE of tes Tuira, Interamerican Highway, Duke | 14568 (MO). River Ganglon, Duke & Bristan 368 (MO). 1— N of Paya, Duke & Kirkbride 14007 (MO). Between Paya and Palo de los Letras, Duke & Leis 14032 (MO). d Stern et al. 461, 470 (both MO). 3 mi i of Santa Fé, Tyson et al. 4696 (MO). Los SANTOS: Headwaters of Rio Pedregal, 25 mi SW of To . Lewis et al. 2900 (MO, UC). PANAMA: Rio =e Maestra, pre 2 (MO). Juan Diaz, Castillo 15 "(MO ). Jenine. Duke 3803, 3877 (both MO). Río Pita, 1-3 mi above confluence with Río Maesus; Duke 4767 (MO). Piria-Canasas Trail near Piria. id pei O). Goofy Lake, Cerro Azul, Dwyer & Hayden 8023 (GH, MO, US). Rio Pasiga, near terfall, Gentry 2281 (MO). 5 mi SW of Cerro Ere lee Lewis et al. 3525 (MO). Chimán, Lewis 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 423 et al. 3235 (COL, DUKE, K, MO, UC, VEN). 6.5 km by road N of Lago Cerro Azul, 650-730 m, Nee 9302 (MO). Cerro Jefe, Wilbur et al. 11336 (MO). SAN BLAS: E of Cangandí-Mandinga Airport, Duke 14770 (MO). Rio Diabolo and Rio Acuati near Nargana, Duke 14874 (MO). 80. Psychotria rosulatifolia Dwyer.!? TYPE: Panama, Gentry & Mori 141 74 (MO, holotype).—Fic. 77 Herbs to 0.5 m tall, the stems smooth, drying brown, the lenticels small, diffuse, pustulate, the nodes moderately well spaced except at the apex of the twiglets. Leaves sessile, obovate oblong or obovate, occasionally subspathulate, 18-31 cm long, 8-9 cm wide, acute or deltoid toward the apex, acuminate, the acumen conspicuous, to 2.5 cm long, falcate, ultimately subacute, somewhat attenuate toward the base or below the middle of the blade, finally truncate or subauriculate basally, the costa subimmersed above, prominent beneath, the lat- eral veins 20-30, spreading at first perpendicular to the costa then widely arcuate, plane above, prominulous beneath, subrubescent when dry, with a conspicuous paralleling vein ca. 2 mm from the margin, the intervenal areas spreading retic- ulate, stiffly membranous, glabrous above, puberulent beneath on the costa and the veins; stipules free, ovate lanceolate, to 1.5 cm long, ca. 1.8 cm wide, acute, truncate at the base, scarious, glabrous, the median vein prominulous. /nflores- cences terminal, solitary, cymose paniculate, the peduncle to 9 cm long, ca. 0.12 cm wide, obtusely angular, the primary basal branches paired, opposite, to 4.5 cm long, the upper primary branches disposed as 2 sets of 3 well-spaced whorls, the secondary branches few, similarly disposed; lowermost bracts ovate oblong, to 3 mm long, spreading, glabrous, resembling the stipules in texture. Flowers sessile or with pedicels to 2 mm long; hypanthium ca. 1.5 mm long, glabrous; calycine cup cylindrical or cup shaped, ca. | mm long, truncate, glabrous, the teeth as 5 scarcely measurable points; corolla white, the tube cylindrical, 3.2—6.0 mm long, 1.2-1.5 mm wide in the middle, glabrous outside, villosulose within near filament attachment, the lobes 5, narrow oblong, 3-4 mm long, ca. | mm wide, narrow toward the apex, erect as bud opens, reflexed at anthesis, with a linear, adaxial spur, 0.3-0.6 mm long, 1.0-1.5 mm below the tip of the lobe; stamens 5(—6), narrowly oblong, ca. 2 mm long, obtuse to truncate, the scarcely measurable filament affixed near the middle of the tube; ovarian disc 0.8—1.0 mm long; style 2.2-5.0 mm long, the stigmatic lobes 2, scarcely wider than the style, 0.2-0.4 mm long. Fruits oblong, to 5 mm long, each pyrene 4-ribbed, glabrous, red when dry, the calyx often persistent, to 0.5 mm long. Psychotria rosulatifolia is known only from Panama. It is distinguished from all other Psychotria by the sessile many-nerved leaf blades disposed in rosette fashion at the apex of the branchlets. The slender subterminal adaxial appendage on each corolla lobe is diagnostic. The auriculate character of the blades is es- pecially striking for a Psychotria. d Men iuri Dwyer, spec. nov. Herbae ad 0.5 m altae. p eigen obova oblonga vel obov subspathulata venis lateralibus 20—30 rigido-membranacea glabra ds Mau stipulis liberis Mine lanceolatis, ad 1.5 cm longis, ca. 0.8 cm latis, a n na media vix prominula praeditis. Inflorescentiae (in fructu visae) terminales cymoso-paniculatae pedunculo ad 9 cm longo, ramis 3 ascendentibus terminato, inferioribus ad 4 cm longis, his dichotome vel trichotome jaa tibus, bracteis inferioribus ovato-oblongis ad 3 mm longis, glabris. Fructus oblongi, ad 5 mm longi, pyrena utraque 4-costata glabra in sicco rubra calyce ad 0.5 mm longo. 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 DARIEN: Ascent of Cerro Pirre, Rio Pirre, S of El Real, 750-1030 m, Duke 5339 (MO). Cerro mossy forest, 2500-4500 ft, Duke & Elias 13788 (MO). Cerro Pirre, vicinity of Rancho Plastico, Folsom 4295 (MO). Trail from Rio Pucro, base camp up W ridge of Cerro Mali, wet forest, 640-1000 ntry & Mori 14174 (MO). 81. Psychotria sanfelicensis Dwyer.!!4 Type: Panama, Mori & Kallunki 5871 (MO, holotype Shrubs to 2 m tall, branching freely, the ultimate branchlets glabrous, pou smooth, the nodes well spaced. Leaves elliptic or oblong-elliptic, 2.5-6.0 c long, 1.3-2.2 cm wide, conspicuously attenuate toward the apex, often mos. acuminate, the acumen to 1.5 cm long, acute, basally attenuate, the costa prom- inulous on both sides, the lateral veins 9—10, joining to form an irregular vein to 1 mm from the margin, slender, arcuate, the smaller veins between pairs of lateral veins soon branching, the intervenal areas spreading reticulate, the margin thinly callous, chartaceous, discolorous, yellow green when dry beneath, glabrous; pet- ioles slender, to | cm long, glabrous; stipules ?connate, the body cupuliform, to 1.5 mm long, the margin undulate, the teeth 4, minute, to 0.3 mm long, similar in aspect to the teeth of the calyx, glabrous. Inflorescences terminal, pyramidal paniculate, to 8 cm long, to 6 cm wide, glabrous, the peduncle to 2.5 cm long, .] cm wide, the branches few, opposite, the lowermost to 3 cm long, the sec- ondary branches occasionally opposite, the lowermost to 3 cm long, the cymules few, unilaterally disposed; bracts and bracteoles subulate, the lowermost bracts subulate, to 6 mm long. Flowers (only one partly seen) with the hypanthium subrotund, | mm long, glabrous, the cystoliths abundant, the calycine cup 0.3- 0.6 mm long, the teeth 5, scarcely visible or to 0.3 mm long, truncate; ovarian disc longer than the calyx. Fruits (?immature) oblong, to 3 mm long, obtuse, glabrous, the cystoliths dense, the calyx persistent. Psychotria sanfelicensis is known only from Panama. CHIRIQUÍ: N of San Félix on Cerro Colorado copper mine road, along continental divide, lower montane rain forest (cloud forest, trees to 5 m tall) 5000-5500 ft, Mori & Kallunki 5871 (MO). 82. Psychotria pg Dwyer.!? TYPE: Panama, Dwyer & Gentry 9565 (MO, holotype).—Fic. 78. "4 Psychotria sanfelicensis Dwyer, T nov. dare es ad 2 m alti bia ultimis glabris. Folia elliptica vel oblongo-elliptica, 2.5-6.0 cm longa, 1.3-2.2 cm lata, acumine ad E 5 cm longo, venis lateralibus 9-10, chartacea omnino rase Sedis ad | cm longis; «usus fortasse HUE vagina ad 1.5 mm longis, margine undulato dentibus 4 ad 0.3 mm longis. didus entia term ad 8 cm i ad x. cm latae, glabrae; pedunculo ad 2.5 cm longo, ramis paucis oppos s inferioribus ad 3 cm longis, cymis paucis unilateraliter dispositis; bracteis ad 6 mm lon ue Flores cu ant a calycis dentibus 5 a visibilibus vel ad 0.3 mm longis, truncatis; disco ovarii quam calyce SE. Fructus oblongi, glabri aS Psychotria ee Dwyer, spec. nov. Arbores ad 20 m altae, ramulis glabris. Folia ovato-rotunda vel lato-oblonga, 5-19 cm longa, 2.2-12.6 cm lata, apicem versus rotunda, venis la- teralibus 8—13 rigido- seit. tenui coriacea in sicco sordido-flava supra minuto-farinosa subtus amdo: puberula; petiolis ad 1.5 cm longis; stipulis ad 5 mm longa truncata “yasa, 2 solum ad 1.2 longis, coriaceis glabris. Inflorescentiae terminales paniculatae, ad 9.5 cm longae, ad 8 cm latae. Flores (in gemmis) sessiles cupula calycis nulla lobis 5 rotundis vel G. 0.5-1.0 mm longis; corolla tubo 3 mm longo, extus puberulo; antheris ca. 2 mm longis. Fructus oblongo-rotundi, ad 0.9 cm longi, glabri. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 425 aa s. El Cope, 9.4 km above El Cope, 750-900 m, Croat 44664 ae 7 km N of El Cope de Veraguas 900 m, dpi Ed (MO). Alto Calvario, 7 km N of El Cope, 700 m, d 2483 (MO). Cerro Pilon. 5 km NE El Valle, 800-1045 m, Mori 6568, 6617 (both MO). : Ric Guanche, ca. 3—5 km above Sia bridge, Croat 37013 (MO). DARIÉN: Cerro Pirre, Bristan 605 (MO): Duke 5337 (MO). Cerro Campamento, S of Cerro Pirre, Duke 15694, 15713 (both MO). Cerro Pirre, Summit, Gentry & Clewell 7035 (MO). Cerro Mali, 1200-1400 m. Gentry & Mori 13703 (MO). Caná Trail between Cerro Campamento and La Escalera to ‘‘Paramo,”’ E of Tres Bocas, Kirkbride 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 455 & Duke 1309, 1332 (both MO). Cana, Williams 832 (US). PANAMA: Cerro Campana, Croat 17176 (MO). Cerro Jefe, Croat 22653 (MO). El Llano-Carti Road, 360-400 m, Croat 25157 (MO); D’Arcy 10612 (MO). Between Cerro Jefe and La Eneida, Dwyer 8238 (MO). Cerro Jefe, Dwyer & Gentry 9458A (MO); Dwyer et al. 7247 (MO). W of El Llano, Gentry 5096 (MO). 5-6 mi N of El Llano, Gentry 5789 (MO). El Llano to Carti Tupile, 150-400 m, Kennedy et al. 3135 (MO). Cerro Campana, Kirkbride & Hayden 270 (MO). Campo Tres, 3 mi NE of Altos de Pacora, Liesner 544 (MO). El Llano-Carti Road, Maas et al. 1769 (MO): Mori & Kallunki 2297, 2330, 4123, 5584 (all MO). Cerro Campana, 870-1000 m, Mori & Kallunki 2499, 3585 (both MO). Gorgas Memorial Labs, 25 km NE of Cerro Azul on Rio Piedras, Mori & Kallunki 3266 (MO). 5-10 km NE of Altos de Pacora, Mori et al. 4200, qe (both MO). El Llano-Carti Road, 400-450 m, Nee et al. 8796 (MO). VERAGUAS: Cerro Tuté 0 km NW of Santa Fé, Mori 6259, 6772 (both MO). NW of Santa Fé, 2 km from Escuela pices yh de Piedra, Mori & Kallunki 5246 (MO). 5. Raritebe trifoliatum (Dwyer & Hayden) Dwyer, comb. nov. PNE “aap Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 139. 1967. TYPE: Panama, Dwyer (MO, holotype). Subshrubs to 1.3(—3.0) m tall, the branchlets angular, densely pubescent, can- aliculate. Leaves usually 3 per node, elliptic or obovate elliptic, 15-33 cm long, 6-11 cm wide, obtuse or widely deltoid at the apex, occasionally falcate, acu- minate, the acumen to 1 cm long, basally cuneate to attenuate acute, often slightly inequilateral, the costa prominulous or immersed above, prominent beneath, the lateral veins ca. 17, widely arcuate, the intervenal areas spreading reticulate, rigidly papyraceous, diffusely appressed pilose to glabrate above, strongly ap- pressed pilose beneath; petioles 0.8-2.5 cm long, ca. 0.3 cm wide, lignose, ie twisted, densely pubescent; stipules ovate lanceolate, to 2 cm long, ca. 0 wide at the base, tapering narrowly at the apex, acute or obtuse, Tongiadinalls striate, appressed pilose, the base of the hairs usually somewhat turgid. /nflores- cences terminal, solitary, thrysoid cymose, 3—9 cm long, the peduncles to 6 cm long, stout, usually twisted, 1.5-3.5 mm wide, densely puberulent, the branches few, the lowermost paired, 2.0-2.8 cm long, 1.0-1.6 mm wide, often somewhat deflexed, to 3 cm apart, the cymules terminal; bracts and bracteoles subulate, densely golden pilose. Flowers (bud) with the hypanthium ca. 1 mm long, the calycine cup elongate, cylindrical urceolate, to 2.5 mm long, the hairs dense, appressed inside and out, carnose, often drying black, the teeth 4—5, often un- equal, triangular subulate to subulate, to 0.5 mm long, ciliate; corolla cream colored or green, to 4.5 mm long, less than 2 times the length of the hypanthium and the calycine tube, the tubes densely appressed pubescent outside, glabrous within, the lobes 4 or 5, short; stamens 5, the anthers narrowly oblong, 2.2 mm long, the apex petaloid, acuminate for 0.5 mm, acute, the filaments short; style slender, the stigmas 2, small. Fruits rotund, to 0.7 cm in diam., orange, drying red or occasionally black, pubescent, delicately ribbed and rugulose, the fleshy surface marcescent; seeds rotund, reticulate, glandular shiny. Raritebe trifoliatum 1s known only from Panama. AS DEL TORO: Quebrada Huron on Cerro pr Kirkbride & Duke 603 (MO). COCLE: El Valle, po 1933 (MO). Cerro Pilon, Dwyer 7925 (MO, US), 8355A (MO). Cerro Gaital Caracoral, Dwyer & Correa 8849 (MO). uo Pilon, Kirkbride p 1119 (both MO); Lallathin 32-B (MO). VERAGUAS: Río Primero Braso. 2.5 km beyond Agricultural School Alto Piedra near Santa Fé. 700- 750 m, Croat 25490 (MO). Road between yn Piedra school and Río Calovebora, ca. 500 m, Croat Folsom 34106 (MO). NW of Santa Fé, 8.8 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 3914 (MO). 456 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 6. Raritebe victoriae (Dwyer) Dwyer, comb. nov. Dukea victoriae Dwyer, Ann. Missouri Bot. Gard. 53: 364. 1966. TYPE: Panama, Duke 6099 (MO, holotype). Shrubs, the branchlets pubescent. Leaves falcate lanceolate, 16-27 cm long, 3.5-8.0 cm wide, narrowly falcate acuminate at the apex, cuneate at the base, the costa prominulous above and beneath, the lateral veins ca. 20, the median veins 1.5-2.0 cm apart, glabrescent above, golden villosulose beneath, especially on the veins; petioles to 4.5 cm long, ca. 1.3 cm wide; stipules narrowly triangular subulate, to 1.5 cm long, acute, pubescent. /nflorescences terminal and axillary, shorter than the terminal leaves, the peduncle either absent and then with 3-4 axes arising from the tip of the twiglet, slender, to 5 cm long, divergent, puber- ulent, or the peduncle to 1.5 cm long, ca. 0.2 cm wide, pubescent, the cymules subflabellate; bracts subulate, to 3 mm long; bracteoles minute. Flowers pedi- cellate, the pedicels to 6 mm long; hypanthium urceolate, to 2 mm long, sparsely pubescent, the calycine cup to 1 mm long, truncate, the teeth 4, much reduced; corolla tube cylindrical, to 2.5 mm long, ca. 0.8 mm wide, glabrous, subcarnose, the lobes a little longer than the tube; stamens 5, the anthers linear, ca. 2 mm long, attenuate toward the apex, ultimately obtuse, the filaments ca. 0.8 mm long, affixed to the base of the tube; style ca. 1 mm long, the stigmas linear, ca. | mm long. Fruits not seen. Raritebe victoriae is known only from Panama. DARIÉN: Cerro Pirre, Duke 6099 (MO). 70. RAVNIA Ravnia Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1852: 49. 1852. TYPE: R. triflora Oerst. Epiphytic shrubs. Leaves fleshy, the lateral veins frequently evanescent; stip- ules interpetiolar, large, oblong, obtuse (fide Standley). Flowers large, 1-3 at the end of the branchlets; hypanthium turbinate, the calycine cup absent or greatly reduced, the lobes absent or 5-6, minute to conspicuous, narrowly oblong, often persisting in fruit; corolla with the tube cylindrical or ampliate at maturity, pe- taloid, the lobes 5-6, valvate, short, finally reflexed; stamens 5-6, the anthers oblong, attached by very short filaments to the mouth; ovary 2-celled, the ovules numerous, attached to a median septum, ascending, the style filiform, elongate, the stigmas 2, short, obtuse. Fruits capsular, elongate and narrowly oblong, split- ting septicidally from a thin septum; seeds small with a distal tuft of elongate hairs. Ravnia is known only from Central America, probably confined to Costa Rica and Panama. It is made up of 3 species. The genus is in need of intensive field study, especially from the viewpoint of floral development. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 457 FIGURE 82. Ravnia triflora Oerst.—A. Habit (x!⁄2). [After Woodson & Schery 559.] —B. Flower opened to show interior (x24). [After Lent 1225, Costa Rica.]—C. Fruit (x%).—D. Seed (Xx 11⁄2). [After Gómez 2230, Costa Rica.] 458 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1. Ravnia triflora Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1852: 49. 1852. TYPE: Costa Rica, collector not given.—Fic. 82 R. pittieri Standley, N. Am. Fl. 32: 114. 1921. TYPE: Costa Rica, Pittier 13461 (NY). R. longifilamentosa Steyermark, Ceiba 3: 21. 1952. TYPE: Panama, Allen 4744 (F, holotype). R. panamensis Steyermark, Ceiba 3: 22. 1952. TYPE: erroneously omitted. Epiphytic shrubs, usually anisophyllous, branching loosely, the stems gla- brous, the nodes well spaced. Leaves occasionally sessile, lanceolate to ovate lanceolate, 4.5-17.0 cm long, 1.7—4.0 cm wide, the larger leaves usually 4 times longer than wide, the small anisophyllous leaves !4 to % the length of the larger leaves, the smallest mostly ovate or ovate lanceolate, the larger often falcate, acute or narrowly deltoid toward the apex, often gradually caudate, basally acute, cuneate or rounded, the costa plane or prominulous above, prominulous beneath, the lateral veins 4—7, not evident or only faintly visible, sharply arcuate ascending, papyraceous to subcoriaceous, concolorous, often farinose, glabrous; petioles absent or to 1.5 cm long; stipules deciduous, free, the body oblong, ca. 2.5 mm long, drying tan, puberulent, with 2 linear oblong awns to 7 mm long, puberulent, 3-veined. Flowers solitary or to 3 at the end of the twiglet, the peduncle absent or scarcely measurable; bracts subtending flowers paired, deciduous, oblong, to 2.2 cm long, ca. 0.8 cm wide, obtuse, chartaceous, glabrous. Flowers sessile or subsessile; hypanthium turbinate, ca. 5 mm long, ca. 2 mm wide, the calycine cup absent, the calycine teeth absent or 5-6, linear oblong, unequal, 1-12 mm long, glabrous; corolla with the tube narrowly cylindrical or widely hypocrater- iform, 3.5-6.5 cm long, 0.8-1.6 cm wide, glabrous, venose, the lobes 5—6, erect when immature, reflexed at maturity, linear oblong or triangular, to 6.5 mm long at maturity, obtuse; stamens 5-6, exserted or included, the anthers oblong, ca. 4 mm long, the filaments 2-3 mm long, attached near the mouth; style slender, 38-50 mm long, ca. 0.3 mm wide. Fruits narrowly oblong or narrowly obovate, often falcate, to 7 cm long, to 0.9 cm wide, each valve often with delicate lon- gitudinal lines, the calycine teeth persistent or deciduous from a somewhat round- ed fruit apex; seeds numerous, narrowly cuneate, ca. 0.2 mm wide, proximally slender, distally thicker, the terminal hairs to 20 mm long. Ravnia triflora is known only from Costa Rica and Panama. The corolla tube in maturing passes from a cylindrical shape to a gibbous state with its lobes becoming reflexed, exposing the anthers. This is why I consider all the collections to represent one species plausible. The calycine lobes may be absent or may range from 1-15 mm in length. Anisophylly is of little importance as a diagnostic character, as well as is the number of flowers at the tip of the twig. The stipules and bracts were described from Folsom 4361 (MO). CHIRIQUÍ: Southern slopes of Quebrada Ma vicinity of Finca Lerida, 1631 m, Allen 4744 ee Cerro Horqueta, Hagen & nie dee 2168 (F). per Rio Chiriqui Viejo, White & White 33 (MO), 3 (F, MO). Bajo Mono and Quebrada Chi Vi. Medan & Schery 559 (MO). cocr É: Cerro d Croat 14341 (MO); Duke & deren 14997 (MO). Asceradera Rivera, Alto de Calvario, 7 km N of El Cope, 700 m, Folsom 2340 (MO). DARIEN: Cerro Pirre, valley between Pirre and mountain directly S, Bane 4361 (MO). Cerro Pirre, Gentry & Clewell 6971, 7032 A (both MO). PANAMA: Altos de Pacora, 15-20 km WNW of Cerro Azul, ca. 800 m, Mori et al. 6913 (MO). VERAGUAS: Road between Escuela Agricola and Alto Piedra, iS Santa Fé and Río Dos Bocas, ca. 5-8 km from Escuela, 730-770 m, Croat en (MO). 0.6 mi beyond Escuela Agricola Alto Piedra, Croat & Folsom 34048 (MO). 5 mi W of Santa Fé, Liesner 876 (MO). N of Santa Fé, ca. 2 km N of Escuela Agricola Altos de Piedras, Mori & Kallunki 2600 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 459 71. RELBUNIUM Relbunium (Endl.) Benth. & Hook. f., Gen. Pl. 2: 149. 1873. Rubia sect. Involucratae DC., Prodr. 4: 590. 1830. TYPE: not selected. Galium sect. Relbunium Endl., Gen. Pl. 523. 1839. TYPE: Relbunium hypocarpium subsp. nididum (H.B.K.) Ehrendf. (fide Ehrendorfer). Herbs, annual or perennial, erect or sprawling. Leaves opposite, together with the stipules disposed verticillately. Inflorescences pedunculate, the flowers solitary to numerous; flowers usually bisexual and sessile, enveloped by an involucre of 2-4 bracts. Flowers with the calycine cup bearing reduced or ob- solete lobes, not persistent in fruit; corolla rotate, the lobes 4, valvate; stamens 4, exserted, the filaments attached at the mouth; ovary 2-celled, the ovules sol- itary in each cell, the style branches 2. Fruits fleshy, didymous, 1 seed per cell or | seed aborting; seeds oblong, the ventral surface concave. Relbunium, a genus of about 30 species, ranges from Mexico to Central America, the West Indies and northern South America. It bears a striking resem- blance to Galium but its flowers are involucrate. Literature: Ehrendorfer, F. 1955. A revision of the genus Relbunium (Endl.) Benth. et Hook. (Rubiaceae-Galieae). Bot. Jahrb. Syst. 76: 516—533. l. Relbunium t dE (L.) Hemsley, Biol. Centr. Amer., Bot. 2: 63. 1881.—Fic. 83. Vaillantia hypocarpia L., Syst. Nat., ed. 10, 1307. 1759. TYPE: eg Browne (?LINN 1219.13). Rubia incana H.B.K., Nov. Gen. Sp. Pl. 3: 338. "9 TYPE: not se R. hispida Willd. ex Spreng., Syst. Veg. 1: 397. 1825. Relbunium hypocarpium var. incanum (H.B.K.) as in Mart., Fl. Bras. 6(6): 113. 1888. Herbs creeping, branching freely, the stems weak, 4-angled, 0.7-1.0 mm wide, the angles thickened, villose, the nodes usually well spaced. Leaves 4 per node, sessile, oblong, obovate oblong, oblong rotund, 0.3—1.3 cm long, 0.2-0.7 cm wide, deltoid to rounded at the apex, minutely apiculate, obtuse at the base, the costa evanescent above, villose, plane and usually evident beneath, the secondary veins ca. 3, evanescent above and beneath, arcuate, stiffly chartaceous, smooth, usually glandular shiny above, often marcescent, white hirsute, the hairs to 0.7 mm long, often confined to the costa above, glandular punctate; stipules probably connate, scarcely measurable. Flowers solitary in the axils, the pedicels articulate with the bracteoles, to 0.8 cm long, filamentous, usually rigid, the bracteoles 4, im- mediately subtending the flower, lanceolate to ovate oblong, 2.8—4.0 mm long, hirsute; hypanthium ovate rotund, the calyx cup and teeth absent; corolla white, the tube cylindrical, short, the lobes 4, similar in shape to the bracteoles, shorter than and alternate with bracteoles, glabrous except usually marginal hairs; sta- mens 4, the anthers oblong, obtuse, the filaments short, attached at the mouth; style short, the stigmas 2, short, obtuse. Fruits fleshy, usually didymous, orange red, rotund, 2-3 mm long, often wider than long, glabrous to pubescent. 75 For more complete synonymy see Ehrendorfer (155). 460 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 WE N N a TE Klein FIGURE 83. Relbunium hypocarpium (L.) Hemsl. Habit (x24). [After Woodson & Schery 442. | 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 461 Relbunium hypocarpium is common in the mountains of Mexico, Central America, West Indies, and South America. CHIRIQUI: Cerro Horqueta, 7000 ft, Blum & Dwyer 2658 (MO). Cerro posi to Las Nubes, 6000 ft, Correa 1344 (MO). N of Cerro Punta, Croat 10454 (MO). Las Nubes, ca. 2000 m, Croat 26417 (MO). Cerro Punta Bajo Grande, Croat & Porter 16002 (MO). Bajo ao Boquete, ca. 6000 ft, Davidson 428 (F, MO). La Popa above Boquete, 5500 ft, D'Arcy & D'Arcy 6361, 6442 (both MO). Cerro Horqueta, ca. 1500 m, Duke et al. 13699-A (MO). Finca Collins, Boquete, ca. 5000 ft, Dwyer & Hayden 7703 (MO). Cerro Punta, Folsom 4019 (MO). Boquete to 3 mi N, Lewis et al. 379 (GH i : , US). El Barú, above 11,000 ft, Tyson & Loftin 6160 (MO). Rio Chiriquí Viejo Valley, White 73 (F, MO). Finca Lerida to Pena Blanco, 1750-2000 m, Woodson & Schery 335 (F, MO). Potrero Muleto to ped Volcán de Chiriquí, 3500-4000 m, Woodson & Schery 442 (MO). Boquete, Woodson & Schery 771 (MO). Casita Alta, Woodson et al. 827 (NY). Loma Larga to summit, Volcán de Chiriquí, Woodson et al. 1076 (F, MO, NY), 1089 (MO). 72. RICHARDIA Richardia L., Sp. Pl. 1: 330. 1753, not Richardia Kunth, Mém. Mus. Hist. Nat. 4: 433, 437, 1818. (=Zantedeschla Sprengel), not Richardia Adans. Fam. 2: 158. 1763. Nomen rejic. Richardsoniana Kunth ex H.B.K., Nov. Gen. Sp. Pl. 3: 350 Schiedia Bartl. Ord. 210. 1830, not Cham. & Schl. 1826; not pom 1834. TYPE: S. martinicensis .) A. Richard. Tertrea DC., Prodr. 4: 481. 1830. TYPE: T. martinicensis DC. Plants herbaceous, the stems terete, usually densely pubescent. Leaves with the blades usually wide; stipules adnate to the petioles, the sheath multisetose. Inflorescences terminal, capitate, sessile, subtended by conspicuous foliose in- volucral bracts. Flowers with the calycine cup turbinate or subglobose, the lobes 4-8, connate. Fruits partly superior, dry, the cocci indehiscent; seeds with the strophiole very narrow and short, the cells of the testa isodiametric to contorted. Richardia is known from the United States, Mexico, Central America, South America, and the West Indies. There are about 5 species in the genus. Literature: Lewis, Walter H. & Royce L. Oliver. 1974. Revision of Richardia (Rubiaceae). Brittonia 26: 271—301 Richardia scabra L., Sp. Pl. 330. 1753. TYPE: Mexico (LINN 451.1) Fig. 4.— FIG. 84 Richardsonia scabra bred faece Pl. Us. Bras. 8: 1. Richardia pilosa Ruiz & P , Fl. Peruv. Chil. 3: 50. Toy TYPE: Peru, Pavón (MA). Richardsonia pilosa (R. QP) H LB. K., Nov. Gen. Sp. Pl. 3: 350. 9. Richardia cubensis A. Rich. in Sagra, Fl. Cub. 2: 31. 1853. Type: Cuba, Franqueville (F, K). Annual herbs, erect to decumbent, often sprawling, branching loosely, hir- sute. Leaves ovate to oblong lanceolate, rarely linear lanceolate, 1.5—6.5 cm long, 0.3-1.5 cm wide, obtuse to acute at the apex, attenuate at the base, scabrous 462 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 84. Richardia scabra L. Habit ( x!⁄4). [After D'Arcy 5387. | above and beneath, more so on the lateral veins; petioles to 5 mm long; stipules with the sheath ca. 2 mm wide, the marginal setae to 5 mm long. Inflorescences terminal, capituliform, the flowers numerous, to 20, the involucral leaves in 2 pairs, the larger pair widely ovate and estipulate. Flowers with the hypanthium rotund, the calycine tube 0.35-0.50 mm long, the lobes 5—6, unequal, subulate or linear oblong, 1.7-2.3 mm long, acute to subobtuse, the margin ciliate, the hairs to 0.7 mm long; corolla white, occasionally lavender, the tube funnel shaped, ca. 4.5 mm long, constricted below the middle, glabrous, the lobes 5, oblong, 0.5— 2.5 mm long; stamens 6, the anthers subrotund, 0.5 mm in diam.; stigmas 3, slender, ca. | mm long, the style ca. 4 mm long, the ovary 3-loculate. Fruits with mericarps 3, the adaxial face closed, with a narrow groove, the abaxial face papillose, more or less strigose. Richardia scabra is a common weed ranging from North America through Central America and the Greater Antilles to northern South America east to Guyana. ZONE: Miraflores, Bristan P-6 (MO). Sosa Hill, Duke 4653 (MO). Frijoles, Mn d 627 (MO). Gee Golf Club, Nee 7298 (MO). cHiRIQUÍ: Volcan, Croat 10406 (MO). El Hato Del Volcan, Croat 10699 dapi Volcan, Pid. 9168 (MO). Llanos Francia, 4 mi from Boquete, ns yer & Hayden 7618 (GH US). Llan ancia, near Boquete, Stern et al. 1178 (MO). 2 mi SE of Guabala, i & He " 4240 (MO). prd Chiriquí Viejo Valley, White 237 (MO). CocLE: 20 mi P of Nata, D'Arcy & à vat 4119 (MO). El Valle, Folsom 3134 (MO). NE of El Valle, Lewis et al. 1810 (GH, MO "US. mi E of Nata at Rio Grande, Tyson 5294 (MO). Las Margaritas and El Valle, Woodson et al. oe eT COLÓN: Rio B. road ne Portobelo, Blum et al. 2547 (MO). Lo NTO Monagre Beach, 5 mi SE of Chitre, Tyson et al. 3012 (MO). PANAMA: Rio ast Duke 11782, 12413 (both MO). Cerro Jefe, Dwyer & Gauger 7326 (MO): Gentry 2139 (MO). i N of Goofy Lake, Oliver et al. 2688 (MO). Río Pacora to Chepo, Porter et al. 5149 (MO, A "UO. Chepo, Wilbur & Luteyn 11797 (MO). vERAGUAS: Laguna La Yeguda, 15 mi N of Calobre, Luteyn 1450 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 463 73. RONDELETIA Rondeletia L., Sp. Pl. 172, 1753. TYPE: R. americana L. Petesia P. Br., Civ. Nat. Hist. Jam. 143, tab. 2 & 3, 1756. TYPE: not designated. Lightfootia Screb., Gen. Pl. 122, 1789, not Swartz 1788, not L’Her. 1788. TYPE: not given. Willdenovia J. F. Gmelin, Syst. Nat. 2: 362, 1791, not Thunb. T TYPE: W. schreberi Gmel. t1 Rogiera Planch., Fl. Serres 5: 442, 1849. TYPE: R. Arachnothryx Planch., Fl. Serres 5: 442. 1849. TYPE: inn " buddleioides Benth. Shrubs or trees. Leaves coriaceous to membranous, often acuminate; stipules interpetiolar, free, mostly entire. Inflorescences terminal or axillary, usually pa- niculate, rarely thrysoid, rarely racemose or spiciform, pedunculate. Flowers with the hypanthium rotund or oblong, the calycine cup with 4—6(—7) lobes, the lobes occasionally foliaceous; corolla tube narrowly cylindrical, hairs present or absent within, occasionally the mouth with a ringlike thickening, the lobes 4-6, imbricate; stamens 4—6, the anthers oblong, dorsifixed, the filaments often un- equal; ovarian disc annular, the style slender, the stigmas 2(—3), the ovary 2(-3) celled, the septum complete or incomplete, the ovules numerous. Fruits capsular, globose or rotund, the valves 2, splitting loculicidally or septicidally, each valve often split; seeds numerous, exalate or winged at one or both poles. Rondeletia, with about 125 species, is found in the West Indies, Mexico, Central America and South America. The author acknowledges assistance from Kirkbride. Literature: Kirkbride, J. 1969. A revision of the Panamanian species of Rondeletia (Rubi- aceae). Ann. Missouri Bot. Gard. 3: 372-391 The key below has been adapted from Kirkbride's key. a. Leaves 0 ————— EE 9. R. odorata var. breviflora e Leaves with dense white indumentum benea Leaves arachnoid-tomentose beneath; ums lobes 10-13 mmlong |... cdm MEE 6. R. darienensis cc. Leaves white tomentose beneath; calycine lobes to 2 mm long... as Danun beneath. d. MN cte FE stipules reflexed 9 l. R. amoena dd. Leave membranous or chartaceous; ae ules erect. Flowers 4-merous; fruit ie (not seen in R. d'arcyi and R. kirkbri- dei), loculicid al in R. secundi f. Calycine lobe ad in length ne lobes inconsistent as to one Mies 2 times E ag T c the Bre . darcyi gg. Calycine lobes unequal i in n length, with one 2 2c or more seca aie than the other 3. The ne ais lobe 2.0-7.5 mm long, capsule strigose or glabro i. Hypanthium densely strigose or puberulen Hypanthium densely strigose, na ‘tube 9-11 me long, white |. . - 4. R. cooperi 464 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 J. Hypanthium puberulent; corolla tube a 20 m long, pink or lavender |... R. kirkbridei ii. Hypanthium arachnoid tomentose. Small calycine lobes linear or narrowly oblong, 1.7-4.0 mm long; capsule loculicidal _____________- OPEET S d l3. R. sec unda kk. Small calycine AN ee 0.3- : 8 mm lon capsule septici - 2. R. hertieroidss hh. The large Saipan lobe 6. 0- 11. 5 mm long. ES 12. R. salicifolia ff. Calycine lobes about equal in length ` ll. R. platysepala ee. Flowers 5-merous; fruits loculicidal. l. Calycine lobes 1-2 mm long; corolla tube 6-9 mm long . PE 7. R. hamelifolia ll. Calycine lobes ca. 8.5 mm nad corolla tube 11- 18 mm E CUR . R. panamensis 1. Rondeletia amoena (Planch.) Hemsley, Diag. Pl. Nov. 26. 1879. e amoena Planch., Fl. Serres 5: 442. 1849. TYPE: not seen. R. menechma Planch., Fl. Serres 5: 442. 1849. TYPE: not seen. Rondeletia versicolor J. Smith, Bot. Mag. tab. 4579. 1851. TYPE: not s R. aa Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1852: 43. 1852. TYPE: not R. Wisaa Bentham ex Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn 1852: 43. 1852. TYPE: not s Rogiera latifolia Decne., Rev. Hort. sér. 4, 2: 121. 1853. TYPE: not seen. R. versicolor Lindley & Paxton, Fl. Gard. x 69. 1853. TYPE: not se Rondeletia apa K. Krause, Bot. Jahrb. Syst. 40: 315. 1908. TYPE: not seen. Trees to 10 m tall, the branchlets terete, drying dull golden tan, cracking, glabrescent. Leaves ovate, ovate rotund, 9.0-21.5 cm long, 4.0-12.5 cm wide, regularly more than 2 times longer than wide, rounded or obtuse at the apex, the acumen deltoid, to 1 cm long, ultimately acute or obtuse, often falcate, subtrun- cate or cordate at base, the costa slender, immersed or prominulous above, prom- inent beneath, the lateral veins 5-10, the median veins to 1.5 cm apart, usually strict, the venules prominently reticulate beneath, stiffly chartaceous to subcoria- ceous, usually bicolorous, rarely subbullate above, glabrous above except along costa and veins or appressed strigose or golden villose above, densely golden villose to glabrate beneath; petioles occasionally minute or to 1.5 cm long, to 0.3 cm wide, stout, golden pubescent; stipules reflexed, deltoid, 4—17 mm long, 3-10 mm wide, obtuse, sericeous outside, glabrous within except toward the margin. Inflorescences terminal or axillary, paniculate, to 17 cm long, the peduncle to 9 cm long, ferrugineous pilose, glabrescent, the bracteoles several. Flowers sessile or subsessile, the pedicels velutinous; hypanthium rotund, ca. 2 mm long, fer- rugineous to white tomentose, the calycine cup ca. 0.5 mm long, the lobes 5-6, unequal, deltoid to deltoid oblong, to 0.5 mm long, occasionally wider than long; corolla white, the tube 9-15 mm long, tomentose outside, yellow pilose within near the mouth, glandular hairy toward the base within, the lobes 5—6, to 3.5 mm long, glabrous within, tomentose outside; stamens with anthers oblong, ca. 2 mm long, attached near middle of tube, the filaments to 2 mm long. Fruits loculicidally dehiscent, rotund to compressed rotund, to 6 mm in diam., tomentose to glabrate. Rondeletia amoena extends from Chiapas, Mexico, to Panama. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 465 RIQUÍ: Volcan de Chiriquí Viejo, Allen 994 (MO). ‘New Switzerland," Valley of Rio Chiriqui Viejo. "Allen 1350 (MO). Llano del Volcan, Allen 3469 (MO). Finca Collins, Boquete, Blum & Dwyer 2553 nee is 1 km from Hato del Volcán, road to Concepcion, Correa & Lazor 1436 (MO). Methodist Camp, Nueva Swissa, Croat 13519 (MO). N of Audubon Cabin, Croat 13610 (MO). Monte Rey above Boquete, po 15718 (MO). 2.5 mi N of Río Chiriqui Viejo, Croat 15885 (MO). Volcán de Chiriquí, Boquete, Davidson 951 (MO). E slope Volcán de Chiriqu uí, WNW Boquete, Davidse & D'Arcy 10180 (MO). 3 km NE EI Hato del Volcán, 1800 m, Davidse & D'Arcy 10413 (MO). Valley of Río Chiriquí Viejo, N of Volcan City, Duke 9067 (MO). Cerro Horqueta, Duke et al. 13650 (MO). Cerro Horqueta, Dwyer et al. 556 (MO). Boquete, Dwyer 7000 (MO). Between Pinola and Quebrada Seco on Chiri- quicito-Caldera Trail, Kirkbride & Duke 1026, 1036 (both MO, NY). Alto Lino, 4000-5000 ft, Maurice 839 (MO). W slope Volcan de Chiriqui, trail to summit, 2500-3400 m, Mori & Kallunki 5740 (MO). W slope Volcan de Chiriqui, lava flow, ca. 1500 m, Mori & Kallunki 5700 (MO). 17 km NE of San orado, 33773 (MO). La Mesa, Tyson 818, 6057 (both MO). Rio Chiriqui Viejo Valley, White & White 19, 35, 231 (all MO). Finca Lerida, Woodson & Schery 227, 311 (both MO). Casita Alta, Volcán de Chiriquí, Woodson et al. 810 (MO). 2. Rondeletia bertieroides Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 267. 1929. TYPE: Panama, Cooper 598 (F, holotype; NY, isotype). Trees or shrubs, the branchlets ultimately tomentose. Leaves lanceolate, oc- casionally ovate, 5.5-23.0 cm long, 1.5-6.0 cm wide, conspicuously acuminate at the apex, rarely simply acute, cuneate to acute at base, the lateral veins 5-7, arcuate, usually stiffly chartaceous, discolorous. Inflorescences terminal, rarely axillary, paniculate, the peduncle 2-5 cm long or absent, the branches helicoid cymose or with reduced dichasia, the basal branches 1-8 cm long, tomentose to glabrate. Flowers 4-merous, the pedicels minute or to | mm long, tomentose, the hypanthium rotund, to 1.5 mm long, the calycine cup to 0.5 mm long, | lobe elliptic or obovate, (1.3-)2.0—4.2 mm long, 0.8- 2.7 mm wide, acute to occasion- ally obtuse at the apex, the other 3 lobes triangular or oblong, 0.3-2.0 mm long, 0.5-1.0 mm wide, the corolla white, the tube 1.0-1.2 cm long, strigose outside, puberulent within below the anthers, the lobes obtuse, ca. 2.5 mm long; anthers 0.2-2.7 mm long, the filaments short; ovary with the septum entire or incom- pletely fused. Capsule globose, ca. 4 mm in diam., glabrous, septicidal, the ca- lycine lobes persistent. s DEL ToRO: Buena Vista Camp, Chiriquí trail, 3000 ft, Cooper 598 (NY). Between Buena Vista Coffee Finca and Cerro Pilon, Chiriqui Trail, Kirkbride & Duke 712, 713 (both MO). Between Criollo and Quebrada Higueron, uem Trail, Kirkbride & Duke 799 (MO). cocLÉ: Cerro Pilon, Croat 14402, 22944 (both MO). L esa, Dwyer 8316 (MO). Cerro Pilon, Dwyer & Lallathin 8682 (MO). Cerro Gaital Caracoral, b r & Correa 2 (MO). Cerro Caracoral, Kirkbride 1124 (MO). Cerro Pilon, Liesner 788 (MO); Porter et al. 4401 (MO), 4411 (MO, SCZ, UC, VEN). La Mesa, 2 m W of Cerro Pilon. Sullivan 513 (MO). Cerro Pilon, 5 km NE of El Valle, Mori 6557 (MO). 3. Rondeletia buddleioides Benth., Pl. Hartw. 69. 1840. TYPE: Mexico, Hartweg 502 (B, if extant, not seen, photo MO; NY, isotype). Arachnothryx buddleioides (Benth.) Planch., Fl. Serres 5: 442. 1849. Rondeletia affinis Hemsley, Diag. Pl. Nov. 28, 1879. Type: Panama, Seemann 1594 (K, not seen). Trees to 15 m tall or shrubs, the branchlets ultimately white tomentose to glabrescent. Leaves ovate, elliptic or obovate, 2.5-20.0 cm long, 1.0-7.5 cm wide. acute to acuminate at the apex, the acumen short, the base cuneate to attenuate acute, the smaller veins perpendicular, the costa prominulous beneath, the lateral veins 10—14, arcuate, prominent and conspicuous, sparsely webby 466 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 tomentose to glabrous above, often asperous above, densely white tomentose beneath; petioles to 2 cm long, usually white tomentose; stipules polymorphic, generally linear to ovate, 4-10 mm long, 0.5-5.0 mm wide, acute. Inflorescences terminal or pseudo axillary, white tomentose to glabrous, the peduncle 0.5-9.0 cm long, the cymules often helicoid. Flowers sessile or subsessile, 4-merous, the hypanthium rotund, to 1.5 mm long, densely tomentose, the calycine lobes deltoid or oblong, 0.5-2.0 mm long, reflexed; corolla white, cream or pink, the tube 6— 11 mm long, pubescent within, sparsely villous within below the middle, the lobes 2-3 mm long, papillate adaxially; stamens 4, the anthers oblong, ca. 1.5 mm long, the filaments very short, attached at middle of the tube: ovarian disc to 0.5 mm in diam., glabrous. Fruits subrotund, 3-4 mm in diam., septicidal, the hairs few. a. Upper surface of the leaves nonasperous.......... 3a. var. buddleioides aa. Upper surface of the leaves asperous sb var. aspera 3a. Rondeletia buddleioides Benth. var. buddleioides CANAL ZONE: Fort Sherman, Pina Highlands, Hayden 120 (MO). cuiRiQUÍ: Finca Lerida, Que- brada Velo. Allen 4753 (MO). Finca Collins, Blum & Dwyer 2423A (MO). Buena Vista mt Chiriqui Trail, Mie o 615 (NY). Monte Rey above Boquete, Croat & Porter 15675 (MO). Boquete, D'Arcy & D'Arcy 6468 (MO). Bajo Chorro, Boquete, Davidson 231 (MO). Volcán de Chiriquí, Davidson 918 (MO). Bun Davidson 1000 (MO). Cerro Hor rae Duke 13622, A13728 (both MO). Cerro Hor- queta, NW of Boquete T az et al. 551 (GH US). Boquete, Dw zer 6955, 7004 (both. MO). Boquete, Llanos Francia, Dwyer & Hayden n T UC, VEN), 7626 (MO). Finca Collins, Bo- Kirkbride & Duke 864 (MO, NY). Between Pinola and Quebrada Seco, Chiriquicito-Caldera Trail, Kirkbride & eid 1023 (MO, NY). Finca Collins, Boquete, Srern & Chambers 1128 (MO). Llanos Francia, Boquete, Stern et al. 1199 (MO). Finca Collins, Boquete, Stern et al. 2037, 2044 (both MO). Bambito, Dion 3865 (MO). Cerro Horqueta, Hagen & Hagen 2151 (MO, NY). Upper Río Chiriquí Viejo, White & White 28, 30 (both MO). San Ramon, Bajo Mono, Wilbur et al. 13546 (MO). Callejon Seco, Volcán g n ard s & Schery 493 (MO). Casita Alta, Volcan de CE Woodson et al. 930 (MO, NY). c : 4 a from El Valle, Correa & Dressler 1003 (MO). Cerro Pilon and El Valle, Duke & Diver 13945 (G . MO, UC, US). Las Margaritas and El Valle, Woodson etal. 1280, 1757 (both MO). PANAMA: e Campana, Allen 2650 (MO). Cerro Azul, Blum & Dwyer 2423A (MO). Cerro Jefe, E 17363 (MO). Cerro Campana, Duke 5997, 8680 (both MO). Cerro Azul. Dwyer 1880, 2069 (both MO). Cerro Jefe, Dwyer & Ge ntry 9426B (MO). Cerro Campana, Dwyer & Kirkbride 7814 (COL, D MO, UC, VEN). Cerro Azul, Ebinger 394 (MO). zd Jefe, Folsom 3814 (MO). vA Campana, Lewis et al. 1912 (GH, US); McDaniel 6812 (MO). Cerro Azul. Porter et al. 4118 (MO, SCZ, UC, VEN). El Valle to La Mesa, ca. 2400-2600 ft. Spellman > al. 583 (MO). Cerro Azul to Cerro Jefe, Tyson et al. 4328 (MO 3b. Rondeletia buddleioides Benth. var. aspera Kirkbride, Ann. Missouri Bot. Gard. 55: 379. 1968. TYPE: Panama, Kirkbride & Duke 780 (MO, holotype; NY, isotype). CAS DEL TORO: Buena Vista Camp, Chiriqui Trail, 1000 m, Cooper 65 (F, SROI Between Criollo and Quebrada Higueron on Chiriquí Trail, Kirkbride & Duke 780, 796 (both MO, NY). 4. Rondeletia cooperi Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 267. 1929. TYPE: Panama, Cooper 600 (F, holotype; isotype, NY). Trees or shrubs, to 7 m tall, the branchlets terete, strigose, densely so ter- minally, the internodes variable in e Leaves elliptic, 10-22 cm long, 4.0- 9.5 cm wide, acuminate, the acumen to 2.5 cm long, attenuate to cuneate at the base, the costa prominent above, the lateral veins ca. 10, papyraceous, strigose 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 467 on the veins but glabrescent above, strigose on the veins beneath; petioles to 3 cm long, strigose; stipules triangular, often compressed triangular, ca. 4 mm long, sericeous strigose outside, sericeous and with digitiform processes within, to 0.8 mm long. Inflorescences terminal, paniculate, the branches compound dichasia, 10-25 cm long; peduncles variable in length, 3-17 cm long, strigose. Flowers sessile or pedicellate, the pedicels ca. 1 mm long; hypanthium ferrugineous stri- gose, the calycine cup ca. 0.5 mm long, the lobes variable in length, 1—2 lobes ca. 3 times longer than the others, the smaller lobes oblong to triangular, ca. | mm long, strigose outside; corolla white, the tube ca. 10 mm long, strigose out- side, the lobes ca. 4 mm long. Fruits capsular, ovoid, to 4.5 mm long, costate, sparsely strigose, septicidal to about the middle, the calyx lobes persistent. Rondeletia cooperi is known only from Panama. BOCAS DEL TORO: Buena Vista Camp, Chiriquí Trail, 416 m, Cooper 600 (F, NY). CHIRIQUÍ: Punta Pena, ca. 333 m, Lewis et al. 2158 (MO). CocLE: Between Buena Vista coffee finca and Cerro Pilon, Chiriqui Trail, Kirkbride & Duke 703 (MO, NY). 5. Rondeletia darcyi Dwyer.'*° rype: Panama, D'Arcy 4079 (MO, holotype). Shrubs?, the twiglets and leaves dark red, the twigs slender, terete, smooth, glabrescent, the nodes well spaced. Leaves oblong elliptic or ovate elliptic, 2-13 cm long, 1.2-5.0 cm wide, deltoid or acute toward the apex, abruptly acuminate, the acumen to 2.5 cm long, often falcate, the base obtuse or truncate, the costa prominulous beneath, the lateral veins ca. 6, arcuate, prominulous beneath, the submarginal vein ca. 3 mm from the margin, rigidly membranaceous, concolor- ous, glabrous above, beneath with weak and often appressed hairs especially on the costa and the veins, the hairs swollen basally; petioles to 0.5 cm long, the hairs few; stipules free, triangularly subulate, ca. 3 mm long, rigid, erect, the cusp often longer than the body, the hairs similar to those of the leaves. /nflo- rescences terminal, compressed cymose paniculate, to 5.5 cm long, to 3.3 cm wide, the peduncle slender, to 3.5 cm long, to 0.8 mm wide, the hairs weak, appressed, white, the branches slender, to 1 cm long, the inferior branches op- posite; bracts subulate, to 4 mm long, the flowers few. Flowers with the pedicels to 4 mm long; hypanthium oblong rotund or compressed rotund, to 1.2 mm long, drying red, the hairs few, appressed, the calycine cup very short, the lobes 4, narrowly oblong, 2.34.2 mm long, unequal, sometimes with one 2 times longer than the others, unequal in width, 0.25-0.80 mm wide, thick, the hairs few; corolla white. the tube narrowly salverform, to 12 mm long, ca. 0.5 mm wide in the middle, expanded toward the apex, thinly carnose, the hairs few, ascending, glabrous on both sides except puberulent toward the base within, the lobes 4, compressed hemispherical in bud, oblong, to 4 mm long, obtuse; stamens 4, the 130 Rondeletia darcyi Dwyer, spec. nov. Frutices? ramulis glabrescentibus. Folia oblongo-ellip- tica vel ovato-elliptica, 2-13 cm longa, 1.2-5.0 cm lata, acumine ad 2.5 cm longo, venis lateralibus ca. 6, rigido-membranacea supra glabra subtus ciliis debilibus saepe appressis; petiolis ad 0.5 cm longis: stipulis ca. 3 mm longis. /nflorescentiae cymoso-paniculatae, ad 5.5 cm longae, pedunculo ad 3.5 cm longo. Flores pedicellis ad 4 mm longis; lobis calycis angusto-oblongis, 2.3-4.2 mm longis, o 3 m: corolla alba tubo ad 12 mm longo, lobis 4, ad 4 mm longis; antheris ca. 3.2 mm longis; disco ovarii nullo, stylo 4.0-4.8 mm longo. Fructus non visi. 468 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 anthers oblong, ca. 3.2 mm long, obtuse, the filaments short, ca. 0.4 mm long, attached to the tube below the mouth; ovarian disc absent, the style slender, 4.0- 4.8 mm long, glabrous except puberulent toward the apex, the stigmas 2, digiti- form, 1-2 mm long, puberulent. Fruits not seen. Rondeletia darcyi is known only from Panama. It is named in honor of Wil- liam D'Arcy, student of the flora of Panama and one who assisted the author greatly in editing this treatment. The most important diagnostic character of this new species is the unequal length of the calycine lobes. The longest lobe in the complex of 4 is not 2 times longer than the others but is quite variable. Four sets of lobe lengths, measured in millimeters, were discerned: a) 2.3, 2.6, 3.0, 4.2; b) 2.3, 2.5, 4.0, 4.0; c) 2.5, 2.6, 3.0, 3.6; d) 2.8, 3.0, 3.6, 3.8. The new species seems closely related to R. secunda Standley whose calycine lobes are twice as long as the other 3 lobes and are consistently wider than in R. darcyi. The new species has the stamens at- tached just below the corolla mouth rather than below the middle of the tube and the branches of the inflorescence are not helicoid. COLÓN: Road bank just W of Portobelo, D'Arcy 4079 (MO). 6. Rondeletia darienensis Standley, N. Amer. Fl. 32: 53. 1918. TYPE: Panama, Pittier 4684 (US, holotype; F, isotype). Shrubs?, the branchlets terete, webby tomentose. Leaves ovate to elliptic, 7.5-17.0 cm long, 2.5-7.0 cm wide, acuminate, basally cuneate or obtuse, the costa prominent beneath, the lateral veins prominent beneath, the blades arach- noid tomentose; petioles 0.3-2.0 cm long, tomentose; stipules ovate, ca. 0.8 mm long, the sheathing base 5-6 mm wide, adaxially glabrous, tomentose along the margin and sericeous in the center. Inflorescences axillary, thrysoid, 4-5 cm long, the peduncle 1.0—1.5 cm long, arachnoid tomentose; bracts 2, 3-4 mm long; bracteoles | or 2. Flowers with the pedicels to 1 cm long, white tomentose: hypanthium oblong, ca. 3 mm long, ca. 2 mm wide, the calycine lobes 4, lanceo- late, 1.0-1.3 cm long, 0.2-0.4 cm wide, acute, glabrous within; corolla densely tomentose. Fruits capsular, ca. 4 mm long. Rondeletia darienensis is known only from a single collection in Panama. DARIÉN: Boca de Paurandó, on Sambü River. southern Darién, 20 m, Pittier 4684 (F. US). = Rondeletia hamelifolia Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 144. 1967. TYPE: Panama, Dwyer 1350 (MO, holotype; NY, isotype). Shrubs or trees to 8 m tall, the branchlets terete, diffusely white pilose, the nodes often somewhat swollen. Leaves elliptic or elliptic rotund, 5-18 cm long, 2.3-6.0 cm wide, deltoid and widely acuminate at the apex, basally subcuneate, the blade somewhat decurrent on the petiole, the costa prominulous, the lateral veins 6—9, strongly arcuate ascending, prominulous beneath, stiffly papyraceous, discolorous, tending to dry gray green beneath, pilose above and beneath, the hairs above few, white, appressed, densely crowded on the veins; petioles oc- casionally absent, 0.2-2.0 cm long; stipules triangular, 4-10 mm long, 2-4 mm 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 469 wide, the cusp 2—5 mm long, sericeous inside and out, bearing several oblong glands within, these to | mm long, drying black. Inflorescences terminal, cymose paniculate, 4-13 cm long, to 10 cm wide, the peduncle 1-2 cm long, the primary branches paired, few, well spaced, 0.5-1.2 cm long, the branchlets ternately disposed above the middle of the branches, the cymules few flowered or often with the flowers solitary; bracts and bracteoles persistent, triangular subulate, golden pubescent. Flowers sessile or on pedicels to 3 mm long, golden pubescent; hypanthium rotund, ca. 2 mm long, densely golden pubescent, the calycine lobes 5, oblong or triangular subulate, subequal, ca. 1.5 mm long, obtuse at the apex; corolla white, the tube salverform, to 5 mm long, ca. | mm wide at the apex, yellow within, the mouth scarcely annular, thickened, the lobes 5, rotund, ca. 4 mm long, rounded or obtuse at the apex; stamens 5, the anthers linear oblong, ca. 2 mm long, the filaments short, attached near the middle of the tube. Fruits oblong rotund, to 0.8 cm long, loculicidally dehiscent, the ribs scarcely promi- nulous, puberulous. Rondeletia hamelifolia is known only from Panama. LOS SANTOS: Canofistulo, Dwyer 2458 (MO). Los Asientos, Wendehake 38 (MO). VERAGUAS: San Francisco near Rio San Francisco, Dwyer 1272 (MO). Santiago, Dwyer 1350 (MO, NY). San- tiago, 14 mi from Panamerican Highway toward Atalaya, Dwyer & Kirkbride 7410 (GH, MO, US), 7422A (MO). 2 mi W of Santiago on Panamerican Highway, Dwyer et al. 7556 (GH, MO, US). Between Tolé and Santiago, Kirkbride & Hayden 164 (MO). Alto Piedra, Lao 471 (MO). Canazas, Tyson 3638 (MO: SCZ). 8. Rondeletia kirkbridei Dwyer.'*! rype: Panama, Mori & Kallunki 6112 (MO, holotype). Trees small, to 5 m tall, the branchlets angular, minutely pubescent, the nodes well spaced. Leaves ovate lanceolate, 10-16 cm long, 4.5-5.5 cm wide, often slightly inequilateral, attenuate apically, often deltoid or acute, acuminate, the acumen wide or narrow, to 2 cm long, acute to cuneate at the base, the costa prominulous above and beneath, the lateral veins 7-8, strongly arcuate, evanes- cent near the margin, the smaller veins often plane or immersed, papyraceous, concolorous, glabrate above, moderately puberulent on the costa and the veins; petioles to 2.5 cm long, to 0.2 cm wide, rigid, glabrescent; stipules connate, appressed to the stem or spreading, ovate triangular or triangular, to 0.7 cm long, the awn to 2 mm long, coriaceous, puberulent outside, within densely white villose. Inflorescences terminal, paniculate, 20-24 cm long, to 9 cm wide; pe- duncle 9-12 cm long, to 0.2 cm wide, minutely puberulent, the rachis with 6-8 pairs of branches, these opposite or subopposite, the lowermost branches to 2.5 cm long, angular, spreading. Flowers with the hypanthium oblong, ca. 2 mm long, puberulent, the calycine cup ca. 0.5 mm long, puberulent outside, with a minute gland in the interdental sinus, the lobes 4, subequal or unequal, the 3 shorter lobes triangular, 0.7-1.2 mm long, the fourth lobe oblong lanceolate, 2.5-4.5 mm 131 Rondeletia kirkbridei Dwyer, spec. nov. Arbores ad 5 m. Folia pin lanceolata, 10—16 cm longa, ad 5.5 cm lata, acumine ad 2 cm longo, venis lateralibus 7-8 subtus osta venisque puberula; petiolis ad 2.5 cm longis, stipulis connatis ad 0.7 cm longis seta ad 2 mm longa. /nflorescentiae ee 20-24 cm longae, ad 9 cm latae; aoe ad 12 cm longo. Flores cupula calycis ca. 0.5 onga, extus puberula lobis 4, 3 parvis a 1.2 mm longis, uno magno 2.5-4.5 mm longo; corollae ee ad 2 cm longo; antheris ca. 3 mm long 470 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 long, ca. 1.5 mm wide, venose; corolla pink or lavender, the tube infundibuliform, to 2 cm long, in the middle to 1.2 mm wide, puberulent outside, within glabrous above, puberulent below the middle, the lobes (3—)4, ovate oblong rotund, ca. 5 mm long, obtuse, glabrous; stamens 3-4, the anthers sessile, included, rectangular oblong, ca. 3 mm long, obtuse, attached at the middle of the anther; style linear, 6-8 mm long, the stigmas 2-4, linear oblong, 1.3-2.0 mm long. Fruits not seen. Rondeletia kirkbridei is known only from Panama. It is named in honor of Joseph Kirkbride, Jr., an authority on the genus Rondeletia. While the new species is closely related to R. cooperi Standley, it is distinguished by its elongate pink or lavender corolla tube and stipules which are externally glabrate. It is also related to R. skutchii Standley, a red-flowered species in Guatemala which has few lateral veins in the leaves, a pendent inflorescence, and a much shorter corolla. VE apa NW of Santa Fé, 1 km from Escuela, Cerro Tute, Mori & Kallunki 4780 (MO). d of Santa Fé, Mori & Kallunki 4849 (MO). 16 km NW of Santa Fé, road to Calovébora, 300-50 Mori d "Kallunki 6112 (MO). 9. Rondeletia odorata Jacq., Enum. Pl. Carib. 16. 1762. TYPE: not seen. 9a. Rondeletia odorata Jacq. var. breviflora Hooker, Curtis Bot. Mag. tab. 6350. 1878. TYPE: not seen. Shrubs, the branchlets terete, ultimately ferrugineous hirsute. Leaves ovate, obovate or elliptic, 1.7-5.0 cm long, 1-3 cm wide, acute at the apex, subcordate basally, the intervenal areas spreading reticulate, coriaceous, scabrous, bullate, sparsely hirsute beneath; petioles absent or to 0.2 cm long; stipules broadly del- toid, 2-4 mm long, 1.5-4.0 mm wide. Inflorescences terminal, narrowly thrysoid; peduncles 0.3-0.7 cm long; ferrugineous hirsute; basal bracts 2, foliar, 3-8 mm long, to 5 mm wide. Flowers pedicellate, the pedicels ca. 2 mm long, ferrugineous tomentose; calycine lobes 6, oblong, ca. 4 mm long, ca. | mm wide, glabrous within; corolla bright red, the tube 5-7 mm long, scarcely tomentose outside, glabrous within, the lobes 5, ca. 2.5 mm long, obtuse, the callosity at the mouth ringlike, ca. 0.4 mm thick, ca. 2 mm long; stamens 5, the anthers oblong, ca. 2 mm long, the filaments ca. | mm long, attached at the middle of the tube; stigmas 2-3, the style thick and glabrous. Fruits not seen. Rondeletia odorata var. breviflora is known only from Panama. Rondeletia odorata Jacq. var. odorata is not found in Panama. CANAL ZONE: Matachin to Las Cascadas, Cowell 359 (NY). 10. Rondeletia panamensis DC., Prodr. 4: 408. 1830. TYPE: Panama, Seemann (BM, not seen; MO, photo) Shrubs or trees to 6 m tall, the branchlets terete, white tomentose to glabrous. Leaves elliptic to subovate, 2-15 cm long, 1.3-7.0 cm wide, acute to subacumi- nate at the apex, attenuate basally, the lateral veins 6-9, strongly arcuate as- cending, stiffly papyraceous to subcoriaceous, discolorous, the tertiary veins per- pendicular; petioles scarcely measurable or to 1.2 cm long, white tomentose to 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 471 glabrate; stipules deltoid, 3-5 mm long, 2.0-6.5 cm wide, the cusp ca. 2 mm long, tomentose to glabrate. Inflorescences terminal, axillary, the peduncle to 4 mm long, the dichasia compound, tomentose to glabrate. Flowers pentamerous, ped- icellate, the pedicels to 7 mm long; hypanthium small, rotund, ca. 2.5 mm long, truncate, white tomentose, the calycine cup ca. 1.5 mm long, the lobes 5, subu- late, ca. 8 mm long, ca. 2.5 mm wide, petaloid, densely sericeous within at the base; corolla white, the tube 11-18 mm long, petaloid, villosulose outside, mod- erately so within except dense near the mouth, the latter with a ringlike callosity, the lobes 4, subrotund, ca. 5 mm in diam., 1 lobe ca. 3 mm in diam., anthers oblong, ca. 2 mm long, the filaments ca. 2 mm long; ovarian disc ringlike, ca. 3 mm deep, the style linear, ca. 8 mm long. Fruits rotund, to 1 cm in diam., sparsely tomentose, dehiscing loculicidally, the calycine lobes conspicuous, persistent. Rondeletia panamensis extends from Panama to Colombia. CANAL ZONE: Quebrada Fea, Quebrada Pura and Canon of Rio Chagres, Dodge & Allen 17413 ). Road C-21, N of Corozal, Dressler 3143 (MO). Albrook-Ft. of Clayton Dump Road, Duke & Mussell 6622 (MO). Quarry Heights, Dwyer 2607 (MO). Albrook Site, USATTC, Dwyer 7123 (MO). Curundu, Thunder Hill, McDaniel 5189 (MO). Balboa, Standley 26996 (MO). Curundu, Tyson & of Chepo, D’Arcy 6031 (MO). Cerro Azul, hd Arcy & D'Arcy 6221 (MO). Jenine, Duke 3843, 4482 (both MO). Río Charco-Espirito on Tocumen Highway, Duke 5706 (MO). Río Mamoni, Duke 5686 (MO). Cerro Azul, Dwyer 1485 (MO). Río iidem Waterfall, 2nd main fork, Gentry 2284 (MO). Icanti, Tyson 6839 ( MC, 11. Rondeletia platysepala Standley, Ann. Missouri Bot. Gard. 27: 343. 1940. TYPE: Panama, Allen 1791 (F, holotype; GH, MO, NY, isotypes). Trees to 6 m tall, the branches with numerous branchlets, the twiglets slender, terete, smooth, puberulent. Leaves narrowly elliptic, 2.8-8.0 cm long, 0.7-2.2 cm wide, acute to deltoid at the apex, acuminate, the acumen to 1.5 cm long, ultimately acute, basally acute, the costa lightly prominulous above and beneath, the lateral veins 4—5, strongly arcuate, chartaceous, lustrous and sparsely to- mentose to glabrescent above, glabrous to minutely farinose beneath except for small appressed hairs; petioles 1-3 mm long; stipules triangular, to 3 mm long, the cusp ca. 0.75 mm long with several digitiform processes ca. 0.5 mm long, drying black. Inflorescences terminal, cymose paniculate, puberulent, the pe- duncle 1-4 cm long, ca. | mm wide, slender, flexuous; bracts narrowly oblong or ovate oblong, to 3 mm long, acute, pubescent, deciduous. Flowers 4-merous, the pedicels absent or to 5 mm long; hypanthium oblong, ca. 1.5 mm long, to- mentose, the calycine lobes equal elliptic, oblong elliptic or ovate elliptic, 1.5— 4.2 mm long, to 2.5 mm wide, tomentose outside, glabrate within; corolla pale pink, the tube 7-11 mm long, tomentose outside, pubescent within below the middle; stamens included, the anthers oblong, 2 mm long, the filaments very short, attached near the middle of the tube. Fruits sessile, oblong or oblong rotund, to 0.6 cm long, 6-costate, glabrous, drying dark brown, the ribs thin, 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 white, woody, the intercostal area white venose, the 2 carpels septicidally dehis- cent to base, each valve slightly loculicidally dehiscent, the calyx finally decid- uous. Folsom et al. 4711, juvenile and in mature fruit, seems to belong to R. pla- tysepala. The sepals are equal in length, which supports this. The sepals of Fol- som 4711, located only on the young fruit, are about twice the length of those found on the type collection, Allen 1791. The fruits seem much larger than those of R. bertieroides Standl., a species easily confused with R. platysepala. Rondeletia platysepala is known only from Panama. CHIRIQUÍ: Cerro Colorado, 1500-1750 m, Folsom et al. 4711 (MO). cocLé: El Valle, North Rim, Allen 1791 (F, GH, MO, NY). 12. Rondeletia salicifolia Dwyer & Hayden, Phytologia 15: 58. 1967. TYPE: Pan- ama, Tyson et al. 3319 (MO, holotype).—Fic. 85. Trees 4—8 m tall, the branchlets terete, smooth, the hairs slender, golden, the nodes well spaced. Leaves lanceolate to elliptic, 3-20 cm long, 2-9 cm wide, the larger blades 3-4 times longer than wide, deltoid to subobtuse at the apex, acu- minate, the acumen to 2 cm long, 0.15-0.2 cm wide at the middle, usually finally acute or subacute, basally cuneate to subobtuse, the costa prominulous to grooved above, subprominent beneath, convex, the lateral veins ca. 7, arcuate, prominulous, stiffly chartaceous, concolorous, drying chocolate brown, minutely golden puberulent, especially on the midvein and lateral veins: petioles moder- ately slender, 0.5-1.0 cm long; stipules deltoid to widely triangular, to 4 mm long, the terminal awn shorter than the body, arising from a ringlike thickening, curving arcuately, the body stiff, subcoriaceous, puberulent. Inflorescences terminal or axillary, paniculate; peduncle to 6 cm long, the branches cymose and occasionally secund, to 12 cm long, tomentose. Flowers sessile or shortly pedicellate, tomen- tose; hypanthium rotund, ca. 2 mm long, the calycine cup ca. 0.5 mm long, sparsely hairy outside, glabrous within, the lobes 4, with 1 lobe 3—4 times longer than the others, the larger lobe elliptic or ovate, 6-12 mm long, to 7 mm wide, acute, the remainder elliptic to deltoid, 0.7-3.5 mm long, to 3 mm wide, acute; corolla white, the tube 8-15 mm long, glabrous within, glabrate outside; anthers ca. 3 mm long, the filaments to 2 mm long; style much shorter than or equal to the corolla tube. Fruits rotund, to 4 mm in diam., costate, the hairs deciduous. 12a. Rondeletia salicifolia var. salicifolia is known only from Panama. There is a so-called type photo of R. salicifolia H.B.K. in the Missouri Botanical Gar- den; the original specimen is in the Herbier H&B in Paris, but this H.B.K. binomial is unpublished. Gentry 9449 (MO). Cerro Azul to Cerro Jefe, Foster 1139 (MO). Finca Indio, slopes of Cerro Jefe, Gentry 2154 (MO). Cerro Jefe, Gentry & Mori 13445 (MO). Cerro Jefe, 800 m. Gomez-Pompa et al. 3069 (MO). 16 km above Panamerican Highway, road from El Llano to Carti-Tupile, Kennedy et al. 2432 (MO). Altos de Rio Pacora, 833 m, Lewis et al. 2315 (MO). Finca Indio, slopes of Cerro Jefe, Webster et al. 16478 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 473 FiGure 85. Rondeletia salicifolia Dwyer & Hayden.—A. Habit (<'%).—B. Flower (x12).—C. Flower opened to show interior (x2!2).—D. Ovary cross section (x6). [After Gomez-Pompa 3069. } 12b. Rondeletia salicifolia subsp. brevicolla Kirkbride, Ann. Missouri Bot. Gard. 55: 383. 1963. TYPE: Panama, Kirkbride & Duke 597 (MO, holotype; NY, isotype). Stipules with cusp 3.0-6.5 mm long; leaves sparsely tomentose on the veins beneath, the tertiary veins disposed at right angles to the lateral veins. Flowers 474 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 with the corolla tube ca. 8 mm long; septum of the ovary altogether connate, the placenta elliptic. Rondeletia salicifolia subsp. brevicolla is known only from Panama. BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 98 (F). e Teribe between Quebrada Huron and Quebrada Schlunjik, 100 m, Kirkbride & Duke 467 (MO, NY). 13. Rondeletia secunda Standley, Contr. U.S. Natl. Herb. 18: 141. 1916. TYPE: Panama, Pittier 4279 (US, holotype; NY, isotype). Shrubs, the branchlets terete, tomentose to glabrous. Leaves narrowly ellip- tic, rarely subovate, 7-16 cm long, 3-7 cm wide, often acuminate, the acumen to 1.7 cm long, basally rounded, truncate or acute, the lateral veins 7-8, arcuate, prominent beneath, stiffly chartaceous, concolorous, glabrate above and beneath, occasionally with sparsely distributed, weak, appressed hairs; petioles 1.6—6.0 mm long, glabrous to pubescent; stipules connate, the free portions triangular, 3-6 mm long, 2.0-3.5 mm wide, cuspidate, the cusp to 3.5 mm long. /nflores- cences terminal, paniculate, the peduncles 4.0—6.5 cm long, the branches helicoid cymose, to 10 cm long, the flowers secund, well spaced. Flowers subsessile or with pedicels to 0.5 mm long, sparsely arachnoid tomentose; hypanthium rotund, ca. 1.5 mm in diam., sparsely arachnoid tomentose, the calycine cup to 0.3 mm long, the lobes 4, unequal, | lobe narrowly obovate, 3.6-7.5 mm long, 2 times the length of the other 3 lobes, the latter linear or narrowly oblong, 1.7-4.0 mm long, 0.2-0.6 mm wide; corolla white, the tube funnel shaped, 9-12 mm long, sparsely villose outside, the lobes 2-3 mm long, obtuse, papillate inside; ovarian disc ca. 0.1 mm long, glabrous. Fruits capsular, rotund, ca. 4 mm in diam., scarcely costate, glabrous, rugulose, the seeds rectangular, the calyx persistent. Rondeletia secunda is known only from Panama. The fruits of Dwyer 5138 are smaller than those of Duke & Bristan 337 and Duke et al. 3657. It may be a different species: it alone of all the material cited was collected on the Pacific side of the Isthmus. : Cocalito, Dwyer 5138 (MO). SAN BLAs: Perme, Cooper 229 (F, NY). E of Puerto Ob- aldia. Gan 16919 (MO). Ailigandi River, Duke & Bristan 337 (MO). oe of Rio Cuadi along river, Duke et al. 3657 (MO). Near Puerto Obaldia, Pittier 4279 (NY, 74. RUDGEA Rudgea Lum Trans. Linn. Soc. London 8: 327. 1807. TYPE: R. lanceaefolia Salis Strempelia A. Rich. ex DC., Prodr. 4: 498. 1830. TYPE: S. guianensis A. Rich. ex DC. Shrubs, occasionally small trees. Leaves often coriaceous or subcoriaceous, petiolate or sessile; stipules interpetiolar, often with subulate or prickly teeth on the dorsal surface or on the margins, the margins occasionally laciniate. /nflo- rescence terminal, paniculate, cymose, umbellate or capitate, occasionally I-flowered. Flowers with the calycine cup obsolete or obvious, the teeth present or absent; corolla salverform or funnelform, the lobes valvate; ovary 2-celled, the ovules connate, attached to a common basal column. Fruits baccate; seeds vertical, commonly only one present in the mature fruit. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 475 Rudgea occurs throughout the American tropics and in the West Indies. It is a genus of about 150 species. Literature: Steyermark, J. 1967. Rudgea. Botany of the Guayana Highland—Part VII. Mem. New York Bot. Gard. 396—425 a. vibe sessile, at least some auriculate and/or amplexicaul at the base; largest blades wide I. R penus aulis aa. icd petiolate, not wipe or ampem; largest blades 4.5—-10.0 cm wide. argest leaf blades 5-11 cm lon 2. R. chiriquiensis - [o] = c 2 c N 3. R. cornifolia ee. Leaves with 8-11 lateral veins; stipules ca. | cm long _____- 4. R.« coronicarpa Fruits oblong, | cm or more long 5. R. isthmensis cc. Largest ae blades 20-22 cm long. f. ves coriaceous; ae to 1.5 cm long 6. R. pittieri ff. Leaves chartaceous; stipules 0.6-0.9 cm long --------------- 7. R. skutehi 1. Rudgea amplexicaulis Dwyer.'*? TYPE: Panama, Croat 33133 (MO, holotype). Scandent shrubs to 1.5 m tall, the branches terete, moderately slender, smooth, glabrous, the nodes well spaced, scarcely turgid. Leaves sessile, nar- rowly obovate oblong, 5-15 cm long, 2-4 cm wide, acute to deltoid at the apex, truncate at the base, at times vaguely auriculate and amplexicaul, the costa prom- inulous on both sides, the lateral veins arcuate, the undulate submarginal vein only to 3 mm from the margin, about 2 smaller veins between 2 lateral veins, arising at right angles from the costa and soon branching, the intervenal areas patulous reticulate, chartaceous, scarcely discolorous, glabrous; stipules decid- uous, connate, cup shaped, to 1.5 mm long, scarious, scarcely swollen, glabrous with a few oblong glands, ca. 0.5 mm long. /nflorescences terminal, few flowered, glabrous, cymose paniculate, to 4 cm long, ca. 0.8 mm wide, the branches dis- posed as 2 well spaced, opposite pairs, each bearing one 3-flowered cymule; bracts patulous, ovate subulate, to 1.3 mm long. Flowers with the pedicels to 3 mm long; hypanthium oblong, ca. 2 mm long, glabrous, black when dry, the calycine cup ca. | mm long, the lobes triangular, to 0.35 mm long, acute, glabrous: corolla white, the tube narrowly cylindrical, scarcely attenuate basally, glabrous outside, the lobes 5, oblong, 5-6 mm long, minutely apiculate and vaguely cu- cullate. Fruits not seen. U? Rudgea amplexicaulis Dwyer, ae nov. Frutices scandentes ramulis glabris. Folia sessilia angusto-obovato-oblonga, 5-15 cm lon 2—4 cm lata, basi truncata interdum modice au p et amplexicauli, chartacea vix discoloria . stipulis deciduis connatis cupuliformibus hic ad 1.5 mm longis, vagina sine dentibus scariosa vix inflata glabra glandulis oblongis paucis, ca. 0.5 mm longis, ornata. /nflorescentiae terminales cymoso-paniculatae glabrae, ad 4 cm longae, pedunculo ud ad | cm longo, cymulis trifloris; bracteis patulis ovato-subulatis, ad 1.3 mm longis. Flores pedic v H 3 mm longis: hypanthio glabro cupula calycis ca. 1 mm longa, lobis 5 pes: triangularibus, ad 0.3 longis, glabris; corolla alba tubo cylindrico extus glabro, lobis 5, 5-6 mm longis. Fructus non visi. 476 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Rudgea amplexicaulis is known only from Panama. It is immediately distin- guished by its narrow leaves, some of which are amplexicaul at the base. The sprawling habit of the shrub is unique at least for the Central American species of Rudgea. HIRIQUÍ: Above San Félix along mining road 18-27 mi off Panamerican Highway above Chame or hsa to Escopeta, 1200-1500 m, Croat 33133 (MO). 2. Rudgea chiriquiensis Dwyer.'? rype: Panama, Croat 37017 (MO, holotype). Shrubs to 4 m tall, the branchlets terete, glabrous, the nodes moderately turgid, well spaced. Leaves oblong, 5-11 cm long, 1.5-5.0 cm wide, acute to subobtuse at the apex, acuminate, the acumen often falcate, ultimately acute, the base acute to vaguely obtuse, slightly inequilateral, the costa and the veins above plane, scarcely prominulous beneath, the lateral veins 6-8, arcuate, the submar- ginal vein scarcely distinguishable, the intervenal areas patulous reticulate, thinly chartaceous, somewhat discolorous when dry, glabrous; petioles slender, to 1.5 cm long, ca. 0.1 cm wide, glabrous, often rigid; stipules perhaps connate basally, widely triangular or subhemispherical, to 3.5 mm long, the marginal or submar- ginal glands few, oblong, to 1 mm long, the sheath drying black. Inflorescences terminal, cymose paniculate, glabrous, 3-7 cm long, 2-10 cm wide, the peduncle slender, 1.0-1.2 cm long, with 3 branches arising at the apex of the peduncle, these arcuate or angular ascending, 1-4 cm long, each branch terminated by 3 short and spreading branchlets, the cymules 3-flowered; bracts subulate, to 2 mm long, the bracteoles ovate, to 1.2 mm long. Flowers sessile, the hypanthium ca. | mm long, black, glabrous, the calycine cup ca. 0.5 mm long, truncate, the margin undulate or with 5 widely triangular, scarcely measurable teeth; corolla (in bud) ca. 7 mm long. Fruits not seen. Rudgea chiriquiensis is known only from Panama. It is distinguished by its small leaves and reduced stipules. CHIRIQUÍ: Cerro Colorado, along road above San Félix, 29 km above bridge over Río San Félix, 7.9 km above turnoff to Escopeta, disturbed primary forest, 1500 m, Croat 37071 (MO) 3. Rudgea cornifolia (H.B.K.) Standley, Field Mus. Publ. Bot. 432. 1931.—Fic. 86 Psychotria cornifolia H. °. “Q Nov. Gen. Sp. Pl. 3: 362. 1819. rvPE: Venezuela, Humboldt & Bon- pland 893 (P), not se P. s" ss: Benth., Hook. London J. Bot. 3: 226. 1841. Type: British Guiana, Schomburgk 51, not en. Rudge a micrantha Muell.-Arg., Flora 69: 454. 1876. rype: Brazil, Martius 3007 (Photo type from M R. ceratopetala Donnell Smith, Bot. Gazette 35: 3. 1903. rype: Guatemala, Tuerckheim 7904 (F. olotype). R. fimbriata (Benth.) Standley in Standley and Calderon, List PI. Salvador., 274. 1920. 199 . a C: Dwyer, spec. nov. Frutices ad 4 m alti ramulis glabris. — oblonga, 0 5-11 cm lon 1.5-5.0 cm lata, costa venisque supra planis venis lateralibus 6-8, tenui-chartacea glabra; denm ad 1.5 cm poc ix ies fortasse connatis, ad 3.5 mm longis. Inflores scentiae termi- nales cymoso-pan niculatae, ad 7 cm longae, ad 10 cm latae, cymulis 3- Lliorbus: bracteis subulatis, ad 2 mm lo n Flores hypanthio ca. | mm longo, glabro, cupula calycis truncata margine undulato vel dentibus 5 minutis ornato 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 477 EM ay’ (SES Bie d E 86. e s b iae (H. B. K.) Standley.—4A. Habit (x'2).—B. Flower, part of peri- anth d (x915).— ary, cross section (X9!2).—D. Fruit (x'2). [After Allen 288.] Strempelia ceratopetala (Donnell Smith) Brem., Recueil Trav. Bot. Néerl. 31: 303. 1934. S. fimbriata (Benth.) Brem., Recueil Trav. Bot. Néerl. 31: 303. 1934. Trees to 7 m tall, the branchlets slender, smooth, glabrous, nodose, the nodes well spaced. Leaves oblong to lanceolate, occasionally obovate, 8-14 cm long, to 7 cm wide, obtuse or rounded at the apex, occasionally subacute, acuminate, the acumen to 1.7 cm long, to 0.3 cm wide, obtuse to cuneate at the base, oc- 478 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 casionally vaguely auriculate, the costa slender, prominulous above and beneath, occasionally with pores (‘‘domatia’’) in the vein axils, the lateral veins 7-8, at first strict, then arcuate, joining adjacent veins to form an undulate submarginal vein to 1.5 cm from the margin, chartaceous, concolorous, glabrous; petioles absent or to 0.7 cm long, glabrous to minutely pubescent; stipules connate, com- pressed rectangular, to 3 mm long, truncate, subcoriaceous, glabrous, the margin thickened with ca. 6 triangular subulate lobes, to 0.75 mm long. Inflorescences terminal, solitary, glabrous, paniculate; peduncles 3.5—4.0 cm long, usually longer than the remainder of the inflorescence, either with 4 upright branches umbel- lately disposed and once ramose or the rachis with 2 pairs of opposite branches, to 2 cm long, dichotomous, each branchlet with a single few flowered cymule. Flowers glabrous; hypanthium oblong or rotund, the calycine cup cylindrical, ca. 2 mm long, carnose, the teeth as evanescent points; corolla white, to 7 mm long, the tube basally attenuate, the lobes 5, cucullate, longer than the tube; anthers 5, narrowly oblong, ca. 3 mm long, obtuse, dorsifixed, the filaments short; stigmas 2, capitate, the style narrowly cylindrical, thicker above the middle, the ovary wall and septum thick. Fruits white, rotund or compressed rotund, to 5.5 mm long, to 6.5 mm wide, smooth, glabrous, minutely farinose often vaguely 2-lobed, the calycine scar annular, ca. 1 mm in diam. Rudgea cornifolia occurs in Middle America and in South America south to Bolivia. AS DEL TORO: Fish Creek Mts., Wedel 2301 (US). Chiriquí Id d Mn (US). CANAL ZONE: " Frijoles, Standley 27506, 27591 (both US). COLON: Santa Rita Ri m . 8.5 km Boyd- lu Highway, Mori E "aypay 2081 (MO). Santa Rita Ridge mi “pss, L ua Highway, Porter et Hi 4758 (MO, SCZ). DARIEN: Pinogana and Yaviza, vis 288 (MO), 927 (US). El Real. Duis 4836 (MO). Eine Piriaque, Duke 8085 (MO). Santa Fé, Duke 8407, 10249, 10697, 10700, 14262, 14290 (all MO). Río Perecenico and Rancho Frio, Duke & Elias 13878 (MO). Casi-Cana Trail, Cerro p» ampamento E of Tres Bocas, Kirkbride & Duke 1221 (MO). N slope of Cerro Pirre, ca. 300-700 m, Mori & Kallunki 5394, 5514 (both MO). El Real, Stern et al. 120 (MO, US). Cerro Ad IUE et d 3840 (MO). Río Lara, Tvson & pe 3851 (MO). Santa Fé, Dion * A 4718 (MO). P MÁ: Pas o Jefe, Dwyer et al. 7240 e SCZ, US); Dwyer & Gauger 7369 (G , MO, AS: "Rio Diabolo and Río Asuati near Margana, Duke 14872 (MO). 3-4 s a up Río Mui tape: Kirkbride 232 (MO). El Llano- Cai Road, 24.5-25.0 km Hote Interamerican Highway, Mori & Kallunki 5530 (MO) 4. Rudgea coronicarpa Dwyer.'* TYPE: Panama, Gentry & Dwyer 3647 (MO, holotype). Small trees or shrubs, the twiglets angular, smooth, glabrous, the nodes well spaced but crowded at the very tips of the twigs. Leaves oblong or obovate oblong, 12-18.5 cm long, 4.5-10 cm wide, obtuse or widely deltoid at the apex, with a conspicuous acumen, to 1.5 cm long, often falcate, ultimately subacute, basally obtuse, truncate or scarcely auriculate, occasionally the blade constricted on lower !⁄4, the costa prominulous above, prominent beneath, the lateral veins 8-11, arcuate, subprominent beneath, drying somewhat rubescent, the intervenal Rudgea coronicarpa Dwyer, spec. nov. Arbores parvae, ramulis glabris. Folia oblonga vel d ara 12-18.5 cm longa, 4.5-10 cm lata, venis lateralibus 8-11, rigido-chartacea: petiolis nullis ad 0.5 cm e stipulis liberis oblongis, ad 1 cm longis, obtusis calloso-fimbriatis fimbriis crebris b 0.4 mm longis. /nflorescentiae ad 8.5 cm longae; pedunculo 4.0-5.5 cm longo, ramis 1—2 arcuato- soa Aenea unilateraliter dispositis. un tus oblongi vel rotundi, ad 0.9 cm longi 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 479 areas spreading reticulate, stiffly chartaceous, glabrous, the margin somewhat thickened; petioles obsolete or to 0.5 cm long, thick, to 2.5 mm wide, glabrous; stipules free, persistent only apically, erect, oblong, to 1 cm long, ca. 0.4 cm wide, obtuse, callose fimbriate at the apex, the fimbriae crowded, to 0.4 mm long, coriaceous, glabrous. Inflorescences terminal, glabrous, to 8.5 cm long; pedun- cles 4.0-5.5 cm long, slender, ca. 0.3 cm wide, angular, glabrous, dark green when dry, the peduncle terminated by 1—2 arcuate, ascending branches, the latter unilaterally disposed, each terminated by a sessile or subsessile fruit (flowers not seen), or with pedicels to 3 mm long, the one or two fruits arising below the terminal flower on slender, glabrous pedicels to 1.4 cm long. Fruits sessile or on slender pedicels to 1.4 cm long, oblong or rotund, at maturity to 0.9 cm long including the persistent calyx, smooth, glabrous, minutely farinose, the persistent calyx absent or to !4 the length of the fruit. Rudgea coronicarpa is known only from Panama. It is related to R. cornifolia but it has much larger stipules and a larger fruit. : Lumber camp at Alto ^ d 7 km N of El Cope, Folsom 1309 (MO). Cerro Pilon, ca. 2700 P "Gentry & Dwyer 3647 (M 5. Rudgea isthmensis Standley, Ann. Missouri Bot. Gard. 26: 313. 1929. TYPE: Panama, Woodson et al. 1618 (MO, NY, isotypes). Trees to 4 m tall, the branchlets slender, the nodes well spaced. Leaves lance oblong, 5-13 cm long, 1.5-7.5 cm wide, deltoid at the apex, acuminate, the acu- men usually falcate, cuneate basally, the lateral veins 6-8, the submarginal vein ca. | cm from the margin, stiffly papyraceous, drying yellow, glabrous; petioles slender, to 1.3 cm long, glabrous; stipules ca. | cm long. Inflorescence cymose paniculate, glabrous, to 3.5 cm long, much shorter than the terminal leaves, pedunculate, the rachis with 2 sets of widely divergent opposite or whorled branches. Flowers not seen. Fruits subsessile, large, oblong, to 1 cm long, round- ed at the apex and base, sulcate, glabrous. Rudgea isthmensis is known only from Panama. CANAL ZONE: Salmanca Hydrographic Station, Rio Providencia and ridge S of river, Gentry & Nee 8699 (MO). Fort Sherman, Tyson & Dwyer 1191 (MO). Pequeni, Woodson y v ` I8 (F. MO. NY). SAN BLAS: Between Río Diabolo and Río Acuati near NAE Duke 14882 6. Rudgea pittieri Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 211. 1940. TYPE: Panama, Pittier 3855 (F, holotype, US).—Fic. 87. Shrubs 5—8 m tall, the branchlets 4-angled, ashen gray when dry, the nodes conspicuous, swollen, the internodes to 2 cm long. Leaves widely oblong or rotund, 13-22 cm long, 5.0-10.5 cm wide, deltoid or rotund toward the apex, the acumen absent or when present to 0.5 cm long, attenuate acute at the base, the costa prominulous above, prominent beneath, to 1.3 mm wide proximally, the lateral veins 10-12, arcuate, somewhat prominulous, the margin revolute, cori- aceous, concolorous, shiny, glabrous; petioles thick, 1.5-3.0 cm long, to 0.25 cm wide; stipules free, compressed rotund, to 1.5 cm long, thick, coriaceous, gla- brous, with an elevated rectangular portion extending beyond the body, rounded 480 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 87. Rudgea pittieri Standley. Habit (x'2). [After Porter 4803.] at the apex, and bearing 6—12 oblong glandular fimbriae to 2.5 mm long. Inflo- rescences terminal, solitary, the flowers congested into a rotund mass, to 3 cm in diam., sessile or the peduncle to 1.3 cm long, to 0.45 cm wide; external bracts ovate oblong, ca. 1.5 cm long, acute, glabrous, stiffly carnose, dark brown, the interior bracts similar but narrowly lanceolate. Flowers with the hypanthium short, the calycine cup short, the margin with 5 irregular, scariose, erose, trian- gular teeth, 0.5-2.5 mm long; corolla white, the tube cylindrical, ca. 1 cm long, thickly petaloid, glabrous outside, glabrous within except for the throat, the hairs 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 481 dense, the lobes 5, narrowly oblong, ca. 8 mm long, acute or obtuse, cucullate; stamens 5, the anthers oblong, 1.8 mm long, the filaments short; style linear, 15 mm long, the stigmas 2, | mm long. Fruits sessile, oblong, to 1.5 cm long, sub- truncate or rotund, orange, black when dry, smooth, glabrous, often marcescent. Rudgea pittieri is known from Panama and Colombia (Choco). Standley’s origi- nal description was based on poor material, although foliage and stipule characters are enough to suggest the genus Rudgea and are diagnostic at the species level. In some ways the plant resembles a Cephaelis but the character of its stipules pre- cludes that genus. The thick coriaceous leaves are unlike those of any Rudgea of Middle America. ANAL ZONE: Near Maru Towers NW part of Canal Zone, Johnston 1553 (MO). Santa Rita ee road, 15 km E of Colón, Dressler 3801 (MO). COLON: Tres Brazos sawmill, Icacal between Salud and Bocas de Rio Indio, Howell 88 (MO). Santa Rita Ridge, ca. 10 mi from the Interamerican Highway, Porter et al. 4803 (MO). Achiote, Tyson et al. 4537 (MO); Tyson 4532 (MO). PANAMÁ: E Llano-Cartí Road, 7-18 km from Interamerican Abas Correa et al. 1857 (MO); Croat 25125 (MO); Folsom 2556 (MO). W of El Ee 4.8 mi N of highway, Gentry 5102 (MO). Cerro Jefe, 18.2 km E of Goofy Lake, 800-900 m, Gentry & Mori 13427 (MO). El Llano-Cartí Road, 16-1812 km N of Panamerican Highway, Nee & Tyson 11000. 10955 (both MO). 7. Rudgea skutchii Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1374. 1938. TYPE: Costa Rica, Skutch 2836 (F. holotype; isotypes NY, US). Trees 2-5 m tall, the branchlets smooth, lustrous, glabrous, the nodes well spaced. Leaves oblong to obovate oblong, occasionally salicoid, 8.5-20.0 cm long, 3-10 cm wide, obtuse or rounded at the apex or occasionally truncate, the acumen narrowly deltoid, conspicuously falcate, to 2 cm long, to 0.25 cm wide in the middle, ultimately acute or obtuse, apiculate, the apiculum occasionally to 0.3 mm long, obtuse, rounded or truncate at the base, occasionally vaguely au- riculate, subequilateral, the costa plane to prominent above, to 0.2 cm wide, the lateral veins 7—9, arcuate, the larger veins to 2.5 cm apart, prominulous above and beneath, the intervenal areas reticulate, chartaceous to papyraceous, scarcely discolorous, often with minute subepidermal raphides above, glabrous beneath; petioles 0.3-0.8 cm long, ca. 0.3 cm wide, minutely puberulent; stipules free, appressed, oblong or rotund, 0.6—0.9 cm long, about as long as wide, obtuse with about 25 crassate, linear oblong, fimbriate excrescences at apex, these to | mm long, glabrous. /nflorescences terminal, solitary, glabrous, cymose paniculate, to 6 cm long, to 6 cm wide, the peduncle to 4.5 cm long, 0.1 cm wide, glabrous, often slightly curved, angular, the primary branches 2-3, to 1.5 cm long, the cymules few, 3-flowered. Flowers sessile or pedicellate, the median flower ses- sile, the lateral flowers on pedicels to 1 cm long, glabrous, the floral parts gla- brous, the hypanthium obconic, ca. 2 mm long, the calycine cup to 5 mm long, truncate or with 5 widely triangular teeth, to 0.4 mm long, the margin scarious; corolla white, the tube cylindrical, to 7 mm long, stiffly carnose, the lobes 5, narrowly oblong, 7-8 mm long, with a subapical adaxial projection, narrowly oblong, ca. 2 mm long, the stamens 5, the tips of anthers barely exserted, the anthers narrowly oblong, to 3.5 mm long, obtuse, the filaments short, inserted in the upper 25 of the tube, the stigmas included in tube, the style slender, the stigmas erect (at least in bud), subulate. Fruits not seen. 482 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Rudgea skutchii is restricted to Costa Rica and Panama and Choco, Colombia. It resembles Rudgea killipii Standley from Colombia (holotype Killip & Smith 15246 [US]). While the inflorescences and calycine cups of the two species are strikingly similar, the leaves of R. killipii are scarcely acuminate in contrast to those of R. skutchii. Croat points out (27207) that the flowers are ‘‘sweetly aro- matic.” BOCAS DEL TORO: Río Tuquesa ca. 2 km by air from Continental Divide, near Tyler Kittredge gold mining camp, Croat 27187, 27209 (both MO). Forest above railroad stop at Milla 7.5, Croat & Fag 16271 (M - Punta Pena, near Chiriquicito, Lewis et al. 2143 (COL, MO, UC, VEN), 2166 , UC). cocLé: La Mesa near El Valle, Allen 2699 (MO). N of El Cope, 500 m, D'Arcy 11302 e Rivera seni 600—800 m, Folsom 3149 (MO). PANAMÁ: Cerro Campana, Dressler 4127 (MO). 75. RUSTIA Rustia Klotzsch in Hayne, Arzn. Gew. 14: pl. 14. 1846. LECTOTYPE: R. formosa (Cham. & Schlecht.) Kl. (Exostem(m)a formosum Cham. & Schlecht.). Trees or shrubs, the branchlets glabrous. Leaves glandular punctate; stipules large, deciduous. /nflorescences terminal, paniculate. Flowers pedicellate, the pedicels bracteolate; hypanthium campanulate or turbinate, the calycine cup with 5 lobes or teeth; corolla funnelform or campanulate, the tube glabrous to hairy within, the lobes 5, valvate; stamens 5, the anthers basifixed, opening by a ter- minal slit or pore, the filaments attached at the throat; ovarian disc terete or sulcate, the style dilated medially, clavate at the apex or the 2 stigmatic lobes conspicuous, the ovary 2-celled, the ovules numerous, attached vertically to the septum. Fruits capsular, coriaceous, oblong ovoid, the 2 valves separating loc- ulicidally; seeds numerous, minute, horizontal, the testa membranous. Rustia, a genus of about 15 species, ranges from Mexico, Central America to South America. It also occurs in the West Indies. From a floral viewpoint it is the easiest genus to recognize among the Panamanian Rubiaceae, as the anthers open by terminal pores. The pellucid-punctate leaves are also unique. Literature: Simpson, D. 1976. Studies in Neotropical Rubiaceae. 1. Rustia. Phytologia a. Largest leaves not exceeding 9 cm in width; corolla purple or red; fruits ca. 10 mm long . R. occidentalis aa. Largest leaves 20-19 cm wide; corolla white; fruits ca. 5 mm long |... 2. R. panamensis 1. Rustia occidentalis (Benth.) Hemsl., Biol. Centr. Amer. 2: 14. 1881. Exostemma i ei EE Benth., Bot. Voy. Sulphur 104. 1844. type: Panama, Barclay and/or Hinds (K, holotype). Shrubs or small trees, to 5 m tall; branchlets terete, smooth, rimose, often cracking, glabrous. Leaves narrowly oblong to obovate oblong, often falcate, regularly 3 times longer than wide, to 22 cm long, to 9 cm wide, acute or deltoid at the apex, acuminate, the acumen to 3 cm long, usually falcate, basally acute, 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 483 the costa plane or slightly depressed above, prominent beneath, the lateral veins ca. 15, arcuate, those in the middle of the blade 1.0-2.5 cm apart, stiffly papy- raceous, scarcely discolorous, the intervenal areas with scattered glandular punc- tations, glabrous; petioles slender, to 4 cm long, to 0.2 cm wide, usually angular, the lamina decurrent alate, glabrous; stipules not seen. Inflorescences terminal, solitary, paniculate, occasionally racemiform, shorter than the uppermost leaves, the peduncle absent or 0.5-2.0 cm long, the lateral branches short or to 6.5 cm long, angular, opposite (at least the 2 lowermost pairs), the flowers solitary on reduced ultimate branchlets or in spreading cymules on the upper !^ of the branchlets. Flowers with the hypanthium cuneate, to 5 mm long, ca. 2.5 mm wide, glabrous, the calycine cup ca. 0.5 mm long, the margin erose, toothless or with 5 vague, undulate areas; corolla purple or red, the tube compressed cylin- drical, ca. 7.5 mm long, ca. 3 mm wide at the base, slightly constricted medially, stiffly carnose to coriaceous, glabrous except densely white villose at the filament attachment point, the lobes 4, oblong, slightly shorter than the tube, acute or obtuse, glabrous; stamens 5, the anthers oblong, ca. 6.8 mm long, ca. 2 mm wide, obtuse and porocidal at the apex, obtuse basally, dorsifixed near the base, the filaments short, ca. 0.8 mm broad; stigmas 2, erect, concave adaxially, ca. | mm long, the style cylindrical, ca. 0.9 mm long, ca. 0.7 mm broad, constricted basally. Fruits briefly pedicellate, capsular, obovoid, ca. 1 cm long, to 0.8 cm wide, truncate at the apex with an annular scar, drying dark brown, thin but lignose, delicately longitudinal striate, glandular punctate, splitting to base into 2 valves, each with a conspicuous, median, longitudinal, seminiferous septum. Rustia occidentalis ranges from Costa Rica to Colombia. The fact that the flowers are purple red, the leaves pellucid punctate, and the anthers open by a terminal pore allows for ready identification of the species. Simpson (1976) re- cently treated the genus Rustia as found in Colombia, Ecuador, and Peru. He noted R. occidentalis for Colombia, but he did not mention the Central American distribution of the species. NAMA: BOCAS DEL TORO: Old Bank Island, Wedel 298 (F). CANAL ZONE: Mojinga Swamp, mouth of Río Chagres, Allen 860 (MO). Chagres, Fendler 289 (F, MO). Chagres River, | mi above mouth, Johnston 1745 (MO). COLON: Miguel de la Borda, Croat 9828 (MO). Along Cano Rey, vicinity of Cocle del Norte, Dressler 4207 (MO). DARIEN: Pinas, Duke 10576, 10605 (both MO). Puerto St. Dorothea, Dwyer 2231 (MO), 2290 (F, MO). coLoMBia: 3-7 mi S of Curice, Duke 9653 (MO). 2. Rustia panamensis Dwyer." TYPE: Panama, Croat 27508 (MO, holotype).— Fic. 88 Trees to 12 m tall, the branchlets terete, smooth, glabrous, the scars at the nodes conspicuous, the nodes well spaces. Leaves oblong or obovate oblong, occasionally falcate, 15-45 cm long, 10—19 cm wide, the costa prominulous above, prominent beneath, the lateral veins ca. 15, at first strict then arcuate toward the 75 Rustia paname nsis Dwyer, spec. nov. Arbores ramulis glabris. Folia oblonga, 15—45 cm longa, venis lateralibus ca. 15, rigido-papyracea pellucido-punctata; petiolis ad 6 cm longis. Inflorescentiae cymoso- y eei ad 45 cm longae, ad 26 cm latae. Flores sessiles; calyce dentibus nullis; corolla tubo 0.4-1.0 cm longo, extus glabro; antheris ad 2 mm longis poro dehiscentibus, filamentis ad 1.2 mm longis. Fruc tus subrotundi vel oblongo-rotundi, ad 0.5 cm longi, glabri; seminibus ad 0.8 mm ongis. 484 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 88. Rustia panamensis Dwyer.—A. Inflorescence ( x?/;).—B. Flower opened to show interior (x*/,).—C. Ovary cross section (x1'/,). [After Busey 464. ] margin, prominulous beneath, the veinlets irregularly pinnatiform, stiffly papy- raceous, glabrate or glabrous, pellucid punctate under magnification; petioles to 6 cm long, expanded toward the base, abaxially concave, lignose, glabrous; stipules not seen. Inflorescences terminal, solitary or 2-3, cymose paniculate, spreading, puberulent, to 26 cm long, to 20 cm wide, often with 3 strongly as- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 485 cending branches terminating the peduncle, these 3.0—4.5 cm long, angular, gla- brous, the uppermost branches opposite, well spaced, rigidly patulous or de- flexed, simple or often with short branches, the flowers well spaced, the lowermost bracts deciduous, triangular, to 1.8 cm long, to 0.7 cm wide at the base, elongate attenuate, acuminate toward the apex, rigid, glabrous, the upper- most bracts subulate. Flowers sessile, the hypanthium oblong rotund, glabrous, the calycine cup short, to 0.5 cm long, scarious, petaloid, glabrous and eglandular within, the margin undulate, the lobes virtually absent; corolla white, the tube short, cylindrical, somewhat constricted at the middle, to 10 mm long, to 1.5 mm wide, thickly petaloid, glabrous on the outside and within except near insertion of the filaments, the lobes 4(—5)—6; ovate lanceolate, upright (at least in bud) to 2 mm long, yellow brown; stamens 5, wholly or in part exserted, the anthers oblong, to 2 mm long, dehiscent by a terminal pore, the filaments short, to 1.2 mm long, expanded at attachment point; disc of the ovary strongly compressed rotund, ca. 0.6 mm long, the style to 4 mm long, the stigmas 2, exserted, widely oblong, concave. Fruits scarcely pedicellate, subrotund to oblong rotund, to 0.5 cm long, glabrous, drying purple black, marcescent, lightly venose; seeds com- pressed, falcately fusiform, rhomboid fusiform, ca. 0.8 mm long, shiny, reticulate. Rustia panamensis is known only from Panama. Its pellucid-punctate leaves and porocidally dehiscent anthers immediately distinguish it as a Rustia. The large paniculate inflorescence, extraordinarily large leaves, and very reduced fruits distinguish it from R. occidentalis Hemsley. The variation in the number of stamens (4—6) is noteworthy. CHIRIQUÍ: San Bartolo Limite, 21 km WNW of Puerto Armuelles, Busey 464 (MO). Burica Peninsula, San Bartolo Limite, 12 mi W of Puerto Armuelles, 400-500 m, Croat 2219] (MO). PANAMA: El Llano-Carti Highway, 10-12 km N of El Llano, Dressler 4749 (MO). VERAGUAS: Valley of Río Dos Bocas, 11 km from Escuela Agricola Alto Piedra, 450 m, Croat 27508, 27551 (both MO). Atlantic slope N of Santa Fe, 11 km from Escuela Agricola Alto Piedra, Mori & Kallunki 3086 (MO). NW of Santa Fe, 11 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 3086 (MO). 76. SABICEA Sabicea Aublet, Hist. Pl. Guiane 1: 192. 1775. TYPE: S. aspera Aubl. Climbing shrubs. Inflorescences axillary, sessile or pedunculate, the flowers disposed in bracteate heads or in corymbose or umbellate cymes. Flowers with the hypanthium somewhat ellipsoid, the lobes 3—6, linear or oblong; corolla usu- ally funnel shaped, the lobes 4-5, shorter than the tube, valvate in bud; stamens 4—5, usually included, the anthers dorsifixed, inserted in the tube or in the throat of the corolla; style usually included, the stigmas 4-5, linear; ovules numerous on axile placentae. Fruits fleshy or leathery, 4—5-celled; seeds numerous, minute. Sabicea, a genus of about 125 species, is found in tropical America, Africa, and Madagascar. Literature: Wernham, H. F. 1914. A Monograph of the genus Sabicea, 1-82. 486 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 a. Calycine lobes 2-4 mm long; stipules 3—6(—7) mm long. b. Stems and leaves with hairs closely appressed. i fl h l nflorescences short pedunculate 1. S. panamensis cc. Inflorescences sessile 2b. S. villosa var. adpressa bb. Stems and leaves with hairs spreadin 2a. S. villosa var. villosa aa. Calycine lobes 3-6 mm long; stipules ca. 8 mm long 3. S. stellaris 1. Sabicea panamensis Wernham, Monogr. Sabicea 30. 1914. TYPE: Panama, Fendler 181 (K, holotype).—Fic. 89. S. costaricensis Wernham, Monogr. Sabicea 31. 1914. sYNTYPES: Costa Rica, Pittier 6712, 2904 (both BM). Shrubs or sprawling woody vines, the stems terete, densely pubescent, the nodes well spaced. Leaves elliptic, 7-13 cm long, 3.0-5.5 cm wide, deltoid, rarely obtuse at the apex, often contracted and short acuminate, the acumen widely triangular, cuneate to occasionally obtuse at the base, the costa prominulous above and beneath, the lateral veins 10—14, arcuate, chartaceous, often stiffly so, not strongly discolorous above, the hairs in the intervenal areas stigillose, well spaced or absent, those on the costa and veins stiff, appressed, dense, the hairs in the intervenal areas beneath sometimes shorter than above, the costa and the veins with elongate, appressed hairs; petioles to 2 cm long, to 0.15 cm wide, densely hairy; stipules free, ovate rotund, rotund or compressed rotund, to 7 mm long, often as wide as long at the base, drying red, reflexed, strigose outside, glabrous within. Inflorescences axillary, the peduncle 0.5—1.5(—3.5) cm long, the flowers congested; bracteoles narrowly oblong. Flowers pedicellate, the pedicels slender, 0.3-1.0 cm long; hypanthium densely appressed golden hairy, the caly- cine cup short, the lobes 5, equal or unequal, narrowly ovate, to 4 mm long, the longest often exceeding the hypanthium in length, usually reflexed, glabrescent within; corolla white or yellow, occasionally red or purple pink, often turning brown with age, the tube narrowly funnel shaped or cylindrical, 5-6 mm long, ca. | mm wide in the middle, the hairs outside similar to those of the hypanthium, glabrous within, the lobes 5, subulate or narrowly triangular, equal or subequal, to 2.5 mm long; stamens 5, the anthers sessile or subsessile, oblong, ca. 2 mm long, attached above the middle of the tube; style linear, 2-10 mm long, the stigmas 2, each about as wide as the style, to 1 mm long. Fruits pedicellate, rotund to oblong, 0.5—0.9 cm in diam., pink or red at maturity, the hairs weak, elongate, well spaced, appressed strigose. Sabicea panamensis extends from Guatemala to Colombia. Mori & Kallunki 5603 point out that the flowers are visited by skippers; Duke 9368 indicates that flowers are visited by hummingbirds. Mori & Kallunki 2063 remark that "freshly cut twigs smell like peppermint.” CAS DEL TORO: Santa Catalina, Blackwell et al. 2742 (MO). Old Bank Island, Wedel 1916, 1975 (both MO). Little Bocas, Wedel 2558 (MO). Big Bight, Wedel 2889 (MO). CANAL ZONE: Pipeline Road near Gamboa, Clewell & Tyson 3320 (MO). Hill C-6, Ft. Sher man, Duke 4395 (MO). Coco NW shore of Gatün Lake, Lewis et al. 1831 (GH, M peline Road, Lewis et al. 5465 (MO, SCZ) Mori & Kallunki 2063 (MO). Navy Reservation N of Gamboa, Robyns 65-49 (MO). Fort Sherman Pena Road, Tyson & Chu 1704 (MO). cHiRIQUÍ: 17 km NE of S i Nee 10668 (MO). . I ; ( COCLÉ: Cerro Pilón, 700-900 m, Duke 12109 (MO). El Valle to La Mesa, Dwyer et al. 585 pr Ta Valle, Dwyer 1945, 1946 (both MO); Dwyer & Correa 7987 (COL, DUKE, K, MO, UC, VEN). 7 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 487 J 89. Sabicea panamensis Wernham.—A. Flowering branch (x'2).—B. Fruiting branch (x15). [After Oliver et al. 2674.]—C. Flower opened to show interior (x212).—D. Young fruit and seeds, cross section (x212). [After D'Arcy & D'Arcy 6244. ] N of El Cope, 900 m, Folsom 1337 (MO). La Mesa, Folsom 3650 (MO). La Mesa, El Valle de Anton, Folsom & Butcher 3818 (MO). Summit of Alto Calvario, 900 m, Folsom & Robinson 2429 (MO). La Mesa, N of El Valle, Gentry 6840 (MO). El Valle, Gentry & Dwyer 3631 (MO). N of El Valle, 2500- 3000 ft, Lewis et al. 1743 (GH, MO, US). 8 mi N of El Valle, Luteyn & Kennedy 1722 (MO). Cerro Pilón, 2000 ft, Porter et al. 4359 (MO, SCZ, UC), 4579 (MO, SCZ, UC), 4587 (MO). El Valle, ca. 1000 m, Wilbur et al. 11134 (MO). COLON: Río Miguel de la Borda near Guasimo, Croat 9917 (MO). Santa Rita Ridge, Croat 13836 (MO). María Chiquita, Dwyer & Kirkbride 7801 (MO, UC). Santa Rita 488 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Lumber Road, 5.4 km E of Transisthmian Highway, Folsom 3674 (MO). Rio Piedras and Puerto Pilon, Lewis et al. 3230 (MO, UC). Santa l. roe E of Transisthmian Highway, Lewis et al. 5271 (MO, SCZ, UC), 5284 (MO, SCZ, UC, N). 7 km NE of Puerto Pilón, road to Portobelo, Nee 6677 (MO). Achiote, Tyson et al. 4552 Mu. 8 km NE of Santa Rita, Wilbur & Weaver 10863 (MO). Santa Rita Ridge, Wilbur p Luteyn 11848 (MO). DARIEN: Rio Tuquesa, gold mining camp of Tyler pena Croat 27138 (MO). PANAMA: Cerro Jefe, ca. 2700 ft, Blum & Duke 2191 (MO). Cerro Azul, 2000 ft, Carrera 5 (MO). Goofy Lake to Cerro Jefe, Correa & Dressler 269 (MO). 3.5 mi W of Goofy Lake, Cerro Azul, Croat 11564 (MO). Cerro Campana, Croat 14760 (MO). Goofy Lake to Cerro Jefe, Croat 15233 (MO). 6 mi Goofy Lake, Croat & Porter 15233 (MO). Cerro Azul, Croat 17305 (MO). El Llano-Carti Road, i km from Interamerican Highway, 1100-1200 ft, Croat 25142, 33747 (both MO). Cerro Jefe, D'A & D'Arcy 6244 (MO); Duke 8019 (MO); Duke 8384 (MO). Cerro Campana, 2700-3000 ft, nom 8656 (MO). Cerro Jefe. Duke 9384, 9435, 9437 (all MO). Goofy Lake toward Cerro Jefe, Dwyer 7074 (MO). Cerro Jefe, Dwyer & Hayden 8091 (GH, MO. US). Cerro Azul and Cerro Jefe, Elias & Hayden 1791 (GH, MO). Cerro Jefe, Elias & Hayden 1795 (MO, UC). El Llano to Carti Road, 8 km N of Panamerican Highway, 350—500 m Cy 1429 (MO). Cerro Campana, a 1830 (MO); Hayden 121 (MO); Kirkbride & Crebbs 32 M O): Mod ds Dwyer 7844 (GH, MO, US); Lewis et al. 1915 (GH. MO, US). Altos del Río Pacora, 2500 ft, ] et al. 2339 (COL, NO UC, VEN). Cerro Campana, Lewis et al. 3035 (COL, MO, UC, VEN). pie Jefe, Liesner 588 (MO). Cerro Jefe and La Eneida, Luteyn & Foster 1131 (MO). Cerro Campana, McDaniel 6860 (MO). El Llano-Cartí Road, 18 km from Highway, Mori et al. iid E Mori & Kallunki 5603 (MO). 1 mi N of Goofy Lake, Oliver et al. 2674 Desin Cerro Cam . Porter et al. 4178 (COL, DUKE, MO), 4895 (MICH, MO), 4967 (MO, SCZ). Cerro Jefe, B et v^ 5061 (COL, DUKE, MO, SCZ, UC, VEN). Cerro Azul, Tvson 2150 (MO). Ceno] efe, Tyson et al. 3333 n SCZ). E slope of Cerro Jefe, Tyson 3437A, 3535 (both MO). 4 mi E of Cerro Jefe, Tyson 3576 (MO Cerro Jefe, Tyson et al. 3576 (SCZ), 4404 (MO). Cerro Jefe beyond Cerro iie 2700—3000 ft, us & Teeri 13608 (MO). SAN BLAS: Airport Mandinga, Duke 8915 (MO). vERAGUAS: Río Primero Braso, 2.5 km beyond Agriculture School Alto Piedra near Santa Fé, 700-750 m, ies 25469 (MO). Between Escuela Agricola and Alto Piedra above Santa Fé, 730—770 m, Croat 25927 (MO). I2 km NW of Santa Fé, D'Arcy 10334 (MO). 2 km N of Escuela Agricola Alto de Piedra, Mori & Kallunki 2567 (MO). iid of Santa Fé, 8.8 km from Escuela Agricola Alto de a Mon: gf al. 4017 (MO). 3-4 km by oad W of Santa Fé, ca. 2500 ft, Nee 11298 (MO). COLOMBIA: CHOCO: Top of Serrania del Darién, Pose with Panama, Cerro Mali, 1400 m, Gentry et al. 16972 (MC ». 2. Sabicea villosa R. & S., Syst. Veg. 5: 265. 1819. TYPE: Venezuela, Bonpland, not seen S. hirsuta H.B.K., Nov. Gen. Sp. Pl. 3: 417. 1820. TYPE: not seen. 2a. Sabicea villosa R. & S. var. villosa. Woody vines, the branchlets terete, villose, the hairs spreading. Leaves ellip- tic to elliptic rhomboid, rarely oblong rotund or obovate, 4-14 cm long, 1.8—6.0 cm wide, deltoid, widely deltoid to subobtuse at the apex, cuneate to obtuse at the base, the costa prominulous above and beneath, the lateral veins 8-12, ar- cuate, stiffly chartaceous, concolorous or discolorous, the hairs elongate above, often restricted to the costa and the veins, densely hairy beneath, the hairs spreading; petioles slender, to 1.5 cm long; stipules ovate oblong, to 1.3 cm long, the hairs spreading outside, glabrous within. Inflorescences axillary, usually found in all the axils of the ultimate twiglets, epedunculate, the flowers congested, the heads to 1.2 cm in diam. Flowers sessile; hypanthium densely pilose or hir- sute, the calycine cup ca. 1 mm long, the lobes 5, narrowly ovate oblong, to 4 mm long, glabrous or with a few elongate hairs within, pilose or hirsute outside; corolla white, the tube narrowly infundibuliform, to 5 mm long, ca. 0.6 mm wide, densely or sparsely pilose outside, the lobes 5, triangular, 1.0-1.8 cm long, acute. Fruits sessile, rotund or oblong, to 0.8 cm long, pink, purple, lavender, densely to sparsely golden pilose, the calycine lobes persistent, often reflexe 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 489 Sabicea villosa is widely distributed in Middle America and northern South America. BOCAS DEL TORO: Almirante, Blum 1368 (MO); Cooper 75 (MO). Punta Pena near Chiriquicito, 1000 ft, Lewis et al. 2161 (GH, MO, US). 4-6 mi N of Almirante, McDaniel 5107 (MO). Water Valley, Wedel 1497, 1590, 1782 (all MO). Chiriquí Lagoon, Wedel 2681 (MO). CocLE: Boca del Toabre, confluence of Río Toabre and Río Cocle del Norte, Lewis et al. 5497 (MO, SCZ, UC, VEN). COLÓN: Guasimo, Croat 9974 (MO) 2b. Sabicea villosa var. adpressa (Wernham) Standley, Publ. Field Columbian ot. Ser. 7: 52. 1930. Woody vines, the branchlets with appressed hairs. Leaves elliptic to elliptic rhomboid, 3-13 cm long, 2-6 cm wide, the hairs appressed above and beneath; stipules with the hairs appressed. /nflorescences, flowers and fruits as in S. villosa var. villosa. “Ibia ina" (Stier 232). Sabicea villosa var. adpressa 1s known only from Panama, Colombia, and ru OCAS DEL TORO: Between Almirante and Ojo de Agua, Croat 38204 (MO). Milla 5, railroad tracks, Croat & Porter 16494 (MO). Little Bocas, Chiriquí Lagoon, Wedel 2558 (MO). CANAL ZONE: Río Pedro Miguel, Madden Forest, D'Arcy & D'Arcy 6095 (MO). Ft. Kobbe, Duke 3939 (MO). Coco Solo, Duke 4285 (MO). Military Road K-9, Ebinger 521 (MO). Chagres, Fendler 180 (MO). Pipeline Road, 14.4 km from Chagres Airport Road, Mori & Kallunki 2055 (MO). Cerro Pelado, 1 km N of Gamboa, 200—220 m, Nee 7426 (MO). BARRO COLORADO ISLAND: Aviles 226 (MO); Croat 6703, 9431, 11725 (all MO); Ebinger 281 (MO); Mid aaa 14 (MO); Shattuck 704, 1130 (both MO); Woodw d & Vestal 524 (MO). DARIEN: Río Uruti, Bristan 231 a (MO). Río Pirre, Duke 4963 (MO). Peria Hill, Río Sabana and Lara, Tyson p al. 4732 (MO). PANAMA: 6 mi E of Chepo on Enn Highway, Duke 4076 (MO). El Llano and Río Mamoni, Duke 5599 (MO). San Jose Island, Johnston 1135 (MO). Chiman, ei et al. 3290 (MO). Cerro Campana, Luteyn & Kennedy 1798 (MO). Ar- raijan, Woodson et al. 3 (MO). SAN BLAS: Hills SE of Puerto Obaldia, Croat 16708 (MO). Isla de Ailigandi, Dwyer 6840 io Mainland, opposite Ailigandi, Lewis et al. 160 (MO, SCZ, VEN). Molia, Stier 42 (MO). Tappakanti, Stier 232 (MO). 3. Sabicea stellaris Dwyer.136 TYPE: Panama, Dressler 4887 (MO, holotype). Vine, branched, the branchlets terete, drying red, villose to hirsute, the nodes well spaced. Leaves elliptic, 5-13 cm long, 2.0-5.5 cm wide, apically deltoid, acuminate, the acumen vague or duct to | cm long, usually minutely apiculate, cuneate or acute at the base, the costa slender, prominulous, drying pink below, the lateral veins ca. 15, arcuate, somewhat whitish proximally beneath, papyra- ceous, concolorous, with scattered thin weak white appressed 1-2 mm hairs above and beneath, these more dense beneath especially along the costa; petioles slender, ca. 1 cm long; stipules ovate rotund, to 8 mm long, to 5.5 mm wide, acute, membranous, venose, pilose outside and at base within, with 6—10 subu- late, red glands at base within, these 0.1—0.2(—1.5) mm long. Inflorescences ax- illary, sessile, to 1 cm long, the flowers densely congested; bracts several per inflorescence, similar to stipules in texture and pubescence. Flowers sessile; hy- 136 Sabic s stellaris Dwyer, spec. nov. Vitex ramulis villosis ad Ed ns pis elliptica 5-13 cm longa, 2 e 5 cm lata, venis lateralibus ca. 15; petiolis ad 1 cm lon stipulis ovato-rotundis, ad 8 mm longi ad 5.5 cm p extus pilosis. d idi sillares ad 1 pics ngae. donis lobis calycis oblongis foliosis. 3-6 mm longis, inaequalibus s villosis; corolla tubo ca. 4 mm longo. Fructus sessiles rotundi, ca. 3.5 m mm in diam. costati villosi. 490 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 panthium short with densely appressed, elongate silky hairs outside; calycine cup short, appearing ringlike at the apex within (top view) because of the ovarian disc, the calycine lobes spreading in starlike fashion, oblong, foliose, 3-6 mm long, unequal, acute, villose outside, with a minute linear oblong gland between the lobes; corolla white, the tube ca. 4 mm long, carnose, drying red, villose in upper 25 outside, the lobes triangular, ca. | mm long, ovarian disc doughnut shaped, 0.3 mm long. Fruits sessile, rotund, ca. 3.5 mm in diam., ribbed, the ribs crassate, obtuse, villose above the middle, the raphides abundant. Sabicea stellaris is known only from Panama. The collector notes that the fruits are pink. On dissecting a mass of flowers a number of small insects were seen; it is unusual to find insects in inflorescences of the Rubiaceae, at least in Panama. This probably has some relation to the small glands found at the base of the stipules as well as the junction of the calycine teeth. The new species is characterized by its large stipules and calycine teeth. PANAMA: Río Pequeni, 10-15 minutes upstream from hydrographic station by motor, Dressler 4887 (MO). 77. SCHRADERA Schradera Vahl., Eclog. Amer. 1: 35. 1796. Nomen conserv. TYPE: S. capitata Vahl, nom. illeg. = S. a ees (Swartz) K. Schumann. Fuchsia Swartz, Prodr. Veg. Ind. Occ. 62. 1788, er Fuchsia L. 1753 Urceolaria Willd. in Cothen. Disp. Veg. Meth. 10, 1790. Nome en rejie. TYPE: U. exotica. Uncariopsis Karst., Linnaea 30: 152. 1859. TYPE: U. suaveolens Kar Epiphytic shrubs. Leaves usually leathery. Inflorescences terminal, pedun- culate, the flowers capitate, surrounded by a broad involucre. Flowers with the calyx turbinate, the calycine cup cupuliform, truncate; corolla salverform, the tube broad, villous within, the lobes 5—10, narrow, spreading or reflexed, valvate; stamens 5-10, the anthers subsessile, dorsifixed, included or exserted, the fila- ments adnate to the corolla tube; ovary 2-4-celled. Fruits a syncarp, fleshy, baccate; seeds numerous, small, compressed, suborbicular, horizontally at- tached Schradera occurs in Panama, IBS South America, and in the West Indies. It is a genus of about 16 species Literature: Steyermark, J. 1963. Schradera. Mem. New York Bot. Gard. 10: 259—278. 1. Schradera blumii Dwyer & Hayden, Phytologia 15: 59. 1967. TYPE: Panama, Tyson et al. 3218 (MO, holotype).—Fic. 90 Scandent s/irubs, the branches quadrangular, smooth, glabrous, rimose, the pith hollow in thicker twigs. Leaves narrowly oblong, often falcately oblong, 7- 15 cm long, 3-5.5 cm wide, acute at the apex, scarcely acuminate, basally acute to rounded, often slightly inequilateral, the costa prominulous above and beneath, the lateral veins 10-12, arcuate, evanescent above, prominulous beneath, the 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 491 "et NOS SEEN = ds ERAN C FIGURE 90. Schradera blumii Dwyer & Hayden.—A. Habit ( x1⁄).—B. Flower opened to show interior (x 112).—C. Ovary cross section (x2). [After Correa et al. 1865.] intervenal areas spreading, reticulate, glabrous, fleshy, drying black, coriaceous; petioles 1.5-2.5 cm long, rigid; stipules caducous, presumably connate for % of the length, the stipular scars conspicuous, ca. | mm long. Inflorescences terminal, black when dry, to 4.5 cm long, either pedunculate for 2 cm or peduncles absent and then the tip of the twig with 3 ternately disposed branches, to 2 cm long, ca. 0.3 cm wide, the flowers aggregated into globose heads or cymules; bracts black when dry, the outermost united into a cupule ca. 2.5 cm wide, the margin un- 492 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 dulate. Flowers with the hypanthium oblong rotund, to 10 mm long, glabrous, the calycine cup urceolate, to 8 mm long, truncate, the teeth absent, coriaceous, crassate, glabrous; corolla white, the tube slender, cylindrical, falcate in bud, to 30 mm long or longer, about the same length as the corolla lobes, stiffly carnose, glabrous outside, villose within below the middle, the lobes 4, narrowly subulate, triangular in cross section, subcoriaceous, glabrous; stamens 5, the anthers nar- rowly oblong, ca. 5.5 mm long, obtuse, the filaments ca. 1 mm long, plane, ca. 0.25 mm wide; style slender, the stigmas 2, free for ca. | mm, connate for ca. 2 mm. Fruits disposed as a rotund mass, to 3.5 cm in diam., the persistent calycine cup cylindrical or pipelike, to 1 cm long, to 0.4 cm wide, truncate, glabrous, as long as the rotund body, shiny white, black when dry. Schradera blumii is known from Panama and Colombia. Kirkbride & Duke 1300 notes that "Choco use (it) to stain tongue black." É: La Mesa above El Valle, Croat 13404, 14390, 25379 de a Sullivan 469 (MO). Cerro Pilon, pu 12190 (MO); Dwyer 8608 (MO). Alto Calvario, summit, 900 m, Folsom 2488 (MO). Cerro Pilon, Kirkbride 1091 (MO); Luteyn & Kennedy 1624 (MO); Mori iet (MO); Porter et al. 4613 A (MO); Wilbur et al. 11157 (MO). DARIÉN: Cerro Pirre, 1000—4000 m, Gentry & Clewell 6892 (MO). Cuasi-Caná Trail between Cerro Campamiento and La Escalera, E of Tres Bocas, Kirkbride & Duke 1300 (MO). PANAMÁ: El Llano-Carti Road, Correa et al. 1865 (MO); Mori & Kallunki 2898 (MO); Mori et al. 4594 (MO). Cerro Jefe, Ni NE of summit, ca. 1000 m, Mori & Kallunki 6090 (MO). Cerro Jefe in Clusia forest, 2700-3000 ft, Tyson et al. 3218 ere VERAGUAS: meee Rio Tres Brazos, 9 km from Santa Fé, Croat 2 (MO). Cerro Tute, ca. 10 km NW of Santa Fé, above 1000 m, Mori et al. 7600 (MO). COLOMBIA: CHOCO: 11 km E of e on Quibdo- Medellin Road, 100- 200 m, Gentry & Fallen 17545 (MO). 78. SIMIRA Simira Aubl., Hist. Pl. Guiane 1: 170. 1775. Not Simira Raf. 1836 TYPE: S. tinc- toria Aublet. Sickingia Wills., Ges. Naturf. Freunde Berlin Neue Schriften 3: 445. 1800. TYPE: S. erythroxylon tease Lu: Sitzb. Akad. Münch. 323. 1860. LECTOTYPE: A. rubra Mart. Trees or shrubs, the wood turning red to purple on cutting. Leaves large; stipules persistent or caducous, usually large and glandular basally. /nflores- cences terminal or axillary, paniculate, the flowers in cymes or capitate. Flowers pedicellate or sessile, bibracteolate; calycine cup cupuliform or bell shaped, trun- cate, edentate or with 4—6 teeth; corolla variable in color, fragrant, tubular to campanulate, the lobes 4—5, imbricate; stamens 4—6, exserted, the anthers dor- sifixed, sagittate at the base, the filaments connate basally, attached near the middle or at the base of the tube, villous; ovary 2-celled, the ovules numerous, horizontally biseriate. Fruits capsular, usually globose, loculicidally dehiscent; seeds numerous, large, winged horizontally. Simira ranges from Mexico through Central America and South America. It is a genus of about 35 species. Literature: Bremekamp, C. E. B. 1954. The identity of Simira tinctoria Aubl., Acta Bot. Neerl. 3: 150-153 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 493 Steyermark, J. 1972. Simira. The Botany of the Guyana Highland. Mem. New York Bot. Gard. 23: 299-309. a. Leaves ovate or oblong rotund; fruits 3-6 cm in diam. b. Trees to 5 m tall; corolla 3.5-5.5 mm long (fide Steyermark); fruits ca. 3 cm in diam. 2. S. klugei . Trees to 20 m tall; corolla to E mm long: fruits ca. 6 cm in diam. ---------- 3. S. maxonii aa. Leaves oblong; fruits ca. 1.3 cm in diam. l. S. darienensis 1. Simira darienensis Dwyer.!?? TYPE: Panama, Bristan 1187 (MO, holotype). Small trees or shrubs, the wood hard, the tips of the branchlets angular, glabrous, the bark smooth, cracking. Leaves oblong, 13-38 cm long, 6.5-20.0 cm wide, often widest above the middle, deltoid or obtuse at the apex, short acu- minate, the acumen to 3 cm long or longer, widely cuneate basally, ultimately truncate, ? vaguely auriculate, the costa slender except to 0.3 cm wide proximally, prominulous above, prominent only proximally beneath, the lateral veins 15-20, well spaced, prominulous beneath, arcuate, stiffly chartaceous, smooth and gla- brous above, sparsely appressed villosulose on costa and veins beneath, the in- tervenal areas open reticulate; petioles crassate, to 1 cm long, expanded at the base, 0.2—0.4 cm wide basally; stipules not seen. Inflorescences terminal, cymose paniculate, to 11 cm long, the peduncles 1-3, 5.0-8.5 cm long, angular, rigid, golden puberulent, the median peduncle about 2 times longer than the lateral peduncle, the lateral branches few, well spaced, usually opposite, to 2.5 cm long, angular ascending, the flowers few, terminal, the smaller bracts subulate, 3-5 mm long, the larger bracts confined to the median peduncle, ovate, to 8 mm long, suberect, coriaceous, concave, puberulent outside. Flowers with the hypanthium narrowly cylindrical, expanded at the apex, drying black, the calyx cup chalice shaped, ca. 4 mm long, puberulent, the lobes 5, irregular, obtuse, to 2.2 mm long, stiffly petaloid, puberulent; corolla with the tube cup shaped, ca. 4.5 mm long, glabrous except golden puberulent at the mouth, the lobes oblong, to 3.5 mm long, somewhat expanded, scarcely cucullate at the apex, the lower 25 giving the impression of a wide claw, to 1 mm wide, puberulent marginally; stamens 4(?), the anthers sagittate, ca. 3 mm long, densely golden villosulose above the middle, attached ca. 1 mm from the base of the tube; ovarian disc cushion shaped, 4-lobed, drying black; stigmas 2, narrowly oblong, to 2.3 mm long, reflexed; ovary 2-celled, the septum thin, the ovules numerous. Fruits (juvenile) sessile, oblong, to 1.8 cm long, ca. 1.3 cm wide, terete in cross section, truncate to rounded at the apex, minutely granulose, the persistent ovarian disc lignose, compressed, buttonlike, surrounded by a delicate ring of persistent calyx lobe bases. Simira darienensis is known only from Panama. One of the important diag- nostic features of the genus, i.e. the wood being pink after exposure to air, is not 137 Simira darienensis bug. er, spec. nov. Arbores ramulis ultime angulatis glabris. Folia oblonga, 13-38 cm longa, 6.5-20.0 nig inia venis lateralibus 15-20, rigido- -chartacea i in costa venisque subtus longis, floribus paucis terminalibus, bracteis maioribus ovatis, a 8 mm longis, coriaceis extus pu- berulis. Flores cupula calycis ca. 4 mm longa, lobis irregularibus obtusis, ad 2.2 mm longis; pye tubo ca. 4.5 mm longo, glabro praeter faucem, lobis ad 3.5 mm longis; antheris sagittatis, ca. 3 m longis. Fructus sessiles oblongi in ambitu teretes extus minuto-granulosi 494 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 evident in the two collections cited below. The pink color may fade in herbarium material. The fruits of the new species are oblong and much reduced in contrast to those of S. klugei (Standley) Steyermark and S. maxonii (Standley) Steyer- mark. Simira klugei has much smaller leaves than the new species. The leaf shape and the width of the petals serve to distinguish S. darienensis from S. maxonii. DARIEN: Rio Ucurganti, Bristan 1187 (MO). Rio Chucunaque between Rio Membrillo and Rio Subcuti, Duke 8606 (MO). 2. Simira klugei (Standley) Steyermark, Mem. New York Bot. Gard. 23: 302. 1972. Calderonia klugei Standley, J. Wash. Acad. Sci. 15: 6. 1924. rvPE: Panama, Kluge 19 (US). Sickingia klugei (Standley) Standley, Trop. Woods 22: 10. 1930. S. parvifolia Pittier, Bol. Mus. Com. Ind. Venez. 1: 71. 1926. TYPE: Venezuela, Saer 24, not seen. Trees to 5 m tall, the branchlets terete, ultimately angular, drying ashen grey, often rimose, short and angular ascending, to 9 cm long, glabrous, branching dichotomously. Leaves ovate, 4.0-7.5 cm long, 2.0—4.5 cm wide, deltoid or ob- tuse at the apex, at the base cordate, obtuse or rounded, usually auriculate, the costa prominulous or plane above, prominent beneath, the lateral veins 7-8, widely arcuate, the intervenal areas reticulate, papyraceous, shiny and glabrous above, moderately white villous beneath or with a few elongate hairs on the costa and the veins; petioles slender, 3-6 mm long, pubescent; stipules ovate triangular, 7 mm long, drying brown, puberulent outside. Inflorescences terminal, the flow- ers aggregated into a hemispherical head to 1.5 cm wide. Flowers (buds only seen) subsessile; hypanthium short, the teeth evanescent: corolla 4 mm long, rounded at the apex. Fruits sessile, subrotund, glabrate, 3 cm in diam.: seeds semiorbicular. In Venezuela a similar species, S. rubescens (Benth.) Bremekamp, occurs. In his key to Simira, Steyermark (1972) distinguished S. rubescens from S. klugei in its: ‘flowers usually 4-merous; corolla 3.5-5.5 mm long; calyx 1.0-1.5 mm long, the lobes 0.5-1.0 mm long; fruit 1.5 cm long by 1.8-2.0 cm broad; leaf blades as broad as long or to 1.7 times longer than broad, usually noticeably broadest around the middle, usually 2.5-8.0 cm long; lateral nerves 5-8 on each side." The common name of the species in Panama is ‘‘Palo Colorado." PANAMÁ: Chepo, Kluge 19 (F. photo US). Alhajuela, Pittier 3499 (F. US). Uu Simira maxonii (Standley) Steyermark, Mem. New York Bot. Gard. 23: 306. 1972.—Fic. 91. Genipa maxonii Standley, J. Wash. Acad. Sci. 8: 642. 1918. TYPE: Panama, Maxon 4848 (US, ho- otype). Sickingia maxonii (Standley) Standley, Trop. Woods 14: 30. 1928. Trees to 20 m tall forming a spreading crown, the twiglets ultimately 4-angled, usually turning red on exposure to air, glabrous, the nodes with subrotund scars, drying red. Leaves oblong rotund, 12.5-32.0(—60.0) cm long, 10—26(—40) cm wide, widely cuneate at the apex, acuminate, the acumen 2.0-3.5 cm long (probably longer in larger leaves), tapering obtusely toward the base and often slightly 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 495 R AA X p AN ^N \ E 91. Simira maxonii (Standley) Steyermark.—A. Habit (x!⁄).—B. Leaf venation -— ied Flower opened to show interior (x2). [After Howell 51.] auriculate, the costa immersed above, prominent beneath, often conspicuously lenticellate, the lateral veins 16—20, at first strict, finally arcuate near margin, the secondary veins irregularly pinnatiform, thinly coriaceous or stiffly papyraceous, deep red brown above, often reddish beneath, glabrous above, glabrescent to 496 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 densely white villose beneath; petioles absent or to 0.5 cm long; stipules not seen. Inflorescences terminal, solitary, paniculate, to 10 cm long; peduncle, if present, 1-3 cm long, the branches few, radiate from the tip of the twiglet or disposed in 2 pairs along the rachis, the branches opposite, to 2.5 cm long, drying red, the hairs stiff and short, a large bract located at the base. Flowers with pedicels 3 mm long; hypanthium to 8 mm long, pubescent, the calycine lobes 5, longer than the cup, compressed oblong, wider than long, the margin ciliolate, the cup with a number of glands within, these oblong, 0.1—0.2 mm long, brown; corolla yellow green turning orange, funnelform, to 8 mm long, minutely puberulent outside, villosulose at point of filament attachment, coriaceous to thickly carnose, the lobes oblong rotund or ovate oblong, to 2.5 mm long, obtuse; stamens 4, the anthers slightly or conspicuously exserted, oblong, 2.3-3.0 mm long, obtuse, the filaments 3-5 mm long, pubescent, carnose, each dilated at the point of attach- ment; ovarian disc doughnut shaped, ca. 1 mm long, the style to 10 mm long, somewhat thickened, pubescent, the stigmatic lobes oblong rotund or ovate ro- tund, to 2.5 mm long, obtuse. Fruits subsessile, rotund or oblong, ca. 6 cm in diam., the walls thick and lignose, not ribbed; seeds to 2.5 cm long, broadly winged. Simira maxonii is known from Costa Rica and Panama. Foster notes on his collection 2345 that the flowers are visited by hummingbirds. at appears to be a stipule was found on Witherspoon & Witherspoon 8651 ; it is narrowly deltoid oblong, to 2 cm long, ca. 0.5 cm wide at the base, acute and glabrous. ‘‘Guaiatil,’’ ‘‘Guaiatil Colorado," ‘‘Jagua de Montana,” and ‘‘Ja- gua Colorado." BOCAS DEL TORO: Without specific locality, Wedel 443 (MO). ae Cooper & Slater 3 (US). ONE: ed FRA Rio Gatun, Maxon 4848 (F, photo US). cuiRiQUÍ: Progreso, Cooper & Pom 284 woa Río Guanche, Foster et al. 2345 (F. US). Tres Brazos sawmill, Icacal, between Salud a Fin de Indio, Hos 'ell 51 (MO). Río Guanche, ca. 3-4 km above bridge, road to Portobelo, POERA & Withe wasu 8651 (MO). SAN BLAS: Puerto Obaldía, Gentry 1507 (MO). 79. SOMMERA Sommera Schlecht., Linnaea 9: 602. 1834. TYPE: S. arborescens Schlecht. Shrubs or trees, the branchlets pubescent, entire, mostly elliptic, membranous to chartaceous, the intervenal areas conspicuously lineolate; petioles present; stipules interpetiolar, deciduous. Inflorescences axillary, pedunculate, the flow- ers disposed in cymes, corymbs or racemes; bracts and bracteoles present. Flow- ers with the hypanthium turbinate, the calycine cup 4—5-lobed, the lobes broad, often foliose; corolla funnelform or campanulate, hairy outside and within; an- thers dorsifixed, the filaments short, pubescent; ovarian disc cup shaped, the style slender, short, the stigmas linear, often hairy, the ovary 2-celled, the ovules numerous. Fruits baccate, usually globose; calyx cup and lobes persistent; seeds many, obtusely angular. Sommera, a genus of about a dozen species, occurs throughout Central America and tropical South America. It is readily recognized by the lineolate character of the intervenal areas, coupled with the broad calycine lobes. The 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 497 E 92. Sommera donnell-smithii Standley.—A. Habit (x .45).—B. Leaf venation — A [Ar poen 15760.]—C. Flower, part of perianth removed to show interior (x2*/;).—D. Stigm (x2*/;3).—E. Calyx (x2*/;5).—F. Ovary cross section (x SS) [After Allen 1733.]—G. Fruit (x 49. [After Ebinger 969. | 498 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 genus commemorates C. N. Sommer, an entomologist and a friend of D. F. L. Schlechtendal who authored the genus. 1. Sommera donnell-smithii Standley, Contr. U.S. Natl. Herb. 17: 436. 1914. TYPE: Costa Rica, Donnell Smith 6592 (US, holotype).—Fic. 92. S. mesochroa Standley, Contr. U.S. Natl. Herb. 17: 439. 1914. Type: Panama, Maxon 4941 (US, holotype). Trees to 40 ft tall, the branches terete, glabrous to puberulent drying black brown, often ultimately angular. Leaves oblong to obovate oblong, to 25.5 cm long, to 14 cm wide, cuneate at the apex, cuspidate, the cusp to 2 cm long, 0.3- 0.4 cm wide in the middle, acute or cuneate, often slightly inequilateral at the base, the costa prominulous above, prominent beneath, to 0.25 cm wide proxi- mally, the lateral veins 10-17, strict then arcuate, prominulous beneath, the in- tervenal areas lineolate, papyraceous, appressed strigose villose above, villose beneath, often densely so on the costa and secondary veins; petioles to 3.5 cm long, to 0.25 cm wide, pubescent; stipules connate, lanceolate, 1.5-3.5 cm long, acute, thin caducous. Inflorescences axillary, to 6 cm long, the peduncles to 2 cm long, strongly ascending, golden villose, the rachis often with 1—2 pairs of opposite branches, the flowers numerous in cymules, corymbs or racemes. Flow- ers on pedicels to 8 mm long; hypanthium broadly rotund, truncate, the 5 calycine teeth united into a short tube, unequal, oblong, to 7 mm long, obtuse, petaloid, venose, marginally pubescent; corolla white, the tube compressed cylindrical, to 4 mm long, ca. 2 mm wide, puberulent outside, the lobes oblong, as long as the tube, villose adaxially; stamens 5, the anthers small, oblong, to 0.8 mm long, obtuse, dorsifixed, the filaments short, attached above the middle of the tube; ovarian disc cushionlike, the style to 2.5 mm long, glabrous, the stigmas 2, sub- ulate, erect, crassate, pubescent, ca. 1.5 mm long, the ovary thickly walled, bilocular, the septum thin, the ovules numerous on 2 intrusive placentas. Fruits baccate, rotund or oblong, ca. 1 cm in diam., the calycine tube and lobes persis- tent, cylindrical, the pericarp densely villose, turning red or red black at maturity. Sommera donnell-smithii is known only from Costa Rica and Panama. BOCAS DEL TORO: Río Teribe, e & Duke 456, 550 (both MO). cHiRIQUÍ: Boqueté, Finca Co lins. Blum s Dwyer 2543A (MO). Monte Rey above Boquete, Croat 15760 (MO). Boquete, Da- vidson 815 (MO, US). Cerro Horqueta, a A13715 (MO). Between Guardia Post and Volcan City ( Piedra, Río Monte Road at stream, Folsom 3976 (MO). Boquete, Maxon 4941 (US); Pittier 3137 (US). Cerro Horqueta, Quebrada Horqueta, 1600 m, Webster 16701 (MO). COCLE: El Valle, Allen 1575 (MO), 1733 (MO, US); Duke & Dwyer 13208 (MO). Cerro Pilon, Duke & Dwyer 13893 (MO); Dwyer & Correa 7921 (MO). El Valle, Ebinger 968 (MO). Road to La Mesa, El Valle, 2200 ft, Tyson 6898 (MO). DARIEN: Rio Tacarcuna, old Tacarcuna Village, 850 m, Gentry & Mori 13581 (MO). PANAMÁ: Cerro Campana, Duke 8681 (MO). 80. SPERMACOCE Spermacoce L., Sp. Pl. 102. 1753; Gen. Pl. ed. 5: 44. 1754. LECTOTYPE: $. tenuior L. Herbs, annual or perennial, the stems angled. Leaves linear or narrowly lan- ceolate; petioles short; stipules united to the petioles, the sheath with few to 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 499 numerous slender appendages. Inflorescences axillary, sessile, the flowers glo- merulate; bracts setaceous and filiform, subtending the inflorescences. Flowers small, the calycine cup scarcely measurable, the calycine lobes 4; corolla white or lavender, hypocrateriform to infundibuliform, the lobes 4, valvate; stamens 4, affixed variously in the corolla tube; stigmas connate or distinct. Fruits dry, the mericarps 2, unequal, each with a single seed, the smaller mericarp opening, the larger remaining closed; calyx persistent; seeds oblong. Spermacoce, a genus of about 100 species, occurs in the tropics of the New World and the Old World. Few collections have been made in Panama. Literature: Rendle, A. B. 1934. Linnaean species of Spermacoce. J. Bot. 72: 329. . 1936. Spermacoce remota Lam. J. Bot. 74: 10. Steyermark, J. 1972. Borreria. The botany of the Guayana Highland. Mem. New York Bot. Gard. 23: 813-814. a. Leaves scabrous above; calyx lobes ciliate; fruits pubescent l. S. confusa aa. Leaves glabrous above; calyx lobes glabrous on the margins; fruits glabrous ____ 2. S. tenuior l. Spermacoce confusa Rendle, J. Bot. 74. 12. 1936. LEcTOTYPE: Mexico, Galeotii 6211 (K, not seen). New name for S. tenuior sensu auct., non L S. tenuior sensu authors, not L. Subshrubs or herbs, to !^ m tall, branching, the stems angular, scarcely thick- ened on the angles, scabridulous, the nodes well spaced, the uppermost leaves often narrower or shorter. Leaves sessile or subsessile, elliptic to narrowly ob- long, 0.7-3.5 cm long, 0.1-1.0 cm wide, acute at the apex, basally acute or attenuate acute, the lateral veins 4, strongly ascending, stiffly papyraceous, con- colorous, drying dull green, scabridulous above, scabrid beneath on the costa and the veins; stipules with the sheath short, scabridulous, the setas few, 5-8, longer than the sheath. /nflorescences axillary, those in lower axils to 0.7 cm wide, to 0.3 cm wide in the uppermost axils. Flowers not seen. Fruits oblong, ca. 2 mm long, scabridulous outside, the calycine teeth often persistent, acute. Spermacoce confusa is a common weed in the American tropics, and it ex- tends northward into Florida. Steyermark (1974) has a good description of the flower: **Flowers 7-13, each one of these 0.3-1.0 cm long, calyx tube narrowly turbinate, 1.5 mm long; lobes triangularly lanceolate to linear lanceolate, acuminate, 0.4-1 mm long, conspic- uously setulose ciliolate on the margins, or glabrous . . . ; corolla white, red or lilac, hypocrateriform, 1.9-2.5 mm long, the tube 1.5 mm long, 0.7 mm wide at the base, 1 mm wide above, the lobes ovate oblong, obtuse or subobtuse, 0.5- 0.7 mm long, 0.4-0.5 mm wide, shorter than the tube, sparsely puberulent ex- ternally in the apical region, sparsely papillate within; anthers subglobose or oblong, 0.3 mm long; filaments 0.2 mm long." CANAL ZONE: Thatcher Ferry Road, Duke 5765 (MO). Chagres, Fendler 147 (MO). 500 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 In describing Spermacoce confusa Rendle (1936) referred to two elements as typifying his concept, S. tenuior sensu Gaertner (see Rendle, 1934), and the lectotype designated above. Because Rendle apparently had this specimen in hand when he provided the new name, it is chosen as lectotype. 2. Spermacoce tenuior L., Sp. Pl. 102. 1753. Type: Not seen, see Rendle, 1934.— IG S. riparia Cham. & Schlecht., nae 355. 1828. TYPE: Brazil, Sellow, not seen. . glabra sensu authors, not Mic Subshrubs to 1⁄2 m tall, the stems smooth, usually glabrous, the nodes well spaced. Leaves elliptic, narrowly oblong or obovate oblong, 3.0-5.5 cm long, 1.0-1.5(-1.9) cm wide, acute or obtuse at the apex, basally attenuate acute or acute, the costa prominulous beneath, the lateral veins (3—)4, strongly ascending, evanescent, rigidly papyraceous, glabrous, smooth, black when dry; petioles slen- der, 2-10 mm long; stipules with the sheath reduced, glabrous, the setas 4-5, exceeding the sheath in length. Inflorescences (in fruit) axillary, to 1 cm long. Flowers not seen. Fruits subsessile, oblong to turbinate, to 3 mm long, truncate, glabrous, smooth, black, the teeth tending to persist, to 0.3 mm long, acute. Spermacoce tenuior is a common weed throughout tropical America. It is probably common in Panama but has not been well collected. Steyermark (1974) has a good description of the flower: **Calyx and hypanthium 3.2-3.5 mm long at anthesis; hypanthium narrowly turbinate, 2.2 mm long, 1 mm wide, glabrous. Lobes of the calyx 4, unequal, lanceolate to linear lanceolate, acuminate or acute, one slightly larger, 0.5-0.9(-1.2) mm long, ciliolate on the lower margins where they join the lobes of the calyx but glabrous above and on most of the margins; corolla white, rotate, 1.5-2 mm long; lobes narrowly ovate to lance oblong, obtuse at the curved, cucullate apex, 0.8-1.2 mm long, 0.4 mm wide, glabrous outside, pilulose within toward the middle, longer than the tube; stamens included, reach- ing the mouth; the anthers ovate oblong, rounded at the base, 0.5 mm long, the filaments attached near the middle of the corolla tube, 0.2-0.3 mm long; style 0.5 mm long.” CANAL ZONE: ages Colorado Island, Woodworth & Vestal e Mee PANAMÁ: 5-6 mi E of Chepo, Panameri Highway, Duke 4030 (MO). Chagres, Fendler 147 (MO). Between Pacora and Chepo, Woodson. be al 1660 (MO). SAN BLAS: E of Puerto Obaldia, ie 16930 (MO). 81. STACHYARRHENA Stachyarrhena Hook. in Hook., Ic. Pl. 1068. 1870. TYPE: S. spicata Hook. f. Dioecious trees, unarmed, the branchlets glabrous. Leaves lanceolate, the lateral veins scarcely evident, strongly coriaceous, shiny, smooth, glabrous: pet- ioles rigid, stipules connate, collar shaped, coriaceous, truncate, without teeth or lobes. Flowers terminal, the female flowers solitary, the male flowers sessile, pedicellate, disposed in a stiff narrow spike of well spaced, several flowered clusters, calycine cup thick, glabrous, the lobes obsolete, the margin erosulose; corolla short, carnose, glabrous, the lobes carnose, shorter than the tube. Fruits capsular, the pedicels nodding, elongate; capsule large, oblong, round in cross 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) e ZNN NY ! I i És LE Sec FIGURE 93. Spermacoce tenuior L. Habit (x .65) [After Croat 16930. | 501 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 94. Stachyarrhena dichroa Standley, staminate plant. Habit ( x V). [After Lao et al. 1.] section, resembling a smooth squash, the wall fleshy, lignose on drying; seeds numerous in a jellylike pulp. Stachyarrhena is known from Panama, Venezuela, Colombia and Brazil. The genus has about 8 species. The plate appearing with the original description of the genus beautifully illustrates the fruit. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 503 =, xD, 77) 7 j FIGURE 95. Stachyarrhena dichroa Standley, pistillate plant.—A. Habit with fruit (x1⁄).—B. Flower bud with leaves (x!1⁄4).—C. Fruit opened to show seed (x!2). [After Gentry 1894.] 504 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 1. Stachyarrhena dichroa Standley, Contr. U.S. Natl. Herb. 18: 142. 1916. TYPE: Panama, Pittier 4078 (US, holotype).—Fics. 94, 95 Trees 7-15 m tall, the branchlets glabrous, the wood hard, the twiglets smooth, glabrous, the nodes well spaced, the pairs of scars spirally disposed. Leaves lanceolate, 9-25 cm long, 5.0-9.5 cm wide, deltoid or obtuse toward the apex, acuminate, the acumen to 1.5 cm long, falcate toward the apex, basally cuneate, often slightly inequilateral, the costa prominulous to subplane above, prominent beneath, often with a ridge, the lateral veins 8-15, prominulous, at first arising strictly, then arcuate toward the margin, with a strongly undulate submarginal vein ca. 0.2-1.2 cm in from the margin, the intervenal areas lightly reticulate, the margin slightly revolute, strongly coriaceous, shiny, smooth, slightly discolorous, often drying brown, glabrous; petioles woody, rigid, to 2.5 cm long, to 0.35 cm wide, slightly turgid; stipules persistent, conspicuously connate, collarlike, rigid, strongly appressed to the stem, to 0.4 cm long, conspicuously truncate. Male inflorescences terminal, glabrous, spikelike, to 23 cm long, to | cm wide; pedun- cles 1.5-3.0 cm long, the rachis contorted, the flowers in sessile, well spaced clusters, 2-4 per cluster, occasionally solitary. Female inflorescences terminal, solitary flowers on rigidly lignose pedicels, these slightly flexuous, black, gla- brous, to 2.5 cm long, angular and twisted, ca. 0.2 cm wide, slightly expanded at the apex, the basal bract urceolate, stipule-like, ca. 0.4 cm long, 0.2 cm wide, glabrous. Male flowers sessile; hypanthium scarcely measurable, the calycine cup bowl shaped, to 1.5 mm long, glabrous, coriaceous but somewhat thin walled, truncate, the teeth absent, eglandular within; corolla white, green, pink or lav- ender outside, often purplish pink within, to 8 mm long, the tube longer than the lobes, the lobes 4(—5) triangular oblong, obtuse or subacute, coriaceous, glabrous outside, densely pubescent within. Female flowers with the hypanthium oblong, to 1.5 cm long, ca. 0.8 cm wide, obtuse toward the base, hard, black, constricted toward the apex, the calycine cup urceolate, glabrous, the lobes absent or with the margin slightly erosulose or the lobes to 0.2 mm long; corolla with the tube yellow or white, short, carnose. Fruits on rigid, deflexed, woody, smooth pedi- cels, to 7.5 cm long, to 0.35 cm wide, black brown when dry, the capsule purple to brown in the living state, oblong, to 7 cm long, to 5 cm wide, round in cross section, generally resembling a small squash, smooth, glossy, black brown when dry, with a round, craterlike depression at the apex, to 1 cm in diam., ca. 2 mm wide, the wall lignose, to 1.5 mm thick; seeds many in a jellylike pulp. Stachyarrhena dichroa is known only from Panama. The inflorescence bearing male flowers is strikingly different from the female twiglets with their solitary flowers. Pittier, who collected the type of S. dichroa, describes the fruit as the color of a cherry.’ : Santa Rita Ridge, Croat 15351 (MO); Gentry 1894, 6567 (both MO); Lao et al. 1 (MO). El bane Cae Road, 8-11 km from the Interamerican Highway, Mori 7714 (MO). Santa Rita Ridge, 17-35 km from Boyd-Roosevelt Highway. Mori & Crosby 6304 (MO). Forest, Loma de la Gloria near Fato, Pittier 4078 (F, US). DARIEN: Demartaganala (Summit Camp), Duke 15833 (MO). PANAMA: Road from El Llano to Carti, about 14 km from highway at Chepo, Correa et al. 1834, 1836 (both MO). SAN BLAS: El Llano-Cartí Road, 24.5-25 km from the Interamerican Highway, Mori & Kallunki 5519 (MO) 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 505 82. TOBAGOA Tobagoa Urban, Feddes Repert. Spec. Nov. Regni Veg. 14: 341. 1916. TYPE: T. maleolens Urb. Herbs, the stems somewhat angular, glabrous. Leaves lanceolate, the few lateral veins strongly ascending near the margin, glabrous; petioles obvious; stipules connate, the sheath with 3 setas. Inflorescences axillary, capitate; bracts suborbicular with numerous segments. Flowers with calycine teeth 4, subequal; corolla rotate infundibuliform, densely pubescent outside, the lobes 4, valvate: stamens 4, the anthers oval; ovary 2-celled, with a single ovule per cell, attached to the upper part of the septum. Fruits vaguely didymous, the calyx persistent; seeds obovate, concave ventrally, delicately reticulate. Tobagoa, a monotypic genus, is known from Panama, Tobago, and the north coast of Venezuela. 1. Tobagoa maleolens Urban, Feddes Repert. Spec. Nov. Regni Veg. 14: 343. 1916. TYPE: Tobago, Eggers 5783 (B, not seen).—FIG. 96. Diodia ottonis Schum. ex Pittier, Pl. Usual. Venez. 338. 1926. TYPE: not seen. Herbs to | m tall, the stems strawlike, glabrous, angular, delicately winged, the nodes well spaced. Leaves lanceolate or ovate lanceolate, 6.5—10.0 cm long, 1-2 cm wide, attenuate at the apex, ultimately acute, cuneate at the base, the costa plane to immersed above, prominulous beneath, the lateral veins ca. 7, scarcely evident above, prominulous beneath, at first leaving the costa widely and arcuately then toward margin strongly ascending, the submarginal vein ca. | mm from the margin, inconspicuous, the margin delicately revolute, the inter- venal areas smooth, the veinlets not evident, chartaceous, sometimes discolo- rous, glabrous; petioles slender but rigid, to | cm long, glabrous, attached but not conspicuously so to the stipular sheath; stipules connate, to 3 mm long, each part of the sheath with 3 setas exceeding its length. Inflorescences compressed capitate, axillary, to 0.5 cm long. Flowers with the hypanthium and calycine cup short, the calycine lobes 4, ovate triangular, | mm long, acute, glabrous, with a minute gland in the sinus; corolla white, the tube and lobes 3 mm long, the tube puberulent outside, villose upwards within, the lobes 4, longer than tube; stamens 4, the anthers oblong, | mm long, obtuse, minutely apiculate, the filaments short; style 2 mm long (in bud), papillose, the stigmas 2, curved and reflexed, to 0.4 mm long. Fruits dry, indehiscent, oblong, ca. 2 mm long, vaguely didymous, glabrate, the calyx persistent; seeds ventrally concave, finely reticulate. Tobagoa maleolens is known from Panama, Tobago, the West Indies and Venezuela. Only one Panamanian collection was available for study. The fruit seems glabrate. Steyermark’s (1974) recently published plate on p. 1836 shows the ovules attached basally while his generic description (loc. cit. p. 1835) says the ovule is adherent to the upper part of the septum. COCLÉ: North rim of El Valle de Anton, 600-1000 m, Allen 1666 (GH, MO). 