ANNALS ' OF THE MISSOURI BOTANICAL GARDEN ERRATA Page 31, line 2—1939 should read 1839; line 3—1848 should read 1844. Page 48, following line 18 — Insert Piper aequale var. latirifolmm CDC. in Candollea 1:233. 1923, name only. Page 106, line 19 — ^Syst Veg. 4. Cur. Post. 21 should read: Sysc. Veg. 4. Cur, Post. 20. Page 108, line 24 — 1788 should read 1790. Page 110, line 14 — P Williamsii should read P. flavispica. Page 116, line 7 of key — P. Williamsii should read P. flavispica. Page 118, Species 69 should read as follows: 69. Peperomia flavispica Trel. in Contr. U. S. Nat. Herb. 26:48. 1927. Peperomia Williamsii Trel. loc. cit. 1927, not CDC. Page 404— The heading LOGANICACEAE should read LOGANIACEAE. Page 418, line 23 — uniia should read urtitus. Page 439, line 10, delete '^English." Page 453, line 2 of caption — fungi should read sori. Page 457, line 4 — ^Transpose sentence beginning "Although" to last line on page 454. Page 457, paragraph 5, line 6 — fig, 6 should read fig. 7. f\H C -- Annals o fth e Missouri Botanical Garden Volume XXXVII 1950 With 14 plates, and 124 figures Published quarterly at Galesburg. Illinois, by the Board of Trustees of the Missouri Botanical Garden. St. Loms, Mo. Entered as second-class matter at the post-office at Galesburg, Illinois. Hpder th? Act of March 3, 1879. GARDEN LIBBAR Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Henry Shaw School of Botany of Washington University in aflfiliation with the Missouri Botanical Garden. Information The during the calendar year: February, May, September, and November. Four numbers constitute a volume. Subscription Price -$10.00 per volume Single Numbers 2.50 each Content Company. W. Wilson STAFF OF THE MISSOURI BOTANICAL GARDEN Director George T. Moore Hermann von Schrenk, Pathologist Jesse M. Greenman, Curator Emeritus of the Herbarium Carroll W. Dodge, Mycologist Edgar Anderson, Geneticist Robert E. Woodson, Jr. Curator of the Herbarium Henry N. Andrews, Paleobotanist Robert W. ScHERr, Research Associate GUSTAV A. L. Mehlquist^ Res.c.irch Horticulturist RoLLA M. Tryon, Assistant Curator of the Herbarium George B. Van Sciiaack, Honorary Curator of Gr^i-'^ycs Julian A. Steyermark, Honorary Research Associate Nell C. Horner, Librarian and Editor of Publications Gerald Ulrici, Business Manager BOARD OF TRUSTEES I THE MI 'President Richard J. Lockwood Vice-President Daniel K. Catlix Second Vice-President Eugene Pettus L. Ray Carter Dudley French Henry Hitchcock John S. Lehmann George T. Moore A. Vessel Shapleigh Ethan A. H. Shepley EX-OFFICIO MEMBERS Arthur H. Compton, Chancellor of Washington University Joseph M. Darst, Mayor of the City of St. Louis R. Harris Cobb, President of the Academy of Science of St. Louis William Scarlett, Bishop of the Diocese of Missouri Will L. Schvehr President of the Board of Education of St. Louis Gerald Ulrici, Secretary TABLE OF CONTENTS PAGE Flora of Panama. Part IV, Fascicle 1 (Piperaceae) Robert E. Woodson, Jr., Robert W. Schery, and collaborators 1-120 Flora of Panama. Part V, Fascicle 2 (Resedaceae to Leguminosae, in part) Robert E. Woodson, Jr., Robert W. Schery, and collaborators 121-314 The American Species of Triumfetta L .-Ko Ko Lay 315-395 Miscellanea Taxonomica. I -^ ~ Some American Carboniferous Fern Fructifications Sergius H. Mamay 409-476 The American Species of Sarcostemma R. Br. (Asclepiadaceae) .._ . Richard W. Holm 477-560 The Grain Amaranths: A Survey of Their History and Classification Jonathan D. Sauer 561-632 General Index to Volume XXXVII 633-637 Volume XXXVII Number 1 Annal of the Missouri Botanical !^-^«fi ^ FEBRUARY, 1950 Flora of Panama. Part IV. Fascicle 1 (Piperaceae) . Robert E. Woodson, Jr., Robert W- Schery and Collaborators 1-120 PUBLISHED QUARTERLY AT GALESBURG, ILLINOIS^ B7 THE BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN. ST. LOUIS, MISSOURL Entered as second-class matter at the post*office at Galesburgy lUinoiSt under the Act of March 3* 1879. Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the* Henry Shaw School of Botany of Washington University in affiliation with the Missouri Botanical Garden. Information The Annals of the Mis during the calendar year: Fel numbers constitute a volume. and November. Four Subscription Price $10.00 per volume Single Numbers 2.50 each Contents of previous issues of the Annals of thb Missouri Botanical re listed in the Agricultural Index, published bv the H. W. Wilson Company J FLORA OF PANAMA Part IV. Fascicle 1 PIPERACEAE 1 By T. G. YUNCKER Herbs, sKrubs, or small trees, not infrequently repent or scandent, often with nodose stems, glabrous or pubescent, vascular bundles scattered; leaves mostly alternate, sometimes opposite or vcrticillate, simple, usually entire, ranging in size from less tban 1 cm. up