506 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 96. Tobagoa maleolens Urban.—A. Habit (x34).—B. Node (x2!4). [After Allen 1666. ] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 507 83. TOCOYENA Tocoyena Aubl., Hist. Pl. Guiane 1: 131. 1775. TYPE: T. longiflora Aubl. Trees or shrubs. Leaves opposite or ternate, often drying black; stipules interpetiolar, free or connate, persistent. Inflorescences terminal or subterminal, cymose corymbose or capitate, the cymes often 1 sided. Flowers with the hy- panthium obovoid or cylindrical turbinate, the calycine cup short, the teeth 5-6, reduced, persistent in the fruit; corolla large funnelform or hypocrateriform, usu- ally hairy at the mouth, the lobes 5, contorted in bud; stamens exserted, the anthers dorsifixed, acuminate at the apex, the filaments very short, attached at the corolla mouth; ovary 2-celled, the ovules numerous, the placenta axile. Fruits baccoid, oblong or ovoid, the pericarp thick, the seeds numerous horizontal and embedded in a pulp, compressed, the testa foveolate. Tocoyena is known from Panama and tropical South America. It also occurs in the West Indies (fide Willis, 1966). There are about 20 species in the genus. Literature: Dwyer, J. D. 1969. Borojoa and Tocoyena (Rubiaceae) in Panama. Phytologia 17(6): 445—449. 1. Tocoyena pittieri (Standley) Standley, Contr. Arnold Arbor. 5: 151. 1933.— Fic. 97 Posoqueria pittieri Standley, J. Wash. Acad. Sci. 18: 167. 1928. Type: Panama, Pittier 6635 (US, olotype). hol Tocoyena cuatrecasasii Steyermark, Act. Biol. Venez. 4(1): 113. 1964. TYPE: Colombia, Cuatrecasas 1918. Trees to 10 m tall or much taller (fide Croat), the branchlets terete, smooth but often nodose, glabrous, the nodes usually 2-3 cm apart. Leaves elliptic, to 32 cm long, to 17 cm wide, widely deltoid to somewhat rounded at the apex, shortly acuminate, the acumen to 1 cm long, ultimately obtuse, cuneate toward the base, often somewhat inequilateral, the lateral veins ca. 10, arcuate, the lesser veins slender, irregular and spreading, pinnatiform, plane, stiffly chartaceous, black brown when dry, glabrous except the minutely pubescent axils of the prin- cipal veins beneath, minutely papillate under magnification, lightly marcescent above; petioles to 3 cm long, ca. 0.25 cm wide, glabrous; stipules free, triangular, 5-10 mm long. /nflorescences cymose paniculate, resembling a candelabra, to 13 cm long, ca. 11.5 cm wide, the flowers numerous, erect; bracts and bracteoles triangularly subulate, 0.5-1 mm long. Flowers pedicellate, the pedicels 1-3 mm long; hypanthium oblong, to 4 mm long, truncate, glabrous, the calycine cup ca. I mm long, eglandular within, the teeth 5, widely triangularly subulate, to 0.5 mm long, glabrous; corolla yellow, the tube narrowly cylindrical, 8-12 cm long, to 0.6 cm wide, thickly carnose, glabrous outside, within white villous at the base of the tube, the lobes 5, forming an ovate rotund mass in bud, golden farinose outside, ovate elliptic, obovate or elliptic rotund, 1-1.5 cm long, glabrous; sta- mens 5, exserted, sessile, the anthers elliptic, ca. 6 mm long, ca. 2 mm wide, 508 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FIGURE 97. Tocoyena pittieri (Standley) Standley.—Inflorescence with leaves (x?/;). [After Kirkbride & Duke 1386.]—B. Flower ( x?/,).—C. Ovary cross section (x 2?/,). [After Croat 4636.]— D. Fruit (x2). Fruit opened to show seeds (x). [After Duke 15599. ] attached at the corolla mouth; style slender, ca. 0.6 mm wide, the stigmas ovate- lanceolate, crassate, ca. 5 mm long. Fruits sessile, solitary, subrotund, 6-10 cm in diam., woody when dry, the wall thick, to 1.3 cm wide, drying black outside, tan within, the pulp blue black when dry, smooth externally except longitudinally costate, the ribs 10—15, well spaced, slender or thick, prominent but not ridge- 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 509 like, often evanescing along their length, the surface often marked by lenticels; seeds flat, ovate trapeziform, to 2 cm long, to 1.3 cm wide, slimy to touch. Tocoyena pittieri is known only from Panama. Kirkbride and Duke (1368) note: ‘‘anthers white, reflexed with the petals; seeds embedded in a brownish- black pulp with somewhat the odor and taste of molasses.” CANAL ZONE: Summit Garden, Lindsay 322 (MO, US). Trinidad River near Hydrographic station, Pittier 6635 (US). BARRO COLORADO ISLNAD: Aviles 14 (MO); Croat 4636, 8529 (both MO); Hladik 138 (MO), 299 (MO, US); Shattuck 671 (MO); Starry 131 (MO); Wetmore & Abbe 197 (MO); Wood- worth & Vestal Zetek 3536 (MO). DARIÉN: Rio Ucurganti, Bristan 1185 (MO). La Boca de Pirre, Bristan 1246 (MO). Río Yape, Bristan 1399 (MO). Trail from Cerro Mali to Río Pucro, 650-1200 m, Gentry et al. 16970 (MO). Río Balsa near Río Cuasi, Kirkbride & Duke 1386 (MO). Río Balsa near Cerro Campamento, S. Cerro Pirre cloud forest, Duke 15999 (MO). PANAMÁ: La Cuesta, Panama City, Holdridge 5614 (MO). 84. UNCARIA Uncaria Schreb., Gen. Pl. 1: 125. 1789. Nomen cons. TYPE: U. guianensis (Aubl.) J. F. Gmelin. d Aubl., Hist. Pl. Guiane I: 177. 1775. TYPE: O. DN Aubl. gylophora Neck., Elem. Bot. 1: 145. 1790. Type: not indicated. E ap Reichbach, Nom. 230. 1840. TvPE: not given Vines or sprawling shrubs, armed with recurved spines (aborted peduncles). Leaves opposite, the stipules interpetiolar. Inflorescences axillary and terminal, numerous, in globose heads. Flowers 5-merous, the hypanthium spindle shaped or turbinate, the calyx lobes valvate; corolla funnelform, pubescent outside, gla- brous within, the lobes 5, valvate; stamens 5, the anthers oblong, sagittate at the base, the filaments short, inserted at the mouth; ovary 2-celled, the style slender, the stigmas small, capitate, the ovules numerous, ascending, the ovarian disc obsolete. Fruits capsular, septicidally dehiscent, each valve splitting at the apex; seeds numerous, imbricate, winged, the wing frequently split. Uncaria, a genus of about 60 species, occurs in the tropics of the Old and New Worlds. In the New World it occurs in Central America, Trinidad, and tropical South America. 1. Uncaria tomentosa (Willd.) DC., Prodr. 4: 349. 1830.—Fic. 98. Nauclea tomentosa ies in Roem. & Schult., Syst. Veg. 5: 221. 1819. N. aculeata H.B.K., . Gen. Sp. Pl. 3: 382. 1819, not Willd. 1797. Ourouparia tomentosa MI ) Schum. in Mart., Fl. Bras. 6(6): 132. 1889. Vines, often in forest canopy, the stems quadrangular, minutely puberulent, armed with subnodal spines, the spines uncinate, stiff, lignose, to 2 cm long, 0.4- 0.6 cm wide proximally. Leaves oblong or ovate oblong, 9-17 cm long, 4.3-9.0 cm wide, tapering acutely or occasionally widely deltoid or rounded at the apex, obtuse, truncate, often vaguely cordate at the base, the costa prominulous above, prominulous to prominent beneath, the lateral veins 8-10, arcuate, angular, char- taceous or papyraceous, usually discolorous, minutely pubescent on the veins above, beneath usually densely and finely golden puberulent with fewer larger hairs intermixed, or often sparsely puberulent, the hairs confined to the costa, 510 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 E m. e ‘if [W ur A AN jp o . Uncaria d (Willd. B. Inflorescence (x!4).—C. Flower (x214).— ) rA n prre stem (x 5). [After Croat 8288. |— a opened to show interior (x'2).—E. Style (xX3V3).—F. Ovary cross section (X293). i ay Gentle 1578, Belize.] 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 511 veins and venules, the intervenal areas patulous reticulate; petioles to 1.5 cm long, 0.1-0.35 cm wide, tomentose; the stipules triangular, ca. 1 cm long, ca. 1.2 cm wide at base, acute, stiff, drying rubescent, with appressed white hairs. /n- florescences axillary, to 9 cm long, the peduncles 1.0-3.5 cm long, 0.8-3.0 mm wide, simply terminating in a globose capitulum of flowers, or the axes with 2 pairs of branches, these opposite, perpendicular, well spaced, puberulent, each terminated by a rotund capitulum, ca. 2 cm in diam. Flowers yellowish; hypan- thium oblong, to 1.5 mm long, carnose, puberulent, the calycine cup urceolate, ca. 0.65 mm long, densely white villose within, pubescent outside, the 5 teeth disposed as points; corolla with the tube funneliform, ca. 5 mm long, ca. 0.6 mm wide, petaloid, puberulent inside and out, the lobes 5, reflexed, oblong rotund, 1.4-1.7 mm long, glabrous on the adaxial face; stamens 5, the anthers oblong, 1.1-1.5 mm long, sessile, attached near the mouth; ovary 2-loculate, the septum moderately thin, the style linear, far exserted, ca. 4.5-6.0 mm long, the stigmas clavate, 1.0-1.5 mm long. Fruits septicidally bivalvate, narrowly ovate oblong, to 6 mm long, the persistent calyx coroniform, ca. 1 mm long and wide, drying tan, vaguely longitudinally striate, covered with dense white hairs, the seeds linear fusiform, to 4 mm long, the body ca. 1 mm long, the wing narrower than the body, membranous, 1 end linear oblong, the other deeply cleft and biaristate. This species is known from Nicaragua, Panama, Colombia, Venezuela, Guiana, Ecuador, and Trinidad. Duke (14554) notes that the vine is called ‘‘Bejuco de Agua" . . . from which we drank water. ‘‘Tua juncara’’ (Choco). ‘‘Rangaya’’ (Changuinola Valley). NAMA: BOCAS DEL TORO: Changuinola River, Stork 150 (US). CANAL ZONE: Pipeline Road, 4 mi N of Gamboa, Gentry 6051 (MO). Fort Sherman, Hayden 108 (MO). Gatün River Valley, Pittier 6509 (US). Fort Sherman, ariga up (US). Pipeline Road, N of Gamboa, Wilbur & Weaver 11228 (MO). BARRO COLORADO ISL : Carleton 82 (US); Croat d 5504, 6013A, 8288 (all MO); Hladick s (MO); Kenoyer 543 (US): s Sandi 31274 (US); Wetmore & Abbe 133 (MO); Wilson 72 MO). DARIEN: Interamerican Highway, ca. 1 mi SE of Rio Tuira, Duke 14554 (MO). PANAMA: Rio Tapia, Standley 26161 (US). COLOMBIA: CHOCO: Hydro Camp 11, Río Salaqui, 6 days upstream from Rio Sucio, Duke 11385 (MO). 85. WARSZEWICZIA Warszewiczia Klotzsch, Monatsber. Kónigl. Preuss. Akad. Wiss. Berlin 496. 1853. TYPE: W. coccinea KI. Trees or shrubs. Leaves large, petiolate; stipules large, glandular within at the base. Inflorescences terminal, cymose, the cymes arranged in racemose panicles. Flowers 5-merous, the hypanthium oblong ovoid, the calycine cup with lobes subequal, or 1 disposed as a stalked foliaceous limb; corolla funnelform, villous within, the lobes imbricate, 2 of them exterior; stamens with the anthers dorsi- fixed, versatile, exserted; ovarian disc annular, the style included or exserted, the stigmas 2, short, the ovary 2-celled, the ovules numerous, the placenta adnate to the septum. Fruits capsular, septicidally bivalvate; seeds numerous in 2 cells, horizontal, 3-angulate, marginate, reticulate. Warszewiczia occurs in Mexico, Central America and tropical South America. It is a genus of 4 species. 512 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FiG 9. Warszewiczia coccinea (Vahl) Klotsch.—A. Habit (x!2).—B. Flower opened to show cie m [After Shattuck 979. | l. Warszewiczia coccinea (Vahl) Klotsch, Monatsber. Kónigl. Preuss. Akad. Wiss. Berlin 497. 1853.—Fic. 99 Macrocnemum coccineum Vahl, Symb. Bot. 2: 38. 1791. TvPE: not s Mussaenda coccineum (Vahl) Poir. in Lam., E Méth. Bot. 4: 394. 1797. Calycophyllum coccineum (Vahl) DC., Prodr. 4: 367. 1830. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 513 Warszewiczia alls Klotsch, Monatsber. ae Preuss. Akad. Wiss. Berlin 497. 1853. TYPE: Panama (‘‘Verugua’’), Warszewicz, not see Trees to 12 m tall, the branches stout, angular, drying brown or black brown, relatively smooth, rimose, minutely puberulent to appressed pilose. Leaves ob- long, 5—45(—60) cm long, 10-20(-23) cm wide, tapering acutely or deltoidly at the apex, acuminate, the acumen to 2 cm long, the costa plane or prominent above, prominent beneath, the lateral veins 15-24, widely arcuate or arising from the costa almost perpendicularly, stiffly papyraceous, glabrate above, puberulent on the veins beneath; petioles 2-5 cm long, 0.8-1.1 cm wide, acute, golden puber- ulent, striate. /nflorescences terminal, to 60 cm long, the cyme usually subrotund or flabellate in appearance, disposed as a racemiform panicle, the clusters 1.5- 5.5 cm apart; peduncles suppressed or to 2.5 cm long. Flowers with the hypan- thium compressed rotund, ca. | mm long, the hairs dense, tawny, appressed, the calycine cup somewhat shorter than the hypanthium, its hairs often deciduous, the lobes 5, rotund, ca. 0.5 mm long, at least 1 flower in the cyme with an enlarged lobe, the lobe red, foliose, long stipitate, elliptic calycine to 12 cm long, to 4 cm wide, the blade usually slightly longer than the stipe, venose, glabrescent; corolla orange red, yellow or orange, 6-8 mm long, the tube short, cylindrical, expanded toward the apex, a little longer than the lobes, thickly petaloid, glabrous or gla- brescent outside, pubescent within at the point of filament attachment, the lobes 5, oblong or oblong rotund, 3 mm long, rounded, truncate or attenuate toward apex; stamens 5, the anthers oblong, ca. 1.6 mm long, petaloid, acuminate at the apex, the filaments subulate, 3 mm long, villose at the base; style crassate, to 5.5 mm long, the stigmas 2, rotund, exserted from the corolla. Fruits capsular, septi- cidally dehiscent, pilosulose; seeds to 1 mm long. Warszewiczia coccinea ranges from Nicaragua south to Bolivia. It is also found in Trinidad. Stern et al. (89), in noting the common name ‘‘Sangrenaria,”’ add that the plant is used for hemorrhage. The flowers are protogynous and develop acropetally. Folsom 2599 notes visitation by hummingbirds. ‘‘Guna,”’ "Crucero," ‘‘Sangrenaria.”’ L ZONE: | mi N of Summit Garden, Blum et al. 2336 (MO). Pipeline Road near Gamboa, Clew sell E Ty son 3318 (MO). Summit Garden W of Highway, Croat 15010 (MO). Pipeline Road 3 mi from Ga m PN Croat 16605 (MO). Agua Sucia, Fairchild (F). Frijoles, Ebinger 316 (F). C-29 road, ne arvey 5281 (F). Gamboa, Stephens 1073 (F, US). BARRO COLORADO ISLAND: Aviles 14, I5 pe v Moh 979 (F); Croat 6032, 734 0, 9923, 10278, 15010, 15562 (all MO); Hayden 48 (MO); Shattuck 979 (MO, US); Starry 271 (F, MO). cHIRIQUI: San Félix to Cerro Flor, Allen 1954 (F, MO). Vicinity Remedios pes 3664 (F). ue Transisthmian Highway, ca. 50 mi W of Santiago, Dicver & Hayden 7761 (G , MO, UC, US). cocLé: El Valle, Dwyer 1912 (MO, US), 2/09 (MO). Boca del Toabre, Le wis 2 P oe (MO, SCZ, UC). El Valle, Luteyn & Kennedy 1686 (F); Seibert 427 (F). COLON: Vicinity Guasimo, Croat 9923 (MO). Rio Salud, Howell 127 (MO). 2-3 mi up Rio Guanche, Kennedy & peus 2131 (MO). Río Guanche, 3-4 km above bridge, road to Portobelo, Witherspoon & Witherspoon 8677 (MO). DARIEN: Rio Chico, near Yaviza, Allen 5091 (F). El Rio ape, Bristan 1390 (MO). Gold mine Cana, 500-600 m, Croat 37672 (MO). Río Ganglon, Duke & Bristan 374 (MO). Without other locality, Duke 5112 (MO, US). Río Tucuti, ca. 2 hours by Piragua above Tucuti, epo D (MO). Río Perrecenico off Río Pirré, Duke & Briston 8228 (MO). Pinas, Duke 10592 (MO). ene to mouth of Río Cuasi, Kirkbride & Bristan 1493 (MO). N slope of Cerro Pirre, Mori & Pu and 5426 (MO). Patino, Pittier 6674 (US). Bahia de Pinas, Stern & Chambers 171 (F, MO, US). Paya, Río Paya, Stern et de 371 (MO, US). Cerro Piriaque, Tyson et al. 3831 (MO). PANAMÁ: El Llano to Cartí Road, 14 km N of Panamerican Highway, 350—500 m p et al. 1482 (MO). 5 mi SW of Cerro Brewster, . et al. 3474 (MO, SCZ, UC, VEN). SAN B : El Llano to Cartí Road, 16.5 km N of Panamerican Highway, Folsom 2599 (MO). VERAGUAS: W of So na, Allen 1023 (MO). Sona, Allen 1026 (F). 55 mi W of Santiago, D'Arcy & D'Arcy 6666 (MO). 514 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 FiGURE 100. Wittmackanthus standleyanus (Schomb.) Kuntze. Habit ( x34). [After Duke 6329.] 86. WITTMACKANTHUS Wittmackanthus Kuntze, Rev. Gen. Pl. 1: 302. 1891. TYPE: W. stanleyanus (Schomb.) Kuntze.—Fic. 100. sag Klotzsch, Monatsber. Kónigl. Preuss. joa Wiss. Berlin 498. 1853; not Pallasia Hout. 11: 1775, Cucurbitaceae; not Pallasia Scop. Introd. 72. 1777, Gramineae; not Pallasia L.f., is 37. 1781, Polygonaceae: not Pallasia L’ Tn X Ait., Hort. Kew III: 498. 1789, Compos- itae. Trees or shrubs. Leaves opposite, petiolate; stipules free, small, inter- petiolar. Inflorescences terminal and axillary, spicate. Flowers 4—6-merous; hy- panthium short; calycine cup short, the lobes 4-6, variable in length; corolla tubular, variable in length, the lobes imbricate; stamens 5, unequal, the anthers versatile, attached near the mouth; ovary 2-celled, the ovules numerous, verti- cally imbricate, alate, the placentas carnose. Fruits capsular, septicidally dehis- cent, the calyx persistent, the seeds winged. Wittmackanthus is known from Panama and northern South America south to Peru. The genus is monotypic. 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 515 Literature: Steyermark, J. A. & J. Kirkbride, Jr. 1975. The genus Wittmackanthus (Rubi- aceae). Ann. Missouri Bot. Gard. 62: 504—509. 1. Wittmackanthus stanleyanus (Schomb.) Kuntze, Rev. Gen. 1: 302. 1891. Calyocophyllum stanleyanum Schomb., J. Dus 3: 622. 1844. TYPE: Guyana, Schomburgk 356/411 B , not seen; photo, MO; NY, isot Pallasia stanleyana (Schomb.) Klotzsch, Monaten Konig]. Preuss. Akad. Wiss. Berlin 498. 1853. Rondeletia dukei Dwyer & Hayden, Ann. Missouri Bot. Gard. 54: 144. 1967. TvPE: Panama, Duke 9 (MO, holotype; NY, isotype). Shrubs or trees, the branchlets purple or red, glabrous but often ciliolate above the nodes. Leaves elliptic oblong or widely oblong, 4—13 cm long, 2.0-5.5 cm wide, deltoid, acute, obtuse or rounded at the apex, if acuminate the acumen to 1 cm, cuneate at the base, the costa slender, prominulous beneath, the lateral veins ca. 9, widely arcuate, patulous, forming an undulate vein near the margin or the submarginal vein not conspicuous, the intervenal areas reticulate, stiffly membranous to chartaceous, glabrous or hirsute on the costa beneath, drying grey, the margin delicately callose; petioles 1.0—4.5 cm long, wiry, stiff, glabrous, drying black; stipules triangular, to 0.6 cm long, to 0.4 cm wide, acute, glabrate or golden pubescent. /nflorescences terminal or axillary, spikelike, cymose; pe- duncle to 6.5 cm long, ca. 0.1 cm wide, glabrous, minutely farinose, the flowers either solitary or disposed in few flowered, well-spaced clusters. Flowers sessile or pedicellate, the pedicels to 0.3 cm long; hypanthium subrotund, ca. 1.5 mm long, densely appressed golden pilose; calycine lobes 5, oblong rotund, ca. 2 mm long, about as long as wide, the margin thin, ciliolate, the lowermost flowers with l or 2 calycine lobes expanded as a wide foliose ovate blade, to 4 cm long, obtuse, drying yellow; corolla white, the tube slender, to 5.5 mm long, ca. 0.8 mm wide in the middle, carnose, densely puberulent outside, sparsely ciliate at the mouth within, the lobes 5, oblong, 2-3 mm long, pubescent inside and out, obtuse, spreading at anthesis; stamens 5, the anthers oblong, 1.0-1.5 mm long, sagittate basally, versatile, the filaments unequal, as long as the anthers, attached at the mouth; ovarian disc shallow, doughnut shaped, the style slender, ca. 9 mm long, expanded just below the stigma, the latter saucer shaped, ca. 0.5 mm wide. Fruits sessile, narrowly oblong, to 1.2 cm long, to 0.4 cm wide, covered with a gray indumentum, somewhat plano compressed; seeds membranous, drying yellow, fusiform, to 5.5 mm long. Wittmackanthus stanleyanus is known from Panama, Guyana, Colombia, and Peru. Steyermark and Kirkbride (1975) reduced Rondeletia dukei Dwyer & Hay- den to synonomy under Wittmackanthus stanleyanus. They note: `The type collection of R. dukei from Panama has blades obtuse to rounded at the apex, but may have been collected from an abnormal plant. Finally, the size of the leaf blades is quite variable and has no taxonomic significance.” DARIEN: Rio Pirre, Duke 8329 (MO, NY). Rio Tuquesa, at the middle Tuquesa Mining Company Camp called Charco Peje, tropical wet forest, ca. 250 m, Mori 7001 (MO). 516 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 87. XEROCOCCUS Xerococcus Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjgbenhavn 1852: 52. 1852. TYPE: Costa Rica, X. congestus Oerst. Subshrubs. Leaves opposite, acuminate, petiolate. Inflorescences axillary, epedunculate, the flowers densely conglomerate. Flowers with the calycine lobes slender, elongate, persistent; corolla lobes 4, erect; stamens 4, the anthers ovate, the filaments short, inserted near the mouth; ovary 2-celled, the ovules numerous, horizontal, anatropous, the style slender, the stigmas 2, thick. Fruits baccate, ovoid, 2-celled; seeds numerous, angular. Xerococcus is a genus of 2 species confined to Costa Rica and Panama. — . Xerococcus congestus Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjg- benhavn 1852: 52. 1852. TYPE: Costa Rica, Oersted 11776 (F, photo MO). Subshrubs, unbranched, to 12 m tall, the stems terete, smooth below, blotchy terminally, the nodes well spaced. Leaves obovate or occasionally ovate rotund, 15-32 cm long, 10-13 cm wide, rounded at the apex, acuminate, the acumen triangular, to 0.5 cm long, ultimately obtuse, attenuate cuneate at the base, the lamina decurrent on the petiole, the wing 5-7 mm wide, the costa slender above, prominulous, plano compressed beneath, to 0.25 cm wide, the lateral veins ca. 12, arcuate, prominulous, with 1-3 small intermediate veins between adjacent lateral veins, these soon branching, the smallest veins pinnatifid, widely spaced, chartaceous, glabrous above, beneath with curled hairs, these often collapsed and giving a glandular webby appearance to costa and secondary veins; petioles narrowly winged, 5-7 cm long; stipules apparently persisting only at the apex of the stem, free, triangular, to 18 mm long, to 10 mm wide at base, coriaceous, glabrous. Inflorescences axillary, epedunculate, capitate rotund, to 1 cm long, to 2 cm wide. Flowers with pedicels short, to 2.5 mm long, golden villose; hypan- thium ca. 2 mm long, the calycine cup scarcely measurable, the lobes 4, elongate, ascending digitiform, to 5.5 mm long, about equal, to 0.4 mm wide, stiff, villose outside, glabrous within, often with 1—2 glands between adjacent lobes, the tips often curved, obtuse, often with | or 2 pairs of glands on the basal margin, drying red; corolla tube cylindrical, ca. 3.5 mm long, ca. 1 mm wide, glabrous inside and out, the lobes 4, triangular oblong, to 3.5 mm long, to 1 mm wide at the base, thick, puberulent on the back; stamens 4, the anthers subsessile, ca. 1.2 mm long, attached at the mouth; style 3.5 mm long; stigmas 2, apparently upright, some- what triangular, wider than the style; ovary 2-celled, the ovules numerous. Fruits oblong rotund, to 6 mm long, smooth, white, also on drying, the calyx persistent; seeds compressed, angular, square, hemispherical or rhomboidal, ca. 0.3 mm long, reticulate, drying red. Xerococcus congestus is known only from Costa Rica and Panama. Four collections in the Missouri Botanical Garden from Costa Rica have the wing of the petiole much reduced or absent: Burger & Liesner 6282; Jiménez 622; Burger & Burger 7659; Skutch 3261. The petioles of Burger & Liesner 6282 are to 10 cm in length. As no fruit was seen from Panama, the above description of fruits was taken from Croat 44491 (MO) from Costa Rica. VERAGUAS: l4 km NW of Santa Fe, road to Calovebora, Mori & Bolten 7653 (MO). 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) 517 INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (?) refer to names incidentally mentioned. Agylophora 509 Faramea 376+ Annona 289+ Ficus 346+ Appunia Fuchsia 3017, 490 parviflora 374 Galium Arachnimorpha 463 —sect. Relbunium 459 Arachnothryx 463 Gardenia buddleioides 463+, 465 armata 444 Arariba 492 maritima 446 mussaenda 446 Asclepiadaceae 3671 . Basanacantha 442 maxonii 494 lasiantha 448 Gomosia 290 Bertiera panamensis 454+ Gomozia 290, 2901 Blepharodon granadensis 2907, 292 mucronatum 367+ Gramineae 5141 Cald i Hedy klugei 494 lancifolia 295 Calycophyllum longifolia 295 coccineum 512 stanleyanum 515 striata 39 Carmenocania Howardi porphyrantha 331 caracasensis 331 Cascarella 263+ grandiflora 331 Cascarilla 263, 363+ Ixora 257 bullata 264 coccinea 257+, 258 macrocarpa 266 findlaysoniana 258 kasama pui 3351, 4817 floribunda 260 ti ——— E A 261 aeai 374+ wolfioides 261 plagiantha 373 Joosi: sia polycephala 351 an psychotriaefolia 417 umbellifera. ate 2627 rigidifolia 313 Ladenbergia 263 tetragona 359 ul Chrysoxylon 329, 3297 colombiana 267+ febrifugum 3297 dichotoma 263+ Cinchona laurifolia 265 macrocarpa 266 macrocarpa 266 Citrus 4097 Lasionema Compositae 5141 glabrescens 275 Conotrichia 277 Lauraceae 266+ alba 2777 PNN 401* Coussarea 434 Lightfootia 463 nis 4 Limnosipanea 267 euryphylla 453 palustris 268 Cucurbitaceae 355+, 514+ spruceana 267+ Diodia Lindenia 270 ottonis 505 acutiflora 270 Dukea 451 rivalis 2707, 270 ii 452 vitiensis 2721 euryphylla 453 Lygi 27 darienensis 453 Machao 7 panamensis 4517, 454 acuminata 272, 272+ victoriae 456 martinicensis 273 Eugenia 3761 rotundata 273 Exostemma — var. dodgei 273 occidentale 482 tysonii 272 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 Macrocnemum 275 lancifolia 295 cocci longifolia 295 glabrescens 375 Onagraceae 301+ jamaicense 275+ Ouratea 311 pastoense 277 Ourouparia 509 speciosus 331 guianensis Ped Manettia 27 tomentosa bocataurensis 278 Palicourea Jd 295, 295+, 2991, 3087, 3127, estrellae 3187 flexilis 280 angustifolia 297 hydrophila 280 ella 299 reclinata 277+, 281 brenesii 308+ stenophylla 278+ caerulea 351 Mapouria 335 carpirostrata 299 guianensis 335+ chiriquina 300 micrantha 396 costaricensis 303 obovata 403 crocea 300 subsessilis — var. riparia fo. riparia 3031 —var. angustifolia 374 fastigiata 301 Megaphyllum 319 galeottiana 2987, 303 Melastomataceae 2681, 270+ gibbosa 304 Microlicia 268+ grandibracteata 305 Mitracarpium guianensis 2957, 306 breviflorum 282+ intermedia Mitracarpus 282 lanceolata 297 breviflorus 282 lasiorrachis 307 brevifolius 284 macrosepala 308 hirtus 284 mexicane 3 scaber 282+ montensis 3 villosus 284+ Ronny de 310, 3677 Montamans 284 moritzii panamensis 2841, 286 ochnoides 310 Morinda 288 panamensis 311 citrifolia 28 pauciflora 312 namensis 289 rigidifolia 313 rojoc salicifolia 310 yucatanensis 290 stevensonii 354 Mu subrubra 313 coccineum 512 triphylla 315 formosa 446 tubuliflora 316 rotundifolia 443 tumidonodosa 3097, 317 Myristicaceae 2667 veraguensis Myristiphyllum 335 vestita 318 undatum 401 Palicurea 295 Myrtaceae 3767 Pallasia 514, 5141 Nacibea 277 stanleyana 515 Nauclea Pentagonia 319, 321+, 3267, 3277 aculeata 509 alba 321 tomentosa 509 bocataurensis 321 Nertera 290 brachyotis 322, 3271 depressa 2907, 292 gymnopoda 322 granadensis 290+, 292 macrophylla 319%, 20. 324 Nonatelia 335 magnifica 3207, 322 officinalis 335+, 404 nuciformis 326 pinnatifida 326 Noterophila 268+, 270+ pittieri 326 inundata 2687 tinajita Notopl veraguensis 328 uliginosa 434 wendlandii 329 Ochnaceae 3117 Petesi Oldenlandia 292 lygistum 277+ callitrichoides 294 Pogonopus 329 opu corymbosa 2927, 294 caracasensis 331 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) ottonis 3297, 331 dispersa 414 speciosus 321 dosbocensis 370 Polygonaceae 5141 dukei 371 Posoqueria 333 durilancifolia 372 decora 333 emetica 37 latifolia 333 erecta 374 longiflora 333+ eugenifolia 375 panamensis 333, 334+ fendleri 376 pittieri 507 fimbriata 476 svi 285+, 299+, 3121, 335, 3467, 3471, flexuosa 369 , 355+, 3607, 3621, 3667+, 371+, 3727, fluctuans x 386+, 390+, 396+, 406+, 407+, 413+, —var. angustifolia 374 4237, 4281, 4427 fockeana 356 acuminata 343 foveolata 356 —subsp. acuminata 343 fruticetorum 377 —subsp. boraginoides 344 furcata 377 aggregata 34. gatunensis 352 alba 356, 357+ glaucescens 384 albonervia 354 glo 39 alboviridula 345 goldmanii 379 allenii 3357, 346 graciliflora 378 angustiflora 348 granadensis 380 anomothyrsa 391 grandicarpa 352 asiatica 335+ grandis 381 axillaris 374, 3747 haematocarpa 382 bakeri 347 hammelii 383 bella 299 hebeclada 418 berteriana 348 hindsii 396 —subsp. luxurians 390 irta 401 bimea 384 —— 384 boquetensis 349 rr E: 384 brachiata 351 8 brachybotrya 350 ca 386 caerulea 351, 3517+ insignis 387 calophylla 347+, 352 inundata 354 capacifolia 353, 371+ involucrata 404 capitata 354, 3561 iquitosensis 350 —subsp. amplifolia 355+ aevis —subsp. capitata 354, 3551, 356+ lanceolata 401 —subsp. fissistipularis 355 liesneri 387 bsp. rubra limo is 389 carnosocarpa 3477+, 356 lon s. 389 carthaginensis 356 lupulina 36 catillicalyx 358 luxurians 2 3687, 390 chagrensis 358, 358+ macrophylla 372+, 391 chiapensis 347+, 359 —var. 2 391, 3927 chiriquina 3481, 360 magna 392 chloritica 361 ie pi 393, 3167 —a obovata 3621 maxonii —B bahiensis 361, 362* mexicana en 303+ — y lanceolata 362+ micrantha 3 cincta 362 microdesmia 439 compressicaulis 392 microdon 397 cooperi mima copensis 365 morae 403 copeyana 366 m cornifolia 476 mortoniana 401 crebrinervia 348 nervosa 3807, 401 crocea 300 —subsp. rufescens 401 croceovenosa 367 —subsp. rufescens var. hirta 401 cuspidata 3437 nubiphila 403 davidsei 368 obovata 403, 4307 deflexa 369 oerestediana 403 520 ANNALS OF THE MISSOURI BOTANICAL GARDEN officinalis 383+, 404 —subsp. orenidearum 2 407+ —s subsp. persedens phytolacca 434 pilosa 412, 421+, 4271 pinularis 397 pithecobia 414 pittieri 414 —subsp. oinochrophylla 416 psychotriaefolia 417 racemosa 422, 422 remota 3467 rosulatifolia 423 tacarcunensis 431 taurina 432 tenuifolia 380+ Sa was ai 437 ensis 438 gentryi 447 karstenii 448 DN 449 mira 450 mitis 442+, 443 darienensis 453 euryphyllum 453 palicoureoides 4511 —subsp. dwyerianum 4527, 454+ panamensis 4 trifoliatum 455 victoriae 456 Ravnia 456 longifilamentosa 458 panamensis 458 triflora 4567, 458 ma 464 versicolor 464 Rona 335 ere cta 374 latifolia 3357, 374 Rondeletia 463, 4707 americana 463* amoena bertierioides 465, 472* buddleioides 465 — var. buddleioides 466 odorata 470 — var. breviflora 470 — var. odorata 4707 1980] DWYER—FLORA OF PANAMA (Family 179. Rubiaceae) panamensis 470 platysepala 471 salicifolia 472, 472+ —var. salicifolia 472 subsp. brevicolla 473 MR versicolor 464 —sect. Involucratae 459 hispida 459 incana Rubiaceae eh 267+, 2707, 294+, 3351, 3931, —trit e Psychotriae 2857 Rudgea 474, 476%, 4817 Rustia 482, 483+, 4857 formosa 482+ occidentalis 482, 485+ is 4 —var. adpress —var. villosa ry 489+ Schiedia 461 martinicensis 273, 461+ ra 490 n 25 ri ca 2n 260, 2607 Sideroxyloides 2 maxonii 4944, 494 rubescens 494+ arborescens 496+ donnell-smithii 498 mesochroa 498 Stachyarrhena 500 i 04 Tobagoa Uncinaria 509 chloritica 361 Vaillantia aii wien a 521 522 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 67 gymnopoda 322 Willdenovia 463 macrophylla 3197, 324 schreberi 4637 magni Wittmackanthus pittieri 326 stanleyanus 5147, 515 pubescens 324 Xerococcus 4427, 516 tinajita 327 congestus 4427, 516 wendlandi 329 Zantedeschla 4617 ANNALS OF THE MISSOURI BOTANICAL GARDEN FLORA OF PANAMA! By RoBERT E. Woopson, JR. AND ROBERT W. SCHERY AND COLLABORATORS Part V, Fascicle 52 FAMILY 83. LEGUMINOSAE? SUBFAMILY PAPILIONOIDEAE (CONCLUSION) JOHN D. DwvER* & COLLABORATORS? Trees, shrubs or herbs, occasionally armed. Leaves often alternate, pari- or imparipinnate, frequently 3-foliolate, rarely 1-foliolate; stipules usually present; stipelles mostly present. Inflorescences usually racemose or paniculate, often axillary. Flowers papilionaceous, rarely actinomorphic; hypanthium more or less 1 Assisted by National Science Foundation Grant DEB 77-04300 (W. G. D’Arcy, principal in- vestigator, Flora of Panama panoan, pa in parenthesis at the bottom of pages of early treatments, was discontinued after Part V. The tents uel this and other parts of the Flora should be cited as: Ann. Missouri Bot. Gard. p ia number, pagination and date of the A m 3 Continued from Ann. Missouri Bot. Gar "1 52: 54. February [March 31] 196 4 St. Louis University and Missouri Botanical Garden, Box 299, St. Louis, E 63166. milies rt W. P4 and d ger in the Flora of Panama, Part V, Fascicles 2 and 3; Ann. Missouri Bot. that time. Of the tribe Dalbergieae, the genus Lonchocarpus was contributed by F. J. Her more than a decade of intensive botanical aa and collecting in Panama passed before it was eae p to consider resumption of publication of Dwyer’s treatment, it was decided to revise the earlier treatment completely. This was done under the direction of W. G. D'Arcy working under the guidance of John D. Dwyer. This revision of the earlier manuscript enlisted the help of a number of spe ecialists, cisions e- sponsible for his treatment. Each author is acknowledged under the genera he contributed. In yes e Michael O. Dillon, Peter S. White and James Lackey contributed largely to preparation of the ar published parts of the Papilionoideae treatment, i.e., that of the Dalbergieae, the subfamily descrip- tion, the relevant literature, the synoptical key to tribes are repeated here with some revision. Also, 524 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 campanulate, the teeth 5, the carinal tooth often the longest; petals 5, disposed as an upper standard or vexillum, 2 wing petals or alae, and 2 keel or carinal petals coherent apically, occasionally united in part with the wing petals and enclosing the androecium and the gynoecium, occasionally rostrate and coiled at the apex; stamens 10, rarely 9 or less, occasionally free, usually monadelphous or diadelphous, the free filaments equal or often alternating long and short, the anthers monomorphic or dimorphic; ovary l-carpellate, often surrounded by a glandular disc, 1- to several-ovulate, the stigma capitate or lateral, often enlarged, occasionally bearded. Fruits mostly dehiscing along 2 sutures, often pluri-loculate y the development of septa, occasionally indehiscent and samaroid, drupaceous or lomentaceous; seeds I-several, often arillate, exalbuminous, the radicle of the embryo curved. Of the subfamilies of Leguminosae, the Papilionoideae, whether regarded on a world-wide basis or locally in Panama, contains the greatest number of genera and species and largest representation of plants economically important, as food, drugs, timber or dyes. few genera of Papilionoideae, e.g. Clitoria, Phaseolus, bear uncinate hairs. These minute, usually hooked trichomes usually require the aid of a lens and are best seen in silhouette by viewing the edges of surfaces. The conspectus of the Papilionoideae presented here has been moded from the traditional concepts of older workers. Recently however, a number of inno- vations have been recommended. For example, Lackey (1977) recommends re- moval of Pachyrhizus from the Phaseoleae-Phaseolinae to the Diocleinae, and R. M. Polhill of Kew in a memorandum preparatory to the volume, Advances in Legume Systematics to be published in 1980 by the Royal Botanical Gardens Kew, recommends acceptance of several tribes not recognized here and of several shifts of genera from one tribe to another, e.g. of Dipteryx from the Dalbergieae to a tribe Dipteryxeae, and of Centrolobium, Lonchocarpus, Piscidia, Muellera from the Dalbergieae to a tribe Tephrosieae, together with Tephrosia, Barbieria and Willardia from the Galegeae; elevation of subtribe Galegeae-Indigoferinae to tribe Indigoferae; removal of Dalea from the Galegeae to a tribe Amorpheae; removal of Sesbania from the Galegeae to a tribe Sesbanieae; separation of Des- modium and Alysicarpus from the Hedysareae (Aeschynomeninae) to a tribe Des- modieae; removal of Abrus from the Vicieae to a monotypic tribe Abreae; removal of Crotalaria from tribe Genisteae to a tribe Crotalarieae, and elevation of sub- tribe Galegeae-Robiniinae to a tribe Robinieae but removing Cracca from this. Polhill does not separate Stizolobium from Mucuna but separates Cymbosema from Camptosema. the genera of the Dalbergieae appearing in the already published portion have been incorporated into the key to genera. Many of the illustrations presented here were prepared in the period before 1965 and, in some cases, vouchers could not be relocated to verify determinations, localities and magnifications. De- terminations for these collections were reviewed by the senior author (Dwyer) at the time the illus- trations were made and it is presumed that they accord with the concepts of the various collaborating authors. All illustrations are based on material available at the time at MO. —W. G. D'Arcy 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 525 Literature: Amshoff, G. J. H. 1939. On South American Papilionaceae. Meded. Bot. Mus. Herb. Ryks Univ. Utrecht 52: 1-71, fig. 1-4. . 1939. Papilionaceae in Pulle, Flora of Suriname 2(2): 1-257. Auctores. 1954. Papilionaceae in Flore du Congo Belge et du Ruanda-Urundi (Publ. I.N.É.A.C.), Spermatophytes 4. 1953, 5 & 6. Baillon, H. 1870. Légumineuses in Baillon, Hist. Pl. 2: 21-384, fig. 15-157. Bentham, G. 1837. Commentationes de leguminosarum generibus. Vienna. . 1841. De leguminosarum generibus commentationes. Phyt. Ann. Wiener Mus. Naturgesch. 1 & 2: 63-142. . 1862. Papilionaceae in Martius, Flora Brasiliensis 15(1): 1-216, tab. 1— 56. 1859. 217-350, tab. 57-127. Burkart, A. 1943. Las Leguminosas Argentinas Silvestres y Cultivadas. Dalziel, J. M. 1937. The useful plants of West Tropical Africa. An appendix to the Flora of West Tropical Africa. 227 pp. London. DeCandolle, A. P. 1825. Mémoires sur la Famille des Légumineuses. 825. Leguminosae in DC., Prodr. 2: 93-524. 1825. Ducke, A. 1949. Las Leguminosas da Amazonia Brasileira (Notas sóbre Flora Neo-trópica—II). Bol. Técn. Inst. Agron. Notre (Belém) 18: 3:248. Heywood, V. H. & P. W. Ball, eds. 1968. Leguminosae in T. G. Tutin et al., eds., Flora Europaea 2: 80-191. Hutchinson, J. 1967. Genera of Flowering Plants. Vol. 2. 659 pp. Clarendon Press, Oxford. Jacquin, N. A. 1763. Selectarum stirpium Americanarum historia. Vienna. Johnston, I. M. 1949. The botany of San Jose Island. Sargentia 8: 1—306. Lackey, J. A. 1977. A synopsis of Phaseoleae (Leguminosae Papilionoideae). Ph.D. dissertation, Iowa State Univ., unpubl. Macbride, J. F. 1943. Mimoseae in Macbride, Flora of Peru. Publ. Field Mus. Nat. Hist., Bot. Ser. 13(3): 4-113. Pellegrin, F. 1949. Les Légumineuses du Gabon. Mém. Inst. Étud. Centrafr. Brazzaville 1: 1—248. pl. 1-8. Perkins, J. 1907. The Leguminosae of Porto Rico. Contr. U.S. Natl. Herb. 10: 133-220. Pittier, H. 1928. Arboles y Arbustos del Orden de Las Leguminosas. III. Papi- lionaceas. Bol. Min. R.R.E.E. 4-7: 149-229. Rudd, V. E. 1972. Family Leguminosae, subfamily Faboideae, tribe Sophoreae. N. Amer. Fl. II(7): 1—36. Rydberg, P. A. 1923-1929. Fabaceae, Indigofereae and Galegeae. N. Amer. Fl. 24(3-7): 137-462. Standley, P. C. 1928. Flora of the Panama Canal Zone. Contr. U.S. Natl. Herb. 27: I-X, 1-416, fig. 1-7, pl. 1—66. . 1933. The Flora of Barro Colorado Island, Panama. Contr. Arnold Ar- bor. 5: 1-17, pl. 1-21, map. & J. A. Steyermark. 1946. Leguminosae in Flora of Guatemala. Fieldiana Bot. 24(5): 1-367. Taubert, P. 1891-1894. Leguminosae in Engler et Prantl, Nat. Pflanzenfam. III(3): 70-288, fig. 38-136. 526 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Taylor, A. S. 1976. Las Leguminosas papilionaceas herbaceas y arbustivas de anama. Editorial Universitaria (Panama). p Thuan, N. van. 1979. Légumineuses-Papilionoidées Phaséolées. in A. Aubréville & J. F. LeRoy, eds. Flore du Camboge du Laos et du Viét-Nam 17: 1-218. Verdcourt, B. 1971. Phaseoleae in E. Miln-Redhead & R. M. Polhill eds. Flora of Tropical East Africa Part 4(2): 503-807. CONSPECTUS OF TRIBES AND SUBTRIBES*® a. Stamens free; leaves mostly pinnate, leaflets numerous; shrubs or trees _____- Tribe Sophoreae 2. Acosmium 5. Ateleia 21. Dussia 33. Myroxylon 34. Ormosi 40. Sophora aa. Stamens united, at least in part; leaves and habit various. b. Fruit articulating between the seeds Tribe Hedysareae Anthers all alike Subtribe Desmodiinae 4. Alysicarpus 8. Desmodium cc. Anthers dimorphic. d. Stamens 10, the filament tube closed Subtribe Stylosanthinae 2. Stylosanthes 9. Zornia dd. Stamens 5 + 5, the filament tube open ____________ Subtribe Aeschynomeninae 3. Aeschynomene 11. Chaetocalyx bb. dae 2- beide or indehiscent Leaves entire or 3-5 digitate. Leaflets denticulate Tribe Trifolieae 45. Trifolium ff. Leaflets enti Tribe Genisteae Stamens ; 10 with open filament tube _______________. Subtribe Crotalariinae 15. Crotalaria 30. Lupinus gg. Stamens 10 with closed filament tube. Subtribe Genistinae 46. Ulex ee. Leaves pinnate h. Leaf terminating in a bristle Tribe Vicieae 1. Abrus hh. Leaf terminating in a leaflet. i. Pod indehiscent Tribe Dalbergieae Published previously (Geoffroeae) Andira Centrolobium Dalbergia a 1m Dipteryx Fissicalyx € Adapted in part from Taubert (1891), Dalla Torre and Harms (1907), and Melchior (1964). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 527 I" mostly dehiscent. Ova t: k. kk. pp. Lonchocarpus (Paramachaerium) Machaerium Mueller. Piscidia Platymiscium Platypodium Pterocarpus y surrounded by a disc Tribe Phaseoleae Style poe (except Clitoria), rarely pubescent below — — ary stamen le eee with the others. tals about e n. fie ee rachis nodose ______________ ubtribe Diocleinae 9. Canavalia 19. Dioclea nn. Inflorescence rachis not nodose ---------- Subtribe Glycininae 10. Centrosema 13. Cologania 44. Teramnus mm. Tas the standard or the keel much larg, than the other ---------------- Subtribe inks 23. Erythrina 32. Mucuna 41. Stizolobium Vexillary stamen mostly completely fre o. Inflorescence rachis nodose; leaflets eglandu- lar ubtribe Galactiinae 8. mi 16. Cymbosem 25. Galactia oo. Inflorescence rachis not nodose; leaflets glan- dular beneath Tribe Cajaninae 7. Cajanus 22. Eriosema 24. Flemingia 38. Rhynchosia Style pubescent along the inner edge or around Subt th ribe Phaseolinae N . Lablab 31. Macroptilium 35. Pachyrhizus 36. Phaseolus 47. Vigna zt lacking a dis Tribe Galegeae Anther dior with gland, bristle or apiculum _________- ubtribe Indigoferinae 27. Indigofera Anther connective unappen q. ndaged. Ovules mostly 1-2, "ib glandular punctate ________ Subtribe Psoraleinae 17. Dalea 528 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 qq. "aqa mostly numerous; plants not punctate. Inflorescence terminal or . opposing IT. a leaf 6. 43. Barbieria Tephrosia Inflorescence always axillary 14. 20. 26. Cracca Diphysa Gliricidia 29. Lennea 39. 48. Sesbania Willardia tribe Tephrosiinae Subtribe Robiniinae 529 DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. 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WS og uey} sso] APOOM Jt ‘sqiay Io SIULA snuefgO | ^ Ə1e[onəd saava] ‘z uey} 23Joui spass :3uo| WD ç uey; BOW *oje3uo[o pod `,9ç PUUƏSOLI "7C e[rssosqns soAe9| ‘z spass :3uo| WD ç uey} sso| “ous pod '96 "pounds jou sjoqjue ‘pim uiui ç uey} aow “utu1 pod :qjyeasuoq ponop puer? saava] :poxuiseq siteH `,çç e19josipu[ '/z ^ pounds sioque :ƏpiA WW e uey} SSƏ[ *urgj pod :1e[npue[8o soAeo| :poxyrpoul SIEH "c6 "duo| ui» Ç`] ULY} sso[ *1o[[euis s12^0[4 `,£ç uiros feo ee a SUUS MMC NIMES DUE ILES pol jou *x1ep spəəs ‘quaoseqnd ə[Á1s +4133} peugop [[2^ ç YUM xÁ[e» ‘Burysew o[dund ym ustAo||ƏÁ JO INYM S19^0[] BULIWJAIA ‘EZ — pal Udo spəəs :snoJqe[3 [Ais :poqo[ 1eadde oj 3unjids souimouios əuou JO u]991 2jnurmu YIM XxÁ[83 *o3ug1o 10 Pl SIOMOL "pg 'Suo[| WD ç’; uey} 21oui *oSre| SIIMO EÇ "ues 533 DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 1980] TANCOS A EES ENTE uə||oAs JOU səpou o2uo5soaJogut :221j u1991 xA[e5 1oddn ‘asidasues pJoj [29x oui ‘seld 13470 Əy} ueu) 1o31e| yonw s3ut `ç/ "dunue^ sey ojeur»un :suonn[oA2a1 ç—ç u83noug) uou) poj[oo» Jt ‘snoueA o[Ájs :snonoidsuoour suə| əy} 'LC-9 SpƏƏs ‘Zumo 210joq snoonpeo so[oo) -9eJq pu? sj5*q :3uo| WW /[-] Spd Sung ^, p snjoəseyd `9ç sirey ojeuroun 3uipn[our 22uoosoqnd < Sano -9e1q pue sioe1q :8uo| ww Jg- t s[oorped 3uniniq ‘pL "ÁIeAO ƏY} JO PY uioJj IISIP o[Á1s jo əouəosəqnd :sjueuuio1 UOTYSNd INYM YIM pa19A02 Ajjensn wnpy :suonoujsuoo əsiəAasuen oym pod ' snzmÁugoed 'cg 7777 ` woyoq 01 do} wouy snonunuoo adUadsaqnd 9[A1s :Ə|qISIA DAOOIZ 1EY BY} ‘pataaooun uinjnq ‘opts -INO SUONSLSUOS 3sJoASUEJ] YIM possouduioo pod ‘¢/ `919191 a]Ajs :3uo[ WW çz—ç ‘/7—-¢ spoos ‘snondidsuoou! 10 3unueA UOTYsNd “jəjəuiuəd pass oui £L se Suo| se £4 uey} sso[ unu :ə|Á1s juojsis1od Nom pod `,Z¿ eT g = RR ne U COE poupe ejas :3uo| ww 7[-6 ‘9-¢ spəəs :uorgsno ÁJuods “əJuA *jusunuo4d e ym pƏ4Ə9A02 awd poos ay) se 3uo| se 74-54 un|jiu :3uo| ww 0[-8 luəlsisiəd o[A1s YIM pod "essa AIWAO :291j sjo€Jq :3uo| WW ç uey) sso] *oje[nueduieo xÁ[e) ',]/ €LO]I[) ‘ZI 9je1dns ÁK1eA0 :9j:?uuoo sjoeiq Joddn :3uo| ww ç uey} o1oui “zea xÁ IL "^O[[9Á JOU Apsow *en[q “əldind ‘yurd ‘pas siamoy ‘z uey} o1oui Ansoui spəəs :3uo| wə z uey sow pod ‘juasaid s|ədns :(snoonpeo souinouios) 1u3səid s13eiq aUIDATeD `,69 BISOYDUAYY "QE rrr rrr nnn nn nnn nnn nnn ne AO||ƏÁ SIOMOY ‘Z spass :3uo| Wd z uey} ssə| pod :Sunue^ sjadns ‘Zunuem sjoe1q IUDA ‘6g "exe s1oqiue ƏY} *0I—-8 suouiejs AMAA [VoL. 67 ANNALS OF THE MISSOURI BOTANICAL GARDEN 534 snuweloay ‘pp ^ ^ — juajsis1od Á[isoui sj2eJq :3uo[ uiu g uet) SSƏ[ e[[oJoo :sopouruiejs ç ui^ Sureuloj[e ç s1oqjue tHous ÁIƏ9A 3JÁ}S :3uo| uiu ç uey} sso[ *oje[jnuedureo xÁ ',08 plue3o|oO `€ snonpioəp s35eJq :3uo| un [ uey} 2a1ou) e[[oJ05 :uuogmun ‘QO, s1ouqjue :3uo[ ww ç'I uet Əiouu əƏ[Áis ‘Zuo ww ç uey} aow *1Te[nqn] xÁ[eO “Og 'osopou JOU saousosoaJopgu[ ',9/ Enen “SZ Suo| uio | uey} sso| ‘AUN siəmog :spoos əu) punojge [eljojeu! Əso| -n|[93 YUM pod :3uo| ww 9 uey} sso[ *19[[euss spəəS :,6/ pə|oolG '6I adie] s12^0p :Apoom *eje1dos pod :3uo[ ww g uey) Əioui 'oSre| Sp99g `6¿ ‘ayeao JO jendi SIAL :ç—p S9qo[ xA[e5 {pases ou) SurjoioSuəo 10 uoys ‘Teau umpu Iy} YUM PI ',// uiniuo3odo[e;) *g snoonpes ‘TEU sj2eJq [LIN OAU! :uAOJq 10 u9918 SUIAIP spod `,8/ euelong ` / £ SutiəƏmoj YsNoIY) juojsis -19d *aje[oooue| saq [e1onjoaur *x»e[q BUIAIP SpOg ‘g/ `31quioui SOABI] fÇ SOQO] xÁ[e5 TEMIN ‘ews WHY I4} YM p99S "// ‘asopou Á[snonoidsuoo əoSuəosəiogul :9/ *snoiqe[3 Ə[Á1S `,0¿ eUSIA “LP -7 uo[[o^s 1e[npue[8 ÁA[snonoidsuoo sopou Əouəosəioguir :93jeuuoo Ápued Á[rensn u1ə91 xÁ[|e» 1oddn :3unueA JO [e19je[ p[oJ [99* ‘səd 19y}0 oy) Sun[penbo jnoqe s3ulA :',6/ 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 535 1. ABRUS W. G. D'Arcy? Abrus L., Syst. Nat., ed. 12. 2: 472. 1767. TYPE: A. precatorius L. Mostly twining vines, seldom shrubby. Leaves even pinnate; leaflets mostly numerous, mucronate; rachis grooved, mucronate; stipels minute; stipules per- sistent. Inflorescences axillary or lateral pseudoracemes, terminating leafy or leafless branches, with 1—several sessile flowers at a node; bracts and bracteoles often caducous. Flowers with the calyx campanulate, lobed or short toothed; standard apically notched and with a short claw, the wing petals shorter than the keel petals; stamens 9, monadelphous, the staminal tube adnate to the standard; style curved, glabrate. Legume oblong to linear, much compressed or not, the valves pubescent, often becoming indurated, septate, the short slender beak downturned; seeds ovoid, sometimes compressed, arillate, sometimes with an annulus around the hilum. This genus resembles members of the Phaseoleae in its many leaflets which are borne equal pinnately. The 9 stamens with united filaments are distinctive, as is the lack of a terminal leaflet. The genus is circumtropical and, according to circumscription, includes 4-24 species. Only 2 of these species occur in the New Wor Literature: Breteler, F. J. 1960. Revision of Abrus Adanson (PAP.) with special reference to Africa. Blumea 10: 607-624. Verdcourt, B. 1970. Studies in the Leguminosae-Papilionoideae for the ‘Flora of Tropical East Africa’: II. Kew Bull. 24: 235-307. 1. Abrus precatorius L., Syst. Nat., ed. 12. 2: 472. 1767. LECTOTYPE: ae i Hermann, Fl. Zeylan. 284 (BM, not seen), fide Breteler 1960.—Fic Slender vine; stems twining, pubescent with appressed white hairs, glabres- cent. Leaves pinnate with 5—13 pairs of leaflets; leaflets oblong to ovate, mostly 9-15 mm long, 5-7 mm wide, apically rounded or truncate and mucronulate, sometimes acute, basally obtuse, rounded or truncate; petiolules 0.5-1 mm long; petioles shorter than the width of a leaflet; rachis mostly 7-10 cm long, sparingly pubescent with ascending hairs; stipels minute, following the rachis; stipules sub- ulate, 1-2 mm long. /nflorescences leafless pseudoracemes; peduncles stouter than the petioles and often stouter than the stems, 6—10 cm long, bearing ca. 4 pairs of minute bracts along the length; rachis strongly angled, somewhat con- gested, 3-6 cm long, the flowers 1-15 per node, pedicels 1-2 mm long, glabrous except for the apical swelling. Flowers white or pinkish, ca. 1 cm long, calyx cupular, ca. 2 mm long with low teeth, sparingly short pubescent; standard ob- ovate, 8-9 mm long, the wing petals upcurved, ca. 6 mm long, stamens not exserted. Legume oblong, somewhat oblique, ca. 2.5 cm long, 1.5 cm wide, 7 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 536 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 ? aw Zz (SU) < Z e eRe, SE se a. WAS 4 / GN // f: j N a ^ N ! Ü A FIGURE 1. Abrus precatorius L.—A. Habit (x4). [After Wedel 1731.]—B. Flower (x3).—C. Corolla.—C!. Standard (x2).—C?. Wing petal (x2).—C?. Keel petal (x2).—D. Stamens (x3).—E. Pistil (x3). [After Wedel 1410.]—F. Fruit (x1). [After Gentle 1203, Belize.]—G. Seed (x2). [After Wedel 1308.] hardly stipitate, the beak slender, 3-5 cm long, downturned, dehiscent along 1 margin, the valves softly pubescent, compressed but bulging over the seeds; seeds 5-6, subglobose or slightly compressed ovoid, bright shiny scarlet with a black area near the hilum, ca. 6 mm long, the hilum depressed, ca. 1 mm long. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 537 Abrus precatorius is usually immediately recognizable by its bright shiny red and black seeds. The vine bears leaves with many leaflets, and the congested pseudoracemes of small flowers is also distinctive, but these characteristics may lead to confusion with members of the Phaseoleae. This species occurs in tropical and warm temperate countries on both hemi- spheres. In Panama it is apparently confined to the Caribbean coast. The bright seeds of this species are much used for beads. They contain highly toxic compounds and a single seed may be lethal to a man if the hard seed coat is broken before ingestion. The toxic principle is also found in the roots although the roots are reportedly eaten as food in India. The English name is Precatory Beads, and in Spanish they are known as Bejuco de Peronía or Bejuco de Peonía. Nevertheless many common names have been applied to this plant. BOCAS DEL TORO: Chiriquí Lagoon, or 1304, 1308 (GH, MO). Water Valley, Wedel 1371 (GH), 1410 (GH, MO). Isla Colón, Wedel 2822 (GH). SAN BLAS: Mainland Point, Duke 8981 (MO). Ailigandi, Dwyer 6828 (MO). Mainland opposite Ailigandi, Lewis et al. 77 (GH, MO). Playon Chico, Stier 153 (MO). 2. ACOSMIUM Michael O. Dillon® Acosmium Schott in Spreng., L. Syst. Veg., ed. 16, 4: 406. 1827. TYPE: Acosmium lentiscifolium Schott. Leptolobium Vog., Linnaea 11: 388. Apr.-Jun. 1837, non Bentham, Lu 1837; 1838. LECTOTYPE: eptolobium dasycarpum Vog. = Acosmium dasycarpum (Vog.) Yakovl. Thalesia Mart. ex Pfeiffer, Nom. Bot. 2(2): 1384. 1874, nomen nudum A 484 on Leptolobium Vog., non Raf. 1818, nomen nudum Trees or shrubs, to 40 m, unarmed. Leaves alternate, imparipinnate; leaflets alternate to subopposite, (3—)5—21, lanceolate to ovate, apically retuse or emar- ginate, basally obtuse to truncate, subcoriaceous or membranaceous; stipules small, caducous or inconspicuous; stipels minute or absent. Inflorescence of ra- cemes or panicles, terminal, rarely completely axillary; bracts and bracteoles narrow, small, often caducous. Flowers small, sessile or pedicellate, calyx tur- binate-campanulate, lobes usually 5, unequal, valvate or rarely narrowly imbri- cate, corolla yellowish white, petals 5, all similar, free, erect patent; stamens 10(5), free, subequal, exserted; filaments inflexed; anthers uniform, ellipsoid, dor- sifixed; ovary sessile or short stipitate, I-4-ovuled; style filiform; stigma small or truncate, terminal. Fruit oblong to broadly linear, plano-compressed or rarely turgid, indehiscent, coriaceous or woody, narrowly winged, reticulate: seeds ovate or orbicular, compressed, brown to reddish brown, the hilum apical, ellip- tic. Chromosomes: n = 9 Acosmium sensu Yakovlev (1969) is composed of some 17 species, primarily distributed in tropical and subtropical regions of the Western Hemisphere. Its center of diversity is in Brazil, with only Acosmium panamense extending north into Central America to southern Mexico. While most species are rare and of * Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 538 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 2. Acosmium panamense (Benth.) Yakovl.—A. Habit (x1). little economic importance, several of the more widespread species are valuable timber trees used in general construction and the manufacture of charcoal. Literature: Mohlenbrock, R. H. 1963. A revision of the leguminous genus Sweetia. Webbia 17(2): 223-263. Rudd, V. E. 1968. Leguminosae of Mexico—Faboideae (I. Sophoreae and Po- dalyrieae). Rhodora 70: 492-532. Yakovlev, G. P. 1969. A review of Sweetia and Acosmium. Notes Roy. Bot. Gard., Edinburgh 29: 347—355. 1. Acosmium panamense (Benth.) Yakovl., Notes Roy. Bot. Gard., Edinburgh 29: 353. 1969.—Fic. 2. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 539 Sweetia penini Benth., J. Linn. Soc., Bot. 8: 263. 1865. TYPE: Panama, Canal Zone, Paraíso tation, Panama Railroad, S. Hayes 267 (K, not seen; photo, NY 2807). ibana lavigara Sine. Trop. Woods 12: 5. 1927. TYPE: Belize, lower Belize River, Record (US, n Trees, to 12 m, the crown rounded. Leaves alternate, imparipinnate; rachis slender, canaliculate, puberulent, 6-12 cm long; leaflets 7-13, subopposite, 3-8 cm long, 2-3 cm wide, ovate to elliptic, apically obtuse, usually retuse, basally obtuse to broadly cuneate, coriaceous, nitid above, glabrous, pale beneath, gla- brous, reticulate; petiolules ca. 3 mm long, terete, rugose; stipules linear, ca- ducous. Inflorescence paniculate, branched, axillary to terminal leaves, puber- ulent, primary rachis 8-14 cm long, secondary 2-5 cm long; bracts and bracteoles linear, 1-3 mm long, caducous. Flowers numerous, ca. 6 mm long, pedicellate, pedicels ca. 3 mm long, puberulent; calyx turbinate- campanulate, persistent, 3— 4 mm long, puberulent, 5-lobed, teeth ca. 2 mm long, acute, subvalvate; petals 5. free, white or cream colored, subequal, ca. 4 mm long, 2-3 mm wide, spatulate, emarginate, clawed; stamens 10, filaments equal to petals; anthers ca. 1 mm long, ovoid, dorsifixed, longitudinally dehiscent; ovary glabrous, stipitate, the stigma minutely capitate-penicillate, 2-3-ovulate. Fruit oblong or elliptic, 5-9 cm long, (1.52)2.0-2.5 cm wide, coriaceous, compressed, ca. 5 mm thick at the seeds, reticulate, the ends acute to obtuse, mucronate; seeds 1—3, lustrous, reddish rown, subovate, ca. 8 mm long, 6 mm wide, 2-3 mm thick, the hilum ca. 1 mm long. Acosmium panamense is widespread from southern Mexico to northern Co- lombia and northwestern Venezuela, usually found in moist forests up to 800 m. This species is a valuable timber tree for its wood is strong, durable, and suitable for heavy construction such as bridges, wagons, and woodworking. Its bark is said to be used as an antimalarial in El Salvador. There are many local names (cf. Rudd, 1968) for this species, known as *'malvecino"' in Panama. CANAL ZONE: Ancon Hill, 20 m, niei 6535 (MO). Red Devil drop zone, Howard = Force Base, Tyson 1871 (MO). DARIEN: Rio Con o, Holdridge 6296 (SCZ). Patiño, Pittier 6609 (G H, US) (leaves only). PANAMA: Taboga Island, MeDaniel & Cooke 12701 (FSU, MO). Agricultural Experi- ment Station, Matias Hernandez, Pittier 6793 (F, GH, MO, NY) 3. AESCHYNOMENE Michael O. Dillon? Aeschynomene L., Sp. Pl. 713. 1753. TYPE: A. aspera L. Gajati Rumph. ex Adans., Fam. Pl. 2: 328. 1763. po ti aspera L. Macromiscus Turcz., Bull. Soc. Nat. Mosc. 19(2): š 1846. pe M. brasiliensis Turcz. = A. montevidensis Vog Ctenodon Baill., Adice: 236. 1870. TYPE: C. weddellianum = A. oroboides Benth. yii o dei Hemsley Rose, Contr. U.S. Natl. Herb. 8: 43. 1903. TYPE: C. mexicana Hemsley & e — A. villosa Poi Secula Small, Fl. Miami 90, 200. 1913. TvPE: A. viscidula Michx. — A. oroboides Benth. Herbs, shrubs, or small trees; stems terete, finely striate, unarmed. Leaves Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Td Chicago, Illinois 60605. 540 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 alternate, subimparipinnate, 1-10 cm long; leaflets 5—80, alternate or subopposite, subsessile, 2-40 mm long, 1-20 mm wide, glabrous to sericeous, often minutely glandular-punctate, 1-several costate; petioles 2-40 mm long; stipules peltate, appendiculate below the point of attachment or attached at the base and not appendiculate, membranous or foliaceous, persistent or caducous. Inflores- cences racemose, paniculate, fasciculate, or rarely solitary, axillary or terminal, bracteate, bractlets paired and subtending calyxes, pedicels glabrous to pubes- cent, usually glandular. Flowers 5-15 mm long; calyx bilabiate to campanulate, commonly persistent; corolla yellowish to red or purple, petals usually clawed; stamens 10, essentially diadelphous, 5:5, filaments united into a sheath, open on carinal side, and with splitting on vexillar side delayed or merely incipient; anthers dorsifixed, or occasionally basifixed; pollen ellipsoidal, tricolpate; ovary 2-18 ovulate, rarely 1-ovulate, sessile or short stipitate; style incurved, not bearded; stigma terminal. Fruit a loment, (1—)2-18-articulate, subsessile to long stipitate, laterally compressed, occasionally torulose; seeds reniform, smooth, sublustrous, light brown to black, hilum circular, embryo axile, slightly bent, endosperm pres- ent, cotyledons fleshy. Chromosomes: n = 10 The genus Aeschynomene is composed of about 350 species distributed throughout the warmer parts of the world. In the Western Hemisphere, its pre- sumed center of origin, some 70 species range from the mid-Atlantic states in the United States, west to Baja California, Mexico, throughout Central America, the Antilles and south to the Rio La Plata, Argentina. Two American sections, Aes- chynomene and Ochopodium, are recognized by Rudd (1955), both with repre- sentative species in Panama. Most taxa appear to be of little economic importance; however, the pith from certain South American species is used for corks and for stropping knives and razors. Various Panamanian species are commonly referred to as “pega-pega.” Literature: Rudd, V. E. 1955. The American species of Aeschynomene. Contr. U.S. Natl. Herb. 32(1): 1-172. > “sasa peltate, Sasa passes below the point of attachment: calyx bilabiate, the upper lip 2-merous, the lower lip 3-merous (section Aeschynomene). Lealali with piben costae. c. Ovary glabrous to puberulent; fruit glabrous to para ann and occasionally with scattered tuberculate based trichomes, less than 0.8 ule n length, sutures be- tween articulations well defined, usually more than 5-arti eae 1. A. americana cc. Ovary villous; fruit hispid with tuberculate based trichomes, 0. 8- d 4 B long, sutures not defined between articles, 5-articulate or less |... . A. villas bb. Leaflets with 1 central costa d. Fruit blackening on drying; calyx lips entire to subentir ipe and basal article separated by a suture: fruits submonitfrm with both margins crenate |. nsis var. caribaea ee. spe and basal article continuous, not separated by a iue e ith one margin entire, the other entire or crenate — 1 1 1. . A. sensitiva dd. iiu imn etic parts herd í or straw colored on drying, or remaining gre pue lyx lips definitely inden eaflets usually dee or - serrulate- ciliate, 10-20 mm long, 4-8 mm ue plant generally hispid; fruit 5-7 mm wide, hispid at MMC not muricate, both margins entire or nearly so. . ciliata 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 541 ff. Leaflets entire, 5-7(-10) mm long, 1-2(-2.5) mm wide; plant pe gla- brous or sparsely hispid; fruit 4-5 mm broad, glabrous at maturity or with icd trichomes 0.5 mm long, muricate, upper margin straight and lower 7. A. rudis ar aa. Stipules not Ed Ed below the point of attachment to stem; calyx campanulate A 5 = ubequa al lo i articles 2-3 mm in diameter (section Ochopodium). t 2- or 3-articulate, the stipe 1-5 mm long. Fruit u sually biarticulate; leaves and stems canescent ______ 5. A. histrix var. incana hh. Fruit 3- pom leaves pilose, stems hispid 2. A. brasiliana BE. Fruit 4- to 9-articulate, stipe 8-15 mm long 4. A. elegans Aeschynomene americana L.,'° Sp. Pl. 713. 1753. TYPE: Jamaica, Sloane (BM, not seen). — Herbs to 2 m tall; stems erect to ascending, subglabrous to hispid. Leaves 2— 7 cm long, 20-60 foliolate; leaflets linear-oblong, 4-15 mm long, 1-2 mm wide, subfalcate, 2-several costate, apices and margins usually serrate-ciliate; stipules appendiculate below the point of attachment, striate, glabrous to hispid, attenuate at each end. Inflorescence few flowered, axes flexuous, hispidulous; bracts cor- date, acuminate, 2-4 mm long, 2-3 mm wide, glabrous, margins ciliate; bracteoles linear to linear-ovate, 2-4 mm long, 1.0-1.5 mm wide, acute to acuminate, gla- brous, margins serrate-ciliate. Flowers yellow, (5—)6—8(—10) mm long, calyx 3-6 mm long, glabrous to sparsely hispid; petals 5-10 mm long, clawed, the standard suborbiculate to broadly obcordate, 5-10 mm wide, apices ciliate, wings obliquely oblong, 5-8 mm long, 2 mm wide, ciliate at the apex, keel blades 4-5 mm long, 2.5-3.5 mm wide; stamens 6-8 mm long. Fruit (3—)6-8(-9) articulate, articles 2.5-5.0 mm wide, 3-6 mm long, glabrous to puberulent, sometimes with glandular trichomes, verrucose at maturity, the margins thickened; seeds 2-3 mm long, 1.5-2.0 mm wide, dark brown. Aeschynomene americana is the most widespread and abundant member of the genus in Panama, occurring in nearly every province. It is distinguished from its closest relative, A. villosa, in possessing loments with generally more than 4 articulations, and with these possessing well-defined lateral sutures. Rudd (1955) recognized 2 varieties from Panamanian material based upon differences in fruit pubescence. Since no clear ecological or geographical pattern is apparent, this species is treated as being polymorphic for fruit pubescence with vegetative and floral characters remaining quite constant. OCAS DEL TORO: ers Valley, celia 343 (US). Water Valley, Wedel 1531 (GH, MO, US), 1616 (GH, MO). Colón, Wedel 3 (GH, MO, US). CANAL ZONE: BARRO COLORADO ISLAND: Croat 12817, Bor (both MO), s (DUKE, MO, NY, SCZ). Venado Beach, Correa & Gonzales 523 (DUKE, MO, PMA). Summit Garden, Croat 12830 (MO, NY, SCZ). Río Indio, Dodge A bs Trail, Hunter & ae 737 (GH, MO). 0.5 mi N of Gamboa, Lazor & Tyson 5685 (MO). ad Rio Petitpie on road to Ft. Sherman, Mori & Kallunki 3669 (MO). 6 km E of Gamboa, Nee 900 ). ipods Hig un way between Rodman Marine Base and Chorrera, Nowicke et al. 3597 DUKE. MO). Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, Piper 5190 (US). Río Grande, near Culebra Pittier 2112 (NY, US). Balboa, Standley 25518, 25540 (both US). Las Cascadas Plantation, near 10 For a list of synonyms see Rudd (1955). Only the listed names have been used for Panamanian materi 542 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Summit, Standley 25768, 25776 (both US). Summit, Standley 26921 (US). Near Gatun, Standley 27268 (US). Frijoles, Standley 27643 (US). Gamboa, Standley 28325 (US). Near Fort Randolph, Standley 28735 (US). Mount Hope Cemetery, Standley 28847 (US). Old Las Cruces Trail between on and Corozal, Standley 29059, 29182 (both US). Rio Pedro Miguel near East Paraiso, 6 (U en i i Tyson et al. 2752 (MO, PMA). iss secondary forest, margin of bs pete road N to Gamboa, Wilbur & Teeri 11245 (DUKE, GH, MO). Playa Venado, Wilbur & Teeri 12959 (DUKE). Pipeline road, NW of Gamboa, “asss, & Teeri 13378 (DUKE). cHiRIQUÍ: 2 mi W of Concepción, Baltimer (MO). Near Veladero, t & Koster 128 (MO). Rabo de Puerco, 8 mi W of Puerto Armuelles, Liesner 33 a c 2 Melliza, 6 mi S of Puerto Armuelles, Liesner 491 (MO). Boquete, Lewis et al. 391 (GH, MO, US). 2 mi S of Puerto Armuelles, Wilbur et al. 13579 (DUKE). cocLé: EI Valle, Baltimer 1470 (MO). Ola, Pu. 5092 (NY, US). Río Grande, Troetsch 24 (DUKE, PMA). 10 mi E of Nata, Tyson 5237 (FSU, SCZ). Penonomé, Williams 359 (NY, US). COLON: Calzada Larga, Gracia 48 (PMA). Quebrada Bonita, 3 km NW of Salamanca, 13 km E of Buenos Aires, Nee 9107 (MO). 1.2 mi NE of bridge over Río Piedras on road from Portobelo to Pilón, Nee & Mori 3669 (MO, PMA). Without definite locality, Rose 23997 (GH, NY, US). pARIÉN: Santa Fé, Duke 9481 (NY, US). HERRERA: Chitré, Allen 1104 (GH, MO, US). Ocú, Allen 4087 (MO). 4 mi S of Los Pozos, Tyson 2679, 2695 (both FSU, MO). 10 mi S of Ocü, Tyson 2822 (MO, US). Los SANTOS: Punta Mala, Tyson 2722 (SCZ). PANAMA: Isla Taboga, Allen 1283, 1295 (both GH, MO, US). 1 mi E of Tocumen, Blum & Tyson 1953 (FSU, MO, SCZ). Campo Bi, de Monte Oscuro, Cs 1914 (PMA). Be- tween Capira and Potrero, Dodge & Hunter 8604 (GH, MO). Río Indio, iis i & Allen 17378 (MO). Tocumen Airport, Dwyer 5603 (PMA). Chilibre, Dwyer 11978 (MO). Diete pus 131 (MO, PMA). Without other Locality, Hayes 455, 767 (both NY). El Palorin, Piao hia 255 (NY, US). San Jose Island, Johnston 832, 1201 (both GH, US). Icaco Island, 2 NO (GH, MO, ke Alhajuela, Killip 3226 (US). SE slope of Cerro Campana, Lewis et al. 3153 (MO). Cerro Cam , McDan 8121 (DUKE, FSU, MO). Cerro Penon, 3 km S of Alcalde Diaz, fe 8894 (MO). Matías ener bet E Pittier 6871 (NY, US). Between Río Pacora and Chepo, Porter et al. 5144 (MO). Old Panama, ilie 1933 (NY). Vista del Mar, Porterfield (NY). Arraijan, Porterfield (NY). Las Cumbres, Puga 42 eats 1 mi W of Bejuco, Tyson & Blum 2543 (FSU, MO, SCZ). Bella Vista, Standley 25405 (US). E of Rio Tocumen, Standley 26579, 29477 (both US). Corozal Road near Panama, Standley 26787, 26865 (both US). Toboga Island, . 27110 (US). Río Tapia, Standley 28066 (US). 2 m entrada camino Cerro Azul, Taylor 5, 78 (both PMA). vERAGUAs: 2 km NW of Atalaya, Nee 8251 (MO). 1 mi W of Santiago, Tyson 5171 (FSU, SCZ). Puerto Mutis 12 mi S of Santiago, Tyson 6001 (DUKE, FSU, MO). 2. Aeschynomene brasiliana (Poir.) DC., Prodr. 2: 322. 1825. type: Brazil, Rio de Janeiro, Commerson (P, holotype, not seen; F, fragment) Prostrate or decumbent /ierb, stems to ca. 1 m long, glandular-hispidulous, crisp-pubescent. Leaves mostly 8-12 foliolate, less than 3 cm long; leaflets usually obovate, obtuse to rounded at each end, ciliate, mucronulate, sparsely pubescent, ca. 7 mm long, ca. 4 mm wide; stipules ovate, acuminate, 3-4 mm long, ca. 1- mm wide, glandular-hispidulous, ciliolate. Inflorescence racemose, axillary, 3-8 flowered; peduncles and pedicels hispid; bracts and bracteoles ovate, 1.0-2.5 mm long, 1.0-1.5 mm wide, hispidulous, ciliate. Flowers yellow, 5-8 mm long; calyx 1.5-3.0 mm long, ciliate, hispidulous; standard ca. 6 mm long, the claw ca. 1 mm long, the blade suborbiculate, ca. 5 mm in diameter, pubescent on the outer face, retuse, wings and keel equal to the standard, ca. 2 mm wide; stamens ca. 6 mm long. Fruit 2-3 articulate, the stipe 3-4 mm long, hispidulous with trichomes ca. 0.5 mm long, the articles 2-3 mm in diameter, moderately crisp-pubescent; seeds ca. 2 mm long, ca. 1.5 mm wide, dark brown. Chromosomes: n = 10 This species is widespread throughout tropical America, but is represented in Panama by only a few collections. This species is distinguished from the super- ficially similar Aeschynomene histrix var. incana by the former's more prostrate 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 543 habit, glandular- E stems, and fruits with longer stipes. It is known lo- cally as ‘‘pega-peg CANAL ZONE: Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, Piper 5189 (F, US). Ancón Hill, Standley 25153 (MO, US). Sosa Hill, Balboa, Standley 25275 (US). Balboa, Standley 25586 (US). Cerro Gordo, near Culebra, Standley 26042 (US). Sosa Hill, Balboa, Standley 26442 (US). Corozal, Standley 27369 (US). Along old Las Cruces Trail, between Ft. Clayton and Corozal, Standley 29111 (US). Balboa, Standley 29295 (US). Summit, Standley 30069 (US). PANAMÁ: Bald Hill, San Jose Island, Johnston 880 (GH). Agricultural Experiment Station of Matías Hernández, Pittier 6908 (US). Corozal Road near Panama, Standley 26806 (US). Juan Franco race track near Panama, Standley 27724 (US). Taboga Island, Standley 28022 (US). Tumba Muerto Road near Panamá, Standley 29747 (US). Between Las Sabanas and Matías Hernández, Standley 31866 (US). Between Matías Hernández and Juan Díaz, Standley 32083 (US). Río Mar, Tyson et al. 2296 (FSU, PMA). 3. Aeschynomene ciliata Vog., Linnaea 12: 84. 1838. svNTYPES: Brazil, Para: Belem, Sieber 13736 (B, presumably destroyed; photo US, not seen). Brazil, between Rio de Janeiro and Cabo Frio, Sello (not located, possibly B or BM). A. hispida sensu Standley, Flora of Panama Canal Zone, Contr. U.S. Natl. Herb. 27: 210. 1928. Herb 1.0-2.5 m tall; stems hispid with yellow glandular trichomes. Leaves 10-15 cm long, 30—40 foliolate, petiole and rachis hispid; leaflets oblong with rounded to retuse apices, cordate and slightly asymmetrical basally, 10—15(—30) mm long, 3-5(-8) mm wide, glabrous, costae central, serrulate-ciliate, micro- punctate adaxially; stipules appendiculate below the point of attachment, upper portion acute, lower rounded, 1—2 cm long, serrulate-ciliate, caducous. /nflores- cence racemose, axillary, 5-many flowered; peduncles and pedicels hispid; bracts ovate-subcordate, 4-6 mm long, 2-3 mm wide; bracteoles lanceolate-ovate, acute, ciliate, 3-4 mm long, 1-2 mm wide. Flowers yellow, 8-10 mm long; calyx bila- biate, the upper lip trifid, the lower lip bifid, 5-7 mm long; the standard orbiculate, 5-8 mm in diameter, the claw ca. 2 mm long, serrulate-ciliate, emarginate, wings ca. 8 mm long, the claw 3 mm long, the blade ca. 5 mm long, 4 mm wide, ciliate, keel petals entire, the claws ca. 1.5 mm long, the blades 5-8 mm long, ca. 3 mm wide; stamens ca. 8 mm long. Fruit usually 8—10 articulate, stipe 5-10 mm long, hispid, the margins entire, articles ca. 4 mm long, 6 mm wide; seeds 3-4 mm long, ca. 2 mm wide, dark brown. This species is distributed from southern Mexico to Ecuador, Amazonian Brazil, and Jamaica. In Panama it is known from tropical moist forest in the Canal Zone and the immediate vicinity, usually growing near water. It appears to flower and fruit principally from November to January. NAL ZONE: Barro Colorado Island, Bailey & Bailey 763 (F). "UE Cowell 173 (NY). Chagres, Fendler 99 (MO, US). Barro Colorado Island, Foster 1403 (MO, PMA, US). Without exact locality, Hayes 184? (NY). Between France sige nd Catival, deri 30270 (US). CHIRIQUÍ: Quebrada del Medio, 2 km N of n ye es, Busey 753 Pags : Portobelo, D'Arcy & D'Arcy 6691 (MO). PANAMÁ: Toc rt, Dwyer 5194 (MO, P MA). Mati Hernandez, Pittier 6946 (GH, NY, US). Juan E Ca. 30516 (US). Capitana, Taylor 28 (PMA). 4. Aeschynomene elegans Schlecht. & Cham., Linnaea 5: 583. 1830. TYPE: Vera Cruz, Hacienda de la Laguna, Jalapa, Mexico, Schiede & Deppe (F, GH). Herb, stems to 0.5 m, hispid with tuberculate-based, glandular trichomes, to 1 mm long, sericeous with weak, appressed trichomes, to 0.25 mm long. Leaves 544 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 8-12 foliolate, ca. 2 cm long; leaflets usually obovate, obtuse to truncate, mu- cronulate, oblique, upper surface glabrous, reticulate beneath, costae villous with appressed trichomes, ciliolate, ca. 5 mm long, ca. 2.5 mm wide; petiole hispid; rachises villous. Inflorescence racemose, axillary, 2-6 flowered, longer than the subtending leaves, pedicels and peduncles with indument like the stems; bracts and bracteoles ovate, acute, ca. 1 mm long, less than 1 mm wide, ciliate. Flowers yellow, ca. 5 mm long; calyx ca. 1 mm long, ciliolate, hispidulous; standard ca. 4 mm long, the claw ca. 1 mm long, the blade elliptic-orbiculate, entire, wings and keel equal to standard, 1.5-2.0 mm wide; stamens 4-5 mm long. Fruit mo- niliform, curved or straight, (3-)4—6 articulate, the stipe 7-10 mm long, hispid, the articles ca. 2.5 mm long, ca. 2 mm wide, crisp-pubescent, often slightly gla- brate; seeds ca. 2 mm long, ca. 1 mm wide, dark brown. Chromosomes: n = 10. This taxon is widespread in tropical America, but apparently rare in Panama, only being known from El Boquete, Chiriquí at ca. 1,500 m. It is called ‘‘dor- midea" in Panama, and ‘‘pega-pega” in Costa Rica and Colombia. CHIRIQUI: El Boquete, Maurice 726 (US). Alta Boquete, Terry 1263 (F, GH, MO, US). 5. Aeschynomene histrix Poir., Lam. Encycl. Méth. Bot. Supp. 4: 77. 1816. 5a. A. histrix var. incana (Vog.) Benth., in Mart. Fl. Bras. 15(1): 69. 1859. Aeschynomene incana Vog., Linnaea 12: 90. 1838; non G. F. W. Mey. ex DC., 1825. TYPE: Brazil, Sellow (F, fragment). Much branched herb, stems to 0.5 m, prostrate to suberect, canescent. Leaves 8-18-foliolate, to 3 cm long; leaflets oblong-elliptic, rarely obovate, obtuse to rounded, apiculate, entire, ciliate, oblique basally, pubescent, 4-8 mm long, 1.5- 4.0 mm wide; stipules lanceolate, acuminate 4-10 mm long, 1-3 mm wide, hispid, serrulate-ciliate. Inflorescence racemose, axillary, 4-15 flowered: peduncles and pedicels hispid; bracts and bracteoles ovate, acute, 1-2 mm long, 1 mm or less wide, hispid, ciliate. Flowers 6-7 mm long; calyx 2-3 mm long; standard ca. 5 mm long, the claw ca. 1 mm long, blade suborbiculate, 4—6 mm in diameter, the wings and keel about as long as the standard, 1-2 mm wide; stamens 4-5 mm long. Fruit 2-articulate, the stipe 1-2 mm long, hispid with trichomes 2-4 mm long, mostly at the basal article, the articles 2.0-2.5 mm in diameter, glabrous to puberulent; seeds 1.5-2.0 mm long, 1.0-1.5 mm wide, black. Aeschynomene histrix var. incana is readily distinguished from the typical form by its more robust habit and larger leaf size. Fruit and flower characteristics are essentially the same for the 2 varieties and there appears to be intergradation; therefore, only varietal status seems justified. Variety incana is found from Mex- ico to South America, whereas variety histrix is primarily South American in distribution. In Panama it is found in wet sites below 100 m in the provinces bordering the Gulf of Panama from Los Santos to Panama. CANAL ZONE: Ancón Hill, Greenman 5127 (MO). S side of Summit Gardens, Nee 9549 (MO). COCLE: Río Hato airstrip, Burch et al. 1150 (GH, MO, US). Aguadulce, Pittier 4863 (GH, NY, US). Penonomé, Williams 102 (NY, US). Los SANTOS: Chitre/Las Tablas, Burt & Rattray 55A (MO). PANAMÁ: San Carlos, Río Hato, Burt & Rattray 46 (MO). Hills NE of Hacienda La Jo oya, Dodge et al. 16887 (MO). Savannas near Chepo, Duke 6040 (MO). Without exact locality, Seemann 216? (NY). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 545 6. Aeschynomene pratensis Small, Bull. New York Bot. Gard. 3: 423. 1905.—Fic. 3F 6a. Aeschynomene pratensis var. caribaea Rudd, Contr. U.S. Natl. Herb. 32: 47. 1955. TYPE: Cuba, Isla de Pinos, near Nueva Gerona, Curtiss 300 (F, GH, isotypes). Herbs; stems 1-2 m long, becoming suffruticose, drying dark, glabrous or sparsely hispid. Leaves 4-7 cm long, commonly 20-25 foliolate; leaflets oblong, obtuse to subacute, obliquely rounded basally, 5-10 mm long, 1.5-2.5 mm wide, micropunctate adaxially, the costae central, petioles and rachises glabrous or with few glandular trichomes; stipules entire, acuminate above, truncate below, erose peltate, appendiculate below the point of attachment, caducous. Inflores- cence racemose, few flowered, axillary; peduncles and pedicels glabrous to his- pid; bracts stipuliform, 4-5 mm long, 1-2 mm wide, margin entire, hyaline, some- times ciliate; bracteoles ovate, subacute, 3-4 mm long, 1.5-2.0 mm wide, hyaline margined. Flowers yellow, 7-10 mm long; calyx bilabiate, 4-5 mm long, vexillary lip subentire to 2-denticulate, carinal lip 3-denticulate; standard suborbiculate, 7— 10 mm long, 5-9 mm wide, the claw 2-3 mm long, wings 6-10 mm long, 3-4 mm wide, the claw 1 mm long, keel falcate, 6-9 mm long, ca. 3 mm wide, the claws ca. 1 mm long; stamens 7-10 mm long. Fruit 5-9 articulate, stipe 8-10 mm long, glabrous, articles 5-6 mm long, 4-5 mm wide, glabrous, reticulate veined, ver- rucose at maturity, basal article separated from stipe by a suture, both margins crenate, or dorsally subentire; seeds 4-5 mm long, 2.5-3.0 mm wide, dark brown. This variety is found chiefly in the Caribbean area and southward to central South America, and it has been introduced into the Old World. In Panama it has been collected from wet areas from the provinces around the Gulf of Panama. Variety caribaea is distinguished from variety pratensis by the former's smaller flowers and fruits, and also by the latter's restricted distribution in the everglades of Florida. Aeschynomene pratensis is distinguished from A. sensitiva var. sen- sitiva, its closest relative, by its slightly larger flowers and the presence of a suture between the stipe and the first article. cocLÉ: Penonomé, Ebinger 1016 (MO, SCZ, US); Williams 130 (NY). Between Las Margaritas and El Valle, Pie et al. 1768 (A, MO, NY). HERRERA: Ocú, Ebinger 1049 (MO, US). PANAMÁ: Road between Panama and Chepo, Dodge et al. (GH, MO). Without exact locality, Seemann 203? (GH). Nuevo San Ple Sidley 30753 (MO). 7. Aeschynomene rudis Benth., Pl. Hartw. 116. 1843. TYPE: Ecuador, Guayas, Guayaquil, Hartweg 649 (K, not seen). Shrubby herb, 1-2 m tall, sparsely hispid. Leaves 4—10 cm long, 30—40 fo- liolate, petiole and rachis hispidulous; leaflets oblong, rounded to truncate; slight- ly asymmetrical basally, 8-10 mm long, 2-3 mm wide, glabrous, the costae cen- tral, entire, micropunctate adaxially; stipules 7-15 mm long, 2-3 mm wide, the upper portion acute, the lower 2-3 mm long, rounded, entire. Inflorescence ra- cemose, few flowered, axillary; peduncles and pedicels hispidulous; bracts sub- ovate, acute, 3-4 mm long, 1.5-2.0 mm wide, ciliate; bracteoles ovate-oblong, acute, 2-3 mm long, ca. 1 mm wide, ciliate. Flowers yellow, (8-)10—15 mm long; 546 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 calyx 5-8 mm long; the standard ca. 14 mm long, the claw 2 mm long, the blade orbiculate, ca. 12 mm in diameter, retuse; the wings ca. 10 mm long, the claw ca. 1 mm long, the blade ca. 9 mm long, 5-6 mm wide; the keel petals ca. 10 mm long, 3 mm wide; stamens ca. 12 mm long. Fruit 6-10 articulate, the upper edge essentially entire, the lower edge crenate to subentire, the stipe glabrous, 3-6 mm long, the articles 4-6 mm in diameter, moderately hispid, usually muricate or verrucose at the center; seeds ca. 3 mm long, 2 mm wide, black. Aeschynomene rudis is principally found in moist or wet places in tropical and warm temperate America, ranging from the United States to Argentina. It is apparently rare in Panama, with no collections known from the last 25 years. It might be confused with A. ciliata from which it is distinguished by several char- acters as indicated in the key to species. PANAMA: NE of Hacienda la Joya, Dodge et al. 16913 (MO). oo . Aeschynomene sensitiva Swartz,'! Prodr. Veg. Ind. Occ. 107. 1788. TvPE: based upon Plumier's pl. 149, fig. 2, Icones 1693 (MO).—Fic. 3A-E. Herbs to 4 m tall; stems glabrous to glabrate below, fruiting branches glabrous to moderately hispidulous with glandular trichomes. Leaves 2-10 cm long, 10—40 foliolate, petiole and rachis hispidulous; leaflets oblong, obtuse to truncate, 4-15 mm long, 1.5-3.0 mm wide, the costae central, the nerves inconspicuous, mi- nutely punctate adaxially; stipules appendiculate below the point of attachment, caducous, 5-20 mm long, 1.5-5.0 mm wide, subentire, acute to acuminate above, truncate below, erose. Inflorescence racemose, axillary, 4-8 flowered; peduncles and pedicels subglabrous to densely hispidulous; bracts stipuliform, entire to ciliate-laciniate; bracteoles ovate, subacute, 1.5-5.0 mm long, 1-2 mm wide, en- tire to ciliate. Flowers yellow, 5-6 mm long; calyx 2-4 mm long, usually ciliate, vexillar lobe emarginate or subdenticulate, the cardinal lobe obscurely 3-dentate, standard 6-8 mm long, the claw 1.5-2.0 mm long, the blade suborbiculate, 4.5— 6.0 mm wide, ciliate, sometimes retuse, wings and keel subequal to the standard, the claws 0.5-1.0 mm long, blades 3.5-7.0 mm long, 2.5-3.5 mm wide; stamens 5-7 mm long. Fruit a loment, 5-8 (rarely —10) articulate, stipe 4-8 mm long, glabrous, articles 5-7 mm long, 4-7 mm wide, glabrous to sparsely hispid, smooth to verrucose, the upper margin of loment entire, the lower crenate to subentire; seeds 3-4 mm long, 2.5-3.0 mm wide, brown. Chromosomes: n = 10. Aeschynomene sensitiva is found primarily in wet sites at low elevations from the Antilles, southern Mexico, Central and South America, and has been intro- duced into the Old World tropics. It is distributed throughout Panama and is distinguished from its closest relative, A. pratensis var. caribaea by its smaller flowers and the absence of a suture between the stipe and the first article of the fruit. This species was not typified in the original protologue, but the Plumier illus- tration is presumably an authentic representation of the species and was proposed as a lectotype by Rudd (1955). !! For a list of synonyms see Rudd (1955). Only the listed names have been used for Panamanian material. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 547 SSH ZA P < SSSS LEE LL. FIGURE 3. Aeschynomene—A-E. Aeschynomene sensitiva Sw.—A. Habit (x1).—B. Flower (x314).—C. Corolla —C!. Standard (x 3).—C2. Wing petal (x3).—C?. Keel petal (x3).—D. Stamens x4).—E. Pistil (x4). [After Dwyer 2867.]—F. Aeschynomene pratensis var. caribaea Rudd. Fruit (x2). [After Woodson et al. 1768.] BOCAS DEL TORO: Chiriquicito to 5 mi S along Río Guarumo, Lewis et al. 2091 (GH, MO, US). CANAL ZONE: Gamboa, Allen 1976 (GH, MO, US). Frijoles, Baltimer 1565, 1567 (both MO). Juan Mina, Bartlett & Lasser 16766 (GH, MO). Gamboa, Kennedy et al. 2324 (MO). Frijoles, Piper 5182 (US). Between Corozal and Ancón, Pittier 2647 (GH, NY, US). BARRO COLORADO ISLAND: Bailey & Bailey 300 (F); Bangham 505 (A, P); Croat 4199 (MO, SCZ), 13232 (MO); Kenoyer 386 (US); 548 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Shattuck 624 (F, MO, po bd 256 (F); Wentmore & Abbe 215 (A, F, GH, MO); Woodworth & Vestal 401 (A, F, MO). cociÉ: Ca. 2.5 km S of Antón on old road to Puerto Obaldia, Lasseigne 4296 (MO). Aguadulce, in savannas near sea level, Pittier 4918 (US). Between Las Margaritas and El Valle, Woodson et al. 1768 (US). COLON: Miguel de la Borda, Croat 10019 (MO, PMA). Mouth of Rio Piedras, Lewis et al. 3180 (MO). Penonomé and vicinity, 50-100 ft, Williams 130 (US). DARIEN: Without other locality, Macbride 2706 (F, MO, US). HERRERA: Ocü, Ebinger 1049 (SCZ). PANAMA: ías dley 31964 (US). Las Sabanas, Standley 40770 (US). Capitana, Taylor 26 (PMA). 9. Aeschynomene villosa Poir., Lam. Encycl. Méth. Bot. Suppl. 4: 76. 1816. TYPE: Puerto Rico, Ledru (P, not seen). Herbs; stems usually decumbent, to 2-6 dm long, hispid. Leaves 2-7 cm long, 20-50 foliolate; leaflets linear oblong, subfalcate, mostly 3-costate, 3-6 mm long, 0.5-1.5 mm wide; stipules appendiculate below point of attachment, striate, subglabrous, (5—)10-15 mm long, 1.0-1.5 mm wide, usually somewhat hispid at the point of attachment, margins ciliate, attenuate at each end. Inflorescence often paniculate, few flowered, peduncles and pedicels hispid, bracts cordate, acuminate, 1.5-6.0 mm long, 1-2 mm wide, ciliate; bracteoles ovate-lanceolate, acute to acuminate, 1-4 mm long, 0.5-1.0 mm wide, ciliate. Flowers yellow, 3— 5(-7) mm long; calyx 2-4 mm long, hispid; standard 5-7 mm long, claw 1-2 mm long, blade suborbiculate, 4-5 mm wide, entire, wings 5-7 mm long, 1-2 mm wide, keel 4-5 mm long, claws 1.0-1.5 mm long; stamens 4-5 mm long. Fruit 3— 5(-7) articulate, articles 2.5-3.0(—4.0) mm in diameter, villous-hispid, with tri- chomes tuberculate based; seeds 2.0-2.5 mm long, 1.5-2.0 mm wide, blackish. Aeschynomene villosa is distributed from southwestern United States, through Central America into northern South America. It appears to be rare in Panama, having only been collected from two localities. As with other species in this genus, it is called ‘‘pega-pega” locally. AL ZONE: Madden Forest Road at entrance to Boy Scout road, Croat 12907 (MO). Sosa Hill, Balboa, Standley 25267 (US). Old Las Cruces Trail between Fort Clayton and Corozal, Standley 29163 (US). PANAMÁ: Abalaba, Killip 3367 (US). Near Matías Hernández, Standley 28893 (US). Río Tocumen, Standley 29399 (US). vERAGUAS: Divisa/Santiago, Burt & Rattray 82 (MO, SCZ). 4. ALYSICARPUS Michael O. Dillon? Alysicarpus Desvaux, Journ. Bot. (Desvaux) sér. 2. 1: 120. 1813. TYPE: A. bu- pleurifolius (L.) DC. — Hedysarum bupleurifolium L., type cons. Herbs; stems ascending to procumbent, terete, striate. Leaves alternate, 1-foliolate (rarely 3-foliolate); leaflets linear-lanceolate to oval, reticulate-venose; stipules scarious, acuminate, free or connate; bistipellate. Inflorescence race- '2 Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 549 mose, terminal or rarely axillary, bracts scarious, deciduous, pedicels paired, short. Flowers small; calyx 5-lobed, deeply divided, lobes unequal, the upper 2 connate nearly to the apex, glumaceous; corolla reddish purple, or rarely orange; the standard obovate to orbicular, clawed; the wings obliquely oblong; the keel slightly incurved, obtuse; stamens 10, diadelphous, upper connate only basally; anthers uniform; ovary sessile or shortly stipitate; ovules numerous; style fili- form, incurved at the apex, stigma broadly capitate, terminal. Fruit a loment, terete to compressed, submoniliform, indehiscent; seeds suborbicular or globose, smooth, lustrous, estrophiolate. Chromosomes: n = 8, 10. Alysicarpus is a genus of about 30 species, primarily of the Old World tropics, of which we have one, A. vaginalis (L.) DC. The genus appears to have no economic importance in the New World except as a noxious weed. However, in Africa Dalziel (1937) reports that various species including A. vaginalis are used locally as ‘‘a fodder for all kinds of domestic stock, used fresh, but preferably cut after fruiting and stored as hay; regarded as an excellent fodder for horses, but it is said to cause mucous diarrhoea if given in excess in the young and immature state during the rains.’ 1. Alysicarpus vaginalis (L.) DC., Prodr. 2: 353. 1825. non Hochst. ex Baker (1871); non Wall. (1831)—Fic. 4. Hedysarum vaginale L., Sp. Pl. 746. 1753. TYPE: Ceylon, Hermann (?BM, not seen). Herbs, suberect to prostrate, to 0.5 m tall; stems straw colored to black, glabrescent. Leaves unifoliolate, elliptic-orbicular, to 2.5 cm long, ca. 1.5 cm wide, apex rounded, apiculate, obtuse, truncate or subcordate basally, drying green, glabrous above, strigose-villose beneath; stipules erect, often amplexicaul, subulate, to 1.5 cm long, ca. 2 mm wide basally, acute, scarious, ciliolate. /nflo- rescence racemose, terminal, to 10 cm long. Flowers purple, ca. 5 mm long; the calyx deeply 5-lobed, the lobes unequal, acuminate, striate, puberulent; the stan- dard obovate, ca. 5 mm long; the wings oblong, ca. 4 mm long, the claw ca. 1.5 mm long, the keel connivent, oblong, ca. 4 mm long; stamens diadelphous, the staminal sheath terete, to 7 mm long, ca. 0.6 mm wide, the filaments alternating in length, the longer filament to 0.6 mm long, the anthers rotund, ca. 0.3 mm long; ovary linear, ca. 3 mm long, ca. 0.5 mm wide, appressed villosulose, the style compressed, geniculate, swollen, the stigma subdiscoidal, truncate. Fruits strongly ascending, terete, drying black, puberulent, ca. 6-articulate, articles ca. 2 mm long; seeds ca. 2 mm long, suborbicular, lustrous, reddish brown. This species is widespread in Panama and easily distinguished by its elliptic- orbicular, unifoliolate leaves and erect, terete loments. BOCAS DEL TORO: Almi ei eis 403 (GH). Milla 5, Croat & Porter 16460 (MO). Bocas, Lazor et al. 2376 (FSU). CANAL E: Gamboa, Busey 842 (MO). Ft. Kobbe, Duke 3914 (MO). Toro Point, Ft. Sherman, Duke 4336 TMO). Sosa Hill, Duke 4655 (GH, MO). Ft. Clayton, Dwyer 1001 (FSU, MO, SCZ). Curundu, Dwyer 1764 (MO). Frijoles, Ebinger 309 (MO, US). Madden Dam, Maurice 823 (US). Miraflores Lake, Tyson 1342 (FSU, MO, SCZ). Ft. Amador, Tyson 2014 (MO, SCZ). Curundu, Tyson 6631 (MO). Miraflores Lake, White 263 (MO, US). cHIRIQUí: David airport, Lewis et al. 739 (MO, NY). Puerto Armuelles, Liesner 141 (MO, NY). cocLÉ: NE of Río Hato, 550 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 os ESR AS W SA Y N coaer MY Q N Wye 7 {} , o NA J We x Z S Ë Y / Hh (4 ) pu. 4 X 9 A, V S 7 E 4. Alysicarpus vaginalis (L.) DC.—A. Habit (x?/s). [After White 263.]—B. diss er (x6). ae, "Con olla.—C'. Standard (x3?/;).—C?. Wing petal (x3/5).—C?. Keel petal (x33/5).— mens —E. Pistil (x6). [After Duke 3914.]—F. Fruit (x 14/s).—G. Sections of fruit “mm ies (x 33/s). "Afer White 263. Lasseigne 4293 (MO). COLON: Río Pie pies x et al. sa (MO, SCZ). Colon, Correa 1216 (MO). Portobelo to Pilon, Nee & Mori 3653 (MO, WIS). HERRERA: Los Pozos, Burt & Koster 94 (MO). PANAMA: Rio Pacora and Chepo, Duke 22 (MO). “wasata Duke 12416 (MO, NY); Ebinger 496 (MO, US). San Jose Island, Jona 858 (GH, US). Panama City, Mendez 112 (US). Between Río 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 551 Pacora and Chepo, Porter et al. 5102, 5134 (both MO, SCZ). Riomar, Tyson et al. 2319 (FSU, SCZ). SAN BLAS: Point opposite Ilsa Mosquito, Duke 8993 (MO, NY). vERAGUAS: La Mata, Koster 118 (MO). 5. ATELEIA Michael O. Dillon? Ateleia (DC.) Benth., Comm. Legum. Gen. 37, 37. Jun. 1837 (preprint); Ann. Wiener Mus. Naturgesch. 2: 101. 1838. LECTOTYPE: Ateleia pterocarpa Moc. & Sesse ex D. Dietr. (Pittier, 1918). Pterocarpus section a DC., Prodr. 2: 419. 1825; Mém. Légum. 393. 1826. Ateleia Ses. & Moc. ex DC., Prodr. 2: 419. 1825; Mém. Légum. 394. 1826, nom. in syn. Unarmed trees or shrubs. Leaves imparipinnate; leaflets alternate to subop- posite, subcoriaceous; stipules minute and caducous or none, reduced to tufts of hairs; stipels none. Inflorescences racemose, sometimes paniculate, terminal or axillary, 5-20 cm long; bracts small, deltoid or linear, persistent or caducous; bracteoles none. Flowers 10-several hundred, 5-14 mm long; calyx regular, cam- panulate, valvate or subimbricate in bud, truncate or shortly dentate, 5-lobed, the teeth ca. 0.5 mm long; petal 1, white or yellowish, clawed, cucullate, some- times expanded, spatulate at anthesis, glabrous or pubescent, the margin erose or sinuate; stamens free, the anthers uniform, ellipsoid, ca. 1 mm long, dorsifixed; ovary 1-2 ovulate, the stigma subsessile, ovate, obliquely terminal, inflexed. Fruit indehiscent, samaroid, semiorbicular, a narrow wing along the upper suture, 2 valved, usually 1 seeded, compressed stipitate; seeds reniform, compressed, reddish brown to dark brown, the hilum lateral, orbicular or elliptic. Chromo- somes: n — 20. Ateleia is composed of about 17 species distributed throughout tropical and subtropical Central and South America. All the species are known from limited ranges with the exception of A. gummifera, which ranges throughout the West Indies and the eastern coast of Central America. Literature: Mohlenbrock, R. H. 1962. A revision of the leguminous genus Ateleia. Webbia 17: 153-186. Pittier, H. 1918. A new species of Ateleia from Colombia. Contr. U.S. Natl. Herb. 20: 113. 1. Ateleia gummifera (DC.) D. Dietr.,'* Syn. Pl. 4: 219. 1847.—FiG. 5. Mii gummifer Bert. ex D.C., Prodr. 2: 419. 1925; Mém. Légum. 395. 1826. TYPE: Santo ingo, Bertero (G- DC. 1 MO, isotype). Shrub or small tree to 7 m tall; branchlets thick, knotty, lenticellate, usually glabrous. Leaves alternate, 7—11(—13) foliolate, rachis puberulent, 5-10 cm long, 13 Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 14 For additional synonymy see Rudd (1972). 552 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 NINN WW) H ` n e E FiGURE 5. Ateleia gummifera (DC.) D. Dietr.—A. Habit ( x?/s). [After Allen 4083.]—B. Flower (x7!/5).—C. Stamens and petal (<7!/s).—D. Pistil (x7!/s). [After Schipp 705, Belize.]—E. Fruit (x1!/3).—F. Seed (x4^*/:). [After Allen 4083.] terete; leaflets subopposite, ovate to elliptic, obtuse, or rarely emarginate, cu- neate to obtuse, 3-7 cm long, 1.5-3.0 cm wide, the upper surface glabrate, the lower surface dull white, puberulent, reticulate; petiolules ca. 2 mm long, puber- ulent. Inflorescences racemose, densely flowered, 6-17 cm long; bracts deltoid, commonly 1 mm long. Flowers pedicellate, the pedicels 1-2 mm long, villous; calyx campanulate, 2-3 mm long, tomentulose; petal yellow, 5-6 mm long, 2-4 mm wide, cucullate, the claw ca. 3 mm long, the blade suborbicular, irregularly sinuate; stamens 10, unequal; ovary stipitate, compressed, villous at the base, 2 ovulate. Fruit glabrous, 2.0-3.5 cm long, 1.0-1.5 cm broad, the wing 1-2 mm wide along the upper margin, the stipe 0.5-1.3 cm long; seed 5-7 mm long, 3.0- 3.5 mm broad, ca. 2 mm thick, the hilum ca. 1 mm in diameter. Chromosomes: n — 20. This species is the most widespread element of the genus, occurring through- out the West Indies and from the Yucatan south to Panama, from sea level to 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 553 1,000 m. It is infrequently collected and appears rare in Panama. There is con- siderable variation in leaflet shape and degree of pubescence, but flower and fruit characteristics appear constant throughout its range. No local names or uses are known from Panama; however, it is called ‘‘balsamo hediondo” or ‘“‘cerezo” in Cuba and ‘‘tush che" in Belize. HERRERA: Ocu, Allen 4083 (MO). 6. BARBIERIA Peter S. White!’ Barbieria DC., Prodr. 2: 239. 1825. Mém. Légum. 241. pl. 39. 1825. TYPE: B. polyphylla DC., nomen illeg. (Galactia pinnata Pers., Barbieria pinnata (Pers.) Baillon). Barbiera Spreng., Gen. Pl., ed. 9. 2: 587. 1831. orth. mut. Woody vine or low shrub, branchlets hirsute, but pubescence sometimes becoming sparse on older stems. Leaves alternate, imparipinnate, many foliolate, the leaflets entire; stipels persistent, long subulate, stipules persistent, narrowly triangular, acuminate. Inflorescence of relatively few-flowered racemes, terminal and from upper axils, nodes 2-5 per raceme, 2-3 flowers per node; bracts narrow, acuminate, bracteoles 2 subtending the hypanthium, narrow, acuminate, flowers short pedicelled. Flowers large, red; hypanthium tubular, pubescent, persistent in fruit, the teeth 5, long, narrow, acuminate, nearly equal, stiff, at least when dry; petals red, long clawed, glabrous, the standard oblong, the wings oblong, adherent to and shorter then the keel; stamens diadelphous, the vexillary stamen free, the anthers uniform, the ovary sessile, the ovules numerous, the stigma small, terminal, the style elongate, barbate on upper margin. Fruits linear, straight, flattened, septate and transversely impressed outside between the seeds, pubescent, the two valves coiling in dehiscence; seeds oblong reniform. Barbieria is a monotypic genus of tropical America. 1. Barbieria pinnata (Pers.) Baill., Hist. Pl. 2: 263. 1870.—Fic. 6. Galactia pinnata Pers., Syn. Pl. 2: 302. 1807. type: West Indies, not seen Clitoria polyphylla Poir. in Lam., Encycl. Méth. Bot. Suppl. 2: 300. 1811. TYPE: Puerto Rico, not seen. Barbieria polyphylla pues Prodr. 2: 239. 1825. Mém. Legum. 242. pl. 39. B. maynensis Poepp. & Endlich., Nov. Gen. Sp. Pl. 3: 58, pl. 264. 1845. TYPE: E Poeppig 2367 (not seen, F frag.). Woody wine, twining, trailing, or suberect, branchlets hirsute. Leaves alter- nate, imparipinnate; leaflets (9—)15-21(—23), elliptic with base and apex rounded, the apex mucronate, 1.5-7.0 cm long, 0.6-2.0 cm wide, surface glabrous to pu- bescent with minute scattered hairs above, the midrib hairy above, densely hairy and lighter beneath; paired stipels to 4 mm long, narrow; stipules 6-10 mm long, narrow; petiolules 1-3 mm long; petioles 1-4 cm long, hirsute, grooved. /nflo- 15 Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. [VoL. 67 ANNALS OF THE MISSOURI BOTANICAL GARDEN URE 6. Barbieria pinnata (Pers.) Baill.—A. Habit (x 3⁄s). [After Skutch 2263.|—B. ayasa FIG (xX?/19).—C. Corolla.—C'!. Standard (x*/). —C?, r a A —C?. Keel petal (x3/5).—D. S ns “Sa —E. Anthers.— 2. Side w.—F. Pistil (x°/io). [After een op view.— 845.]—G. Fruit (x?/10). —H. Seed (x6). [After Klug 3095. Peru.] rescence racemes, occasionally branched, to 25 cm long; flowers 2-3 from paired stipulelike bracts and a third bract between these on the rachis; pedicels ca. 5 mm long; bracteoles ca. 1 cm long at the base of the hypanthium. Flowers large, to 5.5 cm long, red; hypanthium ca. 3 cm long, 6 mm broad, densely pubescent 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 555 and with scattered longer hairs, the 5 teeth ca. 1⁄2 as long as the tube, ca. 1 cm long, narrow lanceolate acuminate; petals glabrous, standard to 5.5 cm long, ca. 1.5 cm wide, wedge shaped at the base, clawed, wings and keel petals all ca. 4.5 cm long, abruptly contracted to a claw, keel oblong; staminal column ca. 5 cm long, open above, glabrous; ovary sessile, ca. 7 mm long, terete, silvery pilose, ovules several, the stigma capitate, the style ca. 4 cm long, heavily bearded on upper margin. Fruit linear, ca. 5 cm long, 0.6 cm wide, compressed, pubescent, septate inside and indented on the outside between the seeds, 2 valves coiling in dehiscence; seeds 4-9, small, dark brown or black. Barbieria pinnata, the only species in the genus, is found from Mexico to South America and in the West Indies. It has not yet been found in Panama, but is known from bordering countries and likely occurs there. Judging from the number of collections in major herbaria, Barbieria pinnata may be infrequent or rare throughout its range. The large red flowers make the plant showy when it is in bloom. Barbieria pinnata is distinctive for its large blood-red flowers, clawed petals, long tubular calyx with sharp, equal calyx teeth, its long and hairy style, its long (to 1 cm) and narrow stipules, and its bracts and bracteoles. 7. CAJANUS Muriel E. Poston!? Cajanus DC., Cat. Hort. Bot. Monspel. 85. 1813. nom. conserv. TYPE: Cajanus cajan (L.) Millsp. Cajan Adans. Fam. Pl. 2: 326. 1736. Based on Cytisus cajan L. = Cajanus cajan (L.) Millsp. Shrub or subshrub. Leaves trifoliolate, leaflets oblanceolate, puberulent; stip- ules lanceolate, caducous. Inflorescence an axillary or terminal panicle or ra- ceme, flowers 6-12 per rachis; bracts small, caducous. Flowers with calyx cam- panulate, 5-lobed, the upper lobe longer; corolla yellow with brown to maroon striation, the standard ovate, auriculate, the wing oblong, the keel falcate; sta- mens diadelphous, the vexillary stamen free, the anthers monomorphic; ovary sessile with 4-6 ovules, the style slender, glabrous, the stigma capitate. Fruit oblong, 2-valved with oblique constrictions between the seeds, yellowish green with brown mottling, puberulent; seeds 4-5, obovoid, hilum oblong, estrophio- late. Cajanus is an Old World genus of 1 or 2 species widely spread through Africa and Asia and has been introduced into the New World in cultivation. 1. Cajanus cajan (L.) Millsp. Publ. Field Columbian Mus., Bot. Ser. 2: 53. 900.—FiG. 7 Cytisus cajan L., Sp. Pl. 739. 1753. TYPE: India, Tab Linnaeus (LINN 912:4, not seen; photo, MO). Cajanus bicolor DC., Cat. Hort. Bot. Monspel. 85. 1813. TYPE: not seen Cajanus indicus Spreng. Syst. Veg. 3:248. 1826. TYPE: not see 16 Department of Biology, Howard University, Washington, D.C. 20059. 556 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 X. S S W Qi) SA} \ ay Sa p. N FIGURE 7. Cajanus cajan (L.) Millsp.—A. Habit (x!4).—B. Flower (x2).—C. Corolla.—C'. Standard (x 1).—C2. Wing petal (x1).—C?. Keel petal (x 1).—D. Stamens (x2).—E. Pistil ( x214). After Hunter & Allen 70.]—F. Fruit ( x 12).—G. Opened fruit with seeds (x'4). [After Greenman & Greenman 5194.] Shrub or subshrub; stems angular, striate, strigose. Leaves trifoliolate: leaflets oblanceolate lanceolate, 4.3-8.2 cm long, 1.2-3.2 cm wide, the terminal leaflet slightly longer than the lateral leaflets, apically acute, basally cuneate, the margins revolute, velutinous and gland dotted beneath, glabrous above; petiolules 0.4—1.0 cm long, the lateral leaflets subsessile; petioles 0.6-1.2 cm long, angled, puber- | ulent; stipules lanceolate, 2.0-5.0 mm long, strigose; stipels linear to subulate, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 557 ca. 1.0 mm long. Inflorescence with racemes 4.0—9.0 cm long, 5-6 (-12)-flowered; peduncles 2.0—4.0 cm long, strigose; pedicel 0.6-0.8 mm long, strigulose; bracts subtending the flowers ovate, 0.5-0.7 cm long, 0.3 cm wide, caducous. Flowers with the calyx 0.8-1.0 cm long, not exceeding the corolla, puberulent, the upper lobes of the calyx partly united; corolla yellow striped with maroon or brown, 1.0-1.6 cm long, the standard obovate, 1.3-1.5 cm long, 1.4-1.5 cm wide, auri- culate, the claw ca. 2.0 mm long, the wing oblong, 1.3-1.4 cm long, 0.5-0.7 cm wide, the claw ca. 4.0 mm long, the keel falcate, not auriculate, 1.2-1.5 cm long, 0.5-0.7 cm wide, the claw 0.5 cm long; stamens 1.3-1.4 cm long, vexillary stamen free; style 1.2-1.3 cm long, the stigma capitate. Fruits oblong, 5.5-7.0 cm long, 0.8-1.0 cm wide, green with brown mottling, puberulent, the beak 0.7-1.0 cm long, the compressions between the ovules oblique; seeds 4—5, ovoid, brown with white mottling, 5.0 mm long, 6.0 mm wide, the hilum linear. Cajanus cajan is cultivated for its edible seeds known locally as Guandu, Guandul or pigeon-pea. It is found in tropical moist forest in Panama or where cultivated. BOCAS DEL TORO: 15 mi S from mouth of Changuinola River, Lewis et al. 929 eye CANAL ZONE: Between Miraflores and Panama, Blum 2077 (MO). Without p locality, Crawford 438 (NY). Barro Colorado Island, Ene 4670 (SCZ), 7238 (DUKE, MO, NY, SCZ, US). Di ee ya her Dec. 7, 1962 (SCZ). Corozal, Greenman & Greenman 5194 (MO). CHIRIQUÍ: 2 km of Punta de Purical, Burica Peninsula, Busey 761 (MO). cocLe: El Valle de Anton, Hammel 1746 E MO); prs et al. 2589 (DUKE, MO). Penonomé, Williams 360 (NY). COLON: Without other locality, Wedel Feb. 23 1940 (MO). DARIEN: Upper Río Tuquesa, Le Clezio 109 (MO). PANAMA: Isla del Rey, Duke 9504 (DUKE, MO). Sabanas near Chepo, Hunter & Allen 70 (MO). SE slope of Cerro Jefe, Tyson 3551 (SCZ). Santa Clara, Tyson & Blum 1884 (FSU, MO, SCZ). Cerro Azul, Tyson et al. 6045 (FSU, SCZ). Cerro Campana, Wilbur 24371 (DUKE). Pacora, Williams 770 (NY). LOCALITY UNKNOWN: Tabernilla, Cowell 283 (NY). Aspinwall, Hayes 642 (NY). 8. CALOPOGONIUM W. G. D'Arcy” Calopogonium Desv., Ann. Sci. Nat. (Paris) ser. 1(9): 423. 1826. TYPE: C. mu- cunoides Desv. Erect or climbing perennial herbs, often robust. Leaves pinnate trifoliolate, stipels and stipules scale-like. Inflorescences axillary racemes or fascicles, the rachis, when present, nodose with flowers in small fascicles on the nodes; pedicels short or long, bracts and bracteoles small, sometimes caducous. Flowers some- times showy, the calyx campanulate tubular, 5-toothed, the upper pair fused partway up, forming a lip, the lower 3 lanceolate; corolla pink, white or violet, narrow, the standard obovate with inflexed basal auricles, the wings and keel narrow; stamens diadelphous, the vexillary stamen free, the other filaments united halfway or less, the anthers 8-9, all alike, ellipsoidal, glabrous, dorsifixed; ovary sessile, hirsute, ovoid, many-ovulate, the style slender, incurved, glabrate, es- pecially above, the stigma capitate globose or compressed. Legume linear oblong, compressed, 2-valved, demarcated between the seeds, elastically dehiscent; seeds several, ovoid, aril wanting. 17 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 558 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Calopogonium is a genus of 6-8 species mainly centered in South America, but extending into Central America and introduced and naturalized in the Old World tropics. There are reports that plants in this genus are toxic to livestock, and in some countries they have been used as insecticides. In Central America, the vines are sometimes used to scour pots. a. Leaflets glabrate above; inflorescences fasciculate, pedicels more than 4 mm long; corolla pilose outside; seeds greenish... . C. galactioides aa. Leaflets pubescent ELE inflorescence racemose, pedicels less than 4 mm long; corolla adir outside: seeds brownish. b. on becoming fine puberulent or glabrate; bracteoles ca. 2 mm long, not s Si the calyx tube, early caducous; pods more than 5 mm wide |... 2. Tr HE MES . C. coeruleum bb. Stems remaining long pilose velutinous; bracteoles 4-6 mm long, Me as long as the calyx lobes, mostly persistent in flower; pod less than 5 mm wide |... 1. Calopogonium coeruleum (Benth.) Sauv., Ann. Acad. Habana 5: 337. 1869.— Fic. 8 Stenolobium coeruleum Benth., Ann. Wiener Mus. Naturgesch. 2: 125. 1838. TYPE: not seen. Vine, sometimes becoming large, yellowish pubescent, stems slender, softly villous with stout hairs to 2 mm long, these soon deciduous leaving a dense soft pubescence of short hairs. Leaves pinnate trifoliolate, leaflets ovate or rhomboi- dal, mostly 6-9 cm long, 4-6 cm wide, the lateral pair oblique, obtuse at each end, mucronate, mostly 3 veined at the base, slightly discolorous, appressed fine pubescent on both sides but more so beneath; petiolules 2-4 mm long; rachis 1- 4 cm long; petiole mostly somewhat shorter than the terminal leaflet; stipels subulate, pilose, ca. 4 mm long; stipules caducous. Racemes elongate, sturdy but flexuous; peduncle mostly 1-5 cm long, puberulent; rachis to 20(—40) cm long, nodose, the nodes sometimes elaborated into short cymes; pedicels tomentose, ca. 2 mm long, inserted on the nodes, 2-5 at a time; bracts 1-3 mm long, pilose. Flowers bluish, the calyx tubular campanulate, ca. 4 mm long, the lobes subulate acuminate, pilose, about as long as the tube; standard glabrous, ca. 1 cm long, the wings narrowly spatulate oblong, about as long as the standard, the keel shorter; anthers oblong ovoid, 0.6 mm long; style ascending, pubescent to the tip. Legume linear oblong, 4-6 cm long, ca. 8 mm wide, impressed between the seeds laterally and slightly marginally, the beak small, curved, softly fine tomen- tose on the sides with longer hairs along the margins; seeds 4-8, squarish discoid, 4—5 mm across, shiny brown, the hilum oval, centered on one edge. BOCAS DEL TORO: Water Valley, Wedel 1814 (GH, MO, US). Old Bank Island, Mag 1981 (GH, ; - CANAL ZONE: Victoria Fill near Miraflores Locks, Allen 1709 (GH, MO US). Galen Pe Atlantic side, Blum & Dwyer 2151 (FSU, MO, SCZ). Pa ipie 2s 7152 (MO). Pipeline road 2/4 m from Gamboa Gate, Croat 9359 (NY). Near Paraiso, Croat 5 (F, MO, NY, SCZ). Summit pepe radio i Croat 14244 (MO). Albrook Tropic Test Site ae yer 7314 (MO). C-12 Road, Dwyer Balboa Heights, Greenman & Greenman 5053 (MO). Paraiso Station, Hayes 113, 118 (both NY). atún, Hayes 491 (NY). Barro Colorado Island, Hladick 136 (MO). Las Cruces Trail, 75 m, Hunter & Allen 680, (GH, MO), 742 (MO). ? Madden dam, 80 m Nee 7774 (MO). Valley of Masambí, road to Las Cascadas Plantation, 20-100 m, Pittier 2669 (GH, NY, US). Quebrada Melgada, Vo Mae 17490 (GH, MO). Barro Colorado Island, Woodworth ^ Vestal 443 (A, F, MO). cHiRIQUÍ: 20 km 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 559 FI E 8. m coeruleum (Benth.) Sauv.—A. Habit (x!2).—B. Flower (x214).—C. Corolla.—C!. Standard (x5).—C?. Wing petal (x2).—C?. Keel petal (x2).—D. Stamens (x3).—E. Pistil (x3). [After Wedel 1956.]—F. Fruit (x14). [After Allen 1709.] of Puerto Armuelles, Busey 408 (MO). Rabo de Puerco, 10.5 km W of Puerto Armuelles, Busey 520 (F, GH, MO, NY). Que Te pi cw gen beyond La Represa, 2 mi SW of Puerto Armuelles, 0-200 m, Croat 22066 (F, GH, Y). 4 mi from Paso Canoas on road to Canas Gordas, pose Da (F, GH, MO). Quebrada Lie dun 0-100 m, Croat 22533 (MO). Las Vueltas-Boca del Monte, 0- 560 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 100 m, Koster 111 (MO). Boquete, Lewis et al. 618 (NY). COCLE: Road to El Valle close to Pana- merican Highway, Burt & Rattray 34 (MO). ca. 2 km de el Copé en la Madera, Taylor 971 (PMA). up : Tyson 2704 (FSU, MO). 10 mi S of Ocü, Tyson et al. 2814 (MO, SCZ). Los SANTOS: 4 mi SE of Posos, Tyson 2704 (MO). PANAMÁ: Límite de Panamá-Zona del Canal, Carrasco 41 (F, MO, PMA). Campo Experimental de Monte Oscuro, 10 m, Correa 1920 (PMA). Cerro Jefe, 2700 ft, Dwyer 8533 (MO). Capira, Foster 2128 (F, MO, PMA, US). Sabanas near Chepo, Hunter & Allen 100 (GH, MO). Agricultural Station at Matías Hernández, Pittier 6956 (NY, US). La Capitana al lado del potrero, Taylor 21 (PMA). Carretera de la Urbanizacion Chanis, Vía Cincuentenario, Taylor 47 (PMA). vER- AGUAS: Divisa-Santiago, 0-100 m, Burt & Rattray 71 (MO). 1-2 mi above Santa Fé, Gentry 3054 (MO, PMA). Canazas, Tyson 3653 (MO). 8 mi S of Santiago, Tyson 60443 (FSU, MO, SCZ). 2. Calopogonium galactioides (H.B.K.) Hemsl., Biol. Centr. Amer., Bot. 1: 301. l Glycine? galactioides H.B.K., Nov. Gen. Sp. Pl. 6:427. 1823. TYPE: Venezuela, not see Stenolobium galactoides Benth.: Ann. Wiener Mus. Naturgesch. 2: 125. 1838. TYPE: not seen. Galactia belizensis Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 133. 1932 fide Standley & Steyer- mark 24(5): 172. 1946. TYPE: British Honduras, Bartlett 11449 (F, not see n). Slender climbing vine; stems villous with ascending yellowish hairs, glabres- cent. Leaves pinnate trifoliolate, the leaflets ovate or elliptical, mostly 2—4 cm long, 1.3-2 cm wide, obtuse at both ends, mucronate, the costa prominent above with ca. 4 arcuate veins on each side, glabrate above, appressed pilose beneath; petiolules 1-1.5 mm long, drying dark, pubescent; rachis to 9 mm long; petiole often longer than the terminal leaflet; stipels to 1 mm long pilose, caducous; stipules pilose. Inflorescences axillary fascicles of 2-5 flowers; pedicels ca. 5 mm long, puberulent; bracteoles 1-2 mm long, subulate, pubescent. Flowers white, pink or purplish, calyx ca. 5 mm long, villous hispid with coarse hairs, divided about halfway, the teeth acuminate subulate; corolla pilose outside, the standard emarginate, about twice as long as the calyx. Fruit linear oblong, 2-3 cm long, ca. 3 mm wide, nearly straight, compressed, falsely septate between the seeds, the beak 1-2 mm long, nearly straight; seeds plump, rectangular, dull greenish, 3 mm long, the hilum circular, located on one edge. Calopogonium galactioides is amply distinct in a number of ways: fasciculate inflorescences, pilose corolla, and glabrous leaflet uppersides as well as in overall appearance which is of a delicate, small leaved vine. It is known in Panama only from western Chiriquí. The species ranges from southern Mexico to northern South America. HIRIQUI: Cuesta de Piedras, 730 m, Burt & Koster 161 (MO, SCZ). San Bartolo Limite Asie Costa Rican border, 12 mi W of Puerto Armuelles, 400-500 m, Croat 22193 (F, GH, MO, NY). Volcán de Chiriquí, Boquete District, 7000 ft, Davidson 963 (GH, M ). 3. Calopogonium mucunoides Desvaux, Ann. Sci. Nat. (Paris) sér. 1. 9: 423. 1826. TYPE: ?Guyana, not seen. Stenolobium brachycarpum Benth. in Seem., Bot. Voy. Herald 109. 1838. type: Panama, Panama to antiago, Seemann (?K, not seen). Calopogonium orthocarpum Urb., Sy ymb. Ant 327. 1899. syNTYPES: um Rico, Krug 366, Sin- tenis 817, 1097 none seen, 3622 (MO), ze Stahl 976, none see 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 561 Slender, sometimes wiry vines, twining or trailing, stems pilose with spreading golden hairs ca. 1 mm long. Leaves pinnate trifoliolate, the leaflets chartaceous, ovate, mostly 2-6 cm long, mucronulate, obtuse at both ends, lateral leaflets sometimes slightly oblique, the major veins prominent beneath, appressed pilose, more so beneath; petiolules ca. 3 mm long, tomentose; rachis mostly 5-15 mm long; petioles mostly longer than the terminal leaflet; stipels ca. 4 mm long, pubescent; stipules ovoid, ca. 3 mm long, pilose. Racemes slender, peduncles short in flower, elongate, 3—12 cm long in fruit, the rachis not expanded, flowers in fascicles of 2—6, becoming distant in fruit, pedicels 1-2 mm long, appressed pubescent; bracts and bracteoles narrowly ovate, to linear, striate, long pilose, ca. 4mm long. Flowers blue or purple, inconspicuous or rarely showy, the calyx 4-7 mm long, deeply 4-fid, the lobes narrow, pilose, the sinuses somewhat ex- ceeded by the bracts, the upper pair fused basally, basally deltoid, apically sub- ulate; corolla with the standard 7-10 mm long, emarginate, slightly exceeding the wings, the keel shorter, curved; ovary villous, the style glabrous. Legume linear oblong, flat, mostly 2-3 cm long, ca. 4 mm wide, mostly straight, the beak 1-2 mm long, softly pilose with coarse reddish brown hairs, sometimes laterally im- pressed between the seeds; seeds 3-8 dark yellowish or reddish brown, com- pressed squarish, 2-3 mm long, the hilum circular, central on one edge. This species is distinct in its hairy pods. They are transversely impressed and falsely septate between the seeds, but the transverse ridges are mostly hidden by the pubescence. Although reported to be sometimes showy in nature, on the herbarium sheet the flowers are seldom well preserved and are inconspicuous. BOCAS DEL TORO: Railroad, Changuinola Valley, Dunlap 385 (F, US). CANAL ZONE: Chiva Chiva, tower site, Correa 490 (MO, PMA, SCZ). Barro Colorado Island, Croat 6925 (MO, PMA, SCZ), 9104 (MO, SCZ). Pipeline Road NW of Gamboa gate, Ea 12746 (MO, SCZ); D’Arcy 10615 (MO). Frijoles near railroad station, Croat 13220 (MO). Barro Colorado Island, Croat 6925 (SCZ), 9104 (SCZ), 13169A (MO). Corrosion lab, Miraflores La Dwyer 3037 (MO). Cocoli, Dwyer 7207 (GH, MO). Chagres, Fendler 79 (GH, MO). Outskirts of Ancón, Greenman & Greenman 5070 (MO). Las Cruces Trail, 75 m, Hunter & Allen 736 (GH, MO). La Boca, Mori & Kallunki 3677 (MO). Road C 29 6 km E of Gamboa, 190 m, Nee 9001 (MO, US). % mi W of Summit Gardens, Nee 9495 (MO). Panamerican Highway between Rodman Marine Base and Chorrera, Nowicke et al. 3587 (MO). Between Corozal and Ancón, 10-30 m, Pittier 2174 (NY, US). Balboa Station, Porterfield 1933 (NY). end of air strip, Albrook Air Force Base, Tyson & Lazor 6010 (FSU), CHIRIQUI: Concepción, Baltimer 1526 (MO). Auge pd 1 km W of dam at La Represa, Busey 486 (F, MO). Puerto Armuelles, Busey 507 (MO). Quebrada de Vuelta, Busey 622 (MO). San Bartolo Limité, 10— 11 mi W of Puerto Armuelles, a 22021 (MO). Puerto Armuelles in bananas, D'Arcy 10075 (MO). David airport, 25 ft, Lewis et al. 766 (GH, MO). Monte Verde, 2.5 km W of Puerto Armuelles, 80 m, Liesner 22 (MO, NY). COCLE: Río Hato ‘airstrip Burch et al. 1130 (GH, MO, NY). Foot of Cerro Pilón above El Valle de Antón, 2000 ft, Porter et al. 4596 (MO, NY, SCZ). CoLON: Santa Rita 4 km desviación de la Transistmica, 150 m, Gomez-Pompa et al. 3220 (MO). HERRERA: Las Minas, 340 m, ) L 93 ( mi S of Macaracas, Tyson et al. 3073 (GH, MO, SCZ). PANAMÁ: I mi E of Tocumen airport on side road off wajar zb: Highway, Blum & Tyson 1959 (FSU, MO, SCZ). Calzada Larga, Carrasco 45 (MO, PMA). Campo Experimental de Monte Oscuro 12 km de Capira, 10 m, Correa 1 1928 (PMA). Between Chepo and El Llano, Croat 14493 (MO). Between Rio Bayano Dam and Canitas, D’Arcy 9406 (MO). Hills between Capira and Potrero, 80-130 m, Dodge & "Hulier $602 (MO), 8608 (MO). Sabanas near Chepo, Hunter & Allen 77 (MO). SE slope of Cerro Campana, Lewis et al. 3102 (MO). Las Cumbres, Muñoz 18 (MO, PMA). 1 km S of Madden Dam, Nee 8902 (MO). Sabanas N of Panama City, Paul 534 (US). Bella Vista, Standley 25400 (US). Bella Vista, i prey eee (US). Las Sa- banas, Standley 25893 (US). La Garrapata, Taylor 14 (PMA). 1 mi W of Bejuco on Panamerican ighway, Tyson & Blum 2545 (FSU, MO, SCZ). VERAGUAS: | mi W of sco along old Inter- american Highway, Tyson 5182 (FSU, PMA). 562 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 9. CANAVALIA W. G. D’Arcy!® Canavalia DC.,!9 Prodr. 2: 403. 1825. TYPE: C. ensiformis (L.) DC. Nom. cons. contra Canavali Adans. Canavali Adans., Fam. Pl. 2: 325, 531. 1763. TYPE: Dolichos ensiformis L. = Canavalia ensiformis L.) DC. nomen rejic. contra Canavalia DC. Wenderothia Schlecht., Linnaea 12: 330. 1838. rype: W. discolor Schlecht. = Canavalia villosa Benth. Twining vines, prostrate, clambering or high climbing, rarely more or less erect subshrubs, mostly perennial, somewhat woody; stems often sturdy, mostly pubescent. Leaves alternate, pinnate trifoliolate, the leaflets mostly nearly sym- metrical, ovate or elliptical, sometimes broad, mucronulate, rarely emarginate, mostly pubescent on both sides, pinnately veined, the principal veins and costa slender and elevated beneath; petiolules 5-7 mm long, mostly pubescent with stiff ascending hairs; petioles equalling or shorter than the terminal leaflet: stipels minute and caducous; stipules caducous. Inflorescences axillary racemes, mostly 10-60 cm long, pendant or prostrate and ascending; peduncles with swollen nodes each bearing 2-6 pedicels; bracts caducous; bracteoles rotund to acute, caducous; pedicels mostly 1-3 mm long, the flowers mostly resupinate. Flowers showy, somewhat fragrant; the calyx tubular or campanulate, bilabiate, the upper lip sometimes forming a short beak, and when pressed appearing either entire or emarginate, the lower lip of 3 minute teeth, the lateral pair often rotund, the lowest tooth mostly pointed; corolla white, purplish, bluish, pink or red, some- times with white or yellow markings, the standard obovate, reflexed, emarginate (subgenus Wenderothia) or entire, the wings free, narrow, auricled, the keel ascending, united, auricled; stamens mostly united, the vexillary stamen some- times more or less free; stigma capitate, the ovary pubescent. Legume 6—40 cm long and 1.5-5 cm wide, oblong, often falcate when young, apically beaked, sometimes stipitate, coriaceous or becoming woody, mostly compressed, with prominent longitudinal ridges and 1 or 2 (3) ridges on the valves, dehiscent, sometimes explosively, the valves often twisting, pubescent or glabrescent; seeds numerous imbedded in a papery endocarp, sometimes buoyant, brown, mottled or white, mostly ellipsoidal and somewhat compressed, the hilum peripheral, short or long, 7-35 mm long. Canavalia is recognizable by its large, often woody oblong pods, and by the sturdy stems. The calyx is diagnostic with its often beaked upper lip and minute 3-toothed lower lip. The genus is pantropical in distribution with about 30 species in the New World and about 15 species in the Old World, mostly from the eastern Indian Ocean area and the western Pacific. A few species also occur in Hawaii. Only one species, C. maritima, is pantropical, a common element of beaches and coastlines, but at least two species are cultivated as experimental forage crops. Canavalia ensiformis was cultivated for food in pre-Colombian America. "5 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166 '? Other synonyms are given by Sauer (1964); only these names relate directly to Panamanian plants. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 563 Literature: Sauer, J. 1964. Revision of Canavalia. Brittonia 16: 106—181. a. Leaflets nearly circular, emarginate; terminal petiolule often more than 5 mm long; pedicels commonly 3 mm long; plants common on beaches and coastal rocks... 7. C. maritima aa. Leaflets narrower, pointed; terminal petiolule mostly less than 5, mm long; pedicels mostly less than 2.5 mm long; plants occurring behind beaches and inland. b. Calyx more than 15 mm long, lowest tooth mostly over 2.5 mm long; leaves mostly copiously pubescent, at least beneat . C. villosa bb. Calyx less than 15 mm long, lowest tooth less than 2.5 mm long; leaves often glabrate beneath. c. Lowest d tooth shorter than the lateral teeth. with 2 extra "sao pee n spaced well away from the sutural i pubescent overall at maturity, upper calyx lip shorter wo the tube, the lowest tooth less than 2 mm lon . C. bicarinata dd. Pod with one extra sutural ridge ae near the sutural ridge, pt ie at maturity; upper calyx lip as long as the tube, the lowest tooth over 2 mm lo 3. C. dictyota ong cc. p calyx tooth longer than the lateral teeth. eeds red or white; pedicels sometimes 2 mm long or more; cultivated species f. Seeds white; hilum less than 4 as long as the seed; the lowest calyx tooth ca. 2.5 mm long . . ensiformis ff. Seeds red or pink, hilum more than !⁄ as long as the seed; the lowest calyx tooth ca. 1.5 mm long ------ 6. C. gladiata ee. Seeds brown; pedicels 2 mm long or less; wild speci g. Pods with extra ridge(s) spaced well away pit ty the sutures; lowest calyx m less than 2 mm long. per two lI be evenly pubescent at m seeds DE C. bicarinata * € oxyphylla Pods with extra ridge near t 6 was the sutural ridges; lowest calyx tooth mostly more than 2 mm long. i. Upper calyx lip ae a distinct beak or umbo |... 5. C. glabra i. Upper calyx lip emarginate, not forming an umbo -------------- 2. C. brasiliensis a ae 1. Canavalia bicarinata Standley, Contr. U.S. Natl. Herb. 18: 106. 1916. TYPE: Panama, Pittier 2354 (US, holotype, not seen; GH, NY, isotypes). Wenderothia bicarinata (Standley) Piper, Contr. U.S. Natl. Herb. 20: 578. 1925. Trailing vine; stems sparingly pubescent with ascending weak, brownish or white hairs. Leaves pinnate trifoliolate, leaflets to 15 cm long, 9 cm wide, ovate, apically obtuse or short acuminate, basally rounded, chartaceous, drying slightly discolorous, sparingly pubescent to villose with short hairs on 1 or r both sides. Inflorescence axillary, 4-15 cm long, bracteoles rotund, ca. 1 mm long, pedicels wanting or to 1.5 mm long. Flowers red or rose; calyx ca. 12 mm long, tubular, pubescent with short, straight hairs, the upper lip short, apiculate, the lowest tooth ca. 1.5 mm long, pointed, slightly longer than the lateral teeth; standard 3 cm long. Legume mostly less than 15 cm long, woody, oblong with a short downcurved beak, sessile or short stipitate, ca. 2 cm wide, longitudinally ridged on the sutures and each valve with 2 additional ridges almost equidistant, pilose when young, maturing with fine even gray pubescence between the ridges; seeds ca. 9 mm long, ovoid or ellipsoidal, dark brown, mottled, the hilum ca. 7 mm long, ca. % the length of the seed. 564 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 This species is distinct in its pods which have 2 longitudinal ridges in addition to those over the sutures, thus giving an appearance of evenly spaced ridges. The pods are usually grayish pilose when young and fine pubescent overall when mature. The upper calyx lip is short in relation to the tube. Canavalia bicarinata occurs in Panama, Colombia, Peru and Ecuador. In Panama it is found at low elevations. ANAL ZONE: Boy Scout road, Madden Dam area, Dwyer 11971 (MO); ides et al. 4069 (WIS). Ft. Clayton, 1 mi on old Albrook Tower road, Tyson & Dwyer 6504 (SCZ). cuiRIquí: Monte Verde 2.5 km W of Puerto Armuelles, 80 m, Liesner 65 (MO, NY). COCLE: Ser 50-1000 ft, Williams 132 (NY, US). DARIÉN: Near Refugio, 15-21 mi N of Santa Fé, 30 m, Duke 10258 (MO). Isla Saboga, Duke 10368 (MO). Los SANTOS: 10 mi S of Tonosí, Tyson et al. 2943 (SCZ). PANAMÁ: Taboga Island, Dwyer 3104 (MO). Punta Paitilla, Piper 5169 (GH, US). Around Alhajuela, 30-100 m, Pittier 2354 (GH, NY). Taboga Island, Standley 27925 (US). Between Las Sabanas and Matías Hernández, Stan- dley 31919 (US). 2. Canavalia brasiliensis Mart. ex Benth., Ann. Wiener Mus. Naturgesch. 2: 135. 1837. TYPE: Brazil, Martius 12798 (M, not seen). Canavalia panamensis Piper, Contr. U.S. Natl. Herb. 20: 569. 1925. Type: Panama, Piper 5168 (US, en). Vine trailing or climbing on herbs and shrubs, rarely high climbing; stems terete or drying slightly angled, puberulent with ascending white hairs. Leaves pinnate trifoliolate, the leaflets to 15 cm long, 11 cm wide, ovate, chartaceous, apically obtuse but often with a short acuminate and sometimes mucronulate tip, basally obtuse, truncate or rounded, mostly pubescent with short scattered hairs on both sides; petiolules 3-4 mm long, ascending pubescent; petioles mostly shorter than the leaflets. Inflorescences axillary racemes 10-20 cm long, the flow- ers mostly towards the apex; bracteoles rotund, ca. 1.5 mm long; pedicels 1—1.5 mm long. Flowers showy, pink, white or purplish, calyx campanulate tubular, 8- 12 mm long, drying with evident veins but seldom mottled, slightly oblique, the upper lip as long as the tube, entire or emarginate, deflected upwards, and the area behind compressed, the area behind this compression expanded into a small hump or bulge, the lower lip consisting of 2 round or pointed lateral teeth and 1 slightly longer, lower tooth 2-2.5 mm long, pointed and often ciliolate; standard 2.25 cm long. Legume woody, to 20(25) cm long and to 2.5 cm wide, oblong with a short downcurved beak, each valve with prominent sutural ribs and one addi- tional rib 4-6 mm from the sutural rib, usually quite glabrescent; seeds ca. 12, ellipsoidal, 12-15 mm long, brown, sometimes marbled, the hilum ca. 7 mm long, forming ca. ?4 of 1 edge. This species is often difficult, especially when dried, to distinguish from other Panamanian species. From Canavalia glabra and C. oxyphylla it is distinguished by a larger lowest calyx tooth and often by the emarginate upper calyx lip. From C. ensiformis it differs in its smaller seed. Canavalia brasiliensis is one of the most widespread and common of the Canavalia species. It has been recorded from Florida, Mexico and Central and South America as far south as northern Argentina. It is a species of disturbed lowlands, mostly below 1,000 m. This species has occasionally been planted in some countries as a cover crop. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 565 BOCAS DEL TORO: Changuinola Valley, Dunlap 253 (US, F). Water p Wedel 809 (MO, GH). j S, WIS). S P WIS). Near Chepo, Hunter & Allen 97 (MO). Debajo del puente de Juan Díaz, Lara 47 (F, PMA). Sabanas N of Panama City, Paul 594 (US). Agricultural station, Matías T med gs (GH, US), 6913 (US). Bella Vista, Standley 25348 (US). Juan Franco Race Track ne ndley 27693 (US). Juan Díaz, Standley 30475 (US). Punta Paitilla, Sun 30818 (US). Beiudon J Matil Hernández and Juan Díaz, Standley 31951 (US). 3. Canavalia dictyota Piper, Contr. U.S. Natl. Herb. 20: 574. 1925. TYPE: Guyana, Jenman 4211 (US, not seen). Twining vine clambering on herbs and shrubs, sometimes high climbing; stems glabrescent. Leaves pinnate trifoliolate, the leaflets mostly to ca. 15 cm long, 6 cm wide, ovate or elliptic, apically obtuse mostly with a short, often blunt acu- men, mucronulate, basally obtuse, slightly coriaceous, sparsely pubescent or gla- brate. Inflorescence to 25 cm long, the peduncle and rachis becoming woody in fruit; bracteoles rotund, ca. 1 mm long; pedicels 1.5-2.5 mm long. Flowers pink or lavender; calyx ca. 13 mm long, tubular campanulate, the upper lip large and broad, nearly as long as the tube, deeply emarginate, forming a small beak, the lower teeth obtuse, 2-3 mm long, the lowest tooth shorter than the laterals; standard 2.5-3 cm long, erect. Legume to 20 cm long, 3.5 cm wide, oblong with a short, apical beak and a short basal stipe; compressed, becoming woody, gla- brate, longitudinally ridged at the sutures and with an additional ridge 3-5 mm from the longitudinal ridge; seeds 6-9, 17-20 mm long, compressed ovoid to ellipsoidal, mottled brown, often buoyant, the hilum extending along most of one edge. Canavalia dictyota may be distinguished by the lowest calyx tooth which is shorter than the 2 lateral teeth. It also has narrower leaves than many other species, and the seed is often buoyant. The pod has a distinct stipe. Plants are less pubescent than most other Panamanian species. most all Panamanian collections of this species are from the same locality, an open area of tropical moist forest. Canavalia dictyota ranges through the Antilles and northern Central America. CANAL ZONE: BARRO COLORADO ISLAND: Bailey & Bailey 290 (F); Brown 53, 107 (both F); Croat 4675 (MO, SCZ), 4794 (F, MO, NY, SCZ), 7062 (MO), 7399 (F, MO, SCZ), 7228 (F, MO, NY, SCZ, US), 7702 (MO), 12944 (MO); Foster 1477 (MO, dp hp 427, 628 (both F, MO); Wetmore & Abbe 45 (A, MO), 45A (F, GH), 366 (A, F, MO), 381 (F); Wilson 77 (F); Zetek 3801 (F, MO). CANAL ZONE: Mindi, Cowell 183 pi^ asa 2 9051, 12607 (both MO), 13224 (F, MO). Miraflores, White & White 63 (MO, U 4. Canavalia ensiformis (L.) DC., Prodr. 2: 404. 1825. Phaseolus maximus, siliqua ensiformis .... yy Cat. Pl. Jamaica 1: 68. 1696. LECTOTYPE (fide Sauer) Sloane, Cat. Pl. Ja ide tab. 114 Dolichos ensiformis L., Sp. Pl. 2: 725. 1753. Based 1 on Phaseolus maximus, siliqua ensiformis .... Sloane. Vine, stems glabrescent. Leaves pinnate trifoliolate, leaflets to 20 cm long, broadly ovate, apically obtuse with a blunt apiculum, glabrate. Inflorescence 566 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 slender in flower, to 30 cm long; bracteoles obtuse, ca. 2 mm long; pedicels ca. 2 mm long. Flowers lavender; calyx tubular, ca. 14 mm long, sparingly pubescent, the upper lip about equalling the tube, emarginate, the lowest tooth slender, ca. 2.5 mm long, longer than the ovate lateral teeth; standard 2.75 cm long. Legume to 30 cm long, 3.5 cm thick, woody, oblong with a short apical beak and short basal stipe, explosively dehiscent, the valves glabrescent, each with a sutural rib and an additional rib ca. 5 mm from the sutural rib; seeds ca. 29 mm long, oblong, compressed, white or ivory, the hilum ca. 9 mm long, about !^ as long as the seed. Canavalia ensiformis may be distinguished by its white seeds, which are not buoyant. The lowest calyx tooth is longer and narrower than any other Pana- manian species except C. villosa. Canavalia ensiformis is also recognizable by its narrow calyx tube. This species is cultivated from time to time for soil binding or fodder, and the plants collected in Panama may have been cultivated. Sauer (1964) indicated that this species was domesticated by pre-Colombian Indians and that current records of the plant all come from cultivation. The plants were cultivated as an annual crop, although plants may live longer than one year. In South America, the seeds and shoots are eaten. CANAL ZONE: Vine on trellis at barracks, Fort Clayton, Greenman & Greenman 5188 (MO). 5. Canavalia glabra (Mart. & Gal.) Sauer, Brittonia 16: 130. 1964. Wenderothia glabra Mart. & Gal., Bull. Acad. Roy. Sci. Bruxelles 10: 192. 1843. TYPE: Mexico, Galeotti 3307 (P, not seen). Canavalia villosa Benth. var. glabra (Mart. & Gal.) Standley, Field Mus. Nat. Hist., Bot. Ser. 11: f 6. C. munda Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 10. 1943. TYPE: Gua- temala, Steyermark 33179 (F, not seen). Vine trailing or climbing on herbs and shrubs; stems glabrate, sometimes with sparse erect brownish hairs. Leaves pinnate trifoliolate, leaflets mostly 10-14 cm long, 4-7 cm wide, ovate, apically acuminate, acute or obtuse with a short acu- men, sparsely pubescent with weak white hairs. Inflorescence to 20 cm long; bracteoles rotund, ca. 2 mm long; pedicels ca. 3 mm long. Flowers white or rose; calyx ca. 13 mm long, pubescent with short, dense hairs, the tube relatively long, the upper lip as long as the tube, forming a short umbo, the top of the calyx with a distinct hump behind the recessed area behind the lip, the lowest tooth ca. 2.5 mm long, pointed, exceeding the obtuse lateral teeth; standard ca. 4 cm long. Legumes to ca. 20 cm long, 3 cm wide, oblong with a pronounced beak and a short narrow stipe, compressed, glabrescent, with prominent longitudinal ridges at the sutures and each valve with an additional ridge near the sutural ridge; seeds to 15 mm long, oblong, compressed, the hilum ca. 7 mm long. Canavalia glabra may be recognized by the hump on the top of the calyx and by the relatively long lowest calyx tooth. In spite of its epithet, the species is more pubescent than some other species in Panama. This species ranges from southern Mexico to Peru, but it is more common in northern Central America. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 567 HERRERA: Chitré, 20 m, Allen 1108 (MO). Los SANTOS: Headwaters of Río Pedregal 25 mi SW of Tonosí, 2500-3000 ft, Lewis et al. 2956 (MO, SCZ). 6. Canavalia gladiata (Jacq.) DC.,”° Prodr. 2: 404. 1825. Dolichos gladiatus Jacq., Collect. Bot. 2: 276, tab. 215, 1788. LECTOTYPE: Jacquin’s plate (design. er 1964). Twining climbing vines; stems terete, glabrate, with a few minute retrorse weak hairs. Leaves pinnate trifoliolate; leaflets ovate, mostly ca. 10 cm long, ca. 6 cm wide; apically acute or short acuminate, basally rounded or obtuse, char- taceous, glabrate, the surface with scattered minute hairs. Inflorescence axillary, ca. 10 mm long; bracteoles caducous; pedicels ca. 1.mm long or obsolete, rachis with swollen glandular nodes. Flowers lavender; calyx tubular, 10-15 mm long, glabrate, the upper lip elongate, emarginate, the lowest tooth ca. 1.5 mm long, pointed, slightly exceeding the lateral teeth, puberulent; standard 2.5-3 cm long. Legume 24-30 cm long, ca. 5 cm wide, oblong, compressed, the beak slightly downturned, longitudinally ridged on the sutures and with an additional ridge 4— 6 mm from the sutural ridges, glabrate or with minute, inconspicuous scattered white hairs; seeds dark red or pink, oblong ovate or elliptical, 3-4 cm long, ca. 2 mm wide, compressed, the hilum ca. 2 cm long, dark, ?4 the length of one edge of the seed. This species may be distinguished by its very large pods and large pink or red seeds. Canavalia gladiata is commonly cultivated in tropical regions of the Old World. The pods are cooked green and the seeds are also eaten. Sauer (1964) reports that it is unknown except in association with cultivation. The species is rarely cultivated in the New World. The single Panamanian record is of a plant cultivated as an ornamental patio climber. The large pods and unusual seeds render it a horticultural curiosity. ANAMÁ: San Francisco de la Caleta, seeds obtained from Camino de la Pintada at Santa María, Coclé Province, Arosemena 26 Oct. 1979 (PMA). 7. Canavalia maritima (Aubl.) Thouars,?' J. Bot. (Desvaux) 1: 80. 1813 'Cana- , 9 vali’ .— Dolichos maritimus Aubl., Hist. Pl. Guiane Fr. 765. 1775. LECTOTYPE: aaa 1964) illustration of aseolus maritimus een .... Pluk., TI tab. 51, fig. 2 91. Dolichos maritim . P. Br., Civ. & Nat. Hist. Jam. 293. pd Based on Phaseolus maritimus cite S.. PI D. roseus Swartz, Prod. Veg. Ind. "Occ. 105. 1788. Based on Dolichos maritimus . . . . Browne. Canavalia rosea (Swartz) DC., Prodr. 2: 404. 1825. Prostrate vine, occasionally climbing; stems sparsely puberulent with short white hairs, glabrescent, sometimes stout. Leaves pinnate trifoliolate, the leaflets mostly ca. 10 cm long, 10 cm broad, broadly obovate, ovate or orbicular, apically rounded or emarginate, occasionally short acuminate, mucronulate, coriaceous, 20 For a list of synonyms see Sauer, 1964. All synonyms listed by Sauer relate to Old World Plants and only the names noted here relate directly to Panamanian 1 For further synonyms see Sauer, 1964. Only these names relate directly to Panamanian plants. 568 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 9. Canavalia maritima (Aubl.) Thouars.—A. Habit (x1⁄4). [After Wedel 2131.]—B. Flower (x 114).—C. Stamens (x 154).—D. Pistil (x 114). [After Wedel 2835.]—E. Fruit (xV2).—F. Seed (X1). [After Tracy 184.] sparsely puberulent with short white hairs on both sides, more so beneath; pe- tiolules 5-9 mm long. /nflorescence axillary, to 30 cm long, prostrate but as- cending at the tip; bracteoles often paired, obtuse, 1-1.5 mm long; pedicels 2-4 mm long. Flowers pale rose or blue; calyx tubular, 10-12 mm long, the upper lip 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 569 emarginate, short, not umbonate, the lowest tooth ca. 2 mm long, slightly ex- ceeding the lateral teeth, moderately pubescent; standard ca. 3 cm long. Legume mostly 9—15 cm long, ca. 2.5 cm thick, oblong, minutely puberulent, longitudinally ridged at the sutures and each valve with 1 additional ridge 3—5 mm from the suture; seeds ca. 14 mm long, ovoid, brown, the hilum short, ca. 7 mm long; buoyant. Canavalia maritima is distinct in its broad, apically rounded or emarginate leaflets, in its often longer petiolules and pedicels, and in the relatively short upper lip of the calyx. This species is mostly restricted to sea beaches, where it is a pioneer on the sand along with Ipomoea pes-caprae. At the edges of beaches it sometimes climbs on rocks or thickets. Canavalia maritima is common to tropical and subtropical seacoasts in both hemispheres. BOCAS DEL TORO: Isla Colón, Wedel 449 (MO). Old Bank Island, Wedel 2131 (MO). Isla Colón, Wedel 2835 (MO). CANAL ZONE: Farfan area, Ft. Kobbe, D'Arcy 9632 (MO); Duke 4208 (MO). E of Gatün Locks, Duke 4320 (MO). Farfan Beach, Dwyer 6768 (MO). Chagres, Fendler 80 (MO). Farfan Beach, Lewis et al. 307 (MO). Balboa, Mell 4 (NY). Fort Sherman, Porter et al. 4982 (MO); Tvson 2253 (MO). First island out Amador causeway, Tyson & Lazor 6148 (PMA); Tyson 7372 (PMA), 7382 (MO, PMA). cuiRIQUÍ: Guanaban, 2-4 km S of pega Armuelles, Busey 499 (MO). Quebrada Melliza 6 mi S of Puerto Armuelles, 0-150 m, Liesner 508 (MO). COLON: Río Piedras on road to Portobelo, Blum et al. 2526 (MO). San Miguel de la Borda, Eos 9862 (MO, NY, PMA). María Chiquita, Dwyer 4467 (MO); Taylor 113 (PMAO). Los SANTOS: Monagre Beach, Dwyer 4136 (MO). PANAMA: San Carlos, Allen 1141 (MO). Punta Chame near S tip, D'Arcy 10223 (MO). Isla Tobaguilla, Duke 5875 (MO). Isla del Rey, Duke 9550 (MO). Isla de Pedro Gonzales, Perlas Archipelago, Dwyer 1715 (MO). Playa Coronado, Gentry 2899 (MO). San Jose Island, ga 98 (US); Kennedy 2283 (MO). Bella Vista, ar 12008 (NY). Playa Coronado, Koster 175 (MO). PANAMA: Vera Cruz, Lewis et al. 2993 (MO, SCZ). Moro Island, Mori et al. 4080 (WIS). | Farallon, Río Hato, Taylor 112 (PMA). San Jose oe Tyson & Loftin 5051 (MO, SCZ). SAN BLAS: Naragana Island, Croat 16848 (MO). Isla Soskatupu, Duke 8932 (MO). Isla Ustupo, Gentry 6336 (MO). 8. Canavalia oxyphylla Standley & L. O. Williams, Ceiba 3: 201. 1953. TYPE: Honduras, Allen 6458, not seen. Vine twining on shrubs and high climbing; stems glabrate. Leaves pinnate trifoliolate, leaflets mostly 10-12 cm long, 5—7 cm wide, ovate, apically acumi- nate, chartaceous, sparingly pubescent on both sides; petiolules 4-6 mm long. Inflorescences axillary, mostly 10-18 cm long; bracteoles obtuse or deltoid, 1— 1.5 mm long; pedicels 1-1.5 mm long. Flowers white, pink or lavender; calyx 10— 14 mm long, often drying mottled, tubular campanulate, the upper lip short, forming a distinct umbo, the lower teeth membranaceous, the lowest acute, 1.75— 2 mm long, exceeding the obtuse lateral teeth; standard ca. 4 cm long. Legume oblong, the apical beak short, the basal stipe terete, stout, 1.5-2 cm long, woody, glabrescent, each valve with sutural ridges and 1 additional ridge situated more than 7 mm from the suture and hence often appearing medial on the valve; seeds compressed ovoid, ca. 17 mm long, tan colored, the hilum forming half the cir- cumference. This species is distinct in its pod which in addition to the sutural ridges has one longitudinal ridge appearing almost medially on the valves, and which has a stout, terete stipe 1-2 cm long. The seeds are quite flat, and the hilum is longer than on other Panamanian species. The lower calyx teeth are membranaceous 570 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 and conspicuously pubescent and the lowest is acute. The short upper lip forms a distinct umbo. In spite of these differences, the species is often difficult to distinguish in the herbarium from C. brasiliensis and C. bicarinata. Canavalia oxyphylla ranges from southern Mexico to northern South America. OCAS DEL TORO: Santa Catalina, Blackwell et al. 2712 (MO). Water Valley, ne 1595 (MO, GH, MO). 1803 (US), 1820 (GH, MO), 1846 (MO, GH). CANAL ZONE: Farfan beach area, Tyson et al. 3161 (SCZ). CHIRIQUí: Monte Verde 2.5 mi W of Puerto Armuelles, 80 m, Liesner 57 (GH). Above San Félix, road to Cerro Colorado, 800-1000 m, Croat 33163 (MO). cocLé: El Valle, Croat 13313 O . Bie Aires, 340-410 m, Nee 9077 (MO, US). P Á: Capira along trail between Ledice and Aguacate, 300 m, Foster 2147 (MO, diu Mai uis Experimental. de Monte Oscuro, 12 km de Capira, 10 m, Correa 1932 (PMA). VERAGU n Canazas and foot of Cordillera Central, headwaters of Río Canazas, 300-600 m, Allen 197 o. US). Hills W of Soná, 500 m, Allen 1032 (GH, MO, US). 9. Canavalia villosa Benth.,22 Ann. Wiener Mus. Naturgesch. 2: 135. 1837. TYPE: Mexico, Karwinsky M12819 (M, not seen). Twining vine climbing on shrubs and trees, stems mostly puberulent with curved brownish hairs. Leaves pinnate trifoliolate; leaflets mostly 10-12 cm long, 5-8 cm wide, ovate or obovate, the lateral leaflets oblique, apically acuminate, basally obtuse or rounded, drying discolorous, pubescent along the veins and margin on both sides, mostly copiously pubescent on the lamina, especially be- neath; petiolules and petioles mostly densely pubescent. /nflorescence mostly 12-20 cm long; bracteoles ca. 2 mm long, often wider, pointed or rounded, mostly tomentose; pedicels obsolete or to 3 mm long. Flowers blue, purplish, or pink; calyx 17-20 mm long, often drying mottled, campanulate, the upper lip as long as or longer than the tube, prominently umbonate or not, sometimes slightly emarginate, glabrate to tomentose, the lowest tooth narrow, 2.5-4.5 mm long, longer than the deltoid lateral teeth; standard ca. 3.5 cm long, often drying brown- ish and striate. Legume mostly 14-17 cm long, ca. 2.5 cm wide, oblong, some- times slightly expanded apically, the beak short and hardly downturned, stipe evident or not, longitudinally ridged at the sutures and with 1 additional ridge 5— 10 mm from the sutures, long shaggy pilose when immature, at maturity evenly pilose with soft erect hairs; seeds 12-17 mm long, elliptical discoid, light brown, mottled, the hilum short, 6-7 mm long. Canavalia villosa is distinct in a number of features; long calyx, long and narrow lowest calyx tooth, elongate upper calyx lip, narrow pointed bracteoles, and usually in its general overall pubescence. This species ranges from northern Mexico to Panama and has been occasion- ally collected in South America. In Panama most collections have been made in upland areas. CHIRIQUÍ: El Barú above Boquete, 1200-1800 m, D'Arcy 9932, 9934 (both MO). Lava flow between El Hato and Bambito, D'Arcy 10034 (MO). Lava fields near town of Volcán, Duke 9138 (MO, NY). From Boquete to 3 mi N, 3300-4200 ft, Lewis et al. 325 (MO, GH, US), 603 (GH, NY, US). 2.5 km W of Puerto Armuelles, 80 m, Liesner 57 (MO). eu deni 0-120 E Pittier 5298 (GH NY, US). Boquete, 3800 ft, Terry 1282 (A, F, MO, US). Baja Mono, 4000 ft, Terry & Terry 1648a (F). Lava flow 4 mi above Volcán, Tyson 7285 (PMA). cocrÉ: Eb Valle de Pede 1000-2000 ft, ?? Several synonyms for this species are given by Sauer, 1964. Only this name has been used for Panamanian plants. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 571 Lewis et al. 2612 (MO, SCZ). HERRERA: 4 mi S of Los Pozos, Tyson 2678 (MO). Los SANTOS: 11 mi N of Tonosi, Tyson 2966 (SCZ). 12 mi S of Macaracas, Tyson et al. 3077 (MO, FSU). PANAMA: Debajo del puente de Juan Diaz, Lara 47 (F). VERAGUAS: Quebrada just S of Santa Fé, 450 m, Nee 8045 (MO). 10. CENTROSEMA W. G. D'Arcy? Centrosema (DC.) Benth., Comm. Legum. Gen. 53. 1837; Ann. Wiener Mus. Naturgesch. 2: 117. 1840 (1839). TYPE: C. brasiliana (L.) Benth. nomen cons. vs. Steganotropis Lehm. Bradburya Raf., Fl. Ludov. 104. 1817. TYPE: B. scandens Raf. = Centrosema virginiana (L.) Kuntze. Nomen rejic. vs. Bradburia Torr. & Gray (1842). Compositae. Cruminium Desvaux, Ann. Sci. Nat. (Paris) sér. 1. 9: 423, Sept. 1826. TYPE: C. giganteum Desv. = Centrosema plumieri (Pers.) Benth. Steganotropis Lehm. I. Sem. Hort. Bot. Hamburg 1826: 18. 1826; Linnaea 3 (Litt.): 11. 1828. TYPE: S ] nth. . coniugata J. C. C. Lehmann. Nomen rejic. vs. Centrosema (DC.) Be Clitoria sect. III Centrosema DC., Prodr. 2: 234. 1825. Mostly twining vines, trailing, or sometimes high climbing, stems often wiry, puberulent. Leaves imparipinnate 1—5-foliolate, but mostly 3-foliolate, the leaflets narrow or broad; petiolules mostly short, drying wrinkled; petioles sometimes winged; stipels equalling or exceeding the petiolules (Panama); stipules promi- nently nervate. Inflorescences several flowered congested racemes, usually only one flower opening at a time; bracts and bracteoles nervate, the bracts often spathaceous or calyx-like, enfolding the bud. Flowers showy, the standard flat, discoid, often emarginate, with a distinct notch or spur near the base, mostly puberulent outside, the keel and wings directed upwards, curved; stamens dia- delphous, the vexillary stamen free or nearly so, ovary linear, sessile, pubescent, the style incurved, sometimes persistent as an elongate beak, glabrous except basally, apically expanded into a blade, the stigma marginal on the style apex, minutely pubescent. Legume linear, dehiscent along the 2 valves, often spiralling, the sutures prominent, the apex sometimes long-beaked; seeds plump, ellipsoidal, numerous, the hilum linear along one edge. Centrosema is a New World genus of perhaps 35 species ranging from the southern United States into Argentina and throughout the Antilles. The genus is distinct in the notch or spur on the standard and in the flat, linear beaked pods. The striate bracts and stipules, and stipels as long as the petiolules are useful recognition features for the Panamanian species. Centrosema virginiana (L.) Kuntze, widely dispersed in tropical America, has not been collected in Panama. Similar to C. pubescens, it has 5 subequal subulate calyx teeth and the pod is less than 4 mm wide. Literature: Barbosa-Fevereiro, V. P. 1977. Centrosema (A. P. de Candolle) Bentham do Brasil—Leguminosae—Faboideae. Rodriguezia 29: 159-219. Fantz, P. R. 1979. A new species of Centrosema (Leguminosae) from Nicaragua and a key to the species in Central America. Sida 8: 152-156. ?3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 572. ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 a. Leaves unifoliolate (simple); petioles conspicuously winged |. 6. C. sagittata aa. Leaves trifoliolate; petioles unwinge b. Leaves mostly linear, less than 8 mm wide; pod less than 4 mm wide aves with secondary venation at right angles to the midrib; flowers more than 2 cm long; stipules less than 5 mm long |. 1. l. C. angustifolia cc. Leaves with secondary venation ascending; flowers less than 1.5 cm long; stip- ules more than 5 mm long |... U a... . C. pascuora bb. Leaves ovate or elliptical, more than 10 mm long; pod more than 4 mm wide. d. Bracts large, persistent 1-3 cm long, 8-15 mm wide; persistent stipules large 7. C. vexillata dd. Bracts smaller, caducous or persistent, less than 1 cm long, narrow, less than mm long. wide; stipules smaller, 8-11 e. Bracts caducous; bracteoles calyx and pod drying dark; seeds 6-9 mm across; calyx lobes undulate; flowers pink and red... . C. plumieri ee. Bracts persistent; bracteoles calyx and pod drying green or brown; seeds less than 6 mm long; calyx lobes pointed; flowers purple, blue or violet. f. i Pod more than 7 mm wide, the margins ca. 1 mm wide: stipels 4—5 mm long; leaves glabrate above, mostly 8-13 cm long, narrowly ovate; petiolules 3-5 mm long |... 2. 2. C. macrocarpa = Pod mostly less than 7 mm wide, the margins less than 1 mm wide: stipels 1-2 mm long; leaves pubescent above, mostly 4-8 cm long, broadly ovate, petiolules 1-3 mm long = 5. C. pubescens 1. Centrosema angustifolia (H.B.K.) Benth., Comm. Legum. Gen. 54. 1837.— Fic. 10. Clitoria angustifolia H.B.K., Nov. Gen. Sp. Pl. 6: 417. 1824. TYPE: Venezuela, not seen. Slender twining vines; stems wiry, minutely pubescent with erect white hairs. Leaves pinnate trifoliolate; leaflets narrowly linear oblong, mostly 3—5 cm long, 5-9 mm wide, gradually long tapering to a blunt tip, basally rounded and minutely cordate, the costa conspicuous, lateral veins ca. 20-30 arising at right angles to the costa, anastomosing irregularly near the margin, shiny and glabrate above but with scattered minute reduced trichomes and a small tuft at the base, glabrous beneath, slightly revolute; petioles 1-1.5 mm long, drying wrinkled with a few hairs; rachis ca. 2 mm long, ridged or channelled dorsally; petioles mostly shorter than the terminal leaflet, sparingly pubescent with erect elongate hairs; stipels subulate, glabrous, exceeding the rachis; stipules narrowly deltoid, ca. 4 mm long, conspicuously striate. Inflorescence a few flowered congested raceme, only 1 flower opening at a time; peduncle slender, glabrate, 1-20 cm long; bracts spathaceous, strongly nervate, 10-15 mm long, the inner bract pointed; pedicels obscure. Flowers showy, purple, calyx cupular, 10 mm long, the sinuses rounded, the lobes deltoid, but the lowest lobe elongate, subulate, ca. 8 mm long, standard orbicular, flat, minutely emarginate, 2.5-3 cm across. Legume linear, 7-10 cm long including the acicular, 10-15 mm long beak, 3-4 mm wide, glabrous; seeds dark brown, discoid, ca. 2 mm long. Centrosema angustifolia may be recognized by its smaller, narrower, sub- coriaceous leaves with venation diverging at right angles from the costa, and by the seeds which are only about 2 mm long. This species ranges from Central America to northern South America. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 573 C" C? FIGURE 10. Centrosema angustifolia (H.B.K.) Benth.—A. Habit (x14). [After Hunter & Allen 37.]—B. Pu (x1).—C. Corolla.—C!. Standard (x!4).—C?. Wing petal (xV2).—C?. Keel petals (x14).—D. Stamens (x2).—E. Anthers.—F. Pistil (x2). [After Dodge et al. 16679.]—G. Fruit with seeds (x4). [After Dodge et al. 16679A CANAL ZONE: Government Ss along Las Cruces Trail, 75 m, Hunter & Allen 707 (MO). Ancon Hill, Greenman & Greenman 5126 (MO); Piper 5148; Standley 25182 (US); 26344 (MO, US). CHI- RIQUí: Sabana de Marcelito near El Vigia, Pittier 2380 (US). cocLE: Nata, 50 m, Allen 814 (MO). HERRERA: to 8 mi SW of Las Minas, D’Arcy 1354] (MO). PANAMA: Pacora, 35 m, Allen 995 (MO). Road between aste te nd Chepo, Dodge et al. 16679 (GH, MO). Cerro Campana, Duke 5960 (MO). Savanas near Chepo, Duke 6054 (MO). Las Sabanas, Heriberto 142 (GH, US). Savanas near Chepo, Hunter & Allen 37 (MO). Juan Diaz, Killip 3162 (US). Taboga Island, Miller 2023 (US). Laguna de Portala near Chepo, 50 m, Pittier 4599 (US). Sabana de Dormisolo near Chepo, Pittier 4650 (US). Big Swamp E of Rio Tocumen, Standley 26539 (US). Tumba Muerta Road near Panama, Standley 29748 (US). 574 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 2. Centrosema macrocarpa Benth., Ann. Nat. Hist. (Paris) ser. 1. 3: 436. 1839. TYPE: Guiana, Schomburgk (K, not seen). Centrosema lisboae Ducke, Notizbl. Kónigl. Bot. Gart. Berlin 11: 548. 1932. TYPE: Brazil, Ducke 2309 (G, not seen; photo MO). Robust vine, stems terete, becoming somewhat woody, pilose with tan colored hairs, glabrescent. Leaves pinnate trifoliolate, the leaflets narrowly ovate, api- cally acute or slightly acuminate, mostly 8-13 cm long, 3-8 cm wide, basally rounded, papyraceous to subcoriaceous, glabrate on both sides or pubescent beneath; petiolules 3-5 mm long, stout; rachis and petioles drying grooved, pu- bescent; stipels linear subulate, about as long as the petiolules; stipules acute, nervate, 4-5 mm long. Inflorescences few flowered pseudoracemes, peduncles 2-6 cm long, the rachis thickened with conspicuous nodes, 2-3 cm long; bracts stipule-like; bracteoles ovate subspathaceous, ca. 10 mm long, ciliolate, glabrate, finely nervate, enclosing the calyx cup but not the longest tooth; pedicels 3-7 mm long, puberulent. Flowers pink and red within, green and pink outside; calyx cupular, the cup 3—4 mm long, the teeth dorsally pubescent, the longest 7-10 mm long, the others 4-5 mm long, the lower pair united about halfway up, the sinus acute; standard orbicular, ca. 3 mm long, puberulent dorsally. Fruit linear, mostly 15-20 cm long, 7-8 mm wide, compressed, smooth, the margins conspicuously thickened, ca. 2 mm wide, the beak 10-13 mm long, acicular, straight, glabrous; seeds dark brown, rectangular, 5-6 mm long, the hilum occupying one side. This species is distinct in its narrow, pointed leaves with small stipels and its large pod. The vine is sturdier with stouter stems than most other Panamanian species of Centrosema. Centrosema macrocarpa is widespread in tropical South America ranging as far as Sao Paulo, Brazil, and Panama. In Panama it has been collected in the Canal Zone and in Bocas del Toro. CAS DEL TORO: Quebrada Huron, Kirkbride & Duke 454 (MO). CANAL ZONE: Near water reservoir W of Cocoli, Croat 9163 (MO). Road C21 near Police Lodge, Croat 12987 (MO). K-9 Road, Dwyer 3017 (MO). '^ km W of Summit Gardens, Nee 9492 (MO). Ims . Centrosema pascuora (Mart. ex Benth., Comm. Legum. Gen. 56. 1837. TYPE: Brazil, Martius, not seen. Delicate vine or erect herb; stems slender, sparingly long pilose with spread- ing white hairs, glabrescent, sometimes reddish, a fine tomentum of minute hairs sometimes persistent near the base of the plant. Leaves trifoliolate, the leaflets linear, narrowly ovate, or oblong, mostly gradually tapering to the apex; apically mucronate, basally abruptly obtuse, mostly 4-10 cm long, 4-7 mm wide, glabrate, the major pinnate veins ascending, arcuate and anastomosing near the margin to form a submarginal vein, the minor venation reticulate, elevated; petiolules ca. 2 mm long with a few elongate white hairs; petioles mostly 2-4 cm long, glabrate, slender, drying angled; stipels subulate, 3-6 mm long; stipules ca. 7 mm long, acuminate deltoid, sometimes slightly reddish, strongly nervate. Inflorescences l or 2 peduncles per leaf axil, each peduncle 1 flowered but with an undeveloped flower at the apex; peduncles 5-10 mm long, sparingly pilose; bracts ca. 8 mm 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 575 long, ciliate, resembling the stipules but narrower; bracteoles resembling the bracts but broader, covering most of the calyx; pedicels about as long as the peduncles, glabrous. Flowers blue, standard ca. 2 cm long, calyx short cupular, the teeth subulate, the uppermost longest, ca. 7 mm long, the others ca. 5 mm long, the sinus between the lowest 2 shorter; standard orbicular, flat, ca. 2 cm long. Legume linear, slightly curved, compressed, mostly 4-8 cm long, 3-4 mm wide, the margins thickened, the beak acicular, to 10 mm long, sparingly pilose; seeds compressed rectangular, ca. 4 mm long, brown. This species is superficially similar to Centrosema angustifolia, but the ve- nation of the leaves is ascending rather than at right angles to the costa; the flowers are smaller and the seeds are larger. Centrosema pascuora ranges from Panama to Ecuador and Brazil. In Pan- ama it has been collected near the Pacific coast. cociÉ: Río Hato airstrip, Burch et al. 1132 (GH, MO, US). PANAMA: Río Mar, Tyson et al. 2303 (FSU, NY, SCZ). 4. Centrosema plumieri (Pers.) Benth., Comm. Legum. Gen. 54. 1837. Clitoria — Pers., Syn. Pl. 2: 303. 1807. Based on Phaseolus ampla flor . Plumier. Descr. Pl. . 5: 108. 1713 [1693]. Type: Plumier’s illustration, based on a Pee: S ellechon from Port- s Paix, Hispaniola. Robust twining vine; stems pubescent just above the nodes, otherwise often glabrate. Leaves pinnate trifoliolate, leaflets broadly ovate to rhombic 5-9(-15) cm long, sharply short acuminate, basally obtuse or cuneiform, shiny and glabrate above except on the midvein, copiously appressed pubescent beneath with straight white hairs; petiolules 3—4 mm long, drying dark, glabrate except tomen- tose ventrally; rachis and petiole drying with a conspicuous dorsal groove near the ends, glabrate to pilose; rachis 10-25 mm long, petiole mostly shorter than the terminal leaflet; stipels subulate, exceeding the petiolules, stipules obtuse, ca. 5 mm long, conspicuously striate, ciliolate. Inflorescence axillary, few flowered; peduncles stiff, drying dark, 2—10 cm long; bracts caducous; bracteoles 2, inserted at the top of the pedicel, spathaceous, 10—20 mm long, conspicuously nervate, apically ciliolate, often drying dark; pedicels 6-10 mm long. Flowers showy; calyx cupular, 4-6 mm long, the teeth undulate except for the slightly longer lowest tooth, glabrate; standard orbicular, emarginate, nearly flat, 5—6 cm across, white outside, purple inside, pubescent outside, wings and keel purple. Legume linear, flat, 10-14 cm long, ca. 10 mm wide, thick margined, mostly drying black, glabrate; seeds ellipsoidal, rhomboidal, 6-9 mm long, dark brown, the hilum along one corner This species is distinct in its broad, sharp pointed leaves and in its large showy flowers which are contrasting white and purple. The pod is larger than other Panamanian species of Centrosema, and the margins are much broader. The appressed pubescence of the leaf undersides is also distinctive when present. Centrosema plumieri ranges throughout tropical America and is introduced to the Old World. In Panama it occurs mainly in moist forest areas. 576 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 OCAS DEL TORO: Río Teribe just below Puerto Palenque, Kirkbride & Duke 547 (MO). See Sige, 11 Nov. 62 (SCZ). Water Valley, Wedel 1629 (GH, MO, US), 1753 (GH, US). CANAL Z Tabernilla, Cowell 269 (NY). Road to hop decies Hill, Dwyer 4004 (MO). Cocoli, Dwyer 7203 (GH. MO), 72/8 (GH). Gamboa, Greenman & Greenman 5166 (MO). Observatory on Gamboa Road, Hladik 235 (MO). Pedro Miguel, Killip 3346 (US). Summit, Lindsay 413 (MO). Chiva Chiva Trail, Red Tank to Pueblo Nuevo, ad 5178 (US). Between Miraflores and Corozal, ded 2194 (N US). Gamboa, Pittier 6517 (GH, NY, US). Old Las Cruces trail between Fort Clayton and Corozal, Standley 29006 (US). genie 1. ws La Pintada, 400-600 m, Hunter ^ Allen 555 (GH, locality, Kennedy 204. (P. Matías Hernández, Pittier 6884, 6897 n US). dons Díaz, Standley 30473 (US). Between Matías Hernández and Juan Díaz, Standley 31934 (US). SAN BLAS: Río Cartí, Kennedy et al. 2014 (MO); Perrin 1967 (MO). Between Río Pialla and Río Aili gandí, Warner 190 (MO). 5. Centrosema pubescens Benth., Ann. Wiener Mus. Naturgesch. 2: 119. 1837. TYPE: Mexico, Keerle, Herb. Martius (?M, not seen). Bradburya pubescens (Benth.) Kuntze, Rev. Gen. 1: 164. 1891. Twining vine scrambling on herbs and shrubs, stems slender, often wiry, puberulent to pilose. Leaves pinnate trifoliolate; leaflets ovate, apically obtuse or acute, mucronulate, basally obtuse or rounded, mostly 4-8 cm long, 1.5-3.5 cm wide, the terminal leaflet largest, pubescent with short weak erect hairs, more so beneath; petiolules tomentose, 1-3 mm long; petioles shorter than the terminal leaflet; stipels subulate, 1-2 mm long; stipules deltoid acuminate, drying striate, 1-2 mm long. Inflorescences axillary, congested several flowered racemes: pe- duncle 2-5 cm long, bracts narrow, nervate, caducous; bracteoles 2, spathaceous, 5-8 mm long, conspicuously nervate, enveloping the lower calyx teeth; pedicels ca. 5 mm long, glabrate. Flowers showy, blue or violet, almost white: calyx cupular, the lower tooth mostly subulate, often pubescent, 2-3 times as long as the others, the other teeth deltoid, 2-3 mm long, glabrate; standard 3-5 cm broad, orbicular, nearly flat, emarginate, puberulent outside, with fine dark lines and often with a purplish spot. Legume linear oblong, greatly compressed, 6-7 mm across, the margins thickened, mostly straight, 5—7 cm long, the beak prominent, slender, 0.5-1.5 cm long; seeds 15-30, compressed rectangular, dark brown, mot- tled, ca. 3 mm long. This species is distinct in its calyx which has only 1 elongate, subulate tooth, the others being short and the calyx margin nearly undulate Centrosema pubescens ranges from the southern United States to Argentina and is found in the Antilles. Two Panamanian collections, e.g. Folsom et al. 7174 and Lewis et al. 677, are conspicuously different in appearance from the mass of collections of this species from most of its range. The difference consists of much greater pubes- cence on bracteoles and bracts, the hairs being much denser and longer than in usual material. A number of other collections like these have been taken from widely scattered plants in Central America, but intermediates between this pu- bescent material and the usual pubescence have not been seen, suggesting that two different taxons are present. Bentham’s original description of Centrosema pubescens notes bracts densely sericeous villous and bracteoles sericeous, which 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) $77 seems to apply more to the pubescent plants than the usual less pubescent plants, so recognition of more than one taxon might require renaming of the commoner plants rather than the unusual plants. A study of variation within the full range of material and study of the type collection will be necessary to clarify the taxons and names to be recognized here. s DEL TORO: Almirante, Cooper 76, 189 (both F). Changuinola to 5 mi S of junction of Rio Terebe, 100- 200 ft, apace et al. 931 (MO). Water Valley, Wedel 1813 (GH, MO, US). CANAL ZONE: BARRO COLORADO ISLAND: Croat 4398 (MO), 4947 (SCZ), 6986 (F, MO, SCZ), 7031 (MO); Shattuck 715 (MO); mare P i 157 (F, GH); Woodworth & Vestal 500 (F, GH, MO), 716 (F). Pipeline road N of Gamboa, Correa & Haines 539 (SCZ). Monkey Hill, Cowell 45 (NY). Matachin, Cowell 197 (NY). Pipeline oa. 2.3 mi from Gamboa Gate, Croat 12737 (MO). Miraflores, Croat 12860 (F, Quebrada La Palma and Cañon of Río Chagres, Dodge & Allen 17344 (MO). France Field, Dwyer 2893 (MO). Cocolí, road to Contractors’ Hill, Dwyer 7218 (MO, NY). France Field, Dwyer 8520 (F). Chagres, Fendler 70 (F, US). Ancón Hill, Frost 2 (F). 0.4 mi SE of junction road K-2 & K-15, Graham 318 (GH). Ancón, Greenman & Greenman 5075 (MO). Hills E of Curundu, Harvey 5194 (F). Paraiso, Hayes 112, 797 (both NY). Gatun, Hayes 8/5 (NY). Pipeline road, Hladik 51 sae Gamboa, Kennedy & Steiner 2467 (F). Pedro Miguel, Killip 3348 (US). Ancón, Macbride & Feath- erstone 17 (F, US). Boy Scout camp near Madden Lake & dam, McDaniel 12657 (FSU). Chiva- Chiva Trail, Red Tank to Pueblo Nuevo, Piper 5144 (US). Empire to Mandinga, Piper 5145, 5156 (both US). Juan Mina, Piper 5158 (US). Ancón, Piper 5177 (US). Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, Piper 5179 (F, US). Gatuncillo, Sg 5180 (US). Old Fort San Lorenzo, Piper 5979 (US). Río Grande near Culebra, Pittier 2078 (NY, US). Ancón, Pittier 2763 (GH, NY, US). Balboa, Stan- dley 25270, 25486 (both US). Cerr o Gordo near Culebra, Standley 25959 (US). Balboa, Standley 26067 (US). Corozal, Standley 27391 (US). Gamboa, Standley 28513, 29323 (both US). Rio Pedro Miguel near East Paraiso, gue 29931 (US). Summit, Standley 30111 (US). Between France Field and Catival, Standley 30305 (US). Balboa, Standley 30887 (US). Darién Station, Standley 31576 (US). Howard I: Force Di ui Red Devil drop zone, Tyson 1841 (FSU, MO, SCZ), 1846 (SCZ). 1 mi N of Summit, Tyson et al. 2743 (GH, MO, SCZ). Miraflores Locks area, Tyson & Dwyer 2741 (SCZ). Without qute E ie ab collector (MO-1090658). CHIRIQUI: David/Concepción, 60 m, Burt & Koster 135 (MO). Near Frontera, 100-200 m, Burt & Koster 147 (MO). Volcán/Cuesta de Piedras, 1000-1200 m, Burt & Koster 158 (MO). Puerto Armuelles, Busey 513 (MO). Quebrada Cana Blanca, Burica Peninsula, 40 m, Croat 22578 (MO). Baja Gualaca, 0-100 m, Koster 171 (MO). Quebrada Guanabanito beyond La Represa 2 mi SW of Puerto Armuelles 0-200 m Liesner 129 (NY). Quebrada Punta de Piedra, 2 mi SW of Puerto Armuelles, 0-100 m, Liesner 346 (MO). Río Dupi, Pittier 5241 (NY, US). COCLÉ: El Valle de Antón, Burt & Rattray 39 (MO); Croat 13290 (MO); Dwyer 11957 (MO, US); Lewis et al. 2566 (MO). Aguadulce, Pittier 4954 (US). 10 mi E of Nata at Rio Grande, Tyson 5216 (FSU, > a 5250 (FSU, uni 4 mi W of Anton on Río Chico, Tyson na Island, Hunter & Allen 136 (GH, MO). San Jose Island, n 888, 1010, 1062 (GH). Chimán, Lewis et al. 3354 (MO). Bellevista, Macbride 2741 (F). o Emperador, Munoz 35 (MO). La Cresta, Sandoval 13 (MO). Big swamp E of Río Tocumen, Bou. 26639 (US). Corozal road near Panama, Standley 26809 (US). Taboga Island, Standley 27905 (US). Río Tapia, Standley 28191 (US). Tumba Muerto Road near Panama, Standley 29703, 29752 (US). Between Las Sabanas and Matias Hernández, Standley 31817 (US). SAN BLAS: Mouth of Ailigandí river to 2.5 mi inland, Lewis et al. 76 (GH, MO, US); Warner 178 (MO). vERAGUAsS: Divisa/Santiago, 0-100 m, Burt & Rattray 74 (MO). Puerto Mutis 12 mi S of Santiago, Tyson 5190 (FSU, MO, SCZ), 6007 (SCZ). 6. Centrosema sagittata (Willd.) Brandegee ex Riley, Kew Bull. 1923: 344. 1923. HUE sagittata H. & B. ex Willd., Enum. Pl. 757. 1809. TYPE: not Centrosema hastata Benth., Comm. Legum. Gen. 56. 1837. TYPE: Rudolohia dubia H.B.K., Nov Gen. Sp. Pl. 6: 432. tb. 591. 1823. TvPE: Colombia, not seen, cites Glycine sagittata Willd. 578 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Centrosema dubia (H.B.K.) Hemsley, Biol. Amer. Centr. 1: 294. 1880. Bradburya sagittata (Willd.) Rose, Contr. U.S. Natl. Herb. 8: 46. 1903. Slender, twining vine; stems wiry, puberulent, glabrescent. Leaves simple, hastate to deltate, the blades mostly 9-15 cm long, apically acute or acuminate, the basal lobes oblique and obtuse, glabrate, with puberulence on the midvein above; petiolule wrinkled, pilose, 3-4 mm long; petiole shorter than the blade, the conspicuous wing 2-9 mm wide, gradually tapering to the base; stipels sub- ulate, 2-nerved, about as long as the petiolules; stipules narrow, acute, promi- nently nervate, ca. 6 mm long. Inflorescences congested few flowered racemes, peduncle slender, mostly 1-2 cm long, becoming somewhat stout in fruit; bracts resembling the stipules, caducous, bracteoles 2, spathaceous, conspicuously ner- vate, to ca. 10 mm long. Flowers showy, mostly white, with some purple; calyx cupular, pubescent, the teeth subulate, ca. 5 mm long; standard orbicular, nearly flat, emarginate, ca. 5 cm long. Legume linear, flat, 12-18 cm long including the acicular beak, the beak 5-10 mm long, the margins evident, narrow, minutely puberulent overall, glabrescent; seeds plump, dark brown, ca. 5 mm long, the hilum along one edge CANAL ZONE: Near Ft. Kobbe, Allen 2026 (GH, MO). Madden Dam, 50 ft, Lewis et al. 17 (GH, MO). DARIEN: Without other locality, Bristan 1506 (MO). Los SANTOS: Between Macaracas and Tonosi, 1.7 mi W of Highway 50 to El Cortezo, Hammel 5299 (MO). PANAMA: Between Bayano Dam and Canitas, D’Arcy 938 (MO). Canita de Chepo, Escobar et al. 274 (MO). VERAGUAS: above Santa Maria along road to Santiago, 250-300 m, Nee 8/22 (MO). 7. Centrosema vexillata (Benth., Ann. Nat. Hist. (Paris) ser. 1. 3: 435. 1839. TYPE: Guiana, Schomburgk (K, not seen; photo MO) Vine, sometimes robust, stems sometimes woody pilose with elongate, 2 mm long, weak yellowish hairs. Leaves pinnate trifoliolate, leaflets ovate, 5-12 cm long, apically acuminate, the acumen mucronulate, basally obtuse, above with minute hairs, sparingly pilose beneath, especially on the veins, petiolules 2-3 mm long, pilose; petioles slender, drying sharply grooved, pubescent; stipels linear, conspicuous, to 12 mm long, much exceeding the petiolules; stipules conspicyous, ovate, 5-11 mm long, finely nervate, sparingly long pilose. Inflorescence few flowered; peduncles slender, elongate, pubescent; bracteoles large, ovate, sub- spathaceous, nervate, ciliate, glabrous or pubescent, 2-2.5 cm long, 8-10 mm wide; pedicels 10-15 mm long, slender, pubescent; bracts stipule-like. Flowers (from Barbarosa-Fevereiro 1970) white; calyx puberulent, the tube 5-6 mm long, the upper teeth completely fused forming a lip 3-4 mm long, 3 mm wide, truncate or emarginate apically, the lower teeth triangular, the laterals 2-3 mm long, the median 5-7 mm long; standard 4.5-5 cm long, 5-6 cm wide, dorsally pubescent. Fruit straight, linear to oblong, compressed, ca. 10 cm long, 6-8 mm wide, co- piously pubescent, ultimately black; seeds dark brown, rectangular 5-6 mm long. Centrosema vexillata is distinct in its large stipels, stipules and bracteoles. The flat pubescent pods are also distinctive. This species ranges from Panama to central Brazil. In Panama it has been found only near the coast. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 579 MÁ: Camino a Río en Chame, Taylor 91 (PMA). SAN BLAS: Lower Río Ailigandí, Duke 9324 (MO. “ Sasardi, 20 m, Duke 10136 (MO, NY). Molia near Playon Chico, Stier 65 (MO). 11. CHAETOCALYX Michael O. Dillon?* Chaetocalyx DC., Prodr. 2: 243. 1825; Mém. Légum. 6: 262. 1826. TYPE: Chae- tocalyx vincentina DC. = C. scandens (L.) Urb. Boenninghausia Spreng., Syst. Veg. 3: 245. 1826. TYPE: B. vincentina Spreng. = Chaetocalyx scan- L rb dens (L. ; Planarium Desv., Ann. Sci. Nat. (Paris) 9: ie 1826. TYPE: P. latisiliquum (Poir.) Desv. = Chae- tocalyx latisiliqua (Poir. ) Benth. ex Hem Rhadinocarpus Vog., Linnaea 12: 108. 1838. me R. brasiliensis Vog. — Chaetocalyx bras- iliensis (Vos) Benth. Isodesmia Gardner, London J. Bot. 2: 339. 1843. TYPE: I. tomentosa Gardner = Chaetocalyx to- fec Harms, Notizbl. Bot. Gart. Berlin-Dahlem 10: 387. 1928. TYPE: R. platycarpus Harms = Chaetocalyx platycarpa (Harms) Rudd. Twining vines, herbaceous to suffrutescent; stems slender, 1-4 mm in diam- eter, subterete, striate, glabrous to densely pubescent. Leaves imparipinnate, 5— 7-foliolate, the rachis glabrous to pubescent, 2.0-12.5 cm long; leaflets oblong, elliptical, ovate or obovate, 1-8 cm long, 0.5-5.0 cm wide, entire, acute to obtuse or retuse, mucronulate, basally rounded, cuneate or subcordate, glabrous to pu- bescent, sometimes micropunctate, pinnately veined; petiolules pulvinate, 1-2 mm long; stipules paired, attached basally, linear to deltoid-ovate, acute to at- tenuate, entire to setose-ciliate or laciniate; exstipellate. Inflorescence terminal or axillary, racemes, panicles, fascicles, or rarely solitary; bracts stipuliform; ebracteolate. Flowers 5-merous, 12-30 mm long; calyx campanulate, 5-lobed, subequal, glabrous to densely pubescent with tubercular-based trichomes, or ab- nt, symmetrical to gibbous, basally articulate; corolla yellow or sometimes red to violet striate, the standard obovate to suborbicular, emarginate, the wings oblong, free, clawed, the keel nearly straight, clawed, free basally; stamens 10, filaments connate into a sheath, splitting above, or the vexillary stamen free, occasionally splitting below with maturity, the anthers dorsifixed, ellipsoidal, ca. 1 mm long; ovary sessile or stipitate, 6—16-ovulate, glabrous to densely pubescent, the style filiform, incurved, glabrous, the stigma capitate. Fruit a loment, 6-16- articulate, submoniliform, subterete to compressed, linear, reticulate to longitu- dinally striate, the articles oblong to quadrate; seeds elongate, 2.5-6.0 mm long, 1-2 mm wide, subcompressed, smooth, sublustrous, reddish brown, estrophio- late. Chaetocalyx includes about 29 species ranging from southern Mexico into the Antilles, through Central America, and in South America in Peru, northern Ar- gentina, southern Brazil and Uruguay. Most taxa occur in more or less mesic habitats up to 200 m. The genus is represented in Panama by only one species, Chaetocalyx latisiliqua. 4 Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 580 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Literature: Rudd, V. E. 1958. A revision of the genus Chaetocalyx. Contr. U.S. Natl. Herb. 32(3): 207—245. — . Chaetocalyx latisiliqua (Poir.) Benth. ex Hemsley, Biol. Centr. Amer. 1: 268. 1880. TYPE: "Peru," herb. Jussieu (P-JU, holotype, not seen; microfiche MO).—Fic. Hedysarum latisiliquum Juss. ex Poir. in Lam., Encycl. ADS Bot. 6: 432. 1804. Poiretia latisiliqua (Poir.) Desv., J. Bot. (Desvaux) 1: 122. 1813. Planarium latisiliquum (Poir) Desv., Ann. Sci. Nat. Cun : 416. 1826. Sparingly branched vines ; stems brownish, puberulent and sparsely pubescent with trichomes ca. 1.5 mm long. Leaves 5-foliolate, rachis 4-10 cm long, sparsely sericeous; leaflets oblong-elliptic to obovate, 1-5 cm long, 0.6-3.0 cm wide, en- tire, obtuse mucronulate, base rounded to cuneate, discolorous, micropunctate, adaxial surface green, puberulent, abaxial surface whitish, lannose and puberu- lent; stipules lanceolate to linear, 3-10 mm long, 1-3 mm wide, attenuate, entire to setose-denticulate, puberulent. /nflorescence racemose, sometimes paniculate or fasciculate, the bracts stipuliform, the pedicels 6-12 mm long, hirsutullous. Flowers whitish yellow, 15-25 mm long; calyx campanulate, symmetrical, sparse- ly setaceous, 5-6 mm long, the tube 4—5 mm long, 3-4 mm in diameter, the lobes deltoid, acute, 1.0-2.5 mm long, usually ciliate; the standard broadly obovate, 10-15 mm long, ca. 15 mm wide, cleft to shallowly denticulate, the wings ca. 15 mm long, ca. 5 mm wide, obovate to oblong, the claw 1-2 mm long, the keel ca. 15 mm long, nearly straight, the claws 1-2 mm long; stamens monadelphous, connate into a sheath, split above, the filaments glabrous, the anthers uniform, versatile. Fruit a loment, 12-15-articulate, 6-8 cm long, 7-10 mm wide, linear, compressed, puberulent to subglabrous, margins longitudinally striate, centrally reticulate-striate, the stipe 5-7 mm long, the articles 4-5 mm long, 7-10 mm wide; seeds 2.5-3.0 mm long, ca. 1.5 mm wide, smooth, reddish brown. This species ranges from Costa Rica to southern Ecuador. It is readily distin- guished from the other species occurring in Central America by its 5-foliate leaves and wide fruits. In the province of Esmeraldas, Ecuador, this species is called ""chupa-chupa,"' and its leaves are used to treat skin eruptions. No local names or uses have been recorded from Panama. Though the type specimen in Jussieu’s herbarium is marked "'Peru," it was probably collected from present-day Ecuador, since no material has been re- corded from Peru. BOCAS DEL TORO: Region of Almirante, d 88 (F, NY, US). Changuinola Valley, Dunlap 369, 400 (both US). Río Teribé, near Quebrada Lukulon, Kirkbride & Duke 510 (MO, NY, SCZ). NA et al. 12731 (FSU). 1 km W of Gamboa, Nee 9439 (GH, MO). Empire to Mandinga, Piper 5155, 5165 (both US). Around Culebra, Pittier 2212 (NY, US). Las Cascadas Plantation, near Summit, Standley 25813, 29540 (both US). Balboa, Standley 27158, 32153 (both US). Gamboa, Standley 28322, 28452 (both US). Darién Station, Standley 31592 (US). cuiRIQUí: Burica Peninsu la, 6 mi S of Puerto Ar- muelles, Busey 578 (MO); Liesner 425 (MO). cocié: 10 mi E of Nata at Río Grande, : son 5281 (FSU, SCZ). DARIÉN: Santa Fé, Duke 9478 (MO, NY, US). Teotuma, Duke 10059 (MO). Río Sabana, ca. 4 mi above Santa Fé, Duke 10216 (MO, NY). Near Canglón river, Duke & Bristan 355 (MO). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 581 MOY URS hy =. Ñ. NES NN =, Xm E — Al FIGURE 11. Chaetocalyx latisiliqua (Poir.) Hemsl.—A. Habit ( g; O £e < [5] v - © gs = ° = < [od d. = e. e £e e y a e = Ic] m n = ° Lad E © Ee ^ E = [el c = e e e S Q = w 1 m=. o O ° = - = O ° B ° = Articles of the loment almost orbicular or saccate in outline, essentially flat, notched on the dorsal suture, not filled by the seed. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 625 d. Articles small, almost as broad (high) as long,?? 10-12 mm broad (includ- ing notch), 12-14 ee long, notch on dorsal suture narrowly V-shaped, article + beaked at the termini of the dorsal suture; stipe + equal to pedicel in length, PS NN glabrous -------------------------- 11. D infractum dd. Articles of the loment much ih usually 2 cm or more long and high; ob I 2 tomentose on abaxial surface; stipe of loment pubescent. f. icles with flattened ud when mature; sn about nee to the stipe in length ---------------- 3. D. macrodesmum ff. Articles with shoulders um partially, if at all, citened. be- coming beaked when fully mature; pedicel usually exceeding SUPE- Sa usu us u es AL HE 18. D. rhynchodesmum ee. Articles without flattened shoulders and not becoming beaked at maturity, the notch forming a sharply acute or obtuse angle; leaf- ea not suerte n abaxial surface; pedicel exceeding the gla- brate stipe in length -------------------------------------- 9. D. saccatum Articles of the pee smaller e not saccate or orbicular in outline, essentially filled by the seed when mature; the dorsal suture straight or only slightly con- cave or invaginated at the center. owers borne in pairs; each pair subtended by 1 large primary bract; each pedicel dieta subtended by a smaller secondary bract, or second- ary bract lackin Each pair of pedicels subtended by a wawa A IT each pedicel usually subtended by a smaller secondary bra s with inconspicuous pubescence q a hooked tri- pele leaflets short pilose on abaxial s j. Leaflets rhombic, rhombic ovate, or Dos kwy obtuse or rounded at apex; stipe of loment 3-4.6 mm long ------------------------ 4b. D. ax ite var. axillare j. Leaflets ovate, rather abruptly short acuminate; stipe of loment 4.6-8 mm long _____----------------------------- NE VE ANT et s c. D. axillare var. stoloniferum ii. Stems with long dense pubescence of straight trichomes; leaflets ovate or elliptic ovate, long p ilose abaxially, acute to gradually (usually long-) acuminate ---------------------------- Seek Soot A T EERIE AET a. D. axillare var. AWAQ hh. Each pair of pedicels subtended by a primary bract only; seco ary bracts lackin Primary bracts early deciduous, inflorescence lax; calyx pu- berulent and slightly short pilose along central tooth of lower lobe. l. Leaflets kame to 2.3 cm long and 1.5 cm wide; ar- ticles of the loment to 5.5 mm long .. 1. D Viro ad ll. Leaflets (terminal to 0.6 cm long and 0.7 cm wide; a ticles of the loment to 4.5 mm long -------- 23. D. c kk. Primary bracts persistent, inflorescence dense; calyx lon silky pilose and ciliate ------------------------------ 5. D. barbatum gg. Flowers borne singly (rarely in D. incanum moret than one in a fascicle), each pedicel subtended by 1 large primary and 2 smaller secondary bracts. Loments indie ut dorsal suture straight; isthmi broad or narrow; ma of leaflets not undulate. n. Isth sey ea between the articles) nearly pns length of article, loments to 7-articulate; leaflets thick, veins prominent ------------------- --- 10. D. incanum ° e 3? The length of the article in species of Desmodium sect. Nephromeria is measured as are those in other sections of the genus, along the long axis of the loment, assuming that more than one article may always occur. This conforms also to the long axis of the seed. The height of the article is, therefore, the width in this sense. 626 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 nn. Isthmi (area between the articles) narrow, loments p: 4- 2 ticulate; leaflets thin, veins not prominent. D. affine mm ibis 2-3-articulate, dorsal suture curved or Snare narrow; margins of leaflets undulate 24. D. wydlerianum bb. Pro with daa ur straight to curved, or obtusely to acutely angulate. Pe l oo. ° dicels borne in d of 2-several, each fascicle subtended by a narrowly ovate, greenish to stramineous primary bract; each pedicel further subtended by a similar slenderer sneer ee bract; young inflorescences with fascicles early distinct. Isthmi between articles central. Plants fey ieee articles slenderly elliptic, sutures scarcely con- q. stricted at isthmi _ 0 0 scorpiurus qq. Plants meg mostly t tall and usually becoming shrubby; artic les chiefly rhombic in outline (excepting occasionally the terminal), — markedly constric Terminal article of each loment regularly much dilated, re- niform to elliptic |... . glabrum IT. bine CH of each loment equaling the others in size an Leaflets thick and with prominent reticulate venation. t Pedicels stiff, ascending to spreading at maturity, borne in pairs ca. 1 cm apart, 1-1.6 cm long; ar- ticles of the loment 3-3.5 mm long, 2.6-3.5 mm wide __ TERRENI EIE ee in tortuosum Pedicels more slender, P borne in pairs less than 1 cm apart, 1 cm (or less) bor articles of the loment smaller, 1.5-2.5 mm long, 1.5-2 mm wide. u. Leaves dece br ipu abaxial surface rough; pedicels m long; at least the terminal pra 3) ae glabrous |... Rue PERDER D . D. distortum uu. Leaves regularly unifoliolate, abaxial sur- face velvety tomentose; pedicels 2.5-3 mm long; all articles of the loment with so Lad > straight pubescence |... 1. . D. Viu ss. ias ra not KEPA reticu P abou not coiled . procumbens var. procumbens Pedicels longer, usually 1.2-2 cm long, flexuous, filiform; bracts persistent; less EAE to be tightly coiled ______ 17a. D. procumbens var. longipes < < pp. Isthmi between articles eccentric —. 1 F D. campyloclados Pedicels borne in ars each pair subtended by a broadly ovate, stramineous olivaceous primary bract; each pedicel usually further subtended by a smaller, sometimes differently shaped and much thinner and paler, i.e., depauperate, secondary bract; young inflorescences with bracts densely imbrica w sthmi between articles centra °: . D. maxonii cels short (2.5-6 mm long), reflexed or erect at maturity, leaf- pie ometimes lustrous y. Stems and et sericeous tomentose; — strongly re flexed in fruit... . seric ophyllum yy. Stems los, leaflets lustrous above; ae erec maturity -ooa 6. D. cajanifolium Xx. Pedicels longer (mostly 1—2 cm long) or, if not longer, then i pedice not sharply reflexed in fruit and leaflets not lustrous a Z. Senden: to erect plants with stout trisulcate ine mM EE PRIORA PONDUS MICRO RR 2. D. intortum zz. Prostrate or decumbent plants with slender terete stems _ "Tc PPM zz: 15. D. mollic ulum 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 627 1. Desmodium adscendens (Swartz) DC., Prodr. 2: 332. 1825. Hedysarum adscendens hee Prodr. 106. 1788; Fl. Ind. Occ. 3: 1263. 1806. TYPE: Ind. Occ. Swartz (S, holotype; GH, phot Hedysarum adscendens var. UA caeruleum Lindl. Bot. Reg. tab. 8/5. 1824. LECTOTYPE: Bot. Reg. tab. 815. TUR NS adscendens var. (8 ?coeruleum (Lindl.) E » Prodr. 2: 332. 1825. Meibomia adscendens (Swartz) Kuntze, Rev. Gen. Pl. 1: 195. 1891, pro parte. Herbs, 6—24 inches tall, repent or forming tussocks, the stem terete, striate with stiff, upwardly appressed to spreading pilosity. Leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate lanceolate with long attenuate apex and slightly auriculate base, 0.5-1 cm long, 1.5-3 mm wide, striate, ciliate, the abaxial surface glabrescent, persistent; petioles sulcate adaxially, pilose, 1-1.8 cm long; leaf rach- is similar to the petiole, 0.3-0.8 cm long; stipels linear attenuate, ciliate, 1-2 mm long; petiolules stouter than leaf rachis and with stiffer pilosity, 1-3 mm long; leaflets elliptic obovate, acute to obtuse and emarginate at the apex, cuneate to rounded at the base, densely pilose on both surfaces, the terminal leaflet 1.7—4 cm long, 0.75-2 cm wide, lateral leaflets 1.3-3 cm long, 0.7-2 cm wide. Inflo- rescence of terminal and axillary racemes, rachis striate to grooved, sparsely to abundantly stiff pilose and finely puberulent with multicellular and uncinulate trichomes; primary bracts ovate acuminate with long attenuate apex, striate, cil- iate, pilose and puberulent on abaxial surface, early deciduous, 3.5-5 mm long, 1.5-2 mm wide, each subtending 2 pedicels, secondary bracts absent or, if (rarely) present, depauperate; pedicels with pubescence similar to that of rachis, 0.5-1 cm long. Flowers with the calyx distinctly bilabiate, the tube minutely puberulent throughout, teeth of both lobes pilose along each midrib and at the apex, teeth of lower lobe 2-3 mm long, upper bifid lobe 2-3 mm long; corolla usually pink, standard obovate, short unguiculate, 4-5 mm long, 2.5-3.5 mm wide, wings ob- long, broader above, short unguiculate, slightly auriculate, 3-3.5 mm long, 1-2 mm wide, keel petals triangular, truncate at the apex, gradually narrowed to a long claw, 3.5-4 mm long, 1 mm wide. Loment stipitate, stipe 0.5-2 mm long, 1—5-articulate, articles oblong, straight on the upper suture, curved below, con- stricted about 4 their width at the isthmi, uncinulate pubescent and puberulent throughout, 3.5-5.5 mm long, 2.5-3 mm wide; seed transversely elliptic, 2.5 mm long, 1.5 mm wide. Desmodium adscendens is widespread in tropical areas of Asia, Melanesia (see Ohashi, 1973), Africa and America. With Desmodium incanum and D. scor- piurus, it is one of the most common species. There is a much larger synonymy for Desmodium adscendens than here in- dicated. It has not, however, entered the literature of Panama. CAS DEL TORO: Ojo del Agua ca. 7 km W of Almirante, 190—220 m, Croat 38133 (MO). an d Valley, Dunlap 214 (US). Río Teribe, mostly 2nd growth near Quebrada Lukulon, : m, Kirkbride & Duke 522 (SCZ). Zigla amsa at junction of Changuinola and Teribe Rivers, Lazor e 2 SA (FSU, MO, SCZ). CANAL ZONE: BARRO COLORADO ISLAND: Croat 5904, 5951, 5989 (all MO). 96 (A, MO), 8509 (MO), 10763 (MO, SCZ), 11060, 11165 (both MO), 11727 (MO, SCZ), 16569, Pid (both MO); Dodge 3479 (G, GH); Ebinger 21 (GH, MO); Luteyn 1348 (DUKE); Netting 33 MO); Robyns 65-22 (MO); Shattuck 142 (MO). Summit Gardens, Bottimer 1508 (DAO). Frijoles, Bottimer 1496 (DAO). Chagres, Fendler 77 (K). 5 mi E of Ferry Thatcher bridge near mangrove swamp, Lazor 3311 (FSU, SCZ). Pipeline road, 16.8 km from beginning of road; 0-100 m, Mori & 628 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Kallunki 1765 (MO). Trail between Gamboa and Cruces, 50-80 m, Pittier 3769 (G, GH, US). Frijoles, Pittier 6835 (G, GH, US). Old Fort San Lorenzo, Tyson 1559 (SCZ). Pipeline road, Witherspoon & Witherspoon 8287 (MO). CHIRIQUI: Roadside above Princess Janca finca, Boquete, D'Arcy & D'Arcy 6334 (MO). Volcán/Cuesta de Piedras, site 84, 1000-1200 m, Burt & Koster 159 (MO). Boquete Dist., 3800 ft, Davidson 711 (GH, MO, US). Just W of the Fortuna camp, 1400-1600 m, Folsom et al. 5373 (MO). Between Concepción and Hato de Volcán, 1080 m, Koster 159 (A). Nueva California, 4000 ft, Tyson 5707 (DUKE, FSU, MO, SCZ). Boquete, 1200-1 1500 m, Woodson & Schery 719 (G, GH, MO, US). Finca Lérida to Boquete, 1300-1700 m, Woodson et al. 1166 (G, GH, MO, US). cocrÉ: Road past Furlong's Finca, N of Cerro Pilon, 880 m, Croat 37512 (MO). Road to El Valle de Anton, D’Arcy & D'Arcy 6717A, 6721, 6742 (all MO). "Top of El Petroso, N of El Copé, 800 m, D'Arcy 11336 (MO). Slopes of Cerro Pilón near El Valle, 700-900 m, Duke 12189 (MO, OS). Foothills of Cerro Pilon near El Valle, 900 m, Duke & Correa 14701 (N Y). Summit of Cerro Pilón above El Valle de Antón, 2700 ft, Dwyer et al. 4496 (MO). Boca del Toabré at confluence of Rió Toabré and Río Coclé del Norte, Lewis et x See (MO, SCZ). El Valle, Ebinger 951 (MO), 1121 (GH, MO). Summit at Alto dein 900 m, m & Robinson 2394 (MO). El Valle de Antón, 1000-2000 ft, Lewis et al. 2610 (M 3100 ft, nei 4054 (DUKE); 2000 ft, Porter et al. 4592 (MO); Wilbur & Luteyn 11755 one Bismark above Penonomé, Williams 139 (US). COLON: Miguel de la Borda, Croat 10044, 10054 (both & Weaver 11178 (DUKE). DARIEN: Río Pirre near crossing of trail from El Real to Tucutí, 20 mi W US). Los saNTOs: Loma Prieta, 800-900 m, Duke 11880 (MO). From 1 mi S to 10 mi N of Tonosi, Duke 12480 pr OS). PANAMA: Cerro Campana, 2400-2700 ft, Duke 8694 (MO). Between Canasas and Sabalo, 100 m, Duke 14477 (MO, NY, OS). Chimán, Lewis et al. 3300 (GH, MO). Cerro Azul, Tyson 2175 (FSU, SCZ). san BLAS: Hills SE of Puerto Obaldía, Croat 16723 (A), 16741 (MO). Puerto Obaldía, Croat 16867 (MO), 16875, 17009 (both A). Trail E of Cangandí-Mandinga airport road, 2-5 mi S of Mandinga airport, Duke 14809 (NY). Puerto Obaldía, 0 to 50 m, Pittier 4368 (US). Molia, Stier 47 (MO). VERAGUAS: Río Primero Braso, 2.5 km beyond Agriculture School Alto Piedra near Santa Fe, 700-750 m, Croat 25473 (MO). 0.6 mi beyond Escuela Agricola Alto Piedra, 730 m, Croat & Folsom 34001 (MO). Mouth of Río o Concepción, Lewis et al. 2825 (GH, MO, UC). Base of Cerro Tute, 6.5 km outside of Santa Fé, Folsom 3008 (M 2. Desmodium affine Schlecht., Linnaea 12: 312. 1838. TYPE: Mexico, Veracruz, Prope Jalapam et Hacienda de la Laguna, Schiede. Not seen. Desmodium albiflorum Salzm. ex Benth. didt Marea, Meddel. Dansk Naturhist. Foren. Kjgb- en vene 1853: 17. 1854, vielen prin nth. n Mart. Fl. Bras. 15(1): 99. 1859. TYPE: Brazil, a, Salzmann (K, not seen; phot MDC affinis (Schlecht) Kune, Rev. Gen. i i i 1891. M. albiflora (Benth.) Kuntze, Rev. Gen. Pl. 1: Procumbent woody herb with erect peduncles; stem uncinulate puberulent and moderately long spreading pilose. Leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate acuminate, striate, puberulent on abaxial surface, ciliate, persistent, 4-6.5 mm long, 2-2.5 mm wide; petioles sulcate on adaxial surface; pubescence similar to that of stem, 2.6-3.2 cm long; the leaf rachis similar, 4 or less the length of petiole; stipels lance attenuate, striate, puberulent and ciliate, persistent, 2.5-4 mm long; petiolules stouter than the leaf rachis, densely spread- ing pilose, 1-2.5 mm long; leaflets thin, uncinulate puberulent and sparsely scat- tered pilose on adaxial surface, paler and with veins more prominent on the sparsely uncinulate puberulent and pilose abaxial surface, the margins ciliate at least near base, the terminal leaflet ovate, acute to obtuse, the base rounded to deltoid, 3.6—4.8 cm long, 2.3-3.2 cm wide, lateral leaflets obliquely oval, acute, 2.9-4 cm long, 1.7-2.5 cm wide. Inflorescences axillary and terminal; the rachis angulate, uncinulate puberulent, remotely if at all pilose; primary bracts ovate 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 629 acuminate, striate, puberulent and ciliate, not long persistent, 1.5-3 mm long, 1 mm wide; secondary bracts similar, 1-1.5 mm long, 0.5 mm wide; pedicels borne singly, each subtended by 1 primary bract and 2 secondary bracts (borne later- ally), densely uncinulate puberulent, 8-11 mm long. Flowers with the calyx bi- labiate, uncinulate puberulent, pilose on the teeth of both lobes, the teeth of lower lobe acuminate, almost equal, 3.5—4 mm long, the upper bifid lobe 3.5—4 mm long; corolla somewhat longer than the calyx, standard obovate, narrowly cuneate at base, 5.5 mm long, 4 mm wide, wings oblong, unguiculate, 6 mm long, 1.5 mm wide, keel petals oblong to remotely scythe shaped, long unguiculate, 5.5-6 mm long, 1.5-2 mm wide. Loments short stipitate, the stipe 1.5-2 mm long, to 5-articulate, articles oblong, the upper suture straight, the lower curved, con- stricted at the isthmi, articles densely uncinulate puberulent, 5.5-6 mm long, 2.5— 3 mm wide; seed transversely elliptical, 3 mm long, 1.5 mm wide, the funicle short. This species occurs in southern Mexico, Central America, the West Indies, and the northern half of South America. The above noted seed measurements were taken from Standley 28941 (GH), Honduras. CANAL ZONE: Balboa, Standley 25417 (US), 26076 (MO, US), 29262 (US). 3. Desmodium angustifolium (H.B.K.) DC., Prodr. 2: 328. 1825.—Fic. 20G. Hedysarum uan H.B.K., Nov. Gen. Sp. Pl. 6: 404 (folio ed.); 6: 517, 518 (quarto ed.). 1824. TYPE: Mexico (P; photo GH). Desmodium linearifolium DC., Prodr. 2: 327. Ls Mem. Légum. 325. 1826, non G. Don, Gen. Syst. : : 2. TYPE: "ex itin. Nee in hb. T Meibomia angustifolia (H.B.K.) Kuntze, xa “Gen, Pl. 1: 197. 1891, non M. angustifolia (T. & G.) Kearney, Bull. Torr. Bot. Club 20: 481. = linearifolia (DC.) Kuntze, Rev. Gen. Pl. I oe Desmodium angustifolium var. typicum Schubert, a Gray Herb. 129: 27. 1940. Tall, erect, somewhat woody herbs to almost 3 m high; stem angulate, gla- brescent to densely uncinulate puberulent. Leaves unifoliolate, stipulate, petio- late; stipules early deciduous, ovate attenuate (the slender tip almost !^ the total length), striate, puberulent, stiffly ciliate, 3-9 mm long, 0.7-2 mm wide; petioles striate, minutely puberulent, 0.15-1 cm long, stipels lance attenuate, persistent, 1-3.3 mm long, petiolules puberulent and stiffly pilose, 0.5—1.7 mm long; leaflets sparsely to densely uncinulate puberulent on both the surfaces and sparsely stri- gose chiefly on the midrib and the veins of abaxial surface, linear lanceolate, acute and mucronate at the apex, acute at the base, (3—)8—16 cm long, 0.2—0.6 cm wide. Inflorescence racemose paniculate, rachis uncinulate-puberulent and somewhat stiffly pilose; primary bracts each subtending a fascicle of 2—several pedicels, ovate acuminate, striate, puberulent on the abaxial surface, stiffly cil- iate, 1.7-3.5 mm long, 0.5-1 mm wide, each pedicel further subtended by a similar smaller lance acuminate secondary bract 0.8-2 mm long and 0.4-0.5 mm wide; pedicels pubescent as on the rachis, 4-7 mm long. Flowers with the calyx pu- berulent and stiffly pilose, the margins of the teeth of both lobes stiffly ciliate, the upper bifid lobe 2-2.4 mm long, lateral teeth of lower lobe 2-2.3 mm long, 630 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 20. Desmodium.—A-F. Desmodium axillare (Sw.) DC. var. acutifolium Sunize) Urb. d Habit (M) —B. delen a —C. Corolla m Standard (x4).—C*?. Wing petal (x4).—C? Keel petal (x4).—D. Stamens.—E. Pistil.—F. Fruit (x2). [After Erlanson 296.]—G. Desmodium plea es re (H.B.K.) DC. Fruit. Note eee with the rim separating. [After Allen 1269.] the central tooth 2-2.5 mm long; corolla with the standard flabellate and short unguiculate, 3—4.7 mm long, the wings oblong, slightly auriculate and short un- guiculate, equaling or slightly exceeding the standard in length, the keel petals less markedly auriculate, longer unguiculate than the wings, 3.4-5 mm long and 1-2 mm wide. Loment short stipitate, 3-8-articulate, the stipe 0.5-2 mm long, the articles essentially orbicular, uncinulate puberulent on the surfaces and stiffly 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 631 pilose on surfaces and suture, 3-4 mm long, 2-2.5 mm wide, the walls of the articles often dehiscing from the suture which remains essentially intact; seed transversely oblong, 1.8 mm long, | mm wide. This species ranges from Mexico and Central America to northern South America. The characteristic falling away of the walls of the articles of the loment, often leaving the suture of the whole loment intact, distinguishes Desmodium angus- tifolium. This character occurs frequently in species of Mimosa but has been observed in only one other American species of Desmodium. Desmodium gramineum Gray, which was treated by me (1940) as a variety of D. angustifolium is not related to this species, and my early treatment of D. gramineum in this way should not be followed. ANAL ZONE: Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, ey 5181 (US). Sosa Hill, Balboa, Standley WG (MO, US), and 26441 (US). cuiRiQUÍ: from Boquete to 3 mi N, 3300-4300 ft, Lewis et al. 649 (MO). Sabana de la Tortuga between Coa gu - Caldera, 300-700 m, Pittier 5733 (US). HERRERA: pe S. Ocu, Tyson et al. 2877 (MO, SCZ). P MA: San José Island, Johnston 370 (GH, US), 1269 (BM). El Valle, 580 m, ud 29 (A); Pisis: 204 (K), Seemann (BM). Las Sabanas, Standley 25898 (MO, US). Between Las Sabanas and Matías Hernandez, Standley 31869 (US). VERAGUAS: Above Río Santa Maria along road to Santiago, 5 km S of Santa Fé, 250-300 m, Nee 8112 (MO). PROVINCE UNKNOWN: Duchassaing (P). 4. Desmodium axillare (Swartz) DC., Prodr. 2: 333. 1825. Herbs, the stem repent and rooting at the nodes, the internodes 1.3-11 cm long. Leaves trifoliolate, stipulate, the stipules connate up to ! their length, early deciduous; petioles to 10 times the length of the leaf rachises, leaf rachis to 12 mm long or occasionally lacking; petiolules of the lateral leaflets usually some- what longer than those of the terminal leaflet. Inflorescence racemose, the ra- cemes arising from the leaf axils; bracts early deciduous; calyx uncinulate pu- berulent over the whole surface with stout straight trichomes along the teeth of both lobes. Flowers with the corolla always exceeding the calyx; standard ob- ovate or more rarely ovate, broadly acute below or narrowing slightly to an obtuse or truncate base, rounded at apex, retuse, the wings obliquely obovate, truncate or obtuse at apex, auriculate at base, short clawed, the keel petals fused, more or less falcate, obtuse at the apex, broadly clawed. Loment 1-3-, mostly 2-articulate, stipitate, the upper suture continuous, essentially straight. This species is composed of three well defined, easily distinguishable vari- eties all of which occur in Panama. 4a. Desmodium axillare var. acutifolium (Kuntze) Urb., Symb. Ant. 4: 292. 1905.— Fic. 20A-F. Meibomia axillaris (Swartz) Kuntze var. B. acutifolia Kuntze, Rev. Gen. Pl. 1: . 1891. Nephromeria axillaris eiu, B. acutifolia [‘acutifoliola’ ] (Urb.) Schindl., E Spec. Nov. Regni Veg. 20: 284. 5 Desmodium axillare var. já angustatum Urb., Symb. Ant. 2: 303. 1900. D. axillare var. B. angustatum fo. peris Urb., Symb. Ant. 2: 303. 1900. D. axillare var. B. acutifolium fo. robustius (Urb.) Urb., Symb. Ant. 4: 292. 1905. Based on D. axillare var. B. angustatum fo. robustius inka kina: aans var. 8. on (Urb. ) Schindl. fo. robustior (Urb.) Schindl., Repert. Spec. Nov. Regni Veg. 20: 284 632 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Hedysarum oblongifolium Bertero ex DC., Prodr. 2: 332. 1825, pro syn. Desmodium oblongifolium Bertero ex DC., Prodr. 2: 332. 1825. Meibomia umbrosa Britt., Bull. Torrey Bot. Club 37: 353. 1910. M. prorepens Blake, Contr. U.S. Natl. Herb. 24: 6. 1922. The variety acutifolium is characterized by the long dense pubescence of its stems in contrast to the short uncinulate pubescence on the stems of the other 2 varieties, and the ovate to elliptic ovate, usually long acuminate leaflets densely long pilose on the abaxial surface. It is found more or less throughout the range of variety axillare but more rarely. This variety occurs in Central America, the West Indies, Colombia, British Guiana, and Brazi For citation of type material and some discussion of the synonymy see Schu- bert 1963. CANAL ZONE: Ft. Kobbe, Bottimer 1446 (A, DAO). Barro Colorado Island, Bottimer 1481 (DAO); Croat 5414, 6353, 7063, Croat 11797 (all MO); Ebinger 22 (GH, MO); Kenoyer 384 (US); Shattuck 1046 (GH, MO). Pipeline Road near Gamboa, Clewell & Tyson 3209 (FSU, MO), 32/0 (MO). Behind Farfan Beach, D'Arcy & D'Arcy 6087 (MO). 6 mi E of Chepo on Panamerican Highway, Duke 4074 (MO). Ft. Kobbe, Duke 4703 (MO). U.S. Army Tropic Test Center, Albrook, Dwyer 6735 (MO). Gamboa, Naval Reservation, Ebinger 488 (GH, MO). Rio Chagres Bridge, ca. 2 km by road SE of Gamboa toward Balboa, Lass dis 4269 (MO). 2 mi N of Paraíso on Balboa Highway, side of hill overlooking Gaillaird Cut, Lazor & Blum 5321 (SCZ). Altos del Río Pacora, 2500 ft, Lewis et al. 2300 (GH, MO). Ancón Hill, ane 25172 (US). 1 mi N of Summit Garden, Tyson & Blum 1980 (MO). Across highway from Summit Garden, Witherspoon & Witherspoon 8696 (MO). CHIRIQUÍ: Between 2600 ft, Vaughan et al. 601 (MO). COLON: Colón Kuntze (NY); Colón Highway Bridge on Chagres River, Lazor & Tyson 3029 (MO), 3063 (FSU, MO). DARIEN: Gold mine at Cana, 500—600 m, Croat Minas and 14 mi S of Ocá, Wilbur et al. 12114 (DUKE). PANAMÁ: Roadside near Chepo, Dodge 10717 (MO). 5- 6 mi E of Chepo on Panamerican Highway, Duke 4045 (GH, MO). Río Pita, 1-2 mi SANT (MO). PROVINCE UNKNOWN: Between Panama and Sabanas, pe 114 (BM, MO, US). 4b. Desmodium axillare (Swartz) DC. var. axillare. Desmodium axillare (Swartz) DC., Prodr. 2: 333. 1825. Hedysarum axillare Swartz, Prodr. 107. 1788; Fl. e. Occ. 3: 1274. 1806. Meibomia m (Sw.) ae Rev. Gen. Pl. 1: 1891 D. axillare var. a genuinum Urb., Symb. Ant. 2: a 1900. Nephromeria axillaris (Sw. Schindl.. Repert. Spec. Nov. s= Veg. i e 1924. Meibomia axillaris var. a. obtusifoliola Kuntze, Rev. Gen P Ao, D. axillare var. a obtusifoliola (Kuntze) Urb., Symb. Ant. 4: Nephromeria axillaris var. a. obtusifoliola (Kuntze) Sa apie Spec. Nov. Regni Veg. 20: 284. 1924 Hedysarum reptans Poir. in Lam. Encycl. Méth. i 6: 422. 1804. Meibomia reptans (Poir.) Kuntze, Rev. Gen. Pl. 1: 198. 1891. D. radicans Macfad., Fl. Jam. 1: 269. 1837. Based on Hedysarum axillare Sw Meibomia andina Rusby, Mem. Torrey Bot. Club 3: 21. 1893, pro parte, as to isotypes (GH, MO, US), not as to holotype (NY). This, the most common and widespread of the three varieties of Desmodium axillare, is distinguished by its rhombic ovate, or rhombic orbicular terminal leaflets, and by the short stipes of its loments (3—4.6 mm) which are exceeded in length by those of the other two varieties. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 633 Desmodium axillare occurs - Se Central America, the West Indies, and the northern 25 of South Ameri For photographs and “maana of the holotypes of Hedysarum axillare and H. reptans see Schubert 1963. BOCAS DEL TORO: Almirante, Barrus 404 (GH); Carleton 68 (GH, NY, US), Carleton 102 (US). Hill above RR station at Milla 7.5, Croat & Porter 16384 (MO). 1.5 mi SE of Banana Plantation of Changuinola, Lazor et al. 2458 (FSU, SCZ). Junction of Rios Changuinola & Terebe, 100—200 ft Lewis et al. 928 (MO, UC). Chiriquicito to 5 mi S along Río Guarumo, Lewis et al. 2138 (MO, UC). Without other location, Wedel 225 (GH, MO), Water Valley, Wedel 621 (GH, MO). CANAL ZONE: Barro Colorado Island, Dodge 3463, 3478 (both GH). Panamerican Highway, Río Canita, Duke 3837 (GH, MO). Near Gatün Dam, Foster & Chong 1352 (DUKE). Gatün Station, Hayes 62 (G), 232 (NY). Gamboa, 20-100 m, Pittier 3763 (NY, US), Pittier 4439 (US). Las Cascadas Plantation near Summit, Standley 25792 (US). cHIRIQUí: San Bartolo Limite near Costa Rican border, 12 mi W of Puerto Armuelles, 400—500 m, Croat 22174 (MO). San Bartolo Arriba W of Puerto Armuelles, 125 m, f 26724 (MO). N side of Cerro Pando, 6000 ft, D'Arcy & D'Arcy 6658 (MO). Quebrada Melliza, mi S of Puerto Armuelles, 0-150 m, Liesner 453 (MO). Between Río Tinta and Río Tabasara, Sedis et al. 420 (MO). cocLÉ: 44 km N of Penonomé on road to Coclesito, 300—500 p Hammel 1668 (MO). DARIÉN: Path from Canazas to Tortí, border Darién and Panamá, Folsom 4962 (MO). Upper Río Tuquesa, Le Clezio 208 (MO). El Real, Río Tuira 1 mi down from Pinogana near Quebrada Chuito, Stern et al. 104 (MO). Bridge over bi ee downto from zug on Panamer- ican Highway, Stern et al. 151 (GH). H : Road between Las as and Pesé, 600 of Fla. "n hn S eae (NY). [Variety uncertain: only ss a availa no stem or leaves.] r Tapia River, Juan Diaz region, Maxon & Harvey 6660 (US). PROVINCE UNKNOWN: Bohio (ed Cowell 241 (NY, US). 4c. Desmodium axillare var. stoloniferum (Rich. ex Poir.) Schubert, J. Arnold Arbor. 44: 289. 1963 Hedysarum stoloniferum Rich. ex Poir. in Lam., Encycl. Méth. Bot. 6: 421. 1804. Desmodium spirale B. stoloniferum (Rich. ex Poir.) DC., Prodr. 2: 333. 1825 D. stoloniferum (Rich. ex Poir.) Steud., Nom. Bot. ed. 2. 1: 4 0 D. axillare var. y. sintenisii Urb., Symb. Ant. 2: 303. 0. Meibomia sintenisii (Urb.) Britt. in Britton & Wilson, Sci. Surv. Porto Rico & Virgin Isl. 5: 402. 1924. Nephromeria axillaris var. y. sintenisii (Urb.) Schindl., Repert. Spec. Nov. Regni Veg. 20: 284. 1924. D. axillare var. sintenisii sensu Schubert, Contr. Gray Herb. 135: 84-86, pl. 1, figs. A1—7. 1941. Meibomia albida Blake, Contr. U.S. Natl. Herb. 24: 5. 1922 The variety stoloniferum is intermediate in aspect between var. axillare and var. acutifolium. From both these varieties it may be distinguished by its ovate, abruptly short acuminate leaflets, and from var. axillare by the denser pilosity on the abaxial surface of the leaflets as well as by its longer pedicels and loment stipes. Central America, the Greater Antilles, and the northern half of South America. CANAL ZONE: Barro Colorado Island, Croat 6290, 6630 (both MO), 10214 (MO, SCZ), 11254, 11264, 11311, 11828, 14019, 14982, 15118 (all MO); Foster 828 (DUKE); Knight (WIS); Netting 47 (MO); Starry 168 (MO); Wilbur & Weaver 10812 (DUKE, GH, MO). Behind Farfan Beach, D’Arcy ; i io Cani í (MO). ie ad bar. Bartolo Rh e sula de Buri -50 m Woodson & Schery 673 (MO, . COLÓN: bridge crossing ot y endis abes ay over Río unu unaque, down river from DN Stern et al. 151 (MO). LOS SANTOS: E Priet Mog Grande, 2400-2800 ft, Lewis et al. 2210 (MO). PAN- AMA: Río Sancanti, 2 mi upstream bn Piria 120 m, Duke 14384 (NY). From Torti to the Pilota del Toro, mountain overlooking Torti Arriba, Folsam et al. 5098 (MO). Juan Díaz, Killip 3161 (US). SAN BLAS: Between Cangandí and Mandibsa Indigena, Duke 14756 (NY, OS). 634 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 5. Desmodium barbatum (L.) Benth. in Miq., Pl. Jungh. 224. 1852; Benth. & Oerst. in Vidensk. Meddel. Dansk Naturhist. Foren. Kjgbenhavn. 1853: 18. 1854 Hedysarum barbatum L., Syst. Nat., ed. 10. 1170. 1759. TYPE: herb. Linnaeus (LINN 921.48; mi- crofiche A-GH). Perrottetia barbata (L.) DC., Ann. Sci. Nat. 4: 96. 1825. Nicolsonia barbata (L.) DC., Mém. Légum. 313. 1826. Urania barbata (L.) Desv., Mém. Soc. Linn. Paris 4: 309. 1826. Meibomia barbata (L.) Kuntze, Rev. Gen. Pl. 1: 195. 1891. Herbs, prostrate to ascending or erect with white to fulvous appressed to spreading pilosity on stems, branches, petioles, and rachis of the inflorescence. Leaves usually trifoliolate, stipulate, petiolate; stipules conspicuous and persis- tent, deltoid attenuate, slightly auriculate at the base, striate and ciliate, some- what pilose to glabrescent on abaxial surface, 6-10 mm; petioles sulcate on adax- ial surface, 0.8-2.2 cm long, the leaf rachis similar, 0.2-0.7 cm long; stipels slender, ciliate, persistent, 1.5-4.5 mm; petiolules somewhat stouter and more densely pilose than leaf rachis, 1-2 mm long; leaflets almost orbicular to oval, elliptic obtuse or obovate, usually mucronate, often retuse, sparsely appressed pilose adaxially, densely so and with venation prominent abaxially, terminal leaf- lets 1.4-5 cm long, 0.5-2 cm wide, the lateral leaflets 1-3.8 cm long, 0.4-1.3 cm wide. Inflorescence of axillary and terminal short, congested, densely flowered racemes somewhat obscured by the long pubescence of bracts and calyces; bracts long persistent, ovate acuminate, striate, pilose to glabrescent on abaxial surface, ciliate, 5-7 mm long, 1.5-2 mm wide, secondary bracts lacking; pedicels uncin- ulate pilosulous, 3.5-9 mm long. Flowers with the calyx becoming reflexed, sub- equally 5-lobed, silky pilose, the lobes ciliate, 4-6 mm long; standard obovate, somewhat unguiculate, 4-5 mm long, 2-3.7 mm wide, the wings oblong, obtuse, short unguiculate, 3-4 mm long, 1-1.5 mm wide, the keel petals scythe shaped, truncate at the apex, long unguiculate, 3—4.5 mm long, 0.8-1 mm wide at the apex. Loment l—S-articulate, straight on the dorsal suture, somewhat curved below between the wide isthmi, uncinulate pubescent on the surfaces; articles separating partially first, the lower suture dehiscing later, the articles 2-3.5 mm long, 2-2.5 mm wide; seed subquadrate to quadrate, 1.5 mm long, 1 mm wide, pale tan. A polymorphic species widespread in tropical areas of the New World and in Africa and Asia. Many variations have been named but in general the characters cannot be correlated with geographical distribution. In Panama there seem to be two forms, one repent and one erect, but there are many intergradations between the two extremes. BOCAS DEL TORO: Almirante, Rowlee & Stork 1000 (NY, US). Chiriquí Lagoon, Wedel 1179 (GH, MO, US). Johns Creek, Wedel 2758 (GH, MO, NY, US). CANAL ZONE: 1 mi from Miraflores, Blum 1853 (FSU, SCZ). Summit Gardens, Bottimer 1507 (DAO). Venado Beach, Correa & Gonzalez 507 ). N i Dwyer 7167 (MO). Chagres, Fender 76 (K, MO, US). Pipeline Road, 2-4 mi N of Gamboa, Gentry 6536 (GH, MO). Empire Station, Hayes 585 (K). Paraiso Station, Hayes 799 (NY). Cerro Ancón, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 635 join 129 (GH, US). 8 mi N of 2 on Pipeline Road, Lazor 5276 (SCZ). V mi N of Gamboa, or & Tyson 5661, 5684 (both MO). 3 km S. Gualaca, McCorkle C-46 (FSU, SCZ). Near Gamboa dd Club, Mori 413 (MO). Ancón d Piper 5116 (US). Culebra, 50-150 m, Pittier 2142 (BM, C, NY, US), 4827 (NY, US). Seemann 229 (BM, K), [Province uncertain]. Sosa Hill, Balboa, Stan- dley 25273 (MO, US). Ancón Hill, Standley 26354 (C, US). Summit, Standley 26948 (US). Frijoles, Standley 27658 (US). Mt. Hope Cemetery, Standley 28788 (US). Cerro Azul, 2000 ft, Tyson 2128 (MO). Curundu, Tyson & Blum 2523 (FSU, MO). 1 mi from Summit on road to FAA radar tower, Tyson et al. 2771 (MO, SCZ). cHiRIQUí: Near Veladero, site oe 0-100 m Burt & Koster 125 (MO). Savanna just below Boquete, 800 m, D' Arcy 9769 MO). Lava fields near town of Volcán, 4600 ft, Duke 9215 (MO). From Boquete to 3 mi N, 3300—4200 ft, pn et al. 602 (GH, MO). Road to Gualaca 0-100 m, Rattray 125 (A). Boqueté Diiia Volcancitos, 5500 ft, Terry 1262 (GH, MO). cocLe: S of El Valle de Antón, 600-800 m, Allen 2810 (G, GH, MO, US); apes 1473 (A, DAO). Savanna 4- 6 mi E of Natá, 10-25 m, Duke 12388 (NY). ne Ebinger 995 (GH, MO); 50-1000 ft, Williams 128 (NY, US). COLON: Portobelo Road ca. 1 mi W of Port obelo, Gentry 1739 (MO, SCZ). DARIÉN: El Real, Burch et al. 1069 (GH, MO, UC). HERRERA: Near Ocú, 100-200 m, Burt & Koster 88 (MO). Between Panamerican Highway and Ocü, 100-200 m, Koster 88 (A). PA ANAMÁ: Pacora, ca. 35 m, Allen 990 (GH, MO, US). Cerro Campana, D'Arcy 9581, 9602 (both MO). Between Panamá and Chepo, Dodge et al. 16701 (MO). Cerro Campana, Duke 5949 (MO). Tocumen area, Dwyer et al. 7231 (GH, ied Las Sabanas, AG AP 138 (US). Tumba Muerto, Heriberto 176 (NY). Sabanas near Chepo, . Hunter & Allen 19 (MO). Chilibre, Jaén 45 (MO). San Jose Island, Johnston 27 (US, MO), 672, 826 (both GH). Taboga Is., 200 m, Killip 3187 (US). Between Tapía and Tocumen Rivers, 75 m, Killip 3239 (US). Cerro Campana, Lewis et al. 3131 (MO). Taboga Island, Longfield 490 (BM, K, O, US). Cerro Campana, 650—700 m, McDaniel 8138 (FSU). Pacora-Chepo highway, vicinity Pacora River, McDaniel 8179 (DUKE, FSU). El Llano-Cartí road, 5 km N of Panamerican Highway, 300 m, Nee 7929 (DUKE, MO). Sabanas, Paul 52 (US). Sabana de Juan Corso, near Chepo, 60-80 m, Pittier 7 (G, GH, NY, US). Matías Hernández, Pittier 6873 (GH, NY, US). Nuevo Emperador, Puga 30 (DUKE, MO). Road to Chepo near sea level, Semple Pa-1 (US). Big swamp east of Río Tocumen, Standley 26505 (US). Near Matías Hernández, Standley 28887 (US). Between Matías pon and Juan Díaz, Standley 32084 (US). Cerro Azul at 2000 ft, Tyson 2128 (FSU, SCZ). W slope Cerro Campana, 2500 ft, Tyson et al. 2347 (FSU, SCZ). Between Pacora and Chepo, 24 m, Woodson et al. — 0.200 m, Gentry 6341 (MO). vERAGUAS: Divisa/Santiago, 0-100 m Burt & Rattray 77, 81 (both MO). Near La Pena, 100-200 m alt., Burt & Koster 107 (MO). ps 10 mi W of Santiago, Duke 13646 (OS), A13647 MO). Panamerican Highway 1⁄2 km W of Hwy. 50 turnoff to Ocú., 30 m, Nee 8000 (MO). Above Río Santa María, road to Santiago, 5 km S of Santa Fe, 250-300 m, Nee 8130 (FSU, GH, MO). Near Santiago, 0-100 m, Rattray 81 (A). 6. Desmodium cajanifolium (H.B.K.) DC., Prodr. 2: 331. 1825. Hedysarum ot m H.B.K. Nov. Gen. Sp. Pl. 6: 410 (folio ed.); 525 (quarto ed.). pt 598. 1824. TYPE: Amérique Equatoriale, Herb. Humboldt & Bonpland (P, holotype; GH, photo). Hedysarum multiflorum Willd. ex. Schlecht. & Cham. Linnaea 5: 586. 1830; G. Don , Gen. Syst. 2: nomen nudum. TYPE: Colombia, Humboldt (B-Willd 13797; microfiche A-GH). Meibomia calaniolid (H.B.K.) Kuntze, Rev. Gen. Pl. 1: 195. 1891. Herbs or shrubs, not much branched, 2-4 m high; stem terete, striate, densely uncinulate pubescent and pilose. Leaves trifoliolate, stipulate, short petiolate; stipules obliquely ovate, long attenuate, striate, ciliate, pilose and puberulent, early deciduous, ca. 4 mm long and 2 mm wide; petioles narrowly sulcate, densely uncinulate pubescent and pilose, 0.7-1 cm long; the leaf rachis similar, 0.6— 0.7 cm long; stipels lance attenuate, striate, ciliate, to 3.5 mm long; petiolules densely pilose to 2.5 mm long; leaflets narrowly ovate, obtuse and mucronate at the apex, cuneate to obtuse at the base, lustrous on the adaxial surface, pilose along the midrib and with scattered puberulence, duller and rather densely pilose abaxially; terminal leaflets 5-9 cm long, 1.6-3.3 cm wide, lateral leaflets 3.3—6 cm long, 1.3— 2.5 cm wide. Inflorescence terminal and axillary, racemose to racemose pani- culate, much branched and densely flowered; rachis somewhat angulate, uncin- 636 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 ulate pubescent; bracts early deciduous, primary bracts ovate acumi- nate, striate, puberulent, ciliate, each subtending 2 pedicels, 2-3.5 mm long, 0.5- -5 mm wide; secondary bracts each subtending 1 pedicel (laterally), subulate, ciliate, to 1 mm long; pedicels slender, ascending, short spreading pilose, 5-6 mm long. Flowers with the calyx bilabiate, pilose and puberulent throughout, the teeth of both lobes ciliate; central tooth of lower lobe lanceolate to lance atten- uate, 3-4 mm long, lateral teeth acute to obtuse, 2.5-3.5 mm long, the upper lobe only slightly bifid, 2.5-3.5 mm long; petals, especially the standard, papillate, the standard obovate, retuse, narrowed to the cuneate base, 5.5-8 mm long, 2-3 mm wide, the wings oblong, auriculate, unguiculate, equalling standard in length and width (above claw), the keel petals obliquely oblong, long unguiculate, equalling standard in length, 1.5-2 mm wide. Loment to 6-articulate, stipitate, stipe pilose to 3.5 mm long, but not exceeding calyx, articles obtusely triangular, isthmi narrow and eccentric, margins pilose, surfaces uncinulate puberulent and pilose, 5-6 mm long, 4—4.5 mm wide; seed obtusely oblong, 2 mm long, 1.3 mm wide. This species is scattered in southern Mexico and Panama, the West Indies and the northern half of South America. The plants are handsome, easily distinguished by their lustrous leaflets, dense- ly flowered inflorescences, and short pedicellate flowers and loments. BOCAS DEL TORO: Carleton 217 (US). CANAL ZONE: W end of Gatun Lake Dam, Blum & Tyson 1973 (FSU, MO), B 1973 (SCZ). Frijoles, Bottimer 1487 (A, DAO). Gamboa, Bottimer 1620 (DAO). Between Goofy Lake in Cerro Azul and Main Highway, Correa & Dressler 441 (DUKE, FSU, 3 SCZ). Gamboa Navy Pipeline, Correa & Haines 531 (MO). Barro Colorado Island, Croat sn-213 (MO); Shattuck 561 (MO). 1 mi N of Paraiso near Gaillard Cut, Croat 12653 (SCZ). 1 mi N of Su i Garden, Croat 12890 (MO). Road to geologic test site north of Paraiso, Croat 12971 (MO). Gaillard Highway near Paraiso, Croat 13148A (MO). Frijoles, Foster 2056 (DUKE, GH, MO). Frijoles, Busey & Croat 251 (MO). Ancón, Greenman & Greenman 5 ( ). Gamboa, Greenman & Greenman 5170 (MO). Near Miraflores Locks, Greenman & Greenman 5182 (MO). Between Summit and Gam- boa, Greenman & Greenman 5253 (MO). Paraiso Station, Hayes 20 (NY). Summit road, Jones 276 (MO, US). 3 mi N of Paraiso on the Balboa Highway, Lazor & Blum 5324 (SCZ). Roadside % mi NW of Gamboa, 70 m, Liesner 1401 (MO). Along RR, 1% km W of Gamboa, 30 m, Nee 947 ). Headwaters of Río Providencia, 7.5 km SW of Gatun D , 25- m, Nee & Gentry 8654 (MO). to Mandinga, Piper 5192, 5193 (both US). Gamboa, Piper 5195 (SU). El Paraiso, 30-100 i Pittier 2519 (U a a, 40-80 m, Pittier 4808 (BM, etween Panamerican Highway and Cerro Campana, Porter et al. 4867 (MO). Cerro Gordo, near Culebra, Standley 26044 (US). Summit, Standley 26947 (MO, US). Frijoles, Standley 27659 (US). Gamboa, Standley 28336 (US). Old Las 22116 (MO). cocré: El Valle, 800-1000 m, Allen 85 (GH, MO, US); 600 m, Allen 2774 (G, GH, MO, US). Between Las Tibias and El Copé, Folsom & Collins 6426 (MO). COLON: Near Salamanca, 8 mi 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 637 7. Desmodium campyloclados Hemsl., Biol. Centr.-Amer. Bot. 1: 276. 1880. TYPE: Nicaragua. Segovia, Oersted (K, holotype; GH, photo). Meibomia campyloclada (Hemsl.) Kuntze, Rev. Gen. Pl. 1: 197. l. M. sylvestris Blake, Bot. Gaz. 78: 273. 1924. TYPE: E ama Huila, Cordillera Oriental, E of Neiva, Rusby & Pennell wee (NY, ME Acus phot Desmodium immerens Macbr., Field Mus. Nat. , Bot. i 11: 23. 1931. TYPE: Peru, Yanano, 6000 ft, Macbride 3725 d dote: GH. p "S D. "m Pittier, [Leguminosas de Vene uela]. gh Tecn. 5, Minist. de Agric. y Cria, Serv. Bot. 1944, without Latin diagnosis or citation of type. (See discussion below.) D. ur Pittier, Bol. Soc. Venez. Ci. Nat. 11: 17. 1947. (See discussion below.) Sprawling herb to slender straggling shrub, to 3 m or more, branches slender and elongate; stem and branches striate, soft pubescent with upwardly appressed pilosity. Leaves trifoliolate, stipulate, petiolate; stipules ovate attenuate, the slen- der apex equaling or exceeding the basal portion, stramineous, striate, pilose on the abaxial surface, ciliate, not persistent, 5 mm long, 1.5 mm wide; petioles sulcate, striate, with upwardly appressed pilosity shorter than that of stem, 2.5— 5 cm long; leaf rachis similar, 1.5-2 cm long; stipels lance attenuate, puberulent and ciliate, 2.5-3.5 mm long; petiolules darker, stouter than the leaf rachis, spreading pilose, 2-3 mm long; leaflets mucronulate, thin, moderately appressed pilose on both the surfaces, the terminal leaflet ovate, obtuse at apex, cuneate at base, 6—6.7 cm long, 3-4 cm wide; lateral leaflets similar or somewhat rounded At the base, 5-5.5 cm long, 2.5-3 cm wide. Inflorescence axillary, racemose, lax; rachis spreading pilose and puberulent; pedicels borne in bracteate fascicles of 3-many with 2 lateral flowers developing first, the central 1 following, etc.; bracts early deciduous, the primary bracts narrowly ovate attenuate, striate, ciliate, puberulent and pilose over the abaxial surface, 4—4.5 mm long, 1-1.5 mm wide; secondary bracts similar, smaller, to 1.8 mm long, 0.2 mm wide; pedicels lax, finely puberulent, 12-19 mm long. Flowers with the calyx somewhat puberulent throughout, teeth of both lobes ciliate and somewhat pilose, especially central tooth of the lower lobe, all teeth of lower lobe 3 mm long, the upper bifid lobe 3 mm long; corolla with standard obovate, retuse, cuneate at the base, to 8.5 mm long and 6 mm wide, wings oblong, obtuse, short unguiculate, 7-7.5 mm long, 2.5 mm wide, keel petals narrowly oblong, truncate at the apex, unguiculate ca. 14 their length, 8-9 mm long, 2 mm wide. Loment to 5-articulate, stipitate, stipe 3 mm long; articles triangular, the upper suture slightly curved, the lower suture rather deeply indented at isthmi, uncinulate pubescent throughout, to 8 mm long and 4.5 mm wide; mature seeds not seen. Desmodium campyloclados ranges from Central America south to Colombia, Venezuela, and Peru. The slender pedicels and relatively large loment articles distinguish Desmo- dium campyloclados. This species is related to D. procumbens and D. tortuosum in the structure of its inflorescence; it differs from both these species in the shape of the loment articles and in having the isthmi between the articles eccentric. The two species of Pittier cited in the synonymy of Desmodium campylocla- dos, D. flexuosum and D. dubium, seem to be equivalent to it. Both species are probably based on a collection from **around Canaguá, 1,500 m, State of Merida, 638 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Venezuela, Nov. 10, 1943, legit V. M. Badillo n. 578.” of which a fragment is in the Gray Herbarium. The type of Desmodium dubium was cited by Pittier as Badillo 598 with the same locality and date of collection as Badillo 578. I think that the latter number was undoubtedly intended. The packet in which Badillo 578 was sent by Pittier bears still another epithet, fortunately never published. CHIRIQUÍ: Cerro Punta, Volcán de Chiriquí, 5500 ft, Bottimer 1531 (A, DAO). Río Chiriquí Viejo north Volcán City, Duke 9014 (MO). cocrÉ: 10 mi E of Nata at Rio Grande, Tyson 5265 (FSU, SCZ), collection very immature, placed here with doubt. 8. Desmodium distortum (Aubl.) Macbr., Field Mus. Nat. Hist., Bot. Ser. 8: 101. 1930. Hedysarum distortum Aubl., Hist. Pl. Guiane 2: 774. 1775. TYPE: French Guiana. Aublet (BM, holotype; GH, US, photo). H. asperum Poir. in Lam., Encycl. Méth. Bot. 6: 408. 1804. TYPE: locality unknown according to Poiret (P-JU, holotype; GH, photo). Aeschinomene aspera (Poir.) St. Hilaire, J. Bot. (Desvaux) 1: 60. 1813. Desmodium asperum (Poir.) Desv., J. Bot. (Desvaux) 1: 122. 1813 D. rubiginosum Benth., Ann. Nat. Hist. 3: 434. 1839. type: British Guiana, Berbice, Schomburgk 217 (K, holotype; G, isotype; GH, photo). D. spectabile Miq., Linnaea 18: 570. 1844. type: Surinam, Para, Onoribo, Focke 880 (U, holotype; GH, photo). Meibomia aspera (Poir.) Kuntze, Rev. Gen. Pl. 1: 197. 1891. M. distorta (Aubl.) Schindl., Repert. Spec. Nov. Regni Veg. 22: 281. 1926, not Schindl., Repert. Spec. Nov. Regni Veg. 20: 148. 1924. Herbs or shrubs, to 3 m tall, branching mostly above, the stem slender, terete to subangulate, reddish, abundantly uncinulate puberulent and moderately long pilose with attenuate trichomes. Leaves trifoliolate, stipulate, and petiolate; stip- ules essentially amplexicaul, obliquely ovate attenuate, long ciliate, striate, pu- berulent and somewhat pilose on both the surfaces to glabrescent, soon reflexed, long persistent, 6-15 mm long, 3.5-6 mm wide; petioles sulcate, densely uncin- ulate puberulent and moderately pilose, 1.3-8 cm long; leaf rachis similar, 0.7-3 cm long; stipels elliptic attenuate, striate, ciliate, puberulent on abaxial surface, 4-11 mm long, to ca. 2 mm broad; petiolules somewhat stouter than the leaf rachis, densely pilose, 1.5-5 mm long; leaflets elliptic to ovate, mostly obtuse and apiculate, obtuse at the base, uncinulate puberulent and appressed pilose on the adaxial surface, prominently reticulate and abundantly soft pilose on the abaxial surface, rather densely ciliate, terminal leaflet 3.7—17 cm long, 2-6 cm wide, lateral leaflets 2-11 cm long, 1.5-5 cm wide. Inflorescence racemose pan- iculate, composed mostly of densely flowered slender, elongate racemes, the pairs of pedicels at maturity usually less than 1 cm apart: rachis ridged and grooved, densely patent pilose with slender trichomes glandular at the base and also un- cinulate puberulent; primary bracts each subtending 2 pedicels, each of these further subtended by a secondary bract, slenderly ovate attenuate, striate, pilose, ciliate, finely puberulent within, soon deciduous, 2.5-5 mm long, 0.5-1.2 mm wide, secondary bracts similar, slenderer, 1-1.7 mm long, 0.2-0.5 mm wide, also quickly deciduous; pedicels slender, ascending, glandular pilose, 6-10 mm long. Flowers with the calyx bilabiate, glandular puberulent throughout and somewhat pilose, the teeth finely ciliate, the upper lobe orbicular, slightly bifid, 2-2.5 mm long, the central tooth of the lower lobe deltoid acuminate with stout pilosity 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 639 along its midrib, 2.5-3.5 mm long, the lateral teeth acute, 2.5 mm long; corolla rose purple to greenish purple, the standard obovate, acute at the base, 5-6 mm long, 3—4.5 mm wide, the wings oblong, obtuse, auriculate, 4.5-6 mm long, 1.5- 2.5 mm wide, the keel petals scythe shaped, long unguiculate, 5-7 mm long, 1- 2 mm wide. Loment stipitate, to 6-articulate, stipe 1.5-2 mm long; articles or- bicular or nearly so but often becoming variously involuted, 1.5-2.5 mm long, 1.5-2 mm wide; seeds quadrate, obtuse at the ends, reddish brown, 2 mm long, ] mm wide. This species occurs from Mexico to northern South America. It is not entirely clear whether Desmodium hirsutum Mart. & Gal. (1843) of Mexico is distinct from D. distortum. More Mexican material must be studied before a satisfactory conclusion can be reached. Should the two elements prove to be the same, Desmodium distortum (Aubl.) Macbr. is the name to be main- tained for them. (See Standley & Steyermark 1946, under Desmodium hirsutum Mart. & Gal.). ANAL ZONE: Across railway track from Summit Gardens, Bottimer 1512 (A, DAO). Cocoli ad. Burch et al. 1391 (GH, MO). Venado Beach, Correa & Gonzalez 506 (A, SCZ). Hill S of Pedro Miguel Locks, Croat 9175 (MO). Summit Naval Radio Station, Croat 14241 (MO). Cocoli, Dwyer ment ie Las Cruces Trail, 75 m, Hunter & Allen 711 (GH). La Boca, Mori & Kallunki 3678 MO). Ancón, Piper 5186 (US). Around Culebra, 50-150 m, Pittier 2233 (US). Balboa, Standley 26400, 27155 (both US). Corozal, ye uid 27393 (US). Río Pedro Miguel pp a Paraiso, Standley 30037 (US). Balboa, Standley 32138 (US). Curundu, Tyson 3478 (FSU, MO, SCZ). Miraflores Lake, White 59 (US). cHiRIQUÍ: 10 mi W of Puerto Armuelles, 100-300 m, Liesner 90 (MO). Near Remedios, 0-100 m, Koster 167 (A). 6 mi W of San Lorenzo, Wilbur et al. 15473 (Duke). cocLÉ: Panamerican Highway and Rio Teta, Blum & Tyson 1890 (FSU, MO, SCZ). 3 mi NE of Antón, D'Arcy & Croat 4102 (MO). Río Hato, 0-100 ft, Rattray 43, 47 (both A). Penonomé, 50-1000 ft, Williams 117 (NY). Ocú, 100 m, Allen 4043 (MO). Road from La Avena to outskirts of Pesé, 200 ft, Burch et e M ) ft, xar 8505 (MO). 500 m del Río Trapichito (La Chorrera), Ad ra 15 (DUKE, MO). Taboga Island, Hinds (K); Macbride 2816 (US). vERAGUAs: 1 mi W of Santiago along old Interamerican Hi amisqa Tyson 5172 (FSU, SCZ). 9. Desmodium glabrum (Mill.) DC., Prodr. 2: 338. 1825. Hedysarum glabrum Mill., = Dict., ed. 8. no. 12. 1768. TYPE: Mexico. Campeachy. Houstoun (BM, holotype; GH, p H. molle Vahl, Symb. ne 2: ra 1791. TYPE: ex insula St. Crucis, misit Dr. Pflug (C, holotype; GH, photo). H. terminale Rich., Actes Soc. Hist. Nat. Paris 1: 112. 1792. Type: Cayenne. LeBlond (G, probable isotype; GH, photo). H. emarginatum Poir., in Lam., uem Méth. Bot. 6: 412 [no. 45]. 1804. TYPE: Martinique, herb. Lamarck (P-LAM. 307; GH, Desmodium molle (Vahl) DC., Prodr. p 332, 1825. M. emarginata (Poir.) Kuntze, Rev. Gen. Pl. 1: 198. 1891. Desmodium campestre Brandegee, Univ. Calif. Publ. Bot. 6: 43. 1914. TYPE: Mexico. Oaxaca, Pi- cacho-San Geronimo, Purpus 6810 (UC, holotype; GH, isotype). 640 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Herb, erect, to 2 m; stem mostly simple, striate, grooved, densely uncinulate pubescent, pilose. Leaves trifoliolate, stipulate, petiolate; stipules ovate, abruptly long attenuate, slightly auriculate at the base, striate, puberulent and pilose on abaxial surface, ciliate, usually turning dark purplish and becoming reflexed, per- sistent, to 6.5 mm long and 3 mm wide; petioles sulcate on adaxial surface, striate to ridged and grooved, densely uncinulate pubescent and somewhat pilose, to 3 cm long, leaf rachis similar, ca. 1 cm long, stipels lance attenuate or slightly broader, ca. 4 mm long; petiolules somewhat stouter than the leaf rachis, densely pilose, 1.5-2.5 mm long; leaflets slightly uncinulate puberulent especially along the midrib above, and abundantly pilose to tomentose on both the surfaces, ter- minal leaflet ovate acute, rounded to cuneate at the base, to ca. 7.5 cm long, 4.5- 4.8 cm wide, lateral leaflets similar, slightly oblique, usually rounded at base, 3.5-5 cm long, 2.5-3 cm wide. Inflorescence racemose paniculate, often much branched, rachis striate, uncinulate puberulent and with somewhat glandular tri- chomes in addition; primary bracts narrowly ovate acuminate, striate, puberulent, and ciliate, 2.5 mm long, 0.5-0.7 mm wide, each subtending a fascicle of 3 to several pedicels; secondary bracts similar, smaller, each subtending a single ped- icel, 0.8-1.3 mm long, 0.3-0.5 mm wide, all bracts early deciduous; pedicels with pubescence similar to that of rachis, 5-7 mm long; calyx puberulent throughout and moderately stiff pilose, the teeth of both lobes ciliate, the central tooth of the lower lobe 2-2.7 mm long, the lateral teeth 2-2.5 mm long, the upper bifid lobe 2-2.5 mm long; corolla with standard narrowly obovate tapering to an acute base, 2.5 mm long, 1.5 mm wide, the wings obliquely oblong, broadest at the apex and slightly auriculate at the base, unguiculate, to 2.5 mm long, 1 mm wide, keel petals scythe shaped, scarcely auriculate, narrowed to the claw, to 2.5 mm long and 0.7 mm wide. Loment almost sessile, to 3-articulate, the terminal article enlarged and flattened, essentially elliptic except for slight indentation over seed, uncinulate pubescent becoming glabrous except on suture at maturity, with re- ticulate venation prominent, subterminal articles quadrangular with alternate mar- gins folded in or back and appearing twisted, uncinulate puberulent; seed trans- versely elliptic, 2 mm long, 1 mm wide. This species occurs in Mexico, the West Indies, the northern half of South America and the Galapagos Islands. CANAL ZONE: Farfan Beach area, Tyson & Blum 2604 (FSU, SCZ). CHIRIQUÍ: Alanje/David, 0— 00 m, Burt & Koster 151 (MO). COLON: Punta Chame, D'Arcy 10237 (MO). PANAMÁ: La Exposición, Heriberto 276 (US). Bella Vista, Standley 25401 (US). Panama City, Seemann 219 (GH, K, S); Seemann (BM). 10. Desmodium incanum DC., Prodr. 2: 332. 1825. TYPE: Plumier, Ic. 149. Fig. 1. 1757. n Plumier, Ic. 149. fig. /, 1757. H. frutescens sensu Jacq., Hort. Bot. Vindob. 3: 47. tab. 89. 1776, non L. 1753 (=Lespedeza). H. supinum Swartz, Prodr. 106. 1788; Fl. Ind. Occ. 3: 1264. 1806, non H. supinum Chaix ex Villars 177 H. incanum Swartz, Prodr. 107. 1788; Fl. Ind. Occ. 3: 1265. 1806, non H. incanum Thunb. (1784). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 641 H. ur J. F. Gmelin, Linn. Syst. Nat., ed. 13. pu e 1791. Based on Plumier, Ic. 149. fig. . 1757. Illegitimate renaming of H. racemosum H. canum Lunan, Hort. Jam. 305. 1814. Renaming Toy incanum Sw Aeschynomene incana (Swartz) G. F. Meyer, Prim. Fl. Esseq. 245. 1818, not Vog. (1838). Desmodium dd a diia um DC., Prodr. 2: 331. 1825. Hos, on Hedysarum racemosum Aubl., non Thunb. (1784), nec Desmodium racemosum DC. (1825). Meibomia e Abre (Swartz) DC. var. 8 incana TM Rev. Gen. Pl. 1: 195. 1891. M. incana (Swartz) Vail, Bull. Torrey Bot. Club 19: 118. 1892. M. supina (Swartz) Britton, Ann. New York Acad. Sci. 7: 83. 1892. Desmodium supinum var. amblyophyllum Urb., Symb. Ant. 7: 229. 1911. LECTOTYPE: Dominican Republic: Santo Domingo prope Constanza, Tuerckheim 3183 (NY). D. canum (J. F. Gmelin) Schinz & Thellung in Schellenb., Schinz & Thellung, Mem. Soc. Neuchatel EI 371. HS Repert. Spec. Nov. Regni Veg. 12: 428. 1913 . F. 24. dm frutescens Schindl., Repert. Spec. Nov. Regni Veg. 21: 9. 1925. Based on Hedysarum frutescens sensu Jacq., non L. 1753 (= Leineden]. Suffruticose to shrubby; stem to 3 m tall, ascending to generally erect, terete to subangulate, uncinulate puberulent to pubescent and somewhat pilose. Leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate acuminate, striate, pu- berulent and pilose on the outer surface, ciliate, at least partially connate on the opposite side of the stem from the petiole, long persistent, 6-10 mm long, 1-2 mm wide; petioles sulcate adaxially, uncinulate puberulent and long spreading pilose, 1.3-3.5 cm long; leaf rachis similar, shorter, to 1.5 cm long; stipels sub- ulate, puberulent, ciliate, 2-5.5 mm long; petiolules stouter, densely stiff pilose, to 3 mm long; leaflets variable, often nearly orbicular at the base of the plant and lanceolate at the apex, mostly elliptic, acute at the apex and rounded at the base, lustrous, darker, and with pilosity along midrib and some uncinulate puberulence on adaxial surface, densely pilose on the paler abaxial surface, terminal leaflets 4-9 cm long, 2.5-4.5 cm wide, the lateral leaflets 3.3-6 cm long, 1.8-2.7 cm wide. Inflorescence terminal and axillary, racemose; rachis angulate, ridged and grooved, densely uncinulate puberulent; pedicels usually solitary, each subtended by 1 primary bract and 2 lateral secondary bracts, all persistent; primary bracts lance acuminate, puberulent on the abaxial surface, ciliate, 2.2-4.5 mm long, 0.5- 0.7 mm wide, secondary bracts similar, 1-1.8 mm long, 0.2-0.5 mm wide; pedicels also uncinulate puberulent, 5-10 mm long. Flowers with the calyx bilabiate, un- cinulate puberulent and pilose, the central tooth of lower lobe 2.5-3.5 mm long, the lateral teeth 2-3.5 mm long, the upper bifid lobe 2.5-3.5 mm long; corolla exceeding the calyx, standard obovate, retuse, unguiculate, 3.5-5 mm long, 4—5 mm wide, the wings oblong, slightly auriculate, unguiculate up to ca. V5 their length, 3.5-6 mm long, 1.5-2 mm wide, the keel petals scythe shaped, obtuse at apex, scarcely auriculate, unguiculate, 3.5-6 mm long, 1.5-2 mm wide. Loment stipitate, to 8-articulate; stipe 1.5-2 mm long; articles essentially straight above, invaginated about 24 their width at the isthmi, uncinulate pubescent throughout, 3.5-4 mm long, 2.5-3 mm wide; seed obtusely oblong, 1 mm long, 0.5 mm wide This species occurs in tropical areas throughout the Americas. It is presum- ably introduced and widespread in the tropics of the Old Wor Desmodium incanum is a widespread more or less shrubby species varying chiefly in the shape of its leaflets. It may be distinguished by its long persistent stipules usually fused and nearly surrounding the stem at least when young, and by its pedicels which are usually borne singly and are each subtended by one 642 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 primary bract and two (lateral) secondary bracts. The extent of the range of Desmodium incanum is partially indicated by the extent of its synonymy, a new name and description apparently having been provided as it was discovered and collected in each successive new locality. The vernacular names of Desmodium incanum are numerous, some having come with the workmen from the islands of the Antilles. Among the names re- corded by collectors in Central America and the West Indies are the following: Pega-pega which is also used for other species of the genus; John Charles: wild pea-nut; strong back; strongbark; and cepa de caballo. The plants are commonly used medicinally, as a tea, and in decoctions as a remedy for inflammation of the stomach and for kidney ailments. BOCAS DEL TORO: Talamanca Valley, Carleton 120 (NY, US). Region of Almirante, Cooper 19] (US). Hill above RR station at milla 7. 5, Croat 16389 (MO). Lincoln Creek, Changuinola Valley, H f CANAL ZONE: Co ló ón, Angermuller (FSU). Barro Colorado Island, Bailey & Bailey 136 (GH): nos gham 565 (A); Croat 4056, 4174 (both MO, SCZ), 5328, 6140 (both MO), 6421 (A), 6684, 6730 (both MO). 6795, 6928 (both A, MO), 7059, 7456 (both MO), 8261, 8675 (both A, MO), 9198 (MO); 9402 (MO, SCZ), 9203 (MO), 11719 (MO, SCZ), ahs O); Ebinger 30 (GH, MO), 154 (MO), Foster 1376 (DUKE); Luteyn 768 (DUKE). Gatun Lake, Netting 36, 46 (both MO); Starry 23, 113, 219 (all 735 (GH). Ft. Kobbe, Bottimer 1445 (A, DAO). Summit, Bottimer 1506 (DAO). Monkey Hill, Cowell 40 (NY). Trail between Rio Indio Hydrographic Station ee Natural Bridge of Rio Puente, Dodge Allen 17488 (GH, MO). Just E of Gatun Locks, Duke 4300 (MO). Chagres, Fendler 78 (K, MO, P, U mboa, Greenman & Green nman 5161 ( T Gatün, Hayes 79, 81 (both NY). Las Sabanas, i Lazor 2199 (FSU). Frijoles, 10-30 m, Maxon 4708 (US). Near Ft. Randolph, Maxon 6513 (US). Road C29, 6 km E of Gamboa, 190 m, Nee 9003 Mie Panamerican Highway between Rodman Marine Base and Chorrera, Dwyer 3584 (GH, MO). G n, Ostenfeld 68 (C). Sabanas NE of Panama it Paul 150 (US). Corozal, Piper 5185 (US). Frijoles. Pos 5191 (US). Ft. Sherman, Piper 5885 (U Fort Lorenzo, mouth of Río Chagres, Piper 5926 (US). Río Grande near Culebra, 50-100 20-75 m, Seibert 383 (A, K, MO, NY). Las Cascadas Plantation near Sum mit, Standley 25705 (US). Cerro Gordo near Culebra, Standley 25973 (US). Hills W of the Canal near Gatun, Standley 27295 (US). Mt. 30029 (US). Fort Clayton, Cardenas Creek area, Tyson 1288 (MO). Fort Sherman near mouth of Chagres River, Tyson 1534 (SCZ). Old Fort San Lor ae Tyson 1559 (MO). Howard Air Force Base near Red Devil Drop Zone, Tyson 1879 (FSU, SCZ). Fort Sherman main post, Tyson 2243 (FSU, O, SCZ). cHiRIQUí: David/Concepion, 60 m, Burt & Koster 134 (MO). Near Frontera, 100-200 m Burt & Koster 145 (MO). Monte Verde, 2.5 km W Puerto Armuelles, 80 m, Busey 415 (MO). Rabo de Puerco 8 km W from Puerto Armuelles, 150 m, Busey 443 (MO). Boquete, 3800 ft, Davidson 596 (GH, MO). Road to Cia 0-100 m, Koster 123 (A). age David and Concepción, Koster 134 (A). 2 km S of Gualaca, McCorkle C-35 (FSU, SCZ). 4 km N of Burica, McCorkle C-140 (FSU, SCZ). weil El Valle, Bottimer 1474 (A, DAO). Weedy field near Santa Clara Beach, Croat 9603 (MO). Savanna 4-7 mi E of Nata, 10-25 m, Duke 12397 (NY, OS). Penon nomé, Ebinger 1001 (MO). Road to El Valle, Ebinger 1101 (MO); eig 1191 (A). Olá, 100—350 m, Pittier 5048 (US). Between Penonomé and Coclé, Stern et al. 988 (MO, UC). Penonomé 50-1000 ft, Williams 116 (NY, US). COLÓN: Río Piedras along road to dice "Blum et al. 2540 (FSU, GH, MO, SCZ). 2 mi E of Rio Piedras, Correa & Haines 239 (MO, SCZ). 1⁄4 mi N of Maria Chiquita on road to Portobelo, wa 11348 (MO, SCZ). Ridge behind Garotte, D'Arcy 9339 (MO). Road to Portobelo between Río Piedra and Portobelo, Elias & Kirkbride 1644 (MO, UC). Colon Highway Bridge on Chagres River, Lazor & Tyson 3029, 3063 (DUKE, FSU). Mouth of Rio Piedras, Lewis et al. 3174a (MO). Santa Rita Ridge, 2 mi E of Transisthmian Highway, Lewis et al. 5244 (MO, SCZ). Portobelo 0-20 m, Pittier 2458 (N US). Palenque near sea level, Piitier 4130 (US); Rose 22074 (S, US). 5-7 mi SW of Portobelo towards María Chiquita, Wilbur & Weaver 11179 (DUKE). 5 mi NE of Sabanita towards Portobelo, Wilbur 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 643 & Luteyn 11621 (DUKE). DARIÉN: Santa Fé, Duke 9493 (MO). Río Pinas, Duke 10571 (MO). Hydro Camp Pico Pendejo in Monsoon Forest on Río Sabana 50 ft, Duke 15453 (OS). El Real, Folsom 4598 Panamerican Highway, Folsom 3447 (MO). Panama City, Parque Lefevre, Harkins 1186 (A). San José Island, Johnston 1006, 1086 (both GH), Erlanson 131 D 592 (NA). Bella Vista, Killip 3117 (US); Koster 145 (A). Chimán, Lewis et al. 3363 (GH, MO, UC). Near Tapía River, Juan Díaz region, Maxon & Harvey 6624 (US). Taboga Island, Miller 1851 (US). Panamerican Highway 2 km W of El Llano, 0-100 m, Nee 7959 (MO); Peralta 23 (GH, MO). Taboga Island 0-250 m, Pittier Wa (US). Chepo, 60 m, Pittier 4761 (NY, US). El Valle, 580 Et Rattray 26 (A). Panamá, Seemann (BM Seemann 227 (K). Bella Vista, Standley 25352 (MO, US). Taboga Island Standley 27087 S. US), 27902 (C, US). Río Tapia, Standley 28184 (US). Cinna, pu a Platanares, Taylor 16 (MO). HERRERA: Chitre/Divisa, roadside, 0-100 m, Burt & Rattray 69 (MO); Vicinity of Ocú, 1⁄2 mi NE open pasture, Stern et al. 1703 (MO). Los SANTOS: 17.8 m e of Macaracas, 1100 ft, Lewis et ici mi S T Wendehake 22 (DUKE, MO). SAN BLAS: Puerto Obaldía, Croat 16973 (MO). Mainland in front of Ustupo, D’Arcy 9520 (MO). Mulatuppu (Río Ibedi), Duke 8482, without other locality 8536 (both MO), Duke 14831 (NY, OS). Between Río Diablo and Río Acuatí near Narganá, Duke 14860 (m Hydro Camp Cuadí on Río Cuadí, ca. 46 ft, Duke 15467 (OS). San Blas-Darien Border, Camp Sum 1200 ft, Mp 15493 (OS). Soskatupu, Elias 1672 (MO, UC). Headwaters of Río Mulatuppu, pm 1745 (MO, UC). Mouth of Ailigandi River to 2.5 mi inland, Lewis et al. 166 (MO). Molia, Stier 60 (MO). VERAGUAS: Savanna 10 mi W of Santiago, Duke 13649 (OS). 5 mi N of Santiago, Santa María River, Blum & Tyson 597 (MO, SCZ). PROVINCE UNKNOWN: Halsted (NY); Hayes 51 (NY); Cuming 1139 (K); Duchassaing [anno 1851] (P); Sinclair (K, MO). 11. Desmodium infractum DC., Prodr. 2: 330. 1825. Based on Hedysarum biar- ticulatum DC.—FIG. 21E-F. Hedysarum biarticulatum Moc. & Ses. ex DC., Prodr. 2: 330. 1825. as synon., non L. Based on Fl. Mex. Icon. (ined.) pl. 270 (G, not seen); volume known as ''Calques des Dessins de la Flore du Mexique” (i.e. tracings of the original drawings, at GH). H. infractum Sprengel, Syst. Veg. 4(2): 290. 1827. Based on D. infractum DC. Desmodium barclayi Bentham, Bot. Voy. Sulphur 83. 1844. TYPE: Centhal America, Barclay (K, holotype; GH, NY, photos). Meibomia barclayi (Benth.) Rose & Standl., Contr. U.S. Natl. Herb. 16: 216. tab. 51, fig. e. 1913. M. infracta (DC.) Blake, Bot. Gaz. 78: 280. 1924. Nephromeria barclayi (Benth.) Schindl., E. Spec. Nov. Regni Veg. 20: 282. 1924. Nephromeria infracta (DC.) Schindl., Repert. Spec. Nov. Regni Veg. 22: 285. 1926. Sprawling, subscandent vine; stem angulate, ridged and grooved, uncinulate pubescent with somewhat spreading stout, hooked, yellowish trichomes mostly on the ridges and with some puberulence of short, straight, pointed, white tri- chomes intermixed. Leaves trifoliolate, stipulate; stipules early deciduous, obliquely ovate, long acuminate, striate, ciliate, and somewhat soft pilose over abaxial surface, 5-7.5 mm long, 3-5 mm wide at base; petioles adaxially sulcate, ridged and grooved, sparsely to densely uncinulate puberulent to pubescent, 3— 7 cm long, the leaf rachis similar to petiole, 1.5-2.5 cm long; stipels linear atten- uate to narrowly deltoid attenuate, striate and ciliate, persistent, 2-3.5 mm long; petiolules greenish to dark brown, rugose, patent pilose with slender tapering trichomes, 2-3 mm long; leaflets ovate, obtuse to acute at apex, the terminal cuneate, the lateral leaflets rounded at base, the margins ciliate, adaxial surface bright to dark green, sometimes brownish in age, uncinulate puberulent along the midrib and the chief lateral veins or this puberulence somewhat scattered, sparse- ly to abundantly spreading pilose with straight tapering trichomes between them, or the 2 types intermixed, abaxial surface always paler, abundantly soft pilose with slender tapering trichomes, the midrib and the lateral veins prominent, the 644 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 21. Desmodium.—A-B. D. macrodesmum (Blake) Standl. & Steyerm.—A. Habit (x%).—B. Article of lome nts salir to show elongate funicle and seed inside (x1). [After Wedel 1773.]—C-D. D. rhy riu i. Dl Standl.—C. Habit (x 2).—D. Opened article showing funi- cle and seed (x1). [After D'Arcy 11241 (A).]—E-F. D. infractum DC.—E. Habit (x). iar ad 6019 (GH).]—F. Opened article showing funicle and seed (x1). [After Tyson 6019 (GH).]—G-H saccatum Schubert.—G. Habit (xV4). [After Dwyer & Correa 7996.]—H. Opened article (x 1). ye Dwyer 5028 (GH).] 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 645 terminal leaflet 6-7 cm long, 4.5-5.6 cm wide, the laterals 5-5.5 cm long, 3-3.5 cm wide. Inflorescence composed of axillary and terminal racemes or highly branched panicles; rachises angulate, striate, densely uncinulate pubescent with stout, yellowish, hooked trichomes underlain by minute, white trichomes; pedi- cels borne in pairs, each pair subtended by a primary bract; secondary bracts each subtending a single pedicel only rarely seen; primary bracts ovate acuminate, striate, ciliate, puberulent on abaxial surface, early deciduous, 5-7 mm long, 2.5 mm wide^*; pedicels uncinulate puberulent, 4-7 mm long. Flowers with the calyx appressed pilose on the teeth of both lobes, quickly reflexing almost completely at flower maturity, then disintegrating, central tooth of lower lobe 5 mm long, lateral teeth 3.4 mm long, upper bifid lobe 3 mm long*!'; corolla? magenta to white, the standard orbicular, scarcely clawed, 4.5 mm long and broad, the wings almost rectangular, short clawed and slightly auriculate, truncate at apex, 5.5 mm long, 4 mm wide, keel petals partially fused, each scythe shaped and tapering to a slender base, 5 mm long, 1.5 mm wide at apex. Loment stipitate, (1—)2—3- articulate, the stipe slightly pubescent, becoming glabrate, 3-4 mm long; articles nearly orbicular, with a narrowly V-shaped notch ca. 2.5 mm deep on the dorsal suture, shoulders becoming somewhat beaked, the articles often overlapping, ciliate with both hooked and tapering trichomes, puberulent to glabrous on sur- faces; seed reniform, reddish to brown, almost centrally placed in the article but not filling it, long funiculate, ca. 3.5 mm long, 2.5 mm wide. In the sect. Nephromeria, of which it is a member, Desmodium infractum is most closely related to D. painteri (Rose & Standley) Standley, a species known only from the states of Mexico and Guerrero in Mexico. Desmodium infractum differs from D. painteri in its larger, more nearly orbicular articles with a nar- rower, deeper notch on the upper suture that usually flattens out into shoulders (at least on one side). It also differs in its ovate leaflets, rough above to the touch and with only moderately prominent venation below, whereas D. painteri has nearly orbicular leaflets with secondary venation prominent on the velvety abaxial surface. In D. infractum the pedicel and the stipe of the loment are essentially equal, in D. painteri the stipe is shorter (only !5—!5 as long) than the pedicel. Desmodium infractum is known from scattered localities in southern Mexico (Oaxaca, Nayarit, Michoacan or Guerrero, Chiapas, and Veracruz) and Central America where specimens have been seen from every country; in Panama it is known from Veraguas, Los Santos, and the Canal Zone; in South America it has been recorded from Colombia, Venezuela, and Ecuador (see Blake, 1924). Much material that was named D. infractum from the province of Bocas del Toro, Panama, is actually D. macrodesmum. CANAL ZONE: Vicinity of Madden Dam, alt. 50 ft, Lewis et al. 7 (GH, MO). Madden Dam area, Dwyer 11969 (MO). Los SANTOS: 10 mi N Tonosi, Tyson et al. 2944 (MO, SCZ). 12 mi S Macaracas, Tyson et al. 3063 (SCZ). VERAGUAS: Puerto Mutis, 12 mi S of Santiago, Tyson 6019 (FSU, GH, SCZ). 4 Measurements taken from Guatemalan material, Standley 77910 (GH). 41 Measurements taken from Guatemalan material, Standley 74508 (GH 42 Measurements of corolla parts taken from Mexican material, Martinez- Calderon 726 (A). Little flowering material of this species has been collected. 646 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 12. Desmodium intortum (Mill.) Urb., Symb. Ant. 8: 292. 1 Feb. 1920; non Fawc. & Rendle, Fl. Jam. 4: 34. 25 Mar. 1920. Hedysarum intortum Mill., Gard. Mid m 8. Hedysarum no. 11. 1768. TYPE: Cultivated, England, seed sent from Jamaica by Hous H. trigonum Swartz, Prodr. 107. 1788; 'Fl des Occ. 3: 1267. 1806. Based on Hedysarum intortum Mill. Desmodium trigonum (Swartz) DC., Prodr. 2: 332. 1825. Meibomia intorta (Mill.) Blake, Bot. Gaz. 78: 286. I Meibomia trigona Gandara, Mem. y Rev. Soc. Cient. "Antonio Alzate” 51: 113. pl. 14. 1931. [De- scribed as a new species from Salvador[?], not opt on Hedysarum trigonum Sw., but belonging here nevertheless.] Herbs, trailing to climbing; stem triquetrous and grooved, pilose and/or un- cinulate pubescent. Leaves trifoliolate, stipulate, and petiolate; stipules ovate acu- minate, striate, puberulent and somewhat to densely pilose, abundantly ciliate, 5.5-6.5 mm long, 2.5-5 mm wide; stipels linear to narrowly ovate attenuate, puberulent and ciliate, 3-4 mm long; petioles sulcate, finely ridged and grooved, long soft spreading pilose, 3.2—5 cm long; leaf rachis similar, slender, 0.5-1.2 cm long; petiolules stouter, more densely spreading pilose, 1.5-3 mm long; leaflets mostly ovate acute, moderately appressed pilose on both the surfaces, the ter- minal leaflet rhombic, acute at the apex and rounded to cuneate at base, 3.5-7 cm long, 2.4-4 cm wide, the lateral leaflets nearly elliptic, acute at the apex, rounded at the base, 3-5 cm long, 1.6-2.4 cm wide. Inflorescence chiefly terminal, racemose to racemose paniculate; rachis ridged and grooved, densely uncinulate puberulent; bracts ovate acuminate, finely striate, puberulent on abaxial surface, ciliate, 5.5-8.5 mm long, 3-4 mm wide; secondary bracts not seen; pedicels un- cinulate puberulent, 4-6 mm long. Flowers with the calyx puberulent throughout with some long pilosity on the ciliate teeth of both lobes, the central tooth of lower lobe 4-5.5 mm long, the lateral teeth 2.5-4 mm long, the upper bifid lobe 2.5-3.5 mm long; corolla with standard 6-7.5 mm long, 4—4.5 mm wide, the wings 5.5-7.5 mm long, 2-2.5 mm wide above, the keel petals 5.5-8 mm long, 2-2.5 mm wide above. Loments stipitate, to 11-articulate; articles uncinulate pubescent throughout, slightly asymmetrical, more deeply indented below, ca. 3.5 mm long, 2.5 mm wide; seed reniform, 2 mm long, 1.3 mm wide. In the treatment of Desmodium for the Flora Fanerogámica del Valle de México*! (Rzedowski & Rzedowski 1979), D. aparines DC. (316, 317) is described as having pedicels from 5.5-16 mm long and as being much confused with D. uncinatum. Schindler, in 1926 (Repert. Spec. Nov. Regni Veg. 22: 274-276) pro- posed that the amalgam of collections relegated in the past to Desmodium un- cinatum were, in fact, representatives of eleven somewhat related species, five of which he described, op. cit. 20: 142—144. 1924; 22: 274—276. 1926. with distinct geographical distribution. The distributions he cited were well defined, but the characteristics of the species were somewhat less so and, in the case of D. aparines, at least two elements are involved as is made clear from his manuscript notes and specimens of loments. According to Schindler the related species of the Desmodium uncinatum complex to be found in Panama are D. aparines and D. intortum. It is also possible that D. sericeum (Schindl.) Standl., D. subseri- + Rzedowski, J., & G. C. de Rzedowski, eds. Flora Fanerogámica del Valle de México. vol. 1: 314-321. 1979 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 647 ceum Malme and D. Ajalmarsonii (Schindl.) Standl., all recorded from Costa Rica, may be found in Panama, too. Schindler is of the opinion that D. uncinatum is strictly South American in distribution, an opinion I question, although it does not seem to be in Panama. Under these circumstances it seems best not to an- notate any collections as D. aparines, but to relate them to D. intortum, as most have been known, for the time being. Two collections designated below with an asterisk (*), which I did name, some years ago, as Desmodium intortum have differences in type of pubescence of stem and pedicels, larger bracts and larger loment articles. They are part of the relationship of D. intortum but not typical. AS DEL TORO: Talamanca Valley, Carleton 125 (US). cHiRIQuí: Concepción, Bottimer 1523 (A, DAO). Volcán de Chiriqui, below village of Volcán, o 1528a (DAO). Volcán Cerro Punta 1460 m, Burt & Koster 153* (MO). Between Hato del Volcán and Concepción, Correa & Lazor 1437 153* (A). Boquete, 990 m, Maurice 752 (US). Pastures around Boquete, 1000-1300 m, Pittier 2963 EE Valley of the upper Río Chiriquí Viejo, White 31 (MO). DARIEN: Agua Fria, ca. 8 mi N of Santa a. 50 m, Duke 10116 (MO). Above Paca, Williams 708 (US). 12a. Desmodium intortum var. apiculatum Schubert, Field Mus. Publ. Bot. 13(3): 4 1943. TYPE: Peru, Mexia 8158 (F) Described many years ago, it can be distinguished from D. intortum var. intortum by its broad, apiculate primary bracts. However, in view of confusion surrounding the whole complex of Desmodium intortum and its relatives I prefer not to cite material at this time. 13. Desmodium macrodesmum (Blake) Standley & Steyermark, Fieldiana Bot. 24(5): 230. 1946.—Fic. 21A-B Meibomia macrodesma Blake, Contr. U.S. Natl. Herb. 24: 6. 1922. Type: Honduras, trail from Hacienda El Limón to El Paraíso, Dept. Copán, Blake 7359 (US, holotype; fragm Nephromeria macrodesma (Blake) Schindl., Repert. Spec. Nov. Regni Veg. Beih. 49: 370. 1928. Scandent herb to many meters in length, stem terete and ridged and grooved, long uncinulate pubescent and shorter spreading pilose. Leaves trifoliolate, stip- ulate, stipules early deciduous, only the broad scars seen; petioles adaxially sul- cate, angulate, uncinulate puberulent, 2-5 cm long; leaf rachis similar 1-2.5 cm long; stipels narrowly elliptic, long acuminate, striate, ciliate, puberulent on abax- ial surface 3.5 mm long, 0.5 mm wide, early deciduous, few seen; petiolules stouter than the leaf rachis, densely stiff pilose 1.5-4 mm long; leaflets acute, the lateral leaflets somewhat obliquely so, the adaxial surface somewhat lustrous, moderately pilosulous, abaxial surface finely silky tomentose with reticulation prominent, terminal leaflet 5.5—10.6 cm long, 3-8.3 cm wide, lateral leaflets 3.2— 7 cm long, 1.9-4.9 cm wide. Inflorescence of axillary and terminal racemes or developing into much branched panicles; rachis angulate, ridged and grooved, moderately to densely uncinulate pubescent; pedicels in pairs, each pair sub- tended by 1 primary bract, secondary bracts occurring rarely if at all; primary bracts ovate acuminate, striate, puberulent and ciliate, 4.5-6 mm long, 1.5 mm 648 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 wide; pedicels uncinulate puberulent and somewhat pilose, 2.5—5(-12) mm long. Flowers with the calyx finely puberulent and long stiff pilose, the teeth of both lobes ciliate, also stiffly short pilose at least on the teeth within, the central tooth of the lower lobe 4-5 mm long, the lateral teeth 3.5-4 mm long, the upper lobe semiorbicular, slightly bifid, 3-4 mm long; corolla exceeding the calyx, violet to purple, standard obovate, deeply retuse, remotely clawed at the base, 5.5-6.8 mm long, 4-5 mm wide, the wings trapeziform, obscurely auriculate, unguiculate at the base, 5.5-6.3 mm long, 3-4 mm wide, the keel petals scythe shaped, long unguiculate (ca. % their length), 5-6.5 mm long, 1.5-2.5 mm wide at apex. Lo- ment stipitate, usually 2-articulate, stipe densely white pilose, 4-7 mm long; ar- ticles almost orbicular, shallowly notched above, the seed slightly above the center, the body of the article flat and papyraceous, reticulate, somewhat pilose on surfaces and velvety to the touch, long spreading pilose on sutures, 2.4-2.9 cm long, 2-2.5 cm wide, the isthmus connecting the 2 articles narrow; seed dark reddish brown, reniform (somewhat immature), 3 mm long, 1.5-2 mm wide. This species is known from Panama, British Honduras, Guatemala and Hon- duras. CAS DEL TORO: Carleton 36 (GH). Water Valley, Wedel 660 (GH, MO), 794, 901, 955 (all MO). Chiriqui CIAM Wedel 1036, 1293, 1773, 2759 (all GH, MO, US). cociÉ: Between Cerro Pilón and El Valle de Antón, 700-900 m, Duke & Dwyer 13899 (MO). El Valle de Antón at the foot of Cerro Pilón, ca. 2000 t Dwyer & Coven 7996 (MO, UC); Ebinger 1114 (MO). PANAMA: Cerro Azul, 600 m, Dwyer 1370, 5143 (both MO). Cerro Campana area, ca. 3000 ft, Dwyer & Kirkbride 7817 (MO). 14. Desmodium maxonii (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 161. 1936. Meibomia maxonii Standley, hee) U.S. Natl. Herb. 18: 108. 1916. TYPE: Panama, Maxon 5370 (US, holotype; GH, photo; NY, isotype). HEE costaricensis Schindler, Repert. Spec. Nov. Regni Veg. 20: 140. 1924. [See note in dis- cussion. Doea costaricense (Schindler) Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 534. 1937. Shrub to 2 m tall; stem terete, striate, finely uncinulate puberulent and when young long white pilose, glabrescent with pilosity only around nodes. Leaves trifoliolate, stipulate, petiolate; stipules obliquely lanceolate to ovate with long attenuate apex, striate, ciliate, puberulent and pilose on the abaxial surface, long persistent, 4.5-8 mm long, 3—4.5 mm wide; petioles sulcate, densely long pilose, 1-3.5 cm long; leaf rachis similar, 0.5-1 cm long; stipels subulate to linear lan- ceolate, striate, pilose and ciliate, 3-5.5 mm long; petiolules stouter than the leaf rachis, long spreading pilose, 2-3.8 mm long; leaflets ovate acute, rounded at the base, mucronate, dark green, puberulent and pilose and uncinulate puberulent on the adaxial surface, paler with prominent venation and long pilosity on the abaxial surface, the margins revolute, the terminal leaflet 3-6.5 cm long, 1.5-3.5 cm wide, the lateral leaflets 2.8-5.5 cm long, 1.2-2.5 cm wide. Inflorescence of many dense, short, terminal and axillary racemes, the crowded bracts at the apex with their tips prominent before flowering; rachis densely uncinulate puberulent and some- what pilose; primary bracts each subtending 2 pedicels, ovate long attenuate, striate, ciliate, pilose on the abaxial surface, 6-10 mm long, 2-5 mm wide; sec- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 649 ondary bracts each subtending 1 pedicel laterally, lance attenuate, ciliate, pilose on abaxial surface, 2.2-6 mm long, 0.2-0.6 mm wide; pedicels uncinulate puber- ulent and somewhat pilose, 6-12 mm long; calyx puberulent throughout, the teeth of both lobes ciliate, the central tooth of the lower lobe densely long pilose, the upper portion of the tube and the other teeth somewhat pilose, the central tooth of lower lobe 5-8 mm long, the lateral teeth 3—5.5 mm long, the upper lobe entire to bifid, 3—5.5 mm long; corolla with standard obovate to flabellate, rounded to retuse at the apex, narrowed to base, 1—1.3 cm long, 0.6-1.1 cm wide, the wings obliquely oblong, truncate at apex, short unguiculate, 7212.5 mm long, 3.5-5 mm wide, keel petals broadly truncate at the apex, long unguiculate, 9.5-13 mm long, 4—5.2 mm wide above. Loment stipitate, 5—6-articulate; stipe 2.2-4 mm long; articles rhomboidal or appearing so because of alternately revolute margins, un- cinulate puberulent, isthmi central, 3.5-4.5 mm long, 2.5-3.5 mm wide; seed re- niform, ca. 2 mm long, 1 mm wide. This species occurs in Central America and southern Mexico (Chiapas). One collection of Desmodium, Bottimer 1527, seems to be a slightly aberrant form of D. maxonii but, unquestionably, is more closely related to this species than to any other. The pubescence of the leaflets, the shape, pubescence, and persistence of the stipules, and the shape but not the pubescence of the bracts are all as in D. maxonii. The elongate petioles and inflorescence rachises, the softer fine pubescence of the bracts and calyces, the straighter dorsal suture of the loments and the slightly eccentric isthmi indicate, perhaps, a lesser relation- ship also with some other species, or may be the result of some sort of ecological disturbance. Included in the manuscript material of the late Dr. A. K. Schindler, purchased by the Arnold Arboretum before his death, are his notes, description, and cita- tions of specimens of Meibomia costaricensis. Wherever possible he added an- notations to the original studies to bring the information up-to-date before sending it on. For this species he indicated that the specimens Tonduz 11769 (in part), Nelson 3599, and Killip 3606 are, indeed, Desmodium maxonii. See my earlier comments (Schubert, 1941). CHIRIQUÍ: 3.4 mi NE of Volcán, Bottimer 1527 (A, DAO). Valley of the Río Chiriquí Viejo N Volcán City, Duke 9058 (MO). Near El Potrero camp, Chiriquí Volcano, 1900 m, Killip 3606 (US). Cuesta de Cerro Quemado, E slope of Chiriquí Volcano, 1800-2600 m, Maxon 5370 (NY, US); Pittier 3112 (US) 15. Desmodium molliculum (H.B.K.) DC., Prodr. 2: 331. 1825. Hedysarum molliculum H.B.K., Nov. Gen. Sp. Pl. 6: 405 (folio ed.); 6: 519 (quarto ed.). 1824. TvPE: Exact locality unknown (P; GH, photo). Heteroloma lanatum Desv. ex H.B.K., Nov. Gen. Sp. Pl. 6: 405 (folio ed.); 6: 519 (quarto ed.). 1824, men nudum in syn. Desmodium mexicanum S. Watson, Proc. Amer. Acad. au 23 (II. 15): 271. 1888. TYPE: Mexico. Chih a, pine plains at the base of the Sierra Madre, C. G. Pringle 1226 (GH, holotype). Meibomia mallicula (H.B.K.) Kuntze, Rev. Gen. Pl. 1: 198. d Herb, stem repent to scandent from a perennial rootstock, stems densely patent pilose with long white trichomes; leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate, acuminate, striate, ciliate, puberulent and pilose on the 650 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 abaxial surface, becoming reflexed at maturity, long persistent, 4-5 mm long, 2 mm wide; petioles sulcate, patent pilose, ca. 1 cm long, the leaf rachis similar, ca. 2-5 mm long; stipels lance attenuate, ciliate, persistent, ca. 2 mm long; pe- tiolules stouter than the leaf rachis, densely pilose 1-1.5 mm long; leaflets ap- pressed pilose on both the surfaces, the terminal leaflet suborbicular, slightly retuse or mucronulate at the apex, rounded to cuneate at the base, to 1.5 cm long and wide, lateral leaflets slightly oblique, cuneate at the base, ca. 1 cm long and wide. Inflorescence generally erect, racemose, rachis ridged and grooved, dense- ly uncinulate puberulent to pubescent and with some pilosity; pedicels borne in pairs (or 3’s), each pair subtended by a primary bract and each pedicel further subtended by a smaller secondary bract, all early deciduous; primary bracts ovate acuminate, striate, ciliate, and somewhat pilose, 3—5.5 mm long and 1.5-2.5 mm wide; secondary bracts linear, ciliate, puberulent on abaxial surface, ca. 1 mm long and 0.2-0.5 mm wide; pedicels ascending, densely uncinulate puberulent, to ca. | cm long. Flowers with the calyx bilabiate, puberulent throughout, the central tooth of the lower lobe long acuminate and long pilose, 2.5-4 mm long, lateral teeth acute, only slightly pilose, 2-3 mm long, the upper slightly bifid lobe 2-3 mm long; standard obovate retuse, cuneate at the base, 5.5-6 mm long, 3-4 mm wide, the wings oblong, obtuse and truncate at the apex, unguiculate, 5-6 mm long, 1.5 mm wide, the keel petals scythe shaped, obtuse and truncate at the apex, long unguiculate, 6.5 mm long, 1.5 mm wide. Loment stipitate, to 5-artic- ulate, the articles curved above and below, the isthmi slightly eccentric, uncin- ulate pubescent throughout, to 5 mm long and 3.5 mm wide. Seed suborbicular to quadrate, 2 mm long, 1.5 mm wide. Desmodium molliculum is known from Panama from 2 incomplete specimens with smaller leaflets than usual. It occurs from Mexico to South America at rather high altitudes. The persistent, soon reflexed stipules, early deciduous bracts, and ascending pedicels in a rather loose raceme serve to distinguish D. molliculum from other species with similar habit and distribution. The seed measurements noted above were taken from Koch 77382 (A), Mexico. CHIRIQUI: Llanos del Volcan, 1120-1200 m, Seibert 345 (MO). 3 mi N of El Volcan on old lava flow, 5000 ft, Tyson 5833 (SCZ). 16. Desmodium perrottetii DC., Mém. Légum. 7: 324. 1825. TYPE: Guiane Fran- çaise. Perrottet 1820 (G, holotype; GH, photo). Meibomia distorta (Aubl.) Schindler var. perrottetii (DC.) Schindl., Repert. Spec. Nov. Regni Veg. 22: 282. 1926. Herb; stem apparently simple, terete to subangulate, finely striate, uncinulate puberulent. Leaves unifoliolate; stipules transversely oblique, ovate attenuate, striate, densely long pilose on the abaxial surface, finely long pilose on the adaxial surface, ciliate, persistent, 8.5 mm long, 4 mm wide; stipels ovate attenuate, striate, pilose and puberulent, ciliate, persistent, 6 mm long, ca. 1 mm wide; petiolate, petiole densely uncinulate puberulent, sulcate, striate, 1 cm long; petio- lule densely stiff spreading pilose, 3 mm long; single leaflet narrowly ovate elliptic 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 651 to ovate, obtuse at the apex and the base, broadest slightly below the middle, uncinulate puberulent and somewhat appressed pilose on the adaxial surface, tomentose with dense, long, appressed pilosity, and with prominent venation on abaxial surface, 6.5-8.5 cm long, 4.5-5 cm wide. Inflorescence much branched, racemose paniculate, rachises angulate, striate, uncinulate puberulent and with long spreading gland based trichomes; primary bracts ovate attenuate, striate, pilose, ciliate, early deciduous, ca. 3 mm long, 1 mm wide; secondary bracts 1 mm long, slender (only few seen); pedicels borne in pairs, ascending, with pu- bescence similar to that of inflorescence rachis, 2.5-3 mm long. Flowers with the calyx bilabiate, the teeth of both lobes all ciliate, the surface puberulent and somewhat pilose throughout, the upper bifid lobe obtuse, 2 mm long, the central tooth of the lower lobe acuminate, 2.5 mm long, the lateral teeth acute, 2 mm long; corolla with standard obovate, slightly retuse at the apex, gradually nar- rowed to the base (which is ca. 2 mm wide), 3.5 mm long, 2.5 mm wide, the wings obliquely oblong, obtuse at the apex, scarcely unguiculate (short clawed), 3.5 mm long, 1.5 mm wide, the keel petals scythe shaped, truncate at the apex, unguiculate (long clawed), 3.5 mm long, 1 mm wide. Loment stipitate, to 5-artic- ulate; stipe 2 mm long; articles orbicular, the subterminal ones twisted or with the margins revolute, pubescent with fine glandular trichomes and long stiff spreading pilosity, 2 mm long, 1.5 mm wide; seeds brown, almost quadrate, 0.7 mm long, 0.5 mm wide. Desmodium perrottetii is known from British Guiana, French Guiana and Peru. It was treated as a variety of Meibomia distorta by Schindler which may be a more reasonable disposition. However, it would be best to see more material before making a final decision. COCLE: Between Paso del Arado and Ola, 20-280 m, Pittier 5022 (US). 17. Desmodium procumbens (Mill.) Hitchc., Ann. Rep. Missouri Bot. Gard. 4: 893 Herb with several to many stems arising from the base and soon becoming procumbent. Leaves trifoliolate or occasionally unifoliolate, stipulate, petiolate; leaflets narrowly to broadly ovate. Inflorescence axillary and terminal, racemose to racemose paniculate and highly diffuse; primary bracts subtending fascicles of 3 to several pedicels, of which 2 mature together first; secondary bracts similar to the primary but smaller. Flowers with the calyx puberulent and scattered short pilose throughout. Loments sessile to short stipitate, to 5-articulate; articles with isthmi central, rhombic in outline appearing slightly to much twiste 17a. Desmodium procumbens var. longipes (Schindler) Schubert, Contr. Gray Herb. 129: 8. 1940 Hedysarum tenellum H.B.K., Nov. Gen. Sp. Pl. 6: 408 (folio e Ar (quarto ed.). 1824, non D. on, Pr odr. Fl. Nepal. 243. 1825, nec Sprengel ex DC., Prodr. 2: 333. 1825, in syn. TYPE: Humboldt & ay em Caracas, 440 hex. 5 holotype; GH, bly Desmodium tenellum (H.B.K.) DC., Prodr. 2: 333. 1825. Cyclomorium caracasanum | Walp., Repert. 2: 890. 1843. Type: Caracas Moritz 25 (B?, not seen). 652 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 Meibomia tenella var. longipes Schindler, Repert. Spec. Nov. Regni Veg. 20: 151. 1924. TYPE: Bernoulli & Cario 1206, Guatemala. Retalhuleu. The variety longipes differs from var. procumbens in its more diffuse and delicate aspect, almost completely glabrous stems, thinner leaflets somewhat broader in proportion to their length and more abundantly pilose on both surfaces, and most conspicuously by its slender and elongate, only slightly puberulent pedicels, 12-27 mm long, and its smaller loment articles, 1.5 mm long and 1.3- 1.5 mm wide, which seem tightly twisted and have essentially glabrous sutures. This species occurs in southern Mexico, Central America, and northern South America. Variety longipes occasionally has its upper leaves unifoliolate and, in some areas (near Zamorano, Honduras), it often bears quinquefoliolate leaves. For an earlier treatment of this variety and a discussion of the synonymy involved see Schubert 1940. CANAL ZONE: Quebrada Fea, Quebrada Pura and Canon of Río Chagres, 70-100 m, Dodge & Allen 17410 (G, GH, MO). Río Grande near Culebra, 50-100 m, Pittier 2126 (NY, US). Las Cascadas Plantation near Summit, Standley 29685 (US). cuiRIQUÍ: Cerro de la Plata near San Félix, 120-150 m, Pittier 5154 (US). cociÉ: Ola, 100-350 m, Pittier 5080 (US). PANAMA: Rio Tocumen, Standley 29427 (US). 17b. Desmodium procumbens var. procumbens. Hedysarum procumbens Mill., Gard. Dict., ed. 8. Hedysarum no. 10. 1768. TYPE: Jamaica. Houstoun, 1730 (BM; GH, ‘ H. spirale Swartz, Prodr. 107. 1788. Fl. Ind. Occ. 3: 1273. 1806. Based on the type of Hedysarum procumbens Mill., which Swartz cited in synonymy. Desmodium spirale (Swartz) DC., Prodr. 2: 332. 1825. D. tenuiculum DC., Prodr. 2: 333. 1825. TYPE: Bertero, Santo Domingo (G-DC; GH, photo), based n Hedysarum tenellum Sprengel, non H.B.K. Hedysarum tenellum Spreng. ex DC., Prodr. 2: 333. 1825, nomen in syn. Desmodium sylvaticum Benth., Pl. Hartw. 1: 116. 1843. TYPE: Hartweg 650 (K, holotype; GH, Isotype). Meibomia spiralis (Swartz) Kuntze, Rev. Gen. Pl. 1: 197. 1891. M. procumbens (Mill.) Schindler, Repert. Spec. Nov. Regni Veg. 20: 151. 1924. M. procumbens var. sylvatica (Benth.) Schindler, Repert. Spec. Nov. Regni Veg. 20: 151. 1924. Stem terete, striate, uncinulate puberulent. Leaves trifoliolate, stipulate, pet- iolate; stipules lance attenuate, obliquely auriculate at base, minutely puberulent on abaxial surface, 4-6 mm long, 1-1.5 mm wide; petioles sulcate, uncinulate puberulent, 1-3.3 cm long, leaf rachis similar to petiole, 0.4-0.7 cm long; stipels linear lanceolate, 1.7-2 mm long; leaflets pilose and uncinulate puberulent on both surfaces, the margins stiffly ciliate, the terminal leaflets narrowly to broadly ovate rhombic, obtuse to acute at the apex, mostly cuneate at base, 2.3-6 cm long, 1-3.6 cm wide, the lateral similar, slightly oblique, 1.7—4.5 cm long, 0.8-2 cm wide. /nflorescence rachis densely uncinulate puberulent; primary bracts, each subtending 2 to more pedicels, essentially linear to narrowly ovate attenuate, striate, ciliate, puberulent over abaxial surface, 1.5-2.5 mm long, 0.5 mm wide; secondary bracts, each subtending a single pedicel, linear, ciliate, and puberulent, 0.5-0.8 mm long; pedicels stiff, uncinulate puberulent, 5-7 mm long. Flowers with the teeth of both calyx lobes ciliate, the central tooth of the lower lobe 1.5— 2 mm long, the lateral teeth 1.3-1.5 mm long, the upper bifid lobe 1.3-1.5 mm long; standard obovate to flabellate, slightly retuse at the apex, mostly rounded 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 653 above and cuneate at base, 2—3.5 mm long, 1.5-3 mm wide, the wings oblong, obtuse, scarcely auriculate, short unguiculate, 2—2.5 mm long, 1 mm wide, the keel petals remotely scythe shaped, truncate at the apex, narrowed to the un- guiculate base, 2.5-3.5 mm long, 0.8-1 mm wide. Loment with articles uncinulate puberulent throughout, 3 mm long, 2 mm wide. This species occurs from northern Mexico southward through Central Amer- ica, the West Indies, and the northern half of South America, in tropical Africa and the Philippine Islands (where probably introduced). Variety procumbens is the most widespread of the four varieties of Desmo- dium procumbens. Although its leaflets may have prominently reticulate venation and an aspect closely approaching that of D. tortuosum to which it is related, D. procumbens var. procumbens may be distinguished by its smaller, much less conspicuous stipules, its shorter pedicels, usually borne at right angles to the inflorescence rachis at maturity, and its loment articles regularly smaller and more consistently rhombic in outline. OCAS DEL TORO: Changuinola, Godfrey 67321 TE CANAL ZONE: Sabanas N of Panama City, Paul 585 (US). Experiment Station, Pittier 6909 (G, K bn Hospital grounds, Ancón, Pittier 6909 (P). Balboa, Standley 25491 (US). Frijoles, Standley 27651 (US). Gamboa, Standley 28472 (US). Near Ft. Randolph, Standley 28757 (US). Darién Station, Standley 31569 (US). PANAMA: | mi E of Tocumen Airport on side road off Panamerican Highway, Blum & Tyson 1958 (FSU, SCZ). San José Island, Johnston 542 (BM, GH), /005 (GH). Bella Vista, Standley 25328 (US). Corozal road near Panamá, Standley 26771 (S, US). Taboga Island, Standley 27081 (C, US). Juan Franco Race Track near Panamá, Standley 27698 (S, US). Taboga Island, Standley 27948 (US). Tumba Muerto Road, near Panamá, Standley 29728 (US). Taboga Island near village, Tyson & Loftin 5144 (FSU, MO, SCZ). VERAGUAS: Puerto Mutis, 12 mi S of Santiago, Tyson 6015 (FSU, MO, SCZ). PROVINCE UN- KNOWN: Camino de Las Sabanas, Heriberto 264 (US). 18. Desmodium rhynchodesmum (Blake) Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 535. 1937 [Fl. Costa Rica].—Fic. 21C- Meibomia rhynchodesma Blake, Bot. Gaz : 278. 1924. Type: Costa Rica. Tuis, 650 m, Tonduz 11448 (US-694364, holotype; isotypes is. 943724, fruit, US-943725, flower). Herbaceous vine to subshrub (?), scandent, festooning on thicket (D'Arcy 11241) or climbing on trees (Shank & Molina 4180, Costa Rica); stem subterete to subangulate, uncinulate pubescent with upwardly directed, yellowish tri- chomes and also minutely puberulent; leaves trifoliolate, stipulate, stipules early deciduous, only the scars seen; petioles subterete to angulate, ridged and grooved, sulcate on adaxial surface, uncinulate puberulent throughout, 2.3-4 cm long, the leaf rachis similar to the petiole, 1-1.9 cm long; stipels linear attenuate, ciliate, early deciduous, 3 mm long**; petiolules stouter than the leaf rachis, ru- gose, spreading pilose with straight tapering trichomes, those of terminal leaflet 2-3.5 mm long, of lateral leaflets 1.5-2 mm long; terminal leaflets ovate acute, usually cuneate at base, apiculate at apex, the laterals elliptic, slightly oblique, rounded at the base, acute and apiculate at the apex, all minutely uncinulate puberulent on the adaxial surface, rather densely soft white pilose on the abaxial surface, the terminal leaflets 4.2-9 cm long, 3—6.7 cm wide, laterals 2.8-5.7 cm long, 1.6-3.1 cm wide. Inflorescence of axillary racemes or these forming a large, 44 Inflorescence/infructescence of isotypes from Costa Rica included in this description. 654 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 lax, spreading panicle*5, rachises angulate, ridged and grooved, minutely puber- ulent and glandular dotted, and with uncinulate pubescence of pale golden tri- chomes; pedicels regularly borne in pairs, but each 2 not always maturing to- gether, each pair subtended by an ovate acuminate striate primary bract4, minutely puberulent over the abaxial surface and somewhat pilose, the margins ciliate, ca. 3.5 mm long and 2 mm wide; secondary bracts**, each subtending 1 pedicel, rare, linear with a tuft of trichomes at the apex, 0.5-1 mm long, 0.25 mm wide; pedicels minutely puberulent with fine, short, white trichomes and sparsely to abundantly uncinulate puberulent with longer, yellowish (or white) trichomes, 4.5-5 mm long. Flowers with the calyx minutely puberulent throughout and with long stiff tapering trichomes along the whole surface of the central tooth of lower lobe, scattered on the other teeth and on the upper lobe, all teeth ciliate, the central tooth of the lower lobe 3.5—4.5 mm long, the lateral teeth 3-3.5 mm long, the upper bifid lobe 3-3.5 mm long; standard** obovate, slightly tapered toward the base, 6.5 mm long, 4 mm wide, the wings remotely rectangular in outline, truncate at the apex, slightly auriculate and short unguiculate at the base, 5 mm long, 3 mm wide, the wings scythe shaped, long unguiculate, obtuse at the apex, 5.5 mm long, 1.5 mm wide; vexillar stamen fused ca. !⁄4 its length, then free from the others. Loment stipitate, 1-2-articulate, the stipe pilose with appressed to spreading or downwardly directed tapering trichomes, 3.5-4.5 mm long; articles saccate, notched at the apex, the shoulders beaked at maturity, the margin ciliate throughout, the surface minutely puberulent, 1.7-2.3 cm long and high; seed reniform, to ca. 4 mm long and 2 mm wide, funicle to ca. 3.5 mm long. Since 1924, when Blake described Desmodium rhynchodesmum (as Meibo- mia), the only other material that has been referred to this species is one specimen from Limon Province in Costa Rica, and the two collections from Panama, one from Bocas del Toro and one from Colón. Decisions about the distinctness of D. rhynchodesmum must await the collection of more material and better field ob- servations. However, it seems this species can be reasonably well distinguished from its closest relatives, D. macrodesmum and D. saccatum by the characters cited in the key. Schindler, Repert. Spec. Nov. Regni Veg. 22: 288. 1926, equates Meibomia rhynchodesma with Nephromeria metallica (Rose & Standley) Schind- ler, which is, in my opinion, incorrect. BOCAS DEL TORO: Trail leading to ridge above Almirante, Gentry 2760 (MO). COLON: Río Iguanita, 390 m, D'Arcy 11241 (A, MO). 19. Desmodium saccatum Schubert*^*. TYPE: Panama, Dwyer 2990 (MO 1800522, holotype).—Fic. 21G-H. Vines, sometimes high climbing; stem terete, uncinulate puberulent to pubes- cent, glabrescent, finely striate, to 1 m (or more) in length. Leaves trifoliolate, * Seen only on flowering isotype from Costa Rica (US 943725). *5 Desmodium saccatum Schubert, spec. nov. A speciebus affinibus differt articulis saccatis, incisura in suturo superiori angulo acuto vel obtuso formanti, in maturitate in humeros non applanata. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 655 stipulate, stipules stramineous ovate acute to acuminate, lobed to auriculate at the base, striate and with scattered long pilosity over the abaxial surface, ciliate, early deciduous (2-)3-4 mm long, (1.5-)2-3 mm wide at base; petioles remotely to obviously sulcate adaxially, somewhat angulate, striate, moderately to abun- dantly uncinulate puberulent, usually glabrescent, 2-5.5 cm long; leaf rachis sim- ilar to the petiole, 1-2.5 cm long; stipels linear lanceolate to linear attenuate, green to stramineous, striate, puberulent, and ciliate, somewhat swollen and slightly auriculate at the base, usually early deciduous, 2.5-4 mm long; petiolules dark green, stout, rugose, densely pilose with yellowish red, straight, tapering trichomes and some short, hooked ones intermixed, those of the terminal leaflets 2-4 mm long, of the laterals 1.5-3 mm long; leaflets triangular to subrhombic in outline, the terminal leaflet usually cuneate at the base, acuminate to the apex, the laterals truncate to rounded at the base, acute at the apex, all usually mu- cronate, moderately uncinulate puberulent and moderately to abundantly ap- pressed pilose with stout trichomes on the adaxial surface, abundantly longer appressed pilose with tapering stout whitish or golden trichomes, the latter es- pecially along the midrib and the margins which are sometimes involute and always somewhat ciliate, the terminal 6-12 cm long, 3-6 cm wide, the laterals 3— 6 cm long, 2-3.3 cm wide. Inflorescence of long, lax, axillary racemes and pan- icles of ephemeral flowers; rachises uncinulate puberulent with golden trichomes; pedicels regularly borne in pairs, each pair subtended by an ovate acuminate, striate, ciliate, early deciduous primary bract, 2.5-3 mm long, 1-2 mm wide, secondary bracts rarely seen; pedicels uncinulate puberulent with slender tri- chomes, 9-12 mm long. Flowers with the calyx puberulent throughout, long stiff, appressed pilose with golden trichomes over the teeth of both the lobes, the central tooth of the lower lobe 3—4.5 mm long, the lateral teeth 2.5-4 mm long, the upper bifid lobe 2-4 mm long; corolla white to blue, standard obovate, scarce- ly clawed, 3.5-6.5 mm long, 3-5 mm wide, wings semi-elliptic, obliquely trape- zoid, short auriculate, short clawed, 4-5.5 mm long, 2.5-4 mm wide, the keel petals partially fused, roughly scythe shaped, tapering toward the base, 3.5-9 mm long, 1.5-3 mm wide at the apex; stamens fused from the base for about !^ their length, then vexillar stamen becoming free. Loment stipitate, the stipe puberu- lent, becoming glabrous, 3-5 mm long; articles green, sometimes with a pink tinge, becoming brown at maturity, densely soft pilose throughout but only mi- nutely puberulent to glabrescent over surface when mature, ciliate on sutures, 2— 2.9 cm long, 1.7-2.3 cm wide from the base of the notch to the base of the article, the notch ca. 5 mm long; seed subreniform, 2.5-6.5 mm long, 1.4-4 mm wide. The four species Desmodium infractum, D. macrodesmum, D. rhynchodes- mum, and D. saccatum are related members of a group Schindler treated as Nephromeria subgenus Nephromeria. He maintained D. axillare in Nephromeria subgenus Swartziella. At present I consider these species as members of Des- modium section Nephromeria Bentham. They are distinguished from the species Schindler treated as the genus Meibomia, which includes most of the remaining species of Panama, by the almost orbicular loment articles, dehiscence of the loment by the upper, seed-bearing suture, a vine-like habit, ephemeral flowers, and long funiculate seeds not filling the articles. Desmodium axillare differs sub- stantially from this group of species and will be discussed in detail elsewhere. 656 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 The four species noted above differ from each other as indicated in the key. Desmodium saccatum is the only member of the group having a straight-sided, broadly V-shaped notch, which does not flatten out into shoulders or become beaked at maturity. COCLE: El Valle de Anton, 860-900 m, Croat 37386 (MO); Duke & Dwyer 13899 (SCZ); Dwyer (MO); Ebinger 1114 (MO); Lewis et al. 2529 (MO); Spellman et al. 593 (MO). COLON: Santa Rita Ridge lumber road, 300-500 m, Correa & Dressler 909 (DUKE, FSU, MO): Gentry 6113 (MO); Wilbur et al. 15029 (DUKE). PANAMA: Cerro Campana, 780-875 m, Croat 25215 (A, MO). La Eneida, region of Cerro Jefe, Dressler & Williams 3954 (MO). Cerro Azul, 600 m, Dwyer 1370 (SCZ, US); Dwyer 2990 (MO). Cerro Campana, Gentry 5781 (FSU). Near Portobelo, Gentry 6325 (DUKE, MO). Cerro Campana, Witherspoon & Witherspoon 8381 (A, MO). 20. Desmodium scorpiurus (Swartz) Desv., J. Bot. (Desvaux) 1: 122. 1813; DC., Prodr. 2: 333. 1825, not D. scorpioides Micheli, Bull. Soc. Bot. Belg. 30: 289. 1891, error for D. scorpiurus. Hedysarum scorpiurus Swartz, Prodr. 107. 1788; Fl. Ind. Occ. 3: 1269. 1806. TYPE: Jamaica. Swartz (S, holotype; GH, photo). Desmodium virgatum Desv., J. Bot. (Desvaux) 1: 122. 1813, not Hedysarum virgatum Thunb. Desmodium arenarium H.B.K., Nov. Gen. Sp. Pl. 6: 412 (folio ed.): 6: 527 (quarto ed.). 1824; DC., Prodr. 2: 331. 1825. type: Venezuela, Atures, Humboldt & Bonpland 851 (P, holotype; GH, photo). Desmodium multicaule DC., Prodr. 2: 331. 1825. type: Peru. Lagasca 48 (G-DC, holotype; GH, p N Desmodium parviflorum Mart. & Gal., Bull. Acad. Bruxelles 10: 185. 1843. TYPE: Mexico, Veracruz. Galeotti 3337 (P, isotype; F, GH, photos). Meibomia scorpiurus (Swartz) Kuntze, Rev. Gen. Pl. 1: 198. 1891. Meibomia arenaria (H.B.K.) Kuntze, Rev. Gen. Pl. 1: 197. 1891. Meibomia multicaulis (DC.) Kuntze, Rev. Gen. Pl. 1: 198. 1891 Nissoloides cylindrica M. E. Jones, Contr. West. Bot. 18: 135. lajara. Jones 3784 (GH, isotype). 1935. TYPE: Mexico, Jalisco, Guada- Prostrate or repent to ascending, much branched herb: stem angulate, striate, sparsely uncinulate puberulent. Leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate, acuminate, auriculate at base, auricles of the 2 stipules usually overlapping on the opposite side of the stem from the petiole base, puberulent, long ciliate, usually persistent, 2-3.5 mm long, 1.8-3 mm wide: petioles sulcate, long pilose along the groove, otherwise angulate, striate, and uncinulate puber- ulent, 0.9-2 cm long, the leaf rachis similar, 0.2-0.5 cm long; stipels lance atten- uate, striate, ciliate, 0.8-2 mm long; petiolules of terminal leaflets 1-2 mm long, of lateral leaflets 0.8-1.5 mm long; leaflets somewhat uncinulate puberulent and appressed pilose on the adaxial surface with trichomes ca. 1 mm long, moderately appressed long pilose and remotely uncinulate puberulent on the abaxial surface, the margins ciliate, the terminal leaflet elliptic, obtuse at the base and the apex or cuneate at base, 0.9-2.5 cm long, 0.7-2 cm wide, the lateral leaflets obliquely elliptic, rounded at the base, acute at the apex, 0.8-2.2 cm long, 0.4—1.5 cm wide. Inflorescence of axillary and terminal racemes; rachis angulate, striate, uncinulate puberulent, remotely pilose with glandular trichomes; primary bracts narrowly ovate acuminate, striate, puberulent and pilose on the outer surface, puberulent on the inner surface, often reflexed at maturity, 1.5-2.5 mm long, 0.8-1 mm wide, each subtending an indeterminate number of pedicels and each pedicel further subtended by a smaller similar secondary bract borne more or less laterally, 0.8— 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 657 1 mm long, 0.3-0.4 mm wide; pedicels somewhat uncinulate puberulent and mod- erately pilose with glandular trichomes, 4-8 mm long. Flowers with the calyx minutely puberulent and stiffly long pilose throughout, teeth of the lower lobe slender, the central tooth 2-3 mm long, the lateral teeth 2-2.5 mm long, the upper lobe 2-2.6 mm long, bifid for 0.5 mm of its length; corolla exceeding the calyx, the standard obovate, deeply retuse, the 2 halves of the apex rounded, narrowed to an obtusish base, 3.5—4.5 mm long, 2.5—4 mm wide, the wings oblong, truncate above, auriculate and short unguiculate, 3.3-4 mm long, 1.5 mm wide, the keel petals scythe shaped, narrowed to a tapering base, 3.5-4.5 mm long, 1-1.5 mm wide. Loment short stipitate, the stipe 1 mm long, 7-8-articulate, the articles narrowly elliptic, isthmi almost as wide as the articles, the surfaces strongly and closely reticulate, densely uncinulate pubescent, 4 mm long, 1.5 mm wide; seed rectangular, 2 mm long, 1 mm wide. Desmodium scorpiurus ranges throughout tropical America, occurring occa- sionally as a weed in tropical areas of the Pacific, and in Africa. Its fruit and stipules serve to distinguish Desmodium scorpiurus from other American species; it seems to have no close relatives in the New World. Des- modium scorpiurus is the conserved type of the conserved name Desmodium. CAS DEL TORO: Almirante, jid ue (GH). Junction of Ríos Changuinola and Terebe, 100— 200 ft, Lewis et al. 962 (MO). CANAL ZONE: Barro Colorado Island, Bottimer 1486 (A, DAO); Croat 6923 (MO), 6972 (A, MO), 13799 (MO, SC. 15580 (A, MO); Hladick 88 oca Netting 7 (MO); Shattuck 120 (MO); Starry 278 (MO). Corozal to Pedro Miguel, Cowell 398 (NY). A ncón, Croat 9236 ( ); Greenman & Greenman 5001 (DUKE). Empire Station, Hayes 504 (BM, K). Las Cruces Trail, 75 m, Hunter & Allen 691 (G, GH, MO). Balboa Heights, Killip 3396 (US). 1⁄2 mi NW of Gamboa, 0 m, Liesner 1395 (MO). Balboa, Macbride & Featherstone 45 (US). La Boca, Mori & Kallunki 3683 (M € d Piper 5183, 5187, 5194 (all US). Río Grande near Culebra, 50-100 m, Pittier 2107 (US). Ancón, 20-80 m, Pittier 6909 (NY). Summit near Gatün, Standley 27317, 30114 (both US). Balboa, Standley 30838 (US). Darien Station, Standley 31531, 31566 (both US). cHIRIQUÍ: Gualaca, 0-100 m, Burt & Koster 122 (MO). Rabo de Puerco 8 km W of Puerto Armuelles, 50-150 m, Croat 21970 (MO). Ridge above Brazo Seco near Costa Rican border, 100-200 m, Croat 22565 (MO). Road to Alanje, 0-100 m, Koster 138 (A). Quebrada Mellize 6 mi S of Puerto Armuelles, 0—150 m, Liesner 432 (MO). cocLé: El Valle, Bottimer 1476 ‘an DAO). Penonomé, 50-1000 ft, Williams 103 (NY). COLÓN: Without other locality, Rodway (K). Colon, Rose 22064 (US). Los saNTOs: Near Sabana Grande, 0-100 m, Burt & Koster 100 (MO). Las Cruces/Macaracas, 100-200 m, Burt & Koster 101 (MO). Between Las Cruces and Macaracas on way to Tonosí, 100-200 m, Koster 101 (A). oe of Rio Pedregal, 25 mi SW of Tonosi, 2500-3000 ft, Lewis et al. 2995 (DUKE, MO, UC). P Near Chepo, 0-100 m, Burt & Rattray 13 (MO). Near Pacora, 0-100 m, Burt & Rattray 19 (MO). Between Chepo and El Llano, Croat 14494 "v Savannas near Chepo, Duke 6051 (MO). Río Pacora just below confluence with Río Corso, Duke 11997 (MO). Between Río Pacora and Chepo, Dwyer et Between Río Pacora and Chepo, Porter et al. 5135 (MO). Near San Carlos, 0-100 m, Rattray 36 (A). VERAGUAS: Atalaya/Ponuga, 100-200 m, Burt & Koster 100A (MO). Between Atalaya and Ponuga, 100-200 m, Koster 100A (A) 21. Desmodium sericophyllum Schlecht., Linnaea 12: 317. 1838. Type: Mexico, Veracruz, "San Andres et Hac. de la Laguna (Schiede)’’ (not seen). Meibomia sericophylla (Schlecht.) Kuntze, Rev. Gen. Pl. 1: 197. 1891. Perennial herb, sometimes procumbent to ascending, becoming 1 m or more long; stems angulate, ridged and grooved, densely white tomentose. Leaves tri- foliolate, stipulate, petiolate; stipules obliquely ovate attenuate, striate, puberu- 658 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 lent and pilose on the abaxial surface, abundantly ciliate, usually becoming re- flexed, often early deciduous, 5-7.5 mm long, 2-3.5 mm wide at the base; petioles sulcate, densely tomentose, 1.3—2 cm long; leaf rachis similar to petiole, 0.4-1 cm long; stipels linear attenuate, pilose, and ciliate, to 3.5 mm long; petiolules stout, densely tomentose, 2-4 mm long; leaflets softly appressed pilose and dark on adaxial surface, paler and tomentose on the abaxial surface with some retic- ulate venation evident, the leaflets ovate to elliptic, acute to obtuse and mucron- ulate at the apex, rounded to cuneate at the base, the terminal leaflet 3.5-6 cm long, 2-3.5 cm wide, the lateral leafleís 3-5 cm long, 1.5-2.5 cm wide. Inflores- cence racemose to racemose paniculate; rachis subangulate, finely ridged and grooved, uncinulate puberulent throughout; pedicels borne in pairs, each pair subtended by an ovate acuminate, striate, ciliate primary bract, puberulent and somewhat pilose on the abaxial surface, 8-9.5 mm long, 3-5 mm wide; secondary bracts minute, if present, not persistent; pedicels densely uncinulate puberulent, becoming reflexed soon after anthesis, 2.5-4 mm long. Flowers with the calyx minutely puberulent throughout, the teeth of both the lobes somewhat ciliate, especially at the apex, and with at least scattered pilosity along the central tooth of the lower lobe and on the upper lobe; central tooth of the lower lobe 4-5 mm long, the lateral teeth 3-3.5 mm long, the upper bifid lobe 3 mm long; standard obovate, slightly retuse, cuneate at the base, 5.5-9 mm long, 4-6 mm wide, the wings oblong, obtuse at the apex, unguiculate at the base, 5.5-10 mm long, 1.5- 3 mm wide, the keel petals scythe shaped, curved and truncate at the apex, long unguiculate, 5.5-10 mm long, 1.5-3 mm wide. Loment stipitate; stipe 1-2.5 mm long, to 7-articulate; articles suborbicular to subrhombic in outline, with the upper suture usually slightly curved, the lower almost angulate, surfaces uncinulate puberulent throughout, 3-4 mm long, 3 mm wide, the isthmi at least slightly eccentric, ca. 1.5 mm wide; seed subreniform, 2 mm long, 1.5 mm wide. This species is known in Panama only from Chiriquí Province. It occurs from Central Mexico southward through Central America, to Venezuela and Colombia. Two collections cited below, Tyson 5824 (FSU, MO, SCZ) and 5831 (SCZ) are somewhat atypical in leaflet shape, but conform reasonably well in other char- acters to this species. It is easy to distinguish Desmodium sericophyllum from other species of the genus in Panama by its leaflets which are velvety to the touch on both surfaces, and by its short pedicels which become reflexed at anthesis or soon after, so that, for a time at least, the loments appear to be imbricated. CHIRIQUÍ: 3.4 mi NE of Volcán, Bottimer 1528 (A, DAO). El Boquete, 1000—1400 m, Cornman 2055 (US); Maurice 715 (US). Between Cerro Vaca and Hato del Loro, eastern Chiriquí, 850-1100 m, Pittier 5386 (NY, US). 3 mi N of El Volcán on old lava flow, 5000 ft, Tyson 5744 (SCZ); 3 mi N E eg on lava flow, 5000 ft, Tyson 5824 (FSU, MO, SCZ), 5831 (SCZ); Davidse & D'Arcy 10340 22. Desmodium tortuosum (Swartz) DC., Prodr. 2: 332. 1825. Hedysarum purpureum Mill., Gard. Dict., ed. 8. Hedysarum no. 6. 1768. TvPE: Mexico, Veracruz, Houstoun 1730 (BM, holotype; GH, photo). Hedysarum tortuosum Swartz, Prodr. 107. 1788. TYPE: Jamaica, Swartz (S, holotype; GH, photo). Meibomia tortuosa (Swartz) Kuntze, Rev. Gen. Pl. 1: 198. 1891. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 659 Meibomia purpurea (Mill.) Vail in Small, FI. S.E. US. 639. 1903. Desmodium purpureum pem ) Fawcett & Rendle, Fl. Jamaica 4: 36. 1920, non Hook. & Arn. Bot. Beechey's Voy. 62. 1832. Woody herb or shrub to 3 feet tall, branching from the base; stems terete, striate, uncinulate pubescent and somewhat pilose. Leaves trifoliolate, stipulate, petiolate; stipules obliquely ovate attenuate, long persistent, often reflexed, striate, ciliate, puberulent on the abaxial surface, becoming glabrous, 3-12.5 mm long, 1-7 mm wide; petioles sulcate, ridged, pubescent as the stem, 0.75-5 cm long; leaf rachis similar, 0.45—2 cm long; stipels lance attenuate, striate, ciliate, puberulent, 1-7 mm long; petiolules stouter than leaf rachis, densely stiff pilose, 1-2.5 mm long; leaflets elliptic to ovate, mostly obtuse at the apex, cuneate at the base, with prominently reticulate venation, sparsely to moderately uncinulate puberulent and pilose on both the surfaces, ciliate, the terminal leaflet 2.4-8 cm long, 1-2.7 cm wide, the lateral leaflets 2-5 cm long, 1-2.4 cm wide. Inflores- cences axillary and terminal, racemose to racemose paniculate; rachis striate, uncinulate puberulent, moderately to densely and finely glandular pilose, becom- ing less densely so in age; primary bracts, each subtending 2 to more pedicels, narrowly ovate attenuate, striate, ciliate, minutely puberulent on abaxial surface, 5-6.5 mm long, 0.5-1.5 mm wide; secondary bracts, each subtending 1 pedicel, similar, smaller, ca. 2 mm long and 0.4—0.5 mm wide; pedicels with pubescence similar to that of rachis, stiff and ascending to spreading at maturity, 1-1.6 cm long. Flowers with the calyx minutely puberulent and abundantly pilose through- out, the teeth of both lobes ciliate, the central tooth of the lower lobe long atten- uate, 3-4 mm long, the lateral teeth 2-3 mm long, the upper bifid lobe 2-3 mm long; standard obovate, slightly retuse at the apex, gradually narrowed to the base, 2.5-3.5 mm long, 2 mm wide, the wings obtusely oblong, auriculate and short unguiculate, 2.5-3.5 mm long, 1-1.5 mm wide, the keel petals obliquely oblong, broader above, unguiculate about !^ their length, 3-4 mm long, 1 mm wide. Loment 5-7-articulate, stipitate, the isthmi central; stipe 0.5-1 mm long; articles mostly orbicular, sometimes with the margins alternately revolute and appearing rhomboidal, uncinulate pubescent throughout, 3-3.5 mm long, 2.6-3.5 mm wide; seed reniform, 1.5 mm long, 1 mm wide. Desmodium tortuosum ranges throughout subtropical and tropical America. It is found as an escape from cultivation and becomes weedy in most tropical areas of the world. The thick leaflets with prominently reticulate venation, the large persistent stipules and the long, stiff, ascending to spreading pedicels distinguish Desmo- dium tortuosum most easily from its relatives. This species is grown as a green manure in the southeastern United States and other parts of the world. CAS DEL TORO: Changuinola Valley, Dunlap 197 (GH, US). Lincoln Creek, aa negent Val- ley, push 482 (US). Bocas, Lazor et al. 2325, 2334 (both FSU, pns Without other locality, Wedel 379 (GH, MO). Isla Colón, 0-120 m, Wedel 551 (GH, MO). CANAL ZONE: Coastal Island beyond Fort Amador, D'Arcy 9235 (MO). Ancón Hill, Duke 4620 (MO). Old Miraflores bridge, Dwyer 1013 (MO). Farfan Beach, Dwyer dye (MO). Frijoles, Ebinger 308 (GH, MO). Madden Dam, Ebinger 848 (GH, MO). Ancón, Greenman & Greenman yH laii 5017 (DUKE, MO). Between Frijoles and Monte Lirio, 30 m, Killip 12166 ca NY, US). Balboa, Macbride & Featherstone 46 (US). Gamboa, Piper 5188 (US). Balboa, Standley 25522, 25562, ee (all US), Standley 27001 (MO, US). Near Gatun, Standley 27333 (US). Corozal, Standley 27360 (US). Frijoles, Standley 27636 (US). Gamboa, Stan- 660 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 dley 28345 (US). Rio Pedro Miguel near East Paraíso, Standley 30027 (C, US). Ft. Sherman, Standley 31161 (C, US). Darien Station, Standley 31532 (US). Obispo, d 31783 (US). Balboa, vedi 32106 (US). Barro Colorado Island, Standley 40950 (US). Miraflores Locks area, Tyson 1127 (SCZ). Ft. Amador causeway island, Tyson 5415 (FSU, SCZ). Nea ms Venado, Wilbur & Teeri 12965 (MO). Río Pirre near crossing of trail from El Real to Tucuti, Duke 5193 (MO). Ascent of Cerro Pirre up ees Pirre S s El Real, 750-1030 m, Duke 5363 (MO). EI Real, Lazor & Correa 3372 (FSU). LOS SANTOS: Pocrí, Dwyer 2521A (A), 2521B (MO). PANAMÁ: Punta Chame, D'Arcy 10238 (MO). Taboga Island, Dw. yer 2813 (MO); Killip 3183 (US); Standley 27964 (US). 23. Desmodium triflorum (L.) DC., Prodr. 2: 334. 1825. Hedysarum triflorum L., Sp. Pl. 749. 1753. Type: Herb. Linnaeus (LINN 921.45). 1768. VE ean triflora (L.) Poir. in Lam., Encycl. Méth. Bot. 4: 451. 1798. ("Aeschinomene ") Pleurolobus triflorus (L.) St. Hil., Nouv. Bull. Soc. Philom. 3: 192. 1812. Hedysarum granulatum Schum. & Thonn. in Schumacher, Beskr. Guineiske Pl. 362. 1827; Danske id. Selsk. Naturv. Afhndl. 3: 136. 1829; Schubert, J. Arnold Arbor. 44: 294. 1963. Desmodium triflorum var. T minus Wight & Arn., Prodr. 1: 229, 1834. modi . 1842. DM stipulaceum (Burm.) Hassk., Cat. Pl. Hort. Bot. Bogor. 274. 1844. modium parvifolium sensu Blanco, Fl. Filip., ed. 2. 408. 1845, non DC. fide E. D. Merrill, Species "Blanco anae 183. 1918. Nicolsonia reptans Meisn. Linnaea 21: 260. 1848. Nicolsonia reptans Hook. f. & Benth. in Hook., Niger Fl. 304. 1849. gotia triflo i z . Pl. : . glabrescens Kuntze, Rev. Gen. . 1: 197. 1891. . glabrescens forma coerulescens Ru. Rev. Gen. Pl. 1: 197. 1891. . glabrescens forma purpurea Kuntze, Rev. Gen. Pl. 1: 197. 1891. . glabrescens forma violacea Kuntze, Rev. Gen. Pl. 1: 197. 1891. . pilosa Kuntze, Rev. Gen. Pl. 1: 197. 1891 . pilosa forma violacea Kuntze, Rev. Gen Pl. : 197. 1891. ; eee forma flavescens Kuntze, Rev. Gen. 2 1: 197. 1891. x S = š a ~ z = ` 228 € P ^" — ° ° ° ° esmodium triflorum var. epee Hoehne, Comm. Linh. Telegr. Matto Grosso, ‘Ann. 5. Bot. 8: 73. (191 7) 1919. Low creeping herbs often with several stems from the base or rooting at the nodes, suggesting a species of Trifolium in habit and aspect; stem terete and striate to angulate, moderately to abundantly long white pilose. Leaves trifolio- late, stipulate, petiolate; stipules stramineous, obliquely ovate, long acuminate, truncate at the base, striate, somewhat pilose on the abaxial surface, ciliate, long persistent, 2-3.5 mm long, 1-1.5 mm wide; petioles sulcate adaxially, long pilose and minutely uncinulate puberulent, 3—7.5 mm long; leaf rachis similar 1-2 mm long; stipels inconspicuous, ca. 0.5 mm long, petiolules stouter than the leaf rachis, densely pilose, 0.5-1 mm long; leaflets obovate, retuse and sometimes nearly truncate at apex, cuneate at base, essentially glabrous adaxially, abun- dantly pilose and minutely uncinulate puberulent on the abaxial surface or pilose, terminal leaflet 5-10 mm long, 5-11 mm wide, the lateral leaflets similar or more nearly elliptic, 4-7.5 mm long, 4-8 mm wide. Inflorescences axillary, densely compressed, usually composed of 3-4 pairs of flowers; primary bracts ovate, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 661 short acuminate, striate, ciliate, long persistent, 2.5-3 mm long, 0.7-1.5 mm wide, each subtending 2 pedicels; secondary bracts, when rarely present, depauperate; pedicels densely long pilose or only uncinulate puberulent, 4.5-8 mm long; calyx remotely bilabiate, the teeth of both the lobes equal, each ciliate and with a long pilose, reddish midrib, 2.5-3.5 mm long; corolla with a more or less orbicular but long-unguiculate standard, 3.6—4.5 mm long, 2-3 mm wide above the claw, the wings oblong and long unguiculate, equalling the standard in length, 1 mm wide, the keel petals scythe shaped, long unguiculate, as long as the standard, 1-1.3 mm wide. Loment sessile, to S-articulate, straight or curved on the upper suture, slightly invaginated on the lower; articles uncinulate pubescent throughout, ul- timately separating at the wide isthmi, 2.5-4 mm long and wide; seed nearly orbicular, 1.5 mm long and almost as wide. This species is widespread in tropical areas. It is often introduced and appears as a weed on lawns, but is noted by some collectors as a good lawn plant. It is said to be eaten by cattle. m DEL TORO: Changuinola eda Mad 303 (US). Changuinola, Godfrey 67321 (FSU). ae , Lazor et al. 2332 (FSU, SCZ). Bocas, Lazor et al. 2355 (FSU, MO, SCZ). CANAL ZONE: Frijoles, yee 1515 (A, DAO). Chagres, Fendler 84 (GH, MO, US). Between Summit and Gam- boa, Greenman & Greenman 5264 (MO). Barro Colorado Island, Hladick 89 (MO). Ancón, Piper 5184 (US). Río Grande near Culebra, 50-100 m, Pittier 2077 (US). Barro Colorado Island, Shattuck 195 (MO). Near Gatün, Standley 27325 (US). CHIRIQUÍ: Between David and Concepcion 0-100 m Koster 132 (A). Puerto Armuelles, 20 m, Liesner 142 (MO). CocLE: Aguadulce near sea level, Pittier 4856 (US). COLON: Santa Rita Ridge, 2 mi E of Transisthmian Highway, Lewis et al. 5241 ( a SCZ). Portobelo, 5-100 m, Pittier s.n. (US); Rose 23998 (US). PANAMA: Madden Dam 75 m, Allen 2 (GH, MO). San José Island, Erlanson 589 (GH, US). Sabanas near Chepo, 30 m, Hunter & Allen 2 (GH, MO). Sabanas NE of Panama City, Paul 185 (US). Corozal Road near Panama, Standley iu (US). Taboga Island, Standley 27098 (MO, US). Hospital del Seguro Social, Taylor 64 (PMA). 24. Desmodium wydlerianum Urban, Symb. Ant. 2: 302. 1900; Schubert, Contr. Gray Herb. 129: 29, pl. 2, figs. D1—7. 1940. Type: Portorico parte orientale: Wydler 140 (G, holotype; F, GH, isotypes). D. spirale p. stoloniferum (Poir.) DC., sensu DC., Prodr. 2: 333. 1825. TYPE: Porto Rico, Bertero (G-DC; GH, photo). D. lunatum Huber, Bol. Mus. Paraense Hist. Nat. 4: 568—570, fig. 2. 1906; non Brandegee 1908. : Brazil, Chinganilla dim i del Sacramento), Huber 15 (not seen, but fig. 2 agrees well with material w) jane n lunata died Hoehne. yo Mem. Inst. Butantan Secc. Bot. 1: 39, tab. 14. 1921. M. wydleriana (Urb.) Britt., Sci. Surv. Porto Rico & Virgin Isl. 5: 403. 1924. Slender subshrubs, to 4 dm or more, often repent, the stem rooting at the nodes, angulate, striate, uncinulate puberulent. Leaves trifoliolate, stipulate, pet- iolate; stipules elliptic acuminate, striate, uncinulate puberulent over the abaxial surface, ciliate, to 6 mm long, 1.5 mm wide; petioles sulcate on the adaxial surface, striate, uncinulate pubescent and sparsely pilose, 5—4.5 cm long, the leaf rachis similar, more slender, 1.4—1.6 cm long; stipels lance attenuate, striate, uncinulate puberulent on the abaxial surface, ciliate, 1.5-3.5 mm long; petiolules stouter than the leaf rachis, densely pilose and uncinulate pubescent, 2.5-3 mm long; leaflets thin, dark green and often with a blotch above, much paler beneath, terminal leaflet ovate acuminate, truncate at the base, with undulate margins, uncinulate puberulent to pubescent on the midrib and the chief lateral veins, 662 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 similarly pubescent over the whole surface or glabrescent, pubescence of the abaxial surface similar but less abundant and less conspicuous, 7-8 cm long, 5.5- 6 cm wide, the lateral leaflets similar but slightly oblique, 5-6.4 cm long, 3-3.5 cm wide. Inflorescence of axillary racemes, rachis angulate, ridged and grooved, uncinulate pubescent; bracts early deciduous, none seen on Panama specimens, primary bracts narrowly ovate, striate, puberulent, ciliate, ca. 1.5 mm long; sec- ondary bracts similar, 1⁄4 as long; pedicels borne singly, each subtended by 1 primary and 2 lateral secondary bracts, pubescent as the rachis or with sparsely scattered pilosity in addition, 1.2—1.6 cm long. Flowers with the calyx uncinulate puberulent throughout, teeth of both the lobes somewhat appressed pilose, the central tooth of the lower lobe 2.5-3.5 mm long, the lateral teeth to 3 mm long, the upper bifid lobe to 3 mm long; standard orbicular to obovate, short unguic- ulate, to 4.5 mm long, 3.5-4 mm wide, the wings suboblong and short unguiculate, to 4.5 mm long, 1.5-2 mm wide, the keel petals remotely scythe shaped, long unguiculate, 4.5-5 mm long, 2 mm wide. Loment stipitate to 3-articulate, the stipe 1-1.5 mm long, the articles densely uncinulate pubescent on the surfaces, the dorsal suture slightly invaginated, the ventral suture rounded, isthmi eccentric and narrow, 7-11.5 mm long, 3-3.5 mm wide; seed almost naviculate in outline, 4 mm long, 2 mm wide. Desmodium wydlerianum occurs in central America, the West Indies and the northern half of South America. Originally described from Puerto Rico, this species has now been found in most of tropical America in humid forests at relatively low altitudes. The seed measurements noted above were taken from Skutch 4346 (GH, Costa Rica). CANAL ZONE: Barro deret Island, Croat 5884, 14983, 17395 (all MO). Las Cruces Trail, Tyson 1774 (FSU, MO, SCZ). DARIEN: Río Paya, trail between Paya & Payita, Stern et al. 362 (GH, MO). Río Paya, trail to Pucro, SIR et al. 413 (oH. MO, US). 19. DIOCLEA Richard H. Maxwell?? Dioclea H.B.K.,** Nov. Gen. Sp. Pl. 6: 437, tab. 576. 1824. LECTOTYPE: D. sericea H.B. Vines, woody, in clumps, or high-climbing lianas. Leaves trifoliolate, the lat- eral leaflets usually smaller than the terminal, the margins entire, revolute, with variable pubescence; stipels setaceous to filiform; stipules lanceolate or triangu- late, produced below insertion or non-produced. Inflorescences erect, usually single and axillary, fasciculate racemose, tuberculate; bracts linear to triangulate, persistent to caducous; bracteoles ovate, flabellate to orbicular, persistent or caducous. Flowers shades of violet, purple and blue; calyx tube 4-5 lobed, pu- bescent within, glabrous or variously pubescent without, the upper lobes partially or completely fused; lamina of the standard oblanceolate to somewhat orbicular, 47 Indiana University Southeast Herbarium, 4201 Grant Line Road, New Albany, Indiana 47150. 48 Several generic synonymies are listed by Hutchinson, Genera of Flowering Plants 1: 426. 1964. None of these names applies to Panamanian materia 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 663 emarginate, basally biauriculate and usually bicallose, glabrous or puberulent, carnose or membranous, the wings free, the lamina obliquely oblong to obliquely ovate, auriculate, occasionally with a spur, the keel petals fused distally, the lamina triangulate to somewhat obliquely oblong, auriculate; stamens 10, the vexillary filament free ca. 3 mm, basally, then fused to the staminal sheath, all the filaments free distally, the anthers dimorphic or monomorphic; pistil com- pressed, geniculate, the ovary villous, the upper style glabrous, the stigma cap- itate, glabrous. Fruits oblong, flat, compressed or turgid, with variable dehis- cence, pubescence variable; seeds large, few, soft-cuboidal or hard-turgid; if small, then many, hard and flat, oblong to sub-orbicular; the hilum linear or short- oblong, encircling ca. '/, to nearly 12 or to ca. % the testa. Dioclea is a pantropical genus of about 50 species with the great majority in tropical South America. Literature: M H. 1944. p preis de Venezuela, I. Papilionáceas. Min. Agric. Cria. . Bot. Bol. Téc. 5. Caracas. Buen A. 1970. Las e. vt Faseólas argentinas de los géneros Mucuna, Dioclea y Camptosema. Darwiniana 16: 175-218 a. Large coarse vines, frequently high-climbing forest lianas; stipules large, produced; anthers dimorphic (5 perfect and 5 imperfect); bracts from 6-20 mm long, caducous or occasionally semipersistent; seeds large, turgid or compressed, 1—4(—5); lamina of the standard carnose, glabrous; keel petals rostrate. b. Fruits turgid, fleshy; seeds soft, cuboidal, the hilum encircling ca. 1⁄2 the testa; leaflets with the primary lateral veins in 10—14 pairs, the apices abruptly elongate; bracts 6— 10 mm long; keel petals sharply iios. or hooded. c. Leaflets papyraceous, non-rugose above, sparsely pubescent below; calyx yel- lowish, sparsely pubescent wid short, dark brown hairs; pedicels ca. 9 mm long; stipules ca. 15 mm long 5. D. pulchra cc. Leaflets coriaceous, rugose above, densely pubescent below; calyx darker, with dense ferruginous pubescence; pedicels ca. 5 mm long; stipules ca. 25 mm long . D. aurea Fruits coriaceous to ligneous; seeds hard, globose, the hilum encircling ?/,— " the testa; leaflets with the primary lateral veins in ca. 8(—10) pairs, apices usually acute; bracts 15-30 mm long; keel petals with obtuse or truncate beaks. Flowers 2.3-3.0 cm long, borne on pedicels ca. 7 mm long; fruits ca. 3 cm thick, the upper suture paralleled by thick ribs, the lower margin oo constricted between the seeds, the exocarp obliquely fissured; the bracts spreading, t 30 mm long, pred seeds ca. 2 cm thick, reddish, frequently with a scale- like iece of rim aril persisting megacarpa dd. Flowers 1.5-2.0 cm long, borne on pedicels 2-6 mm long; fruits ca. 2 cm thick, the upper suture paralleled by thin wings, the lower margin non-constricted between the seeds, the exocarp without distinct oblique asao bracts erect eds = c or reflexed, 9-20 mm long, with appressed pubescence; see . 1.5 cm thick, dark, frequently with a woody piece of funiculus persisting. e. Inflorescences, flower buds and young fruit indumentum nigrescent fer- ruginous; bracts erect, linear, caducous; fruits with a pedicel attachment at the upper suture, the upper margin usually straight; bracteoles owe 8. cadu D. wilsonii ee. Inflorescences, flower buds and young fruit indumentum fulvous, occa- sionally ferruginous; bracts reflexed, phang” lanceolate, semipersistent; fruits with ie pedicel attachment median, the upper margin arse downcurved distally; bracteoles usually Deristelil UE. IK LT DNE reflexa 664 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 aa. Smaller woody vines (except D. paniculata), frequently in clumps in open scrub or forest edges; stipules small, non-produced; anthers uniform (10 perfect); bracts ca. 3 mm long, persistent; seeds small, flat, 5-12 (except 2-4 in D. s lamina of the standard membranous, puberulent or glabrous; keel petals not rost f. Smaller woody vines, frequently common in open are s: eden papyraceous; flowers 15-25 mm long, standard puberulent; inflorescences cial single; fruits oblong, the upper suture paralleled distinct ribs or wings to either side; seeds oblong rounded, 5-12, the hilum long lin g. Calyx tube glabrous: ; pedicels usually longer than 5 mm long; bracteoles sub- orbicular, 6-10 x caducous; keel petals with the median portion of the upper margin fim s 7. D. virgata gg. Calyx tube codi. pedicels ca. 5 mm long, usually less; bracteoles lanceolate o ovate, ca. 3x 2m rsistent or caducous; keel petals with a wide median portion of the upper ate variously oo or serrate __________ 2. D. guianensis ff. inane climbing lianas, along rivers or in ist forest areas; leaflets rigid, brittle; flowers 10-12 mm long, standard glabrous; T ease s dier double or occasion- ally icd. fruits usually elliptic, the upper suture paralleled by inp di ribs c mm to either side; seeds oval, 2-4, the hilum short oblong |... . D. diy ula 1. Dioclea aurea R. H. Maxwell*?. TYPE: Colombia, Sneidern 5555 (S, holotype). Vines, coarse, twining, climbing; stems terete, with hirsute, fulvous ferrugi- nous pubescence. Leaves with leaflets coriaceous, the lamina broadly ovate, the terminal leaflet to ca. 17 cm long, 14 cm wide, the laterals to ca. 14 cm long, 11 cm wide, inequilateral, the upper surface rugose, glabrous except the midrib, the lower surface with dense, fulvous ferruginous or aureus erect trichomes of various lengths, the apices abruptly acute, the apex ca. 7 mm long, the bases rounded to cordate, the primary lateral veins in 11—14 pairs; petioles to 11 cm long, the rachis to 5 cm long, both fulvous hirsute; stipules to 15 mm long distally, probably 10 mm long below insertion (broken), striate, villous outside, sparsely pilose inside; stipels filiform, 5-10 mm long, pubescent. /nflorescences axillary, ca. 20 cm long, ferruginous tomentose, flowering 2⁄4 of length; tubercles short clavate with globose heads; bracts lanceolate, ca. 6 mm long, hirsute ferruginous outside and inside, caducous; bracteoles oval, ca. 4 x 4 mm, ferruginous pubescent outside, glabrous inside, persistent, with similar bractlets; pedicels ca. 5 mm long. Flowers with buds upcurved slightly at the calyx lobes, flowers ca. 1.8 cm long, calyx tube ca. 8 mm long with dense ferruginous pubescence, fulvous velutinous inside and extending up the lobes, the upper lobe blunt, emarginate, ca. 5 mm long, 10 mm wide, the lateral lobe lanceolate falcate, ca. 7 mm long, 4 mm wide, the lower lobe lanceolate, ca. 8 mm long, 3 mm wide; standard obovate, ca. 15 mm long, 9 mm wide with a claw ca. 4 mm long, the wings narrowly obovate, ca. 18 mm long, 9 mm wide with a claw ca. 6 mm long, the keels semiorbicular, ca. 10 mm long with a claw ca. 4 mm long, the upper margin basally auriculate with a supramedian lobe, the lower margin rising distally ca. 9 mm, culminating in a hooded beak; stamens 10, the base of the vexillary filament free ca. 3 mm, sparse- ly pubescent, the vexillary and alternate anthers of the staminal sheath imperfect, less than 0.5 mm long, perfect anthers ca. 1 mm long; pistil straight, ca. 11 mm, geniculate, rising distally ca. 9 mm, the ovary ca. 7 mm long, subsessile, fulvous + Dioclea aurea Maxwell, spec. nov. Frutex scandens, magnus, grossus, caulibus hirsutis, fer- rugineis. Folia stipulis productis, foliolis coriaceis, pagina supra rugosa, infra dense pubescente, pilis erectis fulvi-ferrugineis, venis latualibus primarius paribus 12-14 disposites. Legumina carnosa, tur- gida, seminibus paucis, mollis 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 665 canescent, the hairs to 2 mm long, 3-4-ovulate, the style frequently ?twisted, the lower style not swollen, with long, strigose hairs, the upper style glabrous, ca. 5 mm, the stigma capitate, terminal. Fruits turgid, fleshy, oblong, to ca. 9 cm long, 4.6 cm wide, 4.5 cm thick, probably indehiscent, the exocarp yellow brown with erect ferruginous hairs, the sutures indistinct; seeds 1 to few, soft, over- grown, probably similar to D. pulchra, with the hilum nearly 2 encircling. Dioclea aurea resembles D. pulchra in the hooded beak of the keel petals, the relatively small bracts and stipules, and the fruits and seeds. The primary differences are in the leaflets and indumentum, with D. aurea possessing coria- ceous and rugose leaflets and a coarser, more dense indumentum. The new species is closely related to other Colombian collections which ex- hibit a kind of indeterminate growth in the inflorescences, bracts and even calyx lobes. This plasticity characterizes Dioclea as a whole. The fruit description is based on the Panama collection and the floral descrip- tion on the Colombian collections cited. MA. DARIEN: Cloud Forest, Casi-Cana trail on Cerro Campamiento, E of Tres Bocas, head- PANA waters of Rio Cuasi, Kirkbride (& ?Duke) 1257, (MO). COLOMBIA. CALDAS: Pueblo Rico, La Selva, 1500 m, Sneidern 5427, 5555 (both S). 2. Dioclea guianensis Benth., Comm. Legum. Gen. 70. 1837. (Ann. Wiener Mus. Naturgesch. 2: 134. 1839.) TYPE: Guyana Parima Mts., Schomburgk 83 (K, holotype; G, F, US, W, isotypes).—Fic. 22 D. panamensis Duchass. & MM ex Walpers, Flora 36: 229. 1853. TYPE: Panama, Duchassaing OET), non Seemann (188 D. comosa (Meyer) Kuntze var. QC EN (Walpers) Kuntze, Rev. Gen. Pl. 1: 179. 1891. TYPE: Panama, Kuntze 691 (NY). Vines of open areas and forest edges; stems with fuscous pubescence. Leaves with leaflets lanceolate, rhomboid to oblong, 7-10 x 4-6 cm, the apex acute, pubescent or glabrous above, usually pubescent below, chartaceous, obtuse at the base, the laterals oblique, primary lateral veins in ca. 6 pairs; petioles 4—6 cm long, with the rachis 1-7 mm long, densely puberulent, the stipels to 2 mm long; stipules triangulate, ca. 3 mm long, not produced. Inflorescences slender, to ca. 60 cm long, flowering ca. 1⁄2 the length, the tubercles sessile or short clavate with globose heads, indumentum canescent fuscous; bracts narrowly ovate to trian- gular, ca. 3 x 2 mm, usually persistent; bracteoles ovate, ca. 3 x 2 mm, mostly persistent; pedicels 2-5 mm long. Flowers 1.5-2.0 cm long, the calyx tube 6-10 mm long, membranous, sparsely sub-strigose without, canescent velutinous or subtomentose within, the upper calyx teeth fused, entire, the upper lobe obtuse, to ca. 9 mm long, the lateral lobes acute to lanceolate, to ca. 8 mm long, the lower lobe linear lanceolate to lanceolate, to ca. 9 mm long; standard orbicular or obovate, membranous, puberulent apically, bicallose or ecallose, 10-20 mm long, weakly biauriculate with a claw 2-7 mm long, the wings obliquely obovate, 10—22 mm long, usually spurred, the claw 3-6 mm long, the keel obliquely oblong, 10—18 mm long, 4-9 mm wide distally, weakly auriculate, the claw ca. 4 mm long, the upper margin variously serrate medially, not rostrate; staminal sheath and pistil subsigmoidal, the anthers perfect, monomorphic; ovary 8-12 mm long, ca. 666 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FiGuRE 22. Dioclea guianensis Benth.—A. Habit (x'4). [After Allen 4149.]—B. Flower (x3⁄4).—C. Calyx (x34).—D. Corolla.—D'. Standard (x1⁄4).—D2. Wing petal ( x 4).—D?. Keel petals (xV$).—E. Stamens (x 1).—F. Pistil (x1). [After Greenman 5133.|—G. Fruit (x14). [After Allen /.] 1.3 mm wide, short stipitate, canescent or sub-velutinous, 8—13-ovulate, the style glabrous, ca. 6 mm long. Fruits compressed, oblong, ca. 10 x 1.0-1.4 x 0.4 cm, the upper suture paralleled by thin wings or slightly thicker ribs, the lower swol- len; seeds 5-11, flat, elliptic oblong, 7-12 x 4-6 x 2 mm, hard, reddish brown, frequently maculate, the hilum encircling nearly !^ the testa. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 667 Dioclea guianensis is distributed through Central America into Mexico, north- ern and northwestern South America, as well as Trinidad. The type of Dioclea guianensis has small flowers. Panamanian specimens show a size ranging from over 1.5 cm to slightly over 2 cm in length. As the flower size increases the orbicular standard shape becomes oblong or obovate and the wings tend to assume shapes oblanceolate rather than obliquely oblong, but the spur or the protuberance opposite the auricle tends to remain. Panamanian material also shows a pubescence on the undersurface of the leaflets that is sparse and erect rather than the denser appressed to dense sericeous pubescence of some Guyanan specimens. There are variants in Belize with flowers approaching 3 cm long that I have not seen in Panama Pittier in 1944 and afterwards attempted to segregate several species from the D. guianensis complex, but he did not supply Latin diagnoses for the names, leaving them not validly published. After an extensive study of the specimens he examined, I conclude that to recognize these taxa is untenable taxonomically and would serve no useful purpose. Our primary area of disagreement is in the weight placed on flower size and indumentum. I place little weight on these characters because of the tremendous variance present. All the D. guianensis specimens I have seen from Panama can be grouped under var. guianensis sensu lato. Dioclea guianensis can be separated from D. virgata by its ovate bracteoles, pubescent calyx, serrate upper keel margin, and smaller, narrower fruits. Dioclea guianensis is distinguished from D. paniculata in its more open habitat, habit, larger flower size, serrate upper keel margin, papyraceous leaflets, somewhat oblong seed shape, and linear hilum. WITHOUT EXACT LOCALITY: Duchassaing (GOET, P); Kuntze 691 (NY); Seemann (BM); See- mann 461 (GOET-mounted sheet, p in envelope). BOCAS DEL TORO: Río Changuinola, Dwyer 4078, 5752 (both MO). CANAL ZONE RO COLORADO ISLAND: Croat 4841, 7147 (both MO); Shattuck 134 (MO); Wetmore & Abbe 61 (F); "Wilson 123 (MO); Woodworth & Vestal 691 (LE, MO), 726 (A, MO). Pedro Miguel, Allen 1 (BR, G, F, MO, U). Between Miraflores & Panama, Blum 2069 (FSU, MO). Cerro Ancón, Celestine 74 (US). Dirt road to Chiva-Chiva town, Correa 493 (FSU, MO). Gaillard Highway NW of Summit Gardens, Croat 8883 (MO). 2.5 mi N of Summit Gardens, Croat 12833 (F, MO). Gaillard Highway near Gamboa, Croat 13139 (MO). 1-2 mi NW of Summit Gardens, Croat 13917, 14045 (both MO). K-10, Duke 15194 (MO). Fort Sherman, beach, Dwyer 6585 (SIU). C-21 Road, Dwyer 8382 (MO, SIU). Ancón Hill, Greenman & Greenman 5133 (MO). ¥2 mi W of Summit Gardens, Hansen 3008 (MO). Summit, Hayes 9 Feb. 1862 (BM). Empire Station, Hayes 562 (BM, M). Hill W of Curundá, Harvey 5233 (F). Gamboa, Heriberto 86 (US). Without other locality, Hicken 1907 (SD). Military Road near Summit Gardens, Hladik 133 (MO). Las Cruces trail, Hunter & Allen 766 (G, MO). Without exact location, Johansen Jul. 1924 (US). Gamboa Road, Jones 307 (US). % mi N of Gamboa, Lazor & Tyson 5659 (MO). 2 mi N on Panamerican Highway towards Nuevo Emperador, Lewis et al. 5222 (MO). Chiva-Chiva Trail 2 mi above Red Tank, Maxon & Harvey 6592 (US). 1% mi N of Summit Gardens, Nee 9515 (MO). Between Rodman Marine Base & Chorrera, Nowicke et al. 3578 (MO). Chiva-Chiva Trail Red Tank to Pueblo Nuevo, Piper 5174 (F, NY, US). Empire to Wege Piper 5175 (US). Culebra, Pittier 2148 (US). Between Corozal & Ancón, Pittier 2176 (NY, US). Summit, Standley 26908 (US). Old Las Cruces Trail between Ft. Clayton & Corozal, Ln e 29107 (US). Balboa, Standley 29296 (US). Río Pedro Miguel near East Paraíso, Standley 30020 (US). Summit, Standley 30097 (A, US). Road K-9 near Río Cocolí, Stern et al. 31 (MO). Quebrada Bonita, Steyermark & Allen 27 Dec 1934 (MO). 5 mi N of Cocolí, Tyson 3878 Toro: Río Changuinola, Dwyer 4078, 5752 (both MO). cHiRIQUÍ: Near Remedios, Burt & Koster 166 (MO). ye mi N of David, Lewis et al. 697 (MO, NY, UC, US). Boca Chica, Pittier 5121 (US). OCLÉ: Antón- Pm roadside, Burt & Rattray p bis 3 mi NE of Antón, D'Arcy 4109 (MO). = Q Dwyer 2030 (SIU). 2 mi E of Río Hondo, along Panamerican Highway, Gentry 2913 (MO). 7.5 mi from Penonomé towards Caimito, McDaniel A Cook 14816 (MO). Penomoné, R. S. Williams 365 (NY, US). coLów: Río Chagres, above Gamboa, Allen 4149 (G, MO). Calzada Larga, de Gracia 668 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 11 (MO). PANAMA: Campana, Allen 4033 (G, MO). Cerro Azul, Cotterman & Klawe P-15 (MO, US). Isla del Rey, Duke 9563 (MO). Between Rio Sulugandí & Rio Espavé, Gentry 4983 (MO). Lidice near the river, Jaramillo 14 (FSU). Savanas of Rio Yguana, J. F. Macbride 2630 (F, US). Panamerican Highway at Continental Divide, Nee 8289 (MO, US). Sabanas N of Panama City, Paul 600 (US). La Chorrera, Paz 20 (MO). Bella Vista, Piper 5176 (US). Sabana de Juan Corso near Chepo, Pittier 4741 (BM, NY, US). Cerro Campana, Porter et al. 4298 (MO). Between Río Pacora & Chepo, Porter et al. 5159 (MO). Punta Paitilla, Standley 26228 (US). Juan Franco Race Track, Standley 27767 (US). Río Tapia, Standley 28182 (US). Matias Hernández, Standley 28912 (US). Tumba Muerto Road, Standley 29786 (US). Juan Díaz, Standley 30551 (US). Between Matías Hernández & Juan Díaz, Standley 32010 (US). Road to Monte Oscuro, Varela 26 (MO). 3. Dioclea megacarpa Rolfe, Kew Bull. 139. 1901. TYPE: Trinidad & Tobago, Trinidad, St. Annes, ?Cangrehol, Hart 6406 (K, lectotype; photos C, F, NY, S, SI of type at K). D. reflexa, sensu Fawcett & Rendle, Fl. Jamaica 4(2): 59, tab. 18. 1920. non Hook. f. See Amshoff loc. cit. Vines, coarse, high climbing; stems pilose. Leaves with the lamina broadly elliptic to obovate orbicular, to ca. 20 cm long, 12 cm wide, the apex with an abrupt acumination, ca. 3 mm long, usually pubescent above, often rough with persistent hair bases, sparsely velutinous below, chartaceous, slightly cordate at the base, the lateral veins oblique, the primary lateral veins in 7-10 pairs; petioles sparsely pilose, the rachis to ca. 5 cm long, stipels 4-7 mm long; stipules lanceo- late, to 35 mm long, including region below insertion, densely pilose outside. Inflorescences stout to ca. 70 cm long, flowering on !^ the length, the tubercles sessile, the indument ferruginous; bracts linear lanceolate, ca. 20 mm long, finally caducous, spreading or somewhat reflexed, pilose; bracteoles flabellate to orbic- ular, 2-4 x 2-4 mm, ferruginous without, persistent; pedicels ca. 7 mm long. Flowers with the buds somewhat incurved, the flowers ca. 2.5 cm long, occa- sionally to ca. 3 cm long; calyx tube 12-15 mm long, carnose, ferruginous pu- bescent without, velutinous within, the upper calyx lobes partially fused, emar- ginate, ca. 7 mm long, 10 mm wide, the lateral lobes acute, ca. 7 mm long, the lower lobe lanceolate, ca. 10 mm long; lamina of the standard obovate orbicular, carnose, glabrous, bicallose, to ca. 25 mm long, the auricles semi-orbicular, re- flexed, the claw ca. 5 mm long, the wings broadly obliquely oblong, to ca. 25 mm long, the claw ca. 5 mm long, the keel triangulate, ca. 8 mm long, ca. 9 mm wide distally, auriculate, the claw ca. 5 mm long, the upper margin slightly undulate, culminating in a truncate beak; stamens 10, the vexillary anther and the inner alternate anthers of the staminal sheath imperfect, the perfect anthers ca. 0.5 mm long; ovary short stipitate, villous, 2—4(—5)-ovulate, the upper style glabrous, ca. 5 mm long, the lower part bulbous. Fruits oblong, frequently somewhat arcuate, ca. 18 x 6 x 3 cm, partially dehiscent, the fuscous pubescence usually persistent, the upper suture indehiscent, paralleled by the thick ribs, the lower margin trun- cate, frequently constricted between the seeds, the suture becoming sulcate, the exocarp with oblique fissures; seeds (1—)3-4(—5), sub-orbicular, ca. 3 x 3 x 2 cm, hard, reddish, the hilum encircling nearly 24 the testa. Rolfe's syntypes included collections from Paraguay. These syntypes were removed by Hassler and others were added to form the basis of Dioclea para- guariensis Hassl., Fedde, Rep. Spec. Nov. 16: 228. 1919. Burkart (1952) placed 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 669 D. paraguariensis in synonymy under the name D. megacarpa Rolfe, but after reexamining the 2 taxa, he restored D. paraguariensis to species rank as separate from the more northern D. megacarpa. See Burkart (1970), in Darwiniana 16(1— 2): 182—186, Figs. 4, 5, for his explanation and excellent illustrations of D. par- aguariensis. I am not as sure that D. paraguariensis and D. megacarpa are distinct taxonomically although I have not found where their distribution over- laps. I have also found that D. violacea Mart. ex Benth. (1839) with Pohl 338 (W) as lectotype is an earlier name for D. paraguariensis Hassl. (Note synonymy under D. wilsonii below). It seems best to maintain D. megacarpa as a good species. The long spreading bracts, large flowers, fruits, fruit sutures, exocarp, and large reddish seeds, frequently with a papery, persistent piece of rim aril, are characters which distinguish this taxon. (Useful reference: Janzen, D. H. 1971. Escape of Juvenile Dioclea megacarpa (Leguminosae) Vines From Predators in a Deciduous Tropical Forest. Am. Natur. 105: 97-112.) Duke reports the native name ‘‘Ojo de Buey.”’ NAL ZONE: Gov. Reservation, Ancón, Bangham 608 (A). cocLÉ: Llano Bonito, N of Marga- ritas, Seibert 540 (A, MO, NY). DARIEN: Chepigana, Duke & Bristan 268 (MO, NY, US). PANAMA: Common along the beach, P 1685 Ns O). San José Island, Erlanson 408 (G, NY, US); Harlow 3 (US); Johnston 53 (US), 360 (GH), 568 (US). Punta Paitilla, Standley 30814 (US). SAN BLAS: Isla Pino near Mulatupo Island, Elias 1707 (MO, SIU, US), 1715 (M + . Dioclea paniculata Killip ex R. H. Maxwell, Phytologia 40: 243. 1978. TYPE: Colombia, Cundinamarca, Quebrada Cabana, Hacienda El Cucharo between Tocaima and Pubenza; 380-600 m, Killip et al. 38329 (US, holotype; COL, US, isotypes). Twining vines, high climbing; stems terete, sparsely pubescent. Leaves with the lamina widely lanceolate, elliptic, ovate, occasionally obovate, 6.5-14 cm long, 2.5-8 cm wide, the lateral leaflets smaller, inequilateral, the apices acute, obtuse or abruptly acuminate with the extension 10-15 mm long, the bases round- ed, occasionally slightly cordate, the upper surface shining or dull, glabrous, sparse pubescent below or with a few hairs on the primary veins, with ca. 6 pairs of primary lateral veins; rachis 7-18 mm long, '/;—'/, the petiolar length, with dense ferruginous pubescence when young, quickly glabrous; stipels setaceous, ca. 1 mm long, usually persistent; stipules linear or lanceolate, sometimes deeply bifurcate or trifurcate, the lobes linear, the middle longest, 1.5-4.0 mm long, not produced, frequently with secondary bracts surrounding the emerging inflores- cence and leaf cluster. /nflorescences terminal or axillary, emerging with the leaves, single, double, occasionally branched, 3-18 cm long; each tubercle with 3-8 flower buds, the tubercles clavate with the heads usually elongate upwards and swollen; primary bract at the base of the tubercle linear lanceolate, ca. 2 mm long, glabrous, caducous; bracteoles ovate to orbicular, ca. 1.8 mm long, semi- persistent; flowers subsessile. Flowers blue violet to deep purple, 10-12 mm long; calyx tube ca. 3 mm long, ferruginous pubescent, the upper lobes entire or emar- ginate, the lobes subequal, 1.5-2 mm long; standard with the lamina obovate, weakly auriculate or not callose, striate, ca. 8 mm long, 6.5 mm wide, glabrous, the claw ca. 2 mm long, the wings oblanceolate or somewhat oblong, auriculate, 670 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 striate, ca. 6 mm long, 2.5 mm wide, glabrous, the claw ca. 2.5 mm long, the keel petals somewhat oblong to obliquely oblong, ca. 6.5 mm long, 2 mm wide, gla- brous, the claw ca. 2.5 mm long, the upper margin entire, not rostrate; vexillary filament often weakly fused to the staminal sheath, the anthers perfect, mono- morphic; ovary ca. 6 mm long, short stipitate, densely villous with rigid, white hairs interspersed with dark brown hairs, ca. 5-ovulate, the style upcurved or ascending, glabrous for ca. 2 mm, shortly exserted through the keel petals. Fruits elliptic, flat, dehiscent, sessile, 5.5-8.5 cm long, 2.1-2.4 cm wide, 2.5-5 mm thick, exocarp of young fruits with dark brown pubescence, finally glabrous, the upper suture slightly raised with shallow parallel ribs ca. 1 mm to either side, the lower margin somewhat swollen; seeds 2-4, flat, oval, brownish, smooth, shining, ca. 13 mm long, 10 mm wide, 3 mm thick, surrounded by papery, red, packing tissue, the hilum oblong, 2.5-3.0 mm long. This species is fairly common in Colombia. The Panama collections are not good specimens and most of the species description is from Colombian material. ANAL ZONE: 4 km W of Gatun Dam, 190-200 m, Nee 8911 (MO, PMA). COLÓN: 15 km E of Colon, Santa Rita lumber road, Dressler 3797 (MO). CA . Dioclea pulchra Moldenke, Phytologia 1(1): 6. 1933. TYPE: Colombia, Boyaca, El Umbo & Mt. Chapon region, Lawrance 528 (A, F, G, MO, NY, S, U, UC, US). Vines, high climbing lianas; stems pilose. Leaves with leaflets broadly elliptic to obovate, to 18 x 11 cm, the apex acuminate, ca. 15 mm long, sparsely ap- pressed strigose above and beneath, chartaceous, slightly cordate truncate to rounded at the base, the lateral veins oblique, primary lateral veins 10—12 pairs; petioles canaliculate, strigose-pilose, the rachis to ca. 5 cm long, stipels ca. 7 mm long; stipules lanceolate, to 14 mm long, produced for ca. 6 mm below insertion, pubescent without. Inflorescences stout, 15-40 cm long, flowering 2⁄4 the length, the tubercles subsessile to clavate; bracts linear, usually emerging from the tu- bercles, occasionally persistent, erect, ca. 6 mm long, fuscous ferruginous pu- bescent; bracteoles broadly ovate or flabellate, 2 x 2-3 mm, fuscous ferruginous without, persistent; pedicels ca. 9 mm long. Flowers ca. 2 cm long; calyx tube yellowish, 7-10 mm long, carnose with sparse fuscous ferruginous pubescence without, velutinous within, the upper lobes partially fused, emarginate ca. 5 mm long, 8 mm wide, the lateral lobes slightly falcate, lanceolate, ca. 6 mm long, the lower lobe broadly lanceolate, ca. 9 mm long; standard broadly obovate rotund, carnose, glabrous, bicallose, to ca. 16 x 17 mm, the auricles semi-orbicular, re- flexed, the claw ca. 3 mm long, the wings obliquely oblong ovate, ca. 16 mm long, ca. 8 mm wide, the claw ca. 5 mm long, the keels triangulate, ca. 8 mm long, ca. 11 mm wide distally, the auricle inconspicuous, the claw ca. 4 mm long, the upper margin usually with 1 prominent lobe, the beak narrow or hooded; staminal sheath sub-sigmatoidal, the anthers dimorphic, perfect anthers ca. | mm long, the middle anther frequently 1.5 mm long; ovary short stipitate, densely argenteous fuscous villous, (2—)5(—6)-ovulate, the upper style glabrous, ca. 5 mm long. Fruits green, fleshy, cylindrical oblong, ca. 15 x 5 x 4 cm, ?indehiscent, fuscous or ferruginous velutinous, the upper and lower suture inconspicuous; 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 671 seeds 2-4, cuboidal, 3.5 x 3.5 x 2.5 cm, soft, light brown, maculate, the hilum encircling nearly 1⁄2 the testa. This species and the next are members of the Dioclea series with fleshy fruits. This species in Panama is considered conspecific with D. pulchra of the Colom- bian Andes flora rather than D. malacocarpa Ducke of the Amazonian flora. These two species are so close I had, at one time, considered D. pulchra a variety, but since then it has become apparent there are several fleshy fruited species that have evolved down the Andean chain as well as extending into Central America. A common name ''Mandeva" is reported from Darién (Kirkbride & Duke 1140). The fleshy fruits with soft seeds, small bracts, thin leaflets with a high number of primary lateral veins, and keel petals frequently with hooded beaks separate this and the following species from other high, climbing, forest lianas in Dioclea. WITHOUT LOCALITY: Carleton 259 (US). BOCAS DEL TORO: Changuinola Valley, Dunlap 101 (F). Río Seo ara Dwyer (MO). Almirante, McDaniel 5110 (MO). Changuinola, island potrero, Stork, 1923 (UC); bai C78 (UC, US). Water Valley, Wedel 786 (MO). Chiriquí Lagoon, Wedel sid (GH, : La Boca de Pirre, Bristan 1274 (NY). Río Tuira, between Río Punusa & Río Mangle, Duke "14590 (MO, NY. El Real, Río Cuasí S of Tres Bocas, Kirkbride & Duke 1140 (MO). El Real, Río Tuira, Stern et al. 121 (MO, U 6. Dioclea reflexa Hook. f., Niger Fl. 306. 1849. Type: W. Africa, Cape Palmas and region of Fernando Poo, Vogel 32 (K, holotype). Canavalia reflexa (Hook. f.) Wright, Sauv. Cub. 535. vagi Fide Sauer, Brittonia 16(2): 179. 1964. Dioclea panamensis Seemann in Hemsley, Biol. Centr. Amer. Bot. 1: 302. 1880, nomen nudum. Based on Fendler 83 (GH, K, MO, US) & penas, 455 (BM), non Duchass. & Walpers (1853). Coarse vines; stems glabrate to fuscous pilose. Leaves with leaflets broadly elliptic to obovate, 5-23 cm long, 3-13 cm wide, the apex acute or shortly acu- minate, usually glabrous except the primary veins, reticulate above, sparsely pubescent below, chartaceous, obtuse at the base, the laterals oblique, primary lateral veins in ca. 8 pairs; petioles pilose, the rachis 2-3 cm long, the stipels filamentous, 3-14 mm long; stipules lanceolate, to ca. 22 mm long, produced to ca. 10 mm below insertion, pilose without. Inflorescences stout, to ca. 45 cm long, flowering 1⁄2 the length, the tubercles clavate, the head elongation variable, the indumentum usually fuscous ferruginous, occasionally the buds nigrescent; bracts lanceolate, 10-20 mm long, ca. 3 mm wide, occasionally persistent, re- flexed, with appressed aureus-cinereous pubescence; bracteoles flabellate, 2-3 x 2—4 mm, fuscous puberulent without, persistent; pedicels 3-6 mm long. Flowers with straight buds, ca. 1.5-2.0 cm long; calyx tube ca. 7 mm long, carnose, fuscous ferruginous pubescent without, fuscous velutinous within, the upper ca- lyx lobes partially fused, emarginate, 2-5 mm long, ca. 7 mm wide, the laterals acute, ca. 5 mm long, the lower acute, ca. 6 mm long; standard rotund orbicular to obovate, carnose, glabrous, bicallose, ca. 15 mm long, the auricles semi-or- bicular, reflexed, the claw ca. 3 mm long, the wings obliquely oblong to obovate, ca. 10 mm long, the claw ca. 5 mm long, the keel triangulate, ca. 9 mm long, ca. 10 mm wide distally, weakly auriculate, the claw ca. 5 mm long, the upper margin undulate or with 1 prominent lobe, culminating in an obtuse beak; stamens 10, 672 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 the vexillary and inner, alternate anthers of the staminal sheath imperfect, the perfect anthers ca. 1 mm long; ovary shortly stipitate, argenteous villous, 2-4 (-5)-ovulate, the upper style glabrous, ca. 6 mm long, the lower part bulbous. Fruits compressed, elliptic oblong, to ca. 17 x 5.5 x 2.4 cm, dehiscent, the pedicel attached medially, somewhat convex above, naviculate below, glabrate, the upper suture paralleled by thin wings, the lower margin swollen, sulcate; seeds (1—)2— 4, semi-orbicular to oblong, ca. 30 x 25 x 14 mm, hard, dark brown, frequently maculate, the hilum encircling ca. ?4 the testa. Dioclea reflexa is distributed pantropically. This is due to a flotation adapta- tion of the seeds which are occasionally found in the drift. These seeds however, cannot be distinguished from those of D. wilsonii and possibly other species, such as D. javanica Benth. of Asia. Dioclea reflexa is difficult to distinguish from D. wilsonii and there may be some gene exchange. In general D. reflexa would be found at low altitudes, but not in the drift flora. Its indumentum ranges from light fulvous to ferruginous, but never as dark as D. wilsonii. The reflexed bract attitude, upon which the epithet is based, is a conservative character, but the width and indumentum are variable. The tendency for bracts and bracteoles to persist is also variable. The lower margin of D. reflexa fruits becomes sulcate and partially opens, but not enough to allow the seeds to disperse. This character can be seen in most Pan- amanian material, but may not hold world wide. The finally glabrous exocarp of D. reflexa is smooth, somewhat thicker than that of D. wilsonii and is frequently constricted somewhat between the seeds and also along the lower margin. The pedicel attachment of D. wilsonii is invariably at the upper margin, which is straight, while that of D. reflexa is variable, and fruits with a median attachment may be obovate in shape. For flower color, seasonal aspects and other field characters of D. reflexa, D. wilsonii (as D. violacea) and D. megacarpa, see Johnston (1949). Johnston, I. M. 1949. The Botany of San Jose Island. Sargentia 8: 150—153, Plate 8, Fig. 4-5. Ducke, A. 1925. Arch. Jard. Bot. Rio de Janeiro 4: 97, tab. 4, D. violacea fide Ducke = D. wilsonii. WITHOUT LOCALITY: Seemann (GOET). BOCAS DEL TORO: Chiriquí Lagoon, Johns Creek, Wedel 2760 (GH, MO); CANAL ZONE: Barro Colorado Island: Croat 5017 (MO, US), 5062, 7387, 8110, 8390, 12762, 13118, 14884 all MO); Wetmore & Abbe 64 (MO). Skunk “Anne Blum & Dwyer 2109 (MO). Westerly arm of Quebrada Salamanca, Dodge et al. 16995 (BR, G, GH, MO, S). Skunk Hollow, Dwyer 5435 (FSU, MO, SIU). Fort Sherman, Dwyer 6621 ao France Field, U.S. Mine Emplace- ment Center, Dwyer 8539 (MO). Chagres, Fendler 83 (GH, MO, US). Near Gatun, Gaillard 1911 (US). Rio Pequeni, a 455 (BM). cHIRIQUÍ: Burica Peninsula, Guanabano, Croat 22526 (MO). COLÓN: near Salamanca, Gentry 6702 (MO). Between Rio Piedras & Puerto Pilon, Lewis et al. 3221 (MO). DARIEN: Rio “Chun naque between Yaviza & junction with the Tuira, Gentry 13484 (MO). PAN- AMA: San José Island, Johnston 615, 653, 711 (all GH), 1207 (GH, S), 1254 (GH); Kennedy 2275 (MO). Drowned forest of Quebrada Ancha, Steyermark & Allen, 21 Dec 1934 (MO). SAN BLAS: Ailigandí, Dwyer 6832 (MO, SIU). 7. Dioclea virgata (L. C. Rich.) Amshoff, Meded. Bot. Mus. Herb. Rijks. Univ. Utrecht 52: 59. 1939. Dolichos virgatus L. C. Rich., Act. Soc. Hist. Nat. Paris 1: 111. 1792. rvPE: French Guiana, not seen. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 673 Mucuna virgata Desv., Ann. Sci. Nat. (Desvaux) 1: 423. 1826. TYPE: French Guiana, L. C. Richard(?) herb. Desvaux (P). Dioclea lasiocarpa Mart Benth., Comm. Legum. Gen. (Ann. Wiener Mus. Naturgesch. 2: 133. 839) 69. 1837. TYPES: Brazil: Minas Gerais, Pohl 3222 (W); Pará, Martius (BR, M); Bahia, Martius 2016 (M). Vines of open areas and forest edges; stems with canescent ferruginous pu- bescence. Leaves with leaflets broadly lanceolate, ovate to narrowly obovate, 7— 10 x 4-6 cm, the apex acute, usually glabrescent above with variable pubescence below, chartaceous, obtuse at the base, the laterals oblique, primary lateral veins in ca. 6 pairs; petioles 4—6 cm long, the rachis 4(-10) mm long, densely puberulent to strigose; stipels to 2 mm long; stipules triangulate, ca. 3 mm long, not produced. Inflorescences slender, to ca. 70 cm long, flowering about V? the length, the tubercles subsessile to elongate, inflorescence indumentum canescent subtomen- tose to glabrescent; bracts triangulate rhombic, ca. 3 x 2 mm, persistent; brac- teoles sub-orbicular, 6-10 x 4-6 mm, caducous; pedicels (5-)7(-15) mm long. Flowers ca. 2.5 cm long, the calyx tube ca. 8 mm long, membranous, glabrous without, canescent velutinous within, the upper calyx teeth fused, entire, the upper lobe acute obtuse, ca. 8 mm long, the lateral lobes falcate, broadly lan- ceolate, ca. 6 mm long, the lower lobe lanceolate, ca. 8 mm long; standard or- bicular obovate, to ca. 25 mm long, membranous, puberulent apically, bicallose, biauriculate, the claw 7 mm long, the wings obliquely obovate, to 22 mm long, usually spurred, the claw 7 mm long, the keel obliquely oblong to nearly oblong, to ca. 20 mm long, ca. 10 mm wide distally, weakly auriculate, the claw ca. 5 mm long, the upper margin with 7-10 medial fimbriae, ca. 2 mm long; stamens 10, glabrous, anthers uniform; ovary ca. 1l-ovulate, canescent velutinous, the style glabrous distally for ca. 7 mm. Fruits flat, oblong, usually obtuse basally, straight above, naviculate below, frequently with a falcate apex, to ca. 11.0 x 2.0 x 0.4 cm, dehiscent, with erect, stiff, ferruginous pubescence, the upper su- ture paralleled by thin wings, the lower swollen; seeds 7-12, flat, elliptic oblong, ca. 10 x 6 x 2 mm, hard, reddish brown, frequently maculate, the hilum encir- cling nearly 1⁄2 the testa. This species is widespread throughout tropical South America and extends through Central America into Mexico. The sub-orbicular, caducous bracteoles, glabrous calyx and fimbriate upper keel margin are characters which readily separate D. virgata from the closely related D. guianensis with its forms and varieties. ANAL ZONE: Pipeline road, Croat 12371 (MO); Gentry & Dvessler 1974 (MO). DARIEN: Near Pinas, Duke 10618 (MO). Rio Areti, Duke & Nickerson 14907 (MC). PANAMA: El Llano-Carti Road, 5 km from Panamerican Highway, Witherspoon 8720 (MO) 8. Dioclea wilsonii Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 310. 1929. TYPE: Honduras, Point Triunfo, near Tela, P. Wilson 336 (F). D. d. ea sensu Mart. Fl. Bras. 15(1): 162. 1859 (based on collec ions such as Guillemin (K)); non n a Mart. ex Benth. (1837), based on Pohl 338 (W lectotype) and Martius (BR paratype). D. Pye ee ‘Rock, Leg. Pl. Hawaii 201, rab. 83. 1920, non Velloso, Fl. Flum. 296. 1825; Icon. 7 tab. 77. 1831. D. us ea auct. mult., non Benth. 1837. 674 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Coarse vines; stems glabrescent to densely ferruginous pilose. Leaves with leaflets broadly elliptic to obovate, 7-16 cm long, 4-11 cm wide, the apex abruptly acute, usually glabrous except the primary veins, reticulate above, sparsely pu- bescent below, chartaceous, obtuse at the base, the laterals oblique, primary lateral veins in ca. 7 pairs; petioles sparsely strigose to pilose, the rachis 2-3 cm long, stipels to 15 mm long; stipules lanceolate, 15-30 mm long, produced to ca. 10 mm below insertion, pilose and ciliate without. Inflorescences to 45 cm long, flowering more than ! the length, the tubercles clavate, the head elongation variable, the inflorescence indumentum ferruginous nigrescent; bracts linear, erect, ca. 8 mm long or longer, caducous; bracteoles broadly ovate to flabellate, 2-3 x 4 mm, caducous; pedicels 2-6 mm long. Flowers with straight buds, ca. 1.5 cm long, calyx tube ca. 7 mm long, carnose, ferruginous-nigrescent pubescent without and within, the upper calyx lobes partially fused, emarginate, ca. 5 mm long, ca. 9 mm wide, the laterals acute, ca. 6 mm long, the lower acute, ca. 7 mm long; standard rotund orbicular, carnose, glabrous, bicallose, to 15 mm long, the auricles semi-orbicular, reflexed, the claw ca. 3 mm long, the wings obliquely oblong to obovate, to ca. 15 mm long, the claw ca. 5 mm long, the keel triangulate, ca. 8 mm long, ca. 9 mm wide distally, auriculate, the claw ca. 5 mm long, the upper margin undulate, culminating in an obtuse beak; stamens 10, the anthers dimorphic, the vexillary anther and the inner, alternate anthers of the staminal sheath imperfect, perfect anthers ca. 1 mm long; ovary stipitate, argenteous vil- lous or bicolored, 3—4-ovulate, the upper style glabrous, ca. 5 mm long, the lower part bulbous. Fruits compressed, oblong, to ca. 10 x 6 x 1-2 cm, pedicel at- tached along the upper suture, straight above, naviculate below, indehiscent, glabrate, the upper suture paralleled by thin wings, the lower margin thin; seeds 2-4(-5), semi-orbicular to oblong, ca. 30 x 25 x 15 mm, hard, dark brown, the hilum encircling ca. 3⁄4 of testa. In trying to match fruits with flowers, Bentham mixed fruits of Dioclea wil- sonii with flowers of D. violacea in his D. violacea description in Flora Brasi- liensis. The taxonomy was confused apparently until Standley named D. wilsonii. At least this was the opinion of the late C. V. Morton of the U.S. National Herbarium when I discussed the nomenclatural problem with him. Dioclea vio- lacea, on the basis of the syntypes Bentham used in his original description, is the same as D. paraguariensis Hassler and very closely related to D. megacarpa. Dioclea violacea is about as closely related to D. megacarpa as D. wilsonii is to D. reflexa, but the latter are sympatric while the former are apparently not. Dioclea wilsonii is found in eastern Brazil, throughout northern South America, in some of the West Indies and throughout Central America into Mexico. It is also found in the Hawaiian Islands and some of the Pacific Islands and I have annotated collections from the Malagasy Republic (Madagascar) as D. wilsonii (as possible introductions?), but Madagascar plants may prove not to be the same species. It is generally assumed that D. wilsonii seeds are distributed as part of the oceanic drift as are D. reflexa seeds from which they cannot be distinguished. Actually collections show D. reflexa to be more widely distributed and that much of the D. wilsonii distribution could be due to man rather than drift. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 675 This species can be separated from the closely related D. reflexa in possessing a very dark inflorescence indumentum, erect bracts, and an indehiscent fruit. BOCAS DEL TORO: Water Valley, Wedel 861 (GH, MO, US). Chiriqui Lagoon, Wedel 1054, 1155 (both GH, MO), 1688 (GH, MO, US), 2147, 2728 (both MO). c:.NAL ZONE: Barro Colorado Island, m (A, j (MO); Shattuck 441 (F, US); Standley 40992a (US, fruit only). COLÓN: T France Field and Catival, per 30390 (US). HERRERA: near Las Minas, along Río Las Trancas, Stern et al. 1802 (MO, US). PANAMÁ: Empire Station, Hayes 313 (BM, K). San José Island, A 418 (BM, GH, MO, P, U, US), 6/2 (GH), //70 (BM, GH). 20. DIPHYSA Peter S. White? Diphysa Jacq., Enum. Syst. Pl. Carib. 7. 1760. TvPt: D. carthagenensis Jacq. Trees or shrubs, unarmed or with spinescent branches; branchlets, pedicels, and/or calyx glandular viscid in some species. Leaves alternate, imparipinnate, petioled; lateral leaflets opposite, subopposite, or 2lternate, entire, short petio- luled; stipels absent; stipules small, caducous. /njlorescence of short axillary racemes or fascicles; pedicels solitary at the nodes, jointed below the calyx, each 2-bracteolate. Flowers yellow; hypanthium campanulate, the 5 lobes unequal, the 2 uppermost lobes relatively broad, the 2 lateral lobes equal to these in length but narrower, the lowest lobe narrow, acute, longer than the others; petals yellow, unequal, the standard the longest, the keel petals ‘he shortest, all petals short clawed; the standard reflexed with 2 basal callosites inside, the wing and the keel petals often auriculate; stamens diadelphous, the cdd stamen free, the anthers uniform; ovary stipitate, the style glabrous, the stigma small, terminal, ovules many. Fruit distinctively inflated, stipitate, oblong, the pericarp splitting into 2 layers, a papery inflated exocarp and a firm endocarp, the legume thus with z bladder on each side; seeds oblong, compressed, at ached near one end. Diphysa is a genus of some 15-18 named speci:s of which perhaps only 1⁄2 will prove to be distinct (Standley & Steyermark, 1946). The species are found from Mexico, where the number of species is greatest, south through Central America to northern South America. Diphysa is reaclily recognized by its papery, reticulately veined, inflated exocarp, which gives the fruit a bladdery appearance. 1. Diphysa robinioides Benth. in Benth. & Oerst., Vidensk. Meddel. Dansk Na- turhist. Foren. Kjøbenhavn 1853: 11. 1854. Type Nicaragua, not seen.—Fic. 23 D. carthagenensis sensu Benth. & Oerst., Vidensk. Meddel. Dinsk Naturhist. Foren. Kjøbenhavn 1853: 13. 1854. not N. ene Jacq., Enum. Syst. Pi. Carib. 28. 1760; Sel. Stirp. Amer. 208. tab. 181. fig. 51. 1763 Medium-sized tree, usually 5—9(—23) m tall, branchlets usually glabrous or sparsely pubescent. Leaves alternate, imparipinnate, to 16 cm long; leaflets 50 Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. [VoL. 67 ANNALS OF THE MISSOURI BOTANICAL GARDEN 676 FIGURE 23. Diphysa robinioides Benth.—A. Habit ( x 1).—B. Fruit (x1). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 677 (5—-)9—15(—21), the lateral leaflets opposite or subopposite, oval or obovate, usu- ally 1.5-3.5 cm long, 0.5-1.0 cm wide, unequal, even on the same specimen, distal leaflets often larger, rounded or retuse at the apex, glabrous and paler beneath, short-petioluled; petioles 1-3 cm long; stipules 2-4 mm long, caducous. Inflorescence of short axillary racemes, 4-7 cm long, 3-6-flowered, appearing with new growth at the end of dry season, pedicels 4-6 mm long. Flowers bright yellow, to 2 cm long; hypanthium funnel shaped, 6-9 mm long, glabrous, the upper teeth partly united, broader than others, the lateral and lowest tooth tri- angular, the lowest tooth slender; standard circular, to 20 mm long, the claw ca. 4 mm long, the wings free, obovate to oblong, auriculate, the keel with basal auricles prominent; stamens diadelphous; ovary with stipe ca. 4 mm long. Fruit a legume with exocarp on either side thin, inflated, conspicuously reticulate veined, (4—)6-11 cm long, 1.5-2.0 cm wide, glabrous; fruiting pedicels 1.5-2.0 cm long, stipe 5-8 mm long; seeds light brown, ca. 6 mm long, ca. 3 mm wide. Diphysa robinioides is found from southern Mexico to Panama. It is appar- ently a common tree in parts of Panama, including the Canal Zone (Standley, 1928). Common names in Panama include ‘‘macano”’ and “‘cacique.”’ It is used as a living fence pos Jacquin (1763) was in error when he cited his plate CLXXX; no figure 51 appears in that plate, but does appear on the next page, plate CLXXXI and the species figured is Diphysa carthagenensis. Most references cite 180, but 181 appears to be correct. Diphysa robinioides is somewhat variable in leaflet size (usually longer than 15 mm, rarely shorter), leaflet texture (usually thin, rarely thick), whether the leaflet apex is rounded or retuse, pubescence (usually none or sparse), and length of legume (usually 6-11 cm long). Collections of flowering specimens have young leaves only. These are, of course, smaller, thus somewhat confusing the situation. Young fruits which may look mature are shorter than fully mature fruits, also complicating the situation. Diphysa robinioides is apparently closely related to D. carthagenensis Jacq. The latter species is distinguished by its smaller leaflets (7-15 mm long), shorter corolla (ca. 10 mm long), and shorter legume (ca. 4 cm long). Diphysa carthagenensis, like D. robinioides, is usually glabrous. Diphysa carthagenensis has been found in Colombia, Venezuela, Guatemala, and Mexico, and is therefore to be looked for in Panama. Diphysa floribunda Rose is also similar to D. robinioides. It has the large leaflets, corolla, and legume of that species but it is densely pubescent on branch- es and peduncles. The leaves are also pubescent, at least when young. Diphysa floribunda has been found in Mexico and Guatemala. It may represent a variety of D. robinioides (Standley & Steyermark, 1946). Duke 9317 from Rio Ailigandi, San Blas, has the short (4 cm) legumes of D. carthagenensis but is leafless. Further collections are necessary for satisfactory determination. Lewis et al. 181, also collected along the Río Ailigandi, is clearly D. robinioides. LM bue ITY: A udis 202 (US). CANAL ZONE: Pacific Saddle Club area, Blum 2055 (FSU SCZ). Road, Correa & Dressler 444 (GH, MO). Gaillard Highway near Pardito: ee ps Erg F, MO, NY, SCZ). Near Paraiso, Cos 13254 (DUKE, F, MO, SCZ). Cerro Galera, 350—400 m, Gentry 6608 (MO). Summit, Johansen s.n. (NY). Gatuncillo, Piper 5/19 (US). Balboa, Standley 27129 (GH, US). Paraiso, Tyson 7401 (MO). CHIRIQUÍ: San Lorenzo, Godfrey 678 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 67334 (FSU, MO). San Félix, 0-120 m, Pittier 5234 (NY, US). Comarca del Barú, Stern & Chambers 134 (US). 3 mi S of El Volcán, 4000 ft, Tyson 886 (FSU, MO). coci£: S rim of El Valle de Antón, 800 m, Allen 2283 (F, GH, MO, NY, US). El Valle de Antón, Folsom & Kauke 2766 (MO). 1 mi from El Valle, Garner 32 (FSU, SCZ). Capellanía, Gonzalez 4 (DUKE, MO). El Valle, White & White 67 (MO, US). Penonomé, Williams 122 (NY, US). DARIEN: Jeniné, Duke 3486 (MO). HERRERA: Chitré, 20 m, Allen 1096 (GH, MO, US). Ocú, 100 m, Allen 4074 (MO). Road from Ocú to Chupampa, nd 224 (GH). Ocú, Stern et al. 1742 (MO, US). 4 mi S of Los Pozos, Tyson 2694 (FSU). Los TOS: 7 mi S of Las Tablas, D'Arcy 4205 (MO). 1-2 mi W of Candelaria, Duke 12432 (MO, NY). qe Tablas, Dwyer 1146 (NY), 2494 (US). Road from Macaracas to Chitré Highway, Tyson et al. 3149 (FSU, MO, SCZ). PANAMÁ: Pacora Road, Correa & Dressler 444 (SCZ, US). Chepo, Gentry & Tyson 1728 F FSU, MO, SCZ). Near Panama City, Hunter (GH, MO). Ancón Hill, Hunter (MO). Chepo, Kluge 28 (US). Cerro Azul, 600 m, Lao & Holdridge 39 (DUKE). Bellevista, MacBride 2751 (F, US). Sabanas, Paul 169 (US). Sabana de Juan Corso, Pittier 4641 (US). rai al Road, Standley 26822 (US). Between Las Sabanas and Matías Hernández, Standley 31826 (GH, US). W of Chepo, Tyson 5371 (FSU, MO, SCZ). Cerro Azul above lake, Tyson & Lazor 6087 x SAN BLAS: Lower Río Ailigandí, Duke 9317 (MO). Ailigandí river bank, Lewis et al. 181 (GH, MO, US). VERAGUAS: E of Santiago, 30 m, Duke 12351 (MO). Between Quebrada El Nance and Santa Fé, 250 m, Nee 8146 (MO). egeo San Francisco and Santa Fé, Stern et al. 1919 (MO, US). El Embalsadero, Tyson 6091 (SCZ). 21. DUSSIA Michael O. Dillon?! Dussia Krug & Urban ex Taubert, Naturl. Pflanzenfam. 3, Abt. 3: 193. 1892. TYPE: Dussia martinicensis Krug & Urban ex Taubert. "d Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 139. 1922. TYPE: Vexillifera micranthera — Dussia discolor (Benth.) Amsh. Cashalia Standley, J. Wash. Acad. Sci. 13: 440. 1923. TYPE: Cashalia cuscatlanica Standley = Dussia uscatlanica *(Standley) Standley & Steyermark. Trees, to 50 m high, to 1 m in diameter, the trunk straight, high crowned, the bases buttressed, the bark smooth, gray, sap blood red; stems terete, striate or slightly angular, puberulent to tomentose, glabrate. Leaves imparipinnate, 5—25- foliolate, the leaflets alternate to subopposite, exstipellate; stipules minute, ca- ducous. Inflorescence axillary, racemose or racemose paniculate. Flowers rarely andromonoecious, pink to purple, 15—25 mm long; calyx campanulate, somewhat oblique, the 5 lobes subequal; standard orbicular reniform, persistent; wing straight, the keel petals similar to wings, dorsally tomentose; stamens 10, sub- equal, the filaments connate at the base, 1 subfree, the anthers dorsifixed, ver- satile, ovate; ovary 1-5-ovulate, pubescent, brevistipitate or subsessile, the style pubescent, incurved, the stigma apical, minute. Fruit narrowly ovoid ellipsoid to obovoid, compressed laterally, 2-valved, dehiscent, orange velutinous, commonly 1-2-seeded; seeds red, ellipsoid or cylindrical, the hilum small, linear, lateral: cotyledons thick and fleshy, radicle short, inflexed. Dussia is an American tropical rain forest element represented by ten species, principally from the Antilles and southern Mexico southward to central Peru and the Amazon basin of Brazil. Each species is essentially endemic, occupying small geographic range and usually quite rare within a community. Since all species are large, buttressed trees, adequate collections are difficult to obtain, and flowering ?' Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 679 or fruiting material is often lacking. Two apparently rare species are found in Panama, each used locally as a lumber source. Literature: Rudd, V. E. 1963. The genus Dussia (Leguminosae). Contr. U.S. Natl. Herb. 32(4): 247—277. w Leaves 11-25- eeri the leaflets predominately oblong to obovate, 7-15 cm long, 2.5- 9.0 cm wide, the lower surface villous with subpatent or sometimes crisp pubescent; bracts larger than ias ug 5- 10 mm long, 2—4 mm wide, the bracteoles 5-7 mm long, 2-3 m wide 1. D. cuscatlanica Leaves 5-9-foliolate, the leaflets predominantely elliptic to ovate or obovate, 9-15 c P^ ad 6-9 cm wide, the lower surface crisp pubescent; bracts smaller than bracteoles, 7-10 m long, 4-7 mm wide, the bracteoles 9-12 mm long, 8-9 mm wide ____-- 2. D. B par A b » — . Dussia cuscatlanica (Standley) Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 341. 1940.—Fic. 24. Cashalia Md Standley, J. Wash. Acad. Sci. 13: 441. 1923. TYPE: = ii gh Finca Colima, Sierra de a, Ahuachapan, Standley 20197 (F, holotype; GH, is pug Kinh Jamel J. Arnold Arbor. 19: 118. 1938. TYPE: votes š e REN Skutch 027 (A, holotype). Tree to ca. 50 m tall; young stems yellowish to reddish pubescent, patent to crispate, glabrate. Leaves 11—25-foliolate; the petioles ca. 9 cm long, the rachises 20-40 cm long; petiolules 4-10 mm long; leaflets 5-11 cm long, 3-6 cm wide, the lower leaflets more oval to ovate, terminal leaflet usually obovate, others usually oblong, the apex acuminate to acute, the base obtuse, truncate or subcordate, the upper surface glabrous, the lower moderately pubescent with patent to slightly crispate trichomes. Inflorescence racemose, yellowish to reddish tomentose; bracts tridentate to rhombic or lanceolate, erose to entire, acuminate, 5-10 mm long, 2-4 mm wide, bracteoles obovate, entire or dentate, acuminate, 5-7 mm long, 2-3 mm wide. Flowers (15—)18—25 mm long; calyx 8-10 mm long, the lobes 4-5 mm long; corolla pale pink with a greenish streak along the standard. Fruit 5-10 cm long, 2.5-3.5 cm wide, dehiscent, the valves curling; seeds 2.0-3.5 cm long, 1.3-1.7 cm in diameter. This species is found infrequently in forests from southern Mexico to Panama. It is distinguished from the following species in having more leaflets, flowers with the bracteoles smaller than the bracts, and usually larger 2-seeded fruits. It is nown by several local names, matabuey" (Mexico), "'cashal" (El Salvador), ‘‘palo de tigre" (Guatemala), and ‘‘pizarro’’ (Panama). CHIRIQUÍ: 8 km N of El Volcán, Little 6048 (MO, US). DARIEN: El Real, Duke 2110 (MO, NY). Río Balsa between Manené and Tusijuanda, Duke 13555 (MO, NY). Summit of Cerro Pirre, Gentry & Clewell 7010 (MO). 2. Dussia macroprophyllata (Donnell Smith) Harms, Repert. Spec. Nov. Regni Veg. 24: 212. 1928. i bale a Donnell Smith, Bot. Gaz. 61: 56. 1913. TYPE: Costa Rica, Las Vueltas, , Cartago, Tonduz 12949 (US, holotype). 680 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 67 «n p P P : Z p WZ 7 / x À S 77 J ý < FiGURE 24. Dussia cuscatlanica (Standl.) Standl. & Steyerm.—A. Habit (x!4). [After Duke 8210.]—B. Fruit (x%). [After Duke 13555.] Cashalia panamensis Standley, Trop. Woods 16: 16. 1928, nomen nudum, Publ. Field Columbian Mus., Bot. Ser. 4: 212. 1929. type: Panama, Cooper 520 (F, holotype). Tree to ca. 40 m tall; young stems yellowish to reddish tomentose. Leaves 7— 9-foliolate, rarely 11-foliolate; leaflets 6-18 cm long, 4-9 cm wide, ovate, elliptic, elliptic oblong, or obovate, the apex obtuse or sometimes acuminate, densely crisp pubescent below, glabrous above. Inflorescence racemose, reddish tomen- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 681 tose, the bracts clawed, rhomboid lanceolate, acuminate, 7-10 mm long, 4-7 mm wide, oblique, erose; bracteoles ovate, clawed, cordate, erose, 9-12 mm long, 8- mm wide, acute. Flowers 17-20 mm long; calyx ca. 10 mm long, the lobes ca. 5 mm long; corolla purplish. Fruit ca. 5 cm long, ca. 2.5 cm wide, 1-seeded. Dussia macroprophyllata is better collected to the north in Costa Rica. It is distinguished by its large leaflets, relatively small fruits, and bracteoles that are larger than the bracts. It is known locally in Panama as “‘citron.”’ CAS DEL TORO: Almirante, Cooper 520 (F). COLON: Ca. 7 km SW of Portobelo along Rio Gian Mori & Kallunki 5203 (MO). 22. ERIOSEMA Muriel E. Poston”? Eriosema (DC.) G. Don, Gen. Hist. 2: 347. 1832. Nomen cons. Rhynchosia sect. Eriosema DC., Prodr. 2: 388. 1825. TYPE: Eriosema rufum (H.B.K.) G. Don. Euriosma Desv., Ann., Sci. Nat. I. 9(1): 421. 1826. Nomen rejic. g d sessiliflora Desv. Euryosma Walp., Rep. 1: 785. 1842. orth. mut. Eroso (DC.) G Perennial herbs or subshrubs; stems angular to terete, striate, pubescent. Leaves trifoliolate or rarely unifoliate; leaflets gland-dotted below; stipules lan- ceolate or oblong, usually persistent. Inflorescence axillary or terminal, race- mose, the flowers sometimes congested at the tips; bracts lanceolate, striate strig- ulose, caducous. Flowers with the calyx campanulate, the teeth 5, gland-dotted; corolla papilionaceous, the standard usually yellow with reddish purple to maroon striations, obovate with auriculate base, the wings narrowly oblong, lobed above the claw, the keel falcate; stamens 10, diadelphous, the vexillary stamen free, the anthers equal; ovary sessile or subsessile, villous, the style curved, slender, the stigma capitate. Fruit oblong, bivalved; seeds 2, ovoid, black, brown or mottled, the striophiole white, the hilum linear or oblong. Eriosema is a pantropical genus of about 500 species with the greatest diversity in Africa. In the Americas 38 species are currently recognized (Grear, 1970). Although Eriosema has often been confused with Rhynchosia the two genera can be separated on the basis of habit, petiolar characters, arrangement of the calyx lobes, and characters of the hilum and the striophiole. Literature: Grear, J. W. 1970. A revision of the American species of Eriosema (Legumi- nosae-Lotoideae). Mem. New York Bot. Gard. 20(3): 1—98. & Leaves trifoliolat b. Leaflets ca. É times long as wide, linear to lanceolate Leaflets ie inflorescence compact, few- flowered; calyx 2-7 mm us TR E. crinitum cc pisei not rund pubescent; inflorescence a spiciform raceme, multirate alyx 3-5 mm lon . E. violaceum š: Department of Biology, Howard University, Washington, D.C. 20059. 682 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 bb. Leaflets less than 5 times long as wide, broadly lanceolate to lanceolate _____________- diffusum aa. Leaves unifoliolate 3. E. pe mi 1. Eriosema crinitum (H.B.K.) G. Don, Gen. Hist. 2: 348. 1832. Glycine crinita H.B.K., Nov. Gen. Sp. j is 421. 1823. TYPE: Venezuela, ‘‘montosis Novae Anda- lusiae, in Cerro del Cocollar," not s Rhynchosia crinita (H.B.K.) DC., “aber sis Fa 389 Eriosema pinetorum Standley, Field Mus. Nat. B. em Ser. 8: 315. 1931. Type: British Honduras, Herb to subshrub; stems erect to ascending from a thick rootstock, puberulent to externally pilose. Leaves trifoliolate, leaflets linear to lanceolate, apically mu- cronate to acute, basally cuneate, the margins ciliate, 4.2-7.2 cm long, 0.7-1.2 cm wide, strigose above, glabrous below, the terminal leaflet exceeding the lateral leaflets; petioles 0.1—0.4 cm long, pilose; stipules lanceolate, connate, strigose externally, 1.0-1.6 cm long, 0.1—0.2 cm wide. Inflorescences not exceeding the leaflets, few-flowered, the peduncle 0.4—1.0 cm long; pedicels 0.4-0.5 cm long; bracts subtending the inflorescence 1.5 cm long, 0.3 cm wide; subtending brac- teoles caducous, 0.5-0.7 cm long, 0.1—0.2 mm wide. Flowers with the calyx 0.4— 0.9 cm long, the lobes 0.2-0.7 cm long, the upper lobe the longest, strigose; corolla yellow, 1.0—1.5 cm long, the standard obovate, 1.0-1.3 cm long, 0.7-0.8 cm wide, auriculate at the base, the claw 1.0 mm long, the wings oblong 1.0-1.1 cm long, 0.3-0.4 cm wide at the largest point, the claw 0.3—0.4 cm long; stamens 0.8-1.1 mm long, the vexillary stamen free; ovary sericeous. Fruit ovate, 1.2— 1.5 cm long, 0.5-0.7 cm wide, externally strigose; seeds ovoid, 2.0—4.0 mm long. 1.0-2.0 mm wide, black brown mottled. Eriosema crinitum appears to have continuous variation throughout its range in Panama and is the most common Eriosema species occurring there. Eriosema crinitum occurs in savannas, campos, cerrados, usually in sandy soils, at 300— 2700 m. Grear (1970) has described three varieties which he considers to have distinct morphological characters well correlated with geographical distribution. Eriosema crinitum var. fusiformis (Rusby) Grear is separated by its smaller stat- ure and its whitish silky pubescence. Eriosema crinitum var. macrophyllum Grear is separated by its smaller fruits and more numerous flowers. Neither of the varieties has been reported from Panama or surrounding regions. CANAL ZONE: Ancón Hill, Piper 5552 (US); Standley 25186 (US). cuiRiQuí: Boquete, Allen 4712 (GH, MO); D'Arcy 6313 (MO); Davidson 760 (F, MO). Sabana de la Tortuga between Boquete and Caldera 300-700 m, Pittier 3301 (US). Llanos del Volcán, 1120-1200 m, Siebert 339 (MO). COCLE: Natá, 50 m, Allen 829 (MO). Cerro Pilón, 2700 ft, Dwyer 1051 (MO). Llanos outside Penonomé, Ebinger 1030 (F, MO, NY). PANAMA: Hills above Campana, 600-800 m, Allen 1312 (MO). Between i M Rio Pacora and Chepo highway, Duke 5897 (MO). Cerro a Duke 6026 (MO); 2400-2700 m, Duke 8679 (MO, OSU, US). Between Río Pacora and Chepo, Dwyer et al. 5091 (MO). Cerro Cam- pana, Ebinger 912 (F, GH, MO). 69 km W of Panamá near Panamerican highway, Folsom 4109 (MO). 7 mi W of Chepo, Gentry & Tyson 1629 (MO, SCZ). amp McDaniel 6820 (FSU); Nee 6846 (MO). Las Sabanas, E 25913 (US). W slope of Cerro Cam- pana, 2500 ft, Tyson et al. 2375 (FSU, NY, SCZ). Near Arraijan, Woodson 1401 (GH, MO). VERA- GUAS: San José, 18 km S of Santa Fé, 400 m, Nee 8173 (MO). e m~ ^ = a ^ ° "ius w © sa, < = 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 683 2. Eriosema diffusum (H.B.K.) G. Don, Gen. Hist. 2: 347. 1832.—FiG. 25. Glycine diffusa H.B.K., Nov. pes i Sp. Pl. 6: 420. 1823. TYPE: Colombia, ‘‘prope fluvium Putes, rov. Popayanensium"' not s Rhynchosia diffusa (H.B.K.) DC. Don 2: 388. 1825. Rhynchosia diffusa var. oblongifolia DC., Prodr. 2: 388. 1825. TYPE: not seen. Eriosema diffusa var. oblongifolia (DC.) G. Don, Gen. Hist. 2: 347. 1832. Eriosema diffusa var. eerie. Seem., Bot. Voy. Herald 110. 1853. TYPE: Panama, Volcano of Chiriqui, Seemann, not s Erect herb or subshrub, the stems ascending from a woody base, terete, densely sericeous. Leaves trifoliolate, the leaflets oblanceolate to obovate, 2.5- 4.7 cm long, 0.8-1.2 cm wide, apically acuminate, basally cuneate, the margins revolute, pubescence strigose above, tomentose silvery beneath, especially ‘gibt the veins, the terminal leaflet longer than the laterals, the petioles 1.0-2.0 m long, densely white sericeous; stipules lanceolate, connate to occasionally split. ting, 0.6-1.0 cm long, 0.3-5.0 cm wide, externally strigose. Inflorescence 1.5-3.8 cm long, not exceeding the leaflets; flowers 5-10 per cluster; peduncles 1.8-3.0 mm long, 1.0-1.5 mm wide; pedicels 0.3-0.5 cm long; bracts subtending the flowers lanceolate, caducous, 2.0—4.0 mm long, 1.0-1.5 mm wide. Flowers with the calyx 3.0—4.0 mm long, the tube 1.0-1.5 mm long, externally pubescent, gland dotted; corolla yellow, 7.0-8.0 mm long, the standard ovate, 8.0-9.0 mm long, 4.0—4.5 mm wide, auriculate at the base, the wings oblong, 0.6—0.7 cm long, the claw 0.2 mm long, the keel falcate, 0.6-0.7 cm long, the stamens 0.7 cm long. Fruit ovate, 0.9-1.1 cm long, 0.5—0.7 cm wide, brown strigose; seeds ovoid, 3.0— 3.3 mm long, 1.5-2.0 mm wide, black or mottled. Eriosema diffusum ranges from Mexico to northern Colombia, occurring on savannas and grassy banks near streams. In Chiriqui, Panama, it occurs on lava elds. Eriosema diffusum is recognized by the silvery pubescence of the under- side of the leaves. Seemann described E. diffusum var. holosericeum based on plants collected in Chiriqui, which have copious pubescence on both the upper and lower leaflet surface. However, this single character difference does not seem to warrant segregation of a variety (Grear, 1970). CHIRIQUÍ: Near Volcán, 1300 m, Burt & Koster 156 (MO). Boquete, 3800 ft, Davidson 728 (GH 1 O of Bambito on lava flow, 1600 m, Lazor & Correa 2710 (FSU, MO, SCZ); Partch 69-28 (MO). Forest near Boquete, 1000- 1300 m, Pittier 2945 (GH, MO, NY). Llanos del Volcán, 1120-1500 m Siebert 340 RO Tyson 5748 (FSU), 5858 (FSU, MO, SCZ). Río Chiriquí Viejo Tv White 85 (GH, M ine Near El Hato del Volcan, Wilbur et ee 17194 (MO), 11934 (F, , NY). PANAMA: Road bet n Panama and Chepo, Dodge 16634 (GH, MO). Pacora, Paul 268 q. Penonomé, 50-1000 ft, Williams 582 (US, NY). VERAGUAS: 18km S of Santa Fé on Santiago road, 400 m, Nee 8173 (MO). 3. Eriosema simplicifolium (H.B.K.) G. Don, Gen. Hist. 2: 348. 1832. Glycine e Virus ay iia Gen. Sp. Pl. 6: 419. 1823. TYPE: Venezuela, "'prope Atures, ad catar: Rise simplicifolia (H. T K i DC., Prodr. 2: 389. Eriosema lanceolatum Benth., Ann. Nat. Hist. 3: 438. Been British Guiana. Schomburgk 651 (US, i ines, Herbs procumbent to decumbent; stems terete, villous to pilous. Leaves un- ifoliolate, the leaflets ovate, 6.0-7.5 cm long, 1.5-3.0 cm wide, apically obtuse, 684 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 67 FIGURE 25. Eriosema diffusum (H.B.K.) G. Don.—A. Habit (x!4).—B. Leaflet underside.—C. Flower (x3).—D. Corolla.—D'. Standard (x2).—D*?. Wing petal (x2).—D?. Keel petal (x2).—E. Stamens (x3).—F. Pistil (x3). [After White 85.]—G. Fruit (x 14%).—H. Fruit opened to show seeds (x 1%).—I. Seed (x3). [After Davidson 728.] basally cuneate, the margins entire, sericeous on both the surfaces, occasionally strigose below; petioles 0.2-0.5 cm long, heavily strigose; stipules lanceolate, 0.4-1.0 cm long, 0.1—0.3 cm wide. Inflorescence few flowered with 2—4 flowers, 3.0 cm long; peduncle 1.5-1.8 cm long; pedicel 0.4-0.5 cm long; bracts lanceolate, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 685 caducous, 0.4—0.8 cm long, 0.2-0.3 cm wide. Flowers with the calyx 0.5-0.6 mm long, the tube 2.0 mm long, strigose; corolla yellow, 0.9-1.0 cm long, the standard obovate, 1.1-1.3 cm long, 0.5-0.6 cm wide, auriculate at the base, the wings oblong, 1.1-1.2 cm long, 0.4—0.5 cm wide, the keel falcate, 1.2-1.3 cm long, 0.4— 0.5 cm wide; stamens 1.0-1.2 cm long. Fruit ovate, 1.5 cm long, 0.7 cm wide, brown strigose; seeds ovate, 4.0 mm long, black. Eriosema simplicifolium ranges from Central America and the Greater Antilles to northern South America. Grear (1970) has segregated E. simplicifolium into two varieties: var. micranthum is restricted to Bolivia, Paraguay, and south- eastern Brazil and is differentiated by its smaller flowers and occasionally erect habit. However, E. simplicifolium appears to have continuous variation through- out its range in Panama and surrounding regions. In Panama E. simplicifolium occurs on llanos, usually in sandy soils. CHIRIQUí: Llanos W of Gualaca, 500 ft, Allen 5050 (GH, MO). San Félix, 0-120 m, Pittier 5257 (NY, US). 4. Eriosema violaceum (Aubl.) G. Don, Gen. Hist. 2: 347. 1832. Cystisus violaceous Aubl., Hist. Pl. Guiane 2: 766. 1775. Type: French Guiana, ''Guianae in pratis Macouria,'' not seen Crotolaria lineata Lam. Encyl. Méth. Bot. 2: 202. 1786. TYPE: not seen. Rhynchosia violacea DC., Prodr. 2: 388. 1825. TYPE: not se Herb or subshrub, few to many branched; branches erect or ascending from a woody base; stems puberulent, villous. Leaves trifoliolate, the leaflets linear lanceolate, 4.7-8.0 cm long, 0.6-1.2 cm wde, apically obtuse, basally cuneate, the margins revolute, the upper and the lower surface strigose, the lower surface gland dotted, terminal leaflet ^ longer than the laterals; petioles 2.0-4.0 mm long, densely villous, appressed, the stipules lanceolate, striate, connate, pilose, per- sistent, 6.0-10.0 mm long, 2.0-3.0 mm wide. Inflorescence 4.7-7.0 cm long, multiflorate with 30-35 flowers, in loose clusters; peduncle 2.2-3.0 cm long; ped- icel 1.8-2.0 mm long; bracts subtending the flowers caducous, ovate, 2.5—4.0 mm long, 1.5 mm wide; bracts subtending the inflorescence lanceolate, 0.6 cm long, 0.2 cm wide, striate, puberulent. Flowers with the calyx 0.3-5 mm long, the lobes 1.5-3.0 mm long, the upper lobe slightly longer, externally glandular, villous; corolla yellow, 6.0-8.0 mm long, the standard obovate, 7.0-8.0 mm long, 4.0-5.0 mm wide, auriculate at the base, the wings oblong, 7.0-7.5 mm long, 2.5-3.0 mm wide, the claw 2.0 mm long, the keel 7.0-7.5 mm long, 2.5-3.0 mm wide, the claw 3.0 mm long; stamens 7.0 mm long. Fruit ovate, 0.7-1.0 cm long, 0.4—0.6 cm wide, densely villous; seeds 3.5—4.5 mm long, oval black. Eriosema violaceum ranges from southern Mexico and the Greater Antilles southward to central and northwestern Brazil. It occurs in savannas, cerrados and palm and pine forests and, particularly in Panama, along streambanks. Eri- osema violaceum is distinguished by its narrow leaflets with revolute margins and its many flowered raceme. L ZONE: Rio Azote, 66-70 m, Dodge et al. 16847 (MO). CocLE: Penonomé, 50-1000 ft, Williams € 95 (NY). PANAMÁ: near Pacora, 0—100 m, Burt & Rattray 18 (MO). Río Puente near junction 686 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 with Río Chagres, Dodge et al. 16846 (GH, MO). Río Pacora and Chepo, Duke 5912 (MO). Orillas del camino ca. 1.5 km del Río Cabra, camino a Chepo, Taylor (MO). 23. ERYTHRINA John D. Dwyer? & W. G. D'Arcyš3 Erythrina L., Sp. Pl. 706. 1753; Gen. Pl., ed. 5. 316. 1754. TYPE: E. herbacea L. Micropteryx Walp., Linnaea 23: 739. 1850. LECTOTYPE: M. poeppigiana Walp. — Erythrina poeppi- giana (Walp.) O. F. Cook. Duchassaingia Walp., Linnaea 23: 741. 1850. LECTOTYPE: D. glauca Walp. — Erythrina fusca Lour. Trees or shrubs, usually armed. Leaves alternate, pinnately 3-foliate, the leaflets large, the terminal leaflet usually larger than the laterals, the lateral leaflets usually inequilateral; stipels small; stipules minute, caducous. Inflorescence ax- illary or terminal; flowers numerous, disposed laterally in pseudoracemes; pe- duncles usually spineless; pedicels thickened distally. Flowers large; calyx in- equilaterally campanulate or tubular, the margin subentire, lobed or vaguely toothed; standard often large, short or long clawed, thick, greatly exceeding the keel, the keel petals often coherent along the lower margin, apically glabrous; stamens 10, monadelphous or diadelphous with 9 stamens alternating long and short, the anthers versatile; ovary stipitate, fusiform, somewhat curved, the style slender, elongate, incurved, glabrous, the stigma capitate. Legumes stipitate, linear oblong, often moniliform, mostly constricted between the seeds, dehiscent, often twisted at maturity; seeds several, ellipsoidal, colored, the hilum lateral. Erythrina includes about 108 species in tropical and subtropical areas of Old and New Worlds. It is distinguished from other Panamanian Papilionatae by its woody habit, red or orange flowers, by the wing petals being much smaller than the keel, and by the elongate stipe below the ovary. The genus has been intensively studied for many years by B. A. Krukoff, New York Botanical Garden, and this treatment relies heavily on his published works and annotations on herbarium sheets. In addition, Mr. Krukoff supplied copious notes and advice which helped form a basis for this treatment. The key to species is modified from Krukoff's key. Krukoff (1939) recognized 10 Species Groups in the New World Erythrina, but later Krukoff and Barnaby (1974) reorganized Erythrina on a worldwide basis, recognizing 5 subgenera and 26 sections. Two subgenera, Micropteryx and Er- ythrina occur in Panama, the first with sects. Micropteryx and Duchassaingia, and the second with sects. Edules, Erythrina, and Gibbosae. Section Cristae- Galli Krukoff (Subgenus Micropteryx) is introduced. Flowers and fruits of all species of Erythrina were well illustrated by Krukoff and Barnaby (1974), and reference to these illustrations can be an aid in deter- mining the plants. Literature: Games, D. E., A. H. Jackson, N. A. Khan & D. S. Millington. 1974. Alkaloids of some African, Asian, Polynesian and Australian species of Erythrina. Lloy- dia 37: 581—588. 58 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 687 Graham, A. & A. S. Tomb. 1974. Palynology of Erythrina (Leguminosae: Pa- pilionoideae): Preliminary survey of the subgenera. Lloydia 37: 465—481. Hargreaves, R. T., R. D. Johnston, D. S. Millington, M. H. Mondal, W. Beavers, L. Becker, C. Young & K. L. Rinehart, Jr. 1974. Alkaloids of American species of Erythrina. Lloydia 37: 569—580. Krukoff, B. A. 1939. The American species of Erythrina. Brittonia 3: 205—337. — —— & R. C. Barnaby. 1974. Conspectus of the genus Erythrina. Lloydia 37: 332-459. Lewis, W. H. 1974. Chromosomes and phylogeny of Erythrina (Fabaceae). Lloydia 37: 460—464. Raven, P. H. 1974. Erythrina (Fabaceae): Achievements and opportunities. Lloydia 37: 321—331. Romeo, J. T. & E. A. Bell. 1974. . of amino acids and certain alka- loids in Erythrina species. Lloydia 68. Toledo, V. M. 1974. Observations on p relationship between hummingbirds and Erythrina species. Lloydia 37: 482-487. a. Seeds dark brown or black; calyx mostly campanulate; introduced and native species. b. Seeds 25-35 mm long, soft, kn z dig 5. E. edulis Seeds 12-19 mm long, hard, dry ooth. Leaflets mostly po ned: pom nis about !⁄2 as long as the keel bb. Jt _ 6. E. E. fusca CG: Leaflets mostly sharp tipped; wing petals minute, less than V5 as E as sp keel. d. Pod papery; stipels forming stalked cups 1-4 mm long . 9. E. poeppigiana dd. Pod leathery or woody; stipels less than 0.5 mm long, not forming stalked 4. E. crista-galli cups aa. IU red; calyx mostly tubular campanulate; native speci Calyx truncate but with a conspicuous bulge on the LU side just below n rim 7. E. gibbosa ee. Calyx 1 or 2 toothed (lobed), lacking a bulge on the side. f Seeds less than 10 mm long; leaflets as broad or broader than long, Lars tipped (sometimes short acumina ay ) . E. berteroana ff. Seeds more than 10 mm long; leaflets longer than broad, sharp poin Ei g. Pod persistently ied | at least between the seeds (lens may be need- ed). h. nee petals united by outer margins; calyx at least twice as long a EE: A D i hh. Kee 1 petals free; calyx less than 1⁄2 as long as wide . EORR: E. lanceolata gg. Pod glabrous at maturity. i. Calyx with 2 subequal lips 2. E. chiriquensis ii. Calyx with 1 lip forming a pronounced beak _______ 10. E. rubrinervia 1. Erythrina berteroana Urban, Symb. Ant. 5: 370. 1908. sYNTYPES: Colombia, Santa Marta, Bertero 2126, 2772 (B, not seen).—FIG. 26 E. neglecta Krukoff & Moldenke, Phytologia 1: 287. 1938. Type: Panama, Williams 372 (NY, not seen). Trees, branchlets smooth, lustrous, armed, the spines occasional, pyramidal, to 0.6 cm long, often obviously reflexed at the apex. Leaves with the leaflets 5— 17 cm long, 5—20 cm wide, obliquely ovate to ovate rotund or the terminal leaflets subrhomboidal, as wide or wider than long, shortly acuminate apically, widely cuneate basally, the margins vaguely irregular, slightly revolute, thick charta- ceous, smaller leaflets often subcoriaceous, glabrous above, minutely puberulent 688 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 E 26. Erythrina berteroana Urban.—A. Habit (x%). [After Dw yer 1237.]—B. Flower en -c yos - —C'. Standard ( x 5).—C?. Wing petal (x 1).—C?. Keel petal (x 1).—D. ignit XV5).—E. Anthers.—E!. Aba xial view (X5).—E*. Lateral view (x5).—E?. Adaxial view (x5).—F Piet E [After Alios 1631.]—G. Fruit ( x 4).—H. Fruit opened to show seeds (x 1).—I. Seed (x 1). [After Dwyer 1237.] beneath, often appearing glaucous, the main veins 5-6, petiolules to 1 cm long, glabrous, rugulose, often a different color than the rachis; petioles 4.5-15 cm long, glabrous; rachises 1.5-5 cm long, glabrous. Inflorescences to 0.5 m long, usually glabrous; pedicels to 10 mm long; bracts ovate, ca. 10 mm long; bracteoles narrowly ovate, acute, ca. 12 mm long. Flowers with calyx narrowly campanu- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 689 late, ca. 16 mm long and 4 mm wide, stiffly carnose, minutely puberulent, the teeth distinguishable as irregular and rounded elevations, standard elliptic, ca. 80 mm long and 20 mm wide, thick petaloid, glabrous, the wings oblong, ca. 10 mm long and 2.5 mm wide, tapering only gradually basally, the claw ca. 2 mm long; keel petals subcircular, ca. 8.5 mm long and 6.5 mm wide; odd stamen free at the base, then united to the middle of the sheath, the other stamens of 2 lengths, the laterally disposed stamens free for ca. 15 mm, the median (carinal position) free for 4-7 mm, the anthers ca. 3 mm long; ovary slender, stipitate for 1.5 mm, linear, ca. 17 mm long and 1.5 mm wide, ferrugineous puberulent, the style ca. 30 mm long, the stigma capitate, ca. 0.3 mm long. Legumes stipitate moniliform, to 15 cm long and 1.5 cm wide, the stipe 2.5 cm long. Erythrina berteroana is distinct in its small black-marked red seeds. This species is well distributed throughout the West Indies and on the conti- nent, ranging from Mexico to Colombia and Venezuela (Zulia). It occurs from low to middle elevations. In Panama, Erythrina berteroana occurs from sea level up to about 1,800 m, with most collections taken from tropical moist forest. It is one of the commonest species in the genus in Panama. BOCAS DEL TORO: Almirante, Sharp 11 Nov. 1962 (SCZ). CANAL ZONE: Miraflores, Blum 542 (FSU, MO, SCZ). Old gun site route C-2 past Pacific Saddle Club, Blum 2056 (FSU, MO, SCZ). are other locality, Blum 2238 (FSU, MO, SCZ). Cocoli Road, Correa & Gonzalez, 513 (MO, PMA, SCZ). N dden Dam, Correa & Stimson 31 (MO, ). Highway near Paraiso, Croat P (MO). Summit Garden, Croat 12313 (F, MO, SCZ). Near Boy Scout Road, Croat 12916 (MO). Gaillard Highway near Paraíso, Croat 13148 (F, MO). Near Summit Naval Radio Station, Croat 14237 (MO). Between Farfan Beach and Vera Cruz, Duke 11737 (MO). Summit Garden, Dwyer 1237 (FSU, MO). T an Beach, Dwyer et al. 4690 (MO). Cocoli Baptist Church, Dwyer 7212 (GH, US). K-2 Road, Dwyer & Hayden 7541 (GH, MO, US). Madden Dam, Ebinger 854 (F, MO, US). Off road from Madden fon to Chilibre, Folsom 3661, 3662, 3663 (all MO). Paraíso, Folsom 3664 (MO). Near Pina, Gentry 8629 (MO). Río Providencia and ridge S of river, 5-170 m, Gentry & Nee 8746 (MO). Back of Curundú, Hee ey 5255 (F). Without other locality, Haynes 501/9305 (A). Farfan Beach and Palo Seco, 5 m, Hunter & Allen 444 (GH, MO). Near Summit Gardens, Kennedy et al. 2207 (MO). 4 mi SE of im on highway to Balboa, Lasseigne 4286 (MO). 1 mi E of Canal Zone boundary, Lazor 2866 (FSU). Madden Dam, 50 ft, Lewis et al. 31 (MO). Cocolí Road, Lewis et al. 776 (GH, MO, US). Sabana of Panamá, 10—50 m, Pittier 2541 (US). . Gatun and Lion Hill, 10-20 m, Pittier "ih (US). Big swamp E of Río Tocumen, Standley 26704 (US). Las Cruces Trail between F on and Corozal, Standley 29037 (US). Balboa, Standley 32125 (US). Madden Dam, Tran- Ape pepe highway. Stern et al. 351 (GH, MO). Venado Beach near Fort Kobbe, Stimson 5247 (SCZ). "e ‘ 3663 (MO, US). Quebrada Guanabanito 1 km W of La Re pen Busey 487 (F). Boquete, 3800 ft, Davidson 735 (A, MO, US). Santa Marta Bugaba, de Caballero 19 (MO). Dolega, Dwyer & Hayden 7758A (MO). Between inspection station and Volcán, Folsom P 3836, 3937 (all MO). 1 mi N of Harias San Miguel, Concepción, Folsom 3957 (MO). Boquete, 3800 ft, King 5320 (US). 1 km E of Remedios, Nee 10107 (MO). S side of Don 1050-1100 m, Nee 10642 (MO). Finca Collins, 5500 r bare et al. 1146 (GH, MO, US). Boquete, 1200-1500 m, Woodson & Schery 751 (GH, MO). : Mountains beyond La Pintada, 400-600 m, Hunter & Allen 582 (GH, MO). El Valle, Folsom z yet 2740 (MO); Kennedy 2114 (MO), MN (F, MO); Nee 9219 (MO). COLON: Quebrada Bonita 2 km NW of Salamanca, 90 m, Nee 69 O). DARIÉN: Slopes of Cerro Pirre, 200-500 m, Gentry P Clewell 7045A (MO). HERRERA: Ocú, Su et al. 990 (MO). Los SANTOS: Between Limón and Punta Mala, D'Arcy & — 4210 (MO). Punta ge Tyson 2719 (MO, SCZ). Road from oo to Chitré Highway, Tyson et al. 3150 (MO, SCZ). PANAMA: Bejuco, Allen 1631 (F, G Río Maje 4-5 mi above falls near Bayano Lake, 100 m, Croat 34673 (MO). Between "A a and Chepo, Dodge et al. 16639 (MO). Río Charco Espiritu on Tocumen Highway, Duke 5694 (MO). Isla 690 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 dley 29783 (US). Juan Díaz, Standley 30489 (US). Between Las Sabanas and Matías Hernández, Standley 31897 (US). SAN BLAS: Mainland opposite Achituppu, Lewis et al. 108 (GH, MO). VERA- i f Santiago near Santa Maria River, Blum & Tyson 593 (MO, SCZ). Between Santa Fé ‘ i from Santiago toward Atalaya, Dwyer et al. 7402 (US, MO). 1-2 mi above Santa F ; Gentry 3050 (MO); Liesner 839 (MO). Just S of Santa Fé, Nee 8023 (MO). 2 mi S of Cañazas, Tyson 3724 (FSU). WITHOUT LOCALITY: Lewis et al. (MO). — x 2. Erythrina chiriquensis Krukoff, Brittonia 3: 322. 1939. TYPE: Panama, White & White 31 (US, holotype, not seen; isotypes, MO). Trees; branchlets smooth, armed, the spines deltoid, flat, to 1 cm long. Leaves with the leaflets ovate deltoid, 6-20 cm long, 4-12 cm wide, the terminal leaflets usually slightly larger, the acumen often to 1.5 cm long, truncate at the base, the margins vaguely revolute, thick chartaceous, glabrous above and below, the main veins 4-8; petioles 10-15 cm long, smooth, glabrous; rachises 3—4.5 cm long; stipules narrowly rectangular. Inflorescences with the peduncles to 27 cm long and 0.6 cm wide, rubescent, sympodium longer in fruit, pedicels 0.3-1.5 mm long. Flowers with the calyx narrowly campanulate, ca. 13.5 mm long and stiffly car- nose to subcoriaceous, rough, glabrous to ferrugineous villosulose; teeth obso- lete; standard subsessile, narrowly oblong, to 60 mm long, 5-15 mm wide, ta- pering gradually toward the apex, concave, subcuneate, basally carnose, glabrous; wing petals oblong to somewhat circular ca. 9.5 mm long, obtuse ba- sally, delicately petaloid, glabrous except for the puberulent margins, the keel petals semicircular, 8.5-10 mm long, 6-7 mm wide, obliquely rounded, truncate basally, irregular undulate on the upper margin, thickened and puberulent on the lower margin, the claw ca. 1.5 mm long; staminal sheath 25-30 mm long, 1.5-3 mm wide, the long anthers ca. 3.3 mm long, glabrous, the short anthers ca. 2.5 mm long, pilose, the odd filament united to the sheath for ca. 8 mm then free for ca. 15 mm; ovary stipitate for ca. 7 mm, vaguely sigmoid and linear fusiform, ca. 13.5 mm long and 1.5 mm wide, the hairs ferrugineous, curled, the style subulate. Legumes stipitate for ca. 3 cm, to 13 cm long, 1.5-2 cm wide, narrowly cuneate apically, the beak to 3 cm long, ca. 0.6 cm wide basally, the seminiferous areas 2-3 per pod, often interrupted by a narrow constriction, 0.5-0.9 cm wide, to 5 cm long, smooth, vaguely reticulate, minutely ferrugineous farinose; seeds ob- long, ca. 1.2 cm long, 0.6 cm wide, dull or bright scarlet. Erythrina chiriquensis is known only from upland Chiriqui, Panama, where it is common. The pods tend to be much shorter than most other species in Panama and they are usually not moniliform or crinkled. i 6 u (MO, GH, SCZ). Cerro Punta, Lao 354 (MO); Mori & Kallunki 5669 (MO); Stern et al. 8I (MO, US); Tyson 872 (FSU, MO), 5823 (FSU, MO, SCZ). Valley of upper Río Chiriquí Viejo, 1300-1900 m, White & White 31 (MO). Finca Lérida to Peña Blanca, 1750-2000 m, Woodson & Schery 284 (GH), 295 (MO). Casita Alta, Volcán de Chiriquí, 1500-2000 m, Woodson et al. 913 (A, MO, US). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 691 3. Erythrina costaricensis M. Micheli, Bull. Herb. Boissier 2: 445. 1894. TYPE: Puntarenas, Costa Rica, Pittier 4804 (BR, not seen). E. panamensis Standley, J. Wash. Acad. Sci. 17: 10. 1927. TvPE: Panama, Pittier 2656 (US, holotype: not seen). E. colombiana Krukoff, Brittonia 3: 325. 1939. TyPE: Colombia, Lawrance 648 (US, not seen). E. costaricensis var. panamensis (Standley) Croat, FI. B.C.I. 471. 1979. Trees, the branchlets often with silver mottling, smooth, occasionally gla- brous, often armed, the spines scattered. Leaves with the leaflets ovate trape- ziform, obliquely ovate to ovate elliptic, 9-22 cm long, 6-20 cm wide, narrowly acuminate at the apex, oblique, rounded or widely cuneate at the base, thin chartaceous to subcoriaceous, reticulate, often glaucous beneath, glabrous to densely puberulent, the hairs appressed and often confined to the costa and sec- ondary veins; petiolules 0.5-1.7 cm long, arachnoid villose to glabrescent; peti- oles 5-22 cm long; rachises 2-20 cm long. Inflorescences with bracts linear lan- ceolate, to 6 mm long, ca. 0.8 mm wide; bracteoles resembling smaller bracts. Flower with the calyx narrowly campanulate or tubular, 15-30 mm long, 8-10 mm wide, chartaceous to subcoriaceous, glabrous or puberulent, bilabiate, the lobes evanescent or to 7 mm long, the teeth scarcely evident; standard obovate elliptic or narrowly oblong, 60-75 mm long, 12.5-16 mm wide, tapering acutely apically, ultimately obtuse, obtuse basally, the claw to 9 mm long, stiff carnose or thin coriaceous, glabrous; wing petals obovate oblong or oblong, 7-9 mm long, obtuse apically and basally, delicately petaloid, the claw indistinct; keel petals circular oblong or inequilaterally ensiform, ca. 8 mm long, 3.5-10 mm wide, briefly broad acuminate apically, oblique basally, petaloid, glabrous, the claw indistinct; staminal sheath 1-2 mm wide medially, the filaments crassate, ca. 0.8 mm wide, the odd filament often briefly united to the sheath, the anthers 3-4 mm long; ovary flat, 10-15 mm long, puberulent, fusiform, sigmatioid, to 60 mm long, ca. 1 mm wide, ferruginous lanose, the stipe filamentous, the style flat basally, cylindrical subulate above, to 20 mm long. Fruit with stipes to 5 cm long, mo- niliform, 15-23 cm long, 1-2 cm wide, the style base to 5 cm long, farinose with minute deciduous hairs, some hairs persisting between the seeds, the seeds scar- let, ca. 1 cm long or longer and 0.7 cm wide. Erythrina costaricensis is a species of the lowlands, occurring commonly in tropical wet and tropical moist forests. The species is known only from Costa Rica, Panama and Colombia. Croat (1979) separated plants from lowland central Panama varietally from the typical Costa Rican plants, saying they have larger leaves, more constricted pods, red instead of orange flowers, and that the Panamanian plants bloom while leafless while those from Costa Rica are reported to bloom while bearing leaves. AS DEL TORO: Potrero, Changuinola Valley, Dunlap 443 (F, US). Río Terebe between Que- brada peas and Quebrada Schlunjik, Kirkbride & Duke 458 (MO, SCZ). 10-15 mi inland from mouth of Changuiola River, Lewis et al. 875 us MO). Water Valley, Wedel 1766 (MO). CANAL ZONE: BARRO COLORADO ISLAND: Aviles 3 (F), 45 (MO); Brown 16 (F); Croat 6286 (MO, SCZ), 6476 (F, MO), 6490 (F, MO, SCZ), 6560, 6625, ied Vis don MO); Frost 212 (F); Harvey 50 (F); Kenoyer 329 (US); Killip 39975 (US), 40036 (MO, US); enheimer 67-1-17-1415 (MO); Salvoza 874 (A); Shattuck 56, 852 (both F, MO); Standley 31352, yi 40843 (all US); Woodworth & Vestal 311 (A, 692 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 F). W end of Gatún Lake and dam, Blum & Tyson 2001 (FSU, MO, SCZ). Gamboa Pipe Line on Tropic Test Center lands 2 km S of main dirt road, Correa & Haines 484 (MO). Pipeline road within 5 mi of Gamboa gate, D'Arcy 9265 (MO). Hill C-6, Ft. Sherman, Duke 4392 (MO). Pipeline road, Folsom 1938 (MO). Río Providencia 3 km SE of Achiote, Gentry & Nee 8657 (MO). Gatún, Goldman 1854 (US). Gatún Station, Hayes 97 (GH). Pipeline Road, Río Agua Salud, Kennedy 1890 (MO). Río Indio de Gatún, Maxon 4808 (US). Río Petitpie from Ft. Sherman-Gatun Locks road, Mori & Kal- lunki 2659 (MO). Río Mendosa above Pipeline Road, Nee 7601 (MO). Río Providencia 7.5 km SW of Gatun Dam, 25-140 m, Nee & Gentry 8667 (MO). Between Gorgona and Gatún, Pittier 2287 (US). Quebrada Bonita, 70-80 m, Steyermark & Allen 1934 (GH, MO). Onebruda Salamanca, 70 m, Stey- between Salud and Boca de Río Indio, arida 63 (MO). Peluca on road to Nombre de Dios, Kennedy 2776 (GH, MO). Río Buenaventura near Portobelo, Kennedy & Gra 2248a (MO). Río Guanche 6 km S of Portobelo, 0-10 m, Nee 7268 (MO). Duc Ancha 4 km E of Buena Vista, Nee 7776 (MO, US). Río Guanche, 50 m, Sullivan 139 (MO). DARIEN: S slope of W peak of Cerro Tacarcuna, 1100— 800 ft, Terry & Terry 1413 (A, F, MO). Can Parcs 782 (US). PANAMÁ: El Jefe, Duke 9398 (MO). Mountains above Tortí Arriba, Folsom et al. 659 4 (MO). Cerro Campana, Gentry 4906 (MO). Without other locality, Hayes (GH). Cerro Campana, Porter et al. 4280 (MO, SCZ). Cerro Campana, collector unknown, 25 Mar. 1969 (MO). 4. Erythrina crista-galli L., Mant. 99. 1767. TvPE: Brasil, not seen. Armed rree, leafy when flowering. Leaves with the leaflets elliptic or ovate, 5-8 cm long, 2-3 cm wide, chartaceous, puberulent but soon glabrous, the costa sometimes armed beneath. Inflorescence of flowers solitary or in 2's or 3's in the leaf axils, often subterminal; pedicels 1.2-6 cm long, pubescent with deciduous hairs; bracts linear lanceolate, 2.4-3 mm long; bracteoles somewhat smaller. Flowers with the calyx papery, broadly campanulate, 10-17 mm long, slightly shorter on the upper side, 10-15 mm wide at the mouth, the apex bilobed, pu- bescent in bud, soon glabrous; standard red, elliptic, sometimes broadly so, ca. 4 cm long, 2.1-2.6 cm wide, the wings variously shaped, sometimes toothed, 7- 12 mm long, the keel petals falcate, 32-53 mm long; stamens 3.6-6.2 cm long, separate for 0.3—0.7 cm; pistil 3.4-6 cm long, the ovary densely pubescent, the stipe sparingly pubescent. Fruit woody, 14-39 cm long, 1.2-1.5 cm wide, slightly constricted between the seeds, the stipe 2-4 mm long, somewhat expanded up- wards, the acumen 0.7-1 cm long, slender but stiff; seeds black with dark mark- ings, 12-19 mm long, 5-8 mm wide, hard, smooth. Erythrina crista-galli is distinct among species of Erythrina growing in Panama in its dark seeds, in its short, wide calyx, and in its wing petals which are much shorter than the keel or standar The species occurs naturally in Brasil and is known in Panama only in culti- vation. It is a favorite cultivated tree in some parts of the world, but only one collection has been made in Panama. The above description is taken mainly from Krukoff (1939). CANAL ZONE: Albrook across from Building 808, Tyson 6261 (MO). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 693 5. Erythrina edulis Triana ex Micheli, J. Bot. 6; 145. 1892. TYPE: Colombia, Triana (K, holotype, not seen). Tree, sometimes armed. Leaves with leaflets ovate to broadly ovate, mostly 8-21 cm long, 5.5-16.5 cm wide, apically acute or acuminate, basally rounded, truncate, obtuse or cuneate, chartaceous or subcoriaceous, glabrous, not waxy, the principal secondary veins 7-12 on each side; costa sometimes armed; petioles puberulent, glabrescent, mostly 11-36 cm long; petiolules 3-11 cm long, 1.5-3.5 mm thick; rachises densely puberulent. Inflorescences with the bracts mostly lanceolate, to 5.2 mm long, 1.5 mm wide, the smaller ones broader; bracteoles resembling the smaller bracts but usually smaller, fruiting pedicels 0.7—1 cm long. Flowers with the calyx thin, papery, broadly campanulate, 8-11 mm long on the lower side, 7-10 mm long on the upper side, 8-10 mm wide at the mouth, the margin irregularly lobed, pubescent; standard broadly elliptic, 2-3.5 cm long, 1- 2.5 cm wide, emarginate or retuse, clawed, the wings variously shaped, 3—6.5 mm geli the keel petals elongate orbicular, often lobed ventrally near the apex, 10-20 mm long; stamens 2-3.5 cm long; ovary 1.8-3 cm long, densely pubescent, stipitate. Legume somewhat woody, thick, mostly 13-25 cm long, ca. 2 cm wide, the acumen ca. 2 cm long, the stipe slender, 3.5—4.5 cm long; seeds 5-6, dark brown, 25-35 mm long, 10-18 mm wide, soft, drying wrinkled. Erythrina edulis is cultivated for its pods which are boiled or fried. It ranges from Panama to Bolivia, occurring mostly at 1,000-2,500 m elevation, but it sometimes occurs nearly at sea level. In Panama it is known only from upland Chiriqui. RIQUí: Finca Collins, Boquete, Blum & Dwyer 2530A (SCZ). Lower road to Bajo Mono, bos et al. 2229 (MO). N slope of Bart E of Bajo Chorro region, 600—6500 ft, Hammel 3017 (MO). Quebrada El Velo near Finca Lérida, 1700 m, Nee 10624 (MO). Bajo Mono ca. 3 km NW of Boquete, 1400 m, Skog et al. 4062 (MO). Boquete above Jaramillo, N slopes of Cerro Palo Alto, 4000 ft, Webster 16663 (MO). 6. Erythrina fusca Loureiro,? Fl. Cochinch. 427. 1790. Based on Gelala aquatica Rumphius, Herb. Amb. 2: 235, tab. 5. 78. 1750 Large trees. Leaves with the leaflets ovate to oblong, 7-14 cm long, 5-11 cm wide, round obtuse apically, obtuse to vaguely cordate basally, thin coriaceous, minutely bullate, usually green, glabrous above, puberulent to minutely farinose below, the principal secondary veins, 6-9, the costa prominulous above, often aculeate; petioles 9-13 cm long, ca. 0.3 cm wide, minutely puberulent to glabrous; rachises 8-25 cm long, pubescent, often aculeate. Inflorescence with bracts and bracteoles ovate or broadly ovate, to 2 mm long; pedicels to 2 cm long and 0.2 cm wide. Flowers with the calyx broadly campanulate, to 17 mm long, asym- metric, longer on the lower side, chartaceous, pubescent, the hairs deciduous, the teeth scarcely evident; standard orange, trapeziform, to 90 mm long and 40 mm wide, widely deltoid apically, obtuse basally, coriaceous, glabrous, the claw For additional synonyms see Krukoff & Barneby (1974). Only this name has been used for ponaienian plants 5 For additional synonyms see Krukoff & Barnaby (1974). Only this name has been used for Panamanian plant 694 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 ca. 20 mm long, the wing petals obliquely semicircular, ca. 25 mm long and 20 mm wide, obliquely rounded apically, the claw scarcely evident, the keel petals connate, ovate falcate, about twice the length of the wing petals, 12-17 mm wide, truncate, oblique basally; staminal sheath ca. 17 mm long, coriaceous, glabrous, the filaments crassate, ca. 1 mm wide, the anthers ca. 4 mm long, the odd stamen ca. 35 mm long; ovary stipitate for ca. 15 mm, linear, ca. 10 mm long, puberulent, the style linear subulate, ca. 15 mm long, erect. Legume with the stipe linear cuneate, ca. 3 mm long, moniliform, ca. 20 cm long and 1.5 cm wide, the beak ca. 2 cm long, falcate, farinose; seeds ca. 1.5 cm long and 0.7 cm wide, opaque, drying brown, smooth, hard, the scar to 0.7 cm long. Erythrina fusca is the most widespread species in the genus, occurring in the American tropics and from Madagascar to Tonga and the Caroline Islands in the Pacific Ocean. It is apparently dispersed by ocean currents, and it is commonly planted for shade. In Panama it occurs in the lowlands, commonly in tropical moist and tropical wet forests. CAS DEL TORO: Farm Six, Almirante, e 471 (MO, US). CANAL ZONE: BARRO COLORADO ISLAND: Croat 8203 (F, MO, SCZ, kd 15159 (MO, SCZ); Foster 2180 (F, GH, US); Kenoyer 395 (US); Woodworth & Vestal 507 (A, F, MO). Miraflores, Blum 2212 (FSU, SCZ). Bohio, Christo- pherson 142 (US). Between Paraiso and country club, Croat 7170 (MO). Near Albrook Air Force Lion Hill, 10-20 m, Pittier 2571 (GH). Ancón, Pittier 2744 (US). Darién Station, Standley 31645 (US). Miraflores Locks, Stern et al. 3 (GH, MO, US). Curundú, Tyson 2632 (FSU, MO, SCZ, US). Between Fort Davis and Fort Gulich, Tison & Lazor 6236 (FSU). Miraflores Lake, White & White 62 (MO, US). cHIRIQUí: Burica Peninsula, Puerto Limones, Croat 22121A (MO). COLON: Juan Mina Plantation, Río Chagres above Gamboa, Allen 4104 (MO). Río Gatuncillo, Santa Rosa, 25 m, Allen oe (MO). 5-7 mi SW of Portobelo towards Maria Chiquita, Wilbur & Weaver 11169 (F, MO, US). NAMA: Arraijan, Allen 1628 (F, GH, MO, US). Tocumen, Cedeno 16 (MO). Without other locality, us & Klawe, 3 July 1960 (US). Panamá Viejo, 5-10 m, Dodge 17518 (GH, MO). Tocumen, Dwyer 31 Dec. 1962 (MO). Between Río Pacora and Chepo, Dixer et al. 5093 (MO, SCZ). San Jose Island, Johnston 955 (GH). Cerro inde les 484 (MO). Panama City, Macbride 2608 (F, US). Nuevo reparto el Carmen a 500 m de la Transistmica, id T Ta Sabanas N of Panama City, Paul 362 (US). Matías Hernández, Pittier 6942 (G H). Big s of Río Tocumen, Standley 26524 (US). Corozal Road near Panamá, Standley 26779 (US). J. s Standley 30503 (US). Between Matías Hernández and Juan Díaz, Standley 31933 (US). 7. Erythrina gibbosa Cufod., Arch. Bot. Sist. 10: 34. 1934. TYPE: Costa Rica, Alajuela, Brenes 828 (NY, fragm.). Small trees or shrubs, usually leafless at anthesis; twigs drying yellow, armed with spines to 0.8 cm long, moderately appressed pilose. Leaves with the terminal leaflets ovate lanceolate or rhombic ovate, 8-30 cm long, 4-15 cm wide, long acuminate apically, cuneate to truncate basally, chartaceous, sparsely pubescent to glabrous; petiolules to 0.8 cm long; petioles 10-15 cm long, to 0.3 cm wide, the spines occasional; rachises 2-3.5 cm long, pubescent; stipules narrowly lan- ceolate, to 2 cm long. Inflorescences ca. 15 cm long, densely villose above; bracts linear lanceolate, to 4 mm long and 1 mm wide. Flowers with the calyx campan- ulate, ca. 15 mm long and 9 mm wide apically, to 3 mm wide basally, conspicu- ously ventricose on the lower surface, chartaceous, puberulent or glabrous, thick- ened apically on the lower side, the teeth scarcely evident; standard subsessile, narrowly oblong, ca. 60 mm long and 12 mm wide, acute apically, cuneate basally, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 695 carnose, glabrous, the wing petals narrowly oblong, ca. 11 mm long and 4 mm wide, obtuse apically, petaloid, the claw scarcely measurable, the keel petals ca. 10 mm long and 5 mm wide medially, obliquely acute apically, obtuse basally, the claw inconspicuous; staminal sheath ca. 2.5 mm wide, the filaments to 20 mm long, the median anthers pubescent, the anthers 2-3 mm long; ovary stipitate for 17 mm, linear falcate, ca. 20 mm long and 1.5 mm wide, the hairs dense, soft, appressed, the style curved, ca. 17 mm long. Fruits subligneous, coiled and twisted when mature, conspicuously and irregularly undulate crenulate on the ventral side; seeds uniformly scarlet, soft, wrinkled when dry. Erythrina gibbosa occurs in Central America from Honduras to Panama. In Panama it has been collected from sea level to about 1,200 m. BOCAS DEL TORO: Chiriqui Lagoon, Wedel 1196 (GH, MO, US). Rio Cricamola between Finca St. Louis & Konkintoe, 10-50 m, Woodson et al. 1930 (A, MO, GH, US). cHIRIQUÍ: Volcán Chiriquí be Río Chiriquí Viejo, 1200 m, Allen 995 (MO). Palo Santo 3 mi N of Volcán, Croat 13580 (FSU, . Upper Río Gariché, 1050-1100 m, Seibert 364 (A, MO). cocrÉ: El Valle, Allen 110 (A, MO), - (MO), 2714 (MO, US), 3622 (GH). Club Campestre, Duke 13232 (MO) EI Valle, Dwyer & Correa 8019 (MO); Dwyer 1839a (US); Folsom & Kauke 2755 (MO); Gentry 6794 (MO); Hunter & Allen 330 (GH, MO, US); Kennedy et al. 2212 (F, MO); Lao 276 (MO). vERAGUAS: Río Tercero Braso above Santa Fé, Croat 27335 (MO). Alto Piedra Santa Fé, Lao 514 (MO). N of Santa Fé, Mori & Kallunki 2537 (MO). 11 km from Escuela Agricola Alto de Piedra, Río Dos Baca, Mori & Kallunki 3125 (MO). WITHOUT LOCALITY: Moore 41 (F) 8. Erythrina lanceolata Standley, Contr. U.S. Natl. Herb. 17: 432. 1974. TYPE: Costa Rica, Wercle (Pittier 3693) (US, not seen). Shrub or small tree, leafy when blooming; often armed. Leaves with the leaflets ovate, apically acute to long acuminate, basally rounded or obtuse, char- taceous, soon glabrescent above, beneath becoming glabrous or remaining spar- ingly puberulent with sparse, weak hairs, and with a minute waxy reticulum visible under a lens, unarmed, the petioles about as long as the terminal leaflet. Inflorescence with the bracts linear lanceolate, 1-1.3 mm long, ca. 0.3 mm wide, the bracteoles slightly smaller; pedicels 0.3-1 cm long, thick and longer in fruit. Flowers with the calyx thin, campanulate, 7.5-17 mm long, slightly shorter on the lower side, 5.5-8 mm wide at the mouth, the apex slightly bilobed, sparsely pubescent with appressed hairs, soon glabrescent; standard scarlet, narrow, ob- lanceolate, 4-8 cm long, 1-2 cm wide, rounded or retuse apically, the wings mostly rounded, not hastate, the keel petals 5-12 mm long, about equalling the wings; stamens 3.6-6.1 cm long, free for 1—2.4 cm; pistil 3.2-5.3 cm long, the stipe and ovary pubescent. Legume linear, terete, compressed between the seeds, 15-32 cm long, the pubescence persistent at least between the seeds, the stipe mostly 4-8 cm long, the acumen 2-8 cm long; seeds scarlet, unmarked, 10-13 mm long. Erythrina lanceolata ranges from Honduras to Panama, occurring mostly at middle elevations. The species is attributed to Panama on the identification of a single collection of the species by Krukoff. As little material was available for study, none in flower, many details of the above description were taken from Krukoff (1939). CHIRIQUÍ: 2.5 km from Questa Piedra along Río Monte road, Folsom 3986 (MO). 696 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 9. Erythrina poeppigiana ( Walp.) O. F. Cook, Bull. U.S. Dept. Agric. Bot. 25: 57. 1901. Micropteryx poeppigiana Walp., Linnaea, 23: 740. 1850. TYPE: Peru, Poeppig (not seen; photos F, GH, Erythrina darienensis Standley, id U.S. Natl. Herb. 18: 108. 1916. TYPE: Panama, Pittier 5578 (US, holotype; isotypes, GH, NY). Large trees, often armed. Leaves with the leaflets rhomboid or deltoid, deltoid ovate to suborbicular, 5-18 cm long, 5.5-15 cm wide, acuminate or obtuse at the apex, the base variable, thinly chartaceous, usually scabrous beneath; stipels forming stalked cups 1-4 mm long; petioles 10-30 cm long; rachises elongate. Inflorescences to 35 cm long; bracts ovate, to 1.5 mm long, ca. 0.8 mm wide, the bracteoles somewhat smaller than the bracts. Flowers with the calyx narrowly cylindrical campanulate, ca. 5 mm long and wide, the teeth absent or present as glandular points, carnose, minutely puberulent; standard scarlet, elliptic, to 50 mm long and 20 mm wide, varying from acute to obtuse apically and basally, the claw scarcely evident, the wing petals obovate or oblong, ca. 12 mm long and 5 mm wide, rounded, tapering obtusely basally, glabrous, the claw indistinct, the keel petals falcate, to 45 mm long and 10 mm wide; staminal tube 30—40 mm long, ca. ] mm wide; pistil to 50 mm long, the stipe glabrous, the ovary linear, finely puberulent, the style filiform sigmatoid. Legume chartaceous, to 25 cm long and 1.5 cm wide, the seeds coffee colored, ca. 12 mm long. Erythrina poeppigiana ranges from Panama to Bolivia. In Panama it has rarely been collected, and all collections are from the lowlands. CANAL ZONE: Summit road, Jones 284 (US). DARIEN: Boca de Pauarando on Samb River, Pittier 5578 (GH, MO-photo, NY, US). 10. Erythrina rubrinervia H.B.K., Nov. Gen. Sp. Pl. 6: 434. 1824. TYPE: Colom- bia, Humboldt & Bonpland 1787 (not seen; photo NY). Shrub or small tree, usually armed, leafy when flowering. Leaves with the terminal leaflet rhombic ovate, mostly 10-20 cm long, 5-12 cm wide, apically acuminate, basally cuneate, unarmed, the secondary veins 8-11 on each side, chartaceous, soon glabrescent, minutely reticulate waxy beneath; petiolules 8-30 cm long, often armed, soon glabrescent; petioles 4.5-10 mm long; 1-2.5 mm thick, soon glabrescent. Inflorescences with the bracts caducous, the bracteoles narrowly elliptic, ca. 1 mm long, ca. 0.3 mm wide, the pedicels 0.3—0.9 cm long, pubescent, becoming 0.9 cm long in fruit. Flowers with the calyx thin, tubular campanulate, 14-30 mm long on the lower side, 10-21 mm long on the upper side, narrowed toward the base, 4-6 mm wide at the top, the margin entire but with the lowest tooth 1.5-2 mm long, glabrescent; standard scarlet, narrowly elliptic oblanceolate, 4.5-7.5 cm long, rounded or subacute, the wing petals api- cally denticulate, mostly slightly smaller than the keel petals, the keel petals abruptly acute or acuminate, 6-12 mm long, 4-5 mm wide; stamens 4.3-6.6 cm long, free for 1-2 cm; ovary and stipe densely pubescent. Legume somewhat woody, 18-36 cm long, ca. 1.5 cm wide, constricted between the seeds, the stipe 3-5 cm long, the acumen 3-6 cm long; seeds numerous, uniformly scarlet, 10—12 mm long, 6-8.5 mm wide. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 697 This species occurs in Belize, Panama, Colombia, Peru and Bolivia. It is usually a species of middle elevations occurring between 1,000-2,500 m. DARIEN: Premontane rain forest E of Tres Bocas, Kirkbride & Duke 1211 (MO). PANAMA: 8 km M SW of Cerro Brewster, 300 m, Lewis et al. 3480 (MO). VERAGUAS: Valley of Rio Dos Bocas 15.6 km NW of Santa Fé, Croat 27763 (MO). 24. FLEMINGIA Muriel E. Poston*® Flemingia Roxb. in Ait., Hort. Kew., ed. 2. 4: 349. Dec. 1812. nomen cons. vs. Flemingia Rottl. (1803, Acanthaceae) et Lourea St. Hil. (1812). TYPE: F. strobilifera (L.) Ait. Shrub. Leaves unifoliate; leaflets oblanceolate, glabrous above, gland-dotted below; stipules lanceolate, caducous, striate; stipels absent. Inflorescences ax- illary racemes; bracts foliaceous, persistent, overtopping the flowers. Flowers with calyx campanulate, the lobes subequal, the upper lobes free; corolla yellow with maroon striation, the standard obovate, auriculate, the wings oblong, the keel falcate; stamens diadelphous, the anthers monomorphic; ovary subsessile, short with 2 ovules, the style slender, the stigma capitate. Fruits oblong, turgid; seeds 1-3, the hilum short. Flemingia is an Old World genus that has been introduced into the New World. 1. Flemingia strobilifera (L.) Ait., Hort. Kew., ed. 2. 4: 350. 1812.—Fia. 27. Hedysarum strobiliferum L., Sp. Pl. 746. 1753. TYPE: India, not seen. Moghania strobilifera (L.) St. Hil. ex Jackson, Index Kew. 2: 252. 1894. Shrub or subshrub; stems terete, strigose. Leaves unifoliate; leaflet ovate to obovate, 9.5—15.0 cm long, 4.0-9.0 cm wide, apically acute, basally truncate to cuneate, glabrous to puberulent above, gland-dotted to villous beneath, the mar- gins revolute; petiole 1.0-2.0 cm long, strigose; stipules lanceolate, striate, 3.0— 4.0 mm long, caducous. Inflorescences exceeding the leaves, racemes or terminal panicles, distichous, 6.0—15.0 cm long, 15-20 flowered; peduncle 1.0-2.0 cm long, strigose; pedicel 2.0—3.0 mm long, strigulose; flowers 1—2 per node; bracts con- spicuous, enclosing the flowers, obovate to cordate, 1.5-2.0 cm long, 0.8-1.2 cm wide, mucronate, basally truncate, strigulose, gland-dotted; bracteoles subtend- ing each flower, ca. 1.0 mm long, strigulose. Flowers with the calyx campanulate, 4.5-5.0 mm long, the tube ca. 1.0 mm long, the upper lobe of calyx somewhat longer than others, corolla 6.0 mm long, the banner orbicular, 5.0-6.0 mm long, 7.5-8.0 mm wide, auriculate, the claw ca. 1.0 mm long, the wing oblong, not auriculate, 5.5—6.0 mm long, 1.0 mm wide, the claw 2.0 mm long, the keel falcate, 8.0 mm long, 2.0 mm wide, the claw 2.0 mm long; stamens 6.0 mm long, vexillary stamen free; ovary sessile to subsessile, the style slender, the stigma capitate. Fruits oblong, turgid, 0.8—1.0 cm long, 0.3—0.5 cm wide, twisting in dehiscence, 56 Department of Biology, Howard University, Washington, D.C. 20059. 698 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 27. Flemingia strobilifera (L.) Ait. —A. Habit (x4).—B. Flower (x4).—C. Glandular hairs.—D. Corolla.—D!. Standard (x3).—D?. Wing petal (x3).—D?. Keel petal (x3).—E. Stamen (x 5).—F. Pistil (x5). [After Wedel 1863.]—G. Fruit (x3).—H. Fruit opened to show seed (x3). [After Allen 2078.] the beak ca. 1.0 mm long, strigose; seeds 2(-3), ovoid, 3.0 mm long, brown black with red mottling, the hilum ovate, 1.0 mm long, estrophiolate. BOCAS DEL TORO: Old Bank Island, Chiriqui Lagoon, Wedel 1863 (MO). CANAL ZONE: Summit, Allen 2079 (F, GH, MO). Transisthmian highway, Atlantic side, Blum & Dwyer 2138 (MO). W end of Gatun Lake dam, Blum & Tyson 1974 (MO, SCZ). Cocoli Road, Burch et al. 1392 (F, GH, MO, NY, SCZ). Contractors Hill, Correa & Gonzales 517 (FSU, GH, MO, NY, SCZ). Between Paraiso and Country Club, Croat 7168 (MO). 1 mi N of Summit Gardens, Croat 9090 (MO, SCZ). Las Cruces 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 699 Trail, Dwyer 1953 (FSU). Albrook, U.S. Army Tropic Test Center site, Dwyer 6590 (MO). Cocoli Road, Dwyer 7227 (GH, MO). Madden Dam, Dwyer 8375 (MO). Albrook, U.S. Army Tropic Test Center site, Tm & Robyns 4 hes K-9 roadside, Gale Sept. 12, 1962 (SCZ). Boyd-Roosevelt highway at Rio Escondido, Gr aham 322 (GH). W bank of Canal opposite Miraflores, Harvey 5226 (F). Without aaa locality, sank 2 (F, MO). 3 mi above Gamboa Bridge, Kennedy et al. 2301 (F, MO, NY). 3 mi N of Miraflores on Balboa highway, Lazor & Blum 5337 (SCZ). V mi N of 7.5 mi SW of Gatün Dam, Nee & Gentry 8675 (MO). Madden Dam, Nee & Mori 3682 (MO). Fort San Lorenzo, Tyson & Blum 3752 (SCZ). 2 mi from Summit Garden SE to Gamboa, Wilbur & Teeri 13381 (DUKE). coLon: Santa Rosa, Río Gatuncillo, Allen 4160 (MO). Salamanca, Gentry 6699 (MO); Nee 9071 (MO). Roadside between Limón and Buena Vista, Wilbur & Weaver 10817 (DUKE, GH, MO). DARIEN: Hydro camp Pico Pendejo on Rio Sabana, Duke 15448 (MO). PANAMA: Nuevo Em- perador, Sandoval 21 (DUKE, MO). 25. GALACTIA W. G. D'Arcy?? Galactia P. Br., Civ. Nat. Hist. Jam. 298. 1756. TYPE: G. pendula Pers. Herbs, trailing or climbing vines or rarely shrubs, perennial. Leaves pinnate (Panama) or digitate trifoliolate or simple; stipels and stipules present. Inflores- cences axillary and terminal, nodose racemes, elongate or short, sometimes sub- sessile; bracts and bracteoles usually present. Flowers mostly pink, purplish or white, the calyx 4-dentate or 4-fid, the upper tooth broader; corolla glabrous or the standard somewhat pubescent; stamens united or the vexilary stamen free, the anthers alike, all fertile, sometimes alternating in height; ovary narrow, mostly pubescent, the style glabrous, curved or nearly straight, the stigma minute. Fruit straight or slightly curved, compressed, sometimes laterally compressed between the seeds or marginally when immature, partially false septate between the seeds, dehiscent by 2 valves; seeds ovoid, dark brown, the hilum small, oval. Galactia includes about 50 species of temperate and tropical regions of the world, the species most numerous in South America and the Antilles. a. Plant erect or ascending, sometimes lax and scandent at the top, not reaching 1 m tall; petiolar rachis less than 5 mm long; leaflets mostly less than 2.5 cm long; pre EIN the wings and keel G. latisiliqua aa. Plant a trailing or climbing vine, mostly exceeding 1 m in length; petiolar Eis more than 5 mm long; leaflets mostly more than 2.5 cm long; standard hardly exceeding or hice than the wings and keel ____ . G. striata — . Galactia latisiliqua Desv., Ann. Sci. Nat. 9: 414. 1826. SYNTYPES: ? Peru, Dombey; Sine loc. ex don. Lavallee (both P, neither seen). Wiry perennial herb or suffrutex, erect or ascending, apically somewhat scan- dent, rooting from a short slender tap root, branching at the ground and only sparingly above; stems densely pubescent, tardily glabrescent. Leaves pinnate, trifoliolate, the leaflets alike, the lateral pair slightly smaller, thick, ovate, mostly (Panama) 1.5-2.5 cm long, apically rounded, mucronate, basally rounded or ob- tuse, the veins 3-5 on each side, prominent beneath, above ascending pilose to glabrous, beneath pilose to appressed pubescent; petiolules 1-1.5 mm long, drying 57 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 700 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 dark, pubescent; rachis to 3 mm long; petioles shorter or longer than the terminal leaflet. Inflorescence lax, few flowered, to 3 cm long, peduncle slender, puber- ulent; pedicels 1-2 mm long, glabrate; bracts ca. 1 mm long, pubescent. Flowers pink, ca. 7 mm long, the calyx pubescent, 4-fid about 24 way down, the standard longer than the wings, erect. Legume flat, oblong, slightly falcate, 3-4 cm long, ca. 5 mm wide, softly pubescent on the sides, the margin straight, the valves slightly compressed between the seeds, tardily dehiscent, ultimately curling; seeds 6-10, brown and mottled, compressed ovoid, 3-4 mm long. This species may be distinguished by its non-climbing habit. Galactia latisi- liqua is a South American species hitherto reaching only as far north as Colombia (Santander) and Guyana. Material from Panama is doubtfully identified with this species as material seen from South America is more pubescent and has some- what larger leaves than the plants from Panama. It is possible that the Panamanian collections represent depauperate individuals of G. striata. They are smaller in most parts than material of G. striata seen from many countries. All Panamanian collections were made in the Pacific lowlands of western Panama during the dry season. CHIRIQUI: Roadside, Veladero, 0-100 m, Burt & Koster 126 (MO). Concepción, Burt & Koster 143 (MO). Gualaca nursery bed, 0-100 m, Koster 174 (MO). VERAGUAS: Trail between Cañazas and Cordillera Central, 300—600 m, Allen 192 (MO). 2. Galactia striata (Jacq.) Urban, Symb. Ant. 2: 320. 1900.— Fic. 28. G. striata Jacq., Hort. Bot. Vindob. 1: 32, tab. 76. M LUE T vov (BM, not seen). G. tenuiflora (Willd.) Wight & Arn., Prod. Fl. Penin. Ind. "a : 761. 184 G. striata var. tenuiflora (Willd.) Burk., Dars iu ua 16: 721. Trailing or prostrate perennial vine, not rooting at the nodes; stems pubescent to tomentose with weak hairs to 1.5 mm long. Leaves pinnate trifoliolate; leaflets about equal or the lateral pair slightly shorter, symmetrical, ovate, mostly 3-6 cm long, 2-3 cm wide, apically obtuse, minutely emarginate and mucronulate, basally rounded, slightly discolorous, thin pilose with loose hairs, more so be- neath, often glabrescent above, the costa and 4 main veins on each side prominent beneath, often glabrescent above, the costa and 4 main veins on each side prom- inent beneath; petiolules ca. 2 mm long, often conspicuously 2-angled; foliar rachis 5-8 mm long; petiole mostly shorter than the leaflets; stipels glabrous, ca. 1 mm long; stipules 1-2 mm long, pilose. Racemes mostly solitary, lax, open, 5- 10 cm long, the peduncle and rachis pubescent; pedicels 1-2 mm long, 1-3 arising at a node, puberulent, the bracts ca. 1 mm long, cucullate, pubescent. Flowers 8-10 mm long, pink, mostly drying white, the calyx puberulent, deeply 4-fid, the lobes narrowly acute; corolla exserted 12-14 way from the calyx, the standard reflexed, glabrate, the wings about as long as the standard, broad; anthers linear, 0.6 mm long, versatile; ovary linear, tomentose, the style glabrous, the stigma minute. Legume flat, oblong, sometimes falcate, mostly 3-5 cm long, ca. 5 mm wide, the margins thick, straight or slightly undulate between the seeds when young, the persistent style forming a small beak, sessile in the calyx, softly pilose on the valves, and on the margins with minute straight whitish hairs, the valves separating and ultimately curling; seeds mostly 5-8, ovoid, oblique, 3.5-5.5 mm 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 701 “ dn cox dresses E FiGuRE 28. Galactia striata (Jacq.) Urban.—A. Habit (x!4).—B. Flower (x215).—C. Corolla.—C!. Standard ( x 2).—C?. Wing petal (x2).—C?. Keel petal (x2).—D. Stamens (x2V2).—E. Pistil (x214). [After Dressler 1861.]—F. Fruit (x'2). [After Gaumer 566.] long, dark brown and mottled, the hilum short, surrounded by the small light colored, usually caducous aril. Galactia striata is common in disturbance in lowland Panama, and it ranges from the United States to Argentina and is naturalized in much of the palaeo- 702 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 tropics. The small but bright pink flowers are usually seen singly on fencerows and in thickets. Several varieties have been distinguished in this species, but their identity and variability are not well understood. Much material from Africa and Central America has been known as Galactia tenuiflora or G. striata var. tenuiflora, but this may not represent a distinct taxon. NAL ZONE: Pipeline road, 1.2 mi NE of Gamboa gate, Croat 12735 (MO, NY). Curundú, D' ec 13428 (MO). Albrook Site, U.S. Army Tropic Test Center, Dwyer 7118 (MO). 0.5 mi W of Summit, McDaniel 12717 (FSU, MO). Río Agua Salud near Frijoles, Piper 5149 (US). Red Tank to eblo Nuevo, Piper 5166 (US), 5167 (MO). Cerro Gordo near Culebra, Standley 26030 (US). Sosa inst area, Tyson 1805 (MO). Howard Air Force Base at Red Devil drop zone, Tyson 1846 (MO). rundu, Tyson 6428 (FSU, MO, SCZ). cuiniQUÍí: Veradero, 0-100 m, Burt & Koster 124 (MO). 