to 30-40 cm. or more long, often with glandular or pellucid dots; stipules lacking or present and commonly more or less adnate to the petiole; flowers very small, usually green, perfect or unisexual, in racemose or more commonly spike-like inflorescences, these usually somewhat fleshy, pedunculate, terminal or leaf -opposed, sometimes axillary or multiple on a common peduncle; the flowers mostly densely crowded, each in the axil of a usually peltate or sub- peltate, sessile or stipitatc, occasionally adnate or concave bract; perianth none; stamens usually 2-6, filaments mostly free, the anthers terminal, erect, 2-ceIlcd; ovary sessile or stipitate, 1 -celled, 1 -ovulate; fruit drupaceous, small. The Piperaceae, one of the larger families of flowering plants, is pantropic in its distribution. Species arc especially abundant in Latin America. They extend from the Keys of Florida through the West Indies, Mexico, and Central America, south- ward to Chile and Argentina. They occur in greatest number throughout northern South America from Brazil to Peru and northward to Panama and Costa Rica. Some species are widespread in their occurrence but each country has a rather large number of endemics. The total number of valid American species is unknown but unquestionably amounts to many hundreds and may approximate 2,000 or more. A number of species of widespread distribution; e.g., Piper aduncum, P, hispidnmy P. margjnattim^ and others appear to exhibit an infinite capacity of varia- tion In minor characters of leaf size, shape, and indumentum. The extremes often appear quite distinct in themselves but study of large numbers of collections may reveal intergradations to such a degree that the extremes lose their seeming sig- nificance. The writer has been reluctant to maintain species or even varieties established by previous workers on such apparent variations when sufficient evi- dence indicates intergradations. It is probable that errors of judgment have been made but it is felt that such errors are on the side of conservatism. On this basis, ^Issued March 31, 1950. (1) (n [Vol. 37 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN a number of previously described species have been reduced to synonymy. At the same time, a number of novelties have been described as new to science. It is believed that with greater abundance of material, especially mass collections, and with correlated field study still further reductions may be in order. On the other hand, such studies may prove the validity of species which now appear to be th same. Small collections had been made in Panama previous to 1800 especially by Nee and by Haenke, and around the middle of the last century larger collections were prepared by Secmann> Fcndler, Hayes, and others. However, collecting in Panama on an extensive basis did not get under way to any extent until after 1900. Miquel, the great Dutch authority and monographer of the Pipcraceae during the first part of last century, studied some of the earlier collections. Casimir DcCandoUe, third In line of the great Swiss family of botanists, following MiqueFs death, published in the DcCandolle Trodromus' in 18 69 a monograph of the Pipcr- aceae. Included In this pubUcation were all of the species then known from Panama, Following this work, there appeared an almost continuous stream of papers by DeCandoUe embodying the descriptions of hundreds of new species based on specimens sent to him by collectors as they explored farther and farther Into botanically unknown areas. Following DeCandolle's death in 1918, Professor William Trelease, of the University of Illinois, undertook the study of the family, especially as it is represented in the Americas, and published a number of papers including the descriptions of many new species. The earliest paper to deal exclusively with the Piperaceae of Panama was pre- -ed 1920. This included the descriptions of a number of new species based on collections made by PIttler and by Maxon In the early part of the century. Subsequently, large collections were made by Standley, Klllip, Piper, and others. From a study of this rather considerable accumulation of material, Trelease In 1927 presented a monographic study of the Panamanian Piperaceae which included 139 species and a number of varieties. Woodson and his associates at the Missouri Botanical Garden have more recently made extensive collections. These have also been studied by Trelease who has pub- lished more than 70 new species and a number of varieties based on this material. Thus, according to Trelease, more than 200 species arc represented in Panama. An attempt has been made In preparing the present paper to account for all species previously reported. Ninety-five species of Piper, 1 of Offonia, 2 of