1 mi i W of airport at Puerto Armuelles, sea level, Croat 21913 (MO). Quebrada Quanabanao, Distrito Guanano, 0-100 m, Croat 22507 (FSU, MO). DARIÉN: Punta Garachiné, Duke 10484 (MO, NY). HERRERA: 10 mi S of Ocú, Tyson et al. 2811 (MO, SCZ). Los sANTOS: Mo onagre Beach 5 mi S of Chitré, Tyson et al. 3009 (GH, MO). PANAMÁ: Without other locality, Duchassaing 1850 (GH). Beach at Nueva Gorgona, Duke 4559 (MO). Bella Vista, Heriberto 227bis (US); Killip peel (US). San Jose ada nie 161, 1103 (both GH). San Jose Island, Johnston 364 64 (GH , US). Between Sav Río Yguana, Macbride 2650 (F). Bella Vista, Standley 25301 den ' Taboga Island, Sanden 27972 (US). Between Matías Hernández and Juan Díaz, Standley 32009 (US). Playa Co- 26. GLIRICIDIA Peter S. White55 Gliricidia H.B.K., Nov. Gen. Sp. Pl. 6: (ed. quarto) 393 in nota; (ed. folio) 309 in nota. 1824. TYPE: Robinia sepium Jacq. = Gliricidia sepium (Jacq.) Walp. Glyciridia DC., Pr odr. 2: 260. 1825. Gliricidia H.B.K. orth. mut t Hybosema Harms, Feddes Repert. Spec. Nov. Regni Ve eg. 19: 66. 1923. TYPE: H. pM (Schlecht.) Harms = Gliricidia ehrenbergii (Schlecht. ) Rydb. Shrubs or medium sized trees, unarmed. Leaves alternate, sometimes subop- posite or opposite, but never exclusively so, odd pinnate; leaflets entire, often drying with purple mottling beneath; estipellate; stipules small, caducous. /nflo- rescences clustered racemes, axillary or cauline, appearing before or with the leaves; bracts inconspicuous, caducous, ebracteolate, pedicels present. Flowers often rose or rose tinged; hypanthium distinctively cup shaped, the calyx teeth short and broad or absent; petals 5, about the same length, free except the basally connate keel petals, short clawed, the standard nearly orbicular, erect or reflexed; stamens 10, diadelphous, the odd stamen free, the anthers all the same shape; ovary short stipitate, slender, straight, flattened, glabrous, the ovules several to 10 or more, the style glabrous, about !4 the length of the ovary, the stigma capitate. Fruit a dehiscent legume, stipitate, flattened, often wider toward the tip, the margin not or only slightly thickened, not septate and not or only faintly indented between the seeds on the outside, glabrous; the 2 valves hard and often coiling in dehiscence. ** Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 703 Gliricidia is a genus of 6—9 species of Mexico, Central America, the West Indies, and northern South America. North temperate botanists often remark on its similarity to Robinia L. There is also a superficial resemblance to Loncho- carpus of the Dalbergieae. Lonchocarpus can be distinguished by its indehiscent fruits and the wing petals which are often adherent to the keel petals. Gliricidia leaves almost always dry mottled with purple beneath. The name Gliricidia appears to have been taken from Jacquin’s (1763) phrase ‘Glis vel Gliricida’’ for the common name ‘‘Raton vel Mata-Raton'' referring to the use of the plant for mouse poison. In first referring to the plant in Nova Genera et Species, Kunth spelled the name ‘‘Gliricida,’’ but in the footnote where he formally described the genus and in all subsequent usage, including the index to the Nova Genera et Species, he spelled the name ''Gliricidia," so the spelling ‘‘Gliricida’’ must be taken as a typological variant which Kunth did not intend to use. Cowan's (1959) argument (Taxon 8: 59) that ‘‘Gliricida,’’ being more etymologically correct and should be used, is not supported. In Kunth's Synopsis Plantarum (1826), published two years after Nova Genera et Species, only the spelling “Gliricidia” appears. Kunth described the genus in a footnote pertaining to Robinia maculata H.B.K. (=R. sepium Jacq.) and it might be argued that that species should be recognized as the type of the genus. However, Kunth was citing the relationship of Robinia maculata H.B.K. to R. sepium Jacq. and was referring to the latter in his description. Literature: Blohm, H. 1962. Poisonous plants of Venezuela. Stuttgart. Cowan, R. S. 1959. L. of the Western Hemisphere. Notes on choices of lectotypes. Taxon Kunth, C. S. 1826. Voyage aux régions équinoctiales du Nouveau Continent, fait en 1799-1804, partie 6. Botanique, sect. 5. Synopsis plantarum. Paris. Walpers, W. G. 1842. Repertorium botanices systematicae. Leipzig. 1: 679. 1. Gliricidia sepium (Jacq.) Walp., Repert. Bot. Syst. 1: 679. 1842.—FiG. 29. Robinia diei Jacq., Enum. Pl. Carib. 28. 1760; Sel. Stirp. Amer. 1: M 2: tab. 179, fig. 101. 1763. : Colombia, Jacquin, not seen. non Robinia sepium Sw. (178 R. Pire H.B.K., Nov. Gen. Sp. Pl. 6: (ed. quarto) 393, (ed. dies 309. 1824. TYPE: Mexico, Humboldt & Bonpland (P, not seen; microfiche MO). L. maculatus ( .) DC., Prodr. 2: 26 Galedupa pungam Blanco, Fl. Filip. (Ed. 1) 558. 1837. TYPE: edis pepe not seen. Robinia variegata Schlecht., Linnaea 12: 301. 1838. TYPE: Mexico, Schiede, Gliricidia maculata (H.B. K.) Walp., Repert. Bot. Syst. 1: 679. 1842 Millettia luzonensis A. Gray, U.S. Expl. Exped., Bot. 1: 456. 1854. TYPE: Philippines, collector unknown, collection not located. Gliricidia maculata var. multijuga rm in Donell S Bot. Gaz. 28: 284. 1895. TYPE: Guatemala, Hyd Lux, D. Smith distrib. no 3296 (US, isoty Gliricidia lambii Fern., Bot. Gaz. 20: 533. 1895. TYPE: QUO Lamb 451 (NY, holotype; isotype, GH Gliricidia sepium f. maculata (H.B.K.) Urb., Symb. Ant. 2: 1900. Robinia hispida L., Mant. Pl. 101. 1767. pro parte, exclus. eii (LINN 913.2, not seen; microfiche MO). 704 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 7 FIGURE 29. Gliricidia sepium (Jacq.) Walp.—A. Habit (x!4).—B. Flower with one wing petal removed (x 1!4).—C. Fruit (x). Gliricidia sepium (Jacq.) Steud., Nom. Bot., ed. 2, 1: 688. 1840. comb. not valid. G. maculata (H.B.K.) Steud., Nom. Bot., ed. 2, 1: 688. 1840. comb. not valid. G. sepium (Jacq.) Griseb., Abh. Kónigl. Ges. Wiss. Góttingen 7: 52. 1857. Redundant combination. Trees to 10 m tall with spreading crowns. Leaves usually alternate, subop- posite or opposite, to ca. 30 cm long; leaflets 5-19, the lateral leaflets mostly opposite, oval to elliptic, 2-7 cm long, 1-3 cm wide, usually glabrous above, sparsely pubescent and lighter beneath, nearly always purple mottled beneath when dry; rachis and petiole lightly to evenly pubescent, petiolules pubescent; stipules minute, caducous. /nflorescences conspicuous, preceding the leaves; ra- cemes clustered from branches or older wood and hence the inflorescence ap- pearing compound; individual racemes 4-15 cm long, the flowers borne singly, uniformly distributed along the axis; bracts minute, soon deciduous. Flowers pink and lilac to white, the petals usually tinged with yellow toward the base; pedicels 5-10 mm; hypanthium cup shaped, ca. 5 mm long, the calyx teeth low to absent; standard nearly round, erect, ca. 20 mm long, 15-20 mm wide, emar- ginate, the claw slender, to 5 mm long, the wings oblong, ca. 20 mm long, 5 mm wide, the keel petals ca. 18 mm long, ca. 5 mm wide, coherent at the base; 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 705 stamens diadelphous, the tube ca. 12 mm long, 3 mm wide at the base, free filaments alternately long and short; ovary linear, ca. 1 cm long, 1 mm wide, glabrous, the stipe to 5 mm long, the style to ca. 4 mm long, the stigma capitate. Fruit light to dark brown, narrow, oblong, ca. 15 cm long, 2 cm wide, the stipe to 1 cm above the persistent cup shaped calyx, margined, surface dull, glabrous, faintly wrinkled or striate; the valves hard, twisting in dehiscence; seeds 3-10, dark brown, nearly round to somewhat oblong, compressed, to 1 cm long. Gliricidia sepium is native from Mexico to Colombia and in the West Indies. It is also widely cultivated in the New World and was early introduced into the Old World tropics where it has become naturalized, at least in some areas, e.g., the Philippines. It is used as a living fencepost, for hedges, and as a shade tree for cocoa and coffee, although it is less suited to the latter crop since it is leafless for part of the year. The ground bark, leaves, and seeds are used for poisoning rats and mice. The plant is reported to be poisonous to dogs and horses, but cows and goats apparently graze the leaves with no harm (Blohm, 1962). Poultices of the fresh leaves are used to treat skin afflictions. In parts of Central America the flowers are fried and eaten. The species is apparently especially common in the woods and pastures of the Pacific plains in Central America. Common names in Panama are ‘‘mata-raton,’’ and ‘‘madre de cacao." (For additional common names and other notes see Standley, 1928.) In describing the genus Gliricidia, Kunth stated that it included Robinia se- pium Jacq. and R. maculata H.B.K., but Kunth did not make the combinations in Gliricidia. Steudel’s 1840 citation is invalid since he used the names as syn- onyms of Lonchocarpus sepium and L. maculatus. Walpers (1842) made valid combinations under Gliricidia. he specific epithet “sepium” is apparently derived from the Latin ‘‘saepes’”’ (genitive plural ‘‘saepium’’ or ‘‘sepium’’) meaning ''of hedges," which is appro- priate since the tree is widely planted as living fenceposts. Thus the epithet does not change in form with gender and the usage in Index Kewensis, Suppl. 13, as “Robinia sepia" is in error. BOCAS DEL TORO: Almirante, Blum 1349 (FSU, MO, SCZ); Carleton 155 (US). Almirante, Cooper 528 (F). Along runway at Bocas, Lazor et al. 2338 (FSU). Bocas del Toro Island airport, Lewis et US). Water Valley, Wedel 2747 (GH, MO, US). CANAL ZONE: Near mouth of Río Chagres, Allen 871 (F, GH, MO). Near Paraíso, Croat 13953 (MO). Banks of the Chagres River, below Gatün, near sea level, Maxon 4838 (US). Ancón, Piper 5133 (US). Summit Road, Stern & Chambers 157 (MO, NY, Road from Macaracas to Chitré Highway, Tyson et al 51 (MO). Fort Sherman, Tyson US). j Blum 3781 (MO). Summit, roadside, Woodworth & Vestal 477 (F, GH). CHIRIQUI: 1.6 mi W of Puerto Armuelles, 50 m, Croat 21927 (DUKE, left-hand specimen; MO, right-hand specimen). Boquete, 3300-4200 ft, Lewis et al. 393 (MO). 2 km NE of Remedios, 15 m, Nee 10124 (MO, US, WIS). COLON: Puerto Pilon, Correa & Dressler 1191 (DUKE). Penonomé, 50-1000 ft, Williams 341 (NY). DARIEN: El Real along trail to Río Pirre, Stern et al. 289 (GH, MO, US). HERRERA: Paris de Parita, ayano, Porter et al. 5176 (MO, SCZ). T Tabog a Island, Standley 27076, 27857 (both US). SAN REA Duke 10179 (MO, OS). vERA RAGUAS: Santiago, Allen 10 77 (MO). Soná, Allen 4261 (MO, US). Canazas, Tyson 3608, 3626 (both MO). 2 mi S of Canazas, Tyson 3723 (FSU, MO, SCZ). 706 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 27. INDIGOFERA Peter S. White59 Indigofera L.,°° Sp. Pl. 751. 1753; Gen. Pl., (ed. 5) 333. 1754. LECTOTYPE: Indi- gofera tinctoria L. (India). Anil Mill., Gard. Dict. Abr., ed. 4. 1754. TYPE: Anil, sive Indigo Americana, siliquis ... Mill. = Indigofera tinctoria L. Anil(a) Kuntze, Rev. Gen. Pl. 1: 159. 2: 938, 965. 1891. Based on Indigofera L. Herbs or shrubs; more or less strigose with pale appressed hairs attached by their centers, these hairs often lending a gray green appearance to the plant. Leaves imparipinnate or pinnately or digitately 3-foliolate, less commonly simple; usually short-petioled; leaflets entire, venation, except for the midvein, obscure: stipels present, often inconspicuous; stipules setaceous, deciduous, apparently slightly adnate to the petioles. Inflorescence of axillary racemes or spikes, often dense, pedicels short or apparently absent, the flowers single in the axil of a caducous bract, the bracteoles absent. Flowers usually reddish or white, the hypanthium campanulate, the 5 calyx lobes subequal, corolla more or less pu- bescent outside, the standard broad, circular, subsessile, strigose outside, the wing petals oblong, somewhat adherent to the keel, auriculate, the keel petals coherent, laterally spurred; stamens 10, diadelphous, the vexillary stamen free, the sheath slender, the anthers with an apical projection; ovary slender, sessile, few to many ovulate, usually strigose, the style short, bent upward, glabrous, the stigma capitate. Fruits oblong or linear, rarely globose, curved or straight, usually terete or 4-angled, not usually compressed, dehiscent or apparently indehiscent, septate between the seeds; seeds globose to cylindric and truncate, compressed or 4-angled, attached at the middle. Indigofera is a genus of some 800 species in the warmer to tropical regions of both the New and Old Worlds and is particularly diverse in Africa. About 10 species occur in Central America, of which 4 are native in Panama. The Old World species Indigofera tinctoria L., formerly in cultivation in Central America, is perhaps to be looked for in Panama as persistent on old sites of cultivation. It has become naturalized in the West Indies (Rydberg, 1923) but apparently has not spread much from cultivation in Central America (Standley and Steyermark, 1946). Indigofera tinctoria L. resembles I. suffruticosa Mill. (described below) but has pods straight or slightly curved, 6-12 seeded, and 2-4 cm long. The 4 species in Panama, weeds in the dry zones of the country, are easily recognized due to their gray green cast which results from pale, appressed, me- difixed hairs. Additional unusual characters are the appendaged anthers and sep- tate legumes. There is a tendency for Indigofera species to darken on drying. The flowers are ‘‘slightly explosive," the stamens and pistil being under some tension within the keel petals and springing outward when the petals are parted. Indigofera was formerly economically important as the source of the blue dye indigo. Before the cultivation of coffee, this was the most important export crop 5 Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. ** For additional generic synonymy see Hutchinson, 1967. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 707 of Central America. Indigo is no longer extracted from the plant, synthetic dyes having supplanted the natural source. The final decline of the industry was prob- ably in the mid- to late 1800’s. The extraction procedure exposed workers to toxic chemicals (Standley, 1928). Cut plants were fermented in vats of water, after which they were trampled by the workers. Eventually the indigo settled to the bottom of the vats as a precipitate (Standley, 1928; Standley and Steyermark, 1946; see also colorful accounts in Miller, 1768, and Lunan, 1814, and also the references cited in Taylor, 1976). The most important species for indigo manu- facture in Central America were the native Indigofera suffruticosa and the intro- duced /. tinctoria. Indigofera species apparently contain compounds with medicinal properties (Blohm, 1962; Standley and Steyermark, 1946), but medicines prepared from the plants can also have poisonous effects (Blohm, 1962). The seeds of several species are said to be poisonous. In Brazil, the roots of Indigofera lespedezioides H. . and I. suffruticosa were used as fish poisons; the macerated roots were also used as an insecticide (Miller, 1768; Lunan, 1814; Blohm, 1962). Literature: Blohm, H. 1962. Poisonous plants of Venezuela. Stuttgart. Lunan, J. 1814. Hortus jamaicensis. Vol. 1. Jamaica. 538 pp. Miller, P. 1768. Gardener’s Dictionary, edition 8. a. Leaflets 7-19, the fruits either strongly constricted between the seeds or strongly curved. Fruits straight or slightly curved and strongly constricted between the seeds -------- 3. I. panamensis bb. Fruits strongly curved, not wm between the seeds -------------- 4. I. suffruticosa aa. a 1-7, the fruits neither strongly constricted between the seeds nor strongly curve Erect plant, leaves 1— An liolate, the piens ascending, widest duri tip, cun at base, fruits 12-20 m SK i s; cc. Sprawling or suberect ia leaves 5(—7) foliolate, the leaflets not sending widest near the middle, rounded at base, fruits (20-)25-35 mm long ------------ . I. jamaicensis 1. Indigofera jamaicensis Spreng., Syst. Veg., (ed. 16) 3: 277. 1826. TYPE: Ja- maica, Bertero (?TO, not seen).—Fic. 30A- I. flaccida Koenig ex Roxb., Hort. Beng. 98. 1814 (without apr op In W. Carey, ed., Fl. Indica, (ed. 2) 3: 375. 1832. TYPE: India, not seen. (fide Rydber 23. I. mucronata Spreng. ex DC., Prodr. 2: 227. 1825. TYPE: fusion. Bertero (TO, not seen); T the same collection as cited for I. jamaicensis Spreng. pn Indigofera mucronata Lam., Méth. Bot. 3: 247. 1789. Type: South Africa (P-JU, not seen; microfiche “1 ae pan im cytisoides L., Mant. 2: 271, in nota. 1771. Nor aa mucronata Willd. ex Spreng., Syst. Veg., (ed. 16) 3: 276. 1826. in syn. = Indigofera lespedezioides H.B.K., Nov. Gen. Sp. Pl. 6: 457, (ed. quarto). 1824. An Indigofera mucronata = e & Moc., FI. Mex., (ed. 1) 189, 190. 1887, (ed. 2) 173, 174. 1894. TYPE: Mexico, not s I. lespedezioides sensu Hook. & Arn., Bot. Beech. Voy. 415. 1840. Non Indigofera lespedezioides H.B.K., Nov. Gen. Sp. Pl. 6: 457, us quarto). 182 I. si diy Presl., Abh. Bóhm. Ges. Wiss. 3: 485. UM Bot. Bemerk. 55. 1844. TYPE: Mexico, n. (fide Rydberg 1923. ) Not Indigofera heterophylla Thunb., Prodr. Pl. Cap. 133. 1800. TYPE: * South Africa, not seen. Nor Indigofera heterophylla Roxb. ex Wight & Arn., Prodr. 200, ots 201. 1834. T I. subulata sensu Gri l. Bri WLI. 181. 1859, et alia auct. Non Indigofera subulata Vahl. ex Poir., Encycl. Suppl. 3: 150. 1813. TYPE: Guinea (P-JU, not seen; microfiche MO). I. iephrasipidas sensu Micheli, Bull. Soc. Roy. Bot. Belgique 30: 286. 1891. Non Indigofera tephro- 708 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 30. Indigofera.—A-C. Indigofera jamaicensis Spreng.—A. Habit (x34).—B. Flower (x33⁄4).—C. Flower with corolla removed (x6).—D-E. Indigofera suffruticosa Mill.—D. Leaf (x1V2).—F. Fruit (x 11⁄2). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 709 sioides H.B.K., Nov. Gen. Sp. Pl. 6: 455, (ed. quarto). tab. 580. 1824. TYPE: Ecuador (B, not seen; photo MO). I. macilenta Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 26. 1940. TYPE: Mexico, Gentry 2335 (holotype F; isotypes GH, MO, US). Perennial herbs or shrubs, trailing, sprawling, or suberect, the branches ridged in narrow lines from the nodes when young, strigose to glabrate. Leaves 5(—7) foliolate; leaflets broadly elliptic to ovate, when mature ca. 18-25 mm long, 8-12 mm wide, rounded apically, mucronate, rounded basally, strigose, the pet- iolules ca. 1.0-1.2 mm long; petioles ca. 15-18 mm long; rachis 2-3 cm long; stipels apparently absent, perhaps represented by tufts of brown hairs at the rachis nodes; stipules setaceous, ca. 2 mm long. Inflorescence of relatively lax and open axillary racemes 6-12 or more cm long, in fruit (8—)15(-18) cm long, nodes ca. 40, not all bearing mature flowers or fruits; flowering pedicels ca. 1 mm long; fruiting pedicels recurved, ca. 1.5 mm long; bracts narrowly triangular, 1 mm long. Flowers reddish, 5-6 mm long, the hypanthium cup broadly campan- ulate, ca. 1.5 mm long, the 5 subequal teeth triangular tapering acuminately to bristlelike tips, 2 mm long; petals subequal, the standard obovate with cuneate base, ca. 5 mm long, 3-4 mm wide, with scattered hairs outside, the wings 4.5 mm long, the blade oblong elliptic, narrower basal part ca. 2 mm long, auriculate, the keel petals fused, obovate, 5 mm long, narrowed to free cuneate bases ca. 1.5 mm long; staminal sheath ca. 3 mm long, the anthers ca. 0.5 mm long; ovary linear, ca. 3 mm long, the style upcurved, 1 mm long, the stigma capitate. Fruits linear, straight or slightly curved, (20-)25-35 mm long, angled, not narrowed above the remnants of the calyx, strigose, strongly reflexed on recurved pedicels, 12-15-seeded; seeds oblong to subreniform, ca. 2 mm long, 0.8 mm wide, light brown, smooth, dull. Indigofera jamaicensis Spreng. is found from southern Mexico to Colombia and Venezuela and also in the West Indies. It has been introduced and is appar- ently established in parts of the Old World tropics. It is a weedy plant, low or suberect and apparently often leaning on or clambering over other plants. It is common in the drier parts of Central America. It is called ''anilillo" in Mexico and Guatemala (Standley & Steyermark, 1946). The names /ndigofera mucronata Spreng. ex DC. and /. subulata Vahl ex Poir. have both been much in use for this species, while the name I. jamaicensis Spreng. has been ignored. In publishing Indigofera mucronata in 1825, de Can- dolle attributed the name to ‘‘Spreng. in Herb. Balb.” Following the description, he indicated the collector (Bertero) and the type locale (Jamaica). De Candolle apparently was unaware of the earlier J. mucronata Lamarck (1789) for a South African plant not resembling the New World plant in question. In the year after de Candolle's publication, Sprengel seems to have recognized this problem. He cited Indigofera mucronata Lam. as a synonym of I. cytisoides L. and published a new name, /ndigofera jamaicensis Spreng., for the Bertero collection from Jamaica. His description is close to that of I. mucronata Spreng. ex DC., and it is likely that they are based on the same collection. Indigofera subulata Vahl ex Poir. (1813) has also been widely used for this species. It is based on a plant from Guinea described as having 1—3-foliolate leaves. A collection in P-JU (microfiche, MO) bearing the legend ''Indigofera 710 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 subulata Vahl. Guinea. 1804" would seem to be the type. It is predominantly 3-foliolate and is clearly not the New World plant under consideration here. In de Candolle’s text under Indigofera subulata Vahl ex Poir. he dropped Poiret’s reference to unifoliolate leaves and cited ‘‘Indigofera mucronata Spreng.," al- though several pages later he described a separate J. mucronata Spreng. ex DC. Similarly, under the description of J. mucronata Spreng. ex DC., he states ‘* Valde accedit ad J. subulatum sed folia bi nec unijuga.’’ This relating of an Old and New World plant and de Candolle's ignorance of I. mucronata Lam. would seem to be the source of the confusion. BOCAS DEL TORO: Almirante, Barrus 405 (GH). Changuinola to 5 mi S at junction of Río Terebe, 100—200 ft, Lewis et al. 819 (GH, MO). CANAL ZONE: Corozal to Pedro Miguel, Cowell 396 (NY). Road to Radar Station, 1 mi N of Summit Garden, Croat 12651 (MO, NY). Summit Naval Radio K Clayton, Greenman & Greenman 5186 (DUKE, MO). Camino de Corozal, Heriberto 275 (NY, US). Las Cruces Trail, 75 m, Hunter & Allen 741 (MO). Frijoles railroad tracks, Killip 3372 (NY, US). 2 mi N of Paraíso on road to Colon, Lazor & Blum 5322 (FSU). Madden Dam, 50 ft, Lewis et al. 26 (GH, MO, US). La Boca, Mori & Kallunki 3682 (MO). Chiva-Chiva Trail, red tank to Pueblo Nuevo, Piper 5108 (US). Gamboa, Piper 5115 (US). Empire to Mandinga, Piper 5135 (US). Río Grande near Culebra, 50-100 m, Pittier 2113 (NY, US). Gamboa, 20-100 m, Pittier 4786 (NY, US). Donato Trail, Barro Colorado Island, Shattuck 562 (MO). Las Cascadas Plantation, near Summit, Standley 25770 (US). Frijoles, diee 27639 (US). Gamboa, Standley 28356 (US). rini si 31779 (US). l mi N of Summit Garden, Tyson & Blum 1989 (DUKE, Z). of mit on road to FAA radar ae Tyson et al. 2753, 2759 (both MO). Fort San paces oy ae & Blum 3691 (FSU, MO). Albrook Air Force Base, E end of air strip, Tyson & Lazor 6014 (FSU). Curundá, Tyson & Lazor 6112 (FSU). cHIRIQUí: Frontera, 100-200 m, dd & Koster 146 (MO). Rabo de Puerco, 150 m, Busey 448 (MO). Boquete, Davidson 795 (MO). Boquete to 3 mi N, 3300-4200 ft, Lewis et al. 249 MO). Monte Verde, 80 m, Liesner 21 (MO). One. Melliza, 0 0-150 m, Liesner 497 (MO). COCLE: El Valle, 582 m, Burt & Rattray 33 (MO). coLoN: Juan Mina plantation, Río Chagres above Gamboa, 25 m, Allen 4116 (MO). 4 km E of Buena Vista, 3 km N of cement plant, road to Sardinilla, 85 m, Nee 7793 (MO). DARIEN: MacBride 2705 (F, MO, US). HERRERA: 10 mi S of Ocü, Tyson et al. (MO, SCZ). 1⁄2 mi N of Macaracas on Río La Villa, Tyson et al. 3141 (MO, SCZ). Los SANTOS: Chitré-Las Tablas, 0-100 m, Burt & Rattray 51 (MO). Headwaters of Río Pedregal, 25 mi SW of Tonosí, 2500— 100 m, Burt & Rattray 37 (MO). Bayano Dam, D'Arcy 9424 (MO). aaa near beach, Panama City, Duke 4007 (MO). El Llano, Duke 5855 (MO). Rio Pacora just below confluence with Rio Corso, Duke 12020 (MO). Qs Pasiga, Gentry 2379 (MO). 1 mi N of Goofy a sil et al. 2685 (MO). Agricultural Experiment Station at Matías Hernández, Pittier 6895 (GH atías Hernández, Sosa (F). Bella Vista, Standley 25330 (US). Corozal Road near Panamá, Sus 26818 (US). Cerro Azul, 2000 ft, m 2058 (MO, SCZ), 2133 (MO), 2143 (FSU, MO, SCZ). vERAGUAs: Divisa-San- tiago, 0-100 m, Burt & Rattray 72 (MO). Atalaya-Santiago, 100-200 m, Koster 115 (MO). Puerto Mutis, 12 mi S of Santiago, Tyson 6012 (FSU, SCZ). 2. Indigofera lespedezioides H.B.K., Nov. Gen. Sp. Pl. 6: 457 (ed. quarto). 1824. TYPE: Mexico (P, not seen; s 1 MO). I. grisea Desv., Ann. Sci. Nat. (Paris) 9: 410. 1826. TYPE: ‘Hab. in Para.," (?P, ?PC, not seen). I. humboldtiana Spreng., Syst. Veg. (ed. 16) 3: 276. EU TYPE: Mexico (holotype, P, a seen; microfiche, MO; paratype, B, not seen; microfiche, MO). I. mucronata Willd. ex Spreng., in syn., Syst. Veg. (ed. is 276. 1826. Non Indigofera mucronata Lam., Encycl. Méth. Bot. 3: 247. 1789. Nor Indigofera mucronata Spreng. ex DC., Prodr. 2: 227. 1825. An Indigofera mucronata Sesse & Moc., Fl. Mex. (ed. 1) 189, 190. 1887. (ed. 2) 173, 174. I. pascuorum Benth., Ann. Nat. Hist. Ser. 1. 3: 431. 1839. type: British Guiana, Schomburgk 96 (G, een; photo, MO). I. pohliana Benth., Ann. Nat. Hist. Ser. 1. 3: 431. 1839. m Brazil, PoA! (PR, not seen). Anila lespedezioides (H.B.K.) Kuntze, Rev. Gen. Pl. 160. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 711 d acasonicae Brandegee, Univ. Calif. Publ. Bot. 7: 326. 1920. TYPE: Mexico, Purpus 8510 type, UC, not seen; isotypes, GH, MO). Perennial herbs or shrubs, stems not much branched, at least above, stiffly erect, appressed strigose, ridged in narrow lines from the nodes when young, later terete, base stout and woody, the whole plant strigose. Leaves ascending, (1-)3-5(-7) foliolate, typically simple below and compound above; leaflets cu- neate at the base, widest toward the tip, mucronate, the largest simple leaflets to 28 mm long, 8 mm wide, other leaflets averaging 20 mm long, 6 mm wide, the smallest 10 mm long, 1.8 mm wide; petiolules to 1 mm long; petioles 1-3 mm long, rachis 3-15 mm long, stipels setaceous, inconspicuous, less than 1 mm long, stipules narrow triangular, acuminate, ca. 3 mm long. /nflorescence of axillary racemes, as long as or longer than the leaves when in full bloom or fruit, the axis to 4 cm long, the nodes 10-30, not all bearing mature flowers or fruits; pedicels single in the axils of narrow triangular caducous bracts, pubescent; bracts to 3 mm long. Flowers with pink or ?yellow corolla, 4-6 mm long, the hypanthium 2-3 mm long including the triangular teeth, campanulate, pubescent, teeth about equal to the cup, the lower longer than the upper; standard circular, ca. 5 mm long, short clawed, the wing petals oblong, ca. 5 mm long, to 1.5 mm wide, the claw about equal to the blade, the keel petals narrowly oblong, ca. 5 mm long, to 1.5 mm wide, the staminal sheath linear; pistil linear, pubescent, the style 2— 3 mm long, the stigma capitate. Fruits reflexed, linear, 122-20 mm long, 1.7-2.0 mm wide, beaked with persistent style base 1 mm long, 2-valved, the valves only slightly twisted on dehiscence, pubescent outside, septate within, 7-10 seeded, the calyx remnants persistent at the base, nonstipitate, fruiting pedicels 1-2 mm long; seeds oblong or subreniform, 1.5 mm long, dull brown, smooth. Indigofera lespedezioides is found from southern Mexico southward to Bolivia and Brazil and also in the Greater Antilles. The plant is distinctive among Pan- amanian species because of its leaves, the lower ones unifoliolate, and all leaflets ascending, short petiolate, and widest bius the tip, and its short, straight le- gumes which are usually densely pubes The original description of this ipm ET to mention the occurrence of unifoliolate and trifoliolate leaves that are so distinctive and conspicuous on Pan- amanian collections. This is perhaps explained by the fact that the type collection (P; microfiche, MO) appears to be the upper !⁄2 of a stem, the unifoliolate and trifoliolate leaves being more common in this species on the lower parts of the stem. However, there are some flowering and fruiting collections from Panama which have unifoliolate leaves to at least the base of the inflorescence. A fruiting collection in the Willdenow Herbarium (B, not seen; microfiche, MO) is clearly conspecific with the Panamanian material. This is one of two sheets cited by Sprengel in his description of Indigofera humboldtiana Spreng. (=Indigofera lespedezioides H.B.K.), the other being the type of I. lespedezioides itself. This strengthens the case for using the name /. lespedezioides for the Panamanian material. CANAL ZONE: Las Cruces Trail, between Fort Clayton and poa Standley 29187 (US). Las Sabanas, Zetek (MO). cuiRiQUÍ: Boquete, 4000 ft, Davidson 748 (F , MO, US). cocrÉ: E of Río Teta and Interamerican Highway y, Blum & Tyson 1880 (SCZ). Near sh. Clara Beach, Croat 9602 (MO). 4-6 mi E of Nata, 10-25 m, Duke 12391 (MO). Penonomé, Ebinger 1027 (MO). Picacho de 712 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Olá, 350-600 m, Pittier 3067 (NY, US). Sob: Penomomé and Coclé, Stern et al. 980 (MO). Pacora, 0-100 m, Burt & Rattray 20 (MO). Río Pacora, Chepo Highway, Duke 5907 (MO). Río Mar, 5-20 m, Duke 12420 (MO, NY). Las Sabanas, Heriberto 139 (NY); Standley 25929 (MO, US). en Saban a Chepo, Taylor (DUKE, F, MO). P a, 25 m, Woodson et al. 732 (GH, MO, US). Between Pacora and Chepo, ca. 25 m, Woodson et uu 1664 (GH, MO, NY, US). 3. Indigofera panamensis Rydb., N. Amer. Flora 24: 147. 1923. TYPE: Panama, Williams 123 (NY). Shrub to 2.5 m, much branched, the stems ascending, slender, slightly lined from the nodes when young, later terete, whole plant strigose with appressed medifixed hairs. Leaves (7-)11—15(-19) foliolate, leaflets narrowly elliptic to ob- ovate, rounded at the apex, mucronate, rounded or broadly cuneate at the base, lowermost leaflets smaller than the terminal ones, 10 mm long, 3 mm wide, the largest leaflets 24 mm long, 8 mm wide, the middle leaflets typically ca. 15-17 mm long, 5-7 mm wide, strigose, greener and less densely hairy above; petiolule 1.5 mm long, the petiole 5-10 mm long, the rachis 3.0—4.5(—6.0) cm long; stipels setaceous, inconspicuous, 0.5-0.75 mm long; stipules setaceous, inconspicuous, ca. 1 mm long. Inflorescences of dense axillary racemes ca. 4 cm long, to 5 cm long in fruit, thus shorter than the largest mature leaves, the nodes ca. 40-70 but not all with mature flowers or fruits; pedicels single, ca. 1 mm long, recurved in fruit; bracts small, inconspicuous, setaceous, barely 1 mm long, early deciduous. Flowers dull red, ca. 5 mm long, the hypanthium campanulate, ca. 1.25 mm long, the 5 teeth subequal, triangular, ca. 0.5 mm long, strigose; petals subequal, the standard obovate, narrowed to cuneate base, 4.5 mm long, hairy outside, the wing petals with blade narrow elliptic, ca. 3.5 mm long, tapered to the claw, ca. 1.0-1.5 mm long, auriculate at the base, the keel petals fused above, obovate, ca. 4 mm long, narrowed to the free clawed bases; staminal sheath ca. 2 mm long, the anthers ca. 0.5—0.75 mm long; pistil ca. 3.5 mm long, scattered hairy, ovary linear, the style upcurved, ca. 1 mm long; stigma capitate. Fruits straight or slightly curved, 15-20 mm long, strongly constricted between the seeds, ca. 1 mm wide between the seeds, 2 mm wide at the seeds, tapering to a slender beak 1-2(-3) mm long, tapering basally to a narrowed portion 1-2 mm long above the persistent remnants of the calyx cup, (2-)3-4 seeded; seeds oblong to subreni- form, 2-3 mm long, 1.0-1.5 mm wide, smooth, dull, brown. Indigofera panamensis Rydb. is endemic to Panama where it is known from three provinces on the Pacific side of the country. It is found primarily in dry to moist climatic areas in fairly open vegetation such as forest edges, roadside dis- turbed areas, and brushy slopes. This species is readily recognized when in fruit: the fruits are 3—4 seeded, more or less straight, and strongly constricted between the seeds, thus bringing to mind short strings of beads. If the similarity of leaves and flowers is any indication, the plant is closely related to Indigofera suffruticosa Mill. The fruits of the latter species also provide easy identification: they are strongly curved, not constricted between the seeds, and resemble miniature ba- nanas. However, when in leaf or flower, the separation of these two species proves more difficult. /ndigofera suffruticosa Mill. tends to have larger leaflets, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 713 longer stipules, and longer bracts. The problem with leaflet size Is, of course, that leaves enlarge as they mature. Herbarium sheets may thus be inconclusive as to mature leaflet size. The bracts are early deciduous in both Indigofera panamensis and I. suffruticosa; and, in fact, stipules may be difficult to find as well. The two species both have stems lined from the nodes when young; these lines are seem- ingly more persistent on /. suffruticosa stems than on I. panamensis stems with age. The older stems of I. suffruticosa are thus noticeably to obscurely angled in cross section. ANAL ZONE: TTC Albrook Tower out C-15 road, Blum & Dwyer 2160 (FSU, MO, SCZ). Chiva- Chiva Trail red tank to Pueblo Nuevo, Piper 5109 (US). Chiva-Chiva Road, Smith & Smith 3339 (F, MO, NY). Sosa Hill, Balboa, Standley 25266 (GH, US). Las Cruces Trail between Fort Clayton and Corozal, Standley 29227 (GH, US). Curundú, road past Survival School, Tyson 3473 (FSU, MO, SCZ). CocLE: 1.5 mi below El Copé, McDaniel & Cooke 14781 (FSU, MO). Penonomé, 50-1000 ft, Williams 123 (NY). HERRERA: 4 mi S of Los Pozos, Tyson 2696 (FSU, MO, SCZ) 4. Indigofera suffruticosa Mill., Gard. Dict., (ed. 9. Indigofera no. 2. 1768. Based n “Colutea affinis fruticosa argentea . . ` Sloane, Cat. Jam. 142. 1696.— Fic. 30D-E. I. tinctoria Mill., Gard. Dict., (ed. 8). Indigofera no. 1. 1768. Not I. tinctoria L., Sp. Pl. 751. 1753. I. anil L., Mant. 272. 1771. TYPE: "Habitat in India," not seen. I. divaricata Jacq., Hort. Schoenb. pl. 365. 1798. TYPE: not located. I. guatimala Lunan, Hort. Jam. 420. 1814. Based on: " Assurgens minusque divisa .. . ," P. Br., Civ. Nat. Hist. Jam. 302. 1756 Indigofera tinctoria var. brachycarpa DC., Prodr. 2: 224. 1825. pro parte (fide Rydberg 1923). TYPE: plant cultivated in Europe, not seen Indigofera ap var. polyphylla DC., Prodr. 2: 225. 1825. TYPE: un Domingo, not seen. I. comezuelo Moc. & Sesse ex DC., Prodr. 2: 225. 1825. F. rrai Berg., in Berg. & Schmidt, Darst. Offiz. lei 4: 30d. 1863. (pro parte fide Rydberg 1923 Anila tinctoria var. vera Kuntze, Rev. Gen. Pl. 160. 1891. Type: St. Thomas (NY). Perennial herbs or shrubs to 3 m tall, much branched, the branches lined and angled from the nodes when young, later more faintly lined and angled to nearly terete; whole plant strigose with pale medifixed hairs. Leaves (9—)13(—17) folio- late, the leaflets elliptic to obovate, rounded apically, mucronate, acute basally, the terminal leaflets largest, to 40 mm long, 15 mm wide, the smaller mature leaflets to 15 mm long, 5 mm wide, average leaflets ca. 30 mm long, 7 mm wide, strigose above, pubescence denser and leaflets paler beneath; petioles to 2.5 cm long, petiolules 1-2 mm long; rachis typically 6—8(—12) cm long; stipels incon- spicuous, setaceous, to 0.8 mm long; stipules inconspicuous, setaceous, 2-3 mm long. Inflorescence of dense axillary racemes ca. 3-4 cm long, the axis somewhat longer in fruit, to 5 cm long, thus shorter than the leaves, nodes ca. 40-70 but not all with mature flowers or fruits; pedicels single, recurved, just under 1 mm long in flower, just exceeding 1 mm long in fruit; bracts setaceous, early decid- uous, to 2 mm long. Flowers rust red, to ca. 4 mm long, with the hypanthium campanulate, ca. 1.5 mm long, the cup 1 mm long, the subequal deltoid teeth ca. 0.5 mm long, strigose; petals subequal, the standard 3—4 mm long, densely hairy outside, obovate, the wings oblong, the claws to 0.8 mm long, auriculate, the keel petals fused, obovate, densely hairy outside; staminal sheath linear, ca. 2 mm long, the anthers ca. 0.5 mm long; ovary linear, the style upcurved, the 714 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 stigma capitate. Fruits to 15 mm long, ca. 2.2 mm in diameter, strongly curved and resembling miniature bananas, the slender beak 1—2 mm long, not narrowed above persistent remnants of the calyx cup; seeds (3-)5-6(-7), rectangular angled, at least when dry, ca. 1.5 mm long, | mm wide, dull, brown, smooth. Indigofera suffruticosa is found from South Carolina, Florida and Texas southward to Bolivia and Argentina, and also in the Bermudas and the West Indies. It is naturalized in tropical Africa, Asia, and Australia. It is common in the drier parts of Central America. The fruits of I. suffruticosa are strongly curved and resemble miniature bunches of bananas. Indigofera suffruticosa is apparently related to the Old World /. tinctoria, which has also become estab- lished or is at least persistent after cultivation in parts of the New World tropics. Differences with J. panamensis have been discussed under that species. Indigofera suffruticosa is the most important species as a source of indigo dye in the New World tropics (see discussion above). Common names are ‘‘anil’’ (for indigo dye) and “‘jiquelite’’ for the plant (Standley, 1928). Without location, Hayes 298 (NY). BOCAS DEL TORO: Changuinola, ie i 67326 (FSU, MO). Runway, Bocas, Lazor et al. 2344 (FSU, MO, SCZ). Changuinola, 5 mi S at junction of Río Terebe, 100-200 ft, Lewis et al. 790 (GH, MO, US). Chiriquicito, 5 mi S ton Rio Guarumo, Lewis et al. 1967 (GH, US). Almirante, McDaniel 5116 (FSU, MO); Wedel 414 (GH, MO). Chiriqui Lagoon, Wedel 1175 (GH, MO, US). Shepherd Island, Wedel 2719 (GH, MO, US). CANAL ZONE: Farfan Beach, Blum & Godfrey 1679 (DUKE, FSU). Venado Beach, Correa & Gonzalez 501 (GH, MO, SCZ), 505 (DUKE). Farfan Beach, Dwyer 11959 (MO). Chagres, in the old fort, Hayes 355 (NY). Madden Road, 80-110 m, Nee 76/3 (MO). Old Fort Lorenzo, mouth of Rio Chagres, Piper 5970 (US). Balboa, Standley 25568 (US). Corozal, Standley 27395 (US). Gamboa, Standley 28464 (US). Fort Randolph, Standley 28746 (MO, US). Rio Pedro Miguel, near East Paraiso, Standley 30035 (US). Balboa, Standley 32145 (GH, US). Curundú, near Tropic Survival School, Tyson & Blum 2525 (FSU, MO). Pipeline Road, Wilbur & Weaver 11231 (DUKE, MO). Playa Venado, Wilbur & Teeri 12976 (DUKE, F, GH, MO, NY). CHIRIQUÍ: Quebrada Punta de Piedra, 0-100 m, Croat 22462 (MO). 2 mi ; al. 15392 (DUKE). cocLÉ: Río Grande, Burch et al. 1167 (GH, MO, US). Aguadulce, near sea level, edt 4958 (US). Penonomé, 50-1000 ft, Williams 108 (NY, US). HERRERA: | mi N of Chupampa, to Ocu, Wilbur et al. 12110 (DUKE). Los sANTOS: Las Tablas-Pedasi, 0-100 m, Burt & Fas 58 (MO). Las Tablas-Chitré, 0-100 m, Burt & Rattray 63 (MO). Pocrí, beach, Dwyer 1189 (MO). Playa de Monagre, 0-5 m, McDaniel n (DUKE, FSU). PANAMÁ: San Carlos, 0-10 m, Allen 1132 (GH, p US). pia Beach, 6 mi E of San Carlos, Croat 14263 (MO). Chepo Highway W of Pacora, Duke 5934 (MO). Río Ailigandí, ae 10853 (MO). Bayano Cuna, Duke 14502 (MO). Chagres, Fendler 101 um Bella Vista, Heriberto 31 (US). Vera Cruz beach, Lewis et al. 3005 (DUKE). Bald Hill Road, San Jose Island, tinh 1060 (GH, MO, US). 2 km W of El Llano, Nee 7968 (MO). Bella Vista, Standley 25395 (US). Corozal Road, near Panama, Standley 26797 (US). Rio Mar, Tyson et al. 2316 (FSU, SCZ). VERAGUAS: Río Santa María along road to Santiago, 5 km S o Santa Fé, 250-300 m, Nee 8138 (MO). 28. LABLAB W. G. D’Arcy®! & Muriel E. Poston® Lablab Adans., Fam. Pl. 2: 325, 550. 1763. Dolichos lablab L. = Lablab pur- pureus (L.) Sweet Dolic i sensu auct., non sensu Verdcourt role oo Code of Botanical Nomenclature 1978, . 352. See Westphal, Taxon 24: 189-192. *! Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. € Department of Biology, Howard University, Washington, D.C. 20059 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 715 Herbs usually twining, stems terete, strigulose. Leaves trifoliolate, leaflets rhomboid to ovate, apically acute, basally cuneate, stipellate, the stipules small. Inflorescences axillary racemes, the rachis elongate, the flowers 1—5 arising from tubercles on the rachis, the bracts caducous. Flowers with campanulate calyx, the lobes 5, the upper teeth united to apices; standard orbicular, auriculate, 2 callosites or elongated appendages on the inner face, the wing obovate, the keel falcate; stamens diadelphous, vexillary stamen free, the anthers monomorphic; style curved, flattened, the stigma capitate. Fruits broadly oblong to falcate, laterally compressed, beaked, seeds ovoid, black, hilum oblong with a white aril. Lablab is endemic to the Old World with cultivated species found in the New World 1. Lablab purpureus (L.) Sweet, Hort. Brit. 481. 1826. [1827].—FiG. 31. Dolichos purpureus L., Sp. Pl., ed. 2. 1021. 1763. TYPE: not determined. Mii eed L., Sp. Pl. 725. 1753. LECTOTYPE (design. Verdcourt): Phaseolus niger lablab Alpini, De plantis Aegypti liber, . . . . 74, tab. 75. 1640. Climbing Aerbs; stems terete, strigose. Leaves trifoliolate, leaflets ovate to broadly rhomboid, 8.0-13.0 cm long, 6.5-12.5 cm wide, the lateral leaflets asym- metric, apically acute, basally cuneate to truncate; petioles 8.0-12.0 cm long, strigose; petiolules 2.5-3.0 cm long, strigose, stipels linear, 4.0 mm long, the stipules lanceolate, 4.0-5.0 mm long, caducous. Inflorescences axillary racemes to 30.0 cm long; flowers 10—15 per rachis, fasicled in groups of 2-3 at tubercles on the rachis; peduncles 18.0—15.0 cm long, the pedicels short 0.3 cm long; bracts subtending the flowers ovate, 3.0 mm long, 2.0 mm wide, strigose, caducous. Flowers with the calyx campanulate, 6.0-8.0 mm long, the lobes 2.5-3.0 mm long, upper 2 lobes fused; corolla white to purple, standard obovate, auriculate with 2 callosites running along the inner face, 1.5 cm long, 1.2 cm wide, the wings oblong to obovate, 1.5 cm long, 1.0 cm wide, the keel falcate, 1.5 cm long, 0.4 cm wide; stamens 1.5 cm long; style 1.3-1.5 cm long curved, flattened, the stigma capitate. Fruits broadly oblong, 5.2-8.0 cm long, 1.5-2.5 cm wide, strigose, the beak falcate, 1.0-1.5 cm long; seeds ovoid, 3-5, 1.3 cm long, 0.8 cm wide, the hilum oblong with white aril. CANAL ZONE: Summit, Johansen May 1924 (US). Balboa, Sosa Hill, Standley 25284, 32151 (both ). 29. LENNEA Peter S. White'? Lennea Klotzsch in Link, Klotzsch and Otto, Ic. Pl. Hort. Berol. 2: 65. tab. 26. 1842. TYPE: Lennea robinioides Klotzsch (Mexico) = Lennea melanocarpa (Schlecht.) Vatke. Trees or shrubs. Leaves alternate, imparipinnate; rachis grooved; stipels nar- rowly triangular or setaceous, persistent; stipules deciduous. Inflorescences of 5! Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. 716 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 t pn EE opa n Load - 2 i he AAA AU lab purpureus (L.) Sweet—A. Sade (X7/10). [After Schultes & Smith 308, . k Pen n [After de la Cruz 4325, Guyana.] 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 717 axillary racemes, frequently clustered at the nodes, generally shorter than the leaves, appearing with or before the leaves; bracts triangular to subulate, decid- uous by fruiting stage; bracteoles absent. Flowers with hypanthium campanulate, the 5 teeth triangular, often acuminate, mostly as long as or shorter than the cup; petals equal, short-clawed, free except the keel petals united along lower margin at the apex, the standard obcordate, the wing and keel petals oblong, curved; stamens 10, monadelphous, the vexillary stamen fused to staminal sheath for a short distance towards the base, free above and below this; ovary sessile or short stipitate, slender, several-ovulate, the style coiled, the stigma terminal, small. Fruits linear, flattened, few to several seeded, 2-valved; seeds compressed, ob- long to nearly circular. Lennea Klotzsch is a genus of five species found in northern Central America, Mexico, Honduras, and in Panama. It is to be expected in Nicaragua and Costa Rica. It is apparently unknown from South America, although several collections have been made in Darien Province in eastern Panama. The genus was described from plants grown horticulturally in Germany from seed collected in Mexico. Like other relatives described here it resembles the temperate genus Robinia. The coiling of the style is distinctive among related genera in Panama. 1. Lennea viridiflora Seem., Bot. Voy. Herald 107 (corrected page). 1857. TYPE: Santiago de Veraguas, Panama, Seemann 1189 (K, not seen; photo F, MO).— Fic. 32. L. rubioides Seem., Bot. Voy. Herald 107 (uncorrected page). 1853. Nomen nudum. Small trees; branchlets hairy with stiff hairs to glabrous, dull, tan to grayish. Leaves with 9-13 elliptic to ovate leaflets, the lateral leaflets mostly opposite or subopposite, obtuse or often retuse, 2-8 cm long, 1—4 cm wide, the first pair often the smallest on the leaf, thin in texture, dull above, sometimes slightly paler beneath, when mature with appressed, fine, stiff, non-overlapping hairs beneath; stipules narrow, to 5 mm long, apparently deciduous, stipels obvious, setaceous, to 2 mm long, persistent, stiff, the petiolules 2-3 mm long. Inflorescence with raceme axis 2—5 cm long, the flowers close, subtended by ovate-triangular bracts, pedicels 2-3 mm long, the axis, bracts and the pedicels pubescent with stiff dark red brown or yellow brown tinged hairs. Flowers ca. 7-9 mm long with the petals extended, the hypanthium tube campanulate, ca. 3 mm long, the 5 teeth narrow triangular, acute, extending ca. 1 mm beyond the rim of the cup, the lowermost tooth longest, ca. 1.2-1.8(-2) mm long, the 2 uppermost united for about ' of their length, the whole calyx pubescent with stiff hairs as on the inflorescence; petals of subequal length, the standard reflexed, circular, ca. 6 mm long, 6 mm wide, glabrous, the claw ca. 1 mm long, wings oblong, ca. 5 mm long, glabrous, the claw ca. 2 mm long, the keel petals fused for part of the length, ca. 7 mm long with scattered hairs or glabrous; stamens 10, the vexillary stamen leaving the sheath near the base, the 9 other stamens leaving the sheath in a regular series, the outermost with longer free filaments, the innermost with shorter fila- ments as seen when the sheath is opened at the position of the vexillary stamen; 718 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 32. Lennea viridiflora Seem.—A. Habit (x<3/s).—B. Flower (x 33/s).—C. Corolla.—C!. Standard (x 33/s).—C?. Wing petal (x 33/s).—C*. Keel petal (x 33/s).—D. Stamens (x3?/s).—E. Pistil (x 33/5). [After Pittier 5101.) ovary compressed, linear, ca. 5 mm long, appressed hairy, the style coiled, some- times forming an almost complete turn, the stigma terminal, capitate. Fruit com- pressed, narrow oblong, tapered abruptly to acute tip formed by the persistent style base, with woody valves, compressed strongly between the seeds, the outer surface dark brown, strongly reticulate wrinkled, at least on drying, usually gla- brous but sometimes with remote hairs on the surface and with scattered hairs on the edges; seeds 3-6, compressed, roundish to oblong, dark brown, smooth, ca. 8 mm long, 7 mm wide. Lennea viridiflora Seem. is apparently endemic to Panama where it was first collected by Seemann in Santiago de Veraguas. In the original printing of the Botany of the Voyage of the Herald, Seemann lists this plant as ‘‘Lennea ru- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 719 bioides” and attributes this name to Link, Klotzsch, and Otto. This is an apparent corruption of Lennea robinioides Klotzsch (=L. melanocarpa [Schlecht.] atke). Seemann soon came to recognize that his collection was distinct from Lennea melanocarpa, and in a correction page printed in 1857 described it as a new species, Lennea viridiflora Seem. Lennea viridiflora is apparently closely related to Lennea salvadorensis Standl., a plant known only from the type locale in El Salvador. Lennea salva- dorensis bears an overall resemblance to L. viridiflora, but the calyx and flowers are smaller, and the fruits are lighter brown in color. Two other species of Lennea also resemble Lennea viridiflora. Lennea brunnescens Standl. of Mexico can be distinguished by the dense pilosity of the legume surface, glabrous or with remote hairs in Lennea viridiflora as treated here. Lennea melanocarpa (Schlecht.) Vatke bears a close similarity in leaf to L. viridiflora, but the inflorescence axis is glabrous and more slender. It should be noted that the inflorescence of L. viridiflora becomes glabrous in fruit. The calyx of Lennea melanocarpa is also glabrous or only puberulent near the margin. In L. viridiflora the calyx is closely and densely pubescent with straight, stiff hairs. The fifth and final species of Lennea, L. modesta Standl. & Steyermark, is found in Guatemala. Its abruptly tapered leaflets are distinctive. Two Panamanian collections possess somewhat unusual features: Pittier 5101 has calyx lobes longer than those on other flowering collections. In this flowering collection, the longest calyx lobe averages 2 mm in length (1.8 mm on younger flowers), whereas for the other collections the longest calyx lobe ranges from 1.2-1.5 mm (maximum 1.8 mm) long when the flower is fully expanded. The Pittier collection is the only one from Cocle Province. The absence of other differences argues that these collections do not represent a species distinct from Lennea viridiflora, as the calyx lobe lengths are somewhat variable depending on flower age. The other unusual collection is Duke 4873. This fruiting collection has scattered hairs along the legume margin and remote hairs on the surface of the legume. CANAL ZONE: Madden m Iu hie 874 (F, MO). Gaillard highway, N of Paraiso, Lewis et al. 5367 (DUKE, MO, NY, SCZ). Boy Scout Road, Madden Dam Area, Porter et al. 4025 (DUKE, GH, MO, NY, SCZ). Farfan Beach, Tyson et al. 3164 (MO). Miraflores Locks, near Fort Clayton, Tyson 5340 (DUKE, FSU, MO, SCZ). N of Miraflores road at 2m Tyson & Gentry 6409 (GH, Y, SCZ). Road to Miraflores Locks, Tyson & Lazor 6064 (FSU). CocLE: Between Paso del Arado Ola, 20-280 m, ded 5101 (US). DARIEN: Hill near Río eect ca. 4 mi below Yaviza, Duke 4873 (GH, MO, NY, SCZ). Near Refugio, 15-21 mi N of Santa Fé, 30 m, Duke 10311 (MO). Rio Sabana above Santa Fé, Duke 14083 (DUKE, F, GH, NY, MO). Patino, Pittier 6609 (GH, — labeled “B”; NY-photo, left-hand specimen; US, left-hand A sasa Rio Congo, Pittier 6 (US). VERAGUAS: "Santiago, Seemann 1189 (not seen, photo-F, NY). EZ 5 30. LUPINUS David B. Dunn Lupinus L., Sp. Pl. 721. 1753; Gen. Pl., ed. 5. 322. 1754. TYPE: L. albus L Mostly erect or ascending herbs, rarely shrubs; roots sometimes stout, some- times with large nodules; stems mostly pubescent. Leaves 5- or more digitate, ^ Professor and Curator of the Herbarium, Division of Biological Sciences, 105 Tucker Hall, cn of Missouri, Columbia, Missouri 65201. 720 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 the leaflets narrow, pubescent, sometimes sericeous, sometimes glaucous, with a single midnerve; petiolules inconspicuous; petioles elongate; stipules mostly narrow, adnate to the petiole. Inflorescences often showy, blue, purple, red or yellow, racemose, the rachis terminal, several-many flowered; pedicels slender. Flowers with the calyx 2-lipped nearly to the base, the upper lip about as long as the lower; corolla glabrous or sparingly pubescent, the standard broad, the wings apically connate, the keel often beaked; stamens monadelphous, the fila- ments unequal, glabrous, the anthers dimorphic; ovary sessile, the style curved upwards, the stigma capitate, often bearded. Legume oblong, somewhat com- pressed and obliquely constricted between the seeds, dehiscing explosively with a spiral tension, 2—12 seeded; seeds plump with a small, sunken hilum. The genus includes about 200 species in western North America but with species throughout the western hemisphere and in the Mediterranean region. Perhaps as many as 200 species occur in South America. Many species grow in full sun and sandy soil. Some species are used for fodder, while others are well known for poisoning cattle. The toxicity varies with species as to part of the plant implicated, type of toxic response, etc. Nothing is yet known about the toxicity of the Panamanian species. In some countries, the seeds have been used as a coffee substitute following leaching of alkaloids from the seeds. "Lupin'"' (En- glish), "Altramuz," "Lupino" (Spanish). a. Plants erect, to pa 100 cm tall, branching above; leaflets 5-7, the largest 3-4 cm long, 8- 11 mm wide, complanate; standard 10-13 mm long, the keels glabrous or occasionally sparsely ciliate. e distally; stipules to 5 mm long |... L. clarkei aa. Plants dec ween in age 15-30 cm long, branching at the base; leaflets 8-11, the largest 2- long, 3-6 mm wide, conduplicate; standard 9-10 mm long, the keels densely ciliate above distally; Goi: 12-15 mm long _ 5. 2. L. costaricensis l. Lupinus clarkei Oerst., Vidensk. Meddel. Dansk Naturhist. Foren. Kjgben- havn 1853. 1. 1854. type: Costa Rica, Oersted 4891 (C, not seen, photo UMO; isotype K).—Fic. 33. L. elegans var. Mea eh re Smith, Sp. Lupulin. 93. 1938, not var. campestris (Schlecht. & Cham.) Smith, Sp. L . 77. 1938. Based on Seemann (K, not seen by Smith or this writer), Maxon 5304 (US) a eri 3605 (herb. Smith, not seen). Normally annual branched herb to 1.5 m tall, mature stems terete, rigid, puberulent to tomentose, the twigs drying minutely angled. Leaves 5-7 digitately compound, the leaflets 3.5-4.5 cm long, 6-10 mm wide, narrowly ovate to ob- lanceolate, apically obtuse or rounded, mucronulate, basally narrowed into the indistinct petiolule, the costa prominent, conspicuously elevated beneath, sub- plane above, the lamina glabrate to sparsely pubescent on both sides, the hairs spreading and curled, to 1 mm long, shorter and denser beneath, the margins pilose, the undersides with a whitish bloom; petiolules to 1 mm long; petioles slender, 3-5 cm long, slightly exceeding the leaves; stipules narrowly deltoid to linear subulate, ca. 3 mm long, dark, pubescent. Inflorescences showy terminal racemes, blue or purple with pinkish or reddish tinges, the rachis tomentose with curved hairs and moniliform hairs to 1 mm long, 6-15 cm long, the flowers un- crowded on pedicels ca. 3 mm long. Flowers with the calyx deeply divided, the lower lip somewhat reflexed, 4-5 mm long, entire or 3-denticulate tomentulose, 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 721 Q UNT. y NNNM WN \ (/ t x | | ANY FIGURE 33. Lupinus clarkei Oerst.—A. Young stem ( x31⁄4).—B. Stem at end of growing season (x314).—C. Leaflet (x124).—D. Stipule (<3%).—E. Bract (x3V5).—F. Flower (x314).—G. Calyx (x314).—H. Standard (x 31⁄4).—I. Wing petal (x 31⁄4).—J. Keel showing outline of gynoecium. [After series of 25 dissections at hand.] the upper lip ca. 5-7 mm long, minutely bifid, applied to the standard; standard blue or purple, 10-13 mm long, broad, the wings apically united, often partly pinkish or red, slightly longer than the standard, the keel glabrous, slightly saccate with narrow beaks, sometimes sparsely ciliate; anthers dimorphic, the lowermost linear, 2 mm long, the uppermost ovoid, ca. 0.5 mm long; ovary copiously as- cending pilose, the style evenly curved upward, the stigma capitate, long bearded. Legume oblong, ca. 4 cm long, 8-10 mm wide, with distinct, oblique constrictions between the 7-8 seeds, drying dark, sometimes slightly reddish, copiously pilose with hairs to 2.5 mm long, minutely beaked; fruiting pedicels stout but not much longer than in flower; seeds plump, mottled, dark, ca. 5 mm long. Lupinus elegans H.B.K. [TvPE: Mexico, herb. H.B.K. (P not seen; microfiche MO)] to which the Cham. & Schlecht., Lupinus campestris was subordinated by Smith is not closely related to either L. clarkei or L. campestris. Lupinus clarkei and L. campestris are part of a closely related complex. This species is known from Panama, Costa Rica and Honduras and is found in open upland sites, commonly above 2,000 m elevation. CHIRIQUÍ: 10 mi above Boquete on road to Volcán, Croat 34819 (MO). Potrero Muleto, Croat 34946 (MO). Roadside to 10 km below summit of El Barü toward Boquete, D'Arcy 9776, 9791, 11055 (all MO). W slope of Chiriquí Volcano, Davidse & D'Arcy 10246 (MO). Boquete District, Volcán de Chiriquí, Davidson 866 (GH, MO, US). Cuesta Grande, E Slope of Chiriquí Volcano, 2900 m, Maxon 5304 (US). Volcán Bart, Mori & Bolten 7419 (MO). Finca Lérida to Peña Blanca, 1750-2000 m, 722 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 os & Schery 326 (MO, US). Boquete District, Volcán de E 11200 n Terry 1328 (MO). n 5 mi NE of El Volcán, 7500 ft, Tyson 6349 (MO). Potrero Muleto summit, Chiriquí Volcano, Woodson & Schery 394 (MO). Casita alto, Chiriqui Volcano. 1500- 2000 | m, Woodson et al. 805 (A, ). 2. Lupinus costaricensis Dunn"’. TYPE: Costa Rica; Dodge & Thomas 6826 (MO- holotype; photo UMO, isotypes GH).—Fic. 34-35. L. aschenbornii sensu Standley, Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 545. 1938, not Schauer, Linnaea 20: 739. 1847. Annual herb, occasionally extending into a second season of growth, then ligneous below, weak stemmed and sprawling to decumbent, to 15-30 cm long, branching mainly at the base, the slender terete stems only 2.5 mm diameter, densely canescent with curved hairs ca. 0.5 mm long, and with numerous spread- ing pilose hairs 1-2 mm long, glabrescent. Leaves 8-11-digitately compound, the larger leaflets 2-3 cm long, 3-6 mm wide, generally conduplicate, glabrous above, thinly pilose beneath; petioles filiform, 4—6 cm long, twice as long as the leaflets, pubescent as on the stems; stipules subulate-caudate, 12-15 mm long, the free tips 6-8 mm long, with both long and short hair types. Inflorescences inconspic- uous, the lower flowers held within the foliage, the rachis with both hair types as on the stems, racemes only 5-12 cm long, the flowers whorled or scattered; bracts 5-10 mm long, caducous by anthesis; pedicels 2.5-3.5 mm long, hispidu- lose. Flowers with the calyx strongly 2-lipped, the lower lip nearly straight, 5— 6.5 mm long, entire or tridentate, densely pilose canescent outside, the upper lip 4-5 mm long, deeply bifid, the notch 1.6-3 mm deep; the standard obovate; 9-10 mm long, 7-8 mm wide, the wings apically connate, slightly longer than the standard, less than 5 mm wide, occasionally with lateral cilia on the veins of the crimped basal portion, the keels densely ciliate above toward the acumen; ovary strongly ascending pilose, the style curved upward, the stigma capitate, bearded. Legume oblong, 2-3 cm long, 6-7 mm wide, canescent and copiously pilose with longer hairs, slightly constricted between the 7-8 seeds; seeds dark, mottled, convex, 3 mm long, 2 mm wide. The taxon described here is quite distinct from L. aschenbornii, which is caespitose, often fistulose, perennial and freezes back annually to a subsurface fleshy taproot. C. P. Smith studied the type of L. aschenbornii and he cited multiple specimens from Nevado de Toluca and Popocatepetl as typical of L. aschenbornii. Smith considered Nevado de Toluca as the most probable type locality. This species is known only from Volcán Chiriquí and Costa Rica. It is found high on the slopes of volcanos in open areas. Plants with only the primary inflo- ? Lupinus costaricensis Dunn, spec. nov. Herbae annuae, caules 2.5 mm crassi, pagina multis pilis parentibus 1-2 mm longis ferenti, canescenti pilis crispis brevibus 0.5 mm longis. Foliola foliorum majorius 8-11, linear- dances conduplicata, supra glabra, inferius tenuiter pilosis, 2-3 cm longis, 3-6 mm latis; petioli filiformibus, 4-6 cm longis; stipulis subulatis-caudatis, 12-15 mm longis, a petioli 6-7 mm ad basim connatis. /nflorescentiae inconspicuae, racemis 5-12 cm longis. Flores calycibus pner iun superiore 4-5 mm longo, bifido, incisura 1.6-3 mm profunda, labio inferiore 5-6.5 mm longo, o vel tridentato, vexillo obovato, 9-10 mm longo, 7-8 mm lato, carina dense ciliata supra distaliter mans dense pilosum, oblongum, 2-3 cm longum, 6-7 mm latum; semina 7-8, fusca, 3 mm longa, 2 mm lata. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae ) Q / À V lI 1 Cy Z çN PP | Z S 4 ZG SA — N > d FIGURE 34. Lupinus costaricensis Dunn. Habit (x14). 723 724 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 IGURE 35. Lupinus costaricensis Dunn.—A. Young stem ( x3!4).—B. Stem at end of growing season (x3!5).—C. Leaflet (x125).—D. Stipule (x124).—E. Bract (x3!4).—F. Flower ( x3!4).— Calyx (x3%).—H. Standard (x3V5).—I. Wing petal (x3%).—J. Keel showing outline of gynoecium (x313). [After a series of 15 dissections at hand.] rescences flowering were generally collected from January through March. Oc- casional collections from August to October were sprawling and ligneous below, with multiple inflorescences in fruit and only occasional late branches with few flowered racemes. COSTA RICA: CARTAGO: Volcán Irazu, 10000-11330 ft, Allen 682 (GH); 2400 m, Blaisdell 2 (MO); 3350 m, Brooks, Jan. 1970 (UMO); Davidse & Pohl 839 (MO); 2400 m, Dodge 3423 (MO). Finca Chiluca, S slope Volcán Irazü, Dodge & Thomas 6826 (GH, MO). Volcán Irazú, 9000 ft, Hunnewell 16632 (GH); 10000 ft, Mart 4798 (GH); 3350 m, Moore, Jr., 6654 (BH); Seibert 1622 (MO); Webster et al. 12093 (MICH). Finca Chiluca, S slope Volcán Irazü, Wilbur & Stone 8734 (MSC). Volcán Irazu, 3350 m, Williams 16024 (MO); Williams & Molina 13905 (GH). PANAMA: CHIRIQUÍ: Cerro Pando, Valley of upper Río Chiriquí Viejo, White 51 (MO) 31. MACROPTILIUM James A. Lackey*5 & W. G. D'Arcy*? Macroptilium (Benth.) Urb., Symb. Ant. 9: 457. 1928. Phaseolus sect. Macroptilium Benth., Comm. Legum. Gen. 76. 1837. TYPE: M. lathyroides (L.) Urban. Vines or sometimes erect or sprawling herbs, sometimes perennial from a thick rootstock. Leaves pinnate trifoliolate or rarely unifoliolate; leaflets glabrous or pubescent but lacking uncinate hairs, stipels ciliate to tomentose; stipules nervate, not prolonged below the insertion. /nflorescences with stiff, elongate °° Department of Botany, U.S. Natural History Museum, Smithsonian Institution, Washington, D.C. 20560. 5" Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 725 peduncles, the flowers mostly congested at the apex; bracteoles narrow, at least distally, nervate, caducous, rachis with small swellings at the nodes, lacking extrafloral nectaries; pedicels equalling or shorter than the calyx. Flowers pur- plish, red, violet, or white, the interior often vivid; calyx campanulate, the teeth free, the upper 2 teeth sometimes reduced; standard orbicular, emarginate, with 2 small basal auricles, lacking median thickenings; wings longer than the standard and the keel, long stipitate, the keel petals apically spiralled, basally adnate to the staminal tube; style apically recurved and thickened, caducous. Legume lin- ear, turgid or compressed, non septate; seeds numerous, small with a short hilum. Macroptilium is a genus of about 20 species, all native to the New World tropics, although at least one species is naturalized in the Old World. The plants are sometimes planted for forage, and the seeds are sometimes used for food. a. Stems conspicuously hollow, easily crushed, often 3-7 mm in diameter; quisa 12- mm long D ndn aa. Stems solid or with a minute, less than 1 mm diameter, lumen, not easily crushed, 1.0-2.5 (4) mm in diameter; standard 8-13 mm . Leaflets linear to lanceolate; pods 3—4 cm long 2. M. gracile bb. Leaflets ovate to broadly lanceolate; pods 4—9 cm long. c. Plants pubescent; leaflets generally bons or mitten- "e stems usually soli MM cc. Plants minutely pubescent to almost glabrous; leaflets generaly entire stem minutely hollow 4. Sieg iaae M 1. Macroptilium atropurpureum (DC.) Urban,® Symb. Ant. 9: 457. 1928. Phaseolus atropurpureus DC., Prodr. 2: 395. 1825. ‘atropurpureus.’ Based on an unpublished plate Sessé & Mociño, Icones Florae Mexicana (G). Twining and clambering vine; stems long pilose. Leaves pinnate trifoliolate; leaflets ovate, elliptical or linear, often 2-3 lobed, the lobes and sinuses mostly rounded; apically acute or obtuse, mucronate, basally obtuse or truncate, the costa and the major lateral veins prominent beneath, obscure above; softly to- mentose on both sides, more so beneath, mostly 2—4 cm long, 8-25 mm wide; lateral petiolules ca. 1.5 mm long, terminal petiolule ca. 2 mm long, petiole and rachis spreading short pilose; stipels subulate, 1 mm long, pubescent; stipules narrowly acute, striate, ca. 3 mm long. Inflorescences to 30 cm long, the flowers distributed along the apex of the peduncle; peduncle retrorsely short pilose; bracts and bracteoles narrowly ovate, acute, 1-1.5 mm long, cucullate, nervate, ciliolate; pedicels 1-2 mm long, puberulent. Flowers deep red to black purple; calyx tu- bular campanulate, densely pilose or merely puberulent, the tube 4 mm long, the teeth lanceolate, ca. 2 mm long, subequal; standard 1.5-2 mm long. Legume linear, 7-10 cm long, 2-4 mm wide, appressed pilose; seeds oblong ovoid, dark, shiny, ca. 3 mm long, the aril white. Macroptilium atropurpureum is distinct in its pubescent, usually lobed leaves. This species ranges widely in the neotropics. In Panama it occurs in Pacific watershed areas with substantial dry season. 68 For a list of other names see Maréchal et al., 1978. Only these names have been used for Panamanian plants. 726 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 CANAL ZONE: Ancon, Greenman & Greenman 5066 (DUKE). La Boca, Mori & Kallunki 3675 (MO). Between Corozal and Ancón, Pittier 2173 (NY). 1 mi N of Summit, Tyson et al. 2744A (SCZ). Contractors Hill, Tyson & Dwyer 6494 (MO). CocLE: Rio Hato Airstrip, Burch et al. 1129 (MO.) Los SANTOS: Chitré- p Tablas, 0-100 m, Burt & Rattray 52, 98 (both MO). Monagre Beach, 5 mi SE of Chitré, Tyson et al. 3011 (SCZ). PANAMÁ: 2 km W of entrance to Lago Mar on Panamerican Highway, Folsom & Lantz 1901 (MO). Sardinilla, Taylor 37 (MO). 2. Macroptilium gracile (Benth.) Urban, Symb. Ant. 9: 457. 1928. Phaseolus gracilis Poepp. ex Benth., Ann. Wiener Mus. Naturgesch. 2: 77. 141. 1838. TYPE: Cuba, Poeppig, not seen. Perennial herb or vine, erect or sprawling; stems slender, wiry, with minute, white retrorse hairs on emerging, soon glabrous, terete; rootstock woody, en- larged. Leaves pinnate trifoliolate; leaflets linear, mostly 3-7 cm long, 2-5 mm wide, apically pointed or blunt, mucronulate, basally narrowed, truncate, rarely ovate and 7-8 mm wide, glabrous or sometimes scabrous above; petiolules in- conspicuous, ca. 1 mm long, yellowish, strigose; petioles slender, about as long as the terminal leaflet, strongly angled, minutely puberulent; stipels 0.5-1.0 mm long, scale-like; stipules subulate, pubescent, ca. 1 mm long. /nflorescences ter- minal, 20-25 cm long; peduncle slender, the nodes swollen, glandular; bracteoles and bracts subulate, ca. 1.5 mm long, ciliate; pedicels 1 mm long, ca. 2 mm long in fruit, disposed along the apex of the peduncle. Flowers red, purple or pink; calyx tubular, ca. 4 mm long, the teeth subequal, deltoid to subulate, much shorter than the tube, evenly minutely puberulent; standard 1.5 cm long, the keel spi- ralled. Legume linear, 3-5 cm long, ca. 2 mm wide; seeds (Standley & Steyer- mark) 2 mm long, shiny. This species resembles Vigna linearis, but the leaves are often scabrous and are more pubescent. Only the major lateral veins of the leaves are prominently at right angles to the costa whereas in V. linearis, much of the minor venation parallels the major lateral veins. The calyces are quite different in the two species. Macroptilium gracile is wide ranging in Central and South America and in the Antilles, occurring in grassy savannahs with seasonal drought. In Panama it has been collected in the Pacific lowlands. CLÉ: Penonomé, . 131 (NY, US). PANAMA: Pacora, 35 m, Allen 815 (MO, NY), 9 aed pipe of lower Río Cabra, Maurice 791 (US). Panama Golf Course, Piper 5146 (US). Sa s na Juan Corso near A 60-80 m, Pittier 4745 (US). Between Rio Pacora and Chepo, Porter et al 5137 (MO). 3. Macroptilium lathyroides (L.) Urban,*® Symb. Ant. 9: 457. 1928.—Fic. 36. Phaseolus lathyroides L., Sp. Pl., ed. 2: 1018. 1763. TYPE: not seen. Erect herb, later sprawling or sometimes twining; stems tough, slender, re- trorsely silver pilose on emerging, often glabrescent. Leaves pinnate trifoliolate; leaflets ovate or elliptical, mostly 2-4 cm long, 1.5-3.5 cm wide, apically obtuse, short acuminate or acute, rarely rounded, basally deltoid, obtuse, or rounded; densely appressed pilose on emerging, soon glabrate; petiolules 1-2 mm long, ** For a list of other names see Maréchal et al., 1978. Only these names have been used for Panamanian plants. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 727 FIGURE 36. Macroptilium lathyroides (L.) Urban.—A. Habit (x!4).—B. Flower (x2).—C Corolla.—C!. Standard (x1%).—C*. Wing petal (x1!/2).—C?. Keel petal (x1!4).—D. Stamens (x2V2).—E. Pistil (x24). [After Hunter & Allen 721.|—F. Fruit (x 14). [Afer Hunter & Allen 8601.] ascending pilose; petiole and rachis slender, drying angled; stipels lanceolate, glabrous, striate, ca. 3 mm long; stipules lanceolate, striate. Inflorescences elon- gate, mostly 10—30 cm long, the flowers mostly near the apex; peduncle narrowing upwards, the nodes enlarged; bracteoles subulate, pilose, caducous; pedicels shorter than the calyx, often twisted, pilose. Flowers scarlet to purple red to nearly black; calyx tubular, 4-6 mm long, pubescent, the teeth lanceolate, much shorter than the tube; standard 12-15 mm long; keel spiralled. Legume linear, straight or slightly curved, mostly 7-12 cm long, ca. 3 mm wide, evenly somewhat turgid, maturing brown, inconspicuously strigose; seeds oblong ovoid, 2-3 mm long, dark, mottled. ANAL ZONE: Mindi, Cowell 186 (NY). Ancón, Greenman & Greenman 5067 (MO). Gamboa, Heriberto 98 (US). Between Farfan Beach and Palo Seco, Hunter & Allen 438 (MO). Las Cruces Trail, 75 m, Hunter & Allen 721, 734 (both MO). Fort Randolph, Maxon & Harvey 6547 (US). Balboa, Standley 25628 (US). Summit, Standley 27324 (US). Gamboa, Standley 28459 (US). Fort Randolph, iniit 28745 (US). Summit, Srandley 30047 (US). Darien Station, ri 31575 (US). COCLÉ: du La Mona, Blum & Tyson 1886 (MO, SCZ). 2.5 km S of Ant En e 4297 (MO). COLÓN: Las Cruces Trail at Portobelo, Ebinger 125 (F, MO). Colón, Rose 22070 (G H, US). HERRERA: Chitré- Davisa, 0-100 m, Burt & Rattray 67 (MO). 2 mi W of Divisa, Tyson 5153 (SCZ). LOS SANTOS: Between Tonosi and Macaracas, Oliver et al. 3576 (MO). PANAMA: Matachui to Las Cascadas, Cowell 337 (NY). Hills between Capira and Potrero, 80-130 m, Dodge & Hunter 8601 (MO). Taboga Island, Dwyer 3102 (MO). Juan Franco Race Track near Panama, Standley 27684 (US). 728 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 4. Macroptilium longepedunculatum (Benth.) Urban, Symb. Ant. 9: 457. 1928. Phaseolus longepedunculatus Mart. ex Benth., Comm. Legum. Gen. 77. 1837. sYNTYPES: Brazil, Paraguay, none seen. P. campestris Benth., Comm. Legum. Gen. 77. 1837. rYPE: non Benth. 1859. Perennial vine; stems pilose with weak spreading or appressed hairs. Leaves pinnate trifoliolate; leaflets ovate, rhombic, elliptical oblong or linear, sometimes shallowly lobed, mostly 2-4 cm long, 1-3 cm wide, sericeous to short pilose, perhaps sometimes scabridulous; apically obtuse, mucronulate, basally truncate; petiolules ca. 1 mm long, tomentose; petioles pilose, drying strongly angled, sometimes longer than the terminal leaflet; rachis short; stipels lanceolate, ca. 1 mm long, long ciliate; stipules lanceolate subulate, 3-5 mm long, nervate, ciliate. Inflorescences 20-30 cm long, the flowers clustered at the apex or distributed along the top half of the peduncle, mostly paired; peduncle glabrate; bracteoles and bracts ovate, acuminate, nervate, ciliolate, 1-2 mm long; pedicels stout, 0.5— 1 mm long. Flowers red to violet; calyx tubular campanulate, pubescent, the tube ca. 3 mm long, the teeth deltoid, the upper pair connate; standard ca. 1.5 cm long. Legume linear, straight or slightly falcate, 3-6 cm long, 2-3 mm wide, turgid CANAL ZONE: Fort Sherman, Piper 5887 (NY). Near Old Fort San Lorenzo, Piper 5961 (NY). COCLE: Aguadulce, eee 4889 (US). Los SANTOS: Chitre-Los Santos, Burt & Koster 97 (MO). PANAMA: Betwee o Pacora and Chepo, Dwyer et al. 5094A (MO). vERAGUAS: Atalaya-Santiago, 100-200 m, Pini 114 (MO). 32. MUCUNA W. G. D'Arcy?? Mucuna Adans., Fam. Pl. 2: 325, 579. 1763. Nomen conserv. TYPE: M. urens (L.) DC. Type conserv. — eiie P. Br., Hist. Jam. 295. 1756. Based on Dolichos urens L. = Mucuna urens (L.) DC. ne ` e Elem . Bot. 3: 1790. Citta Loureiro, Fl. Cochinch. t Pe 1790. T: C. nigricans Loureiro — Mucuna nigricans (Lour.) ackson. Negretia ie Pavón, Prodr. 98, tab. 21. 1794. LECTOTYPE: N. elliptica R. & P. = Mucuna elliptica (R. ibus im Raddi, Mem. Mat. Fis. Soc. Ital. Sci. 18 (fis.): 392. 1820. TYPE: M. pseudo-stizolob- ium Raddi. Literature: Burkart, A. 1970. Las Leguminosas-Faséolas Argentinas de los géneros Mu- cuna, Dioclea y Camptosema. Darwiniana 16: 175—218. High climbing woody vines. Leaves pinnate trifoliolate, the lateral leaflets oblique; stipels scale-like; stipules often caducous. /nflorescences axillary ra- cemes, the peduncles often elongate and the pedicels subumbellate, at least in bud, bracts often subfoliaceous, enclosing the bud, caducous; pedicels arising in 70 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 729 2's and 3's on an expanded portion of the peduncle. Flowers showy, the calyx campanulate, often with irritating hairs, the upper teeth connate, the lower 3 usually unequal; corolla with the standard shorter than the wings, the wings with margins basally ciliate, the keel narrow, apically falcate and indurate; stamens diadelphous, the upper stamen free, the filaments alternately thick and thin, long and short, the anthers sometimes pilose; ovary tomentose, the short stipe sur- rounded by a glandular disc, the style slender, glabrous or pubescent, the stigma capitate, sometimes of a tuft of hairs. Legume oblong, thick or flattened, the margins often winged, undulate between the seeds and somewhat compressed laterally between the seeds, the surface sometimes lamellate with parallel or irregular raised lamellae which may form elongate enations, mostly densely cov- ered with stiff irritating hairs, tardily dehiscent; seeds flat or convex, discoid, the hilum narrow around more than % the periphery. Mucuna Is widely distributed in the tropics, mostly confined to moist or wet forests. The genus includes perhaps a dozen species. a. Inflorescences short, less than 15 cm long; flowering pedicels less than 10 mm long; flowers more than 5 cm long, the wings more than 15 mm wide at the middle __________ 3. M. rostrata aa. e elongate, the peduncle more than 30 cm long; oo pedicels more than m long; flowers less than 5 cm long, the wings less than 15 m passa ey pubescent beneath, at least until flowerin lowers less than 4 cm long, the lowest calyx tooth broad, less than 3 net long ella 1. wings less than 8 mm wide; fruit flat, lacking raised lamellae ________ M. holtonii cc. Flowers more than 4 cm long, the lowest calyx tooth narrow, more than 5 mm long; wings more than 10 mm wide; fruit thick, with conspicuous raised trans- verse lamellae 4. M. sloanei bb. Leaves glabrate beneath. Lowest calyx tooth narrow, 7-13 mm long; lateral leaflets mostly 3-veined from the base; fruit mostly 3-7 seeded 5. M. urens dd. Lowest calyx tooth deltoid, 3-5 mm long; lateral leaflets mostly 4-veined from the base; fruit mostly 1-3 seeded 2. M. mutisiana 1. Mucuna holtonii (Kuntze) Moldenke, Phytologia 1: 7. 1933. iir eee holtonii Kuntze, Rev. Gen. Pl. 1: 207. 1891. syNTYPES: Costa Rica, Kuntze (?NY, not and Colombia, Holton 971 (K, not seen). Mucuna ae Micheli, Morot. J. Bot. 6: 146, tab. 5, 6. 1892. TvPE: Colombia, André 1978, not en. Also cites Holton 971, not seen. High climbing vine, stems slender, slightly angled, minutely tomentulose, tar- dily glabrescent. Leaves trifoliolate, the leaflets about equal, 9-15 cm long, 5-9 cm wide, apically short acuminate, mucronate, manifestly costate, the terminal leaflet with 5-6 major veins on each side, the lateral leaflets strongly oblique, chartaceous, velutinous, later glabrescent especially above, with only a few hair bases remaining; petiolules 7-10 mm long, tomentose, glabrescent and turning black; petioles mostly shorter than the leaves, tomentose, glabrescent; stipules caducous; stipels ca. 3 mm long. Inflorescences pendant, contracted, racemes with peduncle elongate, to 5 m long, slender, flexuous, minutely tomentose ve- lutinous, the raceme enclosed in bud by gray velutinous, papery deciduous bracts and forming a globose ‘bud’ ca. 5 cm across, sinuous; pedicels velutinous, 15—18 mm long upon inflorescence opening, becoming 5 cm long at anthesis and stout in fruit. Flowers greenish or yellowish, the calyx deciduous, cupular, velutinous 730 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 outside with appressed, ascending gray or golden hairs, ca. 10-15 mm long, pilose inside, the teeth small, deltoid umbonate; corolla ca. 4 cm long, drying dark, the standard 18-22 mm long, the wings and keel narrowly tubular, exserted about 4 their length from the standard; anthers basally barbate with elongate hairs, the stigma capitate, a tuft of ascending hairs. Fruit oblong, flat, 1-6 seeded, 14-25 cm long, 5 cm wide, the margin conspicuously undulate between the seeds, softly villous outside, lustrous within and falsely septate between the seeds; seeds dis- coid, black, 2-3 cm wide, ca. 0.8 cm thick, the hilum linear around ?/,-*/, of the periphery, lighter colored. Mucuna holtonii has leaves velutinous on both sides until time of flowering. The inflorescence is greatly contracted, appearing in bud as a ball at the end of a long, slender, pendant peduncle, and later with the flowers on elongate and slightly bent pedicels. The legumes are flat with undulate margins. The legume lacks stinging hairs, but there may be some on the calyx. This species is found in Panama from sea level to at least 1,500 m, occurring in forests and in areas of disturbance. It ranges from Mexico (Chiapas), to Co- lombia. It is apparently closely related to Peruvian and Ecuadorean material going under the name M. elliptica Ruiz & Pavon, which differs in having seeds slightly thicker and more convex, and in the legumes having stinging hairs. The appear- ance of the legume and also of its indumentum is much like that of M. elliptica. BOCAS DEL TORO: Banks of Changuinola River, Dunlap 441 (F, US). Changuinola to 5 mi S at junction Rios ao and Terebe, 100-200 ft, Lewis et al. 943 (GH, MO, US). Almirante Bay, Wedel 8 (MO, US). c - ZONE: Road C-16 NW of Pedro Miguel, Croat 12255 (MO). Pipeline road 3 mi N of Gamboa pA " Croat 13951 (MO, SCZ). Madden Dam, Dwyer 1965 (MO, PMA Between Chilibre and Madden Dam on Transisthmian Highway, Dw ver 8396 (GH, MO). Chagres River 3 mi above Gamboa Bridge, Kennedy et al. 2304 (F, MO). Pipeline Road 2 km NW of Gamboa, 50 m, Nee 7853 (MO, US). Gatuncillo, d ddp: (US). Gamboa, Pittier 4788 (US). cociÉ: Limón, N of Alto Calvario, Pato d 5848 (MO). c N: Quebrada Bonita, 13 km NW of Buena Vista, 3 km of Pirre, Gentry & Clewell 6922 (MO). Río Pucro below Pucro, Gentry & Mori 13531 (MO). Quebrada Camachimuricate cerca casa de Bartolo, Kennedy 2840 (F, GH, MO). Caná, 1750 ft, Stern et al. 482 (GH, MO). veraGuas: Above Santa Fé on slopes of Cerro Tute, 1200-1400 m, Gentry 6280 (MO). 2. Mucuna mutisiana (H.B.K.) DC., Prodr. 2: 406. 1825.— Fic. 37. Negretia mutisiana H.B.K., Nov. Gen. Sp. Pl. 6: 442. 1823. type: Colombia, Mutis, not seen. High climbing vine; stems terete with straight, appressed reflexed hairs, soon glabrous, drying black. Leaves trifoliolate, the leaflets about equal, mostly 7-12 cm long, ovate, apically acuminate, basally obtuse or rounded; lateral pair mostly 4-veined, sparingly pilose with straight hairs on both the surfaces, thin and pa- pery, often drying dark; petiolules 7-10 mm long, sometimes stout, the terminal petiolule articulated near the apex, glabrate, tuberculate; petioles mostly some- what shorter than the leaves; stipels scale-like, 1-3 mm long, glabrate; stipules narrowly ovate, pilose, often caducous. /nflorescences subterminal pendant ra- cemes, the peduncle to 3 m long, slender, terminally fine gray puberulent and minutely angled; pedicels 25-40 mm long becoming slightly longer and stout in fruit; bracts broadly ovate, mostly 15-18 mm long, evenly gray velutinous, often 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 731 IGURE Mucuna mutisiana (H.B.K.) DC.—A. Leaf with 2 leaflets remaining.—B. Fruiting inflorescence (X14). [After Hammel 5442.]—C. Flower (x4). [After Croat 12790.] caducous. Flowers showy, white, cream, green or brown, often with purple mark- ings, the calyx cupular, ca. 10 mm long, evenly gray or brown velutinous, the teeth short deltoid, the longest 3-5 mm long; standard ca. 2.5 cm long, mostly enveloping the wings and the keel, the wings 4—5 cm long, darker, connate, apically broadening, the keel 3.5-5 cm long forming a slender tube, but not con- nate along the upper edge, apically sharply bent and indurated; stamens with the 732 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 filaments united 34 way up, the free portion equal, the united portion biseriate, apically curved; ovary ascending tomentose, the style ascending pilose, less so apically, the stigma minute, peltate. Legume few seeded, 6-15 cm long, 3-5 cm wide, constricted between the seeds and undulate along the margin, transversely or obliquely ridged with narrow, irregular lamellae to 3 mm tall, the entire legume covered with brownish deciduous stinging hairs; falsely septate between the seeds, tardily dehiscent; seeds dull, dark brown, convex-discoid, the hilum linear around ?4 of the periphery, the testa hard, almost flinty. This species is distinguished from other Panamanian species of Mucuna by the short lobes on the calyx and by the lamellae on the fruit which sometimes form elongate enations more than 10 mm long. Mucuna mutisiana occurs in Costa Rica, Panama, Colombia and perhaps far- ther south. RO: Almirante-Changuinola Canal, ra 1407 (FSU, MO, SCZ). Junction Rios Se and Teribe, 100-200 ft, Lewis et al. 935 (GH, MO). Chiriquicito to 5 mi S along Rio Guarumo, Lewis et al. 2094 (MO). Chiriqui Lagoon, a 1003 (GH, MO), 1097 (GH, US), 1106 (GH, MO, US). Water Valley, Wedel 1486 (GH, MO, US). Old Bank Island, num (MO, US). Fish Creek Hills, Wedel 2430 (GH, US). CANAL ZONE: BARRO COLORADO ISL : Bailey & Bailey 348 (F); Croat 4800 (MO, SCZ), 7425 (F, MO), 7426 (MO, SCZ), 10167 (MO. SCD, gi (MO), 12593 (MO, PMA, SCZ); Foster 682 (F, GH, PMA), 1322 (F, PMA), 1424 (F, PMA); Shattuck 286 (F, MO) Wetmore et al (A, F, GH, MO). Road to Pina, 1 sey & Mahler 344 (F, GH, O). Pipeline Road near Gamboa, Clewell & Tyson 3262 (PMA). Dirt road to Chiva-Chiva town, Correa 495 (MO, SCZ). Highway NW of Summit Garden, Croat 12611 (F , SCZ). Near Summit Golf Course, Croat 12790 (MO, SCZ). Allbrook U.S. Army Tropic Test Center, Dwyer 7116 (MO). Chagres, Fendler 459 d E of Fort Clayton, Harvey 5119 (F). K 9 road, Hladik 42 (MO). Chagres River 3 mi above Gamboa Bridge, Kennedy et al. 2306 (MO). Río Grande near Culebra, Pittier 2139 (US). Between France Field and Catival, Standley 30212 (U). George Green Forest Preserve behind Cerro Tigre, Tyson et al. 2511 (FSU, MO, SCZ). 1 mi N of Summit on road to Gamboa, Tyson et al. 2754 (FSU, MO, SCZ). Farfan Beach area, Tyson et al. 3185 (MO). cocié: El Valle, Ebinger 1124 (F). prre 50—1000 ft, Mn ee (US). DARIEN: El Real, 15 m, ea 970 (GH, US). Yavisa, GH Stern et al. 8 ). LOS SANTOS: 10 mi SW of El Cortezo, southern Azuero, Hammel 5442 (MO). PANAMÁ: Río Teribe near El ias Duke 5675 (MO). Río Mamoní, Duke ps ei A (MO). San Jose Island, ou 209 (GH), 245 (GH, MO, US), 677 (GH). Alcalde Díaz, , Nee 8297 (US). Matías Hernández, Pittier 6885 (GH, US). Río Tapía, Standley 28275 tus. SAN BLAS: Opposite Ustupo-Concepción, Mori 393 (WIS). 3. Mucuna rostrata Benth. in Mart., Fl. Bras. 15(1): 171. 1859. TYPE: Brasil, Spruce 1625 (?K, not seen). High climbing woody vine, stems glabrate. Leaves trifoliolate, the leaflets mostly 9-12 cm long, ovate, apically acuminate, basally obtuse or truncate, 3—4 veined from the base, above glabrous except for appressed ascending hairs on the principal veins, beneath sparingly short pilose to glabrous, the principal veins with stout, mostly appressed and ascending yellowish or brownish hairs, often tufted at the very leaf base; petiolules 7-9 mm long, pubescent, drying dark; petioles mostly about as long as the terminal leaflet, glabrate; stipels linear, ca. 3 mm long; stipules 3-4 mm long, ascending pubescent. Inflorescences short racemes, the peduncle 4-13 cm long, the pedicels subumbellate in bud, often becoming well spaced in flower, ca. 10 mm long, appressed pubescent, becoming 10-20 mm long and stout in fruit; bracts subfoliaceous, velutinous on both sides, early caducous, perhaps confined to the lower portion of the inflorescence. Flow- ers showy, yellow or orange, calyx oblique cupular, softly velutinous with minute 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 733 hairs inside and out, and with scattered stout hairs on the cup outside, 2—3 cm long, the upper 2 teeth united into a broadly obtuse hood or into a deltoid lobe 5-8 mm long, the lower 3 teeth narrowly deltoid to linear, ca. 10 mm long; corolla mostly drying black, 6-9 cm long, the standard !2—25 as long as the wings, to 5 cm long, the wings mostly rhomboidal, to 1.8 cm wide, sometimes slightly acu- minate; keel narrowly falcate, the apex indurated and often drying yellowish; anthers copiously pilose with curly hairs; stigma a capitate cone of hairs. Legume oblong, compressed but apparently little compressed between the seeds, with numerous, parallel transverse lamellae to 5 mm tall, 4-5 mm apart, copiously hispid with brownish stinging hairs; seeds not seen. Mucuna rostrata is distinct from other Panamanian members of the genus in its short racemes, short pedicels, and large flowers. The anthers are more pu- bescent, the stigma is bigger than other species, and the wing petals are relatively much wider. Little fruiting material has been seen or described in the literature, and the uniformity of the pod is unknown. The pubescence of the leaflet under- sides is variable in amount. This species has been collected a number of times in Bolivia, Peru, and Ec- uador but is a rare plant in Panama. CANAL ZONE: BARRO COLORADO ISLAND: Netting 25 (F, MO); Shattuck 489 (F, MO, US); Wet- more & Abbe 12 (A, F, GH, MO); Wilson 136 (F); Zetek 15000 (F). 5 km SE of Achiote, Gentry & Nee 8715 (MO). Lion Hill, Goldnan 1881 (US). COLON: Portobelo, Pittier 2450 (US). DARIEN: Rio Chucunaque between Yavisa and Río Tuira, Gentry 13481 (MO). Mucuna sloanei Fawcett & Rendle, J. Bot. 55: 36. 1917. Based on Phaseolus brasiliensis frutescens .... Sloane, Voy. Madera, Barbados, etc. 1: 178. 707. TYPE: Herb Sloane 3: 69 (BM, not seen, designated by Fawcett & Ren- dle). High climbing robust vine; stems appressed tomentose, especially near the nodes, glabrescent, the hairs whitish. Leaves trifoliolate, the leaflets mostly 7—10 cm long, ovate, apically acute obtuse or short acuminate, basally obtuse or trun- cate, the lateral leaflets strongly oblique, the midvein prominent, 3-veined at the base, 3-5 prominent veins on each side higher up, glabrescent above, softly fine pubescent beneath with appressed, whitish hairs; petiole mostly as long as or shorter than the terminal leaflet; petiolules ca. 5 mm long, ascending hispid; stipels linear, ca. 2 mm long, the stipules caducous. /nflorescence pendant ra- cemes, the peduncle ca. 8 cm long, appressed pubescent with whitish hairs, glabrescent but some pubescence usually persistent in lines, the flowers congested on the enlarged 1-3 cm long rachis; pedicels mostly short, to 15 mm long, pu- bescent, becoming stout in fruit; bracts subfoliaceous, 1-2 cm long, ovate, cu- cullate, hispid, early caducous; bractlets wanting. Flowers yellow, showy, the calyx cupular, the cup 6-12 mm long, papery, minutely pubescent inside and out, the hairs finer outside but outside with scattered coarse hairs, the upper teeth united into an upcurved lip 10-15 mm long, the lower pair of teeth obtuse, with linear tips, the lower tooth acute or acuminate deltoid, ca. 8 mm long, slightly exceeding the lower pair; corolla ca. 5 cm long, drying orange, the standard 2.5- 3 cm long, ?erect, the wings and the keel narrow, exserted ca. 2 cm from the um 734 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 standard, oblong falcate; style appressed pubescent, the stigma a small capitate knob. Legume oblong, mostly 2—4 seeded, ca. 10 cm long, 4 cm wide, short beaked, the surface with prominent transverse lamellae 4-6 mm high, copiously covered with deciduous, yellowish or brownish stinging hairs, the surface con- stricted between the seeds, the margin winged, the wings irregular, 4-8 mm wide, drying black; seeds convex lenticular, 2-3 cm across. Mucuna sloanei may be recognized by its pubescent leaf undersides and by its yellow flowers, and there are many minor differences separating the species from other Panamanian elements, e.g. the papery calyx, the erect standard, the orange-drying corolla, the capitate stigma, the caducous bracts and lack of brac- teoles etc. Occasional glabrate individuals, e.g. Tyson et al. 3185 may be confused with M. mutisiana. Mucuna sloanei occurs in the Antilles and in Central America. It is also found, perhaps introduced, in Africa. It has been collected only a few times in Panama, mostly near the sea. CANAL ZONE: Fort Kobbe, Duke 6078 (MO). Farfan Beach, Lewis et al. 56 (GH, MO, US). Farfan Beach area, Tyson et al. 3185 (FSU, MO). coci£: El Valle, Allen 78 (MO, US). PANAMA: San Carlos, 0-10 m, Allen 1135 (GH, MO). San Jose Island, Johnston 248 (GH, MO, US), 530 (GH, US), 957 (GH), /356 (GH). Bella Vista, Standley 25351 (US). Taboga Island, Tyson et al. 5133 (FSU, MO, SCZ). 5. Mucuna urens (L.) DC., Prodr. 2: 405. 1825. Dolichos urens L., Syst., ed. 10: 1162. 1759. Based on Zoophthalmum P. Br., Hist. Jam. 295. 1756. TYPE: Browne, Jamaica, not seen. D. altissimus Jacq., Sel. Stirp. Amer. 203, tab. 182, fig. 85. 1763. TYPE: not seen. Stizolobium altissimum (Jacq.) Pers., Syn. Pl. 299, 1807. Mucuna altissima (Jacq.) DC., Prodr. 2: 405. 1825. High climbing woody vine, stems slender, glabrate. Leaves trifoliolate, leaf- lets mostly ca. 10 cm long, 5 cm wide, ovate, apically acuminate, basally obtuse or rounded, the lateral leaflets oblique, costate with 3 prominent veins at the base and 1-3 prominent veins distally on each side, glabrescent, more so above, be- neath with scattered short hair bases persisting, the leaf base sometimes with a few stouter hairs; petioles somewhat shorter or longer than the terminal leaflet: petiolules ca. 10 mm long, drying dark; stipels caducous, stipules scale-like, 2-3 mm long, pilose. Inflorescence pendulous, the peduncle elongate, slender, gla- brate, the rachis thick, ca. 3 cm long; appressed pubescent, striate, the pedicels slender, 4-5 cm long, stout in fruit, appressed pubescent; bracts not observed. Flowers pale greenish white or yellow, the calyx cupular, the cup ca. 9 mm long, the upper teeth united into a rounded or emarginate crest, ca. 10 mm long, the lower pair of teeth deltoid to acute, 6-7 mm long, the lower tooth linear, 11-13 mm long, appressed velutinous outside and inside, a few coarse ?stinging hairs on the cup outside; corolla ca. 4 cm long, the standard, ca. 2.5 cm long, the wings clavate falcate, ca. 8 mm wide at the widest, exserted 8-12 mm from the standard, drying dark; anthers ca. 3 mm long, basally barbate; stigma capitate tufted. Fruits oblong, becoming 20 cm long, 6 cm wide, the surface and margin with conspic- uous irregular 5-10 mm high lamellae, not constricted between the seeds, copi- ously hispid with reddish brown stout stinging hairs, the beak slender, 1-2 mm 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 735 thick, the stipe slender, 2—3 mm thick, 0—2 cm long; falsely septate between the seeds; seeds not seen. This species is distinct from other Panamanian species of Mucuna in its pod, which is larger and longer with irregular lamellas, and which is sometimes beaked and stipitate. It is also distinct in its calyx, the lower tooth of which is linear deltoid and elongate. Identifying this material with M. urens, which is an Antillean species, is done with some hesitation; no good fruiting material of typical M. urens was seen, and the calyx teeth are longer than on most Antillean material. The pedicels are also much longer than those on Antillean material seen. The above description was made from Panamanian and Costa Rican material at MO. All collections of this species except for that from El Valle by Seibert are from upland Chiriqui, 1,500-2,000 m, an area from where no other species of Mucuna have been reported. CANAL ZONE: Gatun Station, Hayes 134 (GH). CHIRIQUI: Finca Collins, Boquete, Blum & Dwyer 2531A (GH, MO, SCZ). Cerro Horqueta, 4500-7000 ft, Blum & Dwyer 2605 (FSU, SCZ); Dwyer et al. 450 (MO); Dwyer & Lallathin 8738 (GH, MO). Boquete, Ebinger 740 (MO); Kirkbride 156 (MO). Finca Lérida to Pena Blanca, 1750-2000 m, Woodson & Schery 324 (MO). Bajo Mono and Quebrada Chirquero, 1500 m, Woodson & Schery 517 (MO). Casita Alta, Volcán de Chiriquí, Woodson et al. 969 (GH, MO, US). cociÉ: El Valle de Antón, 500-700 m, Seibert 434 (GH, MO). DARIEN: Between Caná and Boca de Cupe, Stern et al. 628 (GH, MO). 33. MYROXYLON Michael O. Dillon?! Myroxylon L.f., Suppl. Pl. 34, 233. 1781. Nom. cons., non J. R. & G. Forster, 1776. TvPE: M. peruiferum Toluifera L., Sp. Pl. 384. 1753; Gen. Pl., ed. 5. 181. 1754. Nom. rejic. TYPE: T. balsamum L. Myrospermum sect. Myroxylon (L.f.) DC., Prodr. 2: 95. 1825. Unarmed trees. Leaves alternate, imparipinnate, 5-15 foliolate; leaflets alter- nate, with pellucid glandular dots and lines; stipules and stipels minute or absent. Inflorescences racemose, terminal and axillary; bracts and bracteoles caducous. Flowers pedicellate, small; calyx turbinate-campanulate, 5-lobed, subequal, val- vate in bud; petals 5, whitish, subequal, free, the standard larger than the wings, the keel narrow; stamens 10, free, subequal, exserted, the anthers uniform, ob- long, acuminate, sagittate, dorsifixed; ovary long stipitate, 2-ovulate near the apex, the style short, incurved, the stigma terminal, minute. Fruits stipitate, sally; seeds subreniform, resinous, the hilum elliptic, subapical. Chromosomes: n = 14 The genus Myroxylon is represented by 2 species distributed from Mexico to northern South America and cultivated in Ceylon and the Congo. In Panama, M. balsamum is known from tropical moist forest in the Canal Zone, Panama and Darien Provinces. Rudd (1972) recognized two varieties in Panama (balsamum 71 Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 736 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 and pereirae (Royle) Harms) with the note that the most important differences between the varieties seemed to be the chemical and physical properties of the balsam. In the absence of morphological characters allowing distinction of these varieties, they are not recognized in the present treatment. Myroxylon supplies the ‘‘balsam of Peru” used pharmaceutically and in per- fumery. Despite its most common name, it was originally discovered and has since been produced commercially in El Salvador. The wood is strong and du- rable, and it is used for cabinet work. In addition to commercial uses, this species is utilized by native peoples for poles in house construction, ‘‘orcones’’ and "pilones." The Choco of Darién call this plant "pido quera," which translates to peccary perfume; the bark is burned in the sun to attract these animals. The fruits are not consumed. Literature: Baillon, M. H. 1873. Sur les toluifera et sur l'origine des baumes de Tolu et du Pérou. Assoc. Franc. l'Avanc. Sci., Lyon, 510-521, plate X. Ruiz, H. 1777-1788. Travels of Ruiz, Pavón, and Dombey in Peru and Chile nA pes ut n translation, Publ. Field Mus. Nat. Hist., Bot. Ser. 21: 176-177. — . Myroxylon balsamum (L.) Harms,” Notizbl. Bot. Gart. Berlin 5: 94. 1908.— Fic. 38 Toluifera balsamum L., Sp. Pl. 384. 1753. TYPE: -aii "in America prope Carthagenam,`` co ector unknown (presumably BAS, not Tree to 40 m tall, d.b.h. ca. 1 m; bark smooth, with abundant lenticels. Leaves usually imparipinnate, 5-10 foliolate; rachis and petiolules pubescent, terete; rachis 5-15 cm long; leaflets alternate, lanceolate to elliptic, acute to acuminate, obtuse at the base, 3-11 cm long, 1.8-4.0 cm wide. Inflorescence racemose, axillary, to 20 cm long, closely cinereous tomentose; pedicels 1.0-1.5 cm long. Flowers oblique on the pedicel; calyx campanulate, 1.5—4.5(-6.0) mm long, finely ribbed, the lobes ca. 1.5 mm long; the standard orbicular, ca. 9 mm in diameter, cordate basally, the claw ca. 1 mm long; wings elliptic to narrowly spatulate, ca. l cm long, ca. 4 mm wide, the claw 2 mm long; keel subelliptic, ca. 8.5 mm long, ca. 3 mm wide, the claw ca. 1.5 mm long; ovary short stipitate for ca. 2 mm, villosulose; style subulate. Fruit a samara, narrowly obovate, to 11 cm long, glabrous, stipitate, ca. 1 cm long; wing to 8 cm long, 1-2 cm wide, the seminif- erous area turgid at the apex of the fruit, obliquely oblong, 2-3 cm wide, ca. 1 cm thic This species ranges from southern Mexico to northern South America, where it is replaced by Myroxylon peruiferum L.f., which ranges south to Argentina. In Panama, it flowers from January to June, fruiting between September and March ?* For a list of synonyms see Rudd (1972). Only the listed names have been used for Panamanian material. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 737 FIGURE 38. Myroxylon balsamum (L.) Harms.—A. Habit (xV2).—B. Fruit (x1⁄2). N: Santa Fé, Duke 8391 (MO); Campamento Buena Vista, Stern et al. 856 (MO). PANAMÁ: TNR ph 4765 (MO). 34. ORMOSIA Michael O. Dillon? Ormosia Jacks., Trans. Linn. Soc. London 10: 360. 1811. Nomen cons. TYPE: O. coccinea (Aubl.) Jacks. (=Robinia coccinea Aubl.). Trees to 60 m, or rarely shrubby; trunks sometimes buttressed, bark gray, rough. Leaves imparipinnate, 3-19 foliolate, occasionally unifoliolate; leaflets coriaceous or subcoriaceous, 1.5-35.0 cm long, 0.5-20.0 cm wide; rachis to 50 73 Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 738 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 cm long; stipules 0.5-15.0 mm long, deltoid to linear, caducous, possibly lacking in some species; stipels rare, minute, usually acicular. Inflorescence racemose, many flowered, terminal or pseudoterminal; bracts stipuliform, deltoid to lanceo- late, 0.5-10.0 mm long, 0.5-4.0 mm wide, pubescent, frequently caducous; brac- teoles paired, subtending the calyx, deltoid to filiform, 0.5-8.0 mm long, 0.5-1.0 mm wide. Flowers yellow, lilac, or dark purple; calyx campanulate, 5-lobed, upper 2 lobes subconnate; petals 5, free, the standard suborbicular, wings oblique, obovate to oblong, the keel similar to wings, incurved, free, often overlapping dorsally; stamens 10, free, alternately subequal; anthers versatile. Fruit 1—6 seed- ed, oblong or rarely elongate, compressed or turgid around seeds, dehiscent or rarely indehiscent, woody or fleshy, 2-valved, continuous or with septa between seeds; seeds ellipsoid, globose or lenticular, unicolored, red, yellow, or black, or bicolored, red and black or yellowish and red, the hilum terminal, elliptic or linear; chromosomes: n = 8. Ormosia is a genus of some 100 species, approximately half of which are American tropical rain forest elements and the remainder of the Old World. This Asian-American disjunct distribution is curious, with no single species common to both hemispheres. In Panama, six species are recognized, two of these endem- ic, O. panamensis and O. cruenta, with the other four distributed south to Brazil. While Ormosia isthmensis Standl., described originally from Oaxaca, has been reported from Panama (Rudd, 1965), all specimens annotated as such are referable to other taxa, usually O. macrocalyx Ducke or O. coccinea var. subsimplex (Spruce ex Benth.) Rudd. While lacking major economic importance, most species do yield wood of good quality. In Panama, various species are used locally for building and fur- niture construction. This genus finds greatest importance among native peoples for personal adornment and medicinal preparations. The bright red seeds are used in necklaces, hence the common name "necklace tree." Leaves, bark, and roots are used alone or in combination with other drug plants for a wide variety of ailments and maladies. In eastern Venezuela, Ormosia monosperma ("'pionia montanero’’) is utilized as follows: “the seed used by doctor; cook seed and drink for pains of the heart; also cooked seed placed in water, given to children to put around their necks for sore throat" (collectors notes, Steyermark 61330 [MO]). Seeds of several species have been tested as possible drug sources. The al- kaloids ormosine and ormosonine have been isolated, possibly from O. avilensis or O. venezolana, with the former reportedly having a physiological effect re- sembling morphine (Hess & Merck, 1919). Ormosia macrocalyx, originally iden- tified as O. panamensis, has yielded the alkaloid panamine, which induced hy- potensive action in test dogs (Lloyd & Horning, 1958). Literature: Hess, K. & F. Merck. 1919. Über Ormosin und Ormosinin, zwei neue Alkaloide aus Ormosia dasycarpa. Ber. 52: 1976—1983. Lloyd, H. A. & E. C. Horning. 1958. Alkaloids of Ormosia panamensis Benth. and related species. Amer. Chem. J. 80: 1506-1510. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 739 Rudd, V. E. 1965. The American species of Ormosia (Leguminosae). Contr. . Natl. Herb. 32(5): 279-384. Mature leaflets glabrous beneath (inconspicuously pubescent in O. panamensis), secondary veins not conspicuously ny mature fruits Ei ae glabrous, seeds unicolored red. . Leaflets 2-6(-8) cm long, 1.0— A - cm wide (Panama) 3. O. cruenta bb. Leaflets 6-15 cm lon ie d "M c Fruit submolli a rm, swan tea between seeds, black or dark brown, shiny, glabrous, septa lacking between seeds (Mexico to Brazil) |. 4. O. macrocalyx cc. as it pn icular, not constricted between seeds, yellow sericeous, glabrate eeds separated by septa (Panama) 3: anamensis Mature jeu ani eneath, secondary veins conspicuously raised; mature fruits rre or glabrous, seeds bicolored red and black. d. Mat £e w N> e fruit densely Serie dd not shiny; leaflets reddish pubescent beneath (Pan- a, Colo mbia gee a) 6. O. tovarensis dd. a fruit glabrous: ue white (Panama, northern South America eaflets with lower surface dud pubescent with minute appressed trichom (Panama, northern South America)... 2a. O. coccinea var. prs ee. Leaflets with lower surface runs and tightly crisped-pubescent durs ma, Am- azonian Basin) . O. amazonica 1. Ormosia amazonica Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 139. 1922. TYPE: Brazil, Para: Obidos, Ducke 14833 (G, holotype, not seen; MO, photo). Ormosia euneura Harms, Notizbl. Bot. Gard. Berlin 9: 972. 1926. TYPE: Peru, Loreto, Iquitos, Tess- mann 3665 (fragment, F; photo neg. #1908, MO). Tree to ca. 20 m; young stems greyish to yellowish tomentulose. Leaves 7— 11-foliolate; rachis 14—40 cm long, tomentulose; petiole 7-12 cm long; petiolules 5-10 mm long; leaflets coriaceous, elliptical, ovate, or obovate, 15-22 cm long, 6.0-11.0 cm wide, the apex obtuse, short acuminate, the base rounded to cordate, the margin sometimes revolute, glabrous above, finely and tightly crisped-pubes- cent beneath. /nflorescence terminal, the axes yellowish tomentulose. Flowers purplish, 15-17 mm long; calyx 8-10 mm long, yellowish to reddish tomentulose, 5-lobed, the teeth 4-6 mm long. Fruit dehiscent, subligneous or coriaceous, gla- brate, black to dark brown, 1—3-seeded, 2-5 cm long, 1.5-2.5 cm broad; seeds bicolored, red with a black spot; 10-13 mm long, 9-11 mm wide, 7-8 mm thick. This species was recently collected in Panama on Barro Colorado Island, Canal Zone. Although sterile, the collection is best referred to this taxon until flowering material is available for study. Previously it had been known only from the Amazonian Basin of Brazil and Peru, and its occurrence north of the equator marks a considerable range extension. Inflorescence, flower and fruit characteristics are taken from Rudd (1965). Ormosia amazonica is distinctive among the Panamanian species with its large leaves and leaflets with upper surfaces glabrous and lower surfaces finely and tightly crisp-pubescent. CANAL ZONE: Barro Colorado Island, Foster 2866 (MO). 2. Ormosia coccinea (Aubl.) Jacks., Trans. Linn. Soc. London 10: 360. 1811. 2a. Ormosia coccinea var. subsimplex (Spruce ex Benth.) Rudd, Contr. U.S. Natl. Herb. 32: 328. 1965 740 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Ormosia subsimplex Spruce ex Benth., Mart. Fl. Bras. 15(1): 316. 1862. TYPE: Venezuela, Amazonas, San Carlos, Spruce 2955 (K, holotype, not seen). Tree, to ca. 30 m; trunk commonly 60(—100) cm d.b.h., outer bark thin, hard, reticulate, flaking easily; stems subflexuose, reddish to yellowish tomentulose, glabrescent. Leaves imparipinnate, 9(—14)-foliolate, 25—40 cm long; rachis 8-30 cm long, tomentulose to subsericeous, the petiole 6-10 cm long; leaflets coria- ceous, elliptic, oblong to obovate, 7-18 cm long, 4.0-7.5 cm wide, abruptly acu- minate, obtuse to subcordate basally, glabrous above, somewhat shiny, the lower surface sparsely pubescent with minute, appressed trichomes, somewhat viscid, the costa tomentulose. /nflorescence cymose-paniculate, terminal, to 30 cm long, axes yellowish to reddish tomentulose, glabrescent. Flowers purple, 10-15 mm long; calyx yellowish or reddish tomentulose, 7—9 mm long, 5-lobed, the teeth 3— 5 mm long; corolla 10-15 mm long. Fruit dehiscent, woody, black or dark brown, glabrous at maturity, shiny, usually 1 or 2 seeded, oblong or obovate, 3.5-5.0 cm long, 2.0-2.5 cm wide, abruptly narrowed at the apex, the valves 2-3 mm thick; seeds bicolored, red and black, 10-15 mm long, 9-12 mm wide, 7-10 mm thick. This variety is distributed in tropical moist forest from Panama to Colombia, Peru, Venezuela, and Brazil, with variety coccinea found in northern Brazil and adjacent French Guiana, Surinam and British Guiana. Variety subsimplex is readily recognized among the Panamanian taxa by its large, coriaceous, glabres- cent leaflets, woody 1—2-seeded fruits and bicolored seeds. It flowers from June to August and fruits mature in 4—6 weeks. The fruits and exposed seeds persist until flowering the following year. It is known locally as ''alcornoque"' or "'per- nillo de monte.’ CANAL ZONE: Near Indio Tower, Johnston 1793 (A). San Pablo, collector xo - 267 (US). BARRO COLORADO ISLAND: Croat 5104 (MO, SCZ, US), 5120 (MO, NY, SCZ, US), 6577 (F, MO, NY, SCZ, US), 8112 (MO), 8440 (MO, US), /0395 (MO, NY), 11306 (MO), ts NY. 12702 (MO); Foster 1114 (PMA); Shattuck 1103 (F, MO, US). vERAGUAS: Isla de Coiba, Dwyer 1617 (GH), 2373 (F, MO, NY, PMA). 3. Ormosia cruenta Rudd, Contr. U.S. Natl. Herb. 32(5): 318. 1965. Type: Pan- ama, Davidson 848 (US, holotype; A, F, MO, isotypes). Tree, to 30 m tall; stems reddish tomentose. Leaves imparipinnate, 5-11 fo- liolate, the rachis 8-10 cm long, velutinous, glabrescent; petioles ca. 2 cm long; leaflets subcoriaceous, elliptic to elliptic-oblong, 2-8 cm long, 1.0-3.5 cm wide, the apex acute, obtuse basally, glabrous above, shiny, glabrous below, somewhat shiny. Inflorescence racemose, terminal, axes reddish velutinous. Flowers 15-18 mm long; calyx reddish velutinous, ca. 10 mm long, 5-lobed, the teeth ca. 4 mm long; petals lilac; ovary subsessile, reddish villous, 3—5-ovulate. Fruit dehiscent, sublignous, glabrous, somewhat shiny, black, l-seeded, 2.0-2.5 cm long, 1.5 cm wide, 1 cm thick; seeds red, 8-10 mm long, 9-10 mm wide, 7-9 mm thick, the hilum ca. 2 mm long, ca. 1.5 mm wide. This species is only known from the mountains of western Panama at 800— 1,300 m. It is distinctive among the Panamanian species with its small leaflets and legume A colam (Lindsay 481 [MO]) from the Canal Zone Experimental Garden 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 741 appears to be Ormosia cruenta; however, its leaflets are more acuminate than other collections examined. CHIRIQUÍ: Boquete, Davidson 848 (A, F, MO, US). cocrÉ: Cerro Pajita, Allen 4499 (F, GH, MO, NY, US). 4. Ormosia macrocalyx Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 137. 1922. TYPE: Brazil, Amazonas: Lago Teffe, Ducke 7345 (BM, isotype, not seen). O. toledoana Standley, Carnegie Inst. Wash. 461: 64. 1935. TYPE: British Honduras, Toledo: Forest o s O. chlorocalyx Ducke, Bol. Técn. Inst. Agron. N. 2: 23. 1944. TYPE: Brazil, Amazonas: Esperanca, Rio Solimoes, ‘‘boca do Javari,” Ducke 1516 (isotypes F, MO). Tree to ca. 40 m; young stems finely pubescent, glabrate. Leaves 7—11-fo- liolate; rachis 15—40 cm long; petiole 4-10 cm long, conspicuously swollen at the petiolules pulvinate, 5-8 mm long; leaflets ovate to ovate-oblong, 6-15 cm long, 4-10 cm wide, glabrous, the apex obtuse to short acuminate, rounded to ane ice basally. Inflorescence terminal, paniculate, to 30 cm long, greyish to yellowish pubescent, the bracts linear, 3-10 mm long, ca. 1 mm wide, the brac- teoles subulate, 1.0-1.5 mm long. Flowers purplish, 18-25 mm long; calyx 8-15 mm long, grayish pubescent, 5-lobed, the teeth 3—5 mm long; petals ca. 2 mm long; standard minutely biappendiculate basally. Fruit dehiscent, coriaceous, cir- cular to oblong, flattened, to ca. 10 cm long, ca. 3 cm wide, 1—4-seeded, the lateral margins straight or submolliform, glabrous; seeds red, ovoid, ca. 1 cm long, ca. 10 mm broad, ca. 8 mm thick, the hilum elliptic. Chromosomes: n = 8. This species is distributed from southern Mexico to the Amazon Basin of Brazil, usually in moist forests up to 100 m. In Panama it is known from tropical moist forest in the Canal Zone, Bocas del Toro, and Veraguas, and from pre- montane wet forest in Chiriqui Province. It appears to flower during mid-rainy season, with the fruits persisting most of the year. A chromosome number of n = 8 was reported by Atchison for O. macrocalyx as O. panamensis. S DEL TORO: Cooper & Slater 125 (A, GH, US). CANAL LI Cascadas Road near Summit, Avilla š 407 (MO, SCZ). Barro Colorado Island, Croat 13217 (F, MO, US). Highway near Fort Kobbe, Dwyer 2666 (MO). Curundu, Dwyer & Robyns 6 (MO). Barro Colorado Island, Foster 2091 (GH, PMA, US). Tropical Test Center Site, Chiva Chiva Trail, Gentry 6335, 8751 (both MO). Corozal area, Mori & Kallunki 3063 (MO, US). Corozal cemetery, Schubert & Lindsay 602 (GH, US). Fort Clayton, Tyson 3472 (SCZ). Corozal area, Tyson & Dwyer 2789 (FSU, MO, SCZ). CHIRIQUÍ: Progreso, Cooper & Slater 243 (F, GH, NY, US). PANAMÁ: Alrededores del Hotel Panamá, 8 Mar 1971, Holdridge (PMA). VERAGUAS: Santiago, Dwyer 1347 (MO, SCZ). 5. Ormosia panamensis Benth., Seem. Bot. Voy. Herald 111, fig. 5. 1854. TYPE: Panama, Seemann 1673 (K, holotype, not seen; BM, isotype, not seen; MO, photo).—Fic. 39. O. stipitata Schery, Ann. Missouri Bot. Gard. 30: 90, fig. 2. 1943. TYPE: Panama, White 306 (holotype, MO). Tree to ca. 15 m tall; young stems yellowish or golden sericeous, glabrate. Leaves 5—9-foliolate, the rachis 9-17 cm long, sparsely pubescent, glabrate, the petiole 5-8 cm long; leaflets subcoriaceous, oblong-lanceolate, gradually long 742 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 E39. Ormosia cruenta Rudd.—A. Habit (<'%2).—B. Flower with one wing petal removed (x i. yee Allen 4999, |—C. Fruit (x4). [After Davidson 848. | acuminate at the apex, obtuse basally, 10-15 cm long, 3-6 cm wide, glabrate above, inconspicuously golden puberulent below. Flowers lilac; calyx densely yellowish sericeous, 8-11 mm long, S-lobed, the teeth 3-4 mm long; corolla ca. 2 cm long. Fruit dehiscent, coriaceous, yellowish sericeous, glabrate; light brown or yellowish, suborbicular when mature, flattened, ca. 4 cm in diameter, not constricted between seeds, the valves thick with an adnate margin, 5-15 mm wide; seeds 1-4, separated by septa, ca. 1.5 cm long, dark red. This species is only known from Panama, and it appears rare locally in Bocas del Toro, Chiriqui, and Barro Colorado Island, Canal Zone. Its dehiscent, cori- aceous legume, lacking constrictions between the seeds, is distinctive among the Panamanian species. It is unique among the American species in possessing sep- tae between the seeds, a character also found in some Asiatic species. It is known locally as ‘‘peronil.”’ AS DEL TORO: Talamaca Valley, near Almirante, cee & Slater 150 (US). CANAL ZONE: Barro Colorado Island, Foster 1128 (F, MO, PMA, SCZ). 1 km N of Gam near summit of Cerro boa Pelado, Nee 9060 (MO). cHIRIQUí: Near David on road to Gualaca, Birdsall (US); Holdridge 2 (US); Roy (US). 6. Ormosia tovarensis Pittier, Bol. Soc. Venez. Ci. Nat. 4: 85. 1938. TYPE: Ven- ezuela, Distrito Federal: El Avila, Delgado 59 (VEN, holotype; F, isotype). Tree to ca. 25 m; young stems yellowish to reddish tomentose. Leaves 7—9- foliolate, the rachis 12-36 cm long, reddish tomentose, the petiole 4-15 cm long, reddish tomentose, the petiolules 3-10 mm long, reddish tomentose, the leaflets 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 743 coriaceous, elliptic, oblong-elliptic or obovate, 8-30 cm long, 1-18 cm wide, rounded at each end, the margins often revolute, the upper surface glabrous, densely reddish pubescent with moderately crispate hairs beneath, the secondary veins raised. Inflorescence terminal, the axes reddish tomentose, the bracts del- toid, 5-6 mm long, the bracteoles linear-deltoid, ca. 2 mm long. Flowers 15-16 mm long, violet; calyx reddish tomentulose, 10-11 mm long, 5-lobed, the teeth 4-5 mm long; ovary yellowish pubescent with crispate hairs. Fruit yellowish to reddish velutinous, 1—3-seeded, 4-9 cm long, 3-4 cm wide, ca. 2 cm thick, the valves ligneous; seeds bicolored red and black, 15-22 mm long, 13-19 mm wide, 10-18 mm thick, the hilum 3.0-3.5 mm long. This species was previously known from the rain forests of the coastal cor- dillera of Venezuela and the Cordillera Oriental of Colombia, and its occurrence on Cerro Tacarcuna, Darien, marks its northwestern extreme in distribution. On Cerro Tacarcuna it occurs in elfin forest between 1,800—1,850 m The specimens cited here are sterile and are only tentatively being assigned to O. tovarensis on the basis of leaflet size, shape and pubescence. Inflorescence, flower and fruit characteristics are taken from Rudd (1965). Fruiting material may prove this to be another or perhaps a new species. DARIÉN: Top of W peak of Cerro Tacarcuna, Gentry & Mori 13988 (MO). Headwaters of Ta- carcuna River, Holdridge (US). 35. PACHYRHIZUS W. G. D'Arcy” Pachyrhizus Rich. ex DC., Prodr. 2: 402. 1825. TYPE: P. angulatus Rich. ex C. = P. erosus (L.) Urb., nomen & typus cons. vs. Caraca Thou. Caraca Thou., Dict. Sci. Nat. 6: 35. 1805 [?1806]. Nomen rejic. vs. Pachyrhizus DC. Based on Dolichos bulbosus L. Taeniocarpum Desv., Ann. Sci. Nat. 9: 420. 1826. TYPE: 7. articulatum (Lam.) Desv. = Pachyrhizus b. Robynsia Mart. & Gal., Bull. Acad. Bruxelles 10(2): 193. 1843. non Drapiez 1841, nec Hutch. 1931. TYPE: R. macrophylla Mart. & Gal. = Pachyrhizus erosus (L.) Urb. Twining annual or perennial vines; stems terete; roots tuberous. Leaves pin- nate trifoliolate; leaflets entire, lobed or deeply dissected, stipels subulate, stip- ules caducous, rachis and petioles with longitudinal ridges running onto the sti- pels. Inflorescences axillary pseudoracemes, the peduncle often longer than the petiole, the flowers somewhat congested apically; bracteoles subulate, shorter than and subtending the calyx; pedicels slender, slightly longer than the calyx. Flowers blue; calyx campanulate tubular, 2-lipped, the upper 2 teeth mostly con- nate, the lower teeth acute, shorter than the tube; standard emarginate with 2 basal incurved auricles, the wings auriculate, medially adherent to the keel, the keel equalling the wings; stamens diadelphous, the anthers 10, elliptical, dorsi- fixed; ovary subsessile, multiovulate, the style incurved, sometimes forming a circle, dorsally glabrous, bearded the full length of the ventral, incurved surface. Legume oblong, compressed, nearly straight, internally septate between the seeds; seeds globose, compressed. 74 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 744 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Pachyrhizus is distinguished by its curved, ventrally pubescent style, often lobed leaves, flowers which are not resupinate, and tuberous roots. Although treated here as monotypic, Clausen (1946) adopted a narrow species concept and recognized six species and several varieties. The genus was widely cultivated in pre-Columbian times and ranges through- out the Neotropics, occurring mainly at lower and middle elevations. It has been introduced to the Old World and is naturalized there. The tubers of Pachyrhizus are cultivated for food in many tropical countries and the plants are sometimes cultivated for forage. The seeds are reported to be toxic. 1. Pachyrhizus erosus (L.) Urban, Symb. Ant. 4: 311. 1905.—Fi. 40. Dolichos erosus L., Sp. Pl. 2: 726. 1753. Type: ?Herb. Linnaeus (LINN 900.11 not seen; microfiche MO). D. bulbosus L., Sp. Pl., ed. 2. 1021. 1763. Renaming of D. erosus L. D. articulatus Lam., Encycl. Méth. Bot. 2: 296. 1786. Based on Plumier, Plant. Amer. tab. 220. 1755. Pac E angulatus Rich. ex DC., Prodr. 2: 402. 1825. Renaming of Dolichos bulbosus L. 26. S. VALUTA Spre . Syst. Veg. 3i 52 5, 1826. Based on Dolichos articulatus Lam. Taeniocarpum SUMUS (Lam.) Desv., Ann. Sci. Nat. 9: 421. 1826. Robynsia macrophylla Mart. a Gal., Bull. Acad. Bruxelles 10(2): 193. 1843. LECTOTYPE: Mexico Galeotti 3278 (BR, not s Pa j ic ia articulatus (Lam. ) Duch. & Walp., Flora 36: 226. 1853. P. bulbosus (L.) Kurz, Asiat. Soc. Beng., J. 45(2): 246. 1876. be seen.) Cacara erosa (L.) Kuntze, Rev. Gen. Pl. 1: 165. 1891. Pachyrhizus panamensis Clausen, Cornell Univ. Agric. te Sta. Mem. 264: 21. 1944 [1945]. TYPE: Panama, Killip 12080 (US, holotype; isotypes GH, NY). P. vernalis Clausen, Cornell Univ. Agric. Exp. Sta. s 264: 23. 1944. [1945]. TYPE: Guatemala, Steyermark 38553 (F, not seen). Herbaceous vines; stems pilose, often drying hollow; rootstock tuberous. Leaves pinnate trifoliolate, the leaflets ovate to broadly ovate, the terminal leaflet broadest and often 3-7 lobed, the sinuses and lobes acute or rounded, mucronate, the lateral leaflets oblique, often less lobed or entire, basally deltoid, 3-veined from the base, drying slightly discolorous, glabrate or densely soft tomentose; petiolules 4-5 mm long, stout, pilose; petioles and rachis slender, drying angled, pubescent; stipels 1-2 mm long, acicular, sericeous, basally extended into definite ribs on the rachis; stipules minute, caducous. Inflorescences axillary pseudora- cemes, mostly 12-20 cm long, the flowers congested near the apex, the peduncle mostly longer and stouter than the petioles, the bracteoles resembling the stipels, subtending and much shorter than the calyx; pedicels ca. 2 per node, slender, ca. 5 mm long. Flowers blue, calyx 5-8 mm long, tubular, 2-lipped, the upper 2 teeth mostly connate, the lower 3 acute or acuminate, shorter than the tube, subequal or the lowermost longest and acuminate; standard ca. 15 mm long, suborbicular, emarginate, the wings and the keel about as long as the standard, sometimes ciliolate. Legume oblong, ca. 10 cm long, 1.5-2 cm wide, compressed, appressed strigose. This species is distinct in its broad, often lobed leaflets and in its tuberous roots. In Panama, upland populations are generally densely pubescent while plants from the lowlands are glabrate. Clausen (1947) distinguished the more pubescent plants as Pachyrhizus panamensis, noting that petals of these plants 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 745 FıGURE 40. Pachyrhizus erosus (L.) n.—A. Habit (x!4). I Flower (x2).—C. Corolla.—C!. Standard (x1).—C?. Wing petal ey Ce. Keel petal (x1).—D. Stamens (x2).—E. Pistil (x2). [After Allen 2027.]—F. Fruit (x12).—G. Seed (x14). [After ssl 5053.) are usually ciliolate. However, these differences are not of specific import. B coincidence, the specimen he selected for the type of his species is an unusually pubescent species from Ancon Hill in the lowlands, thus the distinction between upland and lowland races is not absolute. Pachyrhizus erosus ranges throughout the Neotropics and has been introduced and naturalized in parts of the Old World. It is cultivated for its edible tubers, but many of the Panamanian collections appear to have been taken from the wild. 746 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 The seeds are toxic. The species is known in English as the Yam Bean, and in Spanish as Jícama. BOCAS DEL TORO: Big Bight, Chiriqui Lagoon, Wedel 2981 (GH, MO, US). CANAL ZONE: Fort Kobbe, Allen 2027 (F, GH, MO, NY, US). Ancon, Celestine 28 (US). Fort Sherman, Dwyer & Robyns 132 (MO). Chagres, Fendler 74 (MO), 75 (F, GH, US). Balboa Heights, Greenman & Green- man 5053 (MO). Cerro Ancon, Heriberto 117 (GH, US). Ancon Hill, 100-200 m, Killip 12080 (GH, US). Madden Dam, 50 ft, Lewis et al. 29 (MO). Junction Chagres Boy Scout Road and Madden Dam Road, Mori et al. 4070 (MO, WIS). Madden Dam, Nee 7774 (MO). Cerro Ancon, Standley 25200 (US). Sosa Hill, Balboa, Standley 25271 (US). Gatun, Standley 27281 (US). Mount Hope Cemetery, epo: 28798, 28803 (both US). Between France Field and Catival, Standley 30213 (US). Curundu, Tyson & Blum 2534 (FSU, MO). CHIRIQUI: Quebrada del Medio 2 km N of Punta de Burica, 40—80 m, Ds 747 (MO). Quebrada Guanabanito 2 mi SW of Puerto Armuelles, Croat 2207] (DUKE, F, MO). Burica Peninsula above Quebrada Yerbazales, 200 m, Croat 22586 (MO). Bajo Mono, Bo- quete District, Davidson 496 (US). Boquete, Lewis et al. 618 (MO); bid 2918 (GH, NY, US). COLÓN: Río Guanche, D'Arcy 9721 (MO). Los sANTOS: 16 mi S of Macaracas at Quebrada Bejuco, TAREA et al. 3091 (MO, SCZ). PANAMA: Isla de Pedro Gonzales, ies 1720 (MO). SE side of n Lake near Puente Natural, 90 m Nee & Hansen 14046 (MO, WIS). Ruins of Old Panama, Pio 22878 (GH). SAN BLAS: Mainland opposite Achituppu, Lewis et al. 102 (GH, MO, US). 36. PHASEOLUS James A. Lackey” & W. G. D'Arcy?* Phaseolus L., Sp. Pl. 723; Gen. Pl., ed. 5. 323. 1754. TYPE: P. vulgaris L. Vines, herbs or subshrubs. Leaves pinnate trifoliolate, the leaflets mostly entire, pubescent or glabrate but minute, uncinate hairs present, the stipels often oblong, thin, glabrous; stipules acute, nervate, often pubescent, not prolonged below the insertion. /nflorescences axillary, the flowers congested in fascicles along the rachis, the nodes not swollen, not glandular, the rachis eglandular; bracteoles ovate or greatly reduced and much shorter than the calyx, nervate, puberulent, persistent at least until anthesis; bracts ovate or lanceolate, nervate, puberulent; pedicels mostly longer than the calyx. Flowers blue, purple, violet, yellow or white; calyx mostly 2-lipped, the upper teeth partly united; standard symmetrical, orbicular, basally appendaged, the wings partly spiralled, apically cucullate, the keel in 2—3 spirals; vexillary stamen free, the free part of the others elongate, the anthers nearly uniform; ovary l-many ovulate, the style apically thickened, curved in 1.5-2.5 spirals, hairy inside, distally caducous. Legume linear or oblong, straight or slightly curved, not septate, compressed or turgid, sometimes beaked; seeds I-many, oblong to reniform, the hilum short and cen- tral. Phaseolus includes about 50 species, all of the New World, although under older circumscriptions, the genus may embrace 100—300 species. In Panama, Phaseolus is a genus of both uplands and of low elevations. Various members of the genus are widely cultivated for food in both temperate and tropical countries. a. Bracts conspicuous, foliaceous, more than 5 mm long; inflorescences dense, many flow- ered; pods pubescent ? Department of Botany, U.S. Natural History Museum, Smithsonian Institution, Washington, D.C. 20560. 7% Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 747 calyx tooth conspicuously more tomentose than the others; pod more than 6 cm long; mm e Ss S mor UP bb. Bracteoles minute, shorter than the calyx; flowers red to purple (white), 11- 15 mm long; n est calyx tooth only slightly pubescent, ars more so than the others; pod less than 6 cm long; stipels less than 4 mm long; native species ______ puerum aa. Bracts inconspicuous, minute, less than 5 mm long; eri aeei open, few flowe red: pods mostly glabra c. Pod straight, narrow, mostly 5-10 seeded; peduncle less than 1 cm long; flowers more than 10 mm long, the standard glabrate outside with a few hairs on the midvein; stipels more Wed i l cc. Pod curved, broad, mostly 2—4 seeded; peduncle more than 2 cm long; flowers les sai 10 mm long, the standard ed pubescent outside; stipels less than 1.5 m 2, P. [AE 1. Phaseolus coccineus L., Sp. Pl. 724. 1753. Type: Herb. Linnaeus (LINN 899.2, not seen; microfiche MO). P. formosus H.B . Gen. Sp. Pl. 6: 449. 1823. TYPE: near Toluca, Mexico, not seen. P. obvallatus Schlecht. pies 12: 328. 1838. TYPE: Mexico, Ehrenberg not se en). Phaseolus coccineus ssp. polyanthus (Greenm.) Maréchal et al., Taxon 27: 199. 1978. Herbaceous vine, often large, stems softly hirsute, somewhat glabrescent, becoming stout, ca. 2-3 mm thick. Leaves pinnate trifoliolate, leaflets broadly ovate, the lateral pair strongly oblique, mostly 6-10 cm long, 6-8 cm wide, api- cally acute or acuminate, basally truncate, subcordate or rounded, the venation conspicuous on drying, sparingly soft pubescent with long and short hairs; pet- iolules thick, 3-5 mm long, petioles mostly longer than the terminal leaflet, stipels conspicuous, 4-6 mm long, cucullate, striate and pubescent outside, pubescent basally within. Inflorescences elongate cymose pseudoracemes to 30 cm long, sparingly flowered proximally, somewhat congested apically, the rachis drying sharply angled and tomentulose, especially upwards, bracts subfoliaceous, con- spicuous, to 1 cm long, linear to broadly ovate, often acute, sometimes acuminate; bracteoles 2, ovate, subtending and slightly exceeding the calyx; pedicels mostly straight, 15218 mm long, much longer than the calyx, becoming somewhat woody in fruit. Flowers red to lavender, showy; calyx cupular, 10-13 mm long, glabrate except the lower tooth, the upper teeth united forming a hood, the lower teeth acute, the lowermost longest; standard, ca. 2 cm long. Legume oblong, somewhat falcate, ca. 10 cm long, 15 mm wide, compressed or slightly turgid, beaked, softly short pubescent; seeds 5-8. This species is distinguished by its elongate pedicels, its paired ovate brac- teoles, by its large, conspicuous stipels and by the ciliate to tomentose lowest calyx tooth. This species is usually recorded as having scarlet flowers, but in Panama collectors more often note lavender or magenta flowers. In Guatemala this species is much cultivated for food, and in the United States it is cultivated for its showy flowers as ‘Scarlet Runner Bean. CHIRIQUÍ: Central valley of Río Chiriquí Viejo, 1800-2000 m, Allen 1397 (MO). Nueva Suissa, Croat 13508 (F, MO, NY, PMA). Bajo Mono, Boquete District, 4500 ft, Davidson 496 (MO). Río 748 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 ce "e m — — E4l. Phaseolus lunatus T m Habit (x3⁄s). [After McDaniel & Tyson 12690.]—B. Fruit pa. After Burt & Koster 3 Oct. 1.] Chiriqui Viejo N of Volcan City, ird x ene 9019 (MO, NY). Nueva Suissa near Audubon Society Cabin, Gentry 5986 (MO). Cerro P 000 ft, Tyson 7097 (PMA). 2 km W of La Garita, 3 km WNW of Cerro Punta, 2000 m, Wilbur e et 2 15292 (DUKE). Finca Lérida to Pena Blanca, 1750-2000 m, Woodson & Schery 313 (MO). Casita Alta, Volcán de Chiriquí, 1500-2000 m, Woodson et al. 918 (MO, NY, US). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 749 2. Phaseolus lunatus L.,” Sp. Pl. 724. 1753. TYPE: not determined.—Fic. 41. Erect, ascending or nearly prostrate slender vine, sometimes somewhat scan- dent, sometimes branched; stems pilose with weak hairs, glabrescent. Leaves pinnate trifoliolate, leaflets ovate to deltoidal, sometimes rhombic ovate, apically deltoid, blunt but mucronulate, basally broader, sometimes rounded, mostly 6— 8 cm long, 4—6 cm wide, 3 veined from the base, the lateral veins distally arcuate, somewhat evanescent, mostly drying concolorous, the sparse minute pubescence mostly confined to the costa; petiolules 2-3 mm long, puberulent; rachis mostly 1.5-3 cm long; stipels minute but evident, persistent, mostly 0.5-1.0 mm long; petioles mostly longer than the terminal leaflet, glabrate or sparingly pubescent with weak, subappressed hairs at anthesis. /nflorescences axillary or lateral, short or elongate pseudoracemes to 25 cm long, the nodes often well spaced proximally, more condensed distally, the bracts mostly 2-3 cm long, acute, sericeous, incon- spicuous, the rachis and the peduncle sparingly pilose, glabrescent; pedicels 1— 4 per node, slender, 6-9 mm long; bracteoles mostly ca. 1 mm long, pubescent or not, 3-nervate, spathaceous, subtending the calyx less than !⁄2 as long as the calyx. Flowers greenish or purplish, the calyx 2-3 mm long, the tube campanu- late, the limb somewhat flaring, the lobes rounded or short deltoid, ciliolate; standard ca. 1 cm long, broad, flat, often puberulent externally, keel spiralled. Legume flat, slightly turgid around the seeds, falcate, 3-8 cm long, 1.5-2 cm broad, glabrate; seeds 2—4, reniform, compressed. This species is distinct in its glabrate leaflets and pods, inconspicuous bracts and bracteoles, and short calyx teeth. It is represented by both cultivated and wild varieties which differ in seed size and, according to Marechal et al. (1978), in the wild forms having glabrous standards. The species is believed to be native to Central America and perhaps the Antilles, so some of the populations collected in Panama are probably native to the country. This species is commonly referred to as the Lima bean (English) or as Habas (Spanish). ZONE: Isla Perico near Fort Amador causeway, McDaniel & Tyson 12690 (MO). Sosa Hill, Standley ripe (US). Balboa, Standley 25647, 30894 2nd US). CHIRIQUÍ: E rada Guana- banito beyond La Represa 2 mi o erto Armuelles "ens m, Croat 22076 (MO). Boquete to 3 mi N, 3300-4200 ft, Lewis p Kd 318 (DUKE. F, MO, in A). HERRERA: NO ar Los Pozos, 20 m, Burt & Koster, 10 March 1971 (MO). PANAMÁ: Bella ER joe 25304 (US). Las Sabanas, Standley 25932 (US). Tumba Mudo Road near Panamá, Standley 29792 (US). Camino a Sardinilla, Taylor 38 (PMA). vERAGUAS: Atalaya-Santiago, 100-200 m, Koster 117 (MO). 3. Phaseolus tuerckheimii Donnell Smith, Bot. Gaz. 56: 54. 1913. TYPE: Guate- mala, Tuerckheim 1536, not seen. Paratype: Costa Rica, Pittier 10539 (US). P. chiriquinus Standley, Contr. U.S. Natl. Herb. 18: 109. 1916. rvPE: Panama, Pittier 3111 (US, holotype; isotype NY). Herbaceous vine, sometimes high climbing; stems stout, sericeous, becoming softly short tomentulose, drying angled, to 3 mm thick. Leaves pinnate trifolio- late; leaflets ovate, sometimes narrow, mostly 4—7 cm long, apically acute or acuminate, basally rounded, drying somewhat discolorous, softly densely pubes- 77 For a list of other names see Maréchal et al., 1978. Only this name has been used for Pana- manian plants. 750 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 cent above and beneath, the veins elevated beneath, the minor venation some- times drying reticulate; petiolules ca. 3 mm long, tomentose; petioles mostly shorter than the terminal leaflets, densely puberulent; stipels conspicuous, 2—3 mm long. Inflorescences mostly elongate pseudoracemes to 30 cm long, the basal portion devoid of flowers, the peduncle basally stout, narrowing and becoming more densely pubescent upwards; bracts subfoliaceous, linear to broadly ovate, to 7 mm long, sericeous; bracteoles linear, glabrate, caducous, ca. 1 mm long; pedicels slender, ca. 10 mm long with scattered long hairs. Flowers white to bright violet or purplish; calyx sericeous overall, ca. 4 mm long, the tube ca. 3 mm long drying with umbonate or wing-like longitudinal enations, the limb flaring somewhat, the teeth deltoid, ca. 3 mm long, the uppermost longest, acute; stan- dard 1.5 cm long, the keel convolute, spiralled. Legumes oblong, compressed, slightly turgid around the seeds, ca. 4 cm long, densely tawny hirsute; seeds ca. 4 This species is distinguished by its dense overall pubescence and by its subfo- liaceous bracts and minute bracteoles. The fluting on the dried calyx is an inter- esting feature. Phaseolus tuerckheimii ranges from southern Mexico to Panama. In Panama it occurs only at upper elevations in Chiriqui Province where it is apparently shade tolerant and is found climbing on trees. CHIRIQUI: Volcan de Chiriqui, Boquete District, 7500 ft, Davidson 948 (MO). Rio Chiriqui Viejo N of Volcan City, 5200-5600 ft, Duke 9064 (MO). Cuesta de Cerro Quemado, E slope of Chiriquí Volcano, 1800-2160 m, Pittier 3111 (NY). Cerro Punta, 6000 ft, Tyson 7099 (PMA). 4. Phaseolus vulgaris L.,”* Sp. Pl. 723. 1753. TYPE: ?India, Herb. Linnaeus (LINN 899.1, not seen; microfiche MO). Climbing or trailing vine or erect herb; stems glabrous or pubescent. Leaves pinnate trifoliolate, the leaflets mostly ovate, sometimes broad, the lateral leaflets mostly oblique; glabrate to velutinous, apically acute, or acuminate, the petiolules —4 mm long, stout, pubescent; petioles often exceeding the terminal leaflet; stipels linear, costate, glabrous, as long as or shorter than the petiolules; stipules lanceolate, striate, glabrous. Inflorescences axillary, mostly 3-10 cm long; pe- duncle mostly slender and weak; rachis contracted; bracteoles and bracts spa- thaceous, exceeding the calyx, striate; pedicels slender. Flowers white, yellowish or purplish; calyx campanulate, the teeth rounded; standard 1-2 cm long. Legume linear, turgid, often slightly curved, glabrate, mostly 10-20 cm long; seeds sub- globose to oblong, to 1.5 cm long, variously colored. Phaseolus vulgaris, the well-known string bean or kidney bean in English, the frijol or the frijol negro in Spanish, is cultivated in almost every country in the world, and in some places it escapes and has naturalized. It is a native of the New World, and it was widely disseminated by pre-Colombian man. No collec- tions have been seen from Panama, but the species is grown as a crop plant, mainly in upland areas, to judge from reports of vendors in markets and from fresh vegetables served in restaurants in Panama City from time to time. There 7" No attempt has been made here to research synonyms for this widespread cultivated species. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 751 is also a substantial import of frijoles negros from the United States and other countries for use as foo This species is distinct from other species of Phaseolus in Panama in its long turgid pods and light colored flowers. Many races of Phaseolus vulgaris exist in cultivation, many of quite different appearance, and until specimens are obtained of the Panama elements it is not fruitful to try to amplify details of the above description or to indicate more features of difference. The above description was made from a range of cultivated material from Central America held at MO 37. PUERARIA W. G. D'Arcy” Pueraria DC., Ann. Sci. Nat. 4: 97. 1824. LECTOTYPE: P. tuberosa (Willd.) DC. Neustanthus Benth., in Miq., Pl. Jungh. 234. 1852. TYPE: N. javanicus Benth. Glycine L., Sp. Pl. 753; Gen. Pl., ed. 5. 334. TYPE: G. javanica L. Nomen rejic. versus Glycine Willd. 1802. Robust scrambling vines, sometimes high climbing, ?shrubs; stems herba- ceous or woody, pubescent. Leaves pinnate trifoliolate, the leaflets large, entire, lobed or undulate margined; stipels mostly present, the stipules ovate to linear, often produced below the enlarged point of insertion, the lower element some- times bifid. Inflorescences elongate stout racemes, sometimes branched, the flow- ers somewhat clustered at the end of the rachis; bracts often caducous, bracteoles ovate; pedicels inserted 1—7 at a node. Flowers sometimes appearing before the leaves; calyx campanulate, 5-toothed, the upper pair free or fused into a lip; corolla mostly bluish, pinkish or violet; standard broad, entire or emarginate, minutely spurred, the wings spurred; stamens 10, the vexillary stamen free or united to the others, usually at the middle of the filament, the anthers all alike; ovary pubescent, the style curved or abruptly bent, glabrous, the stigma glabrous. Legume oblong, compressed, usually pilose. This genus of about 20 species is native to southern and eastern Asia. Two species are adventive or naturalized in the New World. Literature: Verdcourt, B. 1968. The identities of Dolichos trilobus L. and Dolichos trilo- batus L. Taxon 17: 170-173 1. Pueraria lobata (Willd.) Ohwi,* Bull. Tokyo Sci. Mus. 18: 16. 1947, not seen.—FiG. 42 Dolichos lobatus ety Sp. Pl. 3: 1047. 1803. Type: Dolichos trilobus Houttuyn, Nat. Hist. ed. Panzer. 8: 560. . 64, fig. 1. 1782, fide Verdcourt 1968. Pachyrhizus NI Ender s se Sieb. & Zucc., Abh. Math.-Phys. Cl. Kónigl. Bayer. Akad. Wiss. 4: 237. 1846. Pueraria thunbergiana (Sieb. & Zucc.) Benth., J. Linn. Soc. Bot. 9: 122. 1865. 79 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 80 For other synonyms see Verdcourt, 1968. Only these names relate to Panamanian plants. 752 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 C? j 1 ! N C 2 c! "P E 42. Pueraria lobata (Willd.) Ohwi.—A. Habit (x). dou pie 2482.]—B. Flowe r T C Corolla.—C'. Standard ( x ).—C?. Wing petal (x 1).—C*. Keel petal (x 1).—D. Stamens (x 11⁄4).—E. Pistil (x 1/2). [After Wedel 2605. ]—F. Fruit (x1). [After Henry 953.] Robust vines clambering on the ground and on shrubs, occasionally high climbing; stems terete, branching, puberulent with appressed short hairs. Leaves pinnate trifoliolate, the leaflets often large and somewhat felty, broadly ovate often 3-lobate, apically short acuminate, the sinuses rounded to obtuse, mucron- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 753 ulate, basally rounded, truncate or subcordate, 3 nerved from the base and pin- nate veined above, discolorous, appressed pubescent on both sides, often with coarse hairs, often ca. 15 cm long and 15 cm wide, the lateral leaflets strongly oblique; petiolules 4-7 mm long, often stout, tomentose; rachis 2-5 cm long; petioles shorter than the terminal leaflet; stipels linear, pubescent, 4-7 mm long; stipules oblong or ovate, puberulent, 4-8 mm long and produced 3-5 mm below the point of insertion, the insertion point ovate. Inflorescences often few, to 30 cm long, the peduncle stout, pubescent, sometimes with 1 or 2 empty stipule-like bracts along its length; rachis slightly thinner, often more than twice as long as the peduncle; pedicels 3-6 mm long, pubescent, inserted mostly in 3's at the nodes; bracts ovate and stipule-like, caducous; bracteoles ovate, ciliate and pu- bescent, 3-5 mm long. Flowers pink or violet, ca. 2 cm long, the calyx campan- ulate, puberulent outside, the cup ca. 5 mm long, the lobes obtuse, unequal, the upper pair mostly or completely fused into a lip, the lowest tooth narrow and usually longest; corolla with the standard broad, emarginate, glabrous, slightly shorter than the wings and the keel, with 2 minute basal auricles, the wings oblong, medially coherent; stamens 10, the vexillary stamen free at the base, perhaps fused to the filament column part way up, the anthers all alike; ovary pubescent, the style and the stigma glabrous, the style bent abruptly upwards. Legume oblong, 4—6 cm long, 4-7 mm wide, long beaked, flat and compressed slightly between the seeds, drying dark, copiously pilose with long, 2-3 mm, yellowish hairs. This species is native to southern and eastern Asia. It has been widely used in other warm parts of the world for forage and erosion control, and it is natu- ralized in many places. In Panama it is very abundant but has been collected only from islands in the Chiriqui Lagoon area of Bocas del Toro and from a few other lowland localities. Pueraria lobata is an aggressive vine which can quickly cover all vegetation and soil over several hectares. In spite of its vigorous vegetative growth it is sometimes seldom lae. and fruits are not always formed. It is known by the Japanese name ''Kudzu BOCAS DEL TORO: Isla Colón, Wedel 136 (MO). Without other locality, Wedel 374 (MO), 511 uy MO), 2482 (MO). Colombus Island, Wedel 2605 (GH, MO), Isla Colón, Wedel 2803 (GH, MO, Y). 38. RHYNCHOSIA Muriel E. Poston*! Rhynchosia Loureiro,? Fl. Cochinch. 425, 460. 1790. nom. conserv. TYPE: K. volubilis Lour. Leycephyllum Piper, J. Wash. Acad. Sci. 14: 363. 1924. TYPE: L. micranthum Piper. Herbs or subshrubs, twining or trailing, erect to ascending; stems terete to angular and striate. Leaves trifoliolate, ovate to rhomboid, gland dotted above *! Department of Biology, Howard University, Washington, D.C. 82 For a list of synonyms see Grear, 1978. Only these names Es "od used for Panamanian plants. 754 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 and beneath; stipules caducous, striate, free; stipels reduced when present; pet- ioles 1.2-8.0 cm long, villous; petiolules 0.4-3.0 cm long, terminal petiolules longer than the laterals. Inflorescence racemose, axillary; bracts caducous or persistent, small, striate. Flowers with the calyx campanulate, the upper teeth at least partly connate; corolla papilionaceous, usually yellow with maroon mark- ings, the standard obovate, auriculate with a short claw, the wings oblong, not auriculate, the claw narrow, the keel falcate, wider than the wings, the claw attenuate; stamens diadelphous, the vexillary stamen free, the anthers mono- morphic; the style slender, the stigma entire, capitate, the ovary sessile, biovu- late. Fruits ovate to oblong, 2-valved, pubescent, gland-dotted; seeds 2 ovate, black often with some red, hilum lateral, ovoid to elongate. Rhynchosia is a genus of several hundred species found in both the Old World and New World tropics. The genus has its greatest diversity in Africa. Literature: Grear, J. W. 1979. A revision of the New World species of ee (Le- guminosae-Faboideae). Mem. New York Bot. Gard. 31(1): 1 a. Calyx lobes 2-3 times longer than calyx tube; seeds never red. Stipules aa to ovate, 0.3-1.0 cm long, 0.1—0.5 cm wide, terminal 1 .R 2.0-11.0 mm long... aaa a a a. . retic ulata bb. Stipules linear " lanceolate, 1.5-3.0 mm long, ca. 0.5 mm wide; porary petiolu les A cm long echt U lu SSS cece R. calycosa aa. . lobes equal to or not more than 2 times longer than calyx tube; seeds red and blac Seeds mostly less than 4 mm E ET Ate ETA Tee EEMS . R. minima cc. Seeds mostly more than 4 m Plants viscid; stipules a hilum Sin IP ss 2. R. edulis dd. DAE not viscid; stipules caducous; hilum tipels absent; pods deeply Padi mna a seeds, glabrous to densely puberulous, shiny black with age -2222222222222222 phe otal Stipels present; pods not iecur constricted, densely puberulous, green o brown, never shiny black. f. naa lanceolate to ovate, 2.0-5.0 mm long; d ca. 5.0 m RON Suyu NENNT n oO oO l. mpya calycosa Hemsley, Diagn. Pl. Mex. 48. 1880. TYPE: Panama, Fen- r 72 (MO, isotype). Dolicholus calycosus (Hemsley) Standley, Contr. U.S. Natl. Herb. 18: 108. 1916. Herbaceous to suffruticose vine; stems weakly angled to terete, strigose to puberulous. Leaves trifoliolate; leaflets ovate to deltoid or ovate rhomboid, 2.5- 7.0(-9.0) cm long, 1.5-5.5 cm wide, apically acute, basally cuneate to obtuse, strigose above, villous and gland dotted beneath; terminal petiolule 1.5-2.0 cm long, the lateral petiolules 0.2-0.3 cm long; petiole 2.5—4.2 cm long, villous; stipels subulate, 1.0-2.0 mm long, the stipules linear to lanceolate, 1.5-3.0 mm long, caducous. Inflorescence exceeding the leaves, 7.0-20.0 cm long; peduncles 1.0— 6.5 cm long; pedicels 1.0-2.0 mm long; bracts lanceolate, 3.0-4.0 cm long, ca- 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 755 ducous. Flowers with the calyx equalling or exceeding the corolla, 7.0-11.0 mm long, the lobes 7.0-9.0 mm long, ca. 2.0 mm wide, upper lobes united halfway up; corolla yellow, 6.0—10.0 mm long, the standard obovate, auriculate, 7.0—10.0 mm long, 3.0-5.0 mm wide, the claw ca. 1.0 mm long, the wing oblong, 6.0-8.0 mm long, ca. 2.0 mm wide, the claw ca. 2.0 mm long, the keel falcate, 6.5-7.5 mm long, 1.5-2.0 mm wide, the claw ca. 1.5 mm long; stamens ca. 7.0 mm long, vexillary stamen free. Fruits ovate to oblong, 1.5-2.5 cm long, 0.5-0.8 cm wide, puberulous to strigose, gland dotted, the beak ca. 2.0 mm long; seeds 2, subor- biculate, ca. 3.5 mm long, 3.0 mm wide, black brown mottled, hilum ovate, 1.0- 1.5 mm long. This species ranges from Central America to Ecuador. In Panama it is found in secondary growth and dry forested areas. CANAL ZONE: Road C-21 near Police wipe Croat 12993 (MO, SCZ). Quebrada La Palma and Canon of Río Chagres, Dodge & Allen 17342 (GH, MO). Las Cruces Trail, Hunter & Allen 761 (GH, MO). cHIRIQUI: Baja Gualaca, Koster 172 Mo. cocLÉ: Río Hato airstrip, Blum & Dwyer 2479 SAN e Koster 121 (MO). PANAMÁ: Hills between Capira and Potrero, Dodge & Hunter 8605 (GH, MO). San Jose Island, Johnston 1106 (GH, MO). vERAGUASs: El Embalsadero, 8 mi W of Santiago, Tyson 6080 (FSU, MO, SCZ). 2. Rhynchosia edulis Griseb., Abh. Kónigl. Ges. Wiss. Góttingen 19: 123. 1874. TYPE: Paraguay, Balansa 1553 (not seen; photo, US) Dolicholus ixodes Standley, Contr. U.S. Natl. Herb. 18: 107. 1916. TYPE: Panama, Williams 119 o R^ uada (Standley) Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 214. 1929. Vine, herbaceous to suffrutescent, stems angular to terete, pubescent. Leaves trifoliolate, ovate to ovate rhomboid, 1.0-3.5 cm long, 1.0-2.3 cm wide, apically acute, basally obtuse, strigulose above, hirsutulose, gland dotted beneath, viscid on both surfaces; terminal petiolule ca. 1.0 cm long, the lateral petiolules obsolete or to 1.0 mm long; petiole ca. 1.3 cm long, viscid; stipels caducous; stipules lanceolate ca. 0.5 cm long. Inflorescence equal to or exceeding the leaves, ca. 2.0 cm long; peduncle ca. 1.0 cm long; bracts lanceolate 2.0-3.0 mm long, ca- ducous; pedicels ca. 0.4 cm long. Flowers with the calyx not exceeding the corolla, viscid, puberulent, ca. 0.4 cm long, the lobes 2.0-3.0 mm long, the upper lobes partially united; corolla purplish, to 6.5 mm long, the standard obovate, auriculate, gland dotted, ca. 5.0 mm long, ca. 3.5 mm wide, the claw ca. 1.0 mm long, the wings oblong, ca. 4.5 mm long, ca. 1.5 mm wide, the claw ca. 1.5 mm long, the keel falcate, ca. 4.5 mm long, ca. 2.0 mm wide, the claw ca. 2.0 mm long; stamens 4.5 mm long. Fruits oblong ovate, strigose, gland dotted, viscid, ca. 1.5 cm long, 0.6 cm wide, the beak ca. 3.0 mm long; seeds ovoid, ca. 4.0 mm long, hilum linear, 1.0-2.0 mm long. This species is known from southeastern Arizona through to South America. In Panama it is found in roadside savannas. PANAMÁ: Between Río Pacora and Chepo; Porter et al. 5136 (MO). 756 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 3. Rhynchosia erythinoides Schlecht. & Cham., Linnaea 5: 587. 1830. TYPE: Mex- ico, inter Misantlem et Nantlam in sylvis. Schiede & Deppe not seen.—Fic. 43 Leycephyllum micranthum Piper, J. Wash. Acad. Sci. 14: 363. 1924. type: Costa Rica: Las Vueltes, Turrique, Tonduz 12951 (isotype, US). Suffrutescent to woody vine, the stems angled, becoming terete, strigulose to puberulose. Leaves trifoliate, the leaflets ovate, ovate rhombic to lanceolate, 3.0— 10.0 cm long, 2.5-8.0 cm wide, apically acute to acuminate, basally rounded to cuneate, sparsely villous, gland dotted below, glabrous to sparsely villosulose above; petiolule of terminal leaflet 1.5-3.0 cm long, petiolule of lateral leaflet 0.2- 0.4 cm long; petiole 2.5-8.0 cm long, villosulose; stipels absent; stipules lanceo- late, 2.0-3.0 mm long, 0.5-1.0 mm wide, caducous. Inflorescence branching, equal to or exceeding the leaves, 5.0-13.5 cm long; peduncles 0.6-1.0 cm long; pedicel 1.0-2.0 mm long; bracts lanceolate, 1.5-3.0 mm long, 0.5-2.0 mm wide, caducous. Flowers with the calyx not exceeding the corolla, 3.0-4.0 mm long, the lobes of the calyx lanceolate, 1.0-2.0 mm long, villosulose, gland dotted; corolla yellow with maroon striations, 5.0-7.0 mm long, the standard obovate, auriculate, 5.0-7.0 mm long, 4.5-6.0 mm wide, the claw ca. 1.0 mm long, the wings oblong, 5.0-7.0 mm long, 1.5-2.5 mm wide, the claw ca. 1.0 mm long, villosulose, the keel falcate, 5.5-7.5 mm long, 2.0-3.0 mm wide, the claw 1.5-2.0 mm long; stamens 5.0-7.0 mm long, vexillary stamen free. Fruits ovate, deeply constricted in the middle, 1.5-2.0 cm long, 0.5-1.0 cm wide, brown to black, hirsutulose, gland dotted, the beak ca. 1.0 mm long; seeds 2, ovoid, ca. 4.0 mm long, ca. 3.0 mm wide, black with a red band around, the hilum ovate, ca. 3.0 mm long. This species ranges from eastern Mexico to Ecuador, occurring in thickets and in clearings and secondary growth. BOCAS DEL TORO: Old Bank d Chiriqui ehe. Wedel 1982 (GH, MO, NY, US), 2084 - CANAL ZONE: BARRO COLORADO ISLAND: ey & Croat 242 (MO); A e 28 August 1971, 10201, 10331, 10928, Pria (al MO), 13817 (F, Mo. NY); Foster 2173 (F, US); Shattuck A (F, p 750 (MO, US), 8/2 (F, MO); Wilson 1217 (F, MO); Woodworth & Vestal 414 (F, MO), 5 (A, F, MO). Chagres, s 66 (F, MO, US). Near Gatún, Hayes 911 (NY). W of Limon E Gatún Locks and Gatún Lake, Johnston 17211 (A, MO). Hills around Agua Clara Reservoir near US). cH [ imi order, Croat 22215 (MO). COLON: Santa Rita Ridge, 5 mi from highway, vun 13188 (MO, NY). María Chiquita, Dwyer 2992 (MO, US). Santa Rita East Ridge, Dwyer 8411 (MO). dps as Cerro Jefe, Duke & Dwyer 15069 (MO). Bismark above Penonomé, Wie p (NY, 4. Rhynchosia minima (L.) DC., Prodr. 2: 385. 1825. Dolichos minimus L., Sp. Pl. 726. 1752. LECTOTYPE: Jamaica, St. Jago de La Vega. Sloane, not seen (selected by Verdcourt, 1971). Herbaceous to suffrutescent vine; stems angular, puberulent to villosulose. Leaves trifoliolate, the leaflets ovate to ovate rhomboid to orbicular, 2.0-3.8 cm long, 1.0-3.0 cm wide, apically acute, basally cuneate to truncate, glabrous to finely strigulose above, puberulose to villosulose, gland dotted beneath: petiolule 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 757 e Sn ng FIGURE 43. Rhynchosia erythrinoides Schlecht. & Cham.—A. Habit (x!2). [After Davidson 927.]—B. Flower (x5).—C. Corolla.—C'!. Standard (x21). —C?, Wing petal (x212).—C?. Keel petal (x214).—D. Stamens (x5).—E. Anthers.—E!. Abaxial view.—E?. Lateral view.—F. Pistil (x5). [Af- ter Wedel 2084.]—G. Fruit (x34).—H. Seed (x2). [After Davidson 927.] 0.3-1.0 cm long, the terminal leaflet with longer petiolule; petiole 1.2—4.0 cm long; strigose; stipels subulate, ca. 1.0 mm long; stipules linear to lanceolate, 3.0— 10.0 mm long, 0.5-1.0 mm wide, persistent, villosulose. Inflorescence equal- ling or exceeding the leaves, 6.8-13.5 cm long, many flowered, the peduncle 1.2- 7.0 cm long, the pedicels ca. 2.0 mm long; bracts lanceolate, ca. 1.0 mm long, caducous. Flowers with the calyx not exceeding the corolla, 2.0—4.0 mm long, the lobes 1.0-3.0 mm long, lanceolate, the upper lobes united part way, puber- ulose, gland dotted; corolla yellow with purplish striations, puberulose, gland 758 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 dotted, 5.0-7.0 mm long, 3.0-5.0 mm wide, the claw 1.0 mm long, the wings oblong, 4.0-5.0 mm long, 1.75-2.0 mm wide, the claw 1.0 mm long, the keel falcate, 5.0-6.0 mm long, 1.5-2.0 mm wide, the claw 1.0-2.0 mm long; stamens .0-7.0 mm long. Fruits oblong, ovate, strigose, gland dotted, 0.9-1.5 cm long, 0.3-0.4 cm wide, the beak 0.5-1.0 mm long; seeds ovoid, brown, black mottled ca. 3.0 mm long, ca. 2.0 mm wide, hilum ovate, 0.5-1.0 mm long, the strophiole lobes narrow. This species is nearly cosmopolitan, occurring throughout Mexico and Central America and south into Argentina. BOCAS DEL TORO: Careening n a of Chiriqui Lagoon, von Wedel 2819 (MO, NY, US). CANAL ZONE: l2 mi NW of Gamboa 973, Liesner 1396 (F, MO); Standley 28527 (US). 2% km W of Gamboa, gravel along RR, ^e us (F, GH, MO). Balboa, Standley 25646 (US), Standley 32143 (US). CHIRIQUI: Corotu, 6 km SW of the airport of Puerto Armuelles, Busey 398 (F, GH, MO, US). 1.6 m W of Puerto Armuelles, ca. 50 m, Croat 21934 (F, GH, MO, NY). Distrito Guanabano; Quebrada Quanabano, 0-100 m, Cina 22512 (F, MO). Quebrada Mellize, 6 km S of Puerto Armuelles; —150 m, Liesner 419 (MO, NY). PANAMA: Campo Experimental de Monte Oscuro, 12 km de Capira, Correa 1929 (MO). Riomar, Dwyer 1844 (MO). El Capitano, 3 mi E of Chepo, Tyson 5357 (MO). Tocumen, Dwyer 2583 (FSU, GH, NY). Forestry Reserve, Flores 28 (F). Los SANTOS: Chitré/Las Tablas, 0-100 m, Burt & Rattray 53 (MO). 5 mi S of Pocri, Croat 9732 (MO, SCZ). 6 mi S of Las Tablas, D'Arcy 4184 (MO). 12 mi S of Macaracas, Tyson et al. (MO, SCZ). vERAGUAs: Atalay/ Santiago, Koster 116 (MO). 5. Rhynchosia precatoria (H. & B. ex Willd.) DC., Prodr. 2: 385. 1825. Glycine precatoria H.B.K., Nov. Gen. Sp. Pl. 6: 425-426. 1823. TYPE: Mexico: prope Acapulca, not e seen. G. precatoria H. & B. ex Willd., Enum. Pl. Hort. Berol. 2: 755. 1809. TYPE: not seen. Indigofera volubilis Wendl., Bot. Beob. 55. 1798. TvPE: not seen Herbaceous to suffrutescent vine, the stems angular becoming terete, villous to strigose, gland dotted. Leaves trifoliolate, the leaflets obovoid to deltoid, 3.5— 8.5 cm long, 2.5-7.5 cm wide, apically acute, basally obtuse to cuneate, sparsely villosulose above, gland dotted, villosulose beneath; terminal petiolule 1.2—2.3 cm long, lateral petiolule 0.2-0.3 cm long; petiole 2.5-7.2 cm long, villous; stipels obsolete; stipules lanceolate, ca. 3.0 mm long, caducous, villous. /nflorescence equal to or exceeding the leaves, 6.5-10.5 cm long; peduncles 1.0-2.0 cm long; pedicels 1.0—2.0 cm long; bracts lanceolate, caducous, 1.5-3.0 mm long. Flowers with the calyx not exceeding the corolla, 3.0-5.5 mm long, the lobes 1.0-2.5 mm long, the upper lobes longer than the others, puberulent; corolla yellowish green streaked with brown, 0.6-0.8 cm long, the standard orbicular, auriculate, gland dotted, ca. 0.8 cm long, ca. 0.4 cm wide, the claw ca. 1.5 mm long, the wings oblong, ca. 0.7 cm long, ca. 0.2 cm wide, the claw ca. 1.5 mm long, the keel falcate, ca. 0.7 cm long, ca. 0.1 cm wide, the claw ca. 0.2 cm long; stamens ca. 0.7 cm long. Fruits ovate oblong, strigose, 1.5-2.0 cm long, 0.6-0.8 cm wide, the beak 2.0-3.0 mm long; seeds ovoid, ca. 5.0 mm long, ca. 3.0 mm wide, red and black, the hilum linear to ovate, 2.5 mm long, the strophiole lobes narrow. This species ranges from Mexico to Colombia. It is found in young secondary growth in thickets PANAMÁ: Pacora, Burt & Rattray 21 (MO). San Jose Island, Erlanson 382 (GH, NY, US); Johnston 588, 1132 (both GH), 1331 (GH, US). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 759 6. Rhynchosia quercetorum Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 555. 1937. TvPE: Costa Rica: “Oak forests, Santa María de Dota" Standley & Valerio 43431 not seen. Rhynchosia picta Seem., Bot. Voy, Herald 110. tab. 20. 1850; Steud., Nom. 1840. TvPE: Panama: * Volcano of Chiriqui, Veraguas” Seemann 1678 not seen. Herbaceous to suffruticose vine, stems angular becoming terete with age, strigose. Leaves trifoliolate, the leaflets ovate to rhomboid, 4.0-10.0 cm long, 4.0-9.0 cm wide, apically acuminate to acute, basally cuneate to obtuse, glabrous to villosulose above, gland dotted and tomentose to villosulose beneath; terminal petiolule 1.5-3.0 cm long, the lateral petiolule 0.2-0.3 cm long; petiole 3.0-8.0 cm long, strigose; stipules lanceolate 0.4-0.6 cm long, caducous; stipels ca. 1.0 mm long, caducous. Inflorescence equal to or exceeding the leaves, branching, ca. 8.5 cm long; peduncles 1.5-3.5 cm long; pedicels ca. 5.0 mm long; bracts lanceolate to ovate, 2.0-5.0 mm long, caducous. Flowers with the calyx not exceeding the corolla, 3.0-4.0 mm long, the lobes 2.0-3.0 mm long, the vexillary lobes united ca. ! to apex, the calyx strigose to villous; corolla yellow streaked with maroon, the standard obovate, puberulose, gland dotted, auriculate, 7.0— 10.0 mm long, 4.5-8.5 mm wide, the claw ca. 1.0 mm long, the wings oblong, 6.0-10.0 mm long, 2.0-4.0 mm wide, the claw ca. 1.5 mm long, the keel falcate, 7.0-10.0 mm long, ca. 3.0 mm long, the claw ca. 2.0 mm long; stamens 7.0-9.0 mm long. Fruits oblong ovate, heavily strigose, viscid, gland dotted, greenish brown but not black, 2.8-4.0 cm long, 0.8-1.0 cm wide, the beak 3.0-4.0 mm long; seeds ovoid to subglobose, red and black with equal areas of coloration, .0—5.0 mm long, 3.0-4.0 mm wide, the hilum ovate, ca. 2.0 mm long, strophiole lobes narrow. This species is distributed throughout Central America to Colombia. Plants of R. quercetorum occur in open forested areas in Panama and have been col- lected only in Chiriquí above 1,500 m. CHIRIQUÍ: Trail from Paso Ancho to Monte Lirio, Allen 1482 (MO, NY, US). Cerro Punta, Allen 3499 (MO). Finca Lerida, S slopes of Quebrado or es 4738 (MO). Boquete, Finca Collins, Blum & Dwyer 2547A (FSU, SCZ); Davidson 555 (A, F, MO, US), 927 (F, MO, US), 1067 (F, GH, US). S slopes of Volcán Chiriquí, Terry 1302 (F, GH EU ). Boquete, von Hagen 2124 (MO, NY). Finca Lerida to Pina Blanca, Woodson & Schery 307 (MO, US). Callejon Seco, Volcán Chiriquí, Woodson & Schery 495 (GH, MO, US). 7. Rhynchosia reticulata (Swartz) DC., Prodr. 2: 385. 1825. Glycine reticulata Swartz, Prodr. 105. 1788. Hp Jamaica, Swartz, not seen. Dolicholus kuntzei Kuntze, Rev. Mag Pl. 3(2): 61. 1898. TYPE: not see Dolicholus hondurensis Rose, Contr. U.S. a E 10: 101. 1906. rype: Honduras, `° Dept. Santa Bárbara near San Pedro iu ' not se Rhynchosia hondurensis (Rose) J. D. Smith, Enum. Pl. Guat. Index 8: 172. 1907. Dolicholus angulatus Standley, Contr. U.S. A 18: 107. 1916. TYPE: Panama, ‘‘between Miraflores and Pedro Miguel, Canal 2c not Herbaceous to suffruticose vine; stems angled to terete, villous to strigose along ridges. Leaves trifoliolate, leaflets ovate to obovate, the lateral leaflets asymmetric, 2.5-6.5 cm long, 2.3-5.5 cm wide, apically acute, basally cuneate to subcordate, finely strigulose above, reticulate, strigulose and gland dotted be- 760 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 neath; terminal petiolule 2.0-11.0 mm long, the lateral petiolules 1.0-2.0 mm long; petiole 2.0-4.5 cm long, angled, villous; stipels linear, 1.0-2.0 mm long; stipules lanceolate to ovate 0.3-1.0 cm long, 0.1-0.5 cm wide, caducous to per- sistent. Inflorescence shorter or longer than the leaves, 5.0-13.5 cm long, dense- ly, ca. 20-30, flowered; peduncles 1.5-2.5 cm long, pedicels ca. 3.5 mm long; bracts lanceolate, 3.0-5.0 mm long, caducous. Flowers with the calyx usually equal to or exceeding the corolla, 0.7-1.1 cm long, the lobes lanceolate, 0.5-0.9 cm long, puberulent, gland dotted; corolla yellow, 0.6-1.0 cm long, puberulent, gland dotted, the standard obovate, auriculate, 8.0-9.5 mm long, 3.0-4.0 mm wide, the claw ca. 1.0 mm long, the wing oblong, 7.5-8.0 mm long, ca. 2.0 mm wide, the claw ca. 1.0 mm long, the keel falcate, 7.5-9.0 mm long, ca. 2.0 mm wide, the claw 1.0-1.5 mm long; stamens 5.0-9.0 mm long. Fruits oblong ovate, 1.5-2.0 cm long, 0.5-0.7 cm wide, the beak 1.0-2.0 mm long, puberulent; seeds 2, ovoid, ca. 3.0 mm long, brown black, hilum ovate, ca. 1.0 mm long. This species ranges from Mexico to South America including the Greater Antilles. It occurs in disturbed areas, moist forests, thickets, and savannas below 1,200 m. Grear (1978) delimited two varieties of R. reticulata but in Panama these varieties do not appear to be distinguishable on the basis of geography and mor- phology. CANAL ZONE: Pipeline road 242 mi from Gamboa gate, Croat 9385 (MO, SCZ). Gaillard highway 1 mi NW of Summit Garden, Croat 13919 (F, MO, SCZ). Pipeline road 4 mi from main gate, Croat 14100 (MO, SCZ). Without other locality, Greenman & Greenman 5135 (MO). Near Summit Garden, Hladick (MO). Balboa, Standley 26086 (MO, US). 1 mi N of Summit on road to Gamboa, jos 2765 (MO). CHIRIQUI: Near Veladero, Burt & Koster 127 (MO). Francis Arriba School, ca. 14 mi N David, Lewis et al. i (GH, MO). coctÉ: Entrance to Rio Hato Air Base, Blum & is 1867 (FSU, MO, SCZ). HERRERA: El Portero, Las Minas, Burt & Koster 90 (MO). Highway N of Las Minas, Croat 9668 (MO, NY). 10 mi S of Ocú, Tyson et al. 2804 (FSU, MO, SCZ). PANAMÁ: La Chorrera, Bejuco, Burt & Rattray 25 (MO, NY). Between Chepo and El Llano, Croat 14487 (MO, NY). Rio Tocumen N of Chepo road, ae & Allen 239 (GH, MO). Between Rio Pacora and Chepo, Porter et al. 4205 (MO, SCZ). vERAGUAS: Between Canazas and the foot of the Cordillera Central, Headwaters of Rio Canazas, Allen 191 (A, MO, US). La Pena, Burt & Rattray 83 (MO). Without locality: Hayes 720 (NY). 39. SESBANIA Peter S. White*? Sesbania Scopoli,“ Introd. 308. 1777. Nomen conserv. contra Sesban Adanson, Agati Adanson. LECTOTYPE: Aeschynomene sesban L. — Sesbania sesban (L.) Merrill (designated by Jones, 1960). Sesban Adans., Fam. Pl. 2: 327, 604. 1763. Nomen rejic. contra Sesbania. Based on Aeschynomene sesban L. — be eros sesban (L.) Merrill. Agati Adanson, Fam. Pl. 2: 326, 513. 1763. Em rejic. contra Sesbania. Based on Robinia gran- iflora L. — d Vide ies (L.) Per Sesbana R. Brow d Hor , (ed. 2) 4: 330. 1812. Sesbania Scop., orth. Daubentonia DC., "Com "Mem. a 285. 1823. TYPE: Piscidia punicea Cav. = S Setaria punicea v.) Resupinaria Raf., Sylva Tell. 115. 1838. rvPE: Robinia grandiflora L. = Sesbania grandiflora (L.) Pers. a pi. Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- tiesaee 3773 4 For imos generic synonymy see Hutchinson, 1967. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 761 Emerus Kuntze, Rev. Gen. Pl. 1: 180. 1891. TYPE: Aeschynomene sesban L. = Sesbania sesban (L.) rrill. Non a lg Mill. (1754) nor Emerus Guettard (1753). Daubentoniopsis Rydb., qu J. Bot. 10: 497. 1923. TvPE: Aeschynomene longifolia Cav. — Ses- bania M Non (Cav. )D Annual or perennial herbs, shrubs, or small trees. Leaves alternate, even pinnate, often relatively narrow in outline and with numerous leaflets; leaflets oblong, entire, obtuse at the apex, rounded at the base, often glaucous, short petioluled; stipules small, deciduous; stipels minute or apparently absent. /nflo- rescence of relatively short and few-flowered axillary racemes; bracts and brac- teoles small, apparently deciduous. Flowers with the hypanthium broadly cam- panulate, as broad as long or broader, the 5 teeth equal, often short, truncate to triangular with the tip acute to acuminate; often thin; petals white, yellow, pur- plish, red, sometimes white spotted with color or variegated; standard longer than other petals, nearly round in outline, reflexed, short clawed and usually appendaged at the base, the wing petals free, the keel petals curved, joined below, with long claws; stamens diadelphous, the vexillary stamen free, geniculate near the base, the united stamens equal or 5 alternate ones somewhat longer; ovary stipitate, linear, the stigma small, capitate. Fruit often long and slender, terete (in extra-Panamanian species sometimes compressed, 4-angled or 4-winged), short stipitate, beaked, septate between the seeds within, the outside indented between the seeds in some species, 2-valved, indehiscent in a few species, the calyx often not persistent in fruit; seeds many oblong, slightly compressed, smooth, brown. Sesbania is a genus of some 50 species of the tropical and warm parts of both the New and Old World. Among its relatives in Panama it is distinguished by the relatively long and narrow even pinnate leaves. Three species are known from Panama, one of which, Sesbania grandiflora (L.) Pers., is an introduced tree. The two other species are native herbs. Sesbania sesban (L.) Merrill, a weedy plant native to the Old World tropics, has become established in parts of the West Indies and South America. It has not yet been collected in Panama, but is to be looked for there. In general it resembles Sesbania emerus and S. exasperata, which are treated below, but can be distinguished by several characters: the calyx teeth are low triangular, less than 1 mm long, and acute; calyx is less than 5 mm long; corolla is less than 1.5 cm long; racemes are 5-12 flowered; outside of the legume is indented between the seeds. In Sesbania emerus and S. exasperata, the calyx lobes are acuminate, longer than 1 mm; calyx is more than 6 mm long; corolla is more than 1.5 cm long; the racemes are usually less than 6-flowered, and the outside of the legume is not or only slightly indented between the seeds. Sesbania sesban, the type species, has received attention in recent literature regarding tautonyms and paratautonyms (Nicolson, 1975; Terrell, 1977). Terrell (1977) argues that names such as Lycopersicon lycopersicum should be regarded as tautonyms since the two words that make up the binomial are orthographic variants of one another. Both the generic name and the specific epithet are in- flected variants of the same stem word. He feels that such names are prone to confusion. On the other hand, he feels that such names as Sesbania sesban are less confusing. Here, the stem of the specific epithet is uninflected. Terrell (1977) 762 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 argues that the two words making up the binomial Sesbania sesban are not orthographic variants and that the binomial is not a tautonym. Terrell’s argument and proposal seem entirely reasonable and with this in mind the name Sesbania sesban (L.) Merrill is used here. The type of Agati Adanson was initially listed in the Code (1961) and in the Index Nominum Genericorum as Robinia caragana L. Later, both were cor- rected to Robinia grandiflora L., i.e., in later editions of the Code list of con- served and rejected names and in a correction card in the ING list. Literature: Jones, K. 1960. The lectotype species selections of Britton and Brown and the standard species of Hitchcock and Green. Taxon 9: 175-189. Nicolson, D. H. 1975. Paratautonyms, a comment on proposal 146. Taxon 24: 389—390. Seemann, B. 1857. The botany of the voyage of the H.M.S. Herald. London. pp. Terrell, E. E. 1977. The name for tomato. Taxon 26: 129-148. a. Flowers large, 6-9 cm long; se (20—)30—50 cm long; an introduced small tree, but som times flowering when quite young |... 0 0 Onda aa. Flowers smaller, ca. 1.5-2. ey cm long; fruit 15-25 cm long; native herbs or with shrubby appearance to several m high. b. Legume 3-4 mm broad; seeds 3-4 mm long; corolla less than 2 cm long; pedicels in both lower and fruit si. slender, rarely more than 1 mm wide, if so then not over 2 mm wide in fruit 11... sore aa a ee 1. S. emerus Legume 4-6 mm broad; seeds 4-5 mm long; corolla 1.8-2.5 cm long; pedicels rela- tively stout, over 2 mm wide in fruit =. a... 2. S. exasperata = T 1. Sesbania emerus (Aubl.) Urban, Feddes Repert. Spec. Nov. Regni Veg. 149. 1919.—Fic. 44. pase alla emerus Aubl., Hist. Pl. Guiane 2: 775; Tabl. noms. 1. 1775. TYPE: Guiana, Aublet, Coronilla occidentalis Willd., Sp. PI., (ed. 4) 1147. 1802. TYPE: West Indies, Humboldt (B, not seen: O). cro Sesban occidentalis (Willd.) Poir. in Lam., Encycl. Méth. Bot. 7: 129. 1806. Sesbania Mad nud) (Willd.) Pers., Syn. PI. 2: 316. 1807. Sesbania macrocarpa sensu Seemann, Bot. Voy. Herald 108. 1857, non S. macrocarpa Muhl. ex Raf., ov. 137. 18 Emerus s shan y ar. occidentalis (Willd.) Kuntze, Rev. Gen. Pl. 1: 181. 1891. Sesban emerus (Aubl.) Britton & Wilson, Sci. Surv. Porto Rico 5: 395. 1924. Erect herbs or shrubs to 2 m tall, glabrous. Leaves long and narrow with numerous regularly spaced, short stalked leaflets; leaflets ca. 70—100, narrowly oblong, rounded at both ends, mucronate at the apex, to 25 mm long, ca. 3 mm wide; petioles 5-10 mm long; petiolules to 1 mm long; rachis to ca. 20 cm long; stipules linear to lanceolate, acuminate, to 5 mm long, apparently deciduous. Inflorescences axillary racemes with 3—5 flowers on thin pedicels ca. 10 mm long. Flowers 12-18 mm long; hypanthium broad campanulate, to 5 mm wide, ca. 5 mm long, including the calyx teeth, the teeth 1.0-1.5 mm long, triangular with abruptly acuminate tips; standard suborbicular, ca. 15 mm wide and long with raised appendages at the base of the blade, short clawed; wing petals oblong, ca. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 763 = W g y (É UL y | < jD | , S SN SNN ZA ES sÇ ` f = FIGURE 44. Sesbania emerus (Aubl.) Urban.—A. Habit ( x2⁄4).—B. Flower, exploded (x2).—C. Corolla.—C!. Standard.—C2. Wing petal.—C?. Keel petal. 764 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 15 mm long, short clawed; keel petals slightly shorter than the other petals, upcurved; stamens with free portion of filaments alternately unequal. Fruit long and narrow, to 25 cm long, 3-4 mm wide, beaked, stipitate, the margins thick- ened, glabrous; seeds numerous, oblong, 3-4 mm long. Sesbania emerus is found in fields and wet ground in Panama and northward throughout Central America. It is also found in the West Indies. The relation of Sesbania emerus to S. exaltata (Raf.) Cory and S. macrocarpa Muhl. ex Raf., both of the southern United States and to S. exasperata H.B.K. awaits mono- graphic study. Sesbania exasperata H.B.K., in material seen by the author, has consistently broader pedicels and fruits and longer petals than S. emerus. See- mann s (1857) reported S. macrocarpa Muhl. ex Raf. from Panama proved to be S. emerus. CANAL ZONE: Río Grande Station, Hayes 772 (NY). Balboa, dien 25648 (US). CHIRIQUÍ: 1 mi W of Puerto Armuelles, Croat 22037 (MO). Los SANTOS: Managre Beach, 5 mi SE of Chitré, Tyson et al. 3018 (MO, SCZ). PANAMÁ: S of Tocumen Airport, D' Arcy 9662 (MO). 2 km W of El Llano, 0-100 m, Nee 7953 (DUKE, MO, US). Between Matías Hernández and Juan Díaz, Standley 31963 (US). Vía Tumba Muerto, Taylor (MO). vERAGUAS: Santiago-San Francisco Road at Río Santa María, 60 m, Nee 8195 (MO). 2. Sesbania exasperata H.B.K., Nov. Gen. Sp. Pl. 6: 534. 1824. TvPE: Venezuela (P, not seen; microfiche, MO). Lotus pega Vell., Fl. a 315. 1825; Atlas 7, tab. 137. 1835. TYPE: Brazil, not seen. Non Willd., . PI. (ed. 4) 1394. d Sesbania dubia Steud., Nom. ., (ed. 2) 2: 572. 1841. TYPE: Brazil, not seen. Non H.B.K., Nov. Gen Pl. 6: 534, tab. Dik 1824 Emerus PN MEN (H.B.K dde dq Rev. Gen. Pl. 1: 181. E Sesban exasperatus (H.B.K.) Rydb., No. Amer. Fl. 24: 203. Sesbania sesban sensu Johnston, Sargentia 9: 161. 1949. S S. sesban (L.) Merrill. Erect herbs or shrubs, to 3 m tall. Leaves 20-30 cm long, leaflets 30—50—70 or more, narrowly elliptic to narrowly oblong, rounded at both ends, mucronate apically, to 30 mm long, to 5 mm wide; petiolules to 1 mm: petioles more than 1 cm long, petiole base slightly sheathing and the stems often noticeably 3-lined from the nodes. /nflorescence of 2-6 flowered axillary racemes, pedicels rela- tively stout, over 2 mm wide. Flowers 1.8—2.5(-3.0) cm long when pressed, the standard not reflexed; hypanthium broad campanulate, 7-10 mm long, 6-7 mm wide when pressed, the teeth 3-4 mm long, triangular, acuminate: petals yellow, standard ovate, 3 cm long, 1.8 cm wide; wings oblong to obovate, to 2.5 cm long, the claw to 0.5 cm long; keel petals obovate, to 2.5 cm long, the claw to 0.5 cm long, upcurved; staminal sheath to 15 mm long. Fruit long and narrow, to 25 cm long, 4-6 mm wide, beaked, stipitate, dull, brown, the flat surface slightly in- dented between the seeds, the margins thickened and also slightly indented be- tween the seeds; fruiting pedicels to 2 mm thick; seeds numerous, oblong, 4—5 mm long. Sesbania exasperata H.B.K. is represented by a single collection from Pan- ama, which was wrongly reported by Johnston as Sesbania sesban. Due to the paucity of Panamanian material, the description above was partly drawn from a 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 765 range of South American material at MO. The species is predominantly South American, although it has also been reported from the West Indies. Sesbania exasperata is similar to S. emerus, but is larger in many dimensions, i.e., in flower length, in thickness of fruits and fruiting pedicels, in the length of the hypanthium teeth and cup. The stems are more noticeably 3-lined from the nodes in S. exasperata than in S. emerus. PANAMÁ: San Jose Island, Johnston 1146 (GH). 3. Sesbania grandiflora (L.) Pers., Syn. Pl. 2: 316. 1807. Robinia grandiflora L., Sp. Pl. 722. 1753. TYPE: India, not seen. Aeschynomene grandiflora (L.) L., Sp. Pl., (ed. 2) 1060. 1763. Dolichos arboreus Forssk., Fl. Aegypt.- -Arab. 134. 1775. TYPE: Yemen, PER lod seen. schynomene coccinea L. f., Suppl. Pl. 330. 1781. TYPE: "Nova Zeelandia, a grandiflora (L.) Willd., Sp. Pl., (ed. 4) 1145. 1802. Non C. grandiflora T. (1844). Coronilla coccinea (L.f.) Willd., Sp. Pl. (ed. 4) 1146. 1802. Sesban lic ed rus (L.) Poir. in Lam., Encycl. Méth. Bot. — E" 1806. S. coccineus (L.) Poir. in Lam., Encycl. jon Bot. 7: 127. Secum coccinea (L.) Pers., Syn. Pl. 2: 316. Agati sabes ae UA AVR GP J. Bot. ( Desvau 1: 120. tab. 4. 1813. A. coccinea (L.) D , J. Bot. (Desvaux) 1: 1813. Agati s. var. pi (L.) Wight & ye Prodr. Ind. Or. 1: 216. 1834. Emerus grandiflorus (L.) Kuntze, Rev. Gen. Pl. 180. 1891. Small tree to 8 m tall, young parts hairy, soon becoming glabrous. Leaves alternate, even pinnate with (18—)20—40(—50) narrowly elliptic to oblong leaflets; leaflets ca. 7-10 mm wide, 25-35 mm long, rounded at the tip and base; petiole ca. 1 cm long; petiolules ca. 2 mm long, the rachis to 25 cm long; stipels minute; stipules narrowly triangular, to ca. 5 mm long, apparently deciduous. /nflores- cence of 2-4 flowered axillary racemes, pedicels to 2 cm long, thin. Flowers large and showy, ca. 6-9 cm long; hypanthium ca. 10-18 mm long, not including the narrowed base, 15-18 mm wide (flattened out); teeth low to absent, the hypan- thium rim variously erose; petals white, rosy white, or red; standard ca. 50-80 mm long, 40 mm wide, claw and basal auricles ca. 15 mm long; wing petals ca. 50-80 mm long, 20 mm wide, the claw ca. 20 mm long; keel petals upcurved, ca. 60 mm long with basal auricles, claw ca. 2 the length of the blade; stamens with the sheath to 80-90 mm long, 3 mm wide, strongly upcurved. Fruits long and narrow, to 50 cm long, ca. 0.6-0.9 cm wide, beaked, the stipe to 4 cm long; surface more or less flat or slightly indented between the seeds, smooth, dull; margins thickened and raised above the valve surface, also slightly indented be- tween the seeds; seeds numerous. Sesbania grandiflora (L.) Pers. is Old World in origin and is widely cultivated in the warmer parts of the world. Most of the Panamanian collections are from cultivated trees; Standley (1928) reports that it has also been found *'wild."" Ses- bania coccinea (L.) Pers., considered here synonymous with S. grandiflora, was proposed for trees with dark red flowers. L ZONE: Cultivated, Culebra, Jp 2573 (US). “Planted but also wild," Balboa, Standley 25513 (US). Cultivated, Balboa, Standley 30824 (US). cocrÉ: Cultivated, Santa Clara, Blum & Tyson 1883 (FSU, MO, SCZ). PANAMÁ: Dd de Panama, Carrasquilla 259 (DUKE, F, MO). 766 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 40. SOPHORA Michael O. Dillon*® Sophora L.,*° Sp. Pl. 373. 1753; Gen. Pl., ed. 5, 175. 1754. LECTOTYPE: S. alo- pecuroides L. (designated by Britton & Brown, 1913). Trees, shrubs, or rarely perennial herbs. Leaves alternate, imparipinnate; leaf- lets alternate to subopposite; stipules linear to deltoid, caducous or absent: stipels linear, minute or absent. Inflorescence racemose or paniculate, terminal or ax- illary. Flowers ca. 1.0-4.5 cm long; calyx campanulate, often gibbous, the lobes 5, subequal or truncate, the upper lobes often connate; corolla white, yellow or blue violet, the standard broadly obovate or orbicular, erect or spreading, the wings oblong, oblique, the keel oblong, usually partially connate or overlapping; stamens 10, free or rarely connate basely into a ring, alternately subequal, the anthers versatile, ellipsoid, dorsifixed, ca. 1 mm long; ovary short stipitate, the ovules numerous, the style incurved, the stigma terminal, minutely capitate-pen- icillate. Fruit moniliform, terete or slightly compressed, somewhat tortulose, fleshy to woody, commonly indehiscent; seeds spherical or subellipsoid, red, yellow or brown, the hilum lateral or subapical; cotyledons thick. Chromosomes: n = 8,9, 11, 14, 18. Sophora contains about 80 species and has a worldwide distribution in tropical and temperate regions. Only one species, Sophora tomentosa, is found in Central America south of Mexico. This geographically widespread and morphologically variable taxon has been separated into four subspecies by recent authors (Brum- mitt & Gillett, 1966; Yakovlev, 1964), with the Panamanian material referable to subspecies occidentalis. Literature: Britton, N. L. & A. Brown. 1913. Illustrated Flora of the Northern United States., ed. 2, 2: 342. Brummitt, R. K. & J. B. Gillett. 1966. Notes on the genus Sophora in Africa, including an Asian species found near Zimbabwe. Kirkia 5: 259-270. Yakovlev, G. P. 1964. Novaiia systema roda Sophora L. i ego filogenez. Trudy Leningrad Khim.-farm. Inst. 17: 50—77. 1. Sophora tomentosa L.,% Sp. Pl. 373. 1753, non Dippel, 1893. TYPE: Ceylon, Hermann (BM, not seen). la. Sophora tomentosa subsp. occidentalis (L.) Brummitt, Kirkia 5: 265. 1966.— Fic. 45 S. occidentalis L., Syst. Nat., ed. 10: 1015. 1759. TYPE: Jamaica, P. Browne (LINN, not seen; microfiche, MO). Shrub, to ca. 3 m tall; stems puberulent to tomentose. Leaves 13—17-foliolate, the rachis 12-25 cm long, puberulent; leaflets opposite or subopposite, broadly ° Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 5 For complete synonymy see Rudd, 1972. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 767 FIGURE 45. Sophora tomentosa L.—A. Habit (x!4).—B. Fruits (x1⁄). 768 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 elliptic or suborbicular, apices rounded to emarginate, the bases rounded, trun- cate or subcordate, 2.5-5.0 cm long, 1.5-4.0 cm wide, the upper surface puber- ulent, shiny; lower surface tomentulose; petiolules ca. 2 mm long, puberulent; stipules linear-deltoid, tomentulose, ca. 3 mm long, caducous: exstipellate. In- florescence racemose, terminal, many flowered; bracts 3-5 mm long; bracteoles absent. Flowers 20-25 mm long; calyx 6-8 mm long, subtruncate, sericeous: corolla yellow, the standard 9-12 mm wide, narrowly obovate, the wings clawed, auriculate, the keel petals clawed, auriculate, fused distally. Fruit stipitate, sub- coriaceous, cinereo-tomentulose, terete, torulose or moniliform, 1—15-seeded, 5-15 cm long, ca. 8 mm in diameter, abruptly constricted between the seeds or sometimes only slightly constricted; seeds sublustrous, light brownish, spherical or subellipsoid, 5-8 mm long, 5-6 mm in diameter. Chromosomes: n = 9. Sophora tomentosa subsp. occidentalis is widespread throughout the West Indies, adjacent southern Texas, northeastern Mexico, the Yucatan, Belize, Pan- ama and northern Colombia. In the Old World, it occurs on the Atlantic coasts of West Africa from Senegal to Nigeria and Sao Tome. In Panama, it is an infrequent element of the beach strand vegetation of the northern Caribbean coastal region, possibly restricted to calciferous soils. Subspecies tomentosa is quite similar to subsp. occidentalis, the latter having leaflets with sparser pubescence of short, adpressed hairs, and the Old World subsp. tomentosa having leaflets with pubescence silky, curved and matted into a short tomentum. BOCAS DEL TORO: Isla Colón, Wedel 504 (GH, seme Old Bank Island, Wedel 2039 (GH, MO, US). pean Cay, Wedel 2807 (GH, MO, US). Nances Cay, Wedel 2864 (GH, MO, US). COLON: Playa L , Correa et al. 3006 (MO). Near Portobelo. Croat 14126 (F, MO). SAN BLAS: Isla i pee 15475 (MO, US). 41. STIZOLOBIUM W. G. D'Arcy*? Stizolobium, P. Br., Civ. Nat. Hist. Jam. 299. 1756. TYPE: S. pruriens (L.) Med. Labradia Swendiatavr, Arznuymitt. 1: 164. 1801. Not seen, fide Index Kewensis. LECTOT YPE: L. ruriens (L.) Med. Mucuna sect. II. Stizolobium (P. Br.) DC., Prodr. 2: 405. Bas Carpopogon Roxb., Fl. Indica 3: 283. 1832. LECTOTYPE SPECIES: C. niveum Roxb. Mucuna subgenus III. Stizolobium (P. Br.) Bauer, Fl. British India 2: 186. 1879. Climbing or trailing vines, puberulent to cinereous. Leaves pinnate trifolio- late, the leaflets mucronate; petiolules pubescent; petioles often elongate; stipels linear; stipules caducous. /nflorescences axillary racemes, sometimes elongate, the flower buds narrow, the calyx with unequal teeth, the upper pair connate, the lowermost teeth longest; standard short, the wings longer than the standard, the keel tubular, apically cucullate, indurated, about as long as the wings; stamens diadelphous, the vexillary stamen free, the other 9 with the filaments connate into a tube above halfway, the free portion alternately thick and short, narrow and long; ovary pubescent, the style glabrate, the stigma minute. Legume oblong, #7 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 769 narrow, several seeded, striate, densely tomentose, usually with stinging hairs, falsely septate, dehiscent; seeds faboid, plump to subglobose, the short hilum surrounded by an elevated elliptical caruncle, the testa thin. Stizolobium includes about 8 species ranging across the Old World tropics to Australia. At least two species are naturalized in the Neotropics. Stizolobium has been included in Mucuna by many writers, usually with hes- itation, but the differences between Stizolobium and typical elements of Mucuna are manifest. Removal of Stizolobium from Mucuna still leaves Mucuna a het- erogenous group (Verdcourt, 1971), at least when Old World species are consid- ered. Most evident characters for separation are in the seeds. In Mucuna the seeds are large, globose to discoid vs. reniform to faboid (S tizolobium), the hilum is narrow and circumferential around more than 4 the seed vs. narrowly elliptical around less than 1⁄4 of the seed. The legumes are seldom more than 3 times as long as wide vs. several times longer than wide. There are some floral differences as well. In Mucuna anthers are sometimes barbate, anthers are dorsifixed, and there is different connation of the filaments. Vegetatively, the eophylls of Mucuna are alternate and reduced vs. opposite, simple and cordate, foliaceous, and in Mucuna germination is hypogeous vs. epigeal in Stizolobium (Moldenke, 1933). Mucuna comprises mostly high climbing forest vines while Stizolobium com- prises vines on the ground or on fence-rows, often of disturbance. In seeds of Mucuna the putamen is resistant to sea water decay, hence seeds are frequently found in beach drift, but seeds of Stizolobium have a less resistant putamen and the seeds are seldom found in beach drift (Standley & Steyermark, 1946). Literature: Burkill, I. H. 1935. A Dictionary of the Economic Products of the Malay Pen- insula. 2 Vols. London. Moldenke, N. 1933. Tropical American Plants. Phytologia 1: 4-18. Piper, C. V. 1917. The Cowhage and related species. Proc. Biol. Soc. Wash. 30: 51-52. 1. Stizolobium pruriens (L.) Medic., Vorles. Churpffálz. Phys.-Ocon. Ges. 2: 399. 1787.—Fic. 46. Dolichos pruriens L., in Stickman, Dios. Herb. Amb. 23. 1754; Syst. Nat. ed. 10: 1162, 1759. TvPE: donesia, Rumphius, Herb. Amb. 5, tab. 142. 175, not seen, fide Verdcourt 1971. 182 M. purita Wight in Hook., Bot. Misc. 2: 348. 1831. Stizolobium pruritum (Wight) Piper, Proc. Biol. Soc. Wash. 30: 54. 1917. Vine sprawling on ground or shrubs, occasionally high climbing, mostly 2-3 m long; stems fine puberulent with white or brownish stiff, curved reflexed or appressed hairs, terete but drying minutely striate. Leaves trifoliolate, the leaflets mostly 5-12 cm long, thin, about equal, apically obtuse, mucronate, basally ob- tuse to truncate, the lateral leaflets oblique, pubescent with soft hairs, densely pubescent to glabrate, mostly more so beneath, the petiolules 3-4 mm long, pilose, sometimes with a few long (1 mm) coarse hairs, petioles exceeding the leaflets, sometimes twice as long, puberulent as on the stems; stipels minute, 770 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 46. Stizolobium pruriens (L.) Medic.—A. Habit (x1⁄2). [After Allen 16675A.]—B. Flower (x1).—C. Corolla.—C!. Standard (X 1).—C2. Wing petal (x1).—C?. Keel petals (x 1).—D. Stamens (x 1).—E. Stamens.—E!. Large anther ( x 10).—E?. Small anther ( x 10).—F. Pistil (x5). [Af- ter Allen 975.]—G. Fruit (x!4).—H. Fruit opened to show seeds (x12). [After Ebinger 81A.] 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 771 scale-like 1-3 mm long; stipules drying dark, narrowly ovoid, apically linear, ca. 4 mm long, mostly pilose with ascending hairs, mostly caducous. Inflorescences axillary, racemes to 20(—30) cm long, mostly 5-15 flowered, the branches grey silver pubescent, the pedicels short, ca. 4 mm long, tomentose, paired on short shoots, the bracts not seen, not elongating in fruit. Flowers ca. 4 cm long, the calyx campanulate, the limb ca. 1 mm across, greyish or silvery velutinous, the teeth linear lanceolate, ca. 7 mm long, the lowermost longest, the upper pair united into a narrowly deltoid lip; corolla purplish and white, the standard 15- mm long, the tip slightly upcurved and obtuse, ovate, the wings ca. 30 mm long, much exceeding the standard, narrow, the keel slightly longer than the wings, tubular, narrow, the apex cucullate and indurated, enclosing the anthers; ovary canescent, the style glabrous, the stigma minute. Legume oblong to fusiform, often irregularly curved, 4-9 cm long, 10-15 mm thick, densely tomentose with stinging hairs, orange, yellow or blackish brown, mostly striate under the hairs, tardily dehiscent with false septa between the seeds; seeds several, faboid, 1—1.5 cm long, plump, variously colored, shiny, with a narrowly elliptical, elevated caruncle around the short lateral hilum, the testa hard but thin. In fruit, this species can be confused with no other member of the Panamanian flora, for the densely woolly pods with stinging hairs often alert the observer before the plant is seen, and the faboid seeds with elevated caruncle around the hilum will separate the species from members of Mucuna which also have stinging hairs on the pods. Some varieties of this species are cultivated in the Old World for forage. The pods are similar to the typical variety but lack stinging hairs. Stizolobium pruriens is widespread in the Neotropics but is probably native to the Paleotropics. The brittle hairs are mechanically irritating and also bear a proteolytic enzyme which is irritating. The English common name ''Cow-Itch" is a corruption of the Hindu name ‘‘Kewach’’ (Burkill, 1935), and does not imply potential for injuring cattle. In Spanish the plant is known as ‘‘Pica-Pica’’ (Standley, 1928). CANAL ZONE: W end of Gatun Lake dam, Blum & Tyson 1999 (FSU, MO, SCZ). W edge of Cocoli, Croat 9180 (F, MO, SCZ). Pipeline Road, 1.2 mi NW of Gamboa gate, Croat 12736 (MO). Farfan, D’Arcy 9628 (MO). Cocoli, Dwyer 7229 (GH, MO, US). Madden Dam Area, Dwyer 8377 (MO). Gamboa, Greenman & Greenman 5164 (MO). Near Fort Clayton, Greenman & Greenman 5185 (MO). Hills E of Curundü, ple 5185 (F). Farfan Beach, Lewis & Dwyer 58 (MO). 1 km S of Madden Dam, 80 m, Nee 8907 (GH, MO, SCZ). Between Miraflores and Pedro Miguel, Pittier 2496 (US). Paraíso, Pittier 2518 (US). Balboa, Standley 27132 (US). Gatün, did 27249 (US). Gamboa, Standley . 28453 (US). Las Cruces Trail between Fort Clayton and Corozal, Epio 29004 14 (MO). Las Vueltas, 1-100 m, Pipe 110 Me Pue rto paw "a , 20 m, Liesner 139 (MO). COLÓN: SANTOS: 1 mi W of To onosí, Croat 298 6 (MO). PANAMÁ: Bej uco, Allen 975 (GH, MO, US). Cerro Cantratista, Buitrago 30 (MO, PMA). Between Gooty Lake. and Panamerican Highway, Correa & Dressler 442 (A, MO, PMA, SCZ). Base of highway to Cerro Jefe, D’Arcy 13440 (K, MO, SCZ). Between Panama and Chepo, Dodge et al. 16675 (GH, MO). Chepo highway W of Rio Pacora, Duke 5893 (MO, WIS). Without j Ie cality, Kennedy et al. 2013 (MO, US). Taboga Island, Macbride 2797 (US). Matías Hernández, Y 6810 (GH ud e Panamá, Sargent 7 (A). 2 km entrada camino a Cerro Azul, Taylor 1 (PMA). Camino después de divisar ay de Sardinilla, Taylor 43 (PMA). VERAGUAS: Puerto Mutis, 12 mi S of rare ‘Tyson 5194 (FSU 772 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 42. STYLOSANTHES Michael O. Dillon** Stylosanthes Swartz, Prodr. Veg. Ind. Occ. 108. 1788. LECTOTYPE: S. procumbens Swartz. Astyposanthes Herter, Revista Sudamer. Bot. 7: 209. 1943. Based on Stylosanthes Swartz. Perennial herbs or rarely subshrubs, 0.1—1.5 m tall; stems erect, ascending or sprawling procumbent, diffusely branched, suffruticose, terete, finely striate, un- armed. Leaves alternate, pinnately trifoliolate, the rachis below terminal leaflet 1-4 mm long; leaflets elliptic to lanceolate, 5-30(-40) mm long, 2-15 mm wide, apices obtuse to acute, occasionally mucronate, basally attenuate, the costas prominent; petioles 1.5-9.0 mm long; stipules amplexicaul, bidentate, pubescent, adnate with the petiole base, 3-11 nerved; exstipellate. Inflorescence spicate, terminal or axillary, I-several flowered, each flower surrounded by a series of bracts and bracteoles, the outermost bracts similar to stipules and giving rise to 1-3 reduced leaflets, similar to cauline leaflets except for size, the bracts enclosing a smaller outer bracteole, usually 3 nerved, ciliate along the margin, within this a densely ciliate axis rudiment sometimes present; within this 1 or 2 inner brac- teoles, deeply cleft, the apices usually long ciliate. Flowers 5-merous, the calyx 4—15 mm long, tubular, 5 lobed, unequal; corolla papilionaceous, yellow or yellow orange and purple striate; stamens 10, monadelphous, the filaments coalescent into a tube, splitting on vexillar side after anthesis; 5 versatile anthers alternating with 5 subbasifixed ones; ovary subsessile, the 2(3) ovules campylotropous, the placentation marginal, the style long, filiform, the stigma minute, terminal. Fruit a loment, biarticulate, distally fertile, proximally abortive or fertile, reticulate or muricate, beaked; seeds ovate or lenticular, compressed, light brown to black, lustrous, strophiolate, Chromosomes: n = 10 Stylosanthes is composed of approximately 50 species ranging throughout tropical and subtropical America, and tropical Africa and Asia. Various species are used as fodder crops, and at least one species, S. guyanensis, has been employed as a cover planting to inhibit soil erosion. Hutchinson and Dalziel (1937) have reported several folk uses for S. erecta Beauv. in Africa including herbal smoke, aphrodisiac, decoction used in baths, and as a charm against injury in battle. No local names or uses have been recorded for the species found in Panama. Stylosanthes hamata (L.) Taub. was reported in error from Panama (Mohlen- brock, 1957), based on Shafer 36 (NY) from Antigua. No representatives of this taxon from Panama have been seen by the present author. Literature: Mohlenbrock, R. H. 1958. A revision of the genus Stylosanthes. Ann. Missouri Bot. Gard. 44: 299-355. "* Department of Botany, Field Museum of Natural History, Roosevelt Road and Lake Shore Drive, Chicago, Illinois 60605. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 773 a. Inflorescence 5 to 20 flowered; beak of loment minute, 0.1—0.5 mm long, inflexed; leaflets 10-30 mm lon 1. S. guyanensis aa. Inflorescence 2 to 5 flowered; beak of loment 1.5-2.5(—4.0) mm long, strongly to weakly hooked; leaflets to 15 m b. Leaflets punctate, bipi pia abaxially; distal articulation of loment ca. 2 mm long, m lon oiled 3. beak ca. 1.5 m , strongly c S. viscosa bb. Leaflets never aes je da setose to glabrous; distal articulation greater than 2 mm long, beak 1.5-3.5(-5.5) mm long, uncinate to coiled ------------------ 2. S. humilis 1. Stylosanthes guyanensis (Aubl.) Swartz,” Kongl. Vetensk. Acad. Handl. 11: 296. 1789.—Fi1Gc. 47. Trifolium guyanense Aubl., Hist. Pl. Guiane 776. 1775. Type: Guiana, Aublet (BM, not seen). Suffruticose herbs, erect or rarely scandent, to 1 m tall, glabrous to puberulent or pilose, often hispid or setose, the trichomes viscid. Leaflets pinnately trifo- liolate or rarely a few unifoliolate, linear lanceolate to elliptic, 10-30(—40) mm long, 2-5 mm wide, the terminal leaflet slightly larger, apically acute, mucronate, puberulent to sericeous with tuberculate based trichomes, nerves 2-8 pairs, prom- inent; petioles 1-10 mm long, canaliculate, puberulent to setose, the rachis 0.5— 1.5 mm long; stipular sheath pilose to setose, 2-15 mm long, 3- to many-nerved, the teeth subulate, 2-10 mm long. Inflorescence spicate, mostly 5—20 flowered; bracts usually subfoliaceous, unifoliate, spinulose toothed, the sheaths glabrous to bristly, 3-7 mm long, 3 to 9 nerved, the outer bracteole 1, lanceolate, 2.5-3.0 mm long, acuminate, glabrous to pilose; axis rudiment none; inner bracteole 1, 2-3 mm long, acuminate, glabrous to pilose. Flowers yellow; calyx tube 4-8 mm long, glabrous to sparsely pubescent, the lobes 3-5 mm long, obtuse to acute, ciliate or sometimes pilose; standard 4-8 mm long, 3-5 mm wide, suborbiculate, the wings 3.5-5.0 mm long, auriculate, spurred within at the base, the keel petals 3.5-5.0 mm long, falcate. Fruit a loment, 1.5-2.5 mm wide, ca. 3 mm long, reticulate, glabrous, ovoid, plump, the beak minute, 0.1-0.5 mm one: strongly inflexed, only the distal articulation fertile. Chromosomes n = 1 Stylosanthes guyanensis is a most variable species occupying a wide range of altitudes and habitats. Numerous subspecies, varieties and forms have been pro- posed; however, intergradations occur between almost all of these. Mohlenbrock (1957) recognized two subspecies within S. guyanensis: ssp. guyanensis occurs from Central America through South America into northern Argentina and also in the Antilles, while ssp. dissitiflora is limited to a small area in southwestern Mexico, between 3,500—5,000 feet. The latter, not occurring in Panama, is easily distinguished by its small, purplish leaflets and inflorescence of only 1-4 flowers. CANAL ZONE: Drowned forest of Río Puente near junction with Río Chagres, Dodge et al. 16835 Cruces Trail in Government Forest, Hunter & Allen 753 (GH, MO). Empire to Mandinga, Piper 5129 (US). Ancón Hill, Piper 5130 (US). Between Corozal & Ancón, Pittier 2165 (NY, US). Sosa Hill, Balboa, Standley 26443 (US). Old Las Cruces Trail d Fort Clayton and Corozal, Standley 29159 (US). Summit, Standley 30068 ios e Road NW of Gamboa, Wilbur & Teeri 13389 (DUKE). 0.5 m from Summit Garden, Wilbur & Weaver 11207 (DUKE, MO). Ancón Hill, Williams 28 (NY). cHIRIQUí: David to Concepción, pre m, Burt & Koster 137 (MO). Volcan to Cuesta de 89 For additional synonyms see Mohlenbrock (1958). 774 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 47. Stylosanthes guyanensis (Aubl.) Sw.—A. Habit (x14). [After diee 16835.]—B. Flower (x6). —C. Corolla.—C'. Standard (x4).—C?. Wing petal (x4).—C?. Kee tal (x4).—D. Stamens (x 6).—E. Pistil (x6). [After Greenman 5125. ]—F. Fruit ( x12). [After mo i Allen 753.) Piedras, 980 m, Burt ga aani 160 (MO). Road to Cerro Punta, ca. NW of Hato de Volcán, Correa 1371 (MO, PMA). La above ^u 4500 ft, D'Arcy & D'Arcy 6366 (MO). E side of Cerro Pando, near Río Chi Ed Viejo. 600 ft, D'Arcy & D'Arcy 6613 (F, GH, MO, NY). Lava fields near town of Volcan, 4600 ft, Duke 9206 Ma. US). Cerro Horqueta, ca. 1500 m, Duke et al. 13648 (MO). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 775 Llanos Francia, Dwyer & Lallathin 8716 (MO). Between Santiago and San Félix, Folsom & Collins MO). Alto Boquete, 1125 m, Partch 69-52 (MO). Boquete, 5500 ft, Terry 1264 (F, GH, MO, US). 5 mi NE of El Hato del Volcán towards Volcán de Chiriquí, Wilbur et al. 11857 (DUKE). 6 m W of San Lorenzo, Wilbur et al. 15393 (DUKE). cocrÉ: Nata, 50 m, Allen 838 (MO, NY, US). 3 mi NE of Antón, D'Arcy & Croat 4112 (MO, PMA). Penonomé, Llanos, Ebinger 1099 (F). 2 m E of Río Hondo along Panamerican Highway, Gentry 2915 (MO). Penonomé, Williams 148, 167 (both NY, US). HERRERA: Between Las Minas and Pesé, Burch et al. 1336 (F). Near Ocú, 100-200 m, Burt & we 87 (MO). Chepo de Las Minas, Folsom et al. 7062 (MO). 10 mi S of Ocü, Tyson et al. 2820 (MO, SCZ). PANAMÁ: Near Chepo, 0-100 m, Burt & Rattray 14 (MO). Near El Valle, 450 m, Burt & Rattray 28 (MO). Road to El Valle close to Interamerican Highway, 0-100 m, Burt & Rattray 38A (M Between Río Pacora and Chepo, Dwyer et al. 5090 (MO). Playa near Río Mar, Ebinger 507 (F). 7 m W of Chepo, Gentry & T 1633 (MO). Sabanas near Chepo, 30 m, Hunter & Allen 75 (GH, MO). Río To ocumen, N of Chepo Road, Hunter & Allen 222 (F, GH, MO, NY). Cerro Campana, wis et al. 3071 (COL, MO, UC). Without exact locality, Maurice 790 (US). Taboga Island, Semple Pa-4 (US). Las Sabanas, Standley 25837 (MO, US). Savanna E of Rio Tocumen, Standley 26510 (US). Juan Franco Race Track, Standley 27726 (US). Taboga Island, x ed 28013 (US). Between Matías WIEN. and Juan Diaz, Standley 32077 (US). Between Cerro Azul and Galaria Barleta, arr : (M MA). Ca. 9 km SE of El Valle de Antón, Wilbur et al. 11166 (DUKE, GH, MO, US). (DUKE). vERAGUAS: Ca. 5 mi NE of La Mesa, Blum & Tyson 631 (FSU, MO, SCZ). Near La Pena, 0-100 m, Burt & Rattray 84 (MO). Without exact locality, Duke 6179, 6198 (both MO). Savanna 10 m W of Santiago, Duke 13647(3) (NY). 270 km W of Panama City, between Santiago & San Félix, Folsom & Collins 6910 (MO). Santiago-Santa Fé Road, 18 km S of Santa Fé, 400 m, Nee 8164 (MO). 2. Stylosanthes humilis H.B.K., Nov. Gen. Sp. Pl. 6: 506. 1823. TvPE: Venezuela, Humboldt & Bonpland, not seen. Non S. humilis Rich. ex Hemsl. Astyposanthes humilis (H.B.K.) Herter, Revista Sudamer. Bot. 7: 209. 1943. Herbs, often suffruticose; stems ascending to prostrate, to 0.5 m tall, branched, puberulent to hirsute. Leaflets lanceolate to elliptic, mucronate, acute, the adaxial surface and the margins sparsely setose to glabrous, 3—4 pairs of conspicuous nerves; terminal leaflet 5-15(-20) mm long, 2-4 mm wide; petioles 2.5-5.0 mm long, canaliform, setaceous, the rachis 1.5—4.0 mm long; stipular sheath 3-5 mm long, short bristly, 5-7 nerved, the teeth subulate, 2.5-3.5 mm long. Inflorescence spicate, 3—4 flowered; bracts 1—3 foliolate, the sheath 2.5—4.5 mm wide, bristly, 5-9 nerved, the teeth 2-3 mm long; outer bracteole 2.5-3.0 mm long, the apex ciliate; axis rudiment none; inner bracteole 1, 2.0-2.5 mm long, the apex ciliate. Flowers yellow; calyx tube 4-5 mm long, the lobes acute, ca. 1.5 mm long; standard suborbiculate, spurred within at the base, 3—4 mm long; wings 3-4 mm long, clawed and auriculate at the base; keel petals 3.0-3.5 mm long, falcate. Fruit a loment, 1.5-2.5(—4.0) mm long, 1.5-2.5 mm wide, reticulate nerved, usually puberulent, rarely pilose, only the distal articulation fertile, the beak strongly uncinate to coiled, 1.5—3.5(—5.5) mm long. Chromosomes n = 10. Stylosanthes humilis is quite variable in size and shape of the leaflets, and in the size of the loments. The species ranges from central Mexico through Panama to Colombia, Venezuela, and Brazil; it also occurs in the Antilles and is adventive in Malaysia and Australia. In Panama, this species generally occurs at elevations below 150 m, and it is 776 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 distinguished by its glabrate, apunctate leaflets and the elongate beak of the ter- minal articulation. . ZONE: Cocolí Road to Contractors Hill, Dwyer 7213 (MO). La Boca, Mori & Kallunki MO). Corozal, Standley 27403 (US). Old Las Cruces Trail between Fort Clayton and Corozal, Standley 2919] (US). Albrook Air Force Base, Tyson & Lazor 6013 (FSU, PMA). CHIRIQUI: bear site 64, Burt & Koster 133 (MO). Road to Divala, site 74, Burt & Moa 149 (MO). Frances cube School, Lewis et al. 655 (GH, K, MO, UC, US). David airport, Lewis et al. 753 (GH, MO). 5 km S of Las Lomas, McCorkle C-25 (FSU, SCZ). David, Rattray 3 (MO). cocLé: Near Divisa, Burt & Koster 96 (MO). Aguadulce, Sri 4837, 4983 (both US). coLON: 0.7 m NE of bridge over Río Piedras on road from Portobelo, Nee & Mori 3650 (MO). oe AMA: Between Panama and Chepo, Dodge et al. 16700 rqa Río Mar, Ps 2321 (FSU, SCZ). ; mi S of Campana, McDaniel 8339 (DUKE, FSU). Las co Sranda 25882, 25914 (both US). Near Punta Paitilla, ues 26293 (US). Corozal Road, . 26794 (US). Vicinity of Juan Franco Race Track, Standley 27730 (US). Río Tocumen, Standley 29474 (US). Las Sabanas, Standley 41178 (US). vERAGUAS: Hills W of Sona, Allen 1054 (GH, MO, US). Davisa/Santiago, 0-100 m, Burt & Rattray 75, 78, 79 (all MO). Panamerican Highway 0.5 km W of Highway 50 turnoff to Ocu, 30 m, Nee 8002 (MO). 2 km NW of Atalaya, Nee 8244 (DUKE, MO, US). m p = n ^ A > ^ hi ~ ~ N © N c “a x 3. Stylosanthes viscosa (L.) Swartz, Prod. Veg. Ind. Occ. 108. 1788. Hedysarum hamatum B viscosa L., Plant. Jamaic. Pug. 20. 1759. Type: Jamaica, Sloane (BM, not seen). Herbs, stems ascending and spreading or prostrate and matted, much branched, to 1 m long, densely pubescent with viscid trichomes. Leaflets lan- ceolate to elliptic, to 15 mm long, 4 mm wide, acute to obtuse, the abaxial surface punctate, reddish, hispidulous, 2—4 pairs of conspicuous nerves; petioles 2.5-5.0 mm long, hispidulous, viscid, the rachis 1-2 mm long; stipular sheath 3.5-5.5 mm long, 3-5 nerved, the teeth subulate, hispidulous, viscid. Inflorescence spicate, 2-5 flowered; outer bracts usually trifoliolate, the inner bracts unifoliolate, the sheath equaling or slightly exceeding the teeth, hispidulous, viscid, 5-7 nerved; outer bracteole 1, 2.5-3.0 mm long, the apex ciliate; axis rudiment none: inner bracteole 1, 2.5-3.0 mm long, the apex ciliate. Flowers yellow, calyx tube 3-7 mm long, glabrous to puberulent; standard suborbiculate, 4-7 mm long, the wings 4-5 mm long, auriculate, spurred within at the base, the keel petals 3-4 mm long, falcate. Fruit a loment, 2-4 mm long, ca. 2 mm wide, the distal articulation fertile, reticulate nerved, puberulent, beak ca. 1.5 mm long, strongly uncinate to coiled. Stylosanthes viscosa ranges from western Mexico through central and north- ern South America to the eastern coast of Brazil. This species is frequently collected in Panama, and it is distinctive with its highly punctate leaf undersurface appearing reddish. cocLe: 3 mi NE of Anton, D'Arcy & Croat 4111 (MO). E of Nata, Duke 12386 (3) (NY). Penonomé, Ebinger 1099a (F, MO). Road to El Valle, Ebinger 1099b (MO). Aguadulce, McDaniel & Cooke 14764 (FSU); Pittier 4983 (GH). Río Hato Military Reserve, Tyson & Blum 2558 (FSU, MO, SCZ). HERRERA: Hills between Las Minas and Pesé, 9-1200 m, Burch et al. 1336 (F, MO). Ocu, ca. 5 m S, Stern et al. 1689 (MO, US). 10 mi S of Oct, Tyson et ul. 2852 (FSU, SCZ). Los SANTOS: Las Cruces to Macaracas, 100-200 m, Burt & Koster 102 (MO). PANAMA: Road to El Valle near Inter- way, 0-1 : (MO). Playa near Río Mar, Ebinger 507 (F, MO). 7 m W of Che epo, Gentry & Tyson 1633 (SCZ). VERAGUAS: Without exact locality. Bristan 795 (MO). La Mesa, con 6049 (FSU, SCZ). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) Hid 43. TEPHROSIA Peter S. White?? Tephrosia Pers.,?' Syn. Pl. 2: 328. 1807. Nomen conserv. contra Needhamia cop., Reineria Moench. LECTOTYPE: T. villosa (L.) Pers. (Cracca villosa L.) Needhamia Scopoli, Introd. 310. 1777. Nomen rejic. contra Tephrosia. TYPE: Vicia littoralis Jacq. — Tephrosia littoralis (Jacq.) Pers. Reineria Moench., Suppl. Meth. Bot. Pl. 44. 1802. Nomen rejic. contra Tephrosia. TYPE: R. reflexa Moench = Tephrosia reflexa (Moench.) DC. Cracca L., Sp. Pl. 752. 1753; Gen. Pl., ed. 5. 1754. Nomen rejic. contra Cracca Benth. TYPE: Cracca villosa L. Not Cracca Hill, Brit. Herb. 285. 1756. TYPE: not designated. Nor Cracca Medic., Vorl. Chrupf. Phys.-Okon. Ges. 2: 359. 1787. TYPE: Cracca benghalensis Medic. = Vicia ben- ghalensis L. Nor Cracca Benth., in Benth. et Oerst. Vidensk. Meddel. Naturhist. Foren. Kjg- benhavn 1853: 8. 1853. Nomen conserv. contra Cracca L. TYPE: Cracca glandulifera Benth. Perennial herbs or shrubs, erect or trailing; base woody; roots usually heavy; pubescence usually close and dense. Leaves alternate, odd pinnate; leaflets (1—)3— 41, almost always hairy, at least beneath; secondary veins parallel, distinctively sharply ascending, ca. 30? to the midrib; rachis usually grooved above; estipellate but tufts of hairs sometimes present in the axils; stipulate. /nflorescence terminal or axillary, sometimes apparently leaf opposed but actually terminal and over- topped by the adjacent axillary branch, racemose, elongate, the flowers in clusters of 2-6 or more at the nodes, each cluster usually with a primary bract at the base and each pedicel with a secondary bract at the base. Flowers red, purple, or white, petals clawed, the standard hairy outside, often densely so, the wings about as long as the standard and usually basally adnate to the keel; stamens usually diadelphous, the vexillary stamen frequently fused to the stamen tube above, free at the base; ovary sessile, slender, usually hairy, the style bearded above in many species, or glabrous. Fruit flat, linear or oblong, straight or slightly curved, sessile, often obliquely contracted distally and beaked on the upper side by the persistent style base, continuous within or slightly septate, the valves usually coiling in dehiscence; seeds several to many, circular to oblong, flattened. Tephrosia is a genus of 250—400 species in temperate and tropical regions of North and South America, Africa, southern Asia, and Australia. It reaches its highest diversity in dry open habitats of Mexico. The four species reported from Panama are also characteristic of open habitats. Many species produce rotenone and related compounds which are used as fish poisons and insecticides, and at least several species have been cultivated in the New World tropics for these purposes. Tephrosia has also been planted as a cover crop and green manure. Chromosomes: 2n — 22 (Wood, 1949). The history of the names Tephrosia Pers., Cracca L., Cracca Benth., and Cracca Medic. may be confusing as presented in the synonymy above and is explained more fully below (see also Wood, 1949). In Species Plantarum (1753), Linnaeus described six species in the genus Cracca L. He subsequently (1759) °° Uplands Field Lab., Great Smoky Mts. National Park, Twin Creeks Area, Gatlinburg, Ten- nessee 37738. s more complete synonymy see Wood (1949). Only the above names relate directly to Panamanian plants. 778 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 submerged this genus in Galega. Soon it was recognized that Galega was a heterogeneous catchall of species, and segregate genera were proposed, one of which was Tephrosia Pers. (1807). At this point Tephrosia included all the original six Cracca L. species plus some other elements. Some of the included elements were later seen as extraneous to Tephrosia and were removed to other genera, leaving Tephrosia Pers. taxonomically equivalent to the original Cracca L. Other species were placed in Tephrosia by several workers. A new use of the name Cracca occurred when Bentham (1853) used the name for some of the original discordant elements in Galega. Confusion was heightened when Otto Kuntze, recognizing the priority of Cracca L., transferred all Te- phrosia Pers. species to Cracca L. For the species of Cracca Benth., Kuntze proposed the genus Brittonamra Kuntze. Yet a third use of the name was Cracca Medic. (1787) which was resurrected by Alefield in 1862 for species of Vicia L. Thankfully, this use of the name was more or less ignored, although Alefield introduced further confusion by proposing the name Benthamantha Alef. for the species of Cracca Benth. Except for Crac- ca Medic., these names were all in current use. Tephrosia Pers. and Cracca L. were in use for one group of plants; Cracca Benth., Brittonamra Kuntze and Benthamantha Alef. were in use for a second group of plants. Resolution oc- curred when Tephrosia Pers. and Cracca Benth. were conserved. Tephrosia Pers. is conserved against Needhamia Scopol. (1777) and Reineria Moench. (1802). Cracca L. is rejected against Cracca Benth., and is unavailable for re- jection against Tephrosia Pers. because Cracca Benth. (1853) is of a later date than Tephrosia Pers. (1807). Hill, in his British Herbal (1756), had also published Cracca (=Vicia L.). Like the later Cracca Medic. (1787; TYPE: Vicia bengha- lensis L.), this use of Cracca was more or less ignored. Cracca Benth. has been confused taxonomically with Tephrosia (=Cracca L.) but in Cracca Benth. the leaflets are stipellate, the wing petals free, the fruit septate, and the flowers often yellowish. Tephrosia is distinctive in the sharp angle of the secondary veins of the leaflets, the hairiness on the outside surface of the standard, and the often bearded style. Literature: Wood, C. E., Jr. 1949. The American barbistyled species of Tephrosia (Legu- minosae). Rhodora 51: 193-231; 233-302; 305-364; 369-384; pls. 1152-1155. a. Low or trailing, weak, herbaceous plants; inflorescences often oe leaf opposed; 3 cm wide style glabrous or only ciliate at the apex; leaflets 1-3 cm long, 0.1-0.3 cm wide... enella w P Upright, stout herbs or shrubs; inflorescence evidently axillary or terminal; style cis above; leaflets 2-8 cm long, 0.5-2.0 cm wide. b. Leaflets 5-13, ren with cuneate base, thick; leaves and hypanthium silvery silky; calyx lobes 3-6 m ng . T. nitens Leaflets 15-41, narrowly elliptic, the base thinner; leaves and hypanthium with usually duis to brownish hairs; calyx lobes less than 3 mm long. Lateral lobes of calyx ais and abruptly acute or acuminate, unlike the lower lobe in shape s aa aaa, . T. dia Lateral lobes of calyx acute to acuminate, symmetrically tapered e appro mately the same shape as the lower lobe _ — . |. . T. multifolia c d [e] a 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 779 1. Tephrosia multifolia Rose, Contr. U.S. Natl. Herb. 1: 320. 1895. rype: Mexico, Palmer 1364 (US, holotype; isotypes GH, NY). Cracca multifolia ea. Rose, Contr. U.S. Natl. Herb. 12: 270. 1909. C. arcuata Rydb., No. Amer. Flora 24: 166. 1923. Type: Mexico, Nelson 4193 (US, holotype; isotype Ç. heydeana Rydb., No. Amer. Flora 24: 166. 1923. Type: Guatemala, Heyde & Lux (D. Smith dist. 6111) (NY, holotype; isotypes GH, US). Tephrosia heydeana (Rydb.) Standley, J. Wash. Acad. Sci. 17: 167. 1927. T. arcuata (Rydb.) Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 213. 1929. Shrubs or erect herbs to 2.5 m tall, much branched; stems, leaves, and inflo- rescence pubescent, pilose, or velvety. Leaves to 35 cm long; leaflets (15—)21— 31(-40), linear, oblong or narrowly elliptic, 2-6 cm long, 0.5-1.2 cm wide, the middle leaflets often the longest, rounded to somewhat acute basally and apically, mucronulate, paler green and more densely pubescent beneath, the main second- ary veins 10-15 on each side, these and the midrib slightly raised beneath; petiole 1.5-3.0 cm long; rachis grooved, 5-25 cm long. Inflorescences terminal and ax- illary, to 20 cm long; peduncle 2-9 cm long, the nodes 20-50, buds to 7 per node; primary bracts 5-10 mm long; secondary bracts 4 mm long; pedicels 4 mm long, slender. Flowers drying to 15 mm long; calyx strigose, 3-6 mm long, the lower and 2 lateral teeth subequal, narrowly triangular to acuminate, the upper teeth connate, shorter, the teeth approximately equaling the tube; petals clawed, ap- parently white to pink, the standard circular to oblong, ca. 15 mm long, the claw to 3 mm long, densely covered with brown appressed hairs on the back, the wings narrow, oblong, ca. 15 mm long, the claw to 3 mm long, slightly adherent to the keel; keel narrow, oblong, to 5 mm wide, the claw to 3 mm long; staminal tube 10-15 mm long, the vexillary stamen connate in part, basally free; ovary silky, linear, ovules 8-10, the style hairy above. Fruit 5-6 cm long, 5 mm wide, widest toward the tip, flattened, slightly upcurved, velvety, the style base persistent, calyx persistent but somewhat torn, the 2 valves twisting on dehiscence; seeds 6-10, brown, oblong, 3 mm long. Tephrosia multifolia is found from Mexico southward throughout Central America, but is apparently absent from South America. It has been collected in open habitats in three provinces of Panama. Tephrosia multifolia is similar in appearance and apparently closely related to T. sinapou. The most easily differentiated character separating the two is the shape of the lateral calyx lobes which are symmetrical and narrowly triangular in T. multifolia and oblique, asymmetrical, and abruptly acute in T. sinapou. The latter also tends to have narrower leaflets with more secondary veins, pubescence more densely velvety and browner in color, and the calyx larger and heavier in appearance. Both species are cultivated for use as fish poisons and insecticides. CANAL ZONE: Chiva-Chiva Trail, Red Tank to Pueblo Nuevo, Vo 5125 (US). cw cA Near Las Lomas, 0-100 m, Burt & Koster 129 (MO). Boquete, Davidson 591 (F, GH, MO, US). Between Hato del Jobo and Cerro Vaca, 700-1000 m, Pittier 5419 (GH, NY, US). HERRERA: Nea r Las Minas, 320 m, Burt & Koster 92 (MO). VERAGUAS: Tienda Santa Ana (La Yeguada), Escobar et al. 291 (MO). El Embalsadero, 8 mi W of Santiago, Tyson 6085 (FSU, MO, SCZ). 780 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 67 9. Sa Ss - = "7205742 MA Nee oe < i p< “3 eM P ntt 5 z = x NA x ir 24 ae M Nm FiGURE 48. Tephrosia nitens Seem.—A. Habit ( x24).—B. Flower (x2). 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 781 2. Tephrosia nitens Benth. ex Seem., Bot. Voy. Herald 107. 1853. TYPE: Panama, Seemann 1036 (K, not seen; photo F, MO, NY; possible isotype at GH).— IG Cracca nitens (Benth. ex Seem.) Kuntze, Rev. Gen. Pl. 1: 175. 1891. Tephrosia nitens var. lanata Micheli in Dur. and Pitt aa Bot. Soc. a 30(1): 286. 1891. Tephrosia albida Brandegee, Univ. Calif. Publ. Bot. 10: 406. 1924. TYPE: tea M PU 9136 (US, not seen). Erect herb or sparsely branched shrub to 3 m tall; stems strigose to villous. Leaves usually 5-10 cm long, leaflets 5-13, ascending, 2-8 cm long, 0.5-2.0 cm wide, the terminal leaflet largest, oblong, retuse, the base cuneate, thick, glabrate above, silvery silky beneath; petiolules 2-3 mm long; petiole grooved, 3-5 mm long; rachis grooved, 1-5 cm long; stipules narrow to triangular, deciduous to apparently persistent, to 8 mm long. /nflorescences terminal and axillary, to 50 cm long, flowers 5-7 per node, apparently never all maturing fruit, nodes to 50 per inflorescence; bracts narrow, deciduous; pedicels to 8 mm long. Flowers drying 1.5-2.0 cm long; calyx 6-9 mm long, the lobes long acuminate, subequal, 4-7 mm long; petals white to rose purple, the standard oblong to circular, 15-20 mm long, 15 mm wide, silky on the back, often notched at the tip, the claw to 3 mm long, the wings oblong, to 2 cm long and 4 mm wide, the claw to 4 mm long, the keel to 15 mm long, the claw to 4 mm long; staminal tube 10—15 mm long, the vexillary stamen connate in part, free at the base; ovary silky, to 15 mm long and 1 mm wide, the margins thickened. Fruit linear, flattened, 5-6 cm long, 0.5 cm wide, evenly tapered to the persistent style base, not septate but the flat surface often indented between the seeds, pubescent; seeds 9-13, nearly circular, com- pressed, ca. 2.5 mm across, dark brown. Tephrosia nitens is found from southern Mexico to Brazil. This is a distinctive and attractive species both in flower and in leaf. All but one of the Panamanian collections seen are from Taboga Island, which is the type locality. Chromo- somes: 2n = 22 (Wood, 1949). MA: Taboga Island, Allen 110 (GH, MO, NY), /276 (GH, MO, US); Killip 3174 (US); PAN aerie 2830 (F, US); McDaniel & Cooke 12903 (FSU); Miller 2026 (US); Pittier 3571 (NY, US); Standley 27999 (US). Cerro Campana, ca. 2500 ft, Tyson 6439 (MO, SCZ) . Tephrosia sinapou (Buchoz) A. Chev., Compt. Rend. Hebd. Séances Acad. Sci. 180: 1522. 1925. (As T. singapou.) Galega sinapou Buchoz, Hist. Univ. Regne Vég., pl. 994. 1775. TYPE: Buchoz diagnostic plate G. sericea Lam., Encycl. Méth. Bot. 2 (part 2): 596. 1788. TYPE: not established. Not Galega sericea m Karan G. sericea Buch.-Ham. (1822), nor Tephrosia sericea (Thunb.) Pers. (1807), oricea Baker 1871). or 7. G. us Saat , Prod. Veg. Ind. Occ. 108. rhe i West Indies, Swartz, not seen. Tephrosia d s (Swartz Pers., Syn. Pl. 2: 329. 1807. T. emarginata H. ., Nov. Gen. Sp. Pl., (ed. P 6: 361; (ed. quarto) 6: 461. 1824. TYPE: Vene- uela, (Pn t seen; microfiche MO; probable isotype B, not seen T. respi Schlecht., Linnaea 12: 299. 1838. TYPE: Mexico, (NY, nui seen; isotype GH). Orobus sericeus Sess. et Moc., santa aa E 1: app. 118. 1889. Type: Mexico, not seen. Cracca toxicaria (Swartz) Kuntze, Rev. Gen. Pl. 1: . 1891. C. AUR (Schlecht) Standley, Contr. U.S. a Herb. 23: 474. 1922. 782 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Shrubs or erect perennial herbs to 1 m tall; stems usually ridged and densely velvety with yellowish-brownish pubescence. Leaves to 30 cm long; petioles 2— 5 cm long; rachis 10-25 cm long; leaflets 17-41, most often 25-35, narrowly elliptic to linear, mucronate, 4-6 cm long, 0.5-1.5 cm wide, pubescent above, more densely pubescent and paler beneath; leaflets with the 15-20 pairs of main side nerves sometimes obscured by pubescence beneath: petiolules ca. 2 mm long; stipules linear, to 15 mm long, apparently persistent. Inflorescences terminal and axillary, to 30 cm long, the peduncle 2-8 cm long, the nodes 15-25 or more, the buds 2-6 per node; bracts linear, to 8 mm long; pedicels 2-5 mm long. Flowers drying to 20 mm long; calyx densely hairy, 5-8 mm long, the lobes about equaling the tube, the lower lobe symmetrically tapered, narrow triangular, the lateral lobes unlike the lower lobe, abruptly and obliquely narrowed to an acute or sometimes acuminate tip; petals apparently white to pink, the standard circular to oblong, ca. 15 mm long, densely hairy on the back, the claw to 3 mm long, the wings narrow, to 15 mm long, the claw 3 mm long, slightly adherent to the keel, the keel narrow, oblong, to 17 mm long, the claw to 4 mm long; staminal tube 10-15 mm long, the vexillary stamen adnate to the tube for part of the length, basally free; ovary silky, linear, the ovules 10—13, the style hairy above. Fruit more or less straight, linear, to 5 cm long and 5 mm wide, hairy; calyx persistent, but somewhat torn; seeds 8-13, oblong, 3.0-3.5 mm long, brown, apparently often faintly mottled. This plant is similar in appearance to T. multifolia (see above under T. mul- tifolia). T. sinapou is found in Mexico, Central America, the West Indies, and South America. T. sinapou has been cultivated for use as a fish poison apparently throughout the New World tropics. The type of Galega sericea Lam. is possibly one of two collections in the Lamarck Herbarium (P-LAM, not seen; microfiche, MO). Lamarck (Encycl. 2 [part 2]: 596) also lists a variety "beta" and refers to having seen a Jussieu specimen. This latter may be the collection that bears the name “Galega sericea” in the Jussieu Herbarium (P-JU; microfiche, MO). The plates in Buchoz's (also spelled Bu'choz and Buch'oz) work are not indexed. They are numbered as follows: "mille" (Arabic numerals, lower left corner); Cent." (Arabic numerals, lower right corner); "Dec." (Arabic numer- als, upper right corner); and Pl." (Roman numerals, upper left corner). There is no ‘‘Cent.” zero nor a "Dec." zero; hence, plate 994, on which the name Tephrosia sinapou is based, is numbered as "Cent. 10," “Dec. 10.," "Pl. IV." This system can be confusing in trying to locate a particular plate. RIÉN: El Real, Duke 4941 (MO). ae St. Dorothea, Dwyer 2274 (GH, MO, SCZ). SAN BLAS: Ailigandí River, Duke & Bristan 338 (MO, NY). 4. Tephrosia tenella A. Gray, Pl. Wright. 2: 36. 1853. TYPE: “San Pedro, Sonora,” now Arizona, Wright 966 (GH holotype; isotypes MO, US). Weak, thin stemmed, trailing herbs with sparse pubescence. Leaves 2-8 cm long; leaflets 5—13, rarely to 21, linear to linear oblong, 1-3 cm long, 1.0-2.5 mm wide, rounded and mucronate at the tip, acute at the base, the secondary veins ascending and inconspicuous; rachis to 6 cm long, grooved, loosely strigose; 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 783 petiolules ca. 0.5 mm long; petioles 0.5-2.0 cm long, grooved, strigose, estipellate but tufts of hairs present in the axils; stipules subulate, ca. 1-3 mm long. /nflo- rescence opposite the leaves or axillary, to 15 cm long, loosely hairy, the nodes distant, 1-2 flowers or fruits per node; bracts ca. 2 mm long, subulate; pedicels 3-5 mm long. Flowers drying to 10 mm long, the calyx 4-5 mm long, the tube campanulate and somewhat longer than the narrowly triangular lobes; standard circular to reniform, 10-12 mm long, puberulent, the claw to 3 mm long, the wings oblong, to 11 mm long and 5 mm wide, longer than the keel, the claw ca. V? the length of the blade, slightly adherent to the keel, the keel to 7.5 mm long, ca. 3.5 mm wide; staminal tube ca. 5.5 mm long, vexillary stamen united briefly to the tube and basally dilated to ca. 1 mm wide; ovary narrowly oblong, short appressed hairy, the style to 3 mm long, glabrous except ciliate towards the apex. Fruit linear oblong, flattened, 3-5 cm long, ca. 0.5 cm wide, pubescent, the valves twisting on dehiscence; calyx persistent; seeds 3-8, oblong, ca. 3 mm long. This species has been reported from the southern United States, Mexico, Central America, and the West Indies. The taxonomy of the glabrous styled Tephrosias awaits monographic study. Several names have been used in this group, at least some of which refer to distinct species. Only a more detailed study than that possible here will elucidate the relationship of this taxon to Tephrosia ascendens (1837); T. brandegei (1923); T. corallicola (1951); T. curtissii (1962); T. leptostachya (1825); T. littoralis (1807); T. vicioides (1838); and T. wallichii (1831). Tephrosia tenella apparently sometimes behaves as an annual. It is unique among Panamanian Tephrosias in this regard. COCLÉ: Ola, 100-350 m, Pittier 5044 (GH, US). HERRERA: Ocú, Ebinger 1079 (MO, SCZ, US). ~ 44. TERAMNUS W. G. D'Arcy” Teramnus P. Br., Civ. Nat. Hist. Jam. 290. 1756. TYPE: T. uncinatus (L.) Swartz. Twining vines, mostly pubescent. Leaves pinnate trifoliolate, the leaflets mostly broadly or narrowly ovate, mucronate; stipels subulate; stipules narrow, striate. Inflorescences axillary racemes; peduncles slender, often elongate, sev- eral flowered; bracteoles 2, subtending the calyx or just below the top of the pedicel; pedicels short, pubescent. Flowers inconspicuous; calyx tubular, lobed about 1⁄2 way down, the lobes subulate, subequal or the upper pair fused and longer; standard emarginate, glabrous, exserted slightly from the upper calyx lobes; stamens pseudomonadelphous, the alternating anthers reduced and sterile; ovary linear, pubescent, the style incurved, glabrous, the stigma capitate. Le- gume linear, mostly straight, sometimes compressed, with a small but prominent often upturned beak, mostly pubescent, explosively dehiscent by 2 valves; seeds numerous, separated by valvular tissue, plump, lenticular. Teramnus is distinguished by its small flowers with subulate-toothed calyx and glabrous standard. The anthers, only 5 fertile, are also distinctive. The genus 92 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. 784 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 includes about 8 species of both Old and New World tropics. Only three species are known in the Neotropics, all three of which occur in Panama. These are mainly plants of recent disturbance, although they may persist as weeds around gardens. Teramnus rhombifolius Beurling, Kongl. Vetensk. Acad. Nya Handl. 1854: 120. 1855, was described from a collection taken near Portobelo by Billberg. This collection has not been located, and the species cannot be determined from the original description. It may be synonymous with either T. uncinatus or T. labialis. a. Leaflets densely sericeous beneath; fruits densely long villous, the beak more than 2.5 m MN aa AE a a ce ea T. nai aa. Leaflets sparsely pubescent beneath, sometimes appressed pubescent; fruits with thin, short pubescence, the beak mostly less than 2.5 mm lon b. Leaflets mostly ovate, only 1.5 times longer than wide; calyx wide 5 neri bap ara . labialis bb. Leaflets ‘mostly 1 narrow, 1,20 or more times longer than wide: ‘calyx with the ae n pair of teeth fused to near the top or higher 3. T. volubilis 1. Teramnus labialis (L.f.) Spreng., Syst. Veg. 3: 235. 1826. did sas L.f., Suppl. Pl. 325. 1781. Lectotype: East Indies, herb Linn (LINN 901.15, not ; microfiche MO). Twining vines climbing on herbs and shrubs, the stems hispid, glabrescent. Leaves pinnate trifoliolate; leaflets ovate, apically acute or obtuse, mucronate, basally rounded, mostly 2-6 cm long, 1.5-2.5 cm wide, mostly copiously pubes- cent on both sides, more so beneath, the hairs sometimes appressed; petiolules 1-2 mm long, hispid; stipels subulate, the stipules subulate, sparingly long hispid. Inflorescences axillary racemes, mostly short, 2—5(—12) cm long, peduncle slen- der, retrorsely pilose, glabrescent in fruit; bracteoles ca. 2 mm long, about as long as the calyx tube. Flowers inconspicuous, white or pinkish; calyx tubular, ascending hispid, the tube ca. 2 mm long, the teeth subequal, about equalling the tube, the standard exserted about 1 mm from the calyx teeth. Legume linear, somewhat compressed, 3-5 cm long, mostly 5-6 mm wide, sericeous; seeds tan with dark mottling, plump lenticular, 4 mm long. This species is similar vegetatively to Teramnus volubilis, but the leaves are usually smaller and broader. The calyx is different. Teramnus labialis is widespread in the Caribbean region, and it is naturalized in the Old World. It appears to be uncommon in Panama. CANAL ZONE: Albrook site, U.S. Army Tropic Test Center, Dwyer 7118 (GH). cHiRIQUÍ: Nea ar Cuesta de Piedras, 730 m, Burt & Koster 161 (SCZ). Los SANTOS: Monagre Beach 5 mi SE of Chitré, Tyson et al. 3009 (SCZ). 2. Teramnus uncinatus (L.) Swartz, Nov. Gen. Sp. Pl. 105. 1788.—Fic. 49. Dolichos uncinatus L., Sp. ed. 2. 1019. 1763. TvPE: not se id^ eis hacophiehm ‘Donnell Smith, Bot. Gaz. 52: 46. 1911. TYPE: Mexico, Heyde & Lux 3742, Long twining vines, mostly on shrubs and herbs; stems brownish to greyish sericeous with elongate and shorter hairs, the shorter hairs reflexed, glabrescent. 1980] DWYER & COLLABORATORS—FLORA OF PANAMA (Family 83. Leguminosae) 785 FIGURE 49. Teramnus uncinatus (L.) Sw. Habit ( a. . Averrhoa bilimbi L., Sp. Pl. 428. 1753. TYPE: Ceylon, Hermann ex herb. Banks no. 177, 1672—1677 (BM). Tree to 15 m tall, 30 cm diam.; branches ascending, new branches and rachis densely yellow or rusty tomentose. Leaves pseudoverticillate at the top of the branches, 7—20 jugate, ovate to 65 cm long; leaflets alike, discolorous, oblong or elliptical, asymmetrical, 2.5—15 cm long, 1-5 cm wide, the lower leaflets smaller, 1-2 cm long 1-1.5 cm wide, acute or acuminate with a mucro 2 mm long, the base rounded or subcuneate, softly pubescent on both sides, more so beneath and above on the midrib; petiolules fleshy, tomentose; petiole pubescent, to 17 cm long, enlarged at the base. /nflorescences axillary or often cauliflorous on aborted branches, sometimes even on the lower part of the trunk, of cymes forming racemes or panicles, 15-20-flowered, or of solitary flowers, pubescent and glandular; bracts ca. 4 mm long, bracteoles 1.5-2 mm long, both subulate, densely tomentose; pedicels 4-20 mm long, articulate near the middle. Flowers with the sepals 5-2.5 mm long, 1.5-3 mm wide, ovate to elliptical, asymmetrical, acute, mucronate, one internal sepal oblong and obtuse, pubescence appressed and glandular inside and out, green yellow to purplish; petals purplish, free or sometimes connate near the middle, linear spathulate, 10-20 mm long, 2.5-4 mm wide, clawed, caducous, apically with a short caducous cilium and minute glan- dular hairs; stamens 10, nearly free, glabrous, the longer 10 mm and the shorter 4 mm long, the anthers suborbicular, the connective conspicuous; pistils mostly mesostylous, 7.5-12 mm long, the ovary cylindrical, 5-lobate, 4-7.5 mm long, acute, lengthened into the styles, the styles cylindrical, ca. 2 mm long, 2-lobed or truncate, densely appressed pilose, the hairs yellow, straight, ascending, the stigma inconspicuous, liguliform or subulate, the carpels with (1—3—)4—7 ovules. Berry oblong, to 7.5 cm long, 3.5 cm in diam., 5-lobate or smooth, in transverse section subcircular or pentagonal, the base roundish, the apex lobulate; seeds 4— 7 per carpel. 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 825 FIGURE 1. Averrhoa carambola L. Habit (x15). [After Chi-en Chang 8567, Taiwan.] Averrhoa bilimbi is a widespread tree in tropical regions. Frequent in the Orient, in America it is known only in botanical gardens and experiment stations. CANAL ZONE: Summit Garden, Steyermark 6, 7, 17.1.1935 (MO). 2. Averrhoa carambola L., Sp. Pl. 428. 1753. TYPE: Ceylon, Hermann ex herb. Banks no. 178, 1672-1677 (BM).—Fic. 1 826 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Tree to 25 m tall, 25 cm diam., abundantly branched; branches sometimes drooping; pubescence of short curved whitish to reddish ascending deciduous airs. Leaves disposed along the branches, 3-8 jugate, the leaflets alternate or subopposite, discolorous, variable in shape, asymmetric, the upper leaflets ellip- tical or ovate, oblong, cuspidate, basally rounded or truncate, the lower leaflets smaller, 1.5-3 cm long, 1-2 cm wide, acuminate or cuspidate, the secondary nerves 4—10 pairs, the pubescence short and even beneath, the nerves pubescent above, the margin appressed ciliate; rachis to 20 cm long, rigid, pubescent; petio- lules thick, 1.5-3 mm long; petioles to 3.5 cm long, pubescent, thickened towards the base. Inflorescences axillary or cauliflorous, of pubescent and glandular cymes united into racemes and these into fascicles or pseudo-panicles; peduncle to 1 cm long; pedicels 3-4 mm long. Flowers with the sepals oblong or obovate oblong, 2.5-3.5 mm long, 1-2 mm wide, obtuse and somewhat emarginate, rarely subacute, a few hairs appressed on the central zone, the margin irregular, hyaline, finely ciliate; petals violaceous, white bordered, linear spatulate, 6.5-9 mm long, 1.5-3 mm wide, clawed, connate at the middle, seldom free, densely glandular inside; stamens enlarged toward the base, connate up to '/;, the anthers orbicular, the connective conspicuous, the longer stamens 2-2.5 mm long, fertile, the short- er stamens without anthers, seldom with rudimentary anthers (D'Arcy & D'Arcy 6799 and others), or rarely with 2 rudimentary thecas separated by a short con- nective, in macrostylous flowers the longer stamens with small glands on the upper half, in mesostylous flowers (most frequent) the short stamens very short, microstylous flowers not seen; pistils 3-4 mm long, sometimes glandular, the ovary ellipsoid, the pubescence scant, appressed, sometimes only on the back of the carpels, the carpels 3-5-ovuled, the styles short, thick, pilose, the stigma enlarged, 2-lobed. Berry greenish or yellow, ovoid or ellipsoid, asymmetric, 8— 12.5 cm long, 5-6 cm wide, the carpels uneven, angular, the base and apex 5- lobed, in transverse section 5-rayed; seeds without outer integument. This species is frequently cultivated in all tropical regions. It probably origi- nated in the Orient and was introduced into America. Fruits are consumed fresh or in conserves, also as refreshments. ZONE: Barro Colorado Island, Foster 2190 (US). PANAMA: Tobago Island, D'Arcy & D' Te 6709 (MO). 2. BIOPHYTUM Biophytum DC., Prodr. 1: 140. 1824. LECTOTYPE: B. sensitivum (L.) DC. Oxalis L. sect. Biophytum Endl., Gen. 1172. 1840. Toddavaddia Kuntze, Rev. Gen. Pl. 1: 96. 1891. Herbs or subshrubs, erect or decumbent, the base sometimes ligneous; stems generally bearing a ring of rigid, retrorse pubescene at the apex. Leaves fascic- ulate at the stem apex, imparipinnate, the terminal leaflet reduced to a bristle- like mucro; leaflets subsessile, the pairs gradually modified in shape and size upwards, the lower pairs usually subtriangular or ovate, smaller, petiolule minute, broader than long, fleshy; petiole short, thick, articulating at the rachis, the bases persistent on the stem. Inflorescences of cymes, globose or cylindric; peduncu- 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 827 late; bracts subulate, situated at the insertion of the peduncles; bracteoles spirally overlapping; pedicels shorter or longer than the bracteoles, articulate near the base, the flowers opening successively, often only 1 or 2 to several seen opened to different degrees. Flowers white to red; sepals almost free, imbricate; petals connate near the middle; stamens short-connate at the base, all fertile or some- times the shorter stamens reduced and sterile; ovary 5-lobed, the carpels 2—6- ovulate, the ovules in two rows alternating in length, the stigmas terminal, en- larged, bifid, subcapitate, papillose. Fruit a capsule covered by the calyx, the carpels 1—6-seeded, loculicidally dehiscent, the valves opening flat over the sepals in a star shape, remaining attached at the base; seeds 2-tegmic, the outer tegument opening and ejaculating the seed at maturity. Biophytum is a pantropical genus of about 75 species. Five species occur in Panama, 3 of which are endemic. a. Cymes glomeruliform or fusiform, 1-2 mm diameter; few-bracteolate. b. Pubescence hirsute, the hairs with thickened bases; bracteoles long se L eRe B. dendroides bb. Pubescence very short, more or less appressed; bracteoles appressed a . B. soukupii aa ren globose or cylidroid, ca. 5 mm diameter; bracteoles subtriangular, keeled. . A Peduncles ca. 10 cm long; calyx and capsule glabrous ___ 3. B. mucronatum cc. Peduncles to 2 cm long; calyx pubescent and glandular, the capsule pubescent at the apex d. ‘Leaflets oblong, the distal border obtuse, mucronate; petals white, the throat yellow 4. anamense dd. Leaflets falcate, acute, mucronate; petals yellow, red-veined ------ 2. B. falcifolium 1. Biophytum dendroides (H.B.K.) DC., Prodr. 1: 690. 1824.—Fic. 2. Oxalis dendroides H.B.K., Nov. Gen. Sp. Pl. 5: Fes deir TYPE: Colombia, near Quebrada de la arbonera, Bonpland 1744 (P, holotype: B, P, iso O. viva Willd. ex Steud., Nom. ed. 2. e i s 1844. TYPE: Colombia, Bonpland 1744 (B, holotype; P, otypes), m. illeg. incl. typ. sp- Toddavaddia Veni (H.B.K.) Ph Tek Gen. Pl. 1: 96. 1891 Biophytum mexicanum KI. ex Knuth, Pflanzenr. 130: 401. 1930, nom. $a in syn. B. longiscapum KI. ex Knuth, Pflanzenr. 130: x 1930, nom. nud. Oxalis dendroides fo. longiscapa Zucc. ex Knuth, Pflanzenr. 130: 140. “1930, nom. nud. in syn. Biophytum mimosoides sensu auct., non (St. Hil. i Guillaumin B. dormiens sensu auct., non Mart. & Zucc Rhizomatous herb, stems obsolete or to 35 cm tall and woody, to 5 cm diam.; rooting up to 20 cm from the base, sometimes 2- or 4-branched; bark rough, appressed retrorse pubescent, glabrescent; root fibrous, to 10 cm long, twisted, branched. Leaves pseudoverticillate at the end of the stems, 15-30, 9-26 jugate, linear in outline, subcuneate; lower leaflets ovate to subtriangular, asymmetric, 1-2.5 mm long, 1-2.5 mm wide, the middle leaflets subrectangular to subfalcate, 2.5-10 mm long, 1.5-7 mm wide, the terminal leaflets semiobovate, asymmetric, basally truncate to subhastate, 6-11 mm long, 2.5-5 mm wide, the leaflets some- times overlapping, glabrous or sparsely hirsute on one or both sides, hirsute ciliate, obtuse or subacute, the mucro sometimes violaceous, the midrib medial or emerging from nearer the posterior angle, secondary nerves ascending, curved, branched, connected to a marginal nerve; rachis 2-10 cm long, hirsute, more so at the insertion of the leaflets; petiolules 1⁄4 mm long, thick, fleshy; petiole 1 mm 828 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 kus QU Wt AM ETE > UE NS Uu P) , SS ⁄ > uL —227 K Z FIGURE2. Biophytum dendroides (H.B.K.) DC.—A. Habit (x 1).—B. Middle leaflet, upper and lower surfaces (x5).—C. Terminal leaflets (x3).—D. Inner bracteole (x6).—E. Outer bracteole (<6).—F. Fruit (x6).—G. Fruit in dehiscence (x6). [After a plant cultivated in Paris and studied by Lourteig. ] 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 829 long, fleshy. Inflorescences subsessile or pedunculate, shortened, glomeruliform or fusiform, only 1 flower appearing at a time; peduncle erect, 1-8 cm long, subappressed pubescent or hirsute, the apex thickened, densely pilose; bracts subulate, to 6 mm long, purplish, pubescent; bracteoles 2-7 mm long, 0.5-1. mm wide, triangular linear to linear, the inner bracteoles prominently keeled, the keels somewhat lengthened, ascending ciliate; exceeding or about equalling ihe pedicels; pedicels 1-5 mm long, articulate in the lower half, glabrous or finely pubescent. Flowers with the sepals linear, 5-7 mm long, 1-1.5 mm wide, acu- minate, 7—10-nerved, glabrous or ascending pubescent, the base densely hirsute, the apex ciliate; petals whitish to pink or lilac, the base obovate or spathulate, 1.5-1.3 times the calyx length, slightly retuse, the base clawed, white; stamens with the filaments basally enlarged, glabrous, the longer stamens 4.5-5.5 mm long, the shorter stamens 3-4 mm long; pistils micro- and macrostylous, the ovary apically pilose, the carpels 1—6-ovuled, the styles sometimes unequal, pilose be- low, the stigma small, bifid. Capsule subglobose or oblong, 2.5-4 mm long, yel- lowish to violaceous, scarious, apically pubescent, or rarely pubescent overall, exceeded 1.75-2 times by the calyx, the carpels 1—-6-seeded, short pubescent within; seeds semiovoid-triquetrous, one face more or less plane with an acute margin and the opposite face convex, 1.5 mm long, acute, longitudinally tuber- culate. This species ranges along the American cordillera from Mexico to Ecuador. It has been introduced with greenhouse plants to botanic gardens on other con- tinents, in some cases becoming subspontaneous, for example in Malasia and Brazil. However, records from Knuth (1930) for Brazil are not this species. CocLE: Mountain beyond La Pintada, 400-600 m, Hunter & Allen 615 (F, GH, MO, NY, P, U, US). 2. Biophytum falcifolium Lourt.? TYPE: Panama, Folsom & Collins 5609 (MO, holotype; P, isotype).—Fic. 3A. Erect or decumbent herbs to 20 cm long; stems single or basally branched, with curved, short pubescence, glabrescent; root fibrous, branched. Leaves 9-20 pseudoverticillate at the end of the stem, linear subcuneate, to 15 cm long, 1.6 cm wide, 13-34 jugate; lower leaflets ovate or subtriangular, ca. 3 mm long, 1.5 mm wide, the middle leaflets falcate, 5.6-11 mm long, 2.5-4 mm wide, basally the posterior side curved, the anterior side slightly curved or truncate at the midrib, central or emerging from nearer the posterior angle, the terminal leaflets elliptical, asymmetric, about equalling or shorter than the middle leaflets, the leaflets opposite or subopposite, subacute, the mucro 0.1-0.2 mm long, fine pu- bescent on both sides or glabrous or glabrescent above, ciliate, the minor venation reticulate; petiole ca. 1 cm long, fleshy; rachis narrowly winged, minutely pu- bescent above, hirsute beneath; petiolules wider than long, 0.1—0.2 mm long, fleshy. Inflorescences 1-3, globose or cylindric, 1—3(—5) flowers seen at different degrees of opening; peduncles 0-7 mm long, violaceous, glabrous or minutely 3 Biophytum falcifolium Lourt. spec. nov. Biophytum panamense Lourt. simile, differt forma foliolorum, colore petalorum (fide collectores), etiam bracteolis pubescentiaque. 830 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 BRACTS FRUITS i Gi Insertion FIGURE 3. Biophytum, diagnostic plate.—A. Biophytum falcifolium Lourt. [After Collins 6500.|—B. Biophytum panamense Lourt. [After Hammel 2528.]—C. Biophytum soukupii Lourt. [After Williams 765; Bristan 1322.]—D. Biophytum mucronatum Lourt. [After Allen 3775.] Stems and leaflets (folioles) (x4), bracts and fruits (x10), calyces (x10) except B. Biophytum panamense (x6). pubescent; bracts subulate, ca. 2.5 mm long, purplish, situated at the insertion of the peduncle, sparsely short pilose; bracteoles spirally overlapping, green, basally violaceous, narrowly triangular, 1.5-3 mm long, l-nerved, strongly keeled, short pubescent and glandular, the hairs I-several celled; pedicels 7-15 mm long, violaceous, fine pubescent. Flowers with sepals linear-elliptic, sharp, 4.5-7 mm long, 0.7-1.2 mm wide, 5—7-nerved, densely glandular, the hairs 1— 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 831 several celled; petals cream, pinkish veined, oblong spatulate, the base clawed, 1.5 times the calyx length; longer stamens unequal, 3—3.5 mm long, white, flat with a few glandular hairs; the anthers ovoid with cordate base, the shorter stamens 1-1.5 mm with sterile anthers, or the entire stamen becoming a glandular relic; pistil equalling or shorter than the long stamens, 2.5-3.5 mm long, the style glandular pubescent, the stigma bifid, the ovary 5-lobulate, apically pilose, the carpels 3-4-ovuled. Capsule globose, ca. 3.5 mm long, shorter than the calyx, apically glandular pubescent, the carpels 3-seeded; seeds too immature to de- scribe. This species is closely allied to Biophytum panamense but is distinguished by the shape of the leaflets and the white petals. Also the bracteoles and the pubes- cence are different. The epithet refers to the shape of the leaflets, curved like a sickle. cocLÉ: Rivera sawmill, Alto Calvario, 7 km N of El Copé, 700-850 m, Folsom & Collins 6509 (MO, P). "Trai from Río San Juan to Río Tife Falls, 1200-2500 ft, Hammel 3352A (MO). 3. Biophytum mucronatum Lourt.* TYPE: Panama, Allen 3775 (US, holotype; F, G, MO, isotypes).—Fic. 3D. Decumbent herb or shrub; stems woody, 50 cm long, 3-4 cm diam., finely short pubescent, glabrescent, decumbent, rooting at the base. Leaves 20—22, pseudoverticillate at the end of the stem, linear subcuneate, to 14 cm long, 2.7 cm wide, 7-27 jugate; lower leaflets suborbicular or ovate, 2-3.5 mm long, the middle leaflets oblong rectangular, oblique, 7-11 mm long, 3-5 mm wide, basally truncate or the posterior side curved, secondary nerves 5-7 pairs, prominent beneath, the terminal leaflets elliptic, 8.5210 mm long, 2.5-5 mm wide, basally truncate, curved on the posterior side, obliquely truncate on the anterior side, 4 Biophytum mucronatum Lourt., spec. nov. Radix fibroso ramosa. Caulis onus (usque 30 cm altus, 3-4 mm diam.), decumbens, basi radicans, pubescentia brevi tenuique, glabrescens. Apex caulis incrassatus, cum annulo saa pilifero, pilis rigidis, retrorsis, pei foliorum inse ionem posito. Caulis supra annulum pilorum foliorum delapsorum cicatricibus instructus. Folia 20-22, pseudoverticillata, in apice caulis disposita, ambitu lineari-subcuneato (usque rere 2.7 cm) 7-27-juga imparipinnata cum mucrone — Petiolus 1-3 cm longus, carnosus. Rachis oe hirsuto-pilosus. Petiolulus 1⁄4 mm longus, carnosus. Foliola opposita, penne ee antice c endo majora, discolora, supra eee qs tenui, adpressa (pilis ca. 1⁄2 mm longis) subtus evi pilosa, glabrescentia; apice insigniter mucronat o (mucro 4% mm lon poe pr a minima, subo rbiculata vel ovata (2-3.5 mm longa); m m oblidi e ida ies (7-1 -5 mm), 5-7 jugis secundariorum nervorum subtus prominentium (paria inferiora parallela, superiora i allela), e truncata vel dimidio eger h plus vel minus curvo et dimidio anteriore obliqu mma elliptica (8.5—10.5 x 2.5-5 mm), basi lobu- lato-curvata in dimidio posteriore, obliquo- ate in in dimi dio anteriore. Cymae 2-5, Jue. glo- boso-congestae (ca. 5 mm diam.) 1-7 floribus adspectabilibus, vicissim apertis. Pedunculi erecti, longi acutissima ( | mm), manifeste (7—)9-11-nervata, glabra. Petala rosea aa ie ), oblong spathulata, basi du Petalorum longitudo et latitude ignotae (flos non apert st). St carina filamentis flavis: longiora (ca. 2 mm) pauci glanduloso-pilosa, antheris ovoideis; hea filamento lato in dimidio inferiora, sursum filiformi, anthera reducta (sterilis?). Pistillum plus minusve aequale longioribus staminibus; stigmata minuta, vix bi-capitata, papillosa; ovarium ellipsoideum, 5-lobulatum, carpello 2—3-ovulato. Capsula ellipsoidea (3-5.5 mm) rite calyce minor, glabra (inma- tura); carpella 1-2(?)-seminata. Semina inmatura. 832 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 the leaflets discolorous with loose appressed hairs ca. 4% mm long above, short pilose and glabrescent beneath, the apical mucro ca. % mm long; petioles 1-3 cm long, fleshy; rachis hirsute; petiolule % mm long, fleshy. Inflorescences 2-5, globose, ca. 5 mm diam., 1-7 flowers seen opened to different degrees, peduncles erect, 7-10 cm long, hirsute; bracts subulate, 3-4 mm long, appressed ciliate, subtending the pedicels; bracteoles spirally overlapping, linear triangular, 2.5—4 mm long, 1-nerved, keeled, fine and long ciliate. Flowers with sepals linear, 5 mm long, ca. | mm wide, acute, (7-)9-11-nerved, glabrous; petals pink oblong spatulate, basally clawed; stamens with filaments yellow, the longer ones ca. 2 mm long, sparingly glandular pubescent, the anthers ovoid, the shorter filaments with reduced (?) sterile anthers, enlarged in the lower half, filiform above: pistil about equalling the stamens, the ovary ellipsoidal, 5-lobed, the carpels 2-3- ovuled. Capsule (immature) ellipsoidal, 3-3.5 mm long, shorter than the calyx, glabrous; seeds 1—2(-?) in each carpel. The size of the petals is unknown to me because the flowers on the collection are not open. The epithet used for this species refers to the conspicuous mucro of the leaf- lets. PANAMA: Cerro Trinidad, 800-1000 m, Allen 3775 (F, G, MO, US). = Biophytum panamense Lourt.? ryPE: Panama, Hammel 2528 (MO, holotype; MO, P, isotypes).—Fic. 3B Erect or decumbent herbs to 20 cm long, the stem branched at the base or not, pubescent with curved, short hairs, glabrescent; roots fibrous, branched. ° Biophytum panamense Lourt., spec. nov. Radix fibroso ramosa. Caulis unus, vel plures, erectus vel decumbens (usque 35 cm altus), pubescentia brevi retrorsa. Apex caulis claviformis, cum annulo dense pilifero, pilis rigidis retrorsis, infra foliorum insertionem posito. Caulis supra annulum pilorum foliorum delapsorum cicatricibus instructus. Folia 9-20, pseudoverticillata, in apice caulis disposita, ug Petiolus (ca. 2 mm longus) carnosus. Rachis canaliculatus, angustissime alatus, hirsuto-pilosus. Pe- tiolulus latior quam longus (1⁄4 mm longus) carnosus. Foliola atroviridia, opposita, aliquam superpos- ita, inaequilatera, mucronata (mucro ca. 4 mm), antice crescendo majora, pubescentia tenuissima et brevissima, glabrescentia, margine c ciliata; nerviatione secundaria et tertiaria ansa infima mini- ma, ovato-triangularia vel suborbiculata (3-4.5 x 3-4 mm); media oblongo-asymmetrica, paulo cur- 8 anteriore obliquo-truncata. Cymae 1-3, abbr eviatae, globoso- vel cylindrico-congestae (ca. diam.), 1-2(-5) floribus adspectabilibus, vicissim apertis. Pedunculi erecti, brevi (ca. 5 mm) EM pubescentia brevi. Bracteae violaceae, subulatae (2-2.5 mm) ad ege culorum insertionibus. Pedicelli longi (10-13 mm) violacei, brevissime pubescentes et glandulosi, ca. basim articulati. Bracteolae ata TI sque 7 5—7-nervata, pubescentia et glandulosa, basim plus minusve violacea, margine subhyalina Petala alba, fauce flava (fide coll.), oblongo-linearia, obtusa, calycis duplo longiora. Stamina longiora 4.5 mm), filamentis pilis glandulosis conspersis, antheris ovoideis basim cordatis; breviora (ca. 1 aes q staminibus: ovarium 5-lobulatum, apice vix pubescens, carpello 2-3-ovulato; stylis pubescentibus et glandulosis; stigmata conspicue bifida, papillosa. Capsula sphaeroidea (ca. 3.5 mm diam.), : vix iloso, rite calyce minor, stylis longis, carpellis l-seminatis. Semina (ca. 1.5 mm, mu. semi- ellipsoidea, fulva, obtusa, verosimiliter longitudinaliter striata vel tuberculata. 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 833 Leaves 9—20, pseudoverticillate at the end of the stem, linear subcuneate, 18 cm long, 2.2 cm wide, 20-32 jugate; lower leaflets ovate triangular, 3—4.5 mm long, 3-4 mm wide, middle leaflets oblong, asymmetric, slightly curved, 5-12 mm long, 6.5—7 mm wide, basally the posterior side curved, the anterior side truncate, the midrib emerging from the center or near the posterior angle, the terminal leaflets narrowly elliptic to semiobovate, 8—11.5 mm long, 4-5.5 mm wide, oblique basally the posterior side lobulate, the anterior side truncate, the leaflets somewhat over- lapping, minutely fine pubescent, glabrescent, ciliate, the mucro ca. 2 mm long, minor venation reticulate. Inflorescences 1-3, contracted, globose to cylindric, ca. 5 cm diam., 1-2(—5) flowers seen at a time, opening successively; peduncle erect, ca. 5 mm long, violaceous, short pubescent; bracts subulate, 2-2.5 mm long, violaceous, situated at the insertion of the peduncles; bracteoles ovate to linear, to 4 mm long, 1 mm wide, green, violaceous at the base, strongly keeled, glabrate; pedicels 10-13 mm long, violaceous, minutely pubescent and glandular. Flowers with the sepals linear, to 7 mm long 1 mm wide, sharp, 5-7-nerved, pubescent and glandular, the base violaceous, the margin subhyaline; petals white, linear oblong, obtuse, twice as long as the calyx; longer stamens ca. 4.5 mm long, sparsely glandular, and the anthers ovoid, basally cordate, the shorter stamens ca. 1 mm long, larger basally, the anthers sterile or obsolete; pistil about equalling the longer stamens, the ovary 5-lobed, apically slightly pubescent, the carpels 2—3-ovuled, the styles pubescent and glandular, the stigma bifid, papillose. Capsule globose, ca. 3.5 mm diam., slightly pilose apically, shorter than the calyx, the style persistent, elongate, the carpels 1 seeded; seeds (immature) tawny, semiellipsoidal, ca. 1.5 mm long, obtuse, perhaps longitudinally striate or tuberculate. The epithet refers to the country where the species was found. COCLE: Coclesito road, continental divide ridge, Hammel 2528 (MO, P). CA Biophytum soukupii Lourt.6 TYPE: Peru, Prov. Mariscal Cáceres, Tocache Nuevo, Quebrada de Cachiyacu de Lopuna, carretera a Progreso, 500-850 m, Schunke 7640 (P, holotype; MO, isotype).—Fic. 3C Herb or subshrub apparently stemless or the stem to 50 cm tall, 4 mm in diam., erect or decumbent, tomentose; root branched, fibrous, elongate. Leaves Biophytum soukupii Lourt., spec. nov. Radix copiose ramosa, longa. Subacaulis vel n uy as erecto vei decumbens, basi radicante, tomentoso-pubescente (usque 50 cm lat diam.). Apex caulis claviformis cum annulo dense pilifero, pilis rigidis, retrorsis, infra fo vile Pi i [o ici ia 13 x 2.5 cm), 10-25-juga oles (c m Rachis canaliculatus, angustissime alato (alis inconspicuis), pubescentia brevi, S pidum inaequali. = mac] & "a £S: 3 Ë O c B = _@ c Ic] 3 B. = Ë as a) Lnd -— D [2l — > s. = i= a Petiolulus (ca. 1⁄4 mm) carnosus, latior quam longus. Foliola disana viridia et purpurea, opposita, oblique, inaequilatera, nervo mediano in tercio basis viciniore an ng posteriore oriundo; — nervis in 20-35 jugis dispositis, tenuibus et crassis alternatis, in nervo n pagina in- feriore prominentibus; pubescentia minutissima recta a, subappressa, sparsa in utraque pagina vel intel. dum glabra in una vel duabus paginis; oe minima, subtriangularia asymmetrica (ca. 3 x 2.5 mm); ud oblongo- rectangularia (6—14.5 x 2.5-4.5 mm) obtusa, basi truncata, angulo posteriore recto parte anteriore oblique truncata angulo anteriore curvato vel raro elongato; summa obovato- elliptica. leviter asymmetrica in medio vix latiora (5.5-6.5 mm), basi dimidio posteriore in angulo 834 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VOL. 67 7-30 pseudoverticillate at the end of the stem, linear subcuneate, to 13 cm long, 2.5 cm wide, 10—25-jugate; lower leaflets subtriangular, 3 mm long, 2.5 mm wide, the middle leaflets larger, 6-14 mm long, 2.5-4.5 mm wide, oblong rectangular, obtuse, basally truncate or oblique on the anterior side, the anterior angle round- ish, the posterior angle square, the terminal pair obovate elliptic, 5.5-6.5 mm, basally truncate, on the posterior side, the angle square, the anterior side oblique, the leaflets green above, violaceous beneath, oblique, the midrib emerging at the base % way to the posterior angle, secondary nerves 20-35 pairs, alternating thick and fine, terminating in a marginal nerve, short appressed pubescent on both sides; petiole ca. 0.25 mm long, broader than long. Inflorescences 1—4(—7), glomeruliform or fusiform, 5 mm long, 2-2.5 mm wide, one flower opening at a time; peduncle 1-2 cm long, shorter than the leaves, pubescent; bracts situated at the insertion of the peduncles, subulate, 2.5-5 mm long, short appressed pu- bescent, purplish; outer bracteoles triangular, 1.0 mm long, 0.5 mm wide, inner bracteoles elongate, subulate, keeled, to 3.5 mm long appressed ciliate, sparingly short pilose; pedicels about equalling the bracteoles, somewhat longer, to 3.5 mm, in fruit, articulate near the base. Flowers with the sepals sublinear, 5.5— 7 mm long, 1-1.5 mm wide, acute, 5-7-nerved, hirsute with long and glandular short hairs; petals white (?pink), oblong spatulate, 1.5 times the calyx length, basally clawed, minutely glandular outside; stamens with filaments enlarged to- ward the base, the thecas subsagittate, the longer stamens ca. 6.5 mm long, pubescent and glandular, the shorter stamens ca. 3 mm long, glabrous; pistils macro-, meso-, and microstylous and as long as the shorter stamens, the style pubescent and glandular, the stigma enlarged, 2-lobed, subcapitate, papillose, the ovary apically pubescent and glandular, the carpels 4—6-ovuled. Capsule oblong, 4.5-5 mm long, about equalling the calyx, apically pubescent and glandular or glabrate, the carpels 4—6-seeded; seeds tawny, semi-ovoid, ca. 1.5 mm long, apiculate, margined, longitudinally tuberculate. This novelty is named after Father Jaroslav Soukup of the Colegio Salesiano of Lima, Peru, editor of the review Biota, enthusiastic botanist who has collected for many years all over Peru, and who has collected this species twice. DARIEN: Cabecera del Rio Pirre, Bristan 1322 (MO). Cana, 200-6500 ft, Gentry et al. 28545 (MO, P), Williams 765 (NY). ecto truncato, dimidio anteriore oblique truncato. Cymae 1-4(-7), congestae (5 x 2-2.5 mm), lome ruliformes vel fusifor rmes plerumque flore uno simul. Pedunculi rite foliis minores (1-2 cm); a siblinearia: (5.5-7 x 1-1.5 mm), 5- 7 nervata, acutiss ssima, uber sa es hirsuta longa m . at: i apice pubescens et glandulosum, carpello 4—6-ovulato. “aaa globosa (4.5-5 mm), calyce aequalis, apice pubescens et glandulosa, carpello 4—6-seminato. Semina fulva, ied cd, -convexa (ca. 1.5 mm) apiculata, marginata, tuberculis longitudinaliter dispositis ornata. 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 835 3. OXALIS Oxalis L., Sp. Pl. 433. 1753. TYPE: O. acetosella L. Acetosella Moehr, Hort. Priv. 4. 1736. Oxys [Tourn.] Adanson, Fam. Pl. 2: 388. 1763. Sassia Molina, Saggio Chile 145. 1782. Oxallis Noronha, Verh. Batav. Gen. 5. Art. IV, 21. 1790. Pseudoxalis Rose, Contr. U.S. Natl. Herb. 10: 116. 1906. Ionoxalis Small, Fl. S.E. U.S. 665. 1903. N. Amer. FI. 25(1): 26. 1907. Monoxalis Small, Fl. S.E. U.S. 665. 1903. N. Amer. Fl. 25(1): 45. 1907. Lotoxalis, Sma ll, Fl. S. E. U.S. 666. 1903. N. Amer. FI. 25(1): 46. 1907. FA nas aml, Fl. S.E. U.S. 665. 1903. N. Amer. Fl. 25(1): 27. 1907. Herbs or subshrubs, rarely lianas; roots branched fibrous or subligneous thickened; stems underground and reduced to an invisible disc covered with bracts and the base of petioles connate to the stipules, or modified in a more or less ligneous, thickened rhizome covered with the base of the decayed leaves, or aerial and climbing, decumbent, trailing or erect, subligneous at the base, or all herbaceous. Leaves alternate, apparently basal or cauline, subopposite or pseu- doverticillate, 3(—4)-foliolate, digitate or pinnate, with or without stipules connate to the petioles. Inflorescences axillary, of 2-branched or umbelliform cymes or solitary flowers; bracteoles 2; bracts small. Flowers varying in size, rarely cleis- togamous; calyx 5-partite; petals 5, connate near the middle; stamens 10, connate at the base, the series of 5 external alternisepalous stamens opposite the petals, shorter, the internal series alternipetalous, longer, usually all fertile; pistils het- erostylous, bi-, trimorphic, the carpels 1-15-ovuled, the styles 5, free, the stigmas capitate. Capsule globose, oblong to cylindric, glabrous or pubescent, the carpels (1—)2-15-seeded, loculicidally dehiscent, the carpels connate to the central axis, persistent; seeds generally ovoid, apiculate or oblong, more or less flattened, the testa crustaceous, longitudinally zig-zag ribbed, transversally striate or sculptured densely verrucate, 2-tegmic, the external integument fleshy, arilliform, breaking elastically and projecting the ripe seeds; albumen fleshy; embryo straight. Oxalis includes more than 800 species distributed around the world in tropical and cold regions, from sea level to the snow-line in the highest mountains. Twelve species have been found in Panama, none endemic. a. Leaves 3-foliolate, pinnat Petals white or E DM at the base, the limb pinkish 2. O. barrelieri bb. = yellow. ymes with foliaceous bracts; leaflets glaucous with white cystoliths . O. dombeii [e] O Cymes with linear, small bracts; leaves green or more or less p Cymes long branched, 10—40-flowered; capsule globose .. 9. ` microcarpa dd. une umbelliform, 3-7(-11-15)- "ewe b ies ovoid or oblong oe O. frutescens en angustifolia £e e ais mta foliolate, digitate Roots fibrous or thickened; stems aerial or rhizomatous; sepals without calli at the n à f. Stem aerial, erect or rooting; flowers yellow, sometimes violaceous veined. iana; leaves exstipulate, the leaflets ovate ---------------- 10. O. rhombifolia 836 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 gg. Herbs; leaves with stipules connate to the petiole, the leaflets commonly obovate with the apex incise h. Stipules truncate or narrowed towards the apex; capsules oblong or n covered with gai (rarely glabrate). Stems decumbent or e j. Petals light lew. translucid; leaflets green, m lobules ascendent; calyx to '/s—'/s of the capsule length __________ 3c. O. co e Rm corniculata jj. Petals bright yellow, orange or pinkish outside; le E aucous, the lobules more or less divergent: calyx 14 ^ of the capsule length |. 3a. O. O aa xai albic ans Simi pied creeping. greenish to violaceous, dark green when dry; cap- sule cylindric, calyx to !/s-!/a its length, slightly ae pubescent to glabrous . O. corniculata var. atropurpurea kk. Stem reddish, decorticating, leaves green: flowers yellow aya oblong, calyx to !2—24 its length, shot pubes- -. = cT SU U E Etc EE O. filiformis hh. ti ih fen acute apex; capsules conic, Ea or with e pluricellular hairs --------------- . spiralis d vulcanicola ff. Plants ks era stemless, rhizomatous to 2 cm diam., ligneous, bearing the bases of decayed leaves; flower . O. articulata ee. Bulbs formed by the reduced stem covered with bracts on the stipules connate to the petioles; sepals usually with - at the apex; flowers p l. Leaflets 3, entire to bilobat I Leaflets subtria irm . O. “a aaa mm. Leaflets obovate, incised, the lobes rounded ____ 4. O. "e var. corymbos Leaflets 4, entire or rarely slightly retuse. aa. . O. fe — . Oxalis articulata Savigny in Lamarck, Encycl. 4. 686-687. 1797. TYPE: Uru- guay, Montevideo, in pascuis, Commerson May 1767 (P-Juss., holotype; P, isotype). Apparently stemless herb to 40 cm; stem ligneous, rhizomatous, brown, dark, more than 15 cm long, to 2 cm diam., tortuose, bearing the scars of the bases of stipules adnate to petioles of decayed leaves; roots tuberous, subligneous, cylin- dric, knobby or articulate, dark, thick, more than 10 cm long, 25 mm diam., branched, fibrous, to 15 cm long, sometimes with a long branch bearing a tuber at its apex. Leaves 3—30-fasciculate; leaflets widely obovate subcuneate or cu- neate, 5-30 mm long, 7-45 mm wide, slightly retuse or acute, incised to !/s of its length, a purplish basal callus frequently present, the pubescence straight, ap- pressed, uniform, dense on both faces or only below, or shorter and loose; orange calli abundant near the margins, unequal, rarely scattered on the rest of the surface; stipules connate to the petiole, ovate or triangular, 4-16 mm long, 2-6.5 mm wide, narrowed towards the apex, the hairs appressed ascendent, ciliate; petioles ascendent, 13-30 cm long, densely appressed pubescent to glabrate with short appressed hairs; petiolule ca. 1 mm long, thick, densely pilose. /Inflores- cences of umbelliform simple cymes, 2—5-flowered, bifid, asymmetric, multiflow- ered with up to 20 flowers or of bifid cymes composed of umbelliform cymes; peduncles taller than the foliage (2.5—40 cm) the pubescence similar to that of the petioles; bracts 2-3 mm long; bracteole 0.5—1(—2) mm long, situated at the base of the inflorescence, oblong or linear, appressed-pilose with 1—2 orange calli at the apex; pedicels long filiform, 1—5.5 cm long, articulate near the base. Flowers with the sepals linear-elliptic to linear 2.5-6.5 mm long, 1-2 mm wide, acute, the internal sepal obtuse, often with 3-4 unequal calli, densely appressed-pubescent, 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 837 the hairs straight; petals violaceous, obovate-oblong, clawed, connate near the middle, 2—3 times the length of the sepals, the areas exposed in bud covered with white sericeous appressed hairs; stamens with linear filaments, basally broad and abruptly narrowed about the middle, the longer stamens 3.5 mm long, pubescent, the shorter stamens 2 mm long, glabrous, connate to about 25, the anthers ovoid; pistils macro-, meso- and microstylous, ca. 5.5 mm long, the ovary pilose at least at the apex and back of the carpels, carpels 4—8-ovuled, the styles densely pilose, the stigma small, capitate. Capsule cylindric or oblong, acute, 8—11 mm long, the calyx to % of its length, densely ascending appressed pubescent overall or only at the apex of the carpels, the carpels 4—8-seeded; seeds light brownish, ovoid or ellipsoid, acute at both ends, 1—1.25 mm long, 8—9-ribbed in zig-zag, transversally striate with 4-8 deep pits. Originally from Uruguay, extreme southern Brazil, and some localities of Buenos Aires (Argentina), this species is introduced and cultivated in other American countries and in the Old World. It may escape and become naturalized. CHIRIQUÍ: Bambito, escaped from cultivation, roadside, Croat 15882 (MO). 2. Oxalis barrelieri L., Sp. Pl. ed. 2. 1763. TYPE: Barrelier, Plant. Rar. Tab. //39. Acetosella barrelieri (L.) Kuntze, Rev. Gen. Pl. 1: 90, 92. 1891. Lotoxalis barrelieri (L.) Small, N. Amer. Fl. 25(1): 49. 1907. Erect herb or shrub to 1 m tall, pubescent; stems green or violaceous, pu- bescence white or yellowish hirsute, in part retrorse, the apex finely pubescent; internodes to 8 cm long, sometimes almost obsolete resulting in subopposite leaves, hairs at the insertion of the petioles straight; roots fibrous, branched; petioles ascendent, to 4 cm long, canaliculate, enlarged to the base, the margin of the groove ciliate; rachis to 15 cm long pubescent; petiolule fleshy, ca. 1 mm long, pilose. Leaves with leaflets varying in shape on the same plant, suborbic- ular, oblong, ovate to elliptic, 10-55 mm long, 8-25 mm wide, obtuse, seldom acute, the base acute, roundish or subpeltate, asymmetric, discolorous, glaucous above, hairs irregularly scattered, below caducous, the margin or only the base ciliate. Inflorescences of cymes taller than the foliage loosely 1-11(—30)-flowered; peduncles to 6.5 cm long, pubescent, bifid with branches to 3 cm long; bracts reddish, lanceolate or triangular, acuminate, ca. 2 mm long, pilose; pedicels to 2.5 mm long, articulate near the base; bracteoles smaller, appressed to the ped- icel. Flowers with the sepals light green, sometimes red-veined, 3—3.5 mm long, 1-1.5 mm wide, linear to ovate or oblong, acute, mucronate, pubescent, the margin hyaline; petals white or yellowish at the throat and base, the limb pink, obovate-subspatulate, clawed, 7-9 mm long, 3.5 mm wide, slightly notched; long- er stamens ca. 3 mm long, pubescent or glabrous, ligulate about the middle, the shorter stamens 1.5-2 mm long, glabrous; pistil 3.524 mm long, the ovary ovoid, glabrous, the carpels 3—4-ovuled, the styles pubescent, the stigma widened, bifid, subcapitate, papillose. Capsule ovoid 7-9 mm long; acute, rarely oblong, gla- brous, membranous, calyx to 1⁄2 its length, the carpels (2—)3(—4)-seeded, inside setose pubescent; seeds brownish, ovoid-compressed, apiculate, 1.5-2 mm long, 8-ribbed in zig-zag, deeply transversally striate, finely tuberculate at the inter- sections. 838 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 This species occurs in tropical and subtropical America. It is introduced into some tropical places of the Old World. CAS DEL TORO: Road to *‘Bomba’’ Almirante, n 1316 (MO). Ojo de Agua, ca. 7 km W o Almirante, 190-220 m, Croat 38149 (MO). Near Milla 7.5, Croat & Porter 16284; Milla 10, 16344; Milla 7.5 16455; Milla 5, 16498 (all MO). Cerro Jefe, in cafetal, D'Arcy 3968 (MO, P). Changuinola, 0-100 m, D'Arcy 11168 (MO). Santa Rita Ridge, Lewis et al. 5245 (MO, P). Runway at Bocas, Lazor 2439 (MO). Zigla Road, junction of Changuinola and Tuibe Rivers, Lazor et al. 2512 (MO). CHIRIQUI: Between Cerro Punta and Bajo Grande, Croat & Porter 16095 (MO). Volcán Chiriquí, Aguacatal 2000 m, Killip 3578 (F). COLON: Santa Rita Ridge, 7 mi from Transisthmian highway, 650 ft, Wilbur 15034 (MO). Without locality: Seemann (BM, K). 3. Oxalis corniculata L., Sp. Pl. 435. 1753. NEOTYPE: Cult. hort. Uppsal. herb. Thunb. 11084 (UPS) Ja. Oxalis corniculata L. ssp. albicans (H.B.K.) Lourt., Phytologia 42: 137. 1979. O. albicans H.B.K., Nov. Gen. Sp. Pl. 5: 244. 1821. TYPE: Mexico, Bonpland & Humboldt 4071 (P, otype; B, isotypes). Herb to 40 cm; stems branched at the base, prostrate or decumbent, rarely erect, pubescence whitish or yellowish to rusty, ascending or appressed to sub- hirsute, curved, sometimes dense; root often ligneous, turnip-shaped, thick, more than 30 cm long, branched. Leaves pseudoverticillate, subopposite or alternate; leaflets greenish, glaucous, largely obovate-cuneate, generally wider than long, incised up !/s-!/3(—!/2) of the length, the lobes divergent, the lateral leaflets asym- metric, the margin often violaceous (conspicuous in dried specimens), the pu- bescence varied, appressed to hirsute-ciliate, commonly glabrous above; petioles ascendent, pilose; petiolules fleshy, violaceous. Inflorescences ascendent, 1-2(— 3)-flowered cymes; generally shorter than the foliage, appressed-pubescent; pe- duncle pilose, to 5 cm long; pedicels appressed-pubescent, longer in fruit, gen- erally the same length or shorter than the fruit. Flowers with sepals elliptic, oblong, commonly greenish-violaceous, uneven, the outer sepals pubescent, the others glabrate; petals yellow striate with red outside, pinkish or salmon inside (fide coll.) the border villous (observed in buds); pistils mostly mesostylous, some pistils about as long as the longer stamens, a few microstylous. Capsules cylin- dric, acute, the calyx to !4—!5 its length, the styles short. The remaining characters are the same as the ssp. corniculata. This subspecies occurs in Western United States, Mexico, Guatemala, Hon- duras and Panama. CHIRIQUí: Río Chiriquí Viejo across from town of Cerro Punta, 6000 ft, D'Arcy 6575 (MO, P). a Popa, above Boquete, 1500-2500 m, D'Arcy 10929 (MO). Paseo Respingo, 8400 ft, D'Arcy & Hammel 12888 (MO). Cerro Punta, 6000 ft, Tyson 5792 (MO). 3b. Oxalis corniculata var. atropurpurea Planchon in van Houtte, Fl. Serres 12: 7—48, t. 205. 1857. TYPE: Planchon's plate. O. corniculata L. var. viscidula Wiegand, Rhodora 27: 117, 121. 1925. rype: Massachusetts, Terry (GH). Small herbs; roots deep, fibrous, profusely branched; stems, stipules and leaves violaceous. Flowers with the sepals green or somewhat violaceous; pistil 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 839 micro- or macrostylous, mostly near the same length as the longer stamens. Capsule slightly pubescent to glabrous. This variety, of horticultural origin from Europe, is easily recognized in the living state. It is found in gardens, greenhouses and often escapes. It is not weedy. CANAL ZONE: Salamanca Hydrological Station, Río Pequeni, 80 m, Woodson et al. 1622 (GH). 3c. Oxalis corniculata L. ssp. corniculata. Prostrate herb, many branched, hirsute; stems to 50 cm long, branched, the tips ascending; internodes variable in length, roots fibrous. Leaves sometimes in long petiolate fascicles; leaflets obovate, cuneate, 4-20 mm long, 7.5-20 mm wide, widely incised up to !/e x !/ of the length, the lobes roundish, obtuse, above glabrous, pubescent below (often only one half), appressed-ciliate; petioles to 7 times the length of the blade; stipules 0.8-3 mm long, 1-3 mm wide, truncate, hirsute-ciliate. Inflorescences of umbelliform 2—7-flowered cymes; peduncles to 21 cm long, ascending, loosely pilose; the bracts linear-triangular 0.5-3 mm long, longer in fruit; bracteoles smaller, linear, ciliate; pedicels 4-15 mm long in fruit and deflexed; sepals linear or narrowly ovate, acute 2.5-5.5 mm long, 4-15 mm wide, greenish or violaceous, sparingly pubescent; petals yellow, oblong-subspat- ulate, about twice as long as the sepals, 5-11 mm long, 1-2.5 mm wide; stamens glabrous, the longer stamens ca. 6 mm long, the shorter stamens ca. 4 mm long, connate up to about !; pistils micro-, meso-, and macrostylous to 8 mm long; the ovary oblong, acute, retrorse-pubescent, the styles appressed retrorse pu- bescent, the stigma 2-lobed, papillose, the carpels 4—15-ovuled. Capsule cylin- dric, acute, 5-20 mm long, 1-3 mm diam., retrorse pubescent with pluricellular glandular hairs, exceptionally glabrous, the carpels commonly 8-10(—14)-seeded, pubescent, pubescent inside, the calyx to !/4-!/s the capsule length, the styles long connate, straight; seed reddish-brown, ca. 1 mm long, ovoid, apiculate, flattened, 6-ribbed, transversally striate and finely tuberculate at the intersections. This species is distributed worldwide, originating in the European Mediter- ranean region. It is well adapted to all climates, requiring only some humidity, and reaching its largest forms in cultivated lands. It can become weedy and hybridize with autochthonous species of the same section. OCAS DEL TORO: Changuinola Valley, Dunlap 393 (F, US). CANAL ZONE: Chagres, between Saas and Alahuela, 30-60 m, Allen 975 (MO). Balboa, Standley 28561 (US). Río Pequeni, Sala- manca cca ad Station, 80 m, Woodson et al. 1622 (MO, NA, NY). curRiQUÍ: Río Chiriquí Viejo w Sw land," 1800-2000 m, Allen 1403 (GH, MO). Boquete, Finca Collins, Blum Dwyer zu (M or oein, Escuela San Benito, Croat 10416 (MO). N of Cerro Punta, Croat 10484 (MO, P). Cerro Punta, IDAAN water tank, 700 m, Croat 26551 (MO, P). 4 mi above Boquete on road to Volcán Barú, 1475 m, Croat 34855 (MO). Between Boquete and Monte Rey, Croat & Porter 15627, 15732 (both MO). Between Cerro Punta and Bajo Grande, Croat & Porter 15994 (MO, P, US). Bambito, 5500 ft, D'Arcy 5341 (MO). Río Chiquero, D'Arcy 11006 (MO). La Popa, Boquete, D'Arcy & D'Arcy 6370 (MO, P). Boquete, 4200 ft, D'Arcy & D'Arcy 6458 d P). Cerro Respingo, Cerro Punta, 8400 ft, D'Arcy & D'Arcy 6551 (MO, P). Cerro Horqueta, 1500 m, Duke et al. 13657 (MO). Boquete, Finca Collins, Ebinger 715 (F, MO, NY). Las Cumbres near ca. Punta, 7200 ft, Hammel 6952 (MO). Boquete, 3300-4200 ft, Lewis et al. p (GH, MO, UC, US); Maurice 685 (US). Finca Collins, “El Velo," 6150 ft, Stern et al. 1967 (MO). Boquete, 4200 ft, Sucre 194 (RB). Nueva California, 4000 ft, Tyson 5709 (MO). Cerro a 6000 ft, Tyson 5793 (MO). Volcán de Chiriquí, Casita Alta, 1500-2000 m, Woodson et al. 990 (MO, NA, NY). Finca Lérida, Woodson & Schery 310 (MO). cocLÉ: Club Campestre, El Valle de Antón, 700 m, Duke 13237 (MO). 840 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 4. Oxalis debilis H.B.K., Nov. Gen. Sp. Pl. 5: 236. r. 466. 1821. 4a. Oxalis debilis H.B.K. var. corymbosa (DC.) Lourt., comb. nov. O. corymbosa DC., Prodr. 1. 696. 1824. LECTOTYPE: Insula Borbona (G-DC). O. martiana Zuccarini: Denksch. Akad. Wiss. Muench. 9: 144. 1825 O. urbica St. Hil., Fl. Bras. Mer. ys 100. 1825. O. bipunctata Graham ex Hook., CMS 54: t. 2781. 1827. TvPE: cult. Roy. Bot. Gard. Edinburgh, seed 2m Brazil, Rio de Dus sent by Harris to Graham April 1827 (E). O. violacea auct. non L. Acet eroe martiana (Zucc.) Kuntze, Rev. Gen. Pl. 1: 90. 1891. Ionoxalis martiana (Zucc.) Small, Fl. S. E. U.S. 665. 1903. Apparently stemless herb; root vertical, thick with fibrous branches; bulb globose, ovoid or oblong, 1-2 cm long, 3 cm diam., formed of nutritious scales and numerous bulbils covered by protective scales and the stipules connate to the petioles; scales brown, ovate, scarious 7-11 mm long, 3-5 mm wide often covered with rigid, acuminate, orange calli, the margin hyaline, ciliate with long smooth hairs, to 4 mm long, rusty, the 3 longitudinal nerves connate below the apex. Leaves with leaflets suborbicular or broadly obovate, 1.25—5.5 cm long, 1.5-4.7 cm wide, incised up to !/9—1/s of the length, cuneate, the hairs short, sparse on both faces or only beneath, with punctiform minute, translucid orange or violaceous oxalate "crystals" scattered on the blade, more visible below, frequently a larger and more prominent row along the border; petioles to 30 cm long with sparse hairs, rarely densely pubescent; petiolules 0.5—1 mm long, fleshy, pilose; stipules completely connate, 13 mm long, 2 mm wide, scarious, brown, linear, 3-nerved, the nerves salient, convergent just below the apex, sometimes with 2 additional thinner lateral nerves running with the central nerve into the petiole, the borders ciliate as the scales; bulbils covered by similar smaller scales. Inflorescences bifid cymes, the branches unequal, sometimes twice bifid, erally asymmetric or umbelliform, loosely 3—15-flowered; peduncles little shorter to longer than the foliage, to 45 cm long, similar to the petioles; bracts opposite, hyaline, largely ovate or rhombic, 2-5 mm long, 2-2.5 mm wide; bracteoles in- serted near the base of the pedicel on the pilose articulation, 1-3 mm wide, 0.25— | mm long, bracts and bracteoles with calli; pedicels 4-20 mm long, similar to the peduncle, articulate 1-3 mm from the insertion. Flowers with the sepals greenish, the margin hyaline, linear to elliptic, 4-7 mm long, 0.75-1.5 mm wide, subequal, obtuse or acute, the apex ciliate, the calli orange, linear or enlarged towards the apex, parallel or oblique, rarely convergent or divided giving 3 or 4 calli; petals pinkish, salmon or red-violet, sometimes with inconspicuous short appressed hairs outside, obovate, clawed, 2-3 times the length of the sepals; stamen filaments enlarged towards the base, the longer stamens 4.5 mm long, pilose towards the apex and with some short glandular hairs mixed; the shorter stamens ca. 3 mm long, glabrous, connate to !^; pistil macro- and mesostylous, rarely microstylous, 7.5 mm long, with scattered ascendent hairs and short glan- dular hairs, the stigma bifid, subcapitate, the ovary glabrous, the carpels 2-12- ovuled. Capsules (immature) cylindric, thin, glabrous to 17 mm long, the calyx to !4—V5 as long as the capsule, styles ca. 2 mm long, the carpels short pilose inside, 3-10-seeded; seeds (immature) brownish, ovoid, flattened, 12-ribbed in zig-zag and 12 transversal striae. 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 841 This is a South American species widespread in the whole world, occasionally fruiting but reproducing through abundant bulbils developing simultaneously. OCAS DEL TORO: Bocas, border mangrove swamp, Lazor et al. 2428 (MO). CHIRIQUI: Boquete 4500 n D'Arcy 5446 (MO); D'Aarcy & D'Arcy 6460 (MO). 3 mi N of Boquete, 3300—4200 ft, Lends et al. 340 (MO). Alto Lino, Maurice 873 (US). 5. Oxalis dombeii St. Hil., Fl. Bras. Mer. 1: 111. 1825. TYPE: Perú, Dombey (P). O. darienensis Woodson, Ann. Missouri Bot. Gard. 27: 312-313. 1940. TYPE: Panama, Allen 881 MO, holotype; GH, NY, P, US, isotypes). O. barrelieri sensu auct. non L. ~ Erect herb to 60 cm, glabrous or puberulent on younger parts; white cystoliths often present on the leaves, bracts, bracteoles, occasionally on the sepals; pu- bescence long transparent, sometimes viscous; stem sometimes subligneous at the base, to 6 mm diam. often with glandular hairs, these inconspicuous when lacking secretion; internodes 1.5-4 mm; roots fibrous, branched; rhizome cylin- dric, slender. Leaves alternate, subopposite or pseudoverticillate; leaflets oblong obovate to suborbicular, the lateral leaflets asymmetric, 9-25 mm long, 5-22 mm wide, glabrous or with hairs on the nerves below and fine ciliate, obtuse or retuse, basally subpeltate or barely auriculate; petiole to 4 cm long, canaliculate, often narrowly winged; rachis to 6 mm long. Inflorescences of cymes taller than the foliage, loosely 10—25-flowered: peduncle to 6 cm long, bifid with long branches, one branch aborted or shortened; bract and bracteoles overlapping, bracts ca. 3 mm long, bracteoles ca. 1.5-2.5 mm, foliaceous, lanceolate, acuminate, greenish or red-striate, appressed to the rachis, subamplexicaul, pedicels ca. 3 mm long, glabrous, rarely pubescent, articulate near the base. Flowers with the sepals greenish, the margins hyaline, ovate and acuminate or oblong 4.5-7 mm long, 1- mm wide, glabrous, acute; petals yellow-orange, obovate-subspatulate, 7-13 mm long; stamens longer, ca. 2.5 mm long, pubescent with a thin ligule, acute in the lower half, the shorter stamens ca. 1.25 mm long, glabrous; pistils ca. 3 mm wide, the ovary oblong, the carpels to 10-ovuled, the styles thin, pubescent, the stigma capitate. Capsule oblong to ellipsoid, ca. 10 cm long, the calyx to % of its length, glabrous, the carpels 7-10-seeded, inside densely and finely pubescent; seeds rusty, ca. 1 mm across, subpyriform, 5—6-ribbed in zig-zag, transversally striate and tuberculate at the intersections. This species occurs on dunes and arid regions, rarely in cultivated fields, of the Pacific coast and Andean slopes of Perú, Ecuador, and Panamá. It also occurs in the Galápagos Archipelago. : Boca de Cupe, Allen 881 (MO, NY, P, US). Río Chico, near Yaviza junction with Río of Psp y e Larsen 215 (G). Camino de El Real a Río Pino, Lazor 1550 (MO). “eet e and a de Cupé, near El Real along Río Pirré, Stern et al. 609 (GH, MO, US). PANAMA: Between Río ines Dam ye Canitas, D'Arcy 9395 (MO, P). Near Bayano bridge, Folsom 3556 (MO). 6. Oxalis filiformis H.B.K., Nov. Gen. Sp. Pl. 5: 190, r. 460. 1821. TYPE: Colom- bia, Guaduas, Santa Ana, Bonpland (P, holotype; B, P, isotypes). 842 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Xanthoxalis filiformis (H.B.K.) Holub., Folia Geobot. Phytol. Praha 8: 175. 1973. Creeping herb to 10 cm tall, pubescent with fine and whitish or reddish simple and pluricellular hairs, 1.5-2 mm long, mixed; roots fibrous, branched; rhizomes and stems filiform, to more than 50 cm long, reddish, decorticating, hirsute, rooting; internodes 1—5 cm long, the nodes 1-leaved or with fascicles of leaves. Leaves with leaflets cordiform or transversely oblong, cuneate, 2-15 mm long, 3-20 mm wide, incised to 4 of its length, both faces pubescent or above glabrous, glabrescent, ciliate; stipules connate, reddish, enlarged at the apex, 2-3 mm long, 0.75-1.75 mm wide, setose-ciliate, concave. Inflorescences of the cymes longer than the foliage, 1-flowered; peduncles 1-7 cm, filiform; bracteoles linear, pilose, 1.5-3 mm long; pedicel 0.5-2.5 cm long. Flowers with the sepals greenish, the margins often violaceous, oblong, 3-5 mm long, 1-2 mm wide, obtuse, the apex densely ciliate, pubescent; petals 2-3.5 times the length of the sepals, rarely with minute glandular hairs scattered on the border and outside; stamens minutely pubescent, the longer stamens to 5.5 mm long, the shorter stamens to 4 mm long, connate to % of its length; pistils all longistylous, 6.5-7.5 mm long, the ovary reddish, carpels with a few short hairs inside, 3—6-ovuled, styles densely pubes- cent, ca. 5 mm long, slightly connate at the base, the stigma bifid, laciniate, papillose. Capsule oblong, 5-12 mm, thick, the calyx to !2—24 the capsule length, acute, minutely pubescent, the styles long, free, the carpels 3-6-seeded; seeds reddish-brown ca. 1 mm long, ellipsoid, flattened, acute, 6-ribbed, transversally striate. This species ranges from Costa Rica along the cordillera to Ecuador, occurring among herbs in prairies and savannas above 1,000 m elevation. CHIRIQUI: 5.4 km del Hato del Volcán en el camino a Las Lagunas, Correa & Lazor 1479 (MO). 7. Oxalis frutescens L., Sp. Pl. 435. 1753. 7a. Oxalis frutescens ssp. angustifolia (H.B.K.) Lourt., Phytologia 29: 463—471, f. 3. 1975 O. angustifolia H.B.K., Nov. Gen. Sp. Pl. 5: 193. ses TYPE: Mexico prope La Venta del Peregrino, Humboldt & Bonpland a (P, holotype; B, isotype). O. coccinea Woodson & Schery, Ann. Missouri Bot. jn 28: 431. 1941. TYPE: Panamá, Woodson & Schery 1029 (MO, boe. NY, US, isotypes). Hirsute subshrub to 40 cm tall or small shrubs branched from a ligneous base, the branches ascending, twisted or decumbent; stems with hirsute, fine long, uneven pubescence; roots twisted, often swollen, to 7 mm diam. Leaves pseu- doverticillate, crowded or the internodes to 5.5 cm long, leaflets green, glaucous or violaceus, the middle one at least twice as long as the laterals, elliptic, subacute or obovate, generally notched, 10-45 mm long, 4-20 mm wide, the lateral leaflets oblong, rarely elliptic, asymmetric, notched, 4-20 mm long, 1.5-10 mm wide; pubescence dense on both faces or above glabrous or glabrescent, some speci- mens only pubescent on the midrib, ciliate; rachis 0.2-10 mm; petiole 10-40 mm, ascendent, pubescence as on the stems. Inflorescences of cymes umbelliform, 2.5 cm, somewhat taller than the foliage, 3-7(—11—15)-flowered; branches short- ened ca. 1 cm, 1 often aborted; bracts 1.5 mm and bracteoles ca. 1 mm, linear, 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 843 acute; peduncles to 3.5 cm; pedicels 5 mm, pilose, articulate 1 mm from the base. Flowers with the sepals elliptic or linear, 3—5.5 mm long, 0.25-1.5 mm wide, acute, mucronate, pilose to glabrous, the interior sepal glabrate, the apex ciliate; petals yellow, 244-3 times the length of the sepals, obovate, clawed, slightly notched; longer stamens ca. 3 mm long, pilose, the ligule acute near the middle, the shorter stamens ca. 1.5 mm long, connate up to 4; pistil 4.5 mm long, pilose, the ovary ovoid, the style pilose, the stigma bifid, capitate. Capsule ovoid or oblong, 5-6 mm long, except those with 1-seeded carpels, rarely to 8 mm when carpels 8-seeded, the calyx usually 1⁄2 (—4/s) its length; the carpels ciliate on the back, inside pubescent, rarely glabrous, 1—5-seeded; seeds brownish 1.5-2 mm long, ovoid, apiculate, 8-ribbed in zig-zag with 10—12 transversal deep striae and fine tubercules on the intersections. This species is frequent in poor sandy, stony soils and savannas and campos from the Texas-Mexican frontier to the northwest of Argentina. OCAS DEL TORO: Nievecita, Woodson & Schery 1029 (MO, NY, US). CocLE: Santa Clara beach, Croat 9587 (MO). 4-6 mi E of Nata, 10-25 m, Duke 12402 (MO). Penonomé, Dwyer 2040 (MO); Ebinger 996 (F, MO). Natá, Seemann 1227 (K). Río Grande 10 mi E of Natá, Tyson 5277 e ERRERA: 10 mi S of Ocú, Tyson et al. 2874 (MO). PANAMÁ: Cerro Jefe, 700 m, Davidse & D'A 10115 (MO, P). Near Playa Río Mar, 10-100 ft, Duke 11783 (MO, US), /2408 (MO, NY); Ebinger 497 (F). 8. Oxalis latifolia H.B.K., Nov. Gen. Sp. Pl. 237. t. 467. 1821. TYPE: Mexico, Campeche, Humboldt et Bonpland (P-HB, holotype; B, isotype). O. intermedia Richard, Essai Fl. Cuba 315. 1841. TYPE: Cuba, de la Sagra (P, holotype; P, isotypes). O. mauritiana Loddiges, Bot. Cab. 178. tab. 1780. 1832. TYPE: Loddiges' plate. O. lilacina Klotzsch in Otto & Dietrich, Allgem. Gartenzeit. 8: 285. 1840. TYPE: Not found. O. chiriquensis Woodson, Ann. Missouri Bot. Gard. 24: 190. 1937. TYPE: Panama, ie 222 (MO, holotype; NY, isotype). O. stylosa Klotzsch ex Knuth in Engler, Pflanzenr. 95(130): 273. 1930. nomen O. prin H.B.K. var. karwinski Progel ex Knuth in Engler, Pflanzenr. 95(130): >. 1930. nomen duse wislae ea (L.) Kuntze ssp. latifolia (H.B.K.) Kuntze, Rev. Gen. Pl. 1: 90. 1891. Ionoxalis latifolia (H.B.K.) Rose, Contr. U.S. Natl. Herb. 10: 113. 1906. I. intermedia (Richard) Small, N. Amer. x 25(1): 43. 1907 Apparently stemless herb; roots deep, vertical, the branches fibrous; bulbs ovoid or globose to 25 mm long, 20 mm diam., outer scales papyraceous, large, sometimes absent; scales (nutritious) brown, elliptic, acute, narrow, 7-15 mm long, 4-6 mm wide, strongly 3-nerved, 1-2 lateral pairs of nerves thinner, excep- tionally yet another shorter lateral pair present; bulbils ovoid, acute, ca. 1 cm, young, protected by several hyaline bracts similar to those described above, smaller. Leaves with the leaflets green, violaceous below, triangular, cuneate 10— 45 mm long, 20-75 mm wide, the distal margin straight, rarely incised to !/s—!/4 the length, short mucronate or broken in a broad, obtuse angle and the lobes divergent, small calli of various shapes often nearby, the distal angles obtuse, glabrous or sparse hairy below on the nerves or rarely overall; the margin with some cilia; petioles ascendent 5-15 cm long, glabrous or somewhat pilose; petio- lules fleshy, ca. 1 mm long, pilose; stipules connate to the petioles, hyaline- papyraceous, rusty-brown, ovate or rectangular, the apex auriculate, ciliate with long, viscid rusty hairs, the margins glandular, 3 prominent nerves arising from 844 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 the base and with orange calli converging just below the apex, lateral pairs gen- erally 2, thinner and shorter, not reaching the apex: the inner pair sometimes bifurcate. Inflorescences umbelliform cymes commonly twice as long as the fo- liage, to 30 cm long, loosely 5-20-flowered; peduncles 10-25 cm long, similar to the petioles; bracts ca. 2 mm long, 2 mm wide, and bracteoles ca. 1 mm, all at the base of the inflorescence, hyaline, broadly ovate to rhombic or elliptic, acute, clasping the peduncle or the pedicel, ciliate, 1-nerved, softly crenate; pedicels thin, 14-20 mm long. Flowers with the sepals green, the margin hyaline, glabrous, uneven, linear-elliptic to oblong, narrowed to the apex, obtuse or subacute, 3-5 mm long, 0.5-1.5 mm wide, with orange or violaceous calli at the apex, 0.3—0.8 mm long, 2-lobate, convergent at the top end, sometimes entire, ovate, the base irregular; petals white at the base, pink to purplish at the limb, 2.5-3.5 times the length of the calyx, obovate, spathulate, clawed; stamens with filaments enlarged towards the base, pilose on the upper half, the longer stamens ca. 4 mm long with an obtuse sometimes inconspicuous ligule near the middle; the shorter stamens ca. 3 mm long, connate !⁄4 way up; pistils micro- or rarely macrostylous, 4.5-5.5 mm long, the ovary ovoid, glabrous, the carpels 4-8-ovuled, the styles fine, gla- brous, the stigma bifid, papillose. Capsule oblong, acute, glabrous 4-8.5 mm long, the calyx to !á—'^ its length, the styles 1.5-2 mm long, the carpels 4—8- seeded, the hairs inside sometimes inconspicuous; seeds brownish, ellipsoid, slightly flattened, ca. 1-1.1 mm long, 8—9-ribbed in zig-zag with 12-13 transversal striae. This species occurs in Mexico, Costa Rica, the Antilles, and the Andes of South America up to 1,800 m elevation and south to Bolivia. It is introduced to the southeast of Brazil, Europe, Africa, Australia, and New Zealand. CHIRIQUI: Volcán, xe San Benito, Croat 10415 (MO). Boquete, 3800 ft, Davidson 703 (F, GH, MO). Volcán Barü, 8000 ft, Hammel 2918 (MO). Valley of upper Río Chiriquí Viejo, Monte Lirio, 1300-1900 m, ie rt 222 (MO, NY). Chiriquí Viejo valley, White 102 (MO). Boquete, 1200— 1500 m, Woodson & Schery 777 (MO). 9. Oxalis microcarpa Benth., Pl. Hartweg. 115. 1839. rype: Ecuador, prope Zam- boronton, Hartweg III . . . (K) O. oe Turcz., Bull. Soc. Imp. Moscou 31: 427. 1858. Type: Ecuador, banks of Guayaquil ver, Jameson 557 (K, Holotype; BM, G, W, isotypes). Herb or subshrub to 1.20 m; stem erect, green or brown, the pubescence fine, curved, generally retrorse, long, hirsute, whitish or yellowish; internodes 2.5-5.5 mm, roots fibrous, branched; rhizome thin, to 4 mm diam. Leaves alternate to subopposite; leaflets ovate, elliptic or subrhombic unequal, 10-38 mm long, 16- mm wide, obtuse or subacute basally cuneate or slightly cordate, the pubes- cence subappressed, even, scattered, above denser, glabrescent beneath, un- evenly ciliate; petiole filiform, ascendent to 4 cm long, pubescent, canaliculate, enlarged to the base, with hairs at its insertion. The inflorescences of bifid cymes, 10-12(—40)-flowered, peduncles filiform, pubescent as on the petioles up to 6 cm long, branches longer in the fruit, to 6 cm; bracts linear, acuminate, hirsute, 1— 1.5 mm; pedicels articulate near the base 2-3 mm long. Flowers with the sepals green, ovate or elliptic, acuminate, 3 mm long, 0.75-1 mm wide, acute or rarely 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 845 subacute, the margin hyaline, often apically ciliate, glabrous; petals yellow, ob- ovate-subspathulate 6-7 mm long; longer stamens ca. 2.5 cm long, ligulate under the middle, glabrous or pubescent; the shorter stamens 1.5—-2 mm long, glabrous; pistil about equalling the longer stamens; styles pilose, the stigma enlarged, 2-lobulate, capitate, the carpels 1—2-ovuled, glabrous. Capsule subglobose ca. 3 mm long, glabrous, about equalling calyx; the carpels membranous, thin, gla- brous inside, 1-seeded; seeds brownish ca. 1.5 mm long, 8-ribbed in zigzag, deeply transversally striate with small tubercules on the intersections. This species occurs along the Pacific coast and western slopes of the cordillera from Ecuador to Mexico. CANAL ZONE: Quebrada La Palma and Canon of Rio Chagres, 70-80 m, Dodge & Allen 17337 (G). Rio Indio, 70-100 m, Dodge & — 17381 (BR, G, K, MO, P). Chagres, 1 mi above Madden Lake, eg 4475 (MO, P). Madden Dam, Ebinger 876 (US). Chagres, Fendler 25 (K, US), 26 (GH, MO). Fort Sherman between dum Batteries and Fort San Lorenzo, Maxon & Valentine 6991 (GH, ex US). Punta de Garachiné, Seemann 1063 (K). Las Cascadas p near Summit, Standley 29618 (US). Río Boquerón, Steyermark & Allen 17230 (BR, G, MO, P, S, US). Quebrada L and Canon Hill, Woodson et al. 720 (F, GH, MO). DARIEN: Río Chico, near Yaviza at junction with Río Chucunaque, Burch et al. 1192 (MO). Mannené, Kirkbride & Bristan -— 1612 (both MO). Río Tuquesa, Camp Charco Peje of Tuquesa Mining Co., 250 m, Mori 7042 (MO). PANAMA: Río Canita, near Jenine, Duke 3838 (MO). petere Río Pacora and Río Corso, 450 m, Duke 11955 (P). Tortí Arriba, Folsom 5163 (MO). La Chorrera, Lewis et al. 5198 (MO). Río Tapia, Standley 28075 (US). VERAGUAS: Piria, 120 m, Duke 14359 (MO). 10. Oxalis rhombifolia Jacq., Oxalis Monogr. 22. tab. 2. 1794. TYPE: Venezuela, Caracas, Bredemeyer ex herb. Jacquin (W). O. maxonii Standley, J. Wash. Acad. 17: 311-2. 1927. Type: Panamá, Maxon 4996 (US, holotype; ,G S, isotypes). O. rhombifolia Jacq. var. pubescens Pittier, Trab. Mus. Com. Venezuela 7: 310. 1930. TYPE: not hte rhombifolia (Jacq.) Kuntze, Rev. Gen. Pl. 1: 93. 1891. Climbing subligneous shrub to 6 m; pubescence yellow to rusty, long hirsute; stems profusely branched, brownish, new branches hirsute, glabrescent; inter- nodes 5-40 cm long, shortened at the apex and on the small lateral aborted branches which bear a fascicle of leaves. Leaflets elliptic to rhombiform or rhom- bic ovate, 14-45 mm long, 10-26 mm wide, obtuse or subacute, the middle leaflet longer, the lateral leaflets somewhat asymmetric, both faces pubescent, glabres- cent above, ciliate; petiolules ca. 1 mm thick, pubescent; petioles filiform, 11⁄— 2 times the length of the blades, pubescent, swollen at the base. Inflorescences with cymes bifid, loosely 3—5-flowered, pubescent; peduncles to 5 cm long; bracts 3-7 mm long; bracteoles 1.5-2 mm long, linear, narrow, pubescent; pedicels 3— 5 mm long, articulate near the base. Flowers with the sepals oblong or linear oblong, 4-5.5 mm long, 1.5-2 mm wide, obtuse, densely ciliate at the tip; petals yellow, obovate, subspathulate, 10-17 mm long, the longer stamens 5-8 mm long, hirsute, the shorter stamens ca. 3 mm long, enlarged towards the base, connate to 4; pistils 7-10 mm long, macro-, meso-, and microstylous, the styles hirsute, the stigma bifid, enlarged, papillose, the ovary densely pubescent, the carpels 3-ovuled. Capsule oblong or ovoid 7-9 mm long, densely tomentose, the calyx 15 as long as the capsule, the carpels setose inside, 2-3-seeded; seed dark colored ca. 2.25 mm long, semiovoid, flattened, 13—14-ribbed, the ribs tuberculate. 846 ANNALS OF THE MISSOURI BOTANICAL GARDEN í M d \ NI N N Y x y ) SN S N \\ ‘A YN M een » "em ) c FIGURE 4. Oxalis spiralis ssp. vulvanicola (D. Sm.) Lourt. Habit (x!4). [After Nee 10001.] This species occurs from Mexico to Venezuela and Colombia in humid and shadowed situations of the forest HiRIQUÍ: Bajo Mono, 4500 ft, D' Arcy 11062 (MO, P); Davidson 495 (F, MO). Without locality, š x: : “rice 888 OL, US). , GH, US); Pittier 2970 (F, US). Quiel Road 12.2 km above Boquete, M tor pee Aa ud Horqueta, 1600 m, Webster in (F). Between Bajo Quiel and Bajo Mono, ca. 6 m of Boquete, 1450 m, Wilbur et al. 12014 (MO). Bajo Mono, mouth of Quebrada Comune. dons ES Caldera, 1500-2000 m, Woodson et al. 1011 (GH, NA, MO); Woodson & Schery 526 (MO). ll. Oxalis spiralis R. & P. ex Don, Gen. Syst. 1: 755. 1831 lla. Oxalis spiralis ssp. vulcanicola (D. Sm.) Lourt O jpa pu D. Sm., Bot. Gaz. 23: 241 , comb. nov. Pittier 869 (US). 1897. TYPE: Costa Rica, Montée au flanc SE du Poás, 1900 Xanthoxalis vulcanicola (D. Sm.) Small, N. Amer. Fl. 25(1): 1907. . "siliquosa" J. Roy. Hortic. Soc. 85: 371. 1960, nom. MAS sine diag. lat. Decumbent or trailing herb (small young plants erect), finely pubescent with yellow or reddish hairs, sometimes glabrate, succulent, to 1.5 m, branched; in- ernodes 2-10 cm long. Leaves with leaflets oblong to suborbicular, cuneate, widely notched, mucronate, 10-35 mm long, 8-23 mm wide, dark green above, violaceous below, the midrib pubescent below, lateral leaflets asymmetric; peti- oles violaceous 2-4 cm long; stipules reddish, hyaline, connate except at the [VoL. 67 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 847 narrow and acute apex, 2.5-5.5 mm long, slightly pubescent. Inflorescences of lateral, axillary and terminal cymes taller than the foliage, (1—)2-17-flowered; peduncles to 12 cm; bracts 1.5—4 mm long and bracteoles ca. 1 mm long, linear, pubescent; pedicels to 16 mm long, articulate near the base. Flowers with the sepals linear, 5-7 mm long, 0.75—1.5 mm wide, narrowed towards the apex, acute, glabrous or somewhat pubescent; petals yellow, obovate, cuneate 12-18 mm long, violaceous-veined; stamens with filaments short, glabrous, the longer stamens ca. 6 mm long with an obtuse ligule near the middle; the shorter stamens ca. 2 mm long, connate halfway up; pistils micro-, meso-, and macrostylous, 6 mm long; the ovary glabrous, the styles very thin, the stigma bifid, capitate papillose. Cap- sule ellipsoid, acute ca. 6 mm long, glabrous, the calyx the same length, the carpels 3-4-seeded; seeds oblong ca. 1 mm long, dark violaceous. Oxalis spiralis ssp. vulcanicola is a common plant of the forest in the sub- paramo and cloudy forest of Central America south to Panama, occurring in volcanic regions up to 3,500 m elevation. It is cultivated as an ornamental in greenhouses, usually in hanging pots, for the beauty of its profusely branched red stems and its abundant bright yellow flowers. Grown in this way the branches are pendulous. CHIRIQUI: Cerro Punta to ago of Rio Caldera, 2250-2500 m, Allen 1425 (F, GH, MO, US). NW of Boquete, Rio Caldera, 6400 ft, Averett 1105 (MO, P). Cerro Punta a Las Nubes, 6000 ft, Correa 1341 (MO). Cerro Punta, pos 10451 (MO, P, US). 2.7 mi NW of Río Chiriquí Viejo, 2200 m, Croat 22362 (MO, P). W of Cerro Punta, 2200 m, Croat 22363 (MO). Road to Volcan Baru, 12 mi above Boquete, Croat 34867 (MO). Alto Respingo, 2800 m, D'Arcy 9959 (MO, P). Potrero Muleto to summit of Volcán Barú, D'Arcy 10155 (MO, um deni Respingo, above Cerro Punta, 3000 m, D'Arcy 10731 (MO). La Popa above Boquete, 1500-2 , D'Arcy 10854 (MO, P). La Nevera, below summit of El um 10,400—10,500 ft, D'Arcy & condi 12447 (MO, P). El Volcan SE of La Nevera, 10,000 ft, D'Arcy & Hammel 12512 (M O). Between Guadalupe and Paso Respingo, 8100 ft, D'Arcy & Hammel 12836 (MO, P). Ridge above Alta Boquete, 7400-8000 ft, D'Arcy et al. 13147 (MO). Volcán Barü, above Boquete, 3100 m, pepe & D'Arcy 10270 (MO, P). Boquete, Bajo Choro, ft, Davidson 418 (F, GH, MO, US). Nubes region 7 km NW of Cerro Punta, 7200 = Hammel 1383 (MO). Volcán Barú, E slope, oer 000 ft, Hammel 2812 (MO). Volcán Barü, slope W of Finca Yen, 8000 ft, Hammel et al. 6470 (MO). Cerro Punta, roadside w of Paso ae 8 : 00 m, Nee 10001 (MO, P, US). Boquete, 10,000 ft, Terry 1342 (F, GH, MO, US). Quebrada Grande from Bajo Grande to Paso de Respingo, 2100-2400 m, Wilbur et al. 15118 (MO). Potrero Muleto to summit, Volcán de Chiriquí, 3500-4000 m, Woodson & Schery 467a (MO). 12. Oxalis tetraphylla Cav., Icones 3: 19-20. t. 237. 1794. TYPE: Cavanilles’ plate. deppei Loddiges, Bot. Cab. 15. t. 1500. 1828. Type: Loddiges' plate. SM wasqan Rd dr pes Allgem. Gartenz. 3: 388. 1835. TYPE: not found. j , Bull. Acad. Sci. Bruxelles 2: 334. 1836. TYPE: Lejeune's plate mid it Knuth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 316. 1919. mucronata (Rose) Knuth, Notizbl. Bot. Gart. Berlin- ers 315. 1919. hayi Knuth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 316 M (Rose) Knuth, Notizbl. Bot. Gart. Berlin- ae 391. 1919. cuernavacana (Rose) Knuth, Notizbl. Bot. Gart. Berlin-Dahlem 7: 291. 1919. . quadrifolia Knuth in Engler, Pflanzenr. 95(4). 130: 288. Lr nom. inval. in synon. Acetosella tetraphylla (Cav.) Kuntze, gs Gen. has n 92. A. deppei (Lodd.) Kuntze, Rev. Gen. Pl. 1: 92. Cond tetraphylla (Cav.) Rose, Contr. E S. AA Herb. 10: 115. 1906. I. deppei (Lodd.) Small, N. Amer. Fl. 25(1): 32. 1907. S 848 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 I. scopulorum Rose in Small, FI. x A d S. 666. 1903. I. divaricata Small, Fl. S.E. U.S. I. mucronata Rose in Small, N. is e 25(1): 40. 1907. Apparently stemless herb; roots vertical, fasciculate, turnip-like or cylindric, sometimes with profuse and very long fibrous branches; bulb globose or ovoid to 4 cm long, 3 cm diam., formed by the reduced conic or discoid stem, the scales and the stipules connate to the petioles; bulbils distant on thin stolons, rarely observed in present material; outer scales scarious, brownish or with hyaline margin, the outermost row linear, 3-5-nerved, 4-18 mm long, 1-12 mm wide, sharp, sometimes absent or lost in collecting, the other ovate or largely ovate- triangular, (S—)7—14-nerved, 15-17 mm long, 5-8 mm wide, abruptly acute; inner nutritious scales yellow, thick rigid, flat-convex or subconcave, 3-5(-7)-nerved, the nerves broad, prominent, 3-7 mm long, 2-7 mm wide; stipules hyaline or scarious, whitish or reddish-brown, connate, broad at the bse, 12-30 mm long, 4—13 mm wide, auriculate at the apex, 1-2 mm long, rarely acute, (5—)10—12- nerved, 5—7 nerves more salient, alternate with the other 5-7 and 2 lateral thinner nerves that do not reach the apex and are sometimes absent; in both bracts and stipules the nerves joining at different levels, only a few reaching the apex or running into the petiole, all having rigid rusty hairs 5-6 mm long inside and long cilia at the margins. Leaves with 4 leaflets, exceptionally in young plants 3, subequal, obovate or subtriangular, rarely suborbicular or with divergent lobules, 20-70 mm long, 25-75 mm wide, truncate, roundish, incised '/10 of its length, the midrib sometimes lengthened with a short mucro, discolorous, with a transverse fringe, purplish to blackish in the middle, the hairs simple or rarely short pluri- cellular, reddish on the nerves below, rarely above and on the margin, generally with small unequal translucid or dark oxalate spots located near the incision on the margin or scattered on the upper part of the blade; petioles 5-30 cm, glabrous or sparsely hairy. Inflorescences of bifid cymes, umbelliform 3—2-flowered, pe- duncles as tall as the foliage or much taller, to 50 cm, similar to the petioles; bracts 2-4 mm long, 2-2.5 mm wide, and bracteoles smaller, hyaline, ovate to rhombic, generally keeled, the midrib lengthened in a mucro, sometimes covered with oxalate calli, irregularly long ciliate; pedicels to 17 mm articulate at the base. Flowers with the sepals green, oblong to elliptic, 4-7.5 mm long, 1-2 mm wide, the margin hyaline, obtuse, truncate to subacute, glabrous or with a few hairs and glandular hairs, the apex ciliate with 2-4 unequal, linear, apically parallel calli, obtuse, truncate or subacute; petals pink to violet, the lower part greenish- white, 2—5 times the length of the calyx, seldom with glandular and short hairs outside; stamens longer, 2.5-4 mm long, with short hirsute, uneven hairs and glandular hairs above, the ligule large, linear, obtuse, near the middle the shorter stamens 1.5-2.5 mm long, enlarged to the base, glabrous, connate to 4—; pistils mostly macrostylous, only 1 mesostylous and 1 microstylous, 5-7 mm long, the ovary glabrous or with hairs at the top of the carpel backs, the styles pilose with curved hairs and a few glandular hairs, the stigma bifid, papillose, the carpels 3— 6-ovuled. Capsule cylindric, acute, 13 mm long, glabrous, the calyx up to '4 of its length, the styles ca. 1.5 mm long, the carpels 4—5-seeded, the interior pilose; seeds tawny, ovoid, flattened, apiculate, 1.25 mm long, 9—14-ribbed in zig-zag, the 12-16 transversal striae prominent. 1980] LOURTEIG—FLORA OF PANAMA (Family 84. Oxalidaceae) 849 This species seems to originate from Mexico. It has been introduced to all the continents, often cultivated as a good luck token. Difficult to eradicate, it does not seem, however, to be weedy. RIQUI: Finca Lérida, S slope of the Quebrada Velo, 5000 ft, Allen 4733 (G, MO, NY). Boquete . 3 mi N of Boquete, 3200-4200 ft, Lewis et al. 428 (GH, MO). Bajo Mona and Quebrada Chiquero, 1500 m, Woodson & Ero 568 (MO). INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (t) refer to names incidentally mentione Acetosella 835 articulata 836 barrelieri 837 barrelieri 837, 841 deppei 847 bipunctata 840 martiana 840 hiriquensis 843 rhombifolia 845 coccine — 847 corniculata 838 violac —ssp. albicans T ios latifolia 843 —ssp. atropurpur Averrhoa 823+, 8 —ssp. conma E 839 bilimbi 824+, 824 —var. viscidu cara 5 corymbosa Biophytum 8231, 826 cuernavacana 847 nd s darienensis 841 dormiens debilis 840 falcifolium 829 —var. corymbosa 840 ongiscapum 827 dendroides 82 mexicanum 827 —fo. longiscapa 827 mimosoides 827 deppei 847 mucronatum 831 dombeii 841 panamense 831+, 832 elegans sensiti 26+ —var. karwinski 843 ouku esculenta 847 Bolboxalis 835 filiformis 841 frutescens Oxalidaceae 823 Oxalis 823+, 8 —sect. Biophytum 826 + acetosella 835 albicans 838 angustifolia 842 842 —ssp. angustifolia 842 Guayaquil 84 intermedia 843 latifolia 843 lilacina 843 violacea 840 850 viva 827 vulcanicola 846 zonata Oxallis 835 Oxynis 8 xys 835 Pseudoxalis 835 Sarcotheca 823+ ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Sassia 835 vulcanicola 846 FLORA OF PANAMA! BY RoBERT E. WoopsoN, JR., AND ROBERT W. SCHERY AND COLLABORATORS Part VI FAMILY 93. MALPIGHIACEAE JosÉ CUATRECASAS? AND THOMAS B. CROAT? Trees, shrubs, or scandent lianas, seldom subshrubs. Leaves simple, opposite or subopposite; petiole often glandular; blades entire, rarely lobate or irregularly toothed when juvenile, often with abaxial or marginal glands; venation pinnate; stipules present; T-shaped trichomes frequently present. Flowers bisexual, acti- nomorphic or usually obliquously zygomorphic principally by modification of 1 petal, and diversification of stamens and styles; in many-flowered, axillary pan- icles, pseudo-racemes, or umbels; pedicels sessile or articulate to bibracteolate peduncles whcch are borne in the axils of bracts; bracts and bracteoles conspic- uous or small, sometimes glandular; calyx of 5 sepals, more or less united at the base, the lobes quincuncial, rarely valvate in bud; often with a pair of conspicuous glands; petals 5, showy, free, often yellow, but also white, pink, red and violet, clawed, fringed or toothed, alternate to the sepals; the interior petal often reduced and of another color, sometimes all equal, the claw articulated with the limb; stamens (8—)10, equal or unequal, sometimes with the 5 epipetalous stamens reduced, sometimes with 4 stamens opposite the 4 glandular sepals reduced to staminodes, sometimes with the 3 stamens opposite the 3 carpels much enlarged (Stigmaphyllon, Banisteriopsis spp.), the filaments more or less connate at the base, rarely free, usually glabrous, the anthers 2-celled, basifixed or dorsifixed, introrse, dehiscing longitudinally, often with a thickened connective, sometimes glandular, sometimes appendaged; ovary superior, mostly 3-locular, (2-)3(—4)- carpellate; placentation axile, the ovules solitary in each locule, pendulous, semi- anatropous; style (2—)3 free, rarely connate, distally terete, triangularly dilated, spatulate or sometimes appendages forming a hood over the stamens, rarely with only 1 style developed, the stigmatic tissue uniform at the tip of the styles or on their adaxial angle. Fruit usually a schizocarp of 3 samaras, sometimes a drupe with 1-3 pyrenes (Bunchosia, Byrsonima, Malpighia), a 2- or 3-locular dehiscent capsule (Galphimia) or separable indehiscent cocci (Pterandra); seeds ovoid, endosperm lacking, cotyledons flat, slightly curved, folded, uncinate or spirally rolled (Byrsonima, Pterandra). The pubescence of Malpighiaceae, though rarely stellate, generally consists ! Assisted by National Science Foundation Grant DEB 77-04300 (W. G. D'Arcy, principal in- vestigator). 2 United States National Herbarium, Department of Botany, Smithsonian Institution, Washing- ton, D.C. 20560. 3 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouri Bor. GARD. 67: 851—945. 1980. 852 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 = c ee ç À g 240 T VAM UNDER — — FIGURE 1. Malpighiaceous trichomes.—A.—K. Trichome types found generally in the family.— L. Unusual warty type found in Baniste riopsis ae Í (Spruce) Morton. See Text for explanation. of unicellular bifurcate, generally T-shaped trichomes. Types A, B, C, G, J and K (Fig. 1) have arms of equal length. Types A and K are sessile, whereas B, C, G and J are variously stipitated. Types D, E, H, and I have unequal arms; in E and J the longest arm is erect, looking superficially like a simple unbranched hair, whereas in type D the arms are less unequal and divergent laterally. Types A, B, C, D, E, have thicker cell membranes and are firm and straight, whereas types G, H, I, J and K, have thinner cell membranes and are flexuous; these types, especially G, I and J, make a lanate indument. Type L is a sessile medifixed trichome with minute warts as in Banisteriopsis caapi (Spruce) Morton. Niedenzu (1928) in the last major revision of the family divided the Malpigh- iaceae in two subfamilies and seven tribes. The subfamily Hiraeoideae (Pyr- amidotorae) includes the tribes: 1. Hiraeeae, 2. Banisterieae and 3. Triccomar- ieae the many genera of which consist mostly of woody lianas, many with winged fruits; of these, 14 genera are exclusive of the Old World, all other genera are restricted to the New World, with the exception of a single African species of the large neotropical genus Heteropteris. The second Niedenzu subfamily Mal- pighioideae (Planitoreae), includes the Niedenzu tribes 4. Galphimieae and 5. 1980] CUATRECASAS & CROAT—FLORA OF PANAMA (Family 93. Malpighiaceae) 853 Malpighieae, and consists mostly of trees, erect shrubs and subshrubs, exclu- sively American. The family consists of about 60 genera with 850 species, mostly in the Amer- ican tropics. Members range from Texas and the Bahamas to northern Argentina and occur in habitats ranging from wet rain forests to arid areas. Some malpighiaceous trees produce fine-grain wood which is used locally for making furniture and in construction. The bark and wood of some species are rich in tannins and are used in small factories as dyeing or tanning material. Many species produce drupaceous fleshy fruits or acidulous juicy berries which are directly eaten or prepared into jellies or preserves. The juice of Malpighia emar- ginata Sesse & Moc. is rich is ascorbic acid (vitamin C). Some species of Ban- isteriopsis contain alkaloids, like harmaline (=yageine, banisterine, telepathine) with dramatic hallucinogenic effects. ‘‘Ayahuasca,”’ “caapi,” ‘‘yagé,” and "'na- tem’’ are the most general common names for B. caapi (Spruce) Morton in Brazil, Colombia, Ecuador, Peru, etc. where the natives have been using it for centuries in ceremonial celebration. Other species of Banisteriopsis and other genera (e.g. Tetrapteris) also have harmaline and several other chemical derivatives with a diversified spectrum of narcotic effects, according to the recent progress of re- search in this field (e.g. Schultes & Hofmann, 1973). The generally zygomorphic flowers of the Malpighiaceae are probably bee pollinated (Croat, 1978) and the fruits are mostly wind-dispersed. A few species, such as species of Spachea, Bunchosia and Malpighia are endozoochorous per- haps at least in part by bats (Croat, 1978). Literature: Adams, C. D. 1972. Flowering Plants of Jamaica. University of the West Indies. Mona, Jamaica. 848 pp. Anderson, W. R. 1976. A new species of Jubelina (Malpighiaceae) from Central America. Brittonia 28: 410—412. De Candolle, A. P. 1824. Prodr. Syst. Nat. 1: 577-592. Croat, T. B. 1978. Flora of — Colorado Island. Stanford University Press. Stanford, California. 943 p Cuatrecasas, J. 1958. Prima [m Colombiana. Webbia 13: 565—588. Hutchinson, J. 1967. The Genera of Flowering Plants (Angiospermae). 2. Di- cotyledons. Oxford University Press. London. 659 pp. Johnston, I. M. 1949. The botany of San José Island. Sargentia 8: 1-306. Jussieu, A. H. L. de. 1840. Malpighiacearum synopsis, monographiae mox ed- enae prodromus. Ann. Sci. Nat. Bot. Sér. 2. 13: 247-291; 321—338. 1843. Monographie de la famille des Malpighiacées. Arch. Mus. Hist. Nat. 3: 5-151; 255-616. MacBryde, B. 1970. A revision of the Galphimiinae Niedenzu (Malpighiaceae). Ph.D. Dissertation. Washington Univ., St. Louis. 248 pp. Niedenzu, F. 1928. Malpighiaceae in A. Engler, Pflanzenreich IV. 141 (91, 93, 94): 1-870. Schultes, R. E. & A. Hofmann. 1973. The Botany and Chemistry of Halluci- nogens. Charles C. Thomas, Publishers. Springfield, Illinois. 267 pp. 854 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Small, J. K. 1910. Malpighiaceae. N. Amer. Fl. 25: 117-171. Standley, P. C. 1928. Flora of the Panama Canal Zone. Contr. U.S. Natl. Herb. 27: 1-416 Triana, J. & J.-E. Planchon. 1862. Malpighiaceae. Jn Prodromus florae Novo- Granatensis. Ann. Sci. Nat. Bot. Sér. 4. 18: 307-338 KEY TO THE GENERA a. Fruit a schizocarp splitting into 3 samaras; receptacle pyramidal; usually lianas or climbing shrubs. Dorsal wing of the samaras well developed, large, conspicuous; lateral wings short or absent. €. Stamens I0, subequal, fertile; style tips blunt, stigma apical, subapical or at the adaxial angle of the styles, or hooked; samara wing thickened on upper or lower argin d. Samara wing thick on the upper margin, thin along the lower margin: styles subulate, the stigma capitulate, apical or subapical |. , uidi dd. Samara wing thin at the upper margin, thickened at the lower mar ill usually thicker and unequal, dilated, spathulate-truncate or hoo ked at apex, the stigma at its adaxial angle — 11 5. Het ee cc. Stamens ig uneg ual, only 4-6 fertile with thicker anthers; styles unequal, t apex enlarged, triangular-truncate, hooked or with foliaceous appendices, the penra at the inner angle; samara wing thickened on upper margin, tuno n low dur mc THREE C 13. Sdpmaphyllon nor dns of the samara obsolete or much reduced; lateral wing or wings well dig en and larger than the dorsal win Flower bud covered until anthesis by 2 large, cochleariform, opposite, embrac- ing bracteoles. Sepals oblong or spathulate, densely pubescent abaxially, soon reflexed RT ERR a tc tence, 10. Mezia ee. Flower bud not covered by 2 large bracteoles before anthesis: sepals linear, oblong, ovate or oblong-ovate. f. Sepals linear or oblong with subvalvate estivation, pubescent on both sides, soon reflexed; 4 sepals with one large subbasal gland; bracteoles linear or chicas ; Samara body with 3 cavities, the median monospe Dus: T c 2 lateral s the dorsal wing developed 1. 7, Juheliha ff. Sepals ovate or oblong-ovate with quincuncial estivation, usually glabrous within, usually erect, each with 2 dorsal glands or eglandular; samara bod unilocular with no lateral empty cavities, the dorsal wing shortly carinate, crested or not developed. ateral samara wing parted into 4 narrower wings disposed ig an X, al equal or with the abaxial pair shorter than the upper pai obsolete aeu cat aM opu ETERNA E EE 14. D gg. Lateral samara wing continued into a disc n the body, part- ed (open) at the adaxial line, or part ted both at the adaxial and abaxial points, thus separating a wing to e Pedicels pedunculate (articulate), the pair of bracteoles at the apex of the floral peduncle. Stipules inserted on the pu close to the base of the petiole — 1. . Mascagnia hh. phas sessile, the bracteoles at the base. old inserted cholo s QR RERO DIR E IEEE 6. Hiraea aa. Fruit drupaceous, a loculicidal . or a smooth 3-coccoid schizocarp; receptacle flat: usually trees or erect shrubs. i. ruit schizocarp splitting into 3-2 indehiscent, smooth, glabrous or pilose cocci. j. Styles short, obtuse at apex. Inflorescences long, pseudo-racemiform, terminal: flowers usually 1-2 per branch; ovary “suwaqa lacking a gynopodium; pedicels pedunculate; anthers not appendiculate — 12. Spachea J. Styles long, subulate. Inflorescences ey subumbellate or fasciculate, axillary or extraxillary; flowers 3 or more per branch; ovary pubescent, sitting on mnt broad gynopodium; pedicels sessile; ie. with dorsal narrow bela? appen- dix _ s mc TERREA = Lll . Pterandra ii. Fruit drupaceous or capsular. 1980] CUATRECASAS & CROAT—FLORA OF PANAMA (Family 93. Malpighiaceae) 855 k. Fruit a 3—2-locular, loculicidal capsule . 4. Galphimia kk. Fruit a drupe or nut, with 3-1 pyrenes. l. Styles subulate, acute; drupe monopyrenous, 3-locular, 3-seeded; cotyle- dons spirally involute; receptacle hirsute . Byrsonima ll. Styles spathulate, obtuse or truncate at apex; drupes (di-)tripyrenous; cot- at apex; styles free; flowers in small axillary corymbs or subum- bellate __ 8. Malpighia Petals yellow; bracteoles with a dorsal gland; pyrenes without crests or ribs; cotyledons straight, plano-convex; styles more or less united or free; flowers in axillary or terminal racemiform inflorescences ------------ . Bunchosia m B 1. BANISTERIOPSIS Banisteriopsis, C. B. Robinson in Small, N. Amer. Fl. 25: 131. 1910. TYPE: B. cornifolia (H.B.K.) C. B. Rob. (see Cuatrecasas, 1958, p. 487). Banisteria sensu auct. non L., H.B.K Lianas. Leaves opposite, (rarely ternate elsewhere), simple, entire, petiolate, coriaceous to membranaceous, commonly glandular along the margin or at the base; stipules interpetiolar, minute, caducous. Inflorescences of umbels, corymbs or rarely racemes arranged in bracteate or leafy, axillary or terminal panicles. Flowers pink, yellow or white; sepals 5, usually ovate, the petals 5, clawed, glabrous or sericeous outside, the limb orbicular or obovate, entire or fimbriate, frequently unequal, the anterior petal being more deeply fimbriate or of different size; stamens 10, frequently unequal, the 3 posterior stamens and the 2 epipetal- ous (lateral) stamens reduced in size, the anterior stamen sometimes much en- larged, the filaments frequently united at the base, the anthers glabrous or pilose (not Panama), the connective thick, glandular; styles generally equal, truncate or capitate, uniformly stigmatic, the ovary trilobate, gibbose, hirsute. Fruit a schi- zocarp of 3 samaras, these with a single dorsal wing, with a thickened ventral margin and a thin dorsal margin, lateral wings absent or poorly developed, the body smooth or with minute wings, tubercles or transverse ridges; embryo curved. Banisteriopsis is a neotropical genus with about 90 species, mostly lianas, more or less frequent in rain forest, subandean forest and savannas, usually below 1,000 m elevation. Some species, e.g. B. caapi (Spruce) Morton, B. rusbiana Ndzu. are well known as popular among natives because of their strong hallu- cinogenous effects. Banisteriopsis caapi especially is cultivated to a limited ex- tent by Indians in Brazil, Colombia, Ecuador and Peru and also by other people who occasionally have spread its cultivation outside of its original area. No spec- imens, spontaneous or planted of this species have been found in Panama. Much confusion has surrounded the nomenclature of plants now called Ban- isteriopsis. Linnaeus, in describing Banisteria in the first edition of Species Plan- tarum (1753), included 7 species which are now in 3 genera. Heteropteris was segregated by Kunth in H.B.K. (1822), Stigmaphyllon by Jussieu in 1833, and Banisteriopsis was later applied to the remaining species in the North American Flora by Small (1910), who based part of his work on an unpublished manuscript 856 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Banisteriopsis cornifolia (H.B.K.) C. B. Robinson ex Small. Fruiting branch ( ictorius (Roxb.) D'Arcy. Stem with leaves (x!⁄2). [After Tyson O).] FIGURE 4. Garcinia.—A. G. pi 6691 (MO).]—B. G. mangostana L. Fruit (xV5). [After D'Arcy & Foster 13308 (M Transfer of Xanthochymus Roxb. from genus to species level. Nom. illeg., cites Xanthochymus pictorius Roxb., X. tinctorius DC. G. tinctoria (DC.) Dunn in Gamble, Fl. Pres. Madras 74. 1915. Medium size or large tree; trunk straight, the branches drooping, angular; sap yellow. Leaves oblong, to narrowly obovate, apically acute or short acuminate, basally rounded, sometimes somewhat dimidiate, the costa broad, plane or im- 1000 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 pressed above, elevated beneath, coriaceous, shiny, often drying yellowish; pet- iole sharply angled with a basal enclosed pit, often drying yellow, the lateral veins arcuate, loop connected to form a submarginal vein 2-3 mm from the margin. Inflorescences several-flowered axillary fascicles; pedicels ca. 2.5 cm long. Flow- ers (from Maheshwari) white, ca. 1.5 cm across; sepals (4—)5, orbicular, concave, fleshy, unequal, apically ciliate; petals 5, ca. 8 mm long, orbicular, spreading, incurved, greenish; stamens in 5 broad bundles of 3-5 each, antipetalous, alter- nating with 5 fleshy glands, the anthers bilocular, pistillate flowers with stami- nodes few, complanate; ovary ovoid, acuminate, 5 locular, stigmas 5. Fruit bac- cate, ca. 6.5 cm across, subglobose, pointed, dark yellow with copious yellow gum when cut; seeds 1-4, imbedded in a yellow, aril like pulp. All collections of this species from Panama are from cultivated trees. On these specimens the fruit has dried conspicuously wrinkled, but it is reportedly smooth in vivo. Garcinia pictorius is a native of eastern India and Burma, and it is cultivated as an ornamental tree. The fruits can be used for making sherbets, preserves and jams. The gummy resins of the fruit have been used to make water colors. In India, the rootstocks are sometimes used for grafting on mangosteen (G. man- gostana). ZONE: Without other locality, Johansen 1924 (US). Summit Gardens, 0-100 m, Mori & Kallunki "1873 (MO). Balboa, Standley 30905 (US). Fort Clayton Officers Club, Tyson 6691 (MO). : Between Las Sabanas and Matias Hernandez, Standley 31800 (US). 6. HAVETIOPSIS Havetiopsis Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 246. 1860. LECTOTYPE: H. martii Pl. & Tr. Dioecious, epiphytic or free standing shrubs and trees. Leaves obovate, co- riaceous, short petiolate. /nflorescences short, mostly branching cymes, each divarication subtended by a pair of minute cucullate bracts. Flowers small, sepals 4, decussate; petals 4, decussate, thick, rotund; stamens 4, opposite the petals, the filaments basally thickened, narrowed under the anther, connate or free, the anthers ovate, 2-loculed, in pistillate flowers the anthers reduced or obsolete; styles 4, short. Fruit a capsule dehiscing into 4 valves; seeds few or many, arillate. This genus includes 3-4 species of Panama and tropical South America, which occur in lowlands or at middle elevations. It is not clear from the material at hand or from the literature that this genus is distinct from Havetia H.B.K. Vesque (1893) argued that the separation of the two genera by Planchon and Triana was based on a misinterpretation of the material and united the two. Engler (1925) however, maintained the two genera as distinct, but the distinctions he saw may not be valid. Should the two genera be united, then the species included here, Havetiopsis flexilis, would be known as Havetia flexilis (Pl. & Tr.) Vesque. 1. Havetiopsis flexilis Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 246. 1860. TYPE: Brazil, Spruce 3294 (P, not seen).—Fic. 5. Havetia flexilis (Pl. & Tr.) Vesque in DC., Monog. Phan. 8: 155. 1893; Epharm. 3, t. 72, 73. 1892. 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1001 KRAGER- PA MER — FiGure 5. Havetiopsis flexilis Pl. & Tr. Fruiting branch (x?/s). [After Folsom 3945 (MO).] 1002 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Dioecious epiphytic shrubs or small trees to 4 m tall; surface of youngest twigs sometimes wrinkling somewhat on drying (see Clusia minor above). Leaves obovate, mostly 6-10 cm long, 3—5 cm wide, apically rounded or emarginate, basally cuneate, the lateral nerves numerous, conspicuous or not, straight, ar- cuate or slightly recurving proximally, arcuate distally forming an even submar- ginal vein, minor venation mostly obscure, coriaceous, the margins drying slightly revolute; petioles ca. 5 mm long. Inflorescences short, mostly 1-3 cm long, sev- eral-flowered dichasial panicles, internodes angled, bracts scale like, cucullate, paired, thick, to 2 mm long or rudimentary; pedicels obsolete. Flower buds glo- bose, sepals 4, decussate, rotund, cucullate, the margins scarious, ca. 5 mm long, the outer pair shorter, basally united; petals 4, rotund, ca. 5 mm long, greenish marked with purplish; stamens 4, ca. 2 mm long, the filaments basally thickened, about as long as the anthers, the anthers introrse; in pistillate flowers the anthers reduced in size, the filaments connivent into a tetragonal column; styles 4. Fruit a fleshy capsule with 4 rounded angles, depressed globose or subcylindrical, to 1.5 cm long, styles persistent, erect, short, dehiscing into 4 valves, seeds nu- merous, orange, ca. 3.5 mm long, immersed in a red-orange matrix. Because of the paucity of material at hand, many details for the above de- scription were taken from Vesque (1893) and Croat (1978). This species ranges from Panama to Brazil, occurring in tropical wet and moist forests of low elevations. AL ZONE: Barro Colorado Island, Croat 6621, 8259, 12505, 14844 (all MO), 16195 (MO, SCZ); Foster 1100 beeen 1767 (DUKE, MO, PMA, SCZ). cHiRIQUí: Haras San Miguel just N of Con- cepción near Rio Mula, Folsom 3945 (MO). COLON: Al este de la zona montanosa de Santa Rita, Correa & Dre »ssler 764 (MO) 7. MAMMEA Mammea L., Sp. Pl. 512. 1753; Gen. Pl. ed. 5. 228. 1754. TYPE: M. americana L. oo Rottb., Abt. Lit. het Hafn. 1: 296. pa n P. mamei Rottb. ?Ochrocarpus Thouars, Gen. v. Madag. 15. 1806 O. madagascariensis DC. “espa sa Wight, Illustr. 1: 130. 1840. TYPE: C. Hae et Wight. Polygamous glabrous, tree; foliage dark green; sap yellow. Leaves opposite, entire, elliptical, coriaceous, the costa prominent, lateral veins numerous in two series, anastomosing only near the margin or forming a reticulum over most of the surface, submarginal vein not defined, minor venation forming a reticulum, evanescent towards the costa, the areoles punctate. /nflorescences of solitary flowers fascicles in the axils of mature leaves or mature wood; pedicels stout. Flower buds globose; sepals 2(-3), coriaceous; petals 4—6, white or pink; stamens numerous, crowded, basally united in several whorls, the anthers longitudinally dehiscent; ovary 2—4-locular, the style short, the stigma peltate 2—4-lobed; ovules 1-2 per locule, erect. Fruit a leathery berry, sometimes large, the exocarp, co- riaceous, the mesocarp orange or yellow, fleshy, sometimes copious; seeds large, oblong or slightly arcuate. Mammea includes one species widespread in the Caribbean area, a new species in Panama, and depending on circumscription, several species in Africa, 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1003 Madagascar and the Indomalay area. The two species treated here may be rec- ognized by their large warty fruits with orange pulp, and by the coriaceous leaves with lateral veins branching near the margin into a reticulum and with punctate areoles. The trees are of minor economic importance. The wood is used for general construction, and the heartwood is reported to have insecticidal proper- ties. The orange pith of the fruits of both species is edible. The seeds are toxic. The two species of Mammea recorded here have the largest fruits of any Guttiferae in Panama. Alexandre (1978) reported that similar fruits of M. africana are dispersed by elephants. The dispersal agent of the New World plants is un- known. The African element of Mammea, M. africana Oliver, has usually been rec- ognized in Mammea, but Vesque (1893) placed it in Ochrocarpus which is typ- ified by a plant from Madagascar where there are other species. Vesque based his placement on structure of the embryos, but he noted that in other respects many of these plants appear to be congeneric. The second species described here, M. immansueta, might fall into Ochrocarpus when data are available to apply Vesque’s criteria. Engler (1925) maintained the African species as Mammea. For controversy about separating Rheedia from Mammea see Grisebach (1861) and Planchon and Triana (1861). Here the justification for separation of the genera is based on the cotyledons and perianth. Rheedia lateriflora is superficially much like a species of Mammea. Literature: Alexandre, Y. D. 1978. Le róle disseminateur des éléphants en forët de Tai, Cóte-D'Ivoire. Terre & Vie 32: 47-72. Pittier, H. 1912. New or noteworthy plants from Colombia and Central America 3. Contr. U.S. Natl. Herb. 13: 431—466. Leaves pointed; leaves mostly more than 7 cm wide; petioles drying yellowish green; fruit seldom more than 7 cm across . immansueta aa. Leaves rounded; a mostly less than 7 cm wide; petioles drying reddish brown: ruits reaching 15 cm acros . M. americana e . Mammea americana L., Sp. Pl. 512. 1753. TYPE: Antilles, herb. Linn. (LINN 675.1, not seen; MO, microfiche). — Polygamous tree to 20 m tall, often narrow and densely leafy; sap yellow; twigs stout. Leaves broadly obovate or elliptical, apically rounded or slightly emarginate, basally rounded or obtuse, the margin minutely indurate, the costa subplane above, proximally canaliculate, conspicuously elevated and drying mi- nutely sulcate beneath; lateral nerves parallel, slightly ascending numerous in 2 similar series, 6-7 per cm (both series), anastomosing at the margin but not forming a submarginal vein, cross veins numerous, forming a rectangular retic- ulum, the areoles 1—2-punctate beneath, ca. 20 cm long, 10 cm wide; slightly discolorous, petiole stout, ca. 1 cm long, canaliculate above, the bases generally not connate or forming a petiolar ridge. Inflorescences 1-several flowers on twigs behind the leaves; pedicels stout, ca. 10 mm long. Flowers fragrant, the sepals 2, rotund, 10-15 mm long, cucullate, sometimes keeled coriaceous; petals ca. 6 1004 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 white, spreading, rotund, shorter or longer than the sepals, stamens numerous, the filaments glabrous, slender, basally expanded and united into several whorls, the anthers of 2 longitudinal thecae separated by the undifferentiated connective; ovary pyriform, the style short, the stigma peltate, repand, centrally depressed. Fruit a large leathery globose berry to ca. 15 cm across, with thick brownish warty rind and thick fleshy orange or yellow pulp (mesocarp); locules 2-4, filled with the large, oblong slightly arcuate seeds 2.5-4 cm long which are covered with reddish brown fibrous material. This species is sparingly cultivated in Panama, and Pittier (1912) reports hav- ing seen it wild in Veraguas. It is common in slightly drier regions of western Central America and in the Antilles, where it is sometimes cultivated as a spec- imen tree or for its fruit. ZONE: Cultivated, Barro Colorado Island, Croat 5787 (SCZ). PANAMA: Taboga Island, Standley "27054 (US). 2. Mammea immansueta D'Arcy.? TYPE: Panama, Mori et al. 4699 (MO, holo- type).—Fic. 6. Tree 30 m tall, twigs drying yellowish. Leaves clustered near the ends of twigs, elliptical, apically short-acuminate, basally obtuse, the margins sharp, sometimes slightly revolute, ca. 15 cm long, 7-8 cm wide, the costa mostly im- mersed above, broad and sulcate proximally, prominently elevated and somewhat sulcate beneath, the lateral veins numerous, parallel, equally spaced, in two se- ries, the primary series ca. 3 per cm, mostly unbranched in the first half, anas- tomosing and forming a reticulum in the region near the margin not forming a distal submarginal vein, obscure but slightly elevated and reticulated above, be- neath more conspicuous, the lateral veins and those near the margin slightly ele- vated, the minor reticulum plane, evanescent towards the costa, the areoles punc- tate, petiole 10-15 mm long, thick, drying yellowish, deeply canaliculate above. Inflorescence and flowers not seen, the fruits borne in pairs 7-8 cm apart along the branch (8-10 mm diam.). Fruits globose with a small apiculum, 4-7 mm across, warty, the pedicel becoming 7-10 mm long, 6-8 mm thick; the rind thick, the pulp scant (?) orange, 4-locular; seeds one in each locule, ca. 5 cm long, arcuate. This species is distinct from M. americana in its smaller fruits which have scant pulp and in its smaller, much narrower pointed leaves. It is closely related to M. africana Oliver, from which it may not be distinct. However, the leaves of M. immansueta are much wider than those of collections seen from the south- s" Mammea immansueta D'Arcy, spec. nov. Arbor. Folia elliptica, brevi-acuminata, coriacea, ca. 15 cm longa, 7-8 cm lata, costa subtus prominenti, nervis lateralibus seriebus duobus, primariis santibus, vena su i a grise: profunde canaliculato. Fructus globosus, brevi-apiculatus, verrucosus, 4—6 cm diametro; semini ibus quattuor, grandis, oblongis, uno in quoque loculo. 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1005 ern Congo and much more coriaceous than those from the. western Congo and Guinea regions. If it is in fact the same as the African species, the time and means of introduction into the New World are enigmatic. The collectors report it is a frequent element in the El Llano-Cartí transect across the isthmus. PANAMÁ: El Llano-Cartí road, Mori et al. 4699 (MO). 8. MARILA Will McDearman? and Sidney McDaniel? Marila Swartz, Prodr. Veg. Ind. Occ. 84. 1788. TYPE: M. racemosa Swartz. Shrubs or trees, the twigs laterally compressed, the plane of compression often decussate from successive nodes, with white, brown, yellow, or clear latex. Leaves evergreen, opposite, simple, entire, glabrous or variously pubescent, co- riaceous or nearly so, nerves impressed above, raised beneath, with more or less parallel secondary veins connecting the lateral, petiolate, estipulate. /nflores- cences racemose or paniculate, axillary at mostly defoliated nodes or rarely ter- minal; pedicels subtended by 3 minute bracts. Flowers bisexual or unisexual, 4— 5-merous; sepals in two series, usually differing in shape, texture and vestiture, persistent in fruit; petals membranous, fugaceous; stamens numerous, the fila- ments free, anthers linear, longitudinally dehiscent with a tubular, rotate or fili- form appendage at the end of the connective; ovary 3-5-celled, ovules numerous in each locule, attached to an axile placenta, the style simple, the stigma conical or nearly so. Fruit a septicidal capsule, in dehiscence the valves often twisting and revealing the papyraceous column; seeds minute, elliptic-oblong, comose at both ends. Marila is a distinctive genus of about 20 species of the neotropics, mostly of moderate to high rainfall areas, from Guatemala and the Antilles to Bolivia. It is questionable that the genus actually belongs in the Guttiferae, and it would prob- ably be better placed in the Bonnetiaceae. The genus is in need of a revision, though the species appear to be well marked. Many specimens of various species from Panama have been determined as Marila macrophylla Benth., which is not known to occur there. This species may be distinguished by the thicker pedicels (to 4 mm wide at base) and larger flowers. a. Leaves with 25-38 pairs of primary veins; anther appendage tubular to rotate. b Rachis of inflorescence with numerous, stalked, stellate trichomes, the surface red- 2. dish-brown; pedicels 4-7 mm long -------------------------------------------- M. lac eens bb. Rachis of inflorescence with mostly ier rarely stalked, stellate Seba t surface yellow-brown; E 11- ye ong |... 4. M. A rd aa. Leaves with 6-17 pairs of primary veins; ree appendage filifor Lower surface of dard vin cons ie cuous punctate glands; ra M: 1-2 mm phe base; capsules 4-7 cm long ------------------------------------------------------ 3. i laiita cc. Lower surface of a. w eni punctate pn META 0.5-1 mm een at base; capsules 1.2-2.5 cm long . n : _ 1. M. domingensis ? [nstitute for Botanical Exploration and Dept. of Botany, Mississippi State University, Missis- sippi State, Mississippi 39762. 1006 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 6. Mammea immansueta D'Arcy.—A. Habit (x!4).—B. Fruit (x!4). [After Mori et al. 4699 (MO).] — . Marila domingensis Urban, Ark. Bot. 22A(10): 17. 1929. Type: Haiti. Massif de la Hotte, ridge of Morne Delcour, Ekman H9001 (S, holotype, not seen; F, isotype). Small or medium sized tree to 7 m, younger branches puberulent or villous, becoming glabrate. Leaves 5-21 cm long, 2.5-6.8 cm wide, apically attenuate, acute or subabruptly acuminate, the acumen to 1 cm long, basally obtuse or rounded, glabrous, primary veins in 6-17 pairs, 0.6-1.5 cm apart, arcuately anas- 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1007 tomosing near the margin; petioles 0.7—2 cm long, longitudinally striate, villous. Inflorescence racemose, axillary at foliated or defoliated nodes, glabrous or pu- berulent to tomentose, 1—3 flowers per cm, rachis 4.5-8.5 cm long, the slender pedicels 0.6-2 cm long, 0.5—1 mm thick at base. Flowers white; sepals 6-7 mm long, ovate, minutely puberulent on the margins or glabrous, reflexed at anthesis, erect in fruit; petals 7-8 mm long, obovate; stamens numerous, anthers ca. 1.3 mm long, appendage filiform, the filaments to 2.5 mm long; ovary ca. 3.5 mm long, style 1.5-2 mm long. Fruit an obconical, often fusiform, brown capsule 1.2- 2.5 cm long; seeds 0.5-0.8 mm long. Marila domingensis is a very distinctive species, easily recognized by the short fruits and very slender pedicels. It has not until now been reported from Panama and was previously known only from Haiti. Specimens suggest that in Panama this species flowers in December and January and fruits in March and April. ANAMA: Carti Road, ca. 16 km, Kennedy 3076 (MO). El Llano-Cartí-Tupile Road, 12 mi above Panamerican Highway, 200—500 m, Liesner 1228 (MO). El Llano-Cartí Road, 10.8 km from Inter- american Highway, 1100-1200 ft, Mori & Kallunki 4115 (MO). El Llano-Cartí Road, 22.6 km from Interamerican Highway, 350 m, Mori & Kallunki 5099 (MO). SAN BLAS: El Llano-Cartí Road, 24.5- 25 km from Interamerican Highway near continental divide, Mori & Kallunki 5520 (MO). N . Marila lactogena Cuatr., An. Inst. Biol. Univ. Nac. México 20: 92. 1950. TYPE: Colombia: Depto. Valle, Río Naya, Puerto Merizalde, Cuatrecasas 14040 (F, holotype; US, isotype). Tree to !3 m, younger branches ferrugineous-tomentose, glabrescent; twigs with sticky white latex. Leaves 14.5—40 cm long, 5-14 cm wide, oblong-elliptic to oblong-lanceolate, apically more or less abruptly acuminate, the acumen to 1 cm long, basally rounded or obtuse, sparsely puberulent above except tomentose near the midvein, beneath reddish-brown with stalked stellate trichomes and mi- nute appressed scales, primary veins 35-38 pairs, 0.5—1.1 cm apart, united into a marginal vein, petioles 2.2-3.5 cm long, ferrugineous-tomentose, grooved above. Inflorescence racemose, axillary from older defoliated nodes or terminal on young branchlets, ferrugineous-tomentose with abundant, minute stalked stel- late trichomes, 2-8 flowers per cm, rachis 15-30 cm long, pedicels 4-7 mm long. Flowers of unknown color; sepals 6-8 mm long, reflexed at anthesis, erect in fruit, mostly oblong-ovate or elliptic, rarely lanceolate, tomentose; petals not seen; stamens numerous, the anthers 0.6-1.2 mm long, the appendage tubular to somewhat rotate, minutely pitted and glandular; ovary ca. 4 mm long, style 1 mm or less long at anthesis, elongating in fruit. Fruit a linear fusiform black capsule 1.1—2 cm long; seeds ca. 1 mm long. This species occurs in Panama and Colombia. This species has not previously been reported from Panama, but the material cited above agrees in all essentials with the type from Colombia. Marila lactogena in Panama would appear to be restricted to wet forest. PANAMA: El Llano-Cartí Road, 7-12 km from Panamerican Highway, 360—400 m, Croat 25113 (MO). 11 km from Escuela Agricola Alto de Piedra, in forest along Río Dos Bocas, Mori & Kallunki 3124 (MO). SAN BLAS: Forest SE of Puerto Obaldía, Croat 16813 (MO) 1008 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 o . Marila laxiflora Rusby, Mem. Torrey Bot. Club 6: 9. 1896. TYPE: Bolivia. Between Tipuani and Guanai, Rusby 1648 (NY, holotype, not seen; MO, isotype). M. verapazensis Donn. Sm. Bot. Gaz. 47: 253. 1909. rype: Guatemala. Alta Verapaz. Cubilquitz, Tuerckheim 1074 (US). Shrub or tree to 20 m, younger branches densely appressed glandular-pubes- cent, glabrescent, the twigs with brown or yellow-brown latex. Leaves 9-28 cm long, 2-12 cm wide, ovate, oblong, elliptic or elliptic-lanceolate, apically abruptly acuminate, the acumen to 1.5 cm long, basally rounded or acute, densely brown velutinous before unfolding, becoming glabrous, glabrate or sparsely pubescent along the veins beneath, primary veins 10-15 pairs, mostly 1-2 cm apart, ar- cuately anastomosing into a marginal vein; petioles 1—2 cm long, finely pubescent, grooved above. Inflorescence racemose, axillary from mostly older defoliated nodes, puberulent, 1-4 flowers per cm, rachis 12-30 cm long, pedicels 5-8 cm long, 1-2 cm thick at base. Flowers green, fragrant; sepals 6-8 mm long, ovate or elliptic, puberulent, reflexed at anthesis, but erect in fruit; petals 6-9 mm long, oblanceolate to oblanceolate-oblong; stamens numerous, the anthers ca. 1.5 mm long, the appendage filiform; ovary ca. 3 mm long, the style ca. 3 mm long. Fruit a linear, yellow-brown capsule 4-7 cm long; seeds ca. 1 mm long. Guatemala to Bolivia. There appear to be no essential differences between plants from throughout the range, those from northern Central America being indistinguishable from those of Bolivia. This species has the greatest distribution of any in the genus both in the neotropics and in Panama. It is a tree of wet forest of low to moderate elevations. ANAL ZONE: River bank between Fort Sherman and Fort San Lorenzo, pis 15437 (F, MO). EE March? 1850, Fendler (MO, US). Barro Colorado Island, Foster s.n. (MO). Pipeline Road, Lao et al. 17 (F, MO, SCZ). coci É: Bismark above Penonomé, Williams 568 m COLÓN: Achiote, Tyson et al. 4517 (DUKE, MO, SCZ). DARIEN: Río Pirre, Bristan 1467 (MO, US). Pinas, Duke 10641 , OS). Puerto St. Dorothea, Dwyer 2225 (MO). Slopes of Cerro Pirre, 1300 m, Mori 370 (MO). Caná, 2500 ft, Stern et al. 525 (MO, US). Periaque camp at river, Tyson et al. 4742 (MO, SCZ). PANAMÁ: Cerro Campana, Croat 14234 (MO). Cerro Jefe, 700 m, Dressler 3681 (DUKE, F, MO); 3100 ft, Dwyer a Gentry 9497 (MO). San José Island, Erisan 52 (US); Johnston 685 (MO, US), 808 (US). Cerro Campana, 2000 ft, Tyson 6435 (F, MO). 4. Marila pluricostata Standley & L. Williams, Ceiba 3: 52. 1952. TYPE: Costa Rica, Skutch 5420 (US, not seen; F, photo).—Fic. 7. Tree to 15 m, younger branches densely brown tomentose with numerous minute appressed lepidote scales, glabrescent, twigs with pale yellow latex. Leaves 11.5-26 cm long, to 40 cm long on sterile shoots, 3.5-9 cm wide, oblong- lanceolate, elliptic or lanceolate, acuminate, the acumen to 1.1 cm long, basally obtuse or somewhat rounded, glabrous above, beneath glabrate or more com- monly with minute sessile stellate hairs and appressed lepidote scales, the major veins with an occasional stalked stellate trichome, primary veins 25-32 pairs, 0.4-1 cm apart, united into a marginal vein; petioles 1-3.7 cm long, grooved above. Inflorescence racemose, axillary at foliated or older defoliated nodes, rarely terminal on younger branches, densely brown tomentose with abundant 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1009 FIGURE Marila pluricostata Standl. & Williams—A. Fruiting branch (x!⁄2). [After Mori & Kallunki 5077 (MO).]—B. Flowers (x15). [After Mori & Kallunki 5129 (MO).] minute sessile stellate trichomes and a few lepidote scales, ca. 4 flowers per cm; rachis 10-18 cm long, pedicels 11-12 mm long, elongating to 15 mm in fruit, ca. 2 mm thick at base. Flowers pink to bright magenta; sepals 9-12 mm long, ovate to ovate-lanceolate, externally densely brown tomentose with minute appressed sessile stellate trichomes and scales, internally glabrate, reflexed at anthesis and in fruit; petals 11-12 mm long, obovate; stamens numerous, the anthers ca. 1.5 mm long, the appendage tubular; ovary 5-6 mm long, the style ca. 2.5 mm long. Fruit a fusiform, frequently bent capsule 1.5-2.5 cm long; seeds ca. 1 mm long. 1010 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 This species occurs in Costa Rica and Panama. This species, like others of the genus in Panama, is confined to wet forest at relatively low elevations. A photo of the type of Marila cespedeziana Tr. & Pl. was examined. The inflorescence appears to be similar to that of this species, but the separate vegetative portion is not this species. It may not even be this genus but Vismia. Therefore the two species are not placed in synonymy. COLON: Santa Rita lumber road, Correa & Dressler 596 (DUKE, MO, SCZ). East Ridge, Duke 15259 (MO). Santa Rita Ridge Road, 9 km from Boyd-Roosevelt Highway, 350 m, Mori & Kallunki 5077 (MO). PANAMA: Between Cerro Azul and Cerro Jefe, Dressler 3271 (DUKE, MO, SCZ). El Llano-Carti Road, 18 km from Interamerican Highway, 350 m, Mori & Kallunki 4580, 5129 (both O). 9. RHEEDIA Rheedia L., Sp. Pl. 1193. 1753; Gen. Pl. ed. 5. 449. 1754. TYPE: R. lateriflora L. Verticillaria Ruiz & Pavon, Prod. Fl. Peruv. Chile 81, t. 15. 1794. TYPE: V. acuminata R. & P. = Rheedia se A & P.) PI. & :7 Chloromyron Pers., Syn. 3. 1807. TvPE: C. verticillatum Pers., superfluous name for Verticillaria acuminata R. & P. = e acuminata (R. & P.) Pl. & Tr. ag ci ror Miers, Trans. Linn. Soc. Lo pud 21: 249. 1854; Proc. Linn. Soc. London 2:883. 185 55]. TYPE: E e eni rai Sp. dubiu MM d & Per . Sci. Nat. Bot. Sér. 9. 11: 256. 1910. TYPE: T. pervillei (Pl. & Tr.) Jum. & Perrier = m sat E Pl. & Tr. Polygamodioecious trees; sap yellowish or white; twigs often angular. Leaves opposite (verticillate), entire, the costa prominent, the lateral nerves numerous, parallel, irregular, sometimes in 2 series, the minor venation reticulate, a sub- marginal vein usually present, at least distally, coriaceous, petioles with a basal wedge of tissue forming an axillary pit. Inflorescences of fasciculate or solitary flowers at the leaf axils, sometimes inserted into a bracteate annulus; pedicels often elongate and slender. Flowers small, the buds globose, sepals 2, imbricate, but somewhat united basally; petals 4, decussate and imbricate, the pairs often of slightly different size, the outer pair alternating with the sepals; the stamens numerous in staminate flowers, free, inserted beneath (abaxial to) or upon a fleshy, hemispherical or lobate disc, in perfect flowers the stamens fewer, the anthers subglobose, dorsifixed, longitudinally dehiscent; ovary wanting in sta- minate flowers, in hermaphrodite flowers 3—4-loculed, partly surrounded by the disc, the ovules 1 per locule, the style forming a stipe or not, the stigmas peltate, discoid or hemispheric, entire or lobed. Fruit baccate, the pericarp leathery, smooth or tuberculate, 1-celled; seeds 1-4, faboid in shape, enveloped in a pulpy aril. Rheedia is a genus of forest trees with perhaps two dozen species ranging through tropical America and Madagascar and is probably congeneric with a number of species in Africa now placed in Garcinia. Recently Robson (1958) and Adams (1970) united Rheedia with Garcinia noting that the affinities of Rheedia are with the African species of Garcinia sect. Teracentrum Pierre and sect. Rheediopsis. These two sections form subg. Rheediopsis Vesque, which is dis- tinct from other species of Garcinia in a number of respects including leaf anat- omy and which has been remarked on by Vesque (1892a, 1892b, 1893) and Engler 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1011 (1925) as being closely related to Rheedia. Rather than uniting Rheedia with Garcinia as was advocated by Robson and Adams, it is probably more correct to separate subg. Rheediopsis from Garcinia and unite it with Rheedia. The other two subgenera of Garcinia, subg. Garcinia and subg. Xanthochymus, typ- ified by G. mangostana and G. pictorius respectively, both represented in this flora, are different from Rheedia in leaf architecture, histology and general ap- pearance and in complexity of the flowers. Lamprophyllum Miers is maintained here under Rheedia following Vesque (1893) and not under Calophyllum L., where it is listed in the Index Nominum Genericorum. In his first discussion of the genus, Miers considered Lampro- phyllum in apposition to Calophyllum, to C. calaba L. specifically, and the fruit he described, which is the basis for the species L. laetum, is like Rheedia in having a '“thick mucilaginous pulpy arillus” and not at all like Calophyllum which has no distinguishable aril. In his second discussion of the genus, Miers included Calophyllum, but he considered C. calaba to be distinct from Lamprophyllum laetum. Vesque may have seen material annotated by Miers. Literature: Adams, C. D. 1970. Miscellaneous additions and revisions to the flowering plants of Jamaica. Phytologia 20: 309-314. Robson, N. 1958. Guttiferae. In New and little known species from the Flora Zambesiaca area. Bull. Soc. Brot. 32: 170-173. Vesque, J. 1892a. Histoire des Garcinia du sous-genre Xanthochymus. Compt. nd. Hebd. Séances Acad. Sci. 114: 562—564. . 1892b. Histoire des Garcinia du sous-genre Rheediopsis. Compt. Rend. Hebd. Séances Acad. Sci. 114: 844-846. » Fruits tuberculate; leaves drying with the submarginal vein at the margin but Pa from it with the naked eye, ion margin thus appearing sharp, lateral veins 5—6 per cm, ca. 12 including ee E 4 R. acuminata aa. Fruits s sm mooth; leaves with e submarginal vein either pipe: ip from the margin when viewed hon ove or more than 2 mm from it, the margin thus appearing blunt or Mind lateral veins fewer than 4 per cm, fewer than 7 se cm including intermediate vei s. Fruit mostly less than 3 cm long; leaves mostly much more than twice as long as 2. R. edulis bb. Fruit mostly more than 3 cm long; leaves mostly about twice as long as c. s drying — Rd angled (Fig. ich leaves drying brown, “asua all but shiny abov mem ny a , not lineolate beneath . R. macrophylla cc. Stems drying modb erat tely sulcate or is (Fig. 8B); leaves drying Dad ve mostly dull above, recht beneath . lateriflora — . Rheedia acuminata (R. & P.) Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 314. 1860 d acuminata R. & P., Syst. Fl. Peruv. 140. 1798. TYPE: Peru, Ruiz (B, if extant, not seen; hoto). Not Gants acuminata Pl. & Tr. (1860). Chloromyron verticillatum Pers., Syn. 2: 73. 1807. Based on Verticillaria acuminata R. & P. Calophyllum madruno H.B.K., Nov. Gen. & Sp. 5: 202. 1822. TYPE: Colombia, Bonpland (P, not i , photo). ?Garcinia floribunda Miq., Stirp. Surinam. Select. 89. 1850. TYPE: Surinam, Kappler 593, not seen. ?Rheedia floribunda (Miq.) Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 319. 1860. 1012 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 R. rostrata Miers ex Vesque, Epharm. 2: t. 72. 1889; in A. DC., Monog. Phan. 8: 510. 1893. R. acuminata var. floribunda (Miq.) Vesque in A. DC., Monog. Phan. 8: 510. 1893. Tree to 10 m tall; sap yellow; glabrous except the fruit; twigs slender, drying green-sulcate. Leaves elliptical or obovate, often drying bluish-green, 10-20 cm long, 4-6 cm wide, acute at each end, sometimes slightly umbonate and falcate apically, the margin appearing sharp, the costa elevated on both sides, sharp beneath, the lateral veins numerous, 5-6 per cm, ca. 12 per cm including inter- mediate veins, the lateral veins parallel to near the margin then becoming retic- ulate, elevated on both sides, mostly conspicuous, the submarginal vein ca. 0.5 mm from the margin, evident on drying; petiole 10-15 mm long. Inflorescences fasciculate in the leaf axils; pedicels slender, 15-20 mm long. Flowers polyga- mous, ca. 6 mm across; sepals 2, ca. 3 mm long; petals 4, white, rotund, cucullate, 5—7 mm long, reflexed; stamens numerous, the filaments free, inserted in several whorls, the anthers yellow, minute, globose to cordate, the thecas opening trans- versely to form an apical slit; ovary inserted in an annular, 1.5-2 mm-thick disc, umbonate, the stigmas forming a hemispherical cap on the style (ovary-umbo). Fruit ellipsoidal, mostly beaked at one or both ends, sometimes oblique, densely covered with soft, tomentulose tubercules, 4-5 cm long, the mesocarp soft, sweet; seeds mostly 2, discoid, 2-3 cm long, enclosed in a reddish brown, chartaceous (dried) septum. This species ranges from Costa Rica to Peru. It is found in tropical wet and moist forests, chiefly as an understory tree. Rheedia acuminata may be distin- guished from other Panamanian species by its tuberculate fruit, its numerous veins, its usually smaller and narrower leaves, and by the marginal vein which may be distinguished with the naked eye only 0.5 mm from the margin. In this species the lateral veins are slightly ascending and slightly arcuate to near the margin. The cross veins appear to be continuous, forming a vein-like series of lines more strongly ascending and arcuate than the lateral veins. A number of African species currently recognized under Garcinia have this series of cross veins more prominent than the lateral veins, an interesting morphological development which emphasizes the affinity of these African elements to neo- tropical members of Rheedia. ZONE: Barro Colorado Island, Croat 8240, 10840 (both Ped SCZ); Foster 1780 (DUKE, PMA): cal 431 (MO, US); Kenoyer 669 (US); Shattuck 668 (MO, US); Standley 31304, 40792 (both US); Starry 46 (MO); Wetmore & Abbé 112 (MO); Zetek 2 (MO), 4627 (MO, SCZ, US). Pipeline Road, Gentry 7398 (MO). Frijoles, Pittier 3750 (US). Quebrada Fea, tributary of Río Chagres, Smith & Smith 3402 (PMA). cuiRiQUÍ: Cerro Horqueta, 1200-1700 m, Pittier 3162 (US). COCLÉ: Santa Rita Ridge, Croat 13847 i ed MO). Salud Hills, Lao & Holdridge 190 (MO, PMA). DARIÉN: Cerro Pirre, Bristan 571 (MO). Paya, Stern et al. 393 (US). PANAMA: Cerro Azul, Dwyer 1344 (MO), 1364 (SCZ). Altos de Pacora, Mori & Kallunki 4971, 2566 (both MO). 2. Rheedia edulis (Seem.) Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 310. 1860.— Fic. 8A eic edule Seem., Bot. Voy. Herald 89. 1853. TYPE: Veraguas, Panama, Seemann (BM, not ; MO, photo). Glabrous trees to 10 m tall; sap yellow; twigs drying conspicuously ridged, older branches often somewhat reddish, sometimes smooth and grayish. Leaves 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1013 ASTA LAO a "< x PER SENNA CNN Gr | N< ç Sky Ns — KS WY” NS S< "FE SUN 2.5 Y LIENS C i ELE FiGuRE 8. Rheedia—A. R. edulis (Seem.) Pl. & Tr. Fruiting branch (x?/s). [After Croat 5602 (SCZ).—B. R. lateriflora L. Stem showing ridging (x?/s). [After Croat 4468 (MO).]—C. R. macro- phylla (Mart.) Pl. & Tr. Stem showing ridging ( x?/3). [After Rodriguez 426 (MO).] 1014 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 coriaceous, narrowly elliptical, mostly 15-25 cm long, 5-7 cm wide, apically and basally acute, the margin drying slightly revolute, the costa prominent, elevated above and beneath, above rounded or slightly flattened proximally, angled be- neath, the lateral veins numerous, parallel, mostly well spaced, ca. 3 per cm, ca. 6 per cm including intermediate veins, the lateral veins anastomosing and forming a reticulum near the margin, the minor venation obscure above, slightly elevated beneath, the submarginal vein not distinguishable from the margin when viewed from above; petiole mostly 10-15 mm long, at first drying reddish brown and fine wrinkled, later darker and somewhat grayish brown, canaliculate above, the mar- gins sharp-angled, sometimes the pairs forming a nodal ridge. Flowers polyga- mous, fasciculate at the nodes of fallen leaves, the fascicles many-flowered; ped- icels slender, 5-8 mm long, longer in fruit. Flowers ca. 5 mm across, sepals 2, coriaceous, cucullate, apically erose to repand; corolla white, the petals 4, the limb reflexing over the sepals, round-keeled and erose margined, 4 mm long, decussate, the outer pair alternate with the sepals; stamens ca. 15 arranged cy- clically or in 2-3 whorls, the filaments free, slender, glabrous, ca. 2 mm long, basally thickened at insertion, the anthers 2-thecate, the thecas ellipsoidal, ca. 0.2 mm long, dorsifixed, not versatile, the thecas dorsally connate, drying red- brown, the longitudinal dehiscence forming an elliptical pore; pistil small, the ovary not evident at anthesis, covered by the sessile stigmas, the stigmas irreg- ularly curved, ca. 1.5 mm across. Fruit a globose or ellipsoidal red or orange drupe 1.5-2.5 cm across, often in large clusters, smooth, minutely apiculate; seeds faboid, dark red, 1.5-2 cm long, 9-10 mm wide, the mesocarp white, ad- hering to the seeds, the seed enclosed in a reddish fibrous material. Rheedia edulis ranges from Mexico to Panama. It is distinguished by its small, smooth, yellow or orange fruits, and by its pointed narrow leaves. The venation appears to be more widely spaced and less prominent than in leaves of comparable size on other Panamanian species. In Panama, Rheedia edulis grows in the under- story of forests in moist and wet lowlands. The fruit is commonly eaten by man and monkeys, but they do not often become a market item. BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 6a (US). CANAL ZONE: Barro Colorado Island, Croat 5602 (DUKE, SCZ), 8354, 11129 (both SCZ), 13921 (DUKE, PMA, SCZ), 14454, 14652 (both SCZ); Foster 561 (PMA), 2184, 2209 (both DUKE, PMA); Hladik 400 (US); Shattuck 1102 (US); Zetek 3758 (MO). Madden cave area, Tyson et al. 6246 (PMA). CHIRIQUI: Progreso, Cooper & Slater 160 (US). Sabana de la Tortuga between Boquete and Caldera, 300-700 m, Pittier 3344 US). Between Cerro Vaca and Hato del Loro, 850-1100 m, Pittier 5399 (US). cocrÉ: Hills S of El Valle de Antón, 700 m, Allen 2474 (US). Penonomé, Williams 378 (US). DARIÉN: Cerro Pirre, Bristan 568 (MO). Rio Uruceca, Bristan 1458 (MO). N of Santa Fé, Duke 10242 (MO). Near Estero Grande off Río Marea, Duke 10959 (MO). Without other locality, Tyson et al. 4752 (DUKE). 3 mi E of Santa Fé, Tyson et al. 4671 (MO, PMA, SCZ, US). HERRERA: 4 mi S of Los Pozos, Tyson 2647 (DUKE, Z). LOS SANTOS: Azuero Peninsula between Jobero and headwaters of Río Pedregal, 300—700 m, Croat 34462 (MO). Tonosí, Stern et al. 33650 (US). 11 mi N of Tonosí, Tyson et al. 2981 (SCZ). PANAMA: El Llano, Duke 5809 (MO). Cerro Jefe, Foster & Kennedy 1825 (DUKE, PMA). 5 mi W of Chepo near ce iie Sey hway, Tyson 6703 (PMA). VERAGUAS: La Yeguada, Calobre 653 m, riis 601 (MO). N of Santa Fé, N of Escuela Agricola Alto de Piedra, Mori & Kallunki 2566 (MO), 3721 (MO, SCZ), 4893 oda 3. Rheedia lateriflora L., Sp. Pl. 1193. 1753. Based on Vanrheedia Plumier, Nov. Pl. 45. 1703, not Garcinia lateriflora Blume (1825).—Fic. 8B. 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1015 Vanrheedia Plumier, Nov. Pl. 45. 1703. TYPE: Martinique, Plumier (P, fide Pl. & Tr., not seen). Mammea humilis Vahl, Eclog. Amer. 2: 40, t. 20. 1798. TYPE: Montserrat, Ryan (? C, not seen; MO, icrofiche). Rheedia sieberi Choisy, Descr. Guttif. Inde 47, t. 5. 1849. type: Trinidad, Sieber 56 (MO). Mammea humilis var. vahlii Griseb., Fl. Brit. W. Indies 108. 1859. Based on M. humilis Vahl. M. humilis var. plumieri Griseb., Fl. Brit. W. Indies 108. 1859. Based on Rheedia Plumier, Pl. Amer. 255, t. 257. 1760. Rheedia d Planch. ex Vesque, Epharm. 2, t. 62. 1889; in A. DC. Monog. Phan. 8: 50. 1893. TYPE: aica, Purdie (K, not seen Garcinia humilis (Vahl) C. D. Adams, Phytologia 20: 312. 1970. Glabrous shrub or tree to 10 m tall; sap yellow; twigs stout, drying moderately sulcate. Leaves coriaceous, thick, ovate or elliptical, 15-28 cm long, 6-12 cm wide, apically obtuse or acute, often umbonate, the tips falcate, basally obtuse or rounded, the margin often slightly revolute, the costa prominent and sharp- angled beneath, plane or slightly raised above, the lateral venation prominent, ca. 3 veins per cm, 5-6 per cm including intermediate veins, little branched to near the margin and then forming a loose, open reticulum, elevated on both sides, the submarginal vein scarcely distinguishable from the margin; petiole 1-2 cm long, stout. /nflorescences fasciculate in the axils of fallen leaves, drying reddish brown; pedicels 7-10 mm long, slightly longer and stouter in fruit. Flowers 9-15 mm across; sepals 2(-3-4), 1.5-2 mm long; petals white or yellowish, petals 4, 6-8 mm long, rotund, cucullate, slightly coriaceous, slightly unequal; stamens 12-25, the filaments free, the anthers minute, opening longitudinally by broad apical slits; style short or obsolete, the stigmas 3—4, covering the young ovary. Fruit ellipsoidal to globose, yellow, acuminate-beaked at each end, 3-4 cm long; seeds 1-3, ca. 2.5 cm long. This species ranges through the Antilles from Hispaniola to Trinidad. It is known to Panama from only a few collections, taken in wet forest near the Ca- ribbean coast. Rheedia humilis may be distinguished by its large thick coriaceous leaves and large, smooth, beaked, yellow fruit. The leaves are similar to those of Garcinia mangostana but only one submarginal vein is present, situated right at the margin, not two such veins situated well in from the margin. Rheedia lateriflora is highly suggestive of Mammea, and in fact, Grisebach united the genus Rheedia with Mammea based on the appearance of this species. In the controversy following (Grisebach, 1861; Planchon & Triana, 1861) Plan- chon and Triana recorded that the cotyledons and the perianths in the two genera are quite different and that this species does indeed belong in a distinct genus, Rheedia, Planchon and Triana placed Rheedia and Mammea in different tribes. COCLÉ: N of El Copé, 750-900 m, Croat 44648 (MO); Folsom 1347 (MO). 4. Rheedia macrophylla (Mart.) Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 309. 1860.—Fic. 8C. Garcinia macrophylla Mart., Flora 24. 2. Beibl. 35. 1841. type: Brasil, Martius, not see Rheedia benthamiantha Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 320. 1860. SYNTYPES: : Guyana, Schomburgk 523, 990 (?P, neither seen). 1016 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 R. — Engler in Mart., Fl. Brasil. 12(1): 460, t. 702. 1908. TYPE: Guyana, Sagot 1182 (W, not n; , photo R. magnifolia oum U.S. Natl. Herb. 13: 454. 1912. TYPE: Costa Rica, Pittier, Aug. 1901 S-578033, not seen). Tree to 10 m tall; twigs drying strongly angled sulcate; sap yellow. Leaves ovate, apically short acuminate, basally truncate or obtuse, 15-40 cm long, 7-15 cm wide, stiff coriaceous, concolorous, mostly drying brown, shiny above, the costa plane or slightly impressed above, elevated and angled beneath, lateral veins mostly 4—10 mm apart, irregularly spaced, arcuate near the margin forming n inconspicuous submarginal vein 1-2 mm from the margin; petioles stout, often triangular, 1-2 cm long but sometimes wanting, with a conspicuous flap of axillary tissue forming a pit, the surface drying yellowish crinkled. /nflorescence fascic- ulate at the nodes or leaf axils; pedicels lax, ca. 2 cm long. Flowers (Pittier) small, pale yellow, sepals 2, 5 mm long; petals 4, ovate, 11-12 mm long; stamens numerous, ca. 30, inserted at the base of the disc, the anthers 0.5 mm long; disc ca. 4.5 mm across, hemispherical, muriculate; pistillate flower with ca. 25 sta- mens, perhaps biseriate, the ovary turbinoid drawn into a pronounced stout style, the stigma flat, peltate. Fruit yellow or orange, turbinoid, lemon shaped, ellipsoid or globose, smooth or rough outside, apiculate or the stigmas depressed, the pulp thick; seeds 1-4, ellipsoidal or faboid. This species is distinct in its large, stiff, coriaceous leaves which usually dry shiny above. In plants from near Puerto Obaldía, some leaves are sessile while others on the same tree have petioles to ca. 2 cm long. The fruit is variable in shape, from globose to lemon shaped, and the seeds vary from 1 to 4 (?5). There is also a strong tendency for the stems to dry angled sulcate and nearly hollow. The name Rheedia macrophylla is used for this species with hesitation, and all the Panamanian material cited here may not be the same species. A photograph of the type collection reveals narrower leaves: the type of Rheedia sagotiana is more like the material described here. This species may be difficult to separate from some collections of Garcinia pictorius which also has similar fruits. The leaves of G. pictorius are usually narrow, mostly oblong rather than ovate. Rheedia macrophylla leaves generally have shorter and stouter petioles than those of R. lateriflora, and the veins are usually farther apart. COLÓN: Santa Rita peek road, 14 km from d Roosevelt Highway, Mori & Kallunki 4923 (MO). PANAMA: Cerro Azul, Rodriguez 426 (MO, A). SAN BLAS: Hills of Sperdi near Puerto Obaldía, 20-200 m, Pittier 4343 (US). Coastal bu gere Puerto Obaldía and Puerto Armila, D'Arcy & Hahn 13673 (MO). 10. SYMPHONIA Symphonia L. f., Suppl. 302. 1781. TYPE: S. globulifera L. f. Trees or shrubs; sap yellowish, glabrate. Leaves opposite, mostly apically acuminate (Panama) or sometimes rounded, pinnately nerved; stipules wanting. Inflorescences few-flowered cymes or solitary flowers, often condensed, terminal or axillary. Flowers perfect, red or pink; sepals 5, imbricate, the outer 2 smaller; petals 5, contorted, arranged in a globose bud; disc extrastaminal, cupulate; fil- 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1017 aments forming a column apically separating into 5 parts opposite the petals, each staminal element bearing 2—5 anthers, the anthers linear, dehiscent by lateral slits; ovary 5-locular, the ovules 2—8 per carpel, ascending, the style elongate, peg-like, apically 5-lobed, the stigmas minute, porate. Fruit pink or red, baccate, lobose or ovoid few seeded, lacking endosperm, the embryo thick, entire and lacking cotyledons. Symphonia, the only genus in tribe Moronobeae occurring in Panama, is dis- tinct in its perfect flowers, contorted sepals, connate androecium, and globose red or pink flowers and fruits. The genus is represented in the New World tropics, Africa and Madagascar by S. globulifera, and in Madagascar more than a dozen endemic species have been recorded, some of which may be valid. The sap is used for caulking and the timber for construction. 1. Symphonia globulifera L. f., Suppl. 302. 1781. TYPE: Surinam, Dalberg (LINN 853.1, not seen; MO, microfiche).—Fic. 9. Tree to 30 m tall; sometimes stilt-rooted; sap yellow. Leaves oblong to lan- ceolate, often slightly wider above the middle, apically acuminate, basally obtuse or acute, 4-6 cm long, 1.5-3 cm wide, the costa impressed above, elevated be- neath, the lateral nerves numerous, straight, parallel, slightly ascending, often abruptly forking near the margin to form an inconspicuous continuous submar- ginal vein, chartaceous, often slightly grayish; petiole short but distinct, 3-5 mm long. Inflorescence condensed and subumbellate, cymose peduncle mostly ob- solete but sometimes 2-3 cm long; bracts wanting; bracteoles obsolete; pedicels straight, ca. 1 cm long. Flowers depressed globose, pink or red, ca. 1.5 cm across; sepals 5, quincuncial, rotund, broader than long, ca. 5 mm long; petals rotund, ca. 10 mm long, imbricate, contorted; filaments exserted, forming a narrowly turbinoid column around the ovary, apically splitting into 5, white strap-like ele- ments, the anthers 3 per segment, linear, abaxial on the segments, dehiscing longitudinally by lateral slits; ovary 5-locular, turbinoid, produced into the style, the style apically divided into 5 porrect linear lobes which alternate with the anther connectives, white, stigmas minute, distal on the style lobes and directed downwards, minute. Fruit subglobose, 2—4 cm long, surmounted by the persistent style and branches, leathery, brownish or yellowish; seeds 1(-3), red, ellipsoid to subglobose. This species occurs as an understory shrub but ultimately reaches large size becoming one of the largest forest trees. The pink or red globose flowers, which are often very abundant, are conspicuous. The yellowish sap of this tree dries into a dark gum which is sometimes used for caulking boats. The timber is suitable for veneers and rough carpentry. Symphonia globulifera is widespread in tropical America, Africa and Mada- gascar, showing little variation throughout its range. In Panama it occurs mostly along the Caribbean watershed at lower elevations. S DEL TORO: Dwedulup Peak N of Rio Terebe across from Quebrada Huron, 300—900 ft, BOCA Kirkbride & Duke 557 (MO). Shores of Laguna de Chiriqui, 150 ft, Stern & Chambers 51607 (MO). Water Valley, Wedel 758, 915 (both MO). Chiriqui Lagoon, Wedel 1030, 1100, 1141, 1210 (all MO). 1018 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 URE 9. Symphonia globulifera L.f.—A. Flowering branch (x ⁄s). Bx Mori & Kallunki 2860 (MO). —B. Fruiting branch (x?/s). [After Dressler & Williams 3964 (MO).] Water Valley, Wedel 1463 (MO). Little Bocas, Wedel 2496 (MO). CANAL ZONE: Barro Colorado Island, Croat 8686 (DUKE, MO, SCZ), 9517 (MO); Foster 943 (PMA). cociÉ: Albo Calvario [Above El Copé], 600—800 m Si 3174 (MO). eei Cerro Santa Rita, 1200-1500 ft, Allen 5101 (MO); aska & Dressler 108 5, 1205 (both DUKE, MO, PMA); Croat 15337 (MO); Dressler 3412 (PMA). Santa Rita lumber road, Kia ossler & DE. 3964 (MO). Cerro Santa Rita, 1200-1500 ft, Dwyer 8580 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1019 (MO, SCZ). Santa Rita lumber road, ae 8584 (MO). Cerro Santa Rita, 1200-1500 ft, Foster 1738 (DUKE, MO, PMA); Gentry 6557 (MO, PMA). Salud, Lao & Holdridge 240 (DUKE, MO, PMA). DARIEN: Cerro Tacarcuna, west ridge ; Gent try & Mori 14116 (MO). PANAMA: Cerro Azul, Dwyer 1507 (MO). 5-6 mi N of El Llano, Gentry 5814 (DUKE, MO). Road from El Llano to Carti, Kennedy & Dressler 2951 (MO). El Llano-Cartí road, 350 m, Mori & Kallunki 2860 (MO), 3539 (MO, PMA). SAN BLAS: Trail E of Cangandi-Mandinga airport road, Duke 14787 (MO, OS). vERAGUAS: Cerro Tute, Mori 6268 (MO). 11. TOVOMITA Tovomita Aubl.,!° Hist. Pl. Guiane Franç. 2: 956, t. 364. 1775. TYPE: T. guianensis Aubl Monoecious or dioecious trees or shrubs, glabrate, stems sometimes but- tressed or stilt rooted, sap often white or yellowish. Leaves at first opposite, soon becoming alternate, entire, simple, drying minutely revolute, the costa el- evated beneath, lateral nerves sometimes forming a submarginal vein; petioles sometimes short, flat or canaliculate above; stipules or pseudostipules wanting, axillary buds not normally conspicuous. Inflorescences cymose panicles, mostly terminal, often pyramidal, basally subtended by a pair of minute to foliaceous bracts which rarely envelop the entire inflorescence in bud, the branching mostly opposite or ternate, bracteoles mostly present, small and cucullate, subtending the pedicels and parts of the rachis; pedicels often thickening but not much elon- gating in fruit. Flowers with the buds minutely apiculate, completely enclosed in the outer sepals which are fused to the top showing only indistinct sutures, split- ting lengthwise into 2-4 opposite, cucullate, coriaceous segments (sepals), an inner whorl of sepals sometimes present, the complete calyx consisting of 2-8 decussate sepals, the inner series imbricate apically and overtopping the bud; corolla sometimes fused in bud and splitting lengthwise to yield the 2-8 segments (petals), or the petals free and distinct in bud, imbricate and overtopping the bud, opposite and decussate; stamens numerous, the filaments free, slender or stout, glabrous, the anthers small, the 2 lobes separated by the connective, each lobe 2-locular with a thin septum, the connective sometimes produced into a minute apiculum; ovary conical or obtuse, sometimes sulcate, 4—5-carpellate, syncar- pous, the styles short or produced into prongs in fruit, the stigmas ovoid to discoid, stigmatic abaxially; ovules | in each locule, axile, medifixed, anatropous or perhaps campylotropous, enveloped in a fleshy, venose aril, the aril and lower portion of the ovule sometimes produced into a spur filling a cavity in the base of the locule. Fruit a fleshy, drupaceous capsule, dehiscent septicidally to the base, often brightly colored within; seeds 1 per locule or fewer by abortion, oblong, ellipsoidal or faboid in shape, enveloped by the fleshy aril, the testa wanting, one end sometimes produced into a basal spur. Tovomita is a genus of one or two dozen species from tropical South America and the Antilles ranging as far west as Panama (?Costa Rica). Although it has been confused with Tovomitopsis, Chrysochlamys, and Dystovomita, it is ap- parently distinct in several features. The outer perianth is complete in preflora- 10 A number of synonyms of Tovomita were published in the last century; only this name has been used for Panamanian plants. 1020 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 tion, that is, it is completely fused and the sepals are schizogenous products of longitudinal tearing. The ovules are reportedly anatropous but appear to be cam- plyotropous instead, a feature which could not be verified at this time. They are inserted about halfway up the carpel wall, and the funiculus joins the ovule at about the middle with the raphe directed downwards. In the case of Tovomita weddelliana and perhaps other species not studied at this time, this lower end of the ovule is produced onto an acute spur which fills an elongation of the locule into the basal region of the carpel. An aril completely envelops the seed, which is fixed at the point of ovule attachment but is otherwise free. This aril is con- spicuously venose, the veins running outwards from the point of attachment. The testa is extremely thin or wanting. According to Engler, to Planchon and Triana, and other earlier writers, the fleshy structure in Tovomita is not a true aril, and they preferred the term arilloid. It is actually a fleshy elaboration of the integu- ments or a sarcotesta and not an outgrowth of the funiculus as in related genera where the term aril may be applied in a strict sense. The term aril is used here in the ecological sense to cover any fleshy, colored seed appendage deployed as an attractant for dispersal. The species of Tovomita occurring in Panama are quite distinct from one another; two of them, 7. stylosa and T. weddelliana, are also distinct in the genus and atypical of the majority of species. In addition to the three species noted below, material of at least two other species has been collected in Panama. These could not be assigned to any known species, but material is not sufficient to warrant description of new species. a. Leaves appearing sessile but at the very base abruptly narrowed into short, stout petioles. eaves narrow, the lateral veins numerous without cross veins, less than 3 mm apart; fruit obturbinoid, smooth 4. T. weddelliana bb. Leaves broad, the lateral veins well spaced with evident cross veins, more than 8 mm apart; fruit globose, warty 2. T. morii aa. Leaves narrowed into slender petioles c. Flower buds d narrow; drying green; leaves membranaceous or chartaceous; stamens fewer than 3. T. stylosa cc. Flower buds n. or ovoid, drying brown; leaves coriaceous; stamens rà 80 . . T. longifolia 1. Tovomita longifolia (L. C. Rich.) Hochr., Ann. Conserv. Jard. Bot. Geneve 21: 66. 1919 Clusia longifolia L. C. Rich., Actes Soc. Hist. Nat. Paris 1: 113. 1792. TYPE: Guiana, Richard (?P, ot seen; ?G, ?G-DC neither seen; MO, photos pase clusioides Choisy, Mém. Soc. Hist. Nat. Paris 1: 224, t. 11, 12. 1822 [1823] pro parte, excl. ty Tovomita richardiana Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 273. 1860. New name for Clusia longifolia L. C. Rich. Stilted tree to 15 m tall, twigs stout, new growth drying yellowish, older growth drying gray, glabrous; latex cream or yellow. Leaves somewhat coria- ceous, obovate, to 20 cm long, 8 cm wide, apically obtuse or rounded, basally acuminate or obtuse, minutely revolute, the costa nearly plane above, elevated beneath, the lateral veins 6-7, arcuate-ascending, wider spaced upwards, as- cending at the margin but not forming a submarginal vein, minor venation ob- scure, consisting of 1—2 ill-spaced intermediate lateral veins and numerous veins 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1021 pinnate on the laterals, linear canals sometimes evident beneath; petiole 10—40 cm long, ca. 2 mm wide, nearly flat above. Inflorescences terminal, pyramidal to rotund, subtended by pairs of normal leaves; rachis stout, drying yellowish brown, to ca. 6 cm long; bracteoles deltoid or obtuse, ca. 2 mm long, the margins hyaline; pedicels 5-8 mm long, stout. Flowers unisexual, the buds ovate to glo- bose, the minute sepal tips forming an apicule, ca. 7 mm long, reddish brown, finely striate, calyx prefloration complete, splitting to the base into 2 outer sepals, the sepals reflexed, boatlike or spathulate, ca. 8 mm long, coriaceous, the 2 inner sepals similar but slightly larger with hyaline margins; petals 4—7, thinner, nar- rower; stamens numerous (ca. 80), the filaments 2-4 mm long, slender, subterete, erect in bud, the anthers elongate, the lobes parallel, the connective slightly emergent and apiculate; ovary not examined (obsolete in staminate flowers). Fruit ultimately subglobose or pyriform, 5 cm long, 3 cm across, sulcate, apiculate, the apex prolonged by the 5 connate styles and sometimes appended by 5 persistent cuneiform patent stigmas, the base drawn into a stout subterete stipe, subtended by the persistent, erect, patent or reflexed outer perianth and stamens; valves oblanceolate, to 5 cm long, 2 cm wide; seeds 5 or by abortion fewer, filling the locules which occupy the globose, central portion of the fruit. The flowers of this species are reported as white or yellow-green, the fruit is reportedly rose-red or bright red with a fleshy, yellow, brick or orange colored covering and the interior red. Tovomita longifolia is sometimes confused with Tovomitopsis nicaraguensis and other species of that genus, but the appearance of the flower buds distinguish it easily. It also shows less tendency to dry brown, and the fruits are much larger. In addition, the inflorescence of Tovomita longifolia is glabrous but is minutely puberulent in species of Tovomitopsis which resemble it. Tovomita longifolia occurs in lowland Panama, and it ranges through the Guianas to Brasil. CANAL ZONE: Barro Colorado Island, Croat 92/4 (MO), 14016 (MO, SCZ), 14452 (MO); Dare 1115 (MO); Foster 763, 2248 (both DUKE, PMA). CoLon: Santa Rita Ridge, Mori & Kallunki 3052, 6310 (both MO). DARIEN: Upper gold mining camp of Tyler Kittredge, Rio Tuquesa ca. 2 air km from continental divide, Croat 27209 (MO). PANAMA: N of Indio Tower, W of Limon Bay, Gatun Locks and Gatun Lake, Johnston 1794 (MO). Saddle on SE slope between peaks on Cerro Trinidad, Kirk- bride & Duke 1647 (MO). Cerro Jefe, Mori 8004 (MO). SAN BLAS: Permé, Cooper 285 (US) 2. Tovomita morii Maguire, Phytologia 39: 76. 1978. TvPE: Panama, Mori et al. 4685 (MO, holotype; NY, isotype, fide Maguire, not seen). Shrub, perhaps epiphytic; twigs slightly compressed, glabrous, stout; sap yel- low, sticky. Leaves obovate, 20-30 cm long, 10-15 cm wide, apically obtuse, minutely umbonate, basally obtuse and abruptly narrowed into the petiole, the costa subplane and evanescent distally above, elevated beneath, the lateral veins nearly straight, arcuate into the margin, not forming a discernable submarginal vein, mostly 1—2 cm apart with numerous wavy cross veins running tangentially to the leaf base; petioles short, stout, 5-10 mm long, with an axillary flap. [nflo- rescences probably much reduced, the main axis 2-3 mm long, the pedicels great- ly enlarging in fruit. Flowers with the bud depressed-globose, 10—15 mm across (after Maguire), sepals 4, decussate, the outer pair longer and including the inner 1022 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 pair, ovate, oblong, 10-12 mm long, subcoriaceous; petals not seen; stamens 3-seriate, free, linear, sublaterally compressed, 3.5—4 mm long, ca. 0.5 mm wide, the anthers lateral at the filament tips, ca. | mm long; pollen not seen; ovary 5-locular, the ovules solitary, basally fixed. Fruit obovoid to subglobose, 4-5 mm long, conspicuously brown flecked; seeds ca. 1 cm across. This species is distinct in its leaf venation. The lateral veins are widely spaced and arcuate at the margin without forming an evident submarginal vein, and the numerous wavy cross veins are somewhat ascending from the lateral veins. De- tails of the flower are taken from Maguire’s description for there are no flowers with the two collections available at MO. It is not certain whether this species belongs to Tovomita or to Tovomitopsis, but the decussate outer sepals which are slightly parted on the one bud present is suggestive of Tovomita. PANAMA: El Llano-Carti road, Busey 370 (MO); Mori & Kallunki 4685 (MO). 3. Tovomita stylosa Hemsl., Biol. Cent.-Amer. 1: 88. 1879. LECTOTYPE: Panama, Fendler 298 (K, not seen; MO, dupla). T. spc te Cuatr., An. Inst. Biol. Univ. Nac. México 20: 104. 1949. rype: Colombia, Cuatrecasas (F, not seen; US, dupla). Glabrous, monoecious, small tree to 8 m tall, apparently lacking milky sap; stems slender. Leaves ovate or elliptical, mostly 10-15 cm long, 3-5 cm wide, membranaceous, apically short acuminate, basally obtuse or acuminate, the costa prominent, narrow, often impressed above, elevated beneath, lateral veins 8-11, arcuate, not anastomosing or forming a continuous submarginal vein, minor ve- nation slightly elevated on both sides, consisting of 1—2 intermediate lateral veins and one or more strongly ascending, somewhat irregular veins pinnate from the major lateral veins, a reticulum of smaller veins also present; petiole slender with narrow wings forming a ventral channel. /nflorescence terminal, a short panicle or fascicle of short cymose panicles, often subtended by a small lead, the peduncle slender, drying strongly angled, to 4 cm long; bracteoles narrow, cucullate, ca. | mm long; pedicels 5-10 mm long. Flower buds narrowly cylindrical, to 10 mm long, 3 mm wide, the pistillate buds slightly shorter and broader, often falcate; calyx ca. 8 mm long, connate tips splitting lengthwise to form 2 narrowly oblong, often reflexed sepals, corolla greenish, the petals 4, connate in bud, narrowly oblong, the margins thin, slightly inrolled; stamens numerous (ca. 47), ca. 6-9 mm long, free, the anther lobes ca. 0.5 mm long. Fruit pyriform, stipitate, ca. 25 mm long, 5-8 mm wide, surmounted by the conspicuous slender styles 4-6 mm long, drying sulcate, dark brown with lighter, warty spots, the valves red inside; seeds 4, narrowly ovoid, smooth, tipped with an orange or red aril. Tovomita stylosa may be recognized by its narrow flowers and buds and by the pronglike, persistent styles on the fruit. The leaves tend to have more inter- mediate veins than other species of Tovomita. This species is closely related to T. tenuiflora Benth. of Venezuela, which has much larger leaves and larger inflorescences. These two species are somewhat isolated from other members of the genus with their elongate buds and only 2 sepals. The connate petals are also unusual. 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1023 Tovomita stylosa ranges from Panama (?Costa Rica) to Venezuela, Ecuador and Brasil. It grows in moist and wet forests. ANAL ZONE: Woods off S-5 road, Atlantic side, Blum et al. 415 (SCZ). Barro Colorado Island, Croat 11164 (MO, SCZ), 11865 (MO); Croat & Oppenheimer 1521 (MO); Ebinger 1140 (MO); Foster 1485 (MO, PMA, SCZ), 1799 (DUKE, PMA, MO); Knight 1026 (MO, SCZ), 2002 (MO). Boy Scout Road, Madden Dam area, Dwyer 6941 (MO). Between Chilibre and Madden Dam, Dwyer & Correa 8409 (MO). Chagres, Fendler 298 (MO). Coco Solo, Gentry 6067 (MO). Between Pipeline Road and Frijoles, Gentry 7463 (MO). Near Maru Towers, W of Limon Bay, Gatun Locks and Gatun Lake, Johnston 1549 (MO). Indio, Madden Lake, Miller 2053 (US). Near summit of Cerro Pelado, 1 km N of Gamboa, Nee 9061 (MO). Between Gatun and Lion Hill, Pittier 2564 (US). Fort Sherman, Tyson 4 Chu 1698 (MO, SCZ); Tyson & Dwyer 1230 (MO, SCZ). cocLE: La Mesa above EI Valle, Croat 37360 (MO). COLON: Santa Rita Ridge lumber road, Correa 767 (MO, PMA); Correa & Dressler 787 (PMA), 1093 (MO, PMA); Dwyer 8409 (MO); Gentry 6569, 8831 (both MO); Mori & Crosby 6415 (MO); Mori & Kallunki 3047, 5061 (both MO); Nee & Hale 9675 (MO). DARIEN: Río Yape, Bristan 1375 (MO). Río Pirre, E 1477 (MO). Old gold mine at Caná, Croat 27305 (MO). Summit camp between Sasardí and Mortí, 400 m, Duke 10041 (MO). Slopes of Cerro Pirre, 500-1000 m, Gentry & Clewell 7120 (MO). chee Tres Bocas and Cerro Campanimento, Kirkbride & Duke 1352 (MO). PANAMA: l mi después de La Eneida, Cerro Jefe, Correa & Dressler 949 (MO). 10 mi from Intera- merican Highway on road to Cerro Jefe, cab 15190 (SCZ). Cerro Azul, Croat 17294 (MO). Campo Tres, 3 mi NE of Altos de Pacora, 500-800 m, Croat 22724 (MO, US). El Llano-Cartí Road, 10 mi from El Llano, Croat 33713 (MO). Cerro Azul, Dwyer 2188 (MO). Cerro Jefe, Dwyer 3431 (NY). El Llano-Cartí road, Folsom & Collins 1508, 1510 (both MO). Valley between Cerro Azul and Cerro Jefe, Foster 1145 (DUKE, PMA). Cerro Jefe, Foster & Kennedy 1831 (DUKE, PMA), 1841 (PMA). 5-6 mi N of El Llano, 1300 ft, Gentry 5818 (MO); Luteyn & Wilbur 4661 (DUKE, PMA); Mori & Kallunki 2893 (MO). 5-10 km NE of Altos de Pacora, Mori et al. 4206, 4970 (both mA s Llano- Cartí Road 12 km N of Panamerican Highway, Nee /0464 (MO). sAN BLAS: Permé, Cooper 255 (US). VERAGUAS: Valley of Río Dos Bocas between Agricola Alto Piedra and Calovebora, nasi 27590 (MO). 4. Tovomita weddelliana Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 277. 1860. TYPE: Bolivia, Weddell 1851 (P, not seen; MO, photo).—Fic. 10 Clusia oblanceolata Rusby, Desc. S. Amer. Pl. 58. 1920. TYPE: Colombia, Smith 1880 (MO). dicis Mi P MA Engl., Bot. Jahrb. 58. Beibl. 130: 7. 1923. TYPE: Peru, Weberbauer 3446 (B, ;M T. Bees Diels, Notizbl Bot. Gart. Berlin-Dahlem 14: 32. 1938. TYPE: Ecuador, Schultze- Rhonoff 1984, not se T. lanceolata Cuatr., An. Te Biol. Univ. Nac. México 20: 102. 1949. type: Colombia, Cuatrecasas 78 (US). T. ligulata Cuatr., An. Inst. Biol. Univ. Nac. México 20: 99. 1949. TYPE: Colombia, Cuatrecasas 949 (US). T. rhizophoroides Ys An. Inst. Biol. Univ. Nac. México 20: 101. 1949. TvPE: Colombia, Cua- trecasas 14280 (U T. glossophylla Cuatr., dius Acad. Colomb. Ci. Exact. 8: 62. 1950. TYPE: Colombia, Cuatrecasas 9194 (US). idco" rani il Standley & L. O. Williams, Ceiba 1: 244. 1951. TYPE: Costa Rica, Allen 5592 (F, PNE je, EE Steyerm., Fieldiana Bot. 88: 399, fig. 82. 1952. TYPE: Venezuela, Steyermark 0475 (US). T. coriacea a irum, oie m 36: 406. 1977. TvPE: Venezuela, Steyermark 94884 (NY, holotype, not ; VEN, pe). Shrub or tree to 30 m tall, supported by conspicuous stilt roots; twigs often stout with conspicuous leaf scars; sap milky white. Leaves oblanceolate to ob- ovate, to 35 cm long and 8 cm wide, apically obtuse or rounded, sometimes short apiculate, basally narrowed into a broad petiolar area, sometimes ultimately trun- cate or slightly auriculate, the margin callose-thickened, slightly revolute, the costa prominently elevated on both sides, more so beneath, lateral veins obscure 1024 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 FIGURE 10. Tovomita weddelliana Pl. & Tr.—A. Twig with inflorescence emerging from p of in b (x3/s). [After Nee & dorus 10973 (MO).]—B. Fruiting branch (x?/:). [After hm 1832 (MO).] or appearing very numerous, ca. 9 per cm, nearly straight, parallel, appearing unbranched except near the margin, butting into the obscure continuous sub- marginal vein, the submarginal vein 0.3-0.5 mm from the margin, often drying glaucous above, minutely (under a lens) whitish-striate, beneath reddish brown, minutely (under a lens) white-punctate; petiole wanting or represented by a small 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1025 pad against the stem. Inflorescence terminal, to 10 cm tall, cymose, ternately branched, many flowered, in bud enveloped completely by a fused narrowly conical spathaceous structure which splits along 2 sides to yield 2 opposite folia- ceous, deciduous bracts; peduncles often stout, drying angled, reddish to green- ish-brown, the divisions subtended by cucullate, deciduous bracts to 2 mm long. Flowers white, globose in bud, the sepals 4, imbricate in 2 opposite pairs, the inner pair slightly larger, rotund, apically cucullate, 11-14 mm long, coriaceous, persistent and altering somewhat in fruit; petals 4-7, membranaceous, drying yellow, rotund, slightly longer than the sepals; stamens numerous in male plants the filaments filiform, free or some inner series sometimes basally fused, the anthers about as narrow as the filaments with 2, 2-locular, linear lobes, ca. 1 mm long, in female plants the filaments broad except at the apex, the anthers narrow, short, perhaps polleniferous; ovary cylindrical, surmounted by 4-6 flat, rotund, padlike stigmas. Fruit a clavate drupe, often oblique, 20—40 cm long, crowned by the sessile stigmas, ultimately splitting into 4—6 valves; calyx lobes erect, ca. 10 mm long, obtuse and more pointed than in flower; seeds 4, black, cylindrical, sulcate, ca. 12 mm long, 4 mm wide, enveloped in a fleshy aril, the seed and aril produced into a narrow basal spur. This species is easily recognized among the Panamanian Guttiferae by its stilt roots and oblanceolate, sessile leaves. The leaves often appear many-nerved, but close examination reveals there are only about 9 per cm. Collectors report the flowers are white with yellow stamens. Tovomita weddelliana ranges from Venezuela and Bolivia to Costa Rica. Sev- eral variants have been recognized as distinct species, but it is not clear whether they are in any way geographically correlated or representative of distinguishable populations. In Panama this tree is conspicuous, occurring mainly in lowland cloud forests and tropical wet forests of the Caribbean drainage area. COCLÉ: La Mesa above El Valle, Croat 25422 (MO); Mori 6611 (MO). COLON: Santa Rita Ridge, Croat 13896 (MO). 10 mi SW of Portobelo, Liesner 1058 (DUKE, MO). Río Guanche, 3 km upriver from bridge on road to Portobelo, Mori & Kallunki 5216 (MO). DARIÉN: Cerro Pirre, Folsom 4304 (MO). PANAMÁ: Cerro Jefe, Correa & Dressler 721 (MO). Camino a Llano a Cartí, Correa et al. 1855 (DUKE, MO). Cerro Jefe, Foster 1832 (DUKE, MO); Gentry 4877, 6146 (both MO). 19 km above Panamerican Highway on road from El Llano to Cartí-Tupile, 200—500 m, Kennedy 2516 (MO). Cerro Azul, 600 m, Lao & Holdridge 31 (DUKE, PMA, MO). Road from El Llano to Cartí-Tupile, 300—500 m, Liesner 1269 (MO). El Llano-Cartí Road, 17.5 km from Panamerican Highway, Mori et al. 4567 (MO). is Llano-Carti Road, 16-18 km from Panamerican Highway, 400—450 m, Nee & Tyson 10973 (MO). VERAGUAS: Cerro Tute near Santa Fé, 3000 ft, Allen 4352 (MO). Between Escuela Agricola Alto E Piedra and Río Dos Bocas, 730—770 m, Croat 25781, 25961, 27492, 27664 (all MO). NW of Santa Fé, 10.8 km from Escuela Agricola Alto de Piedra on road to Calovebora, Mori 6733 (MO). 12. TOVOMITOPSIS Tovomitopsis Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 261. 1860. TYPE: T. paniculata (Camb.) Pl. & Tr ad Spreng., Neue Entdeck. 2: 110. 1820, non Spin (1809) nom. rejic., nec Raddi (1820) nom. Chrysochlamys Poepp. & Endl., Nov. Gen. & Sp. 3: 13, t. 211. TYPE: C. multiflora Poepp. a elt v. Gen. & Sp. 3: 13, t. 211. m Tauomitonsis multiflora (Poepp. & Endl.) D'Arcy, comb. n Dioecious or perfect trees or shrubs, stems terete or angled, trunks sometimes 1026 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 buttressed, sap often milky white or yellow. Leaves at first opposite, later often alternate, simple, entire, the margin usually narrowly differentiated and minutely revolute; costa prominent, mostly elevated beneath, lateral veins several-many, straight or arcuate, sometimes forming a partial submarginal vein, mostly with intermediate veins and secondary lateral veins, glabrate, the costa and major veins sometimes minutely puberulent, secretory canals often forming linear streaks on the leaf undersides; petioles mostly slightly canaliculate; stipules want- ing; axillary buds not prominent. Inflorescences mostly terminal, sometimes be- coming axillary or lateral on the stem, occasionally terminal on short shoots, erect or lax cymose panicles, rarely reduced to 1-3 flowers, the axis mostly unbranched, the nodes branching dichotomously, ternately or in verticels, sub- tended by foliaceous or reduced scalelike bracts, the pedicels bracteolate, the bracteoles mostly paired, small. Flowers monoecious or perfect, the buds mostly globose, the outer perianth imbricate, shorter than the bud or overtopping it, sepals mostly 4—6, rotund, the innermost overtopping the bud, often coriaceous, streaked with canals; petals 4—6, imbricate, mostly overtopping the bud, rotund, often fleshy, streaked with canals; stamens numerous, 25-200, some reduced, fewer or reduced in pistillate flowers, the filaments free or the innermost fused and their anthers reduced or abortive, sometimes crumpled in bud, the anthers 2-lobed, 4-locular, minute, bordering the apex of the filament (connective); ovary ovate, glabrous, surmounted by mostly sessile, discoid styles, reduced or obsolete in staminate flowers. Fruit a leathery or fleshy drupaceous capsule splitting from the apex into (4—)5(—6) valves; seeds 1-5, oblong or faboid, basally fixed, envel- oped at least in part by a loose, fleshy aril or arilloid. Tovomitopsis 1s a genus of several dozen species of tropical America. It may be confused with Tovomita or Dystovomita, but it differs from the first of these in having the perianth completely free and mostly imbricate, and from the second in having the ovules basifixed and ascending instead of pendulous. For routine recognition, Dystovomita differs from Tovomitopsis in having the inflorescence cauliflorous on mature wood and in sometimes having part of the perianth basally connate. Chrysochlamys has traditionally been separated from Tovomitopsis on the number of perianth parts and on the androecium of a few species. In Chryso- chlamys the perianth consists of 5 sepals and 5 petals, but this is also the case in a number of species of Tovomitopsis. It is not always clear which members are sepals and which are petals. The androecium of Tovomitopsis paniculata, the type of Chrysochlamys, and of the closely-related C. pavonii, C. laxa, and C. dependens, have the stamens united into a column while other species which have been traditionally placed in Chrysochlamys have androecia as in Tovomi- topsis. Vasconcellos et al. (1972) reported that pollen in C. dependens is porate, but they did not report on any species then considered to belong to Tovomitopsis. The pollen of Tovomita brasiliensis they found to be colporate. Species of Tovomitopsis occurring in Panama are often difficult to separate, and obvious differences are not always easy to relate in words or in a key. At the same time, in a number of cases, although the number of collections is large, those with good flowers, buds or mature fruits are few, and it has not been possible to make satisfactory comparison with species of South America. 1980] e ° D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1027 Leaves basally truncate, subsessile; flower buds becoming 5-6 mm long and conspicuously striate . T. croatii Leaves basally narrowed, obtuse or acute, petiolate; flower buds not evidently striate or, if so, then less than 4 mm long. Leaves narrow, la dam or elliptical, more than 4 times longer than broad. Flower buds more than Leaves lanceolate, mostly less than 1.5 cm wide; flower buds reddish . angustifolia dd. n broader, mostly more than 1.5 cm wide; flower buds drying brown, n or white ' occasionally pinkish or streaked. ve s, drying noticeably revolute (more than 2 mm of leaf ap te curve Leaves drying with conspicuous, numerous canals beneath; outer sepals as long as the bud; inflorescence glabrous _____. . membrillensis ff. Leaves drying without conspicuous canals; outer sepals only 1⁄4 as long as the bud; inflorescence minutely puberulent __ centistaminibus ee. Leaves membranous or chartaceous, hardly revolute (lens than 0 mm of leaf margin curve Leaves with 16—20 MEE on each side, prominent benea 7. T. m in ea gg. Leaves with fewer than 12 veins on each side, n st obscure 6. T. glauca cc. Flower buds less than 5 mm h Leaves with 5-11 veins on VT side, sometimes glaucous ES Pp evidently streaked T. glauca hh. is aves with ca. 16 veins on each side, not glaucous; buds not E l eak 0. T. micrantha ed IU Tus mostly less than 4 times longer than wide. Flowers solitary (?or in reduced inflorescences); pedicels (in young fruit) si mm long, ds leaves mostly less than 6 cm long, the veins me - 7p c 2 erra fala ii. Flowers in many-flowered inflorescences; pedicels mostly more M 4 mm long, often stout; leaves mostly more than 6 cm long, the veins more widely spaced. Corolla with streaks visible with ibo naked eye; stamens fewer than 40. k Outer sepals almost as long as the flower bud; inflorescence key puberulent T. faucis kk. Duer n less than half as long as the flower bud; inflorescence glabrou 6. T. glauca J. NEA ab not visible with the naked eye; stamens various in number. l. t m. Leaves with fewer than 11 veins on each side. n Upland species (over 1,500 m); flower buds 3-4 mm long; inflorescence branches drying wrinkled and appearing pulverulen , M nn. Lowlands species (below 1,000 m); flower buds ^e long; inflorescence branches drying smooth... ODE . T. myrcioides mm. Leaves with fewer than 11 veins on each side. Leaves coriaceous, less than 4.5 cm wide; fruiting ped- icels slender, 9 mm long |... . centistaminibus oo. Leaves chartaceous, more than 4.5 cm wide: fruitin pedicels stout, 5-7 mm long ______________ 13. T. standleyana Il. Stamens fewer than 50. p. Inflorescence minutely puberulent. Outer sepals almost as long as the flower bud _ 5. T. faucis dd Outer sepals less than !^ as long as the flower r Leaves membra ous Or C eous, most drying green (grayish) beneath; sepals drying whit- ish or pinkish; inflorescence glabrate . T. glauca rr. Leaves coriaceous, mostly drying brown lien: 028 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 sepals drying brown; inflorescence minutely pu- erulen 12. T. nicaraguensis pp. Mie pawaq Landa noU ovate or elliptical with fewer em 12 veins on pes side, coriaceous, more than 9 cm lon I membrilensis Leaves elliptical or ovate with more than 12 vein each side, chartaceous, less than 6 cm long -------------- 7. T. membranacea w w 1. Tovomitopsis allenii Maguire, Phytologia 38: 214. 1978. TYPE: Panama, Allen 5014 (MO, holotype). Tree to 25 m; twigs stout, drying irregularly sulcate often with a granular, yellowish appearance, sometimes compressed; sap reddish. Leaves obovate or elliptical, sometimes narrow, to 20 cm long, 7 cm wide, apically acuminate, ba- sally obtuse or cuneate, the costa glabrous, subplane above, conspicuously rounded, raised beneath, lateral veins 12-15 on each side, evenly spaced and arcuate, loop connected distally and forming a submarginal vein, evanescent and not connected basally, slightly elevated on both sides, inconspicuous above, the minor venation inconspicuous, of intermediate lateral veins running halfway to the margin and numerous secondary lateral veins which interconnect in the mar- ginal 14 of the leaf; petioles stout, 5-20 mm long, canaliculate above, minutely puberulent. Inflorescences terminal or becoming axillary, globose or pyramidal, to 15 cm tall, 12 cm wide, rachis and pedicels minutely puberulent, stout, drying yellowish with a granular appearance, the cymose panicle with the main rachis having ca. 3 nodes, each subtended by foliaceous bracts; pedicels 3-8 mm long, becoming stouter but not much longer in fruit; bracteoles patent, 1-2 mm long, slightly larger in fruit. Flower buds globose, ca. 5 mm long, sepals 4, rotund, coriaceous, the outer pair short, less than !4 as long as the bud, the inner sepals imbricate, one as long as the bud, the other (inner) sepal overtopping it; petals 5 (Maguire), almost twice as long as the sepals, ca. 8 mm long; staminodes nu- merous (79), the filaments slender, varying in length, to 8 mm long, the anthers minute, parallel or convergent, minutely apiculate, the innermost filaments ba- sally connate, polleniferous; ovary truncate, the 5 carpels apically forming a rim around a central depression. Fruits (immature) obpyriform, ca. 14 mm long, the styles conspicuous, short-stipitate, dark; seed ca. 8 mm long, 6 mm wide, ellip- soidal, affixed just above the base, axile, erect, completely but loosely enveloped by an aril, a slightly raised disc present at the chalaza. This species is closely related to Tovomitopsis myrcioides (Pl. & Tr.) D'Arcy, but it differs in the texture of the dried surface of the inflorescence branches and in having generally smaller leaves with more closely spaced veins. It also differs in range, occurring at elevations about 1,000 m above that of T. myrcioides in Panama. It occurs also in Nicaragua (Williams et al. 42981 [US]) and Costa Rica. The above description differs from that by Maguire in noting larger inflores- cences. The unequal, scarious margined outer sepals referred to by Maguire were not confirmed. Maguire described the pollen as spheroidal, tricolporate, 25 wm in diameter and foveolate. BOCAS DEL TORO: Robalo Trail, N slopes of Cerro Horqueta, 6000-7000 ft, Allen 5014 (MO, NY). 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1029 CHIRIQUÍ: Upper Río Chiriqui Viejo 2 km NE of Guadeloupe, 2 km N of Cerro Punta, Wilbur et al. 15331 (DUKE). 2. Tovomitopsis angustifolia Maguire, Phytologia 36: 399. 1977. TYPE: Panama, Mori & Kallunki 5357 (MO) Much-branched shrub to 2 m tall, forming salicoid clumps; sap milky. Leaves narrowly elliptic, to 10 cm long, 2 cm wide, apically acute or acuminate, callose tipped, basally acute, narrowed into the petiole, the margin minutely revolute, bordered by a narrow indurate line, the costa prominent, prominulous above, conspicuously elevated and drying reddish brown beneath, the lateral veins ob- scure, 5-6 on each side, strongly ascending, often with one minor intermediate vein, a few short minor veins pinnate on the lateral veins, not forming a distinct submarginal vein; petiole short, ca. 8 mm long, to 2 mm wide, slightly channeled above, striate and sometimes minutely sharp-angled beneath. Inflorescences short, sparingly branched, cymose panicles to ca. 6 cm long, basal bracts folia- ceous, other bracts narrow, cucullate, 1-2 mm long; pedicels to 10 mm long, longer and stouter in fruit, bibracteolate near the base. Flower buds drying pink- ish or reddish, ellipsoidal or globose, ca. 9 mm long, 8 mm wide; sepals coria- ceous, outer sepals opposite, free, one overlapping in bud but by anthesis shorter than the bud and sometimes spreading, the margins thinned, inner sepals imbri- cate, the margins narrowly thinned, by anthesis slightly shorter than the bud, inconspicuously striate; petals 7, imbricate, overtopping the bud, thinner than the sepals, inconspicuously nerved; stamens numerous (30-40), free, the fila- ments broader than the anthers, narrowing upwards, slightly differentiated below the anthers, anthers slightly converging upwards, the connectives minutely apic- ulate, unequal, the inner longer. Fruit pyriform, 1.5—2.0 cm long, 10-12 mm wide, the body rounded-sulcate, sometimes slightly flattopped, drying pink or red, the styles forming a short apiculus, the stigmas sometimes persistent, basally nar- rowed into an obconical stipe; seed oblong, basifixed, with a discolorous, almost apical chalazal disc, the hilum conspicuous, a loose dorsally venose aril fixed at the base and loosely enveloping the seed. This is one of the most distinctive species in the Panamanian Guttiferae flora having a salicoid growth form, being shrubby with narrow leaves and occurring along streams. In this respect it parallels Solanum intermedium, common in sim- ilar habitats in lowland Darien Province. Tovomitopsis angustifolia is known only from the area around Santa Fé in Veraguas Province. This species displays characters of both Tovomita and Tovomitopsis: the venose aril of the former and the incomplete, imbricate sepal prefloration of the latter. Maguire’s placement of the species in Tovomitopsis in accepted here: the outer perianth parts seem to conform more to Tovomitopsis than to Tovomita. VERAGUAS: Rio Pimero Braso, 2.5 km beyond agricultural school Alto Piedra near Santa Fé, 700—750 m, Croat 25437 (MO). Between Alto Piedra school a nd Rio Calovebora 1.8 mi beyond school, Mori & Kallunki 3179 (MO). Road to Calovebora, 2.7 km from Escuela Agricola Alto de Piedra, Mori & Kallunki 5357 (MO). Road to Calovebora, 16 km NW of Santa Fé, 300—500 m, Mori & Kallunki 6125 (MO 1030 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 3. Tovomitopsis centistaminibus D’Arcy.'! TYPE: Panama, Little 6070 (MO, ho- lotype; US, isotype) ?Dioecious tree to 20 m tall, new growth drying dark reddish brown, minutely puberulent, older twigs gray. Leaves coriaceous, ovate, obovate or elliptical, to 15 cm long, 5 cm wide, apically acuminate or acute, basally acute, the margins minutely revolute, the narrow, indurate border drying reddish, the costa nearly plane above, elevated and drying reddish brown beneath, the lateral veins 6-10, obscure above, evident beneath, parallel, sometimes ill spaced, ascending and arcuate at the margin, not forming a submarginal vein, the minor venation obscure above and beneath, not well developed, consisting of a few intermediate lateral veins and pinnae on the lateral veins, drying discolorous, above slate gray, be- neath yellowish green, canals not evident; petiole slender, 15-20 mm long. /nflo- rescence terminal, soon on a lateral leafy short shoot, ca. 10 cm long, rounded or pyramidal, the primary axis somewhat stout, the axes minutely puberulent; bracts cucullate, ca. 3 mm long, spreading, ?minutely puberulent, drying reddish; bracteoles minute, rounded or blunt, puberulent; pedicels short, 1-3 mm long. Flower buds globose, 3—4 mm long, the basal pair of sepals often wide spreading, inner sepals 3, imbricate, overtopping the bud, drying unlined or with light-col- ored veins, petals 3-4, imbricate, overtopping the bud, thick, sometimes with few fine lines; in male flowers the stamens numerous (90), free, clustered, the longer filaments crumpled in bud, anthers variable in shape and size, some per- haps abortive, the locules lateral at the connective apex, the connective some- times slightly extended; ovary rudimentary consisting of 5 short, digitate, imper- fect carpels. Fruit ca. 20 mm long, globose to pyriform, seed with an orange aril. This species has distinctive subcoriaceous small, elliptical leaves which dry with the venation beneath obscure or somewhat reddish. It is similar to 7. psy- chotraefolia Oerst. of Costa Rica, which differs in having conspicuously maculate sepals and petals and larger leaves. Tovomitopsis centistaminibus is known from Panama, Colombia (Cauca: Entre Aguabonita y Candelaria, 2,280-2,350 m, Cua- trecasas 23563 (US). Meta: 6 km N of Hacienda El Pato, 7,600 ft, Little 8021 (US). Valle: Piedra de Moler, 900—1,180 m, Cuatrecasas 15052 (US). Without Other Locality: Mutis 1069 (US)) and Ecuador (Santiago-Zamora: Between Pailas and El Pan, 2,255-3,445 m, Steyermark 54307 (US)). Panamanian collections are all from upland Chiriqui. CHIRIQUÍ: 8 km N of Camp El Volcán, 6000 ft, Little 6046, 6068 (both MO), 6070 (MO, US). N of San Félix on Cerro Colorado copper mine road along continental divide, 5000-5500 ft, Mori & RE 5970 (MO). ! Tovomitopsis c sos elders D' Arcy, spec. nov. Arbor ad 20 m alta. Ses rm coriacea ribus discolora, venis subtus rubescentibus in sicco, supra obscuris, venis mino , ve at- eralis utraque 6-10, vena DE rginale carenti. /nflorescentia bracteolis minute anne bh ded: icellis 1-3 mm longis. Alabastra globosa 3—4 mm longa, sepalis basalis bu aliquantum patentibus, staminibus numerosis, circa 95, libris, rane longioribus corrugatis. Fructus 20 mm longus, glo- bosus ad pyriformis, semine arillo aurantiaco te 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1031 4. Tovomitopsis croatii (Maguire) D’Arcy, comb. nov. Tovomita croatii Maguire, Phytologia 36: 404. 1977. TYPE: Panama, Croat 14268A (MO). Shrub to 2 m tall, branches terete. Leaves elliptic or obovate, to 25 cm long, 10 cm wide, apically acuminate, basally narrowed, rounded, ultimately truncate or auriculate, the margin with an indurate border 0.25 mm wide, chartaceous; costa and lateral nerves conspicuous, subplane above, elevated and drying brown beneath, the lateral veins numerous, ca. 18 on each side, slightly curved and slightly ascending, parallel and evenly spaced, anastomosing at the margin to form a looping submarginal vein, minor venation obscure consisting of weak intermediate nerves and of cross veins between the lateral nerves, drying above and beneath with a dense sometimes conspicuous array of wavy, linear canals, these somewhat contrasting beneath; petioles to 5 mm long or wanting, often forming a nodal mark or ridge. /nflorescence an open cymose panicle, to 12 cm long, drying reddish brown; bracts conspicuous, smaller and narrower upwards, keeled and clasping the inflorescence branch, apically narrowed; pedicels 5-8 mm long, sometimes minutely puberulent, stouter above the bracteoles; bracteoles ca. 4 mm long. Flower buds globose or ellipsoid, to 7 mm long, mostly conspic- uously striate; outer sepals overlapping the bud, imbricate, the sepals 5; petals 5, all overtopping the bud, conspicuously maculate, especially basally, with a small stipe and large claw; in staminate flowers the stamens numerous (30—40), the anthers minute, convergent at 45°, the filaments free, slightly bent or crumpled in bud; ovary obsolete. Fruit not seen. This species is distinctive in its sessile, basally clasping or auriculate leaves. The leaves and inflorescence dry reddish brown, and the large buds are maculate. Maguire placed this species in Tovomita, which may be correct, but the outer perianth is composed of completely free parts which are imbricate at the top of the bud. Because pistillate flowers or fruit have not been found, it is not possible to examine details of the ovule. Tovomitopsis croatii is known from separated points in western Panama. It is not known from outside of Panama. AS DEL TORO: Between Quebrada Gutierrez and E slope of La Zorra, headwaters of Río Mal on Chiriqui Trail, Kirkbride & Duke 732 (MO). cocrÉ: El Valle, Croat 14268A (MO). Cerro Pilón, is yer et al. 4556 (MO). Cerro Caracoral, Kirkbride 1122 (MO). NW of Santa Fé 0.8 km from Escuela ricola Alto Piedra, on road to Calovebora, Mori 6725 (MO). 5. Tovomitopsis faucis D’Arcy.!? TYPE: Panama, Pittier 3039 (US). ?Tree, twigs at first strongly angled and sulcate, soon terete, the new growth drying olive green, slightly older growth drying yellowish green, hollow, the nodes 1.5-3.0 cm apart. Leaves elliptical or obovate, 10-20 cm long, 4-7 cm wide, m Miri d waw faucis D'Arcy, spec. nov. Species venis ad initium minute puberulentibus mox glabris ca. 11 utraque arcuatis ascendentibus, venam submarginale carens, alabastro roseo 5 m longo je i staminibus numerosis circa 28 libri 1032 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 apically obtuse, short acuminate, basally obtuse, acute or acuminate, the margin narrowly stramineous, drying minutely revolute and thus concealed except at the apex, the costa elevated beneath, the lateral veins ca. 11 on each side, slightly arcuate and ascending, abruptly ascending at the margin, the apical veins anas- tomosing near the margin, not forming a submarginal vein, one or more lateral veins divided well in from the margin, the minor venation obscure above, of ill- spaced, unequal intermediate veins (0—)5—6 weak, irregular pinnate veins arising from the laterals and sometimes anastomosing in the intervenal region, ptyxis at least in part involute, the costa and bases of the lateral veins minutely puberulent beneath on emerging, soon glabrous, surface canals not evident; petioles 15-20 mm long, slightly expanded at the base, minutely puberulent dorsally. /nflores- cences terminal on leafy shoots, a cymose raceme or panicle ca. 8 cm long, the axis drying green, the branches subtended by minute scale bracteoles; pedicels ca. 5 mm long. Flowers with the buds ovoid, ca. 6 mm long; sepals petaloid, hardly striate, overtopping the bud, imbricate, membranous; petals 4-5, overtop- ping the bud; stamens ca. 24, some filaments slightly connate basally, ca. 2 mm long; ovary wanting in staminate flowers, pistillate flowers not seen. Fruit not seen. This species is distinct in its venation pattern and in the size of its flower buds. It appears to be related to Tovomitopsis glauca, but the striation of the pink- ish petals is only slightly evident. This species is known only from the type, which is staminate. It is named for the type locality, El Boquete, meaning a defile or gorge. HiRIQUÍ: Flower white, along river, forests around El Boquete, 1000-1300 m, 5 March 1911, Pittier 3039 (US). 6. Tovomitopsis glauca Oerst. ex Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 264. 1860. TYPE: Costa Rica, Mt. Jaris, Oerstedt 3 (=Oerstedt 3589), (?P, not seen; C, not seen; MO, photo).—Fic. 11A-B. ?Tovomita martiana Engler in Mart., Fl. Bras. 12(1): 453. 1888. rype: Brasil, Martius (M, not seen; oto). T. multiflora Standley, Field Mus. Nat. Hist. Bot. Ser. 4: 234. 1929. TYPE: Panama, Cooper & Slater US). T. po (Oerst. ex Pl. & Tr.) L. O. Williams, Fieldiana Bot. 29: 357. 1961. Shrub or tree to 8 m tall, twigs at first deeply sulcate, soon terete, drying greenish or grayish, glabrous, internodes variable, 5-25 mm apart. Leaves ovate or elliptical, sometimes narrow, sometimes oblique, mostly 5-10 cm long, (1—)3- 5 cm wide, apically acuminate, the tip often curved, basally obtuse or acute, glabrous, minutely revolute; costa plane above, sharply elevated beneath, the lateral veins 5-11, obscure, arcuate and ascending, evenly spaced, recurving and anastomosing in broadly crenate fashion near the margin, the looping outer con- nected vein giving rise to minor, peripheral loops, a submarginal vein not formed, minor venation mostly obscure, mostly of about 2, ill-spaced intermediate veins which arise at irregular angles, and short, ill-spaced veins pinnate on the lateral veins, the lower surface with inconspicuous surface canals evenly dispersed in the intervenal areas, ascending slightly more than the lateral veins, often gray- 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1033 a — | KRAGER -PALMER _ Tovomitopsis.—A.-B. T. glauca Oerst.—A. Flowering twig (x 1⁄2). [After Kirkbride & n. "560 | MO).]—B. Flower (x21⁄). [After Smith 162 (MO), Costa Rica.]—C. T. nicaraguensis Pl. & Tr. Fruit (x214). [After Croat 8647 (MO) and photo of same.] — green or glaucous beneath; petioles 6-25 mm long, narrow, canaliculate above, the narrow margins forming sharp angles. Inflorescences terminal, sometimes becoming axillary, sometimes on leafy short shoots, subtended by normal leaves, a pyramidal hemispherical or cylindrical cymose panicle, the peduncle 5-12 cm long, 3-6-noded, the primary axis stout or slender, compressed above, the branches mostly slender, pedicels 4-6 mm long, becoming 10—12 mm long and stouter in fruit, often bibracteolate in the lower half, sometimes minutely puber- ulent. Staminate flowers with the buds often pinkish, 3-5 mm long, conspicuously striate, ovoid or ellipsoidal; outer sepals opposite, ca. 2 mm long, much shorter than half the length of the bud, striate, sometimes pink at the base, the inner 3(— 4) sepals imbricate, overtopping the bud, membranous, striate, ca. 5 mm long; petals 4-5, about as long as the inner sepals, thicker, striate, overtopping the bud, imbricate. In staminate flowers the stamens numerous (28-33), the filaments 1034 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VOL. 67 ca. 0.5 mm wide, free or the innermost slightly connate basally, the inner stamens slightly longer and broader with flat filaments, some rudimentary anthers perhaps not polleniferous; ovary obtuse, longitudinally sulcate, the styles relatively large, 5, subsessile, flattopped, heel shaped, contrasting in color, ?yellowish or orange; pistillate flowers with buds slightly larger, the outer sepals relatively slightly longer; stamens as in male flowers but the thecae reduced; ovary about as long as the stamens, the stigmas 5, subsessile, spreading. Fruit pyriform, 20-25 mm long, sulcate, slightly depressed apically, often strongly asymmetrical or curved by abortion of 1 or more locules, the styles conspicuous or not, subsessile, drying reddish brown; seeds oblong, basifixed, erect, the testa slightly venose, com- pletely enveloped by a basifixed, unveined aril. Tovomitopsis glauca is extremely variable in leaf and flower bud size but is recognizable by its leaves with few inconspicuous veins and striately maculate flower buds. There is also great variation in inflorescence, this ranging from a short, almost congested cymose panicle to a 30 cm long, open, pendulous panicle with slender, elongate, weak inflorescence branches. Collections of this species from two localities in central Panama, Río Guanche on the Caribbean coast and Cerro Campana near the Atlantic, have smaller stature and smaller narrower leaves than usual and are suggestive of a distinct variety or even species. But in each case other specimens show this variation to be continuous with wider- leaved, more normal morphology. This narrow-leaved form is similar to T. an- gustifolia, which occurs in the same area. The flower buds are very different in the two species. S DEL TORO: Duwebdulup Peak N of Río Teribe across from Quebrada Huron, 90- lis Kirkbride & Duke 560 (MO). VAS naqa Calderas trail between Quebrada Higueron and Guttierrez, Kirkbride & Duke 765 (MO). c E: Slope N of Indio Tower, NW part of Canal Zone, y ankaqa 1794 (MO). Drowned a of. Río omnes and Peluca Hydrographic Station, dide & Allen 17269 (US). CHIRIQUÍ: Progreso, Cooper & Slater 171, 274, 304 (all US). 10-11 mi W of Puerto ine ae 300—500 m, Croqt rides (MO). Paseo Canoas to Cañas Gordas, bed 22211, 22348 (both O). 8 mi W of Puerto Armuelles, 200 m, Croat 22472 (MO). 19 km W of Río Chiriqui Viejo on road * San Serano, 1000-2000 m, D' Arc y 10835 (MO). Near Pinola, Kirkbride & Duke 865 (MO). 1 mi E of Canas Gordas near Costa Rican Border, Liesner 268 (MO). Near Las Lagunas W of El Hato del Volcán, 1400 m, Wilbur et al. 11001 (DUKE). cocrÉé: Between Cerro Pilón and El Valle de Antón, 00-900 m, Duke & Dwyer 13936 (MO, NY, SCZ). Base of Cerro Pilon above El Valle, Gentry & ie yer 3654 (MO). La Mesa, El Valle, 1000 m, Gentry 6848 (MO). N of El Valle de Antón, 2500- 3000 ft, Lewis et al. 1769, 1785 (both MO). Cerro Pilon, 800-1045 m n 6565 (MO). 3 km NE of EI Valle, Mori & Kallunki 2960 (MO). COLON: Santa Rita Ridge 8564 (MO). Rio Guanche, Foster & Vocelka 2201 (DUKE); Foster et al. 2330 (DUKE); di 8764 (MO); Kennedy 1174 (DUKE, MO); Kennedy & Foster 2132 (MO); Mori et al. 6443 (MO); Mori & Kallunki 5194, 5212 (both MO); Mori & Witherspoon 7955 (MO); Nee & Gentry 8719 (MO). DARIÉN: Río Yape, qaq 1378 (MO, US). Between Paya and Palo de las s. Duke & M dade 14024 (MO, , SCH) Cer Campanento S of Cerro Pirre, Duke 15646 (US). PANAMÁ: Cerr dad, —1000 Allen 377 73 (MO). El Llano-Carti road, Busey 3777 (MO). pe Campana, a pon gos (boh MO). El Llano-Cartí road, i & Dressler 9371 (MO). Cerro Campana, Porter et al. 4259 (MO, DUKE). VERAGUAS: Between Escuela Agricola Alto Piedra and Calovébora, Croat 27665 (MO); Mori & Kallunki 3224, 3950 (both MO). 7. Tovomitopsis membranacea (Pl. & Tr.) D'Arcy, comb. nov. Chrysoc Miu d ea 5 bs Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 260. 1860. rype: Colombia, Triana (P, not ; MO, to). Tree to 6 m tall. Leaves membranous or chartaceous, obovate, to 25 cm long, ` 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1035 to 7 cm wide, apically acuminate, basally obtuse or acute, sometimes falcate, the costa narrowly and sharply elevated beneath, subplane above, the lateral veins numerous, 17—20 on each side, nearly straight, ascending, branching near the margin and loop connected but not forming a submarginal vein, slightly further spaced near the apex, chartaceous or membranous, drying beneath with con- spicuous, numerous canals parallel to the lateral veins, the costa and lateral veins minutely puberulent near the base; petiole almost wanting or to 25 mm long, minutely puberulent beneath. Inflorescence an open, ?pendulous panicle, the axes elongate, lax, minutely puberulent, the pedicels ca. 6 mm long, becoming 15-20 mm long in fruit. Flowers (after Vesque, 1893) ca. 15 mm across, calycine bracts (outer sepals) lacking, sepals 5, quincuncial, orbicular, the outermost smaller; petals 5, apparently alternating with the sepals, obovate, contracted into a broad limb; stamens to 20, indistinctly 2-seriate, free, the filaments linear, compressed, the anthers small, the connective bordered by the two basally di- verging, laterally and introrsely dehiscent anther locules, the staminodes forming a solid column in the center; pistillate flowers (after Planchon & Triana) with the stamens few or none, the staminodial annulus shielding the ovary; ovary short, stout, surmounted by 5 thick stigmas. Fruit obovoid to pyriform, 25-30 mm long, pale green tinged with red (Croat 34216), the styles forming a small apiculum, the base with 3—4 persistent, coriaceous, minute sepals. This species is distinct in its papery leaves with numerous straight, ascending veins. The pendulous clusters of fruits are also distinctive in appearance. The material at hand is doubtfully identified with the Colombian material on which the name 7. membranacea is named. The leaf venation and shape is quite similar and the original authors refer to the minute puberulence of the nerves. In Panama the species is known only by one collection and accompanying photograph taken in a region of lowland cloud forest in central Panama. UAS: Above Santa Fé beyond Escuela Agricola Interamericana 1.8 mi beyond road fork on Pacific ° slope and on side of Cerro Tute, Croat 34216 (MO). 8. Tovomitopsis membranifolia (Standley) D'Arcy, comb. nov. Tovomita membranifolia Standley, Field Mus. Nat. ei Bot. Ser. 22: 160. 1940. TvPE: Panama, Terry & Terry 1478 (F, holotype, not seen; MO, isotype). Glabrous shrub or tree to 4 m tall, branches slender, solid. Leaves elliptical, to 10 cm long, 3.5 cm wide, apically acuminate, basally obtuse; venation more or less obscure above, the costa slightly elevated proximally beneath, the lateral veins numerous, 16—26 on each side, mostly straight and ascending, branching near the margin and forming a distinct but discontinuous submarginal vein 0.5— 1.5 mm from the margin, minor venation of intermediate veins which sometimes rank with the lateral veins, and of a fine reticulum, membranaceous or charta- ceous, sometimes drying pinkish or reddish; petioles short, 8-10 mm long. Inflo- rescence (from fruiting material) of a solitary flower terminal between a pair of normal leaves on a short axis; pedicel slender, 3 mm long or becoming longer. Flowers unknown. Fruit ovoid or elliptical, ca. 20 mm long, subtended by 5 coriaceous rotund, slightly cucullate sepals 2 mm long, the valves drying maroon within. 1036 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 This species is known only by the type collection and one other cited below, both in fruit. The original description made no mention of flowers or fruiting parts. The type resembles many specimens of T. glauca, but the venation pattern is different and the inflorescence reduced to solitary flowers. These two collec- tions may represent depauperate individuals of some other species. The Pana- manian material described here is quite similar to the type of Tovomita krukovii A. C. Smith!3 of Amazonian Brasil. This species also has reduced inflorescences consisting of only 1-3 small flowers on a short axis. Tovomita krukovii is de- scribed as having 4 sepals instead of the 5 of Tovomitopsis membranifolia. DARIEN: Cuasi-Cana trail between Cerro Campamiento and La Escalera to ‘‘Paramo”’ E of Tres Bocas, Kirkbride & Duke 1312 (MO). Cana-Cuasi Trail, Chepigana District, 4000 ft, Terry & Terry 1478 (MO). 9. Tovomitopsis membrillensis D’Arcy.'* rype: Panama, Duke 10872 (MO- 2124393, holotype). Small tree to 7 m tall, mature twigs terete with numerous longitudinal furrows, young twigs somewhat compressed, finely many-angled, drying lighter. Leaves lanceolate or narrowly elliptic, occasionally widest above the middle, mostly 15- 20 cm long, 4.5-6.0 cm wide, apically acute or slightly acuminate, minutely cal- lose, basally acute or obtuse, the margins minutely revolute, drying discolorous, slate green above, yellowish or reddish green and lighter beneath, the costa slight- ly raised and rounded above, elevated and almost excurrent beneath with sharp angles, the lateral veins ca. 9 on each side, obscure above, conspicuous beneath, arcuate, ascending at the margin, not forming a submarginal vein, sometimes with some weak anastomosing connections in the upper portion, 2—3 inconspicuous intermediate veins between pairs of lateral veins and 1-3 similar veins pinnate from the lateral veins, numerous conspicuous ducts evident rising at a slightly sharper angle than the veins; petiole 10-30 mm long, slightly and narrowly can- aliculate above, striate beneath, drying dark or reddish brown. Inflorescences terminal cymose panicles, appearing axillary in fruit, glabrous, the main axis slender, ca. 7 cm long, bearing ca. 5 pairs of branches each 3-5 cm long, these in turn bearing 1—3 rachises or pedicels, the pedicels 6-15 mm long, slender, stiff, enlarging upwards, drying ridged, in fruit appearing continuous with the penul- timate element of the inflorescence, these two parts together to 20 mm long, the pedicel slightly longer; bracts cucullate, the inflorescence subtended by normal leaves, the bracts 2—3(—6) mm long, the bractlets subtending the pedicels ca. 1 mm long, persistent. Flower buds ca. 5 mm long, ellipsoidal, the outer sepals mostly equal, as long as the bud or only !^ as long, overlapping, spreading wide, conspicuously ribbed; sepals 5, petals 5, opposite, imbricate and apically hooding the interior of the bud, conspicuously nervate, the innermost sometimes rudi- 13 Tovomita krukovii A. C. Smith, Phytologia 1: 122. 1935. rype: Brasil, Terra Firma, basin of Rio Jurua, Krukoff 4887 (MO, US). 7 a Qin membrillensis D'Arcy, T nov. Arbor ad 7 m alta foliis angustis coreaceis minute revolutis canalibus secretoriis numeros gin ee Wa ie paniculata, alabastro bruneo siccante, eT exterioribus imbricatis. pos tus roseu 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1037 mentary; stamens numerous (37), free, the filaments relatively stout, slightly nar- rower upwards, the anthers 0.3 mm long, the filaments crumpled in bud, all somewhat bent to one side; ovary rudimentary. This species is distinct in its narrow, coriaceous leaves with conspicuous canals in the wide areas between lateral veins. The canals are not all parallel but change direction along their length. The buds dry dark brown but the streaks are still visible with the naked eye. Duke 10901 records the fruits are pink. Tovomitopsis membrillensis is similar to some Colombian material labeled as Chrysochlamys ulei Engler, but these specimens have broader leaves and pu- berulent inflorescences. : Upper Rio Membrillo near construction road to San Blas, 100-800 m, Duke 10863, 10872, ^ 0901 (all MO). 10. Tovomitopsis micrantha (Engler) D'Arcy, comb. nov. iii edid micrantha Engler, Bot. Jahrb. 58, Beibl. 130. 1923. TYPE: Peru, Weberbauer 4658a ( en; MO, photo). Tree or shrub to 7 m tall, branches slender. Leaves elliptical, to 12 cm long, 4.5 cm wide, apically acuminate, the acumen ca. 10 mm long, basally obtuse, the costa subplane above, only slightly round raised beneath, the lateral veins incon- spicuous, ca. 16 on each side, straight, slightly ascending, branching at the margin but not forming a distinct submarginal vein, intermediate veins 2-3 between each pair of lateral veins, these sometimes as prominent as the lateral veins, drying green, glabrous; petiole slender, 6-9 mm long. /nflorescence terminal or axillary, cymose, glabrous, short or elongate, the bracts and bracteoles alike, cucullate, spreading, the peduncle and pedicels slender, drying yellowish, 4 mm long, be- coming 9-10 mm long in fruit. Flower buds globose, 2 mm across, outer sepals V4 as long as the bud, inner sepals as long as the bud but not overtopping it, imbricate, unmarked. Fruit ca. 15 mm long, red (Croat 16754), drying reddish, tipped by 3(?4) persistent dark styles, mostly faboid as a result of abortion, one pair of sepals spreading; only 1 seed maturing, black with a white aril (Croat 16754) This species is distinct in its membranous leaves, which appear to be many- nerved, and in the small flower buds. Because only one flowering Panamanian specimen was available and that with only two buds, the flower was not dissected. Hence, identification of this species with Chrysochlamys micrantha is tenuous at best. The specimens at hand have partly aborted fruits and only 3 styles. It is assumed that these specimens are abnormal and that 5 seeds commonly mature and 5 styles are usual in the species. The Panamanian collections have inflores- cences no longer than 5 cm long whereas that of the type collection is 9 cm long. Chrysochlamys tenuifolia Cuatr.!^ of Colombia may be the same species. 15 Chrysochlamys tenuifolia Cuatr., Revista Acad. Colomb. Ci. Exact. 29: 59. 1950. Type: Co- lombia, Cuatrecasas 10880 (US). 1038 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 ARIÉN: Trail from Cerro Mali to Río Pucuro base camp, Serrania del Darién, Gentry & Leon 16832 (MO). Río Tacarcuna near old n village, 580 m, Gentry & Mori 13577, 13586 (both MO). N slope of Cerro Mali on Colombian border, 1400-1500 m, Gentry & Mori 13806 (MO). SAN BLAS: Forest SE of Puerto Obaldía, Croat 16754 (MO). 11. Tovomitopsis myrcioides (Pl. & Tr.) D'Arcy, comb. nov. Chrysochlamys id pidan Pl. & e m Sci. Nat. Bot. Sér. 4. 14: 260. 1860. Type: Colombia, Triana (P, not seen; US, MO, Tovomita mac sere L. O. Williams, "Trop. Woods 111: 16. 1959, non Klotzsch ex Schomburgk (1848), nec Walp. (1842). TYPE: Costa Rica, Allen 2968 (US). T. icti en L. O. Williams, Trop. Woods 112: 96. 1960. New name for Tovomita macrophylla , non Klotzsch ex Schomburgk (1848), nec Walp. (1842). Tree to 15 m tall, twigs robust, drying reddish brown. Leaves elliptical, to 25 cm long, 15 cm wide, apically acute, acuminate or obtuse, basally acute or obtuse, the margins minutely revolute, the costa slightly elevated above, rounded-raised beneath and drying striate, lateral nerves numerous, 17-19 on each side, nearly straight, evenly spaced, arcuate at the margin and indistinctly loop-connected, not forming a defined submarginal vein, coriaceous, glabrous, the lesser venation obscure, intermediate veins few, often short, secondary lateral veins ascending, crossing the lamina between the lateral veins; slightly discolorous, darker above, sometimes drying pinkish above; petiole stout, to 5 cm long, slightly channeled above, especially at the base, sometimes minutely puberulent, drying striate. Inflorescences terminal, to 15 cm long, cymose pyramidal panicle, the lateral branches to 7 cm long, drying sharply compressed, stout, stouter and broader in fruit; bracts to 8 mm long, rarely subfoliaceous and much longer, rachis minutely puberulent; pedicels sharply compressed, to 7 mm long, 3 mm wide, subtended by pairs of broad bracteoles to 2 mm long. Staminate flowers with buds globose, ca. 6-8 mm across, sepals 5-6, coriaceous, the outermost somewhat divaricate, about ! as long as the bud, rotund, basally imbricate, third sepal nearly over- topping the bud, inner sepals overtopping the bud and enfolding the interior portions; petals 4—5, thick, shorter than the inner sepals, overtopping the bud, the innermost overtopped by some of the stamens, inconspicuously nervate; sta- mens numerous (129), the filaments free, slender, the outermost shorter, about as wide as the anthers, the thecae apically converging; ovary obsolete. Fruit (dried) ovoid or obovoid, 3.5 cm long, ca. 2 cm across, dark red, collapsing and irregularly sulcate, slightly narrowed basally, apically obtuse, the subsessile stig- mas 5, persistent, dark, ca. 2 mm long, ca. 2 mm across; perianth persistent, divaricate, the pedicels 10-15 mm long, 5-7 mm thick, the bracteoles persistent, costate, ca. 4 mm long, 3 mm wide; seeds 1-2 per locule, fabiform or ellipsoidal, erect, affixed basally to the carpel septum (axile), enveloped by a large, free basifixed aril, ca. 20 mm long, 3 mm wide, the chalaza discoid, 2 mm across, prominent, the testa ?fleshy, arilloid-like on the basal !⁄4 on one side, venose along one side. This species is characterized by its large coriaceous leaves which sometimes dry pinkish and which have many evenly parallel veins. The fruit with its large, persistent, coriaceous perianth is also distinctive. It is closely related to T. allenii, from which it may be only varietally distinct. The type collection of Tovomita grandifolia is quite glabrous, but other collections seen are minutely puberulent, 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1039 and Planchon and Triana comment that the type collection of T. myrcioides is also puberulent. Tovomitopsis myrcioides ranges from Costa Rica to Colombia and is known in Panama from lower elevation cloud forests, below 1,300 m. COCLÉ: Cerro Pilón, Dwyer 8628 (MO). COLON: Santa Rita lumber road ca. 15 km E of Colon, Dressler 3802 (MO). PANAMÁ: Cerro Campana, Croat 22840 (MO). Goofy Lake, Dwyer 8028 (MO). erro Campana, Mori & Bolton 7683 (MO). vERAGUAS: 5 mi W of Santa Fé on road past Escuela Agricola rim Piedra, 800-1200 m, Croat 23151 (MO). 12. Tovomitopsis nicaraguensis Oerst. ex Pl. & Tr., Ann. Sci. Nat. Bot. Sér. 4. 14: 266. 1860. TYPE: Nicaragua, ad flumen San Juan, Oersted 2 (C, not seen; MO, photo).—Fic. 11C Chrysochlamys eclipes L. O. Williams, Trop. Woods 111: 15. 1959. rvPE: Panama, Standley 31459 (US). Tovomita nicaraguensis (Pl. & Tr.) L. O. Williams, Trop. Woods 111: 18. 1959. Shrub or small tree to 11 m tall, glabrate; twigs terete, minutely puberulent, becoming rough and dark gray; sap yellow. Leaves obovate, to 25 cm long, 10 cm wide, apically acuminate, the acumen 1—3 cm long, basally acute, subcoria- ceous, the costa prominent beneath, subplane above, the lateral veins 8-13 on each side, ascending, straight and ascending near the margin, looping to form an inconspicuous submarginal vein, discolorous, drying slatey with the venation inconspicuous above, greenish or yellowish brown beneath with prominent, red- dish brown veins, the minor venation inconspicuous, consisting of short inter- mediate veins and short secondary pinnate veins; petiole puberulent, 1.5-3.0 cm long, narrowly channeled above. Inflorescence terminal or on short shoots, a pyramidal panicle to 15 cm long, the axis, branches and sometimes the bracts minutely puberulent, the branches spreading, subtended by minute, scalelike bracts; pedicels ca. 15 mm long, bibracteolate about 1⁄4 way along, longer and slightly stouter in fruit. Flowers with the buds globose, 2-3 mm across, drying reddish brown, not evidently marked; sepals 5, the outermost as long as the bud or shorter, imbricate, the innermost overtopping the bud, coriaceous, the canals scarcely evident to the naked eye; petals 5, yellow (Croat, 1978), thick, rotund, ca. 4 mm long; stamens numerous (87), the filaments unequal, compressed, some bent, some anthers reduced; gynoecium obsolete; pistillate flowers (Croat, 1978) with ca. 20 stamens; ovary ca. 2 mm long, the stigmas 5, | ope Fruit red, ca. 2 cm long, ovoid, 5-carpellate, the stigmas sessile, deltoid, ca. 1 mm long, the carey rosette ca. 3 mm across; locules 1-ovulate, l aon separating widely into 5 valves; seeds 5, narrowly faboid, ca. 1 cm long, erect, attached basally ee green or black, enveloped in an orange aril which is covered by a white membrane. Tovomitopsis nicaraguensis is the most common species of the genus in Pan- ama. It may be recognized by its relatively large obovate leaves with strong venation, its minutely puberulent inflorescence, and by the size of the fruit and flower buds. It is similar to 7. myrcioides, but the leaves, flower buds and fruit are smaller. This species ranges from Nicaragua to Ecuador and Peru. In Panama it occurs in lowland forests on both sides of the isthmus. 1040 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 AS DEL TORO: ga Peak N of Rio Terebe across from Quebrada Huron, 300—900 ft, Kirkbride & Duke 568 (MO). h Creek Mountains, Chiriqui Lagoon, Wedel 2300 (US). CANAL ZONE: Barro Colorado Island, ier eli 499 (US); Croat 4093 (MO. SCZ), 4102, 4285, 4606, 4651, 6189, 6626 (all MO), 6653, 7298 (both MO, SCZ), 7379 (MO), 8228 (MO, SCZ), 8320, 8355, 8647, 9262, 10902, 11313, 11479, 12663, 16544 (all MO); Dodge et al. 17002 (DUKE); Dwyer et 8443 (SCZ); Ebinger 233, 255 (both MO); Foster 495, 598 (both DUKE), 1237 (DUKE, MO, PMA), Kenoyer 448, 450 (both US); Knight, 1967-68 (MO); Maxon et al. 6792 (US); Shattuck 533, n. (both MO); Standley 31445 (MO, US), 3/459, 41119 (both US); Starry 300 (MO); Wetmore & Abbe 81 (MO); Woodworth & Vestal 548 (MO); Zetek 3623 (MO). Gamboa Navy Pipeline Road, Correa et al. 536 (DUKE, MO). W arm of Quebrada Salamanca, 70 m, Dodge et al. 17002 (MO). Quebrada La Palma and Canon of Río Chagres, 70-80 m, Dodge & Allen 17364 (DUKE, MO). Pipeline Road, Gentry 1946 (MO, SCZ), 3396 (MO). Headwaters of Río Chinilla above Nuevo Limon, Maxon 6888 (US). Pipeline Road, Mori & Kallunki 2416, 2825 (both MO). Río Mendosa above Pipeline Road bridge 8 km NW of Gamboa, 95 m, Nee 7605 (MO). End of Pipeline Road 19 km NW of Gamboa, Nee & Smith 11113 (MO). Hills N of Frijoles, ip 27445 (MO, US). COLON: Rio Guanche 3-5 mi inland, 10—100 m, Croat 26188 ee Purdue a Ridge, Foster 2 Mad 2218 (DUKE); Gentry 6544 (MO). Near Peluca on road t mbre "n Dios, Kennedy 2794 (MO). Trail behind Peluca meteorological station, Kennedy & Dude 2961 (MO). Río idc 2-5 km upriver, Mori & Kal- lunki 3691 (MO). Santa Rita Ridge, Mori & Kallunki 5049 (MO). Loma de la Gloria near Fató (Nombre de Dios), Pittier 3853 (US). Between Peluca Hydrographic Station and Quebrada Peluca along Río Boqueron, Steyermark & Allen 17227 (MO). DARIÉN: Cerro Pirre, Bristan 1235 (MO); Duke 261 (ECON, MO, OS), 6586 (MO). Without other locality, dob 8208 (MO, OS). Río Aretí and Manené, Was 8796 (MO, OS, US). Cerro Campamento, Duke 15646 (MO). Trail Qs Tacarcuna Village on o Tacarcuna to Cerro Mali, 800-1300 m, Gentry & ra 13617 (MO). Manene to mouth of R um, Kirkbride & Bristan 1395 tg Cerro Pirre, 300-700 m, Mori & Pus 5398 (MO). Cen of Cana-Cuasi Trail, Chepigana District, 5500 ft, Terry & Terry 1603 (MO). PANAMA: Cerro Campana, 2300 ft, Blum et al. 2378 (SCZ). Cerro Jefe and Cerro Azul, Correa & Dressler 718 (MO); Dwyer 2053 (MO); Dwyer & Hayden 8094 (MO); Foster 1157 (MO, DUKE); Foster & Kennedy 1821 (DUKE); Gentry et al. 3396 (MO). e to Cartí 16 km above El Llano, 350—400 m, Kennedy 2543 (MO). Cerro Jefe, 3000 ft, Lewis & Dressler 7581 (MO). El Llano-Cartí road 8-12 km from junction 15-20 km WNW of Cerro Azul, Mori et al. 6915 (MO). El Llano-Cartí road 8 km N of Panamerican Highway, 450 m, Nee & Warmbrodt 10358 (MO). SAN BLAS: Permé, res 247 (US). Between Rio Diablo and Rio Areti, Duke 14884 (MO). 13. Tovomitopsis standleyana (L. O. Williams) D'Arcy, comb. nov. Chrysoc Eg s standleyana L. O. Williams, Trop. Woods 111: 16. 1959. New name for C. pauciflora Stan C. paucifiora Standley, Ei E 214. 1952 [1953]. non Walp. (1842). TYPE: Panama, Wedel 1019 (F, holotype; MO, isoty Tree or shrub to 6 m tall, twigs slender; sap yellow, sticky. Leaves obovate or elliptical, to 20 cm long, 7 cm wide, apically acuminate, the tip minutely callose, basally obtuse, or acute; costa subplane above, elevated and conspicuous beneath, especially proximally, the lateral veins 7—8(—10), wide spaced, arcuate, not forming a distinct submarginal vein, minor venation obscure, of weak irregular intermediate veins more numerous towards the apex of the leaf and well devel- oped secondary lateral veins which form a reticulum; canals rarely evident, the major veins sometimes minutely puberulent beneath; petiole 1-3 cm long. /nflo- rescences terminal but sometimes soon lateral on short shoots, columnar or py- ramidal cymose panicles, the main axis mostly unbranched, lateral branches short, compressed and sharply angled, minutely puberulent, bracts minute, brac- teoles nearly basal, minute, glabrous, pedicels 4-6 mm long, terete, compressed or quadrate, becoming stout, 3 mm thick in fruit but not longer. Flower with buds globose, 4-5 mm across; outer sepals 2, basally imbricate, shorter than the bud, sometimes unequal, spreading, coriaceous, inner sepals 3, imbricate, over- 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) 1041 topping the bud; petals 5, imbricate, quincuncial, overtopping the bud; stamens numerous (70-80), free, the filaments unequal, thicker than the anthers, the cen- tral few apically bent down in bud, the anthers convergent upwards, the connec- tive minutely apiculate; ovary obsolete; pistillate flowers not seen. Fruit drying reddish, subglobose, the body 10-12 mm across, rounded-sulcate, apically flat or depressed, surmounted by the persistent, subsessile styles, 2 mm long, basally narrowed into an obconical stipe 7 mm long and ca. 5 mm wide. This species is easily confused with T. nicaraguensis, which occupies similar habitats in Panama but which is usually a larger tree. The leaves of T. standleyana seldom dry brown, and the mature fruits are smaller with less pronounced apicu- lum but relatively similar stipe. The inflorescence of 7. standleyana is usually less puberulent than that of 7. nicaraguensis, and the mature flower buds are usually slightly larger. Tovomitopsis standleyana occurs in Costa Rica, Panama, Colombia (Chocó) and Peru. It may not be distinct from some South American material going under other names. BOCAS DEL TORO: Changuinola Valley, Cooper & Slater 71 (US). Near Quebrada Lugron and Cerro Boynic near Rio Pire 300-900 m, Kirkbride & Duke 643 (MO). Near Almirante, 100 m, Lao & Gentry 447 (MO); Nee & Hansen 14103 (MO); Slater 68 (US). Chiriqui Lagoon, Wedel 1019 (F, MO). Fish Creek Hills, Wedel 2300 (MO). cocrÉ: N of El Valle, 1000 m, Allen 3683 (MO). La Mesa above El Valle, Say 14394 (MO). Cerro Pilón, 700-900 m, Duke 12153 (MO). La Mesa, 2500 ft, Dwyer & Duke € MO). Cerro Pilón, Dwyer & Lallathin 8680 (MO). Cerro Gaital, Caracoral, 2700-3200 ft, Dw n. 7 (MO). Above EI Valle, 1000 m, Gentry 5651, 5659 (both MO). Cerro Pilón, 800-945 m, Mori i al. 6562 (MO). La Mesa, 875 m, Nee & Dwyer 9159 (MO). COLON: 2-3 mi up Rio Guanche, Kennedy & Foster 2193 (MO). 10 mi SW of Portobelo and 2-4 mi from the coast, 10—200 m, Liesner 1048 (MO). VERAGUAS: xe km NW of Santa Fé, Croat 27581 (MO). 0.6-1.8 mi m Escuela Agricola Alto Piedra (Santa Fé), 730 m, Croat & Folsom 34053, 34265 (both MO). 5 mi W of Santa Fé past Escuela Agricola Alto Piedra, 800-1200 m, Liesner 889 (MO). Slopes of Cerro 1 Tute NW of Santa Fé, Mori & Kallunki 4794 (MO). INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (+) refer to names incidentally mentioned. Asthotheca 993 Chloromyron 1010 Astrotheca 993 verticillatum 1010+, 1011 Bertolonia 1025 ic bc 1019%, 1025, 1026+ Biwaldia 997 clusia si Bonnetiaceae 1005+ —ssp. ANNE 996 Calophyllum 971, 1011+ Pages fe 1026 brasiliens eclipes 1039 —ssp. longifolium 973 laxa 1026+ calaba 971+, 1011+ membranacea 1034 edule 1012 micrantha 1037, 1037+ inophyllum 972 multiflora 1025+ longifolium 971+, 973 myrcioides 1038 madruno 1011 pauciflora 1040+, 1040 nubicola 973 pavonii 10 rekoi 971+ standleyana 1040 soulattri 975 tenuifolia 1037+ Calysaccion 1002 ulei 1037+ longifolium 1002+ Clusia 975, eu a Cambogia 997, 997+ —subg. Cri gutta 997+ —sect. Asta 993 1042 ANNALS OF THE MISSOURI BOTANICAL GARDEN clusioides 990+, 991+ coclensis 977, 990+ congestiflora 978, 980+ i 9761 divaricata 981 dukei 982 erectistigma 987, 988+ flava t fructiangusta 983 liesneri 979+, 980+, 983, 985+ lineata 975+, 984 longifolia 1020, 1020+ longipetiolata 985, 989+ longisty la major 975+, 986 minor multiflora 992+ oblanceolata 1023 odorata 987, 987+ oedematopoidea 989 s 987 cheep 978+ retusa rosea 986, 986+, 987+ rotundata 978+, 979+, 9847, 990 salvinii 980+, 991 — var. cupulata 980 seemanii 984 stenophylla 976+, 992 uvitana 987, 988+ venosa 987 Clusianthemum 997 pedicellatum 997+ Dactylanthera 997 Dystovomita 995, 1019+, 1026+ pittieri 995+, 9 Ficus 986 Garcinia 997, 1010+, 1011+, 1012+ —subg. Garcinia 997+, 1011+ —subg. Rheediopsis 997+, 1010+, 1011+ —subg. Xanthochymus 997+, 1011+ —sect. Rheediopsis 997+, 1010+ —sect. Teracentrum 1010+ umilis 1015 lateriflora 1014+ macrophylla 1015 mangostana 997+, 998, 10007, 1011+, 1015+ pictorius 997+, 998, 1011+, 1016+ tinctoria 999 xanthochymus 998 Guttiferae 969, 1003+, 1005+, 1025+, 1029+ —tribe Moronobeae 1017+ Havetia 1000+ flexilis 1000+, 1000 c een 1000 1000+, 1000 T Lamprophyllum 1010, 1011+ laetum 10107, americana 1002+, 1003, 1004+ humilis 1015, 1015+ —var. plumieri 1015 —var. vahlii 1015 immansueta 1003+, 1004 Marila 1005 cespedeziana 1010+ domingensis 1006 m d 1005+ pluricostata 1008 racemosa 1005+ verapazensis 1008 Micranthera clusioides 1020 Ochrocarpus 1002, 1003+ madagascariensis 1002+ Potamocharis | mamei 1002+ Rheedia 997+, 9981, i 1012+, 1015+ acuminata 1010+, —va E T idu 1015 * lateriflora 997+, eun 1014, 1016+ macrophylla 997+, 10 sagotiana rh 1016+ ieberi | sessiliflora dfi Solanum ducere 1029+ Symphonia EETA T 1017+, 1017 Tovomita 1019, 1022+, 1026+, 1029+, 1031+ —s n an Dystovomita 995 032 Eh 1023 [VoL. 67 1980] D'ARCY—FLORA OF PANAMA (Family 123. Guttiferae) grandifolia rela 1038 longicuneata 1023 longifolia 1020 macrophylla 1038, 1038+ martiana 1032 membranifolia 1035 multiflora 1032 nicaraguensis 1039 sphenophylla 1023 stylosa 1020+, 1022 tenuiflora 1022+ weddelliana 1020+, To ovomitopsis 1019+, UM 1022t, 1025, 1029+ 8 centistaminibus 1030 croatii 1031 faucis 1031 glauca edi n 1036+ 1034 Tsimatimia 1010 areas p. 1011, 1011+ a 1010+ Vis smia sar aap = 997+, 999+ ictorius 998, 9 ata 998, 999+ 1043 FLORA OF PANAMA BY ROBERT E. WoopsoN, JR. AND ROBERT W. SCHERY AND COLLABORATORS PART VI FAMILY 124. CISTACEAE Although listed in indexes to the families to appear in the Flora of Panama, the Cistaceae has not been found in Panama nor is it expected. The family, which includes herbs and shrubs, occurs in temperate regions of the world, especially in the Mediterranean area.—W. G. D’Arcy. ANN. Missouri Bor. GARD. 67: 1045. 1980. FLORA OF PANAMA! BY ROBERT E. WoopsoN, JR. AND ROBERT W. SCHERY AND COLLABORATORS Part VIII FAMILY 151. THEOPHRASTACEAE W. G. D’Arcy? Shrubs or trees, rarely acaulescent; sometimes dioecious, wood often brittle and breaking along enlarged rays. Leaves simple, opposite, mostly coriaceous, entire or serrate, sometimes spinose, the venation pinnate, all species with char- acteristic linear sclereids running under the epidermis oriented without reference to the venation. Inflorescences short or elongate racemes, mostly with minute bracts subtending the short pedicels. Flowers white, orange or green, 3—5 merous, the sepals imbricate; corolla rotate or campanulate, basally united into a short tube, the lobes mostly rotund; staminodes inserted on the corolla near the base of the tube or at the mouth, often resembling the petals; stamens free or united, the anthers with the thecae distinct but coalescent, the ovary 1-locular, the pla- centation free central, ovules numerous, multiserriate, anatropous. Fruit a berry or drupe, the pericarp coriaceous or testaceous, indehiscent; seeds few or nu- merous, often immersed in a soft or gelatinous pulp; embryo straight, endosperm present. This family of about five genera is centered in the Caribbean area with small genera, Deherania (Guatemala to Costa Rica), Theophrasta (Hispaniola), and Neomezia (Cuba) of local occurrence, and the two larger genera, Jacquinia and Clavija of much wider occurrence. Jacquinia occurs mainly in open hapitats of northern South America, the Antilles and and Central America, while Clavija occurs in deep humid forest with species more abundant in South America. In some places species of Jacquinia are used as barbascos to kill fish, while in some places, fruits of Clavija are eaten by man. Although there are two monographs of the family, one based on gross mor- phology (Mez, 1904) and one on foliar anatomy (Votsch, 1904), the taxonomy of the group is far from understood. There are difficulties in the study of this family: the small genera are known by only very scant material; Jacquinia is abundantly represented in herbaria, but plasticity makes species difficult to separate, and herbarium specimens of Clavija usually lack floral information adequate for un- derstanding of species or relationships. ! Assisted by National Science Foundation Grant DEB 77-04300 (W. G. D’Arcy, principal in- vestigator). 2 Missouri Botanical Garden, Box 299, St. Louis, Missouri 63166. ANN. Missour!I Bor. GARD. 67: 1047-1055. 1980. 1048 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Literature: D'Arcy, W. G. 1973. Correlliana (Myrsinaceae), a new palmoid genus of the tropical rain forest. Ann. Missouri Bot. Gard. 60: 442-448. Mez, C. 1903. Theophrastaceae in A. Engler, Das Pflanzenreich 15(IV 236a). Votsch, W. 1904. Neue systematische-anatomische Untersuchungen von Blatt und Achse der Theophrastaceen. Bot. Jahrb. Syst. 33: 502-546. Wood, C. E., Jr. & R. B. Channell. 1960. The genera of the Ebenales in the southeastern United States. J. Arnold Arbor. 41: 1-35. a. Leaves small, less than 4 cm wide, spine tipped; plants freely branching; flowers perfect . Jacquinia aa. Leaves large, more than 6 cm wide, lacking spines; plants unbranched; flowers unisexual and plants dioecious __________________---_-_-------------------------------------------------- l. Clavija 1. CLAVIJA Clavija Ruiz & Pavón, Fl. Peru. Chil. Prodr. 142, tab. 30. 1794. svNTYPES: C. macrocarpa Ruiz & Pavón, C. longifolia Ruiz & Pavón, C. spathulata Ruiz & Pavón and C. pendula Ruiz & Pavón. Shrubs or trees, erect, mostly unbranched. Leaves simple, alternate, clus- tered at the tip of the stem, mostly elongate, entire or serrate, mostly coriaceous and glabrous, with numerous linear sclereids running beneath the epidermis. Zn- florescences dioecious, mostly cauliflorous racemes clustered amongst the leaves; peduncles slender in flower, stouter in fruit; pedicels subtended by minute bracts. Flowers 3—5 merous, the sepals small, rotate, persistent; petals united into a short tube, rotate, the lobes rotund, fleshy, bearing staminodes as thickened append- ages at the corolla mouth; staminate flower with the filaments united into a stipe- like tube, the anthers forming a peltate disc at the mouth of the corolla, dehiscing on the lower surface, the gynoecium rudimentary or wanting; in pistillate flowers, the stamens free, reduced and sterile; ovary glabrous, the stigma subsessile. Fruit a coriaceous berry, mostly globose, the pericarp becoming hard, brittle; seeds several, the embryo straight. This genus includes several dozen species of humid forests of tropical Amer- ica. In the field, they may usually be recognized by their erect, unbranched habit, elongate coriaceous leaves and globose fruits. In the herbarium, the linear scler- eids just beneath the leaf epidermis is a near-diagnostic feature, occurring oth- erwise only in Correlliana D'Arcy, Myrsinaceae, which differs in inflorescence and flora characters. The taxonomy of this genus is extremely difficult to study, for most herbarium specimens have little in the way of floral material suitable for comparison. Leaves of different species often look quite similar, and it is not always possible to determine species on vegetative characters alone. Whether leaves are entire or have spines seems to be a variable character within species and cannot be relied upon for taxonomic separations. Perhaps field studies looking at the nature of plasticity under different growing conditions would be useful. Quite similar to Clavija in aspect are Gustavia, Lecythidaceae, Corelliana, Myrsinaceae, and a number of genera in other families, but only Clavija and 1980] D'ARCY—FLORA OF PANAMA (Family 151. Theophrastaceae) 1049 Corelliana have the linear sclereids in the leaf lamina. For a review of these genera see D'Arcy (1973). a. Leaves glabrous or with minute trichomes when viewed under high magnification. Leaves broadest near the middle; petioles slender 1. C. biborrana bb. Leaves oa well above the middle; pedicels mostly st or venation visibly reticulate and low elevated i on leaves mostly narrow je" than 8 cm wide | spathulata cc. Minor venation mostly inconspicuous; leaves mostly more than 8 cm "wide _ 2. C. jelskii aa. Leaves evenly short pilose beneath with erect hairs C. pubens — . Clavija biborrana Oerst., Vidensk. Meddel Dansk Naturhist Foren. Kjoben- havn 1861: 117. 1861. TYPE: Costa Rica, Oersted 31A (C, not seen; GH, photo). Erect shrub to 4 m tall, unbranched, the leaves born at the stem apex, the leaf bearing portion of the apex mostly 4-5 mm wide. Leaves ovate, acute or acuminate at each end, mostly 15-25 cm long, 5-9 cm wide, entire, stiff and coriaceous, often drying with a light gray cast, the lateral veins nearly straight, only slightly ascending, forking near the margin and forming a submarginal vein and in places a marginal vein, the minor venation indistinct, the tertiary veins more or less at right angles to the secondaries near the center of the leaf; petiole unwinged or inconspicuously so, slender, flat and drying angled, slender, elon- gate, often drying black in the basal 1 cm, glabrous. Inflorescences (ex Mez) few flowered, ca. 10-flowered, laxly racemose, glabrous, much shorter than the leaves; pedicels slender, to 3 mm long; bracts narrowly triangular bearing scales near the base. Flowers (ex Mez) small or to 5 mm diam., the sepals ovate, rotund, fimbriate-crenulate; staminodes thick, subglobose, not joined, the anthers free, exserted, the free part of the tube shorter than the anthers. Fruit (immature) globose, minutely apiculate, subtended by the persistent, greenish, applied sepals, 5-10 mm across. Described from Costa Rica, this species has been found in several parts of lowland Panama. Material cited here is only tentatively determined, and the name used is based on the appearance of the type photo. This species appears to be distinct in its elongate inconspicuously winged slender petiole, and in its entire symmetrical and relatively broadly elliptic leaves which are widest about the middle. All fruit seen was quite immature, and it is not known how large it may grow. BOCAS DEL TORO: Chiriquí Lagoon, Wedel 1024 (MO). cocLé: El Valle de Antón, Duke 13238 (MO, OS). Mountains beyond La Pintada, 400—600 m, Hunter & Allen 580 (MO). EI Valle, Lindsay 411 (MO). DARIEN: S slope of westermost peak of Tacarcuna massif, 800-1200 m, Gentry et al. 16963 (COL). PANAMÁ: Cerro Campana, Croat 17223 (MO); Kirkbride & Hayden 293 (MO, SCZ); Smith & Smith 3378 (US). 2. Clavija jelskii Szyszyl. Diss. Cl. Math. Phys. Acad. Litt. Cracov. 29: 225. 1894. TYPE: Peru, Jelski 260 (W, not seen; MO, photos).—Fic. 1. C. costaricensis Pittier, Contr. U.S. Natl. Herb. 20: 131. 1918. TYPE: Costa Rica, Cook & Doyle 551 (US-474411, holotype). 1050 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 Lidah Syor "C wid NS IN : dk — TEDN FicunE 1. Clavija jelskii Szyszyl. Habit (x'&). [After Dwyer & Gentry 9376 (MO).] Unbranched shrub 1-4 m tall; leafy portion of twigs 5-20 mm thick. Leaves large, 30-50 cm long, obovate, apically acute, basally cuneately attenuate into the petiole, entire or with shallow teeth, the lateral veins prominent, elevated beneath, nearly straight, evanescent towards the margin, branching to form an inconspicous and somewhat irregular marginal vein, the minor venation indistinct beneath, sometimes appearing somewhat reticulate above, coriaceous, mostly drying light green; petioles stout, especially at the darkened base, narrowly winged to 2-3 cm from the base, or conspicuously narrow winged to within 10 mm of the base. Inflorescences cauliflorous racemes clustered amongst the leaves, the peduncles slender, minutely puberulent, drying orange-yellow, to 15 cm long, the flowers evenly spaced; pedicels ca. 1.5 mm long, subtended by a single, cucullate, ciliolate bract 0.5-1 mm long; staminate flowers ca. 5 mm across, rotate, the sepals 4, rotate, 1-1.5 mm long, cucullate, the margins hyaline; petals forming a tube ca. 1 mm long, shorter than the sepals, the petals rotund, fleshy, orange, staminodes reduced to small flaps at the corolla mouth; filaments united into a narrow spike in the floral center, the 4 anthers connivent into a peltate disc, lobed and dehiscing on the underside where impressed by the stami- nodial flaps; pistillate inflorescences apparently much more contracted than the staminate, pistillate flowers not seen. Fruits globose, to 4 cm across, sometimes minutely apiculate, yellowish, the pericarp becoming hard, brittle, minutely muri- cate; seeds numerous, drying dark reddish brown. 1980] D'ARCY—FLORA OF PANAMA (Family 151. Theophrastaceae) 1051 This species is apparently widespread in tropical America, occurring in trop- ical wet and moist forests. Although the material cited for this species is super- ficially uniform, it is questionable that it is really all the same species. Therefore, the name employed here must be considered as provisional. CANAL ZONE: Mojinga swamp near mouth of Rio Chagres, Allen 856 (MO). Navy reservation N of Gamboa near Rio Mendosa, Dressler 2899 (PMA). Bank of Rio Agua Salud between Frijoles and Pipeline Road, Foster and Kiester 1945 (DUKE, MO). Crest ridge along road to battery 7, area of Limon Bay, Gatun Locks and Gatun Lake, Johnston 1802 (A, MO). Pipeline Road N of Gamboa, Luteyn & Foster 1557 (DUKE). 12 mi S of Colón on Rio Providencia, Tyson & Blum 3986 (MO). CHIRIQUÍ: Without other locality, Cooper & Slater 191 (MAD). San ie Limité, 12 mi W of Puerto Armuelles, 400-500 m, Liesner 202A (MO). Between Canazo and cos near Puert bib ner me Stern & Chambers 142 (F, MAD, MO). coci É: Below Cerro Pilón, Gta 13472 (MO). COLON: 9 m S of Portobelo, ey 11381 (MO). Santa Rita Ridge, Dwyer & Gentry 9376, 9561 (both MO). DARIEN: Mouth of Rio Yapé, 20 m, Allen 343 (MO). Panamerican Highway ex E inet Rio Canita, Duke 3849 (MO). 10 cf) S of El Real on Río Pirre, Duke 5470 (GH, MO). i N of Paya, Duke & Kirkbride 14002 (NY). Santa Fé, Duke 14273 (MO). Yaviza along Río cutem. Stern et al. 88 (GH). Caná, 2000-6500 ft, Williams 806 (NY, US). PANAMA: Camino a Llano-Cartí, Correa & Sucre 1842 (MO, PMA) 3. Clavija pubens D'Arcy.? TYPE: Panama, Cooper 281 (NY, holotype; US, iso- type). Erect, unbranched "shrub or small tree 2 ft. tall (Cooper). Leaves obovate, broadest slightly above the middle, acute at both ends, entire, 25-35 cm long, 9— 12 cm wide, coriacec 1s, copiously pubescent beneath with erect, soft hairs, more or less concolorous, the lateral venation nearly straight and slightly ascending, forking and anastomosing ca. 1 cm from the margin to form a more or less distinct submarginal vein; petioles slender, basally unwinged, 1—4 cm long, drying basally dark. Inflorescences short, perhaps reduced. Flowers not seen. Fruit globose, minutely apiculate, minutely muricate, drying yellowish; seeds drying dark red, ca. 2.5 cm across. This species resembles C. costaricense, and both sheets of the type collection were annotated with that name by Gunnar Harling in June 1964. However, pu- bescent leaf undersides are unknown to me in that species, and the lateral vein termination into an undulate submarginal vein is much more distinct than in material seen of C. costaricense. This species was recognized in the herbarium by Standley as distinct. SAN BLAS: Permé, Cooper 281 (NY, US). 4. Clavija spathulata Ruiz & Pavón, Syst. Veg. Fl. Peru. Chil. 1: 285. 1798, non Hooker (1844). TYPE: Peru, Pavón (G-DC, not seen; MO, microfiche). Theophrasta i si Jacq., Coll. 4: 136. 1790. TYPE: Venezuela, Jacquin (W, not seen; MO, photo). Calvija ornata D. Edinburgh Philos. J. 10: 236, 1831; Bot eia tab. 1764. 1836. TvPE: Not een. N i ea aa spec. prior., Theophrasta longifolia J C. longifolia (Jac q.) Mez, in Urb., Symb. Ant. 2: 438. 1901, nec s longifolia Ruiz & Pavon (1798) C. longifolia Rusby (ined.). 3 Clavija pubens D' Arcy, spec. nov. Clavija costaricense simile, differt foliis subtus pilosulus; venis foliorum valde distinctis. 1052 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VoL. 67 C. mezii Pittier, Contr. U. x DA Herb. 20: 132. 1918. type: Panama, Pittier 4313 (US-679440, holotype; NY, US, iso C. allenii Lundell, Ann. asp Bot. Gard. 28: 452. 1941. TYPE: Panama, Allen 2671 (MO, holotype: MICH, isotype, fragm. not s Tree to 10 m tall; leafy portion of the stems 5-7 mm wide. Leaves narrowly obovate, widest 34-34 way up, apically acute or acuminate, basally cuneate, 30- 40 cm long, 7-9 cm wide, stiff and coriaceous, glabrous, entire or inconspicuously serrate, drying with the minor venation conspicuously elevated reticulate; cuneate and winged to near the base, the slender petiolar area relatively stout, 1-2 cm long, drying dark. Inflorescences short, becoming stout, not clearly observed, the branches drying dark reddish brown (Lundell) pendulous, few flowered, to 36 cm long, sparingly minutely puberulent. Flowers (ex Lundell) 4-merous; sepals basally united, suborbicular, 1-1.2 mm long, fimbriate; corolla ca. 1 cm diam., 6 mm long, orange, the lobes subrotund; ovary of staminate flowers spike-like. This species is distinct from other Panamanian elements of the genus in the coriaceous reticulation of the leaves which sometimes dry somewhat shiny. It is likely that an older name will be found for this species, perhaps C. reticulata Cuatr., or another much earlier name. It somewhat resembles the type of C mezii Pittier, an earlier name from Panama, but that species is here interpreted to be a synonym of C. spathulata R&P. L ZONE: Madden Dam, 100 m, Allen 2671 (MICH, MO). DARIÉN: 2-3 mi Fs of El Real, Duke 4856 (US). PANAMÁ: Without other locality, Duke 5560 (US); Hayes 804 (NY). SAN BLAS: Isla de Soskatupu, Duke 8961 (MO, OS). Isla Pino near Mulatupo, Elias 1724 (MO). Forests around Puerto Obaldia, 0-50 m, Pittier 4313 (NY, US). 2. JACQUINIA Jacquinia L., Fl. Jam. 27. 1759, "Jaquinia." TYPE: J. ruscifolia Jacq. = J. acu- leata (L.) Mez. Bonellia Bert. ex Colla, Hortus Ripul. 21. 1824. type: B. cavanillesii Colla, nom. illeg. = J. macro- ‘ar, iv. Glabrate shrubs or small trees; wood brittle. Leaves mostly alternate or pseu- doverticillate, entire, spinulose tipped, mostly coriaceous and rigid, short petio- late; stipules acicular, or wanting. Inflorescences short racemes or rarely solitary flowers, mostly terminal or nearly so; stipuliform bracts sometimes present; brac- teoles wanting. Flowers mostly perfect, 5-merous, either white or orange; calyx with sepals free or basally connate, imbricate, rotund, sometimes ciliate or cren- ate; corolla salverform, the tube straight, campanulate or urceolate, the lobes imbricate, rotund, thick but thinning at the margins; staminodes 5, resembling the corolla lobes, connate to the top of corolla tube but free above, sometimes dorsally appendaged; the filaments flat, narrowing upwards, basally connate into a tube which is basally adnate to the corolla tube, the anthers ovate, pointed, exserted, the thecae separated dorsally, longitudinally dehiscent, nearly basi- fixed; ovary conical unilocular, the ovules many, the style not demarcated from the ovary, the stigma peltate, entire or minutely lobed. Fruit a coriaceous berry, ovoid, elliptical or globose, sometimes cuspidate, the pericarp thick; seeds large, I—numerous, immersed in a spongey or mucilaginous pulp. 1980] D'ARCY—FLORA OF PANAMA (Family 151. Theophrastaceae) 1053 FIGURE 2. Jacquinia macrocarpa Cav.—A. Habit (x7/,,).—B. Flower (x2*/,).—C. Fruit ( x7/,,). [After D'Arcy 5236 (MO).] Jacquinia is a genus of about 30 species mainly of Central America and the Caribbean region. Species of Central America and some from South America and the Greater Antilles have orange flowers while most of the others from the Antilles and northern South America have white flowers. The flower color seems to speak for a separation of the genus into two reasonable groups. The orange flowered species are difficult to separate, and while there is con- siderable variation between populations and individuals, and some populations 1054 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 seem to represent quite different species from others, it is nevertheless extremely difficult to find separation points in the more or less continuous variation. Keys produced by Mez (1903) and Votsch (1904) are difficult to use and may not reflect differences between valid species. Jacquinia in Panama is usually easy to recognize by its spine-tipped coria- ceous leaves, bright orange flowers, globose, rotund fruits, and coastal habitat. Only one species is present. Species of this genus have been used in many places as barbascos or fish poisons. l. Jacquinia macrocarpa Cav., Icon. 5: 55. tab. 483. 1799. TyPE: Pacific coast of Panama, Cavanilles (B-WILLD., lectotype, not seen; MO, microfiche).— Fic. 2 J. panamensis Lundell, Ann. Missouri Bot. Gard. 27: 329. 1940. Type: Panama, Allen 985 (MICH, holotype, not seen; MO, US, isotypes). J. aurantiaca mult. auct., ?Ait., Ind. Kew., ed. 2. 6. 1811. TYPE: probably Tres Marias Is., Mexico, Menzies (K). Shrub or small tree to 8 m tall; branches brittle; stems drying dark reddish brown, the twigs scurfy puberulent, soon glabrescent. Leaves mostly elliptical, often narrow, mostly 5-7 cm long, 1.5-2.5 cm wide, apically obtuse or acute with a strong dark spine, basally obtuse or rounded, the margins often revolute, co- riaceous, drying light green, the costa impressed and obscure above, prominent beneath, the lateral veins inconspicuous 5-9, forming a submarginal vein, minor venation not evident; petioles ill defined, 3-6 mm long; stipules linear to lanceo- late, 1-6 mm long, scruffy puberulent. /nflorescences short condensed terminal or axillary racemes, round topped, the peduncle and the rachis 1-3 cm long; pedicels ca. 1 mm long, drying orange, subtended by cucullate, lanceolate, cil- iolate bracts 3-7 mm long. Flowers showy orange; calyx cupular, 5-6 mm long, of 5 rounded, ciliolate, imbricate sepals; corolla salverform, the tube ca. 5 mm long, 3 mm wide, the 5 petals rotund, ca. 5 mm long, thinning at the margin; staminodes resembling the petals, but narrower, with 4 longitudinal adaxial ridges (appendages); stamens 5, the filaments flat, basally fused into a tube narrowing upwards, the anthers yellowish white, ovoid, acute, converging around the style; ovary conical, the style not demarcated, the stigma peltate. Fruit a coriaceous berry, ovoid or ellipsoidal, 3-4 cm across, brown, rugose, the pericarp ca. 5 mm thick; seeds I-several, ca. 2 cm long, faboid, immersed in a soft yellowish matrix. Jacquinia macrocarpa may be recognized by its bright orange flowers which are sometimes showy, and by its sharp spined leaves. This species has brighter, showier flowers, larger fruits and larger leaves than most other species in the genus. In Panama it is confined to the Pacific Coast where it grows along the strand with Hippomane mancinella L. (Euphorbiaceae). The one tree cultivated on Barro Colorado Island and Hunter & Allen 657 from the Río Boquerón dem- onstrate that the species is well able to thrive in inland situations. NAL ZONE: BARRO COLORADO ISLAND: ES uA 4119a, 6904 (both MO); Ebinger 600 (F, MO, US); odes 1677 (MO); Zetek 5011 (MO). c : Aguadulce, Pittier 4990 (US). Los saNTOs: Mon- agre Beach, Lewis et al. 1676 (MO); Tyson et n 2885 (MO). Bejuco, Allen 985 (MO, US). PANAMÁ: 1980] D'ARCY—FLORA OF PANAMA (Family 151. Theophrastaceae) 1055 ueva Gorgona, Allen 3763 (MO); D'Arcy 5236 (MO); D'Arcy & D'Arcy 6281 (MO); Duke 4530 (MO). Isla de La Bayoneta, Perlas Islas, Dwyer 1739 (MO). Río Boquerón above Peluca Hydrographic Station, 90 m, Hunter & Allen 657 (MO). Gorgona Beach, Miller 1796 (US); Woodson et al. 1696 (F ) INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (+) refer to names incidentally mentioned. Bonellia 1052 ekle ve 1054+ cavanillesii 1052+ Gustav g 48+ Clavija 1047+, 1048 allenii 105 ieee 1054+ biborrana 1049 Jacquinia 1047+, 1052 costaricensis 1051+, 1049 aculeata 1052 elskii 1049 aurantiaca 1054 longifolia 1048+, 1051, 1051+ macrocarpa 1052+, 1054 macrocarpa 1048+ panamensis 105 mezii 1052, 1052+ ruscifolia 1052+ ornata Jaquini pendula 1048+ Lecythidaceae 1048+ p 1051 Myrsinaceae 1048 reticulata 1052+ Neomezia 1047+ spathulata 1048+, 1051 Theophrasta 1047+ Correlliana 1048+, 1049+ longifolia 1051, 1051+ Deherania 1047+ Theophrastaceae 1047 FLORA OF PANAMA! BY RoBERT E. WoopsoN, JR. AND RoBERT W. SCHERY AND COLLABORATORS Part IX FAMILY 173. PEDALIACEAE W. G. D'AncY? Mostly herbs, often succulent, seldom shrubs, mostly with glandular hairs. Leaves opposite below, sometimes alternate upwards; petiolate, entire, toothed or lobed, often viscid, estipulate. Inflorescences of solitary flowers or few flow- ered cymes, axillary. Flowers perfect, zygomorphic, the calyx (4—)5 lobed to near the base; rarely saccate; corolla sympetalous, campanulate or tubular, 5-lobed, 2-lipped, the lobes imbricate in bud, sometimes gibbous; stamens 4, didynamous, the staminode posterior, the anthers free but often converging, 2-celled, longi- tudinally dehiscent the connective terminated by a gland; pistil 1, the ovary su- perior, the carpels 2 (3-4), the locules (1—) 2 or 4, the ovules 1-many in each locule, axile, anatropous, the style 1, slender, the stigmas 2. Fruit a loculicidal capsule, often appendaged with hooks, spines or wings; seeds occasionally some- what winged, the embryo straight, the endosperm thin. The Pedaliaceae is closely allied to the Martyniaceae with which it is often united, the major distinctions being that the New World Martyniaceae has parietal placentation and woody, indehiscent fruits while the Old World Pedaliaceae has axile placentation and dehiscent fruits. These distinctions are not convincing. The Pedaliaceae (and Martyniaceae) are similar to the Scrophulariaceae, Bignoniaceae and Acanthaceae, and close alliance has been postulated with the first and last of these families. In Panama, only the genus Sesamum is known, and that from cultivated plants which escape and are perhaps naturalized. The balance of the family, which includes about 12 genera, is confined to the Old World tropics, ranging from Africa to the Philippines. Martynia and Proboscidea of the Martyniaceae have species in Mexico and further south in Central America, while Craniolaria, also Martyniaceae, has species in Venezuela and further south in South America. It would not be surprising to encounter species of these genera in Panama, for they are sometimes cultivated for their curious fruits and have been carried from place to place for ritual purposes by indigenous populations. Members of these genera are similar to the one treated here but tend to be more robust, more viscid pubescent, and they have large, woody, fruits with formidable hornlike processes which can damage cattle. 1 Assisted by National Science Foundation Grant DEB 77-04300 (W. G. D'Arcy, principal in- vestigator). 2 Missouri Botanical Garden, Box 299, St. Louis, Missouri 63166. ANN. Missouri Bor. GARD. 67: 1057-1059. 1980. 1058 ANNALS OF THE MISSOURI BOTANICAL GARDEN (VoL. 67 Literature: Bruce, E. A. 1953. Notes on African Pedaliaceae. Kew Bull. 8: 417—429. Nayar, N. M. & K. L. Mehra. 1970. Sesame: its uses, botany, cytogenetics and origin. Econ. Bot. 24: 20-31. 1. SESAMUM Sesamum L., Sp. Pl. 634. 1753. Gen. PI., ed. 5. 282. 1754. TYPE: S. indicum L. Erect or prostrate herbs, often viscid pubescent with glandular hairs and with subsessile glands. Leaves opposite, but sometimes with alternate or cyclical leaves above, entire, serrate, lobed or dissected and sometimes pubescent with subsessile, peltate 4-celled trichomes and with elongate hairs, mostly petiolate. Inflorescences solitary flowers in the upper leaf axils, pedicellate or subsessile; bracts and bracteoles, wanting or obsolete. Flowers sometimes showy, yellow, red, purple or pink; calyx deeply 5-lobed, the lobes similar; corolla campanulate, 5-lobed, slightly 2-lipped, slightly gibbous basally, glandular with short-stipitate 2-4 celled glands; stamens 4, the anthers oblong, versatile, the locules parallel, the connective dorsally thickened and terminated by a gland; ovary oblong, the locules longitudinally demarcated, hispid, the style abrupt, slender. Capsule ob- long, longitudinally grooved or 4-angled, opening apically; seeds numerous, sometimes compressed, smooth or ridged, sometimes rugose, minutely 4-angled. Sesamum is native to the Old World and Australia with 18-34 species de- pending on classification. The greatest number of species is in Africa and India. Various authors have divided Sesamum into smaller genera. The 4-celled glan- dular trichomes are unusual or perhaps unique in the Panamanian flora. A number of species are important crop plants providing edible oils and highly nutritious meals which are used for humans and animals. In addition to Sesamum indica treated here, S. radiata Schum. & Thonn. has been cultivated in Central America and is perhaps naturalized. It is distinct from S. indica in its usually narrowly spatulate leaves and winged and rugose seeds. 1. Sesamum indicum L., Sp. Pl. 634. 1753. LECTOTYPE: Herb. Linnaeus (LINN 802.3, not seen; MO, microfiche). S. orientale L., Sp. Pl. 634. 1753. TYPE: not seen. Erect, fetid, glandular pubescent, annual herb to 2 m tall, somewhat branched; stems 4-angled. Leaves opposite below, often alternate above, variable in shape, entire, serrate, palmately lobed or nearly compound, 8-15 cm long, 6-10 cm wide, pubescent above and below and with copious 4-celled short-stalked glands beneath, the petioles ca. 5 cm long, often pilose. /nflorescence a single flower or 2—3 flowers in the upper leaf axils; pedicels 1-2 cm long with 2 yellow or black basal nectaries. Flowers white, rose or purple, mostly maculate; calyx 6-8 mm long, deeply 5 lobed, the lobes similar, acute or acuminate; corolla campanulate, basally minutely saccate, slightly 2-lipped, pubescent outside and with 1—4 celled glands; anthers pale yellow with a short white or brownish beak; ovary pilose, erect, brown or purple, grooved, with 4 rounded angles, the style slender, gla- 1980] D'ARCY—FLORA OF PANAMA (Family 173. Pedaliaceae) 1059 brous. Fruit apically dehiscent, to 3 cm long, oblong, glabrate to hirsute; seeds smooth or minutely rugose, to 3 mm long. This species is distinct in its usually broad leaves and smooth unwinged seeds. The plants and corolla superficially resemble members of the Scrophulariaceae or Gesneriaceae, but the 4-celled glandular trichomes are distinctive. Sesamum indicum is cultivated in Panama from time to time for its oil pro- ducing seeds, and sometimes it escapes or Is persistent after planting. Ei is known as ‘‘sesame”’ (English) or ‘‘ajonjoli’’ (Spanish). LOS SANTOS: Cultivated, Monagre Beach, Dwyer 1965 (MO). PANAMA: Planted, Rió Tapia, Stan- dley 30657 (MO). INDEX OF LATIN NAMES Numbers in bold face type refer to descriptions; numbers in roman type refer to synonyms; numbers with daggers (+) refer to names incidentally mentioned. Acanthaceae 1057+ Proboscidea 1057+ Bignoniaceae 1057+ Scrophulariaceae 1057+, 1059+ Craniolaria 1057+ oe 1057+, 105 esneriaceae 1059+ indicum 1058+, 1058, 1059+ Martynia 1 orientale 1058 artyniaceae 1057+ radiata 1058+ Pedaliaceae 1057 FLORA OF PANAMA BY ROBERT E. WoopsoN, JR. AND RoBERT W. SCHERY Missouri Botanical Garden AND COLLABORATORS Introduction and Indexes Preface W. G. D'Arcy The Flora of Panama: historical outline and selected bibliography W. G. D'Arcy The history of plant collecting in Panama (1700-1981) John D. Dwyer Research, tropical ecology and the future of Panama Peter H. Raven Contributors to the Flora of Panama W. G. D'Arcy Indexes to the families in the Flora of Panama M. R. Crosby Systematic index Alphabetical index 1943-1981 FLORA OF PANAMA BY ROBERT E. WOODSON, Jr. AND ROBERT W. SCHER Y Missouri Botanical Garden AND COLLABORATORS Flora of Panama is a compilation of the indigenous and natural- ized vascular plants of the Republic of Panama, following generally the system of Engler and Prantl. The treatments of the various families follow consecutively as nearly as is practicable and are of a critical nature, including synonymy, descriptions, and citation of representative Panamanian exsiccatae for each species. The com- pilers of the Flora have been aided in their work very materially by the extensive contributions of American and European specialists of wide repute in the case of numerous critical families. As is customary, such contributions are credited directly to the authors concerned. Flora of Panama will appear in the ANNALS OF THE MISSOURI BOTANICAL GARDEN as contributions accumulate. Since the com- pleted work will be extensive, it will consist of a number of parts designated numerically, each with separate pagination and pertinent illustrations. The first portion of the work to be published will begin the taxonomic treatment of the Gymnospermae and Mono- cotyledoneae, and will be designated as Part II. General discussions of historical, geographical, physiographic and floristic relations, to be published later, will constitute Part I. Full indices, together with addendae and corrigendae, will occupy the final part. The unusual nature of Flora of Panama will necessitate a change in the policy of the Missouri Botanical Garden regarding reprints, which will not be generally distributed except to collaborators. Reprints of the various parts of the Flora, suitable for permanent binding, will be offered for sale by the Missouri Botanical Garden. Prices upon application. MISSOURI BOTANICAL GARDEN Sr. Louis, Mo. PREFACE Formally begun in 1943, the Flora of Panama is complete with this issue. The broadside distributed by Woodson as an insert in the first fascicle of the Flora and reproduced opposite this page is a prospectus of what was planned. The completed flora follows Woodson’s guidelines remarkably closely, though significant changes and (we hope) improvements were made along the way by a succession of contributors and editors, many of whom were not even born when the project began. Over 100 scientists have contributed to its preparation, and publica- tion has extended over 38 years, from 1943 to the present. A short description of the Flora is provided below, and a history of collecting activities by Dwyer updates his earlier (1964) paper on the subject. A symposium celebrating completion of the Flora was held in Panama in April 1980, and the keynote address given at that event by Peter H. Raven outlines the problems facing future activities in Panama and other developing countries. The Flora of Panama is a beginning in the assessment of plant diversity in Panama; many additions to this body of knowledge have already appeared in the literature, and many more additions and changes will be generated by further field work and study.—W. G. D’Arcy THE FLORA OF PANAMA: HISTORICAL OUTLINE AND SELECTED BIBLIOGRAPHY W. G. D’Arcy! Formally begun in 1943, and prepared with the assistance of 103 contributors, the Flora of Panama includes coverage of over 6,200 species. Publication of the Flora in parts in the ANNALS OF THE MIssouRI BOTANICAL GARDEN ensured its wide distribution to botanical institutions around the world. It is the first attempt to survey all plants growing in Panama, and as such, it is the basis on which botanical studies in the future must rest. Although uneven in content and inad- equate in some parts, it is nevertheless a good starting point for assessing plant diversity in the isthmus: the great increase in known diversity since the appear- ance of early parts of the Flora makes clear the need for further effort. It is likely that a final inventory of vascular plant species occurring in Panama will include 9,000—10,000 species. Before commencement of the Flora, the Missouri Botanical Garden had for a number of years been interested in Panamanian plants beginning with the visit in 1922 by Jesse M. Greenman, then Curator of the Herbarium at the Garden. A tropical station was established in 1926 at Ancon which was at first concerned with providing living plants for show and study in the greenhouses in St. Louis, but which soon generated interest in the collection and study of herbarium spec- imens. Several scientific expeditions were sent to Panama in the 1930's and by the end of this decade several ‘‘Contributions toward a Flora of Panama’’ were published in the ANNALS OF THE MISSOURI BOTANICAL GARDEN. The history of this early period is related by Dwyer (1964), and the contents of the ‘‘Contribu- tions” and the early parts of the Flora are reviewed by Robyns (1965). Croat's (1978) section on History of Botanical Studies also gives a good resumé of plant collecting activities in much of the country. The Flora was initiated by Robert E. Woodson, Jr. (1904—1963) who edited the first parts with the assistance, until 1952, of Robert W. Schery (1917- ). From about the time Schery's departure until the late 1950's there was a hiatus in activity on the Flora. In 1958 the Flora resumed publication with several contributors besides Woodson, most notably Loran C. Nevling, who was on the staff of the the Missouri Botanical Garden for much of 1949, and slightly later (from 1960) James A. Duke, who was a staff botanist at the Garden. The Flora received its first funding from the National Science Foundation in 1957, and support has been continuous since then. In the early 1960's activity on the Flora grew, as André Robyns was hired to direct the project, and momentum was sufficient that with the death of Woodson in 1963, publication continued almost uninterrupted. Walter H. Lewis became Director of the Herbarium in 1964, and he improved the activity and professional level of standards of the Flora project. In 1968, Duncan M. Porter replaced Robyns as editor, and by the time of his departure in 1972, Porter was able to boast that all remaining parts of ! Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 67: v-viii. 1980. the Flora had specialists committed to preparing the treatments. During Porter`s time as editor of the Flora, contributions were made by Joan W. Nowicke who was hired to prepare treatments. Following Porter’s departure the Flora of Pan- ama project, which by now included an active field program, was directed by Thomas B. Croat, and the editorial aspects of the proJect were assigned to W. G. D'Arcy. In 1977, as he became engaged in a major study of the Araceae, Croat relinquished management of the project to D'Arcy who saw the Flora to com- pletion. In April 1980, with all treatments ready in at least manuscript form, a sym- posium was held in Panama City to celebrate completion of the Flora. This was supported by the Missouri Botanical Garden, the National Science Foundation, the University of Panama, the Smithsonian Tropical Research Institute, the Pan- amanian conservation commission (RENARE), the Joyce Foundation, and other organizations. About 350 participants gathered to hear a four day program of papers on Panama’s natural history. The concept of the Flora expanded somewhat over the years. Initially the number of species was thought to be small, and the published Flora was arranged so it could be extracted from the ANNALS for use as a separate manual. Thus early parts had *‘Flora’’ pagination as well as pagination of the ANNALS. In all cases citations should be made using the ANNALS pagination in the upper corners of the pages and not to the ‘‘Flora’’ pagination in lower center. The "Flora" pagination was discontinued after publication of the first parts of the Leguminosae in 1950. Other features of the Flora also expanded over time. From a policy of studying only material available in the Missouri Botanical Garden, it became normal procedure to borrow all Panamanian specimens from all institutions with significant holdings of Panamanian plants, the most noteworthy being: A, DUKE, F, GH, FSU, MO, OS, PMA, SCZ, and US, and especially important specimens were studied from many other institutions. In early parts of the Flora, cultivated and introduced plants were ignored, but later it became the rule to attempt to account for all species. Illustrations at first represented only a few of the families, but it later became the policy to include one illustration of each genus, and in large genera, one illustration for each 10 species. Original illustrations were pre- ferred; only in a few cases were older illustrations republished. When it became evident that much of the flora must, because of the poor state of knowledge about neotropical plants, be revisionary in scope, and that a second edition of the Flora would be unlikely for many years, the Flora began to include information on nomenclatural typification, and many of the treatments became semi-monograph- ic in scope. Over the years there was a large increase in the quantity and quality of col- lections available to the collaborators on the Flora. In the 1930’s most collections studied were those of Woodson and collectors directly concerned with production of the Flora, such as Paul C. Allen (1911-1963) Russell J. Seibert, and Julian A. Steyermark (1909- ) although in some cases the collections of Paul C. Standley and others were reéxamined. In the 1960's a series of expeditions was sent to Panama from St. Louis, and a number of other collectors became important contributors to the available stock of herbarium material; John D. Dwyer and James A. Duke were perhaps most noteworthy at this time. Neither of these was vi directly concerned with preparation of the Flora at the time when they did their most important collecting. Walter H. Lewis led several expeditions to Panama in the late 1960’s and early 1970’s, and his encouragement was of great importance in guiding the flora at that time. In about 1970, the assignment of Thomas Croat to prepare the Flora of Barro Colorado Island wrought great changes in the way that the Panamanian flora was to be perceived and studied. Croat became a resident in the isthmus for several years during which time he established a field station facility with housing, drying facilities and field vehicle capability. Croat and his successors at the station were able to make the first intensive collections of tropical wet forest regions. This ecological zone, which largely follows the Caribbean coast, is much richer in species than any other part of Panama, and it had been the least populated and least accessible. Families with important numbers of forest species were soon found to have twice as many or more species in Panama than had been supposed under even the most expansive earlier esti- mates. Families prepared before the early 1970's are found to have only a token representation of the flora actually present, and recently prepared families seri- ously understate the size of the actual flora. The intensive collecting program maintained since 1970, and especially the investigation of tropical wet areas, has revealed a flora much richer than previously suspected, and it has added excite- ment to the study of Panama's plant life. A result is that the Flora greatly un- derstates the actual situation: continuing study will supplement what has already been published, and the Flora together with new publications of present and future workers will give a reasonable picture of the plant diversity in the Isthmus of Panama. The Flora of Panama deals only with seed plants and includes just over 6,200 species. The pteridophytes are being treated separately by David B. Lellinger, Smithsonian Institution, together with the pteridophytes of Costa Rica and the Department of Chocó, Colombia. Lellinger (pers. comm.) estimates that he will record about 700 species from Panama. The bryophytes have not as yet been treated in floristic fashion, although active studies are under way by Prof. Noris Salazar and assocates at the University of Panama. Marshall R. Crosby, Missouri Botanical Garden, estimates (pers. comm.) that there are about 400 species of mosses in Panama. The total derived from the above estimates—7,300 species including seed plants, pteridophytes, and mosses—greatly underestimates the total flora, for the figure for seed plants omits many species. The total flora is more likely in excess of 9,000 species. THE FLORA OF PANAMA: SELECTED BIBLIOGRAPHY BENNETT, C. F. 1968. Human influences on the zoogeography of Panama. Ibero-Americana 51. 112 CroaT, T. B. 1971. Summ mit Garden, Canal Zone. Taxon 20: 769-772. ——. 1972. The Missouri Botanical Garden in Panama. Missouri Bot. Gard. E 60: 19-22. — — . 1978. Flora of Barro Colorado Island. 943 pp. Stanford University Pres M. R. CnosBv. 1974. Systematics program at the Missouri Botanical Garden. Taxon 23: 444—446. Duke, J. A. 1968. Darien Ethnobotanical Dictionary. Battelle Memorial Institute, ooo Ohio. Dwyer, < 1945. A list of localities botanized in Panama. Phytologia 16: 467—4 . Panama, plant collection, and the Missouri anea Garden. mn Missouri Bot. atm Qu 109-117. ——R 1967. A new herbarium in the Canal Zone. Taxon 16: 158-159 ——.T 1968. A list of localities botanized in Panama. Phytologia 16: 467—486. — —. 1973. Henri Pittier's botanical activity in Panama. Taxon 22: 557-576. ———. 1980. Rubiaceae in Flora of Panama. Ann. Missouri Bot. Gard. 67: 1-522. Vx seb L. R. 1970. Manual oe para 1000 especies arboreas en la Republica de ama. Panama City. Govt. of Pana Bupowski. 1959. Mapa s A "i Panamá (1:1,000,000). Turrialba, Costa Rica: Interamerican tues of Agricultural Science JOHNSTON, I. i . The botany of San José Island. Sargentia 8: 1-306 KENOYER, L. v: 2 C. STANDLEY. Meo Su pron to the flora of Barro Colorado Island, Panama. EM Field ose Mus . Hist., Bot. Ser. 4(6): 143-158. KNiGHT, D. H. 1970. A field guide to the ye of Barro Colorado Island, Laramie, Wyoming. Department of Botany, Univ. of Wyomi ing. Lewis, W. H. 1971. High floristic endemism in low cloud forests of Panama. Biotropica 3: 78-80. MEYER, C. W. 1969. The ecological geography of cloud forests in Panama. Amer. Mus. Novit. 2396: NowICKE, J. W. 1970. Type EM of the Panamanian collections of B. C. Seemann. Ann. Missouri Bot. Gard. 57: 352-358. RoBvNs, A. 1965. Index to the ^ ; Contributions Toward a Flora of Panama” and to the ‘Flora of Panama" through March 1965. Ann. Missouri Bot. Gard. 52: 234-24 ScHERY, R. W. 1945. A few facts concerning ds flora of nba Pp. 284-287 in F. Verdoorn, editor, ''Plants and Plant Science in Latin America." Chronica Botanica 5. Wa tham, Mass. SEEMANN, B. 1854. The Botany of the H.M.S. Herald (Flora s the Isthmus of Panama). 468 pp. STANDLEY, P. C. 1927. The flora of Barro Colorado Island. Smithsonian Misc. Collect. 78(8): 1928. Flora of the Panama Canal Zone. Contr. U.S. Natl. Herb. 27: 16. 1930. A Peri supplement to the flora of Barro Colorado Island, Panama. J. Arnold mn ll: 129. . . The 7A i Barro Colorado Island. Contr. Arnold Arboretum 5: 1-178. STERN, E a um Plant collecting in the land of Balboa. Gard. Jour. New York Bot. Gard. 44. 10: ; Tost, J. A., i 1971. Zonas de vida, una base ecológica para investigaciones silvicolas y inventar- iación forestal en la Republica de Panamá. Rome: Organización de las Naciones Unidas para Agricultura y Alimentación. 123 pp. with map. WiLBUR, R. L. & J. L. LUTEYN. 1978. SU MUR. in Flora of Panama. Ann. Missouri Bot. Gard. 65: 144 viii THE HISTORY OF PLANT COLLECTING IN PANAMA (1700-198 1)! JOHN D. DWYER? This talk is dedicated to the memory of Dr. Robert Woodson, Jr., late Curator of the Herbarium of the Missouri Botanical Garden and Professor of Botany at Washington University, who died in 1963. His name, together with that of Dr. Robert W. Schery, has been on the title page of every issue of the Flora of Panama in the ANNALS OF THE MISSOURI BOTANICAL GARDEN. While this paper is historical in character, a point dealing with the present and the future will be made: that Panama is and will be in need of more intensive collection. I estimate that about 750 botanists or naturalists have collected plants in Panama for permanent deposit in herbaria. This may seem to be an exaggeration, considering that my current unpublished list (see also Dwyer, 1964, 1968) includes the names of only 450 collectors. In defense of this estimate of 750 let me point out that new collectors can be found without much difficulty by examining her- barium specimens and through searching the literature. The list includes the names of all who have collected without regard to the quantity of collection numbers. The term ‘‘number’’ does not include duplicate material included under a specific number. Throughout the world there are about one half million numbers of Panamanian plants. Of these about a quarter of a million are deposited in the Missouri Bo- tanical Garden. In the last decade and a half five botanists from the Missouri Botanical Garden have collected approximately 50,000 numbers, these, for the most part, being gathered in non-team fashion (Table 1). As this is a historical treatment of plant collecting in the Republic of Panama, we start at the beginning when in 1700 James Wallace collected several herbarium specimens in the ill-fated Scotch Colony on the Atlantic Coast of the old Province of Darién. It is appropriate to divide the 280 years of plant collecting into 3 phases: 1. 1700-1914. 2. 1915-1957. 3. 1958-1981. The first phase runs from ¿he period of the Scotch Colony in Darién to the opening of the Panama Canal. The second period is marked initially by a lull period during and immediately after World War I, followed by some field work and considerable herbarium study, this to be again interrupted by a war, World War II. The third period, the golden period, extends from 1958, the year when William L. Stern and Kenton L. Chambers made an exciting collecting trip into Panama, up to the present time when herbarium material is being collected on a year-round basis. ! Paper presented at Symposium on the Botany and Natural History of Panama, Panama City, 14-17 April 1980. Supported by National Science Foundation grants DEB 77-04300 and INT 79-13043 to the Missouri Botanical Garden ? Missouri Botanical Garden, P. O. Box 299, St. Louis, Missouri 63166. ANN. Missouni Bor. GARD. 67: ix-xv. 1980. TABLE |. Collectors with largest numbers of collections from Panama. * Indicates individual was a staff member of, student at or professional collector for the Missouri Botanical Garden. *Croat, T. 23,000 *D'Arcy, W. 5.000 Standley, P. 7,500 Dressler, R. 4,000 *Gentry, A. 7,000 *Dwyer, J. 4,500 uke, J. 6,000 *Hammel, B. 4,500 *Mori, S. 6,000 *Lewis et al. 3,700 *Folsom, J. 5.000 * Allen, P. 3,600 *Nee, M. 5,000 Pittier, H. 3,600 Tyson, E. 5,000 *von Wedel, H. 3,000 A glance at Table 2 shows that in the first period (1700-1914) a shift from domination of plant collecting by Europeans to an eventual takeover by North Americans is obvious. Some collections made in this period may never have been cited as exsiccatae, e.g. those of Joseph Jussieu, Robert Millar, and B. Roezl. Particularly noteworthy are the collections of Berthold Seemann, who wrote the first flora of Panama (Seemann, 1854) as well as those of Henri Pittier, numbering over 1,850, drawn from one end of Panama to the other (Dwyer, 1973). Several Christian Brothers living in Panama City collected several hundred numbers be- ginning in 1912. Interestingly, almost all the Americans who collected in the first period were from the eastern United States. The second period (1915-1957) obviously overlaps somewhat with the first, making it the most difficult of the three to bring into proper perspective. Paul Standley's collection of 7,500 numbers in 5 months (end of 1923 and beginning of 1924, and one month in 1925) served as a basis for his Flora of the Canal Zone (Standley, 1928) and the Flora of Barro Colorado Island (Standley, 1927). He demonstrated his extraordinary ability in the field. His collections, mostly uni- cates, were confined to the Canal Zone and the vicinity of Panama City. As early as 1922 Jesse M. Greenman, Curator of the Herbarium at the Garden, made a few collections of phanerogams in Panama, and in 1923 the Missouri Botanical Garden was attracted to Panama by the living orchid collection of a retired postman, C. W. Powell, living in Balboa. To house this collection, re- ceived as a gift, the Garden maintained a tropical station at Ancon Hill, Canal Zone, this surviving until 1939. Beginning in 1934, up to the beginning of World War II, several botanists, functioning mostly in team fashion, collected throughout the isthmus, especially Paul Allen, Carrol Dodge, A. A. Hunter, Russell Seibert, Robert Schery, Julian Steyermark, and Robert Woodson, Jr. I estimate that their collections in Panama amount to fewer than 10,000 numbers. The figure would be considerably higher if the vast majority of the 5,000 specimens collected by Woodson, Allen, and Seibert in 1947 were not destroyed by fire. In this second period the greatest exhibition of collecting was by C. V. Piper of the U. S. National Herbarium who, in 1923, collected 1,000 numbers in 10 days! From 1948 until his death in 1963 Woodson believed that Panama had been adequately collected, and that the Flora of Panama could be written successfully by relying almost exclusively on the collections in the herbarium cases of the Garden. In light of the experience of the last two decades, this was a great mistake X TABLE 2. Early collectors of Panamanian plants. WALLACE, J. Scottish 1700 JUSSIEU, JOS. DE French 17?? MILLAR, R. English 1734 MEE, L. French 1789 HAENKE, T. Bohemian 1780-1791 BILLBERG, J. Swedish 1826 DAHLIN, E. Finnish 1826-1831 CUMING, H. English 1830-1831 HiNps, B. English 1837 ,G English 1837 etc SINCLAIR, A English 1837 etc , W. English SEEMANN, B. German 1846; 1847; 1848; 1849 WARSZEWIEZ, J. Polish 1848; 1850 B German 1848 FENDLER, A. American 1848-1849 Irish 1852 HALSTED, M American 1850-1854 KUNTZE, O. Germa DUCHASSAING, E. French 1849-1851 HAYES, S. American before 1863 ROEZL, B Czech HART, J. English 1885 CowELL, J. American 1905 WE, M. American 1909-1910 WILLIAMS, R. S American 1908 PITTIER, H. Swiss 1910-1911; 1914-1915 PowELL, C. American 1907-1927 OSTENFELD, C. Dane 1921-1922 Bro. GERVAIS Panamanian 1912 Bro. CELESTIN Panamanian 1912 STEVEN, : American 1924 MAcBRIDE, J American 1918 OSE, American M American 1911; 1923 HiTCHCOCK, A. A American KiLLIP, E. P. American 1917-1918; 1922 PIPER, C. V. American 1923 GOLDMAN, F. A. American 189?-19?? CHRISTOPHERSEN, E. Norwegian 197? in judgement, although the lack of money for field work and the pressure of his academic responsibilities have to be considered. As we shall see in the third and last phase, the numbers of novelties being described from Panama, especially in the last decade, e.g., in the Ericaceae, Araceae, Rubiaceae, strongly support such criticism (Table 3). In 1949 Ivan Johnston’s The Botany of San Jose Island was an outstanding contribution, based essentially on a study of his own collections and those of C. O. Erlanson and J. Harlow. In this period the collections of George P. Cooper and George Slater amounted to about 450 numbers from the little explored prov- ince of Bocas del Toro, the higher reaches of which remain little explored today. In 1958, in my opinion, the field trip of William Stern and Kenton Chambers, both then faculty members at the Yale School of Forestry, initiated the third stage of plant collecting in the isthmus, reópening the door of Panama as a land xi cpus indes in Flora of Panama (1970-1980) and the numbers of new or recently described speci Percentage New or Recently Year Number of Families Total Species Described Species 1970 5 9] l 1975 11 413 12.5 1976 9 207 10 1978 8 217 32 1980 l 415 23 of promise for the systematics of flowering plants and ferns (Standley & Stern, 1960). The following year, Stern, Chambers, John Ebinger, and I collected more than 1,200 numbers, principally in Darien. In 1960, Duke University, under the auspices of the U.S. Army Research Office, initiated a decade of collecting in Panama. In 1963 Dr. Edwin Tyson, then of Florida State University, shifted much of his research in zoology to the systematics of the angiosperms. His collections, amounting to 5,000 numbers, were housed in the U.S. Army Tropic Test Center, Canal Zone (Dwyer, 1967), which later became Summit Herbarium (Croat, 1971). Soon after Woodson died in 1963, Walter H. Lewis became Director of the Herbarium of the Missouri Botanical Garden. He immediately set to work getting rant support for botanical exploration in Panama. Table 4 shows that in 1966 and 1967 13 Garden staff members and students made collecting trips in Panama. At about this time Robert L. Dressler became resident botanist with the Smith- sonian Tropical Research Institute, at first living on Barro Colorado Island. His own collections and personal studies, particularly on orchids, coupled with his willingness to assist all of those botanizing in Panama, has been of inestimable value over the past 17 years. One of those profiting from the grant support generated by Walter Lewis was Thomas B. Croat. His accomplishments in collecting during the past 14 years in Panama defy description. Statistics indicate (Table 1) that he tripled the number of plants collected by Paul Standley. Croat’s total from the isthmus is more than 23,000 numbers. (See Croat, 1978.) A glance at Table 1, which shows those who collected more than 3,000 num- bers in Panama, is revealing. With the exception of Walter Lewis, this list scores TABLE 4. Collectors who accompanied Walter Lewis on field t rips to Puasa (1966-1969). * Indicates individual was staff member of or student at Missouri Botanical Garde Escobar, N. *Blackwell, W. *Hawker, J urch, D Little, C. O Correa A., M *Macbryde, B *Croat, T. *Oliver, R *Crosby, M *Robertson, K Dressler, R. *Robyns, A. Duke, J. *Verho *Dwyer, J. Zaborowski, A. *Elias, T. TABLE 5. Resident collectors at Summit Herbarium, Panama. Antonio, Thomas M. May 1979-July 1980 Croat, Thomas B. March 1970-August 1971 February 1976-September 1976 Folsom, James P. January 1977-January 1978 Gentry, Alwyn H. August 1971—October 1972 mmel, Barry E. January 197 ay 1 Kennedy, Helen September 1972-May 1973 Mori, Scott A September 1974—August 1975 Nee, Michael August 1973-June 1974 Sytsma, Kenneth J. August 1980- Witherspoon, John T. September 1975-November 1975 only those who collected alone. Thus some collectors on the list may also have assisted in team efforts. I have no statistics for the numbers collected by Sydney McDaniel, now in Panama, although they are several thousand. In 1969, in the Canal Zone, the Missouri Botanical Garden established a field station with a house trailer and a modest herbarium (Croat, 1972). This allows pre-doctoral or post-doctoral students to pursue their research in systematics, as well as to collect. The Garden's resident collectors in Panama, who were also curators of Summit Herbarium, are shown in Table 5. In 1973 the Missouri Botanical Garden in Panama and the Smithsonian Trop- ical Research Institute combined their herbarium resources in a restored portion of the former Tivoli Hotel in Balboa, Canal Zone (Croat & Crosby, 1974). at the Missouri Botanical Garden dominated plant collecting in Panama from the 1940's onward is obvious. However when one considers the extensive collecting done by Edwin Tyson in the 1960's, as well as by Sydney McDaniel and Kurt Blum, sent into Panama originally by Robert Godfrey of Florida State University, as well as the extensive collecting of James Duke over the 1960-1970 decade, under the aegis of the U.S. Army and Battelle Memorial Institute, as well as the eight botanical expeditions of Robert L. Wilbur of Duke University and his students: Frank Almeda, James Luteyn, Richard Weaver, Robin Foster and others, a more fair perspective of collecting in Panama emerges. Wilbur himself has collected some 1,500 numbers in Panama. Special tribute is due Novencido Escobar and Mireya D. Correa A. of Panama University for building up, over the past decade, the herbarium at Panama Uni- versity, now numbering over 16,000 sheets. TABLE 6. Collectors who made largest collections from provinces of Panama. Bocas Del Toro von Wedel, H. 3,000 Chiri Croat, 3,640 oclé Folsom, J 1,415 Colón Croat, T 1,052 Darién Duke, J 2,300 H r Folsom, J 120 Los Santos Lewis et al 458 Veraguas Croat, T 1,510 San B Duke, J 00 an 5 Canal Zone Croat, T. 10,737 xiii Today, in Central America, the most exciting country for plant collecting is Panama. Evidence of this is the 15 new species of Rubiaceae which are awaiting description at the Missouri Botanical Garden, these having been encountered since sending off my manuscript of the Rubiaceae of Panama with a total of 415 species at the end of 1979 (Dwyer, 1980). Table 3 shows the numbers of recently described species in parts of the Flora of Panama. There are certain areas of Panama, e.g. the Fortuna Dam area, where a single day of intensive collecting will usually yield one or more species of flowering plants new to science. Ob- viously collectors are penetrating such areas because they and others before them have searched them out, often under great hardship. Also, new roads have been opened in the republic in recent years and local landing strips and helicopter pads have been established in remote areas, often related to industrial or national projects. The collector is aware that pristine collecting grounds of today may fall prey to tomorrow’s ax of destruction. Deference is due the Government of Pan- ama for granting collecting permits to qualified collectors. Many institutions re- ciprocate by forwarding identified duplicate specimens to the Herbarium of the Universidad de Panama, the national repository. In conclusion, it is appropriate to pinpoint certain geographical areas of Pan- ama which are *'rich and recent" from the viewpoint of botanical collecting, at the same time acknowledging the pioneer efforts of the botanists. Such a listing is never complete. Only those localities botanized in realistic fashion for the first time between 1967 and the present are considered. Certain relatively new col- lecting areas, e.g. Cerro Jefe (Prov. Panama from 1965) and Cerro Pirré (Prov. Darién from 1960) have not been included as their ‘‘discovery’’ predated 1967. Table 6 indicates how the various provinces of Panama have benefited from the attentions of different collectors. 1967—James Duke and Joseph Kirkbride, Jr. botanized from Chiriquí Grande (Prov. Bocas del Toro) to Caldera (Prov. Chiriqui). 1967— Walter Lewis, Thomas Croat, and Jon Hawker made important col- lections at Río Concepción (Prov. Veraguas). In the same year Will Blackwell and Mireya Correa A. made important collections at Santa Catalina. 1971—Thomas Croat collected from Guasimo (Prov. Colón) to Miguel de la Borda (Prov. Colón), using a canoe. 1973—Thomas Croat, Ronald Liesner and Philip Busey botanized in the tropical wet forest on the Burica Peninsula at Bartolo Limite-Rabo de Puerco area (Prov. Chiriqui), utilizing a vehicle. Today the road is overgrown. 1978—Robert Dressler pioneered in collecting from La Pintada (Prov. Co- clé) to Coclecito (Prov. Coclé). 1975—Scott Mori was the first to collect at Cerro Colorado (Prov. Chiri- qui), close to the Continental Divide, at an elevation of 1,200—1,500 m. Cerro Colorado is the site of a copper mine, easily accessible by road. Mori, together with Alwyn Gentry, explored botanically the summit of Cerro Tacacaruna, elev. 1,975 m, on the Panama-Colom- bia border. 1976—Thomas Croat collected in the Azuero Peninsula walking for 6 hours to the headwaters of the Río Pedregal (Prov. Los Santos) from Job- ero in the same province, southwest of Tonosi. 1977—James Folsom began an era of rewarding collecting on the divide above El Cope (Prov. Coclé). 1978—Barry Hammel collected at two important sites: at Cerro Bruja (Prov. Colón) and the headwaters of the Río Indio and Río Bo- queron (Prov. Panama); from Gualaca (Prov. Chiriqui) to the For- tuna Dam site, proceeding to Cerro Hornito (Prov. Chiriqui). 1979—Barry Hammel collected at Cerro Pate Macho (Prov. Chiriquí, the Rio Culebra (Prov. Chiriqui), as well as along the Río Concepción (Prov. Veraguas, especially at a one-man operated gold mine). There is, of course, a limit to the number of species in Panama, as well as choice collecting sites. It is strange, however, that despite almost 300 years of collecting, the yield of novelties in our day is so great. Since the opening of the Panama Canal in 1914, more endemics have been described from the isthmus than between 1700 and 1913. The richness of the flora is a reflection of the soil and climate of this great tropical country. It is no less a tribute to the efforts of the botanists who have been so richly rewarded. LITERATURE CITED Croat, T. B. 1971. Summit Garden, Canal Zone. Taxon 20: 769-772. ——— . The Missouri Botanical Garden in Panama. Missouri Bot. Gard. "di 60: 19-22. — 1978. Flora of Barro Colorado Island. 943 pp. Stanford University Press. —— & M. R. CnosBv. 1974. Systematics program at the Missouri Botanical Gadea. Taxon 23: 444—446. Dwyer, J. D. 1964. s plant collection and the Missouri Botanical Garden. Ann. Missouri Bot. Gard. 51: m 67. A new he MN um in the Canal Zone. Taxon 16: 158-1 — V 1968. A lis of localities botanized in Panama. Phytologia 16: dde —. 1973. Henri Pittier’s botanical activity in Panama. Taxon 22: 557—576. 980. Rubiaceae ¿n Flora of Panama. Ann. Missouri Bot. cr i: 1-522. JOHNSTON, I. M. 1949. The botany of San José Island. Sargentia 8: SEEMANN, B. C. 1854. The Botany of the H.M.S. Herald (Flora of dun aie of Panama). 468 pp. London. STANDLEY, P. C. 1927. The flora of Barro Colorado Island. Smithsonian -= gn ii aia 78: 1-32. 1928. Flora of the Panama Canal Zone. Contr. U.S. Natl. Herb. 27: STERN, W. L. 1960. Plant collecting in the land of Balboa. Gard. Jour. SEN Taš Bot. Gard. 10: 44—48, 64. RESEARCH, TROPICAL ECOLOGY AND THE FUTURE OF PANAMA! PETER H. RAvEN? Me es muy grato estar presente esta noche para celebrar la feliz conclusión de la Flora de Panamá, una obra que ha ocupado la atención de científicos de varias instituciones por casi cuatro décadas. Iniciada por el Jardín Botánico de Missouri, la Flora de Panamá surgió de una larga y fructifera colaboración entre los científicos de la Repüblica de Panamá, de los Estados Unidos de Norteamér- ica, así como de otros paises, y esperamos sinceramente que esta obra servirá para estimular un creciente interés y comprensión de las plantas de este hermoso pais. La privilegiada posición geográfica de Panamá, uniendo América del Sur con América del Norte, y banada tanto por el Atlántico como por el Pacífico, ha producido tal profusión y variedad de plantas, que Panamá reüne por sí sola tantas especies de plantas que todo el continente de Europa, un área doscientos veces más grande. Las ponencias que hemos escuchado en este interesante simposio, tan hábil- mente organizado por la Doctora Mireya Correa de la Universidad de Panamá y por el Doctor William D'Arcy del Jardín Botánico de Missouri, dan fé de la gran riqueza biológica de Panamá. Estas ponencias servirán de base para un nuevo libro sobre la historia natural del país, que se editará para celebrar la finalización de la Flora, y con miras a contribuir al aprecio y al uso racional de los recursos naturales del país. Panamá está en condiciones especiales para hacer grandes aportes a la biología y agricultura tropical. Tiene una ubicación geográfica ünica e instituciones bien conocidas tanto nacional como internacionalmente. Por un lado, el futuro bi- enestar de sus ciudadanos dependerá en gran parte del uso inteligente de las informaciones disponibles acerca de los recursos naturales del país. Por otra parte, estas mismas instituciones, distinguidas tanto en ciencias básicas como aplicadas, pueden jugar un papel clave en el desarrollo de conocimientos para el beneficio de los seres humanos de toda la faja tropical del mundo. Unas cuantas estadisticas ayudarán a demostrar cuán importantes serán estos conocimientos. Hace un siglo, los bosques tropicales húmedos ocupaban una superficie mucho mayor que Estados Unidos, México, y América Central juntos. Desde entonces, más de la mitad de estos bosques han sido talados o alterados, y con el actual ritmo de conversión, los bosques estarán totalmente destruidos dentro de treintiün anos. Por supuesto que el ritmo de destrucción es irregular, pero se calcula que en la mayor parte de la zona tropical, no quedarán áreas significativas de bosques hümedos tropicales dentro de veinte anos. Para en- tonces, las dos terceras partes de la población mundial estará viviendo en zonas tropicales. Esta destrucción de bosques conducirá a una extinción masiva de animales y plantas, muchos de los cuales no pueden reproducirse fuera de los bosques primarios. ing address, Symposium on the Botany and Natural History of Panama, Panama, April 1980, supported by National Science Foundation grant INT 79-13043 to the Missouri Botanical Garden. 2 Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166. ANN. Missouri Bor. GARD. 67: xvi-xxii. 1980. Estas transformaciones de bosques, equivalentes a la desaparición, cada dos años, de un área de bosque natural del tamaño de toda la América Central, son la consecuencia del desarrollo de los recursos naturales para el aprovechamiento humano. Si fuese verdad que de estas transformaciones resultase un beneficio humano continuo, éstas serían bienvenidas; sin embargo, para una gran parte del trópico, sobre todo para las zonas actualmente ocupadas por los bosques hú- medos tropicales, éste no es el caso. Nuestros conocimientos actuales simple- mente no alcanzan sino para utilizar una porción minima de estos suelos para la producción agrícola a largo plazo, y de esta manera, hay que considerar a los bosques y suelos como recursos naturales NO renovables. Asi, nuestro gran desafío como científicos e investigadores en el trópico será el de aplicar nuestros conocimientos en el campo practico para disenar sistemas agrícolas y agrofores- tales a largo plazo, mientras luchamos por concentrar los sistemas más produc- tivos en las tierras mas idóneas. Si logramos acelerar el inventario de la flora y fauna tropical a la vez que adelantamos estudios intensivos de ecosistemas crí- ticos, podremos unir la información necesaria para obtener soluciones prácticas, y ofrecer alternativas a las presiones y necesidades urgentes e inaplazables de una población creciente. La tecnología de las zonas templadas, basada en milenios de experiencia de agricultura contínua, no se puede aplicar a la mayoría del trópico; poco se hace actualmente para proveer la información tan indispensable para las zonas tropi- cales. Mientras tanto, la transformación de tierras para usos a corto plazo las hace inútiles para el futuro. De esto comentaré mas adelante. Sin embargo, es precisamente en este area que Panama puede aportar una contribución insusti- tuible al bienestar humano y a su propia prosperidad. Resumiendo, tenemos por delante una tarea dificil: la de armonizar el desar- rollo económico y social de una población en aumento, con la preservación, aunque sea parcial, de un mundo biológico que todavia nos oculta sus mayores riquezas. Panama, where the former island continent of South America was joined to North America about 3.1 million years ago (Keigwin, 1978; Marshall et al., 1979), is biologically one of the most critical and fascinating areas of the world’s tropics. The plants and animals of South America, isolated for more than 100 million years, had an ultimate common origin with those of Africa. As South America converged on North America through the 65 million years of the Tertiary Period, its tropical biota, dispersed across the ever-narrower sea, invaded North America in greater and greater numbers. Panama, which has been occupied primarily by open savanna-type vegetation for much of the past three million years, provide the keystone linkage. For this reason, and because of its geographical situation, its 50,000 square kilometers are occupied by the richest and most diverse assem- blage of plants and animals of any area of comparable size. I have already mentioned that Panama has nearly as many kinds of flowering plants as all of Europe; it may also be pointed out that it has, in just over 52,000 km?, about a third as many species of flowering plants as all of tropical Africa (30,000 species), an area of 20 million square kilometers (Brenan, 1978)! Com- parable figures could be provided for most groups of animals that are well enough xvii known to allow such calculations. They illustrate clearly that even by tropical standards Panama is exceptionally rich biologically. It is in fact this rich diversity of the plants and animals of Panama that is so properly a source of national pride, and the reason we have all gathered here for this most enjoyable symposium. Adding to its special interest, Panama was the area where the last direct connection between the marine biota of the Atlantic and Pacific existed, and the area where this connection was finally severed three million years ago. As a result, Atlantic and Pacific marine plants and animals not only exist in close proximity, but they exist in such a way that three million years of divergence between the derivatives of some formerly continuous populations can be timed accurately—a natural evolutionary experiment without parallel. For these and many other reasons, biologists historically have found Panama an exciting and interesting place in which to carry out their studies. Institutions such as the Universidad de Panama, the Gorgas Memorial Laboratory, and the Smithsonian Tropical Research Institute (STRI) have made significant contribu- tions to our knowledge of the biota of the Republic and their significance for man. A continued dedication to the importance of broadening this knowledge and uti- lizing it wisely in development has been signaled in recent years by the estab- lishment of RENARE,’ by the biological and ecological studies carried out under the auspices of IRHE,* and by the special position accorded to STRI under the terms of the Canal Treaty. The Government of Panama has explicitly recognized the principles of con- servation and wise management of land as the basis for sustainable development by the formation of RENARE, in which conservation and development are linked for the benefit of all the people of the Republic. Among the more notable recent accomplishments have been the active efforts to protect the watershed of the Canal, including both the projected reforestation schemes and the establishment by President Royo of the adjacent 24,000-hectare Parque Nacional la Libertad; the establishment of Barro Colorado Island as a nature monument and its sched- uled enlargement in 1999 at the conclusion of the treaty period; the plans for national parks in Darién along the Colombian border, along the Costa Rican border, and surrounding Volcan Baru, the highest mountain in the Republic, in the western highlands; the consistent efforts by IRHE and other agencies to conduct meaningful biological and ecological inventories before carrying out large-scale public works; and the educational components included in many of these efforts. Notwithstanding the great biological interest in the natural resources of Pan- ama, the government’s clear realization that the prosperity of the people rests upon the wise utilization of these resources, and the current implementation of many outstanding efforts to realize the consequences of this relationship, very serious problems face the Republic. These problems are, for the most part, shared with the other countries of the tropics, where, as already pointed out, some two- thirds of the world’s population will be living by the end of the 20th century. 3 RENARE: Dirección Nacional de Recursos Naturales Renovables. * IRHE: Instituto de Kecursos Hidro-Eléctricos. xvii Although the populations of most tropical countries are growing very rapidly— many at a rate at which they will have doubled by early in the next century— population is by no means the only factor involved in the worldwide ecological crisis facing not only Panama but every other tropical country in the world. Panama itself provides a graphic illustration of this relationship. With just under two million people in some 50,000 square kilometers, Panama may be compared not only with the Netherlands, with some 14 million people inhabiting 41,000 square kilometers, but also with Java, which has some 90 million people living in an area only about two-and-a-half times the size of Panama (129,500 km2). The annual increase in the population of Java is nearly as great as the entire population of Panama! Granted that there are more extensive areas of relatively fertile soil in Java than in Panama, a comparison between the two regions, both located near the equator, illustrates graphically the way in which different patterns of land-use can affect population density. It also must be recognized that the relatively small population of Panama has already destroyed more than half of the original forest resources, and may, unless alternatives fostered by the Government prove effec- tive, make short work of the rest. In Java, the heavy utilization of naturally moist or Irrigated rice-lands located on relatively fertile soils is an important element in the nutrition of the people. In addition, the homestead gardens of the rural people in Java provide a rich source of diverse foods, many of them derived from the native forests. Protein is largely provided by fish, some cultured, chickens, goats, and, in non-Moslem areas, by swine. In contrast, land use in Panama, particularly in the long-occupied and once relatively fertile deciduous-forest lands of the Azuero Peninsula, as Stanley Heckadon has emphasized earlier in this symposium, is based on ‘‘roza’’ grain agriculture followed by cattle pasture. These uses are highly destructive of the fertility of the soil when the density of people practicing them becomes too great, and large numbers of santenos (from Los Santos Province, on the Azuero Peninsula) are now emigrating to other areas, with potentially serious conse- quences to which we shall turn our attention later. Panama is in the fortunate situation of being able to derive much of its income from world trade, but, despite this, the wise use of natural resources is still of basic importance for the country. There is much to be said for regional and local self-sufficiency in a world in which the current energy crisis is just the first symp- tom of worldwide shortage of raw materials of all kinds that fuels inflation every- where. No one can guarantee the continued operation of the world economic system under growing pressures of this magnitude, and if Panama is to feed, clothe, and shelter twice as many people early in the next century at least as well as is possible now, or actually to better their lot, the country will need to make extremely wise and forward-looking decisions about the proper utilization of its resources now. These decisions ideally should be made against a background of continued attention to slowing the rate of population growth as much as possible. This will be extremely difficult at first, with about 43 percent of the population under 15 years of age, but the individual prosperiiy of fewer people utilizing an identical set of resources in the future will obviously be greater than otherwise. For this reason, the continuing efforts of the Government to decelerate the rate of growth XIX of Panama’s population appear to be well justified, despite the apparent paradox of the relatively low population level as compared with those of other areas. The principal elements necessary for improving the lot of the Panamanian people are the same as those necessary throughout the tropics of the world. They are all related to the search for finding ways to implement agricultural, forestry, or other kinds of systems that will sustain human life on areas now occupied with different kinds of natural vegetation, on a diverse array of tropical soils. The most direct reason that the population density (40 people per km?) of Panama cannot, with the application of present technology, equal that of the Netherlands (340 per km?), where the per capita gross national product is nearly six times as high as it is in Panama, is that we simply do not know how to utilize most soils of Panama, or most other tropical soils, in as productive a way as is possible in many temporate regions. National schemes have not governed the patterns of land use in Latin America in the past, any more than they have governed the patterns of land use in other regions of the world. What is a problem, however, is that when forest is cleared in most temperate regions, or prairies are plowed, the resulting fields are pro- ductive and, with proper management, remain so indefinitely. The kinds of ag- riculture that function well in temperate regions, however, cannot be transferred wholesale to the tropics, and it is our collective tragedy that we are just starting to act upon the consequences of this fact. The current estimate of Brazilian officials is that no more than 0.3 percent of the Amazon forest can be put into sustainable agriculture, and although the soils of Panama are generally better, the sad truth is that no one really knows how to manage most of those now occupied by moist forest for sustained yield once they are cleared. There is virtually no tropical deciduous forest left in Panama today, and the agricultural practices which have ruined the fertility of this potentially more productive region are now being transferred to the humid lowland forests of the Darién and of Bocas del Toro on the western part of the Caribbean slope, where they will produce food for only a year or two, instead of a decade or two. There is urgency in attempting to restore the fertility and productivity of the ravaged Azuero Peninsula, but technology known at present will not allow the sustained cultivation of the Darién or the Caribbean forests. Where, then, are the hungry santenos to turn? The moist forests that still persist in the northwestern and eastern portions of the Republic are inhabited by the richest and most diverse array of plants and animals of the entire Central American region. Some of them might be useful directly as sources of human food, shelter, energy, medicine, or for other pur- poses, but the only hope of discovering the extent to which this may be true will be to survey these forests biologically as rapidly as possible and to concentrate in the course of these studies on groups of organisms of known or suspected economic importance. In addition, as much as possible should be learned about the functioning of the undisturbed forest that exists in these regions, because only such knowledge offers hope of utilizing them on a sustainable basis or modifying them into some other form from which useful products could be derived. Mean- while, because these forests cannot, by and large, be put into sustainable agri- cultural use with currently available technology, the forests should be conserved to the greatest extent possible while we are gaining knowledge, and the modifi- XX cations that are allowed meanwhile should be as limited as possible in order to help secure the future prosperity of the country. These deceptively rich and productive-looking forests are simply not a suitable locale for most kinds of de- velopment, and such development can, in most cases, lead only to their total depletion, often with severe economic loss to those responsible. For reasons Just outlined, both the citizens of Panama and those who live elsewhere have genuine reason to applaud the establishment of the Darién Fron- tier National Park and the strategic plans that are being developed for the proper management of this area under the guidance and supervision of RENARE. The cooperation between Panama and Colombia that has made possible the progress to date has also been evident in the development of the 200,000 hectare Parque de la Amistad on the Cordillera de Talamanca between Panama and Costa Rica. Of equal importance are the efforts to reforest large denuded sections of the Panama Canal, and for the management of the Volcán Barú park, including the integral management of the Rio Caldera watershed. Commendable as these schemes for preservation and restoration of natural resources are, however, the real problem facing Panama is how to guarantee people an adequate diet without wrecking the natural resources of the country completely—in other words, how to utilize the land on a sustainable basis, and to make the connotation of the word *'renovable" in RENARE meaningful. These are global themes, and they have been stated with exceptional clarity and force in the World Conservation Strategy, promulgated by the International Union for the Conservation of Nature and Natural Resources (IUCN) in early March of this year. Throughout the tropics of the world, both multinational corporations, which presumably have a choice, and the peasant poor, who for the most part do not, are rapidly destroying the productive ecological systems on which human liveli- hood depends, without the creation of viable alternatives. The solution, both for Panama and throughout the tropics, depends upon a reversal of the forces that are destroying these ecological systems. Some of the elements that might be involved include the following: Afforestation schemes for the protection of watersheds, such as those that are currently being conducted by RENARE in the watershed of the Canal and else- where, and by IRHE around the Bayano Dam, are of special importance. The establishment of forest plantations and of areas devoted to agro-forestry, coupled with soil conservation measures and the improvement of pastures on areas that can sustain cattle grazing, all have immediate desirable consequences, among them the provision of employment for rural populations. National policy can help to prevent the institutional squandering of the national patrimony of forest re- sources for short-term gain, but only the provision of employment and the ability to feed their children properly can divert the santenos from the moist forests of the northwest and east, where they can cut off the trees and grow a crop or two before the soils are depleted. Panama will be well served by learning as much as possible about its existing natural resources and by applying the information gained as directly as possible to their sustainable utilization. In this process, the traditional national institutions such as the universities have a vital role to play, and they should be funded in such a way that they can make the contribution of which they are capable, at the xxi same time that new institutions are established. Meanwhile, the land should be developed as wisely as possible, with highly productive modes of agriculture concentrated in areas, such as perhaps portions of Chiriquí Province, where they can be carried out on a sustainable basis. Large-scale irrigation schemes should be developed elsewhere in appropriate areas to further the concentration of pro- ductive agriculture. Much more attention needs to be paid in Panama and every- where else in the tropics to the improvement of the lot of rural agriculturists, and information needs to be brought as efficiently from other areas of the world with similar climates to be used locally. For example, Brosimum alicastrum is a tree crop well known in southern Mexico as a highly productive source of flour that can be grown in many forest soils. Might it not be just as suitable for cultivation in portions of Panama? There is a worldwide crisis that threatens all portions of the tropics. During the course of the next generation, a majority of all undisturbed tropical forest will be modified for human use, and hundreds of thousands of species of plants and animals will become extinct. Only the prompt accumulation of knowledge and its direct application to human welfare will help to ameliorate the situation; the traditional /aissez-faire outlook on the operation of multinational concerns and neglect of the welfare of rural farmers that characterize all too many tropical countries will only serve to make the situation worse. In this situation, it seems possible that Panama, a country blessed with abun- dant and diverse kinds of plants and animals, relatively fertile soils, a varied topography, the presence of several significant international institutions, the worldwide interest of ecologists and other biologists, a well conceived govern- mental program that links conservation and the wise use of natural resources with development according to ecologically sound principles, and an impressive array of resident scientists and of national institutions, will be able to make a significant contribution. If Panama can continue to expand and to accelerate its efforts at intelligent self-development, this small country, located at the crossroads of North and South America, of the tropical Atlantic and Pacific, and forming one of the most intense zones of contact between Hispanic and Anglo-American cul- ture, has the potentiality of contributing in a significant way to the destiny of other countries throughout the tropics of the world. We who have come here to celebrate the completion of the Flora of Panama pledge ourselves to help as much as we can in striving to attain that transcendently important goal. Acknowledgments—I am especially grateful to W. G. D'Arcy, Pedro Galindo, R. J. A. Goodland, M. Lloyd, W. Meijer, I. Rubinoff, and R. E. Silberglied for their informative discussion of many of the ideas presented in this paper. LITERATURE CITED BRENAN, J. P. M. 1978. Some aspects of the phytogeography of tropical Africa. Ann. Missouri Bot. 78. KEIGWIN, L. D., JR. 1978. Focene S of the Isthmus of Panama, based ^ DARAUS evidence from nearby Pacific Ocean and Caribbean Sea cores. Geology 6: 63( MARSHALL, L. G., R. F. BUTLER, R. E. pred G. H. Curtis, & R. H. TEDFORD. p Calibration of the great American interchange. Science 204: 272-279. xxii CONTRIBUTORS TO THE FLORA OF PANAMA The over one-hundred contributors to families or parts of families to the Flora of Panama are listed below, together with their institutional affiliation when the contribution was published. When multiple authors are listed, the institution is always that of the first: the institution of the other will be found in a listing at the second author's name. Some authors published from two institutions, and an asterisk (*) is used to indicate those families published while at the second ad- dress.—W. G. D'ARCY ALLEN, C. Lauraceae. Missouri d Garden. ALLEN, P. H. Orchidaceae (part). Missouri Botanical Garden. AMSHOFF, G. J. H. Myrtaceae. State University of Utrecht. AUSTIN, DA hd Convolvulaceae*; Polemoniaceae; Trigoniaceae. Missouri Bade cal Garden Florida pete pr in BAILEY, L. H. Palmae [Palmaceae]; Rosaceae: Rubus. Cornell University. BARKLEY, T. M. Compositae: Senecioneae. Kansas State University. BLACKWELL, WILL H., JR. Loganiaceae; Sapotaceae. —— & Dopsos. Anacardiaceae. Missouri Botanical Garden. BuRcH, DEREK & WEBSTER. Euphorbiaceae. Missouri Botanical Garden. BunGER, W. C. Moraceae: Sorocea. Missouri Botanical Garden. Busey, PH Compositae: Elephantopidinae. Calves ai of Florida Agricultural esearch Center, Ft. Lauderdale. CANNE, JUDITH Compositae: Galinsoginae. University of E Guelph, Ontario, Canada. CONSTANCE, L. & MATHIAS. Umbelliferae. University of California, Berkeley. CORREA, A., M.D. & TAYLOR, B. Droseraceae. Universidad de Panama. CROAT, THOMAS B. Gnetaceae; ia lees i Sapindaceae; issouri Botanical Garden. Staphylea & CUATRECASAS. l CUATRECASAS, José. Brunelliaceae: Sterculiaceae: Theobroma. Smithsonian Institution. & CROAT. Malpighiaceae. Danpy, J. Magnoliaceae; Winteraceae. British os (N.H.). ANN. Missouni Bor. GARD. 67: xxiii-xxviii. 1980. TABLE |. CONTINUED D'Ancv, W. G. Basellaceae; Bataceae; Callitrichaceae Missouri Botanical Garden. lina Scrophulariaceae; Solanaceae; Styracaceae; Symplocaceae; Theophrastacea & Dwyer. Leguminosae: Erythrina. & LACKEY. Leguminosae: Macroptilium, Phaseolus, Vigna. & POSTON. Leguminosae: Lablab. & TomB. Compositae: Lactuceae. & SCHANEN. Erythroxylaceae. DEN HARTOG, C. Hydrocharitaceae. Rijksherbarium, Leiden. DEROON, A. C. Marcgraviaceae. State University of Utrecht. DEWoLF, G. D., JR Moraceae: Ficus. University of Cambridge, Cambridge. DILLON, MICH Leguminosae: Acosmium, Aeschynomene, Field kanena of Natural History. Alysicarpus, Ateleia, I. alyx, Dussia, Myroxylon, Ormosia, Zorn DopsoN, CALA Anacardiaceae; Coriariaceae. Missouri Botanical Garden. & ROBYNS. Hippocrateaceae. Duke, J. A. Amaranthaceae: Berbaridaceae; eee Missouri Botanical Garden. Ceratophyllaceae; Chenopodiacea Monimiaceae; Myristicaceae; Nymphaeaceae: Polygonaceae; Ranunculacea UNN, DAviD Leguminosae: Lupinus. University of NS Columbia. s D. H; Acanthaceae. Grinnell College, Grinnell, lowa. Dwyer, D. Leguminoseae: Papilionoideae; Ochnaceae; Missouri Botanical Garden. ubiaceae —— & D'ARCY. Leguminosae: Erythrina. EDWIN, GABRI Field Kum of Natural History. —TT OU. Roosevelt University, Chicago. ELIAS, ae s S. Missiouri Bonai Garden The (om Arboretum, Millbrook, New York.* —— ———* & PORTER”. Aquifoliaceae. Celastraceae. Compositae: Vernoniinae*; Menyanthaceae; Cyrillaceae; Balsaminaceae; Vitaceae. Rutaceae*. XXIV TABLE 1. CONTINUED * & RoBYNS*. PLING, CARL WIC x of California, Los Angeles. EXELL, A. British or (N.H.). FANTZ, North (nen State University. FRIE . E. Universi of Uppsala. GARDNER, ROBERT C. Ohio State University. GENTRY, ALWYN H. Missouri Botanical Garden. poe H. A. w York Botanical Garden. GREGORY, D. P. Missouri Botanical Garden. HAYNES, ROBERT R. Ohio State University. & WENTZ. HERM , F. J. United Pis Department of Agriculture, Beltsville, Maryland. HERRERA-MACBRYDE, OLGA & LEwis. Missouri Botanical Garden Hou, DING & EDWIN Rijksherbarium, Leiden. HowARD, R. A. The Arnold os of Harvard Univers HUNTER, ind —— State College, Murray, ucky. JONK aR, Sie dim of Utrecht. KEIL, DAvID J. Ohio State University. KiLLIP, E. P. Siiithedaian Institution. LACKEY, JAMES A. & D'Arcy. Smithsonian Institution. Lewis, WA H. Missouri Acie Garden. & HERRERA-MACBRYDE. LourTEIG, ALI Laboratoire m | ON Paris. LUNDELL, CYRU Texas ene Institute, Renner. LUTEYN, JAM . & WILBUR. New York osa Garden. Gentianaceae*. Labiatae. Combretaceae. Leguminosae: Clitoria. Annonaceae. Compositae: Cardueae. Bignoniaceae; Buxaceae; Humiriaceae; Rafflesiaceae; Sabiaceae Melastomataceae. Rhizophoraceae. Orobanchaceae. Najadaceae; Potemogetonaceae. Leguminosae: Lonchocarpus. Polygalaceae. Celastraceae. Icacinaceae; Polygonaceae: Coccoloba. Dilleniaceae. Burmanniaceae. Compositae: Tageteae. Amaryllidaceae: Bomarea; Urticaceae. Leguminosae: Macroptilium, Phaseolus, Vigna. Dichapetalaceae. Polygalaceae. Oxalidaceae. Myrsinaceae. Ericaceae. TABLE 1. CONTINUED MaAcBryDE, BRU Geraniaceae. Missouri Botanical Garden. MANNING, W. E. Juglandaceae. Bucknell University. MATHIAS, M. E. & CONSTANCE. Umbelliferae. University of California, Los Angeles. MAXWELL, RICHA Leguminosae: Dioclea. Indiana University E Albany. McCLiNTOCK, ELIZABETH. Saxifragaceae. California A a. of Sciences. McLAUGHLIN, LEO & WINDLER. Leguminosae: Croatalaria. Towson State College, Towson, aryland. CVAUGH, R. Rosaceae. University of Michigan. MEYER, FREDRICK G. Valerianaceae. National pn a a Washington, D.C. MOoLDENKE, H. N. Avicenniaceae*; Eriocaulaceae; Verbenaceae*. New York Botanical Garden 303 Parkside Road, Plainfield, N J.* Morton, C. V. Dioscoreaceae; Smilacaceae. Smithsonian Institution. MULLER, C. H. Fagaceae. iussu. of California. Munz, P. A. Onagraceae. Rancho Santa Ana Botanic Garden. NEV , L. I., JR. Aizoaceae; Araliaceae; Balanophoraceae; Missouri Botanical Garden. Chloranthaceae; Corylaceae; Lacistem Lythraceae; Myricaceae; d Opiliaceae; Portulacaceae; Proteaceae; Ulmac NOWICKE, JOAN W. Apocynaceae; Boraginaceae; Rhamnaceae. Missouri vesti Garden. & EPLING. Labiatae. PFEIFER, H. W. Aristolochiaceae. Missouri Botanical Garden. PORTER, DUNCAN M. Burseraceae; Simarubaceae; Zygophyllaceae. Missouri Botanical Garden. —— —— & CUATRECASAS. Brunelliaceae. National Science Foundation. — ——— & ELIAS. Rutaceae. Virginia Polytechnic Institute, Blacksburg E TON, MURIEL E. Leguminosae: Cajanus, Flemingia, Rhynchosia. ward Ta mais aN Washington, D.C. —— & D'Arcy. Leguminosae: Lablab. PRANCE, GHIL N T. Caryocaraceae. New York Botanical Garden. RAEDER, KATHERINE. Phytolaccaceae. Missouri Botanical Garden. xxvi TABLE 1. CONTINUED RHopss, D. G. Missouri Hotaiieal Garden. RIZZINI, C. T. Jardim Botanico, Rio de Janeiro. ROBERTSON, KEN Missouri Bonnes Garden. Rosson, Norman K. B. British ue (N.H.). ROBYNS, AN Missouri Piel Garden. — & ELIAS. Jardin Botanique National de iion Miese ROLLIN aC, Gray a SCHANEN, NoEL & D’Arcy. Missouri Botanical Garden. SCHERY, ROBERT W. Missouri Bo ed Garden. & SCHUBERT. SCHUBER . G. The penne Arboretum. & SCHERY. SIMPSON, BERYL BRINTNALL. Smithsonian Institution. SMITH, C. EA Crops s Division, Agricultural Research Service, U.S.D.A. Beltsville, Maryland. SMITH, LYMAN B. dnm Institution. SPARRE, BEN Swedish Museum of Natural History, Stockholm, Sweden SPELLMAN, D L. Missouri a Garden. STAND , P. C. Fie eld «fiis of Natural History. STUESSY, Top F. Ohio State University, SVENSON E Eo urs Garden. SWAL , J. R. Suman, Institution. TAYLOR, PETER. Royal Botanic Gardens, Kew. TAYLOR B., A.S. & CORREA. Dd de Panamá. Menispermaceae. Loranthaceae. Clethraceae. Hypericaceae. Bixaceae; Bombacaceae; Cochlospermaceae; Flacourtiaceae; Malvaceae; Sterculiaceae; Theaceae; s Turneraceae; Violaceae: Vochysiac Gentianaceae. Cruciferae. Erythroxylaceae. Mi ena eee Mimosoideae; Pontederi op Leguminosae: Desmodium. Begoniaceae. Compositae: Mutiseae. Elaeocarpaceae; Meliaceae. Bromeliaceae. Tropaeolaceae. Asclepiadaceae. Araceae. Compositae: Ambrosiinae, Melampodiinae, iniinae Cyperaceae. Gramineae. Lentibulariaceae. Droseraceae. xxvii TABLE 1. CONTINUED Toms, A. SPENCER & D'ARCY. Kansas State University. VAN ROYEN, Rijksherbarium, Leiden. RHOEK-WILLIAMS, SU Missouri Boi eee VIVALDI, JOSE Universidad ra Puerto Rico. TER, GRA I oft California. Davis. WENTZ, W. ALAN & HAYNES. Ohio State University. WHITE, F. University of Oxford, Oxford, England WHITE, PETER a ke Field Laboratory, Great moky Mts. National Par Gatlinberg, Tennessee. WILBUR, TL, Duke Tine Durham, N.C. & LUTEYN. WILLIAM Harvard d WINDLER, DON . & MCLAUGHLIN. Towson State i EE Towson, Maryland. WoopsoN, R. E., JR. Missanri Botanical Garden. YUNKER, T. G. De Pauw University. WUNDERLIN, RIC University of Son Florida. Compositae: Lactuceae. Podostemonaceae. Plumbaginaceae; Primulaceae. Malpighiaceae: Malpighia. Euphorbiaceae. Najadaceae; Potemogetonaceae. Ebenaceae. Leguminosae: Barbiera, Cracca, Dalea, Diphysa, Gliricidia, Indigofera, Lennaea, Sesbania, Sophora, Stylosanthes, Tephrosia, Willardia. Campanulaceae. Ericaceae. Orchidaceae (part). Leguminosae: Crotalaria. Alismataceae none pany SRECRE: Batac cadaceae; r ua Haemo dora eae; Moringaceae; Musaceae; pn ctaginaceae; Papaveraceae; Passiflora en eiut Rapate eaceae; a Salicaceae; Taxaceae; Thyme Tovariaceae; T dites ae; idm ceae; Velloziaceae; Xyridaceae; Zingiberaceae. Piperaceae. Cucurbitaceae. xxviii INDEXES TO THE FAMILIES FLORA OF PANAMA The Flora was divided into numbered parts using Roman numerals. The sys- tematic treatment of the families is complete as Parts II-IX and indexes follow. Part I is not missing: it will be used to publish ‘‘general discussions of historical, geographical, physiographic, and floristic relations” and other information per- taining to the Flora of Panama, as outlined in Woodson’s broadside (see pg. ii). Early in the Flora program, families known or thought to occur in Panama were assigned a sequential series of numbers, ‘‘following generally the system of Engler and Prantl.’’ These are the Arabic numbers preceding each family name in the Systematic Index. A few families were found to occur in Panama as the project progressed, and they were assigned numbers and letters within the original se- quence. Other families thought to occur in Panama have not been found there, but notes to this effect were published in the Flora series for completeness. Family names in the Systematic Index are followed by the volume(s), pages, and year(s) of the ANNALS in which they appeared. Family names in the Alphabetical Index are followed by the volume number(s), pages, and year of the volume(s) of the ANNALS in which they appeared. Num- bers in parentheses are the Flora of Panama part numbers (Roman numerals) and systematic family numbers (Arabic numerals) assigned to each family. —M. R. Crosby ANN. Missouri Bor. GARD. 67: xxix-xxxiii. 1980. xxix SYSTEMATIC INDEX TO THE FAMILIES IN THE FLORA OF PANAMA Family names are followed by the volume number(s), pages, and year of the volume(s) in which they appeared. Part II 42. Ulmaceae 47: 105-113. 1960. Cycadaceae 30: 97-98. 1943. Taxaceae 30: 98-99, 1943. - t2 13 — > ° 3 5 = š 6 ° f [c A = £G x ° w > ` 943. , Potamogetonaceae 30: 99. 1943; 62: 1- iiid 49. Balanophoraceae 47: 303-308. 1960. : 1-5. luc IN p. 50. Aristolochiaceae 47: 309-323. 1960. $. Butomaceae 30: 103—104. 1943. 50A. Rafflesiaceae 60: 17-21. 1973. 5A. Hydrocharitaceae 60: 7-15. 1973. i. Polygonaceae 47: 323359, 1560, 6. Triuridaceae 30: 1 52. Chenopodiaceae 48: 1-6. 1961. 7. Gramineae 30: 104—280. 1943 53. Amaranthaceae 48: 6-50. 1961. 8 Cyperaceae 30: 281-325. 1943. 54. Nyctaginaceae 48: 51-65. 1961. 9. Palmae 30: 337-396. 1943. 55. Batidaceae 63: 385-388. 1976. 10 Cyclantbacese 30: 396-403. 1943. 56. Phytolaccaceae 48: 66—79. 1961. Il. Araceae 31: 1260. 1044. 57. Aizoaceae 48: 80-85. 1961. D* Lemnaceae vA 60-62. 1944. 58. Portulacaceae 48: 85-89. 1961. 13. Mayacaceae 31: 62-63. 1944 58A. Basellaceae 66: 109-115. 1979. 14. Xyridaceae 31: 63-64. 1944. 39. Caryophyllaceae 48: 90-106. 1961. I5. Eriocaulaceae 31: 65-71. 1944. 0D. Nymphaeaceae 43: 137-141. 1982. 16. Rapateaceae 31: 71-72. 1944 61. Ceratophyllaceae 67: 819-822. 1980 17. Bromeliaceae 31: 73-137. 1944. me aureus ^ l 18. Commelinaceae 31: 138-151. 1944. ER E cu Es m l I: 151-157. 1944. l =LA, Pr tiii 65. Magnoliaceae 49: 173-176, 1962. Part III 66. Winteraceae 49: 177-178. 1962. 67. Annonaceae 49: 179-213. 1962. + Hine ae " un 68. Myristicaceae 49: 214-225. 1962. : kusaq : 69. imi : 225-249, : 22. Smilacaceae 32: 6-11. 1945. Monimiaceae 49: 420-249; 1962 23. Haemodoraceae 32: 11-12. 1945. 24. Amaryllidaceae 32: 12-25. 1945. PARE Y 25. Velloziaceae 32: 25-26. 1945. 70. Lauraceae 35: 1948. 26. Dioscoreaceae 32: 26-33. 1945. 71. Hernandiaceae P p ies 1948. 27. Iridaceae 32: 34-42. 1945. 72. Papaveraceae 35: 71-75. 1948. 28. Burmanniaceae 32: 42—47. 1945. 73. S 35: 75-99. 1948; 66: 29. Musaceae 32: 48-57. 1945. 117- 30. Zingiberaceae 32: 57-73. 1945. 74. inns 35: 99-106. 1948. 31. Cannaceae 32: 74-80. 1945. 75. Resedaceae 37: 121. 1950. 32. Marantaceae 32: 81-104. 1945 76. Moringaceae 37: 121. 1950. 33. Orchidaceae 33: 1-140, 315-404. 1946; 76A. Droseraceae 63: 389-392. 1976. 36: 1-245. 1949. 77. Crassulaceae 37: 121-124. 1950. PART IY 80. Cunoniaceae 37: 145-147. 1950 34. Piperaceae 37: 1-120. 1950. 80A. Brunelliaceae 62: 11-14. 1975. 35. Chloranthaceae 47: 81-83. 1960. 81. Rosaceae 37: 147—178. 1950. 36. Lacistemataceae 47: 84—87. 1960. 82. Connaraceae 37: 178—183. 1950. 37. Salicaceae 47: 87. 1960; 65: 1—4. 1978. 83. Leguminosae subfam. Mimosoideae 38. Myricaceae 47: 88-89. 1960. 37: 184—314. 1950. 39. Juglandaceae 47: 90—92. 1960. subfam. Caesalpinoidea ae 38: 1-94. 1951. 40. Corylaceae 47: 93-94. 1960. subfum, Papilio pies 52: 1-54. 1965; 41. Fagaceae 47: 95-104. 1960. 67: 523-818. . Humiriaceae 62: 35-4 PART VI Oxalidaceae 67: 823-850. 1980. Geraniaceae 54: 201—205. 1967. Tropaeolaceae 62: 15-20. 1975. Erythroxylaceae 62: 21-33. 1975. 4. 1975. Malpighiaceae 67: 851-946. 1980. Trigoniaceae 54: 207-210. 1967. Euphorbiaceae 2p :21 1-350, 1967. Coriariaceae 54; T 13-15. 1967. nacardiaceae 54: 351—379. 1967. . Cyrillaceae 54: 17-19. 1967. Aquifoliaceae 54: 381- 387. 1967. Celastraceae 62: 45-56. 1975. R 55: 81-92. 1968. aeocarpaceae 52: 487—495. 1965. nee 51: 1-35. 1964. Malvaceae 52: 497—578. 1965. Bombacaceae 51: 37-68. 1964. Sterculiaceae 51: 69-107. 1964. Dilleniaceae 52: “sya 1965. Ochnaceae 54: 25-40. 1967. dias caraceae Aao e —546. 1976. e 57: 29-50. 1970. Viola acea ines 54: 85-94. 1967. Part VII Passifloraceae 45: 1—22. 1958. Caricaceae 45: 22-31. Loasaceae 45: 32-40. 1958. Begoniaceae 45: 41-67. 1958. XXXI Cactaceae 45: 68-91. 1958. Thymelaeaceae 45: 93-97. 1958. Lythraceae 45: 97-115. 1958. Lecythidaceae 45: 115-136. 1958. Rhizophoraceae 45: 136-142. 1958. Combretaceae 45: 143-164. 1958. Haloragaceae 46: 221-223. 1959. Araliaceae 46: 223-242. 1959. Umbelliferae 46: 242—254. 1959. Cornaceae 46; 254—256. 1959. PART VIII Clethraceae 54: 389-392. 1967. Pyrolaceae—included in Ericaceae. Ericaceae 65: 27-154. 1978. Myrsinaceae 58: 285-353. 1971. Theophrastaceae 67: re das 1980. Primulaceae 57: 51—54. Plumbaginaceae 57: 55- A 1970. Sapotaceae 55: 145—169. 1968. Ebenaceae 65: 145-154. 1978. Styracaceae 66: 165-171. 1979. Symplocaceae 63: 547—552. 1976. Oleaceae 63: 553-564. 1976. Loganiaceae 54: 393—413. 1967. Gentianaceae 62: 61-101. 1975. Asclepiadaceae 62: 103— 156. 1975. PART IX onvolvulaceae 62: 157—224. 1975. Polemoniaceae 58: 355-361. 1971. Hydrophyllaceae 54: 415-418. 1967. : 783- 990. 197 9. Lentibuleriaceae 63: 565-580. 1976. Acanthaceae 65: 155—283. 1978. Plantaginaceae 58: 363-369. 1971. Rubiaceae 67: 1—522. 1980. Caprifoliaceae 60: 155-167. 1973. Valerianaceae 63: 581—592. 1976. Cucurbitaceae 65: 285—366. 1978. Campanulaceae 63: 593-655. 1976. Compositae 62: 835-1322. 1975. ALPHABETICAL INDEX TO THE FAMILIES IN THE FLORA OF PANAMA Family names are followed by the volume number(s), pages, and year of the volume(s) in which they appeared. Numbers in parentheses indicate the Part of the Flora (Roman) and systematic family (Arabic) numbers. Acantnaceae 65: 155-283. 1978. (IX: 177) Aizoaceae 48: 80-85. 1961. (IV: 57) Alismataceae 30: 100—103. 1943. (II: 4) Amaranthaceae 48: 6-50. 1961. (IV: 53) Amaryllidaceae 32: 12-25. 1945. (III: 24) Anacardiaceae 54: 351—379. 1967. (VI: 101) Apocynaceae 57: 59-130. 1970. (VIII: 162) Aquifoliaceae 54: 381—387. 1967. (VI: 102) Araceae 31: 1-60. 1944. (II: 11 Araliaceae 46: 223-242. 1959. (VII: 144) Aristolochiaceae 47: 309-323. 1960. (IV: 50) Asclepiadaceae 62: 103-156. 1975. (VIII: 163) Avicenniaceae 60: 149-154. 1973. (IX: 168A) Balanophoraceae 47: 303-308. 1960. (IV: 49) Balsaminaceae 54: 21-24. 1967. (VI: 110) Basellaceae 66: 109-115. 1979. (IV: 58A) Batidaceae 63: 385-388. 1976. (IV: 55) Begoniaceae 45: 41-67. 1958. (VII: 133) Bromeliaceae 31: 3- 137. 1944. d: 17) esp due 62: Hips. 1975. (V: A Buxaceae 65: 5-8. 1978. (VI: 99) t e 45: 68-91. 1958. (VII: 134) Callitrichaceae 67: 947-949. (VI Capparidaceae 35: 75- 99, 1948. (V: 73) °. subfam. Tovarioideae 66: 117— 3 . (IV: 73 Cignitoliacese 60: 155- 167. 1973. e 180) Caricaceae 45: 22-31. 1958. (VII: Caryocaraceae 63: 541—546. 1976. Ar 120) Caryophyllaceae 48: 90-106. 1961. (IV: 59) Celastraceae 62: 45-56. 1975. (VI: 103) Ceratophyllaceae 67: 819-822. (IV: 61) Chenopodiaceae 48: 1-6. 1961. (IV: 52) Chloranthaceae 47: oe 1960. (IV: 35) Cistaceae 67: 1045. (VI: 12 Clethraceae 54: 389—392. dr (VIII: 147) : 64. 1967. (VI: 126) Combretaceae 45: 143- 164. 1958. (VII: 139) Commelinaceae 31: 138-151. 1944. (II: 18) Compositae 62: 835-1322. 1975. (IX: 184) Connaraceae 37: 178-183. 1950. (V: 82) Convolvulaceae 62: 157-224. 1975. (IX: 164) Coriariaceae 54: 13-15. 1967. (VI: 100) Cornaceae 46: 254—256. 1959. (VII: 146) Corylaceae 47: 93-94. 1960. (IV: 40) Crassulaceae 37: 121-124. 1950. (V: 77) Cruciferae 35: 99-106. 1948. (V: 74) ucurbitaceae 65: 285-366. 1978. (IX: 182) Cyclanthaceae 30: 396—403. 1943. (II: 10) Cyperaceae 30: 281—325. 1943. (II: 8) Cyrillaceae 54: 17-19. 1967. (VI: 101A) Dichapetalaceae 54: 9-12. 1967. (VI: 96A) Dilleniaceae 52: 579—598. 1965. (VI: 118) Dioscoreaceae 32: 26-33. 1945. (III: 26) Ebenaceae 65: 145—154. 1978. (VIII: 155) Elaeocarpaceae 52: 487—495. 1965. (VI: 113) Ericaceae 65: 27-154. 1978. (VIII: 149) Eriocaulaceae 31: 65-71. 1944. (II: 15) Erythroxylaceae 62: 21-33. 1975. (VI: 87) Euphorbiaceae 54: 211—350. 1967. (VI: 97) Fagaceae 47: 95-104. 1960. (IV: 41) Flacourtiaceae 55: 93-144. 1968. (VI: 128) Gentianaceae 62: 61—101. 1975. (VIII: 160) Sleruniaesne 54: 201- 205. 1967. PA 85) ae 57: 1-4. 1970. (II: 2A) aemodoraceae 32: 11- 42. 1945. (HII: 23) Haloragaceae 46: 221—223. 1959. (VII: 3) Hernandiaceae 35: 68-71. 1948. (V: 71 Hippocastanaceae 62: 57-60. 1975. (VI: 107) Hippocrateaceae 52: 81-98. 1965. (VI: 104) Humiriaceae 62: 35-44. 1975. (VI: 87A) Hydrocharitaceae 60: 7-15. 1973. (II: SA) Hydrophyllaceae 54: 415-418. 1967. (IX: 166) Hypericaceae 65: 9-26. 1978. (VI: 123A) Icacinaceae 63: 399—418. 1976. (VI: 106) Iridaceae 32: 34—42. 1945. (III: 27) Juglandaceae 47: 90-92. 1960. (IV: 39) Juncaceae 32: 1-2. 1945. (III: 20) Labiatae 56: 71—111. 1969. (IX: 169) Lacistemataceae 47: 84—87. 1960. (IV: 36) Lauraceae 35: 1-68. 1948. (V: 70) Lecythidaceae 45: ag 136. 1958. (VII: 137) Leguminosae (V: 83) subfam. Mimosoideae 37: 184—314. 1950. subfam. Caesalpinoideae 38: 1-94. 1951. subfam. Papilionoideae 52: 1—54. 1965; 67: 523-818. 1980. Lemnaceae 31: 60—62. 1944. (II: 12) Lentibulariaceae 63: 565—580. 1976. (IX: 176) - ° Loganiaceae 54: 393-413. 1967. (VIII: 159) Loranthaceae 47: 263—390. 1960. (IV: 46) XXXII Lythraceae 45: 97-115. 1958. (VII: 136) Marcgraviaceae 57: 29-50. 1970. (VI: 121) Mayacaceae 31: 62-63. 1944. (II: 13) Melastomataceae 45: 203-304. 1958. (VII: 141 Meliaceae 52: 55-79. 1965. (VI: 92) Menispermaceae 49: 157-172. 1962. (IV: 64) Menyanthaceae 56: 29-32. 1969. (VIII: 161) Monimiaceae 49: 225-249. 1962. (IV: 69) Moraceae 47: 114-178. 1960. (IV: 43) Najadaceae 60: 1-5. 1973. (II Oleaceae 63: 553-564. 1976. (VIII: 158) Onagraceae 46: 195—221. 1959. (VII: 142) Opiliaceae 47: 291-292. 1960. (IV: 47) Orchidaceae 33: 1-140, 315—404. 1946; 36: 1— 245. 1949. (III: 33) Orobanchaceae 57: 131-134. 1970. (IX: 174) Oxalidaceae 67: 823-850. (VI: 84) Palmae 30: 337—396. 1943. (II: 9) Papaveraceae 35: 71-75. 1948. (V: 72) Passifloraceae 45: 1-22. 1958. (VII: 130) Pontederiaceae 31: 151- 157. 1944. (II: 19) Portulacaceae 48: 85-89. 1961. (IV: 58) Potamogetonaceae 30: 99. 1943; 62: 1-10. 1975. Primulaceae 57: 51—54. 1970. (VIII: 152) Proteaceae 47: 199-205. 1960. (IV: 45) Pyrolaceae—included in Ericaceae. (VIII: 148 Quiinaceae 67: 965—968. (VI: 121A) Pili parq 60: 17-21. 1973. (IV: 50A) Ranunculaceae 49: 144—153. 1962. (IV: 62) Rapateaceae 31: 71—72. 1944. (II: 16) Resedaceae 37: 121. 1950. (V: 75 Rhamnaceae 58: 267—283. 1971. (VI: 111) Rhizophoraceae 45: 136-142. oi (VII: 138) Rosaceae 37: 147-178. 1950. (V: 81) Rubiaceae 67: 1—522. 1980. (IX: 179) Rutaceae 66: 123-164. 1979. MA 89) Sabiaceae 67: 949-964. (VI: 109) Salicaceae 47: 87. 1960; 65: ru 1978. (IV: ) Sapindaceae 63: 419-540. 1976. (VI: 108) Sapotaceae 55: 145-169. 1968. (VIII: 154) Saxifragaceae 37: 137-145. 1950. (V: 79 Scrophulariaceae 66: 173-274. 1979. (IX: 171) Simaroubaceae 60: 23-39. 1973. (VI: 90 Smilacaceae 32: 6-11. 1945. (III: 22) Solanaceae 60: 573—780. 1973. (IX: 170) Staphyleaceae 63: 393-397. 1976. (VI: 105) Sterculiaceae 51: 69-107. 1964. (VI: 117) Styracaceae 66: 165-171. 1979. (VIII: 156) Symplocaceae 63: 547—552. 1976. (VIII: 157) Taxaceae 30: 98-99, 1943. (II: 2 Theaceae 54: 41—56. 1967. (VI: 122) Theophrastaceae 67: 1045-1056. (VIII: 151) Thymelaeaceae 45: 93-97. 1958. (VII: 135) Tiliaceae 51: 1-35. 1964. (VI: 114) Tropaeolaceae 62: 15- 20. 1975. (VI: 86) Turneraceae 54: 85-94. E. (VI: 129) Zingiberaceae 32: 57-73. 1945. (III: 30 Zygophyllaceae 56: 1-7. 1969. (VI: 88) xxxiii VOLUME 67 1980 ANNALS M THE MISSOURI BOTANICAL GARDEN Published by the Mis i Bot cal Gar den, St. Louis, Missouri 63110. © Missouri Bot otanica al Garden 981 CONTENTS CROAT, THOMAS B. (see Cuatrecasas, José & Thomas B. Croat)... CROsBY,M.R. Flora of Panama: Introduction and Indexes: Indexes to the families in the Flora of Panama ... CUATRECASAS, JOSE & THOMAS B. Croat. Flora of Panama, Part VI: Family 93. Malpighiaceae ecce sss D'Ancv, W. G. Abrus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ... D'Arcy, W. G. Calopogonium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .. __ D'Ancv, W. G. Canavalia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .. D’Arcy, W. G. Centrosema. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .. D'Ancv, W. G. Cologania. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... D'Arcy, W. G. Cymbosema. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) — . D'Ancv, W. G. Galactia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... D'Ancv, W. G. Mucuna. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) _ D’Arcy, W. G. Pachyrhizus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ... D'Arcy, W. G. Pueraria. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .... D'Arcy, W. G. Stizolobium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .... D’Arcy, W. G. Teramnus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) .... D’Arcy, W. G. Trifolium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) . D’Arcy, W. G. Ulex. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) D’Arcy, W. G. Flora of Panama. Introduction and Indexes: Preface _ D’Arcy, W. G. Flora of Panama. Introduction and Indexes: The Flora of Panama: historical outline and selected bibliography ------------------ D'ARCY, W. G. Flora of Panama. Introduction and Indexes: Contributors to the Flora of Panama ` XXIII D'ARcY, W. G. Flora of Panama, Part IV: Family 61. Ceratophylla- ceae D’Arcy, W. G. Flora of Panama, Part VI: Family 98, Callitrichaceae |... D'Arcy, W. G. Flora of Panama, Part VI: Family 121A, Quiinaceae ___. D'Arcy, W. G. Flora of Panama, Part VI: Family 123. Guttiferae _ D'Arcy, W. G. Flora of Panama, Part VI: Family 124, Cistaceae _ D’Arcy, W. G. Flora of Panama, Part VIII: Family 151, Theophrasta- ceae D’Arcy, W. G. Flora of Panama, Part IX: Family 173, Pedaliaceae _ D’Arcy, W. G. & M. E. Poston. Lablab. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) DILLON, M. O. Acosmium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... DiLLOoN, M. O. Aeschynomene. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ___- DILLON, M. O. Alysicarpus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) _.. DILLON, M. O. Ateleia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) _ _ DILLON, M.O. Chaetocalyx. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) DiLLON, M. O. Dussia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... DiLLON, M. O. Myroxylon. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion)... DILLON, M. O. Ormosia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... DILLON, M. O. Sophora. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... DILLON, M. O. Stylosanthes. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) II DILLon, M. O. Zornia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ...... Dunn, D. B. Lupinus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ...... DwYER, JOHN D. Flora of Panama, Part IX: Family 179, Rubiaceae ...... DwYER, JOHN D. Flora of Panama. Introduction and Indexes: The history of plant collecting in Panama (1700-1981) 819 947 965 969 1045 1047 1057 Dwyer, JoHN D. & COLLABORATORS. Flora of Panama, Part V: Fasci- cle 5: Family 83, Leguminosae subfamily Papilionoideae (conclu- SION): Less —Á Dwyer, J. D. & W. G. D'ARCY. Erythrina. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) _ FANTZ, P. R. Clitoria. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ------ GENTRY, ALWYN H. Flora of Panama, Part VI: Family 109, Sabiaceae __ LouRrE!IG, ALICIA. Flora of Panama, Part IV: Family 84, Oxalidaceae .... Lackey, J. A. & W. G. D'Arcy. Macroptilium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ---------------- Lackey, J. A. & W. G. D'Arcy. Phaseolus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ... Lackey, J. A. & W. G. D'Arcy. Vigna. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ... McDANIEL, S. (see McDearman, W. & S. McDaniel) McDEARMAN, W. & S. MCDANIEL. Marila. In D'Arcy, W. G. Flora of Panama, Part VI: Family 123, Guttiferae MAXWELL, R. H. Dioclea. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ------ Poston, M. E. Cajanus. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ...... Poston, M. E. Eriosema. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ..... Poston, M. E. Flemingia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ------ Poston, M. E. Rhynchosia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) ` RAVEN, PETERH. Flora of Panama. Introduction and Indexes: Research, tropical ecology and the future of Panama SCHUBERT, B.G. Desmodium. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) _..... VIVALDI, J. L. Malpighia. In Cuatrecasas, J. & T. B. Croat, Flora of Panama, Part VI: Family 93. Malpighiaceae WHITE, P. S. Barbieria. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion). 523 686 WHITE, P. S. Cracca. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion... WuiteE, P. S. Dalea. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion)... WHITE, P.S. Diphysa. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) WHITE, P.S. Gliricidia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) WHITE, P.S. Indigofera. In Dwyer, J. D. Flora of Panama, Part V: Fam- ily 83, Leguminosae subfamily Papilionoideae (conclusion). White, P. S. Lenna. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) WHITE, P.S. Sesbania. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion). WHITE, P.S. Tephrosia. In Dwyer, J. D. Flora of Panama, Part V: Fam- ily 83, Leguminosae subfamily Papilionoideae (conclusion) | . .... WHITE, P. S. Willardia. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) WINDLER, D. R. & L. MCLAUGHLIN. Crotalaria. In Dwyer, J. D. Flora of Panama, Part V: Family 83, Leguminosae subfamily Papilionoideae (conclusion) 599