Pot/joniorphe, 1 of Sarcorbacbis, 1 of Audcrswulopipcr, 72 of Tcperoviiay 22 varie- ties of Piper and 7 of Pepcromia, making a total of 172 species and 29 varieties, are considered as valid in the present treatment. For the present study the writer has had the loan of the collections at Gray Herbarium of Harvard University, University of Illinois, Missouri Botanical Gar- Bot In addition, he has also had the loan of a number of critical specimens from Kew. He had the use of notes made by Trelease during his studies of type material in (2) 1950] FLORA OF PANAMA (Piperaceac) 3 European herbaria and was also given a grant by the Research Grant Fund of DePauw University- For the loan of specimens and all other assistance given him, the writer expresses his thanks. KEY TO GENERA^' a. Stigmas 2-5 (mostly 3-4); floral bracts various, often more or less pubescent; plants more or less woody, commonly shrubby or vine-like, or less frequently herbaceous. b. Inflorescence leaf-opposed. c. Inflorescence spicate; fruit sessile 1. Piper cc. Inflorescence racemose; fruit stipitate 2. Ottonia bb. Inflorescence axillary, spicate. c. Spikes several on a common peduncle ..._ 3. Pothomorphe cc. Spikes one, or, if more than one, not on a common peduncle. d. Spikes solitary 4. Sarcorhachis dd. Spikes 2 (or more?), superposed, axillary 5. Andi:rssoniopip£R aa. Stigmas 1 (may occasionally be cleft to simulate 2), stamens 2; floral bracts mostly round-peltate, glabrous; plants herbaceous, mostly com- paratively small e. Peperomia 1. PIPER Linnaeus Piper L. Gen. PL ed. 1. 3 3 3. 173 7. Shrubs or trees, occasionally subherbaceous, rarely scandent, often nodose; leaves alternate, simple, with rare pel pet to rugose or bullate above and lacunose beneath, glabrous or sparsely to densely hairy, often scabrous, frequently sparsely to densely glanduLar-dotted, palmately nerved or more commonly pinnately nerved from the lower half or two-thirds, or throughout, usually somewhat venulose with finer cross-connecting and anasta- mosing nervulcs; petiole of various lengths, rarely lacking, commonly more or less vaginate-grooved at least at the base and often to the blade, the margins of the groove wing-margined, the wing very narrow or apparently lacking, or broad and conspicuous; spikes leaf-opposed, cylindrical or rarely globose or subglobose, 1-10 mm. or more thick and up to 50 cm. or more long; peduncle from less than 5 mm. up to several centimeters long; rachis commonly somewhat ridged between the flowers^ the ridges low and smooth or papillate to fimbriate; flowers sessile and perfect (American species) or imperfect (Old World species); usually densely congested, in the axis of variously shaped bracts, the bracts sometimes glabrous but more commonly pilose or fimbriate; stamens 2-5; ovary 1, glabrous or sometimes papillate or pubescent; stigmas 2-4, rounded to filiform, sessile or on a short and thick to elongated and slender style; fruit variously shaped, one-seeded, drupe-like, with a thin pericarp and somewhat hardened endocarp. PRIMARY KEY TO SPECIES OF PIPER a. Ovary and fruit stylose I. Spp. 1-18 (pp. 5-14) aa. Ovary and fruit not stylose. b. Leaves scarcely peltate. c. Leaves palmately nerved, or pHncrvcd within the lowermost 5 mm II. Spp. 19-25 (pp. 15-20) cc. Leaves pinnately nerved. d. The principal branches of the midrib arising from somewhat below the middle to the upper third of the blade. *All keys in this paper are to be considered as artificial. P; 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN I Vol. 37 bb. e. Leaves mostly 20-25 cm. or more long X 10-20 cm. or more wide, base equi- or uncquilatcrally lobcd (shallowly so in a few species) HI. Spp. 26-34 (pp. 20-26) ec. Leaves mostly smaller or, if large, not lobcd at the base IV. Spp. 3 5-82 (pp. 3 0-60) dd. The midrib more or less equally branched to the upper fourth or throughout V. Spp. 83-93 (pp. 61-69) Leaves distinctly peltate VI. Spp. 94-96 (pp. 70-71) I. KEY TO SPECIES HAVING STYIOSE OVARIES AND FRUIT (In a few species Included here the styles may be very short or easily deciduous. Such question- ably stylose species have been also included in the non-stylose section of the key.) a. Leaves rugose-bullatc L P. pesarhsanum aa. Leaves not rugose-buUate. b. Leaves peltate 95. P. AIaxonii bb. Leaves not peltate. c. Spikes 15-20 cm. or more long; leaves 10 X 20 cm. or more; styles short. d. Petioles, twigs, etc. more or less warty-vcrrucose -- 50. P. impi-riale dd. Petioles, twigs, etc. not warty-verrucose. c. Twigs glabrate; nerves 6-S on each side f. Leaves elliptic, base oblique, cordatc-auriculate 2. P. albopunctu- LATISSIMUM ff. Leaves ovate, base not oblique or auriculate 34. P. Pit rn.Ri ec. Twigs densely brown-hairy; nerves 5-6 on each side 31. P. TARnrNS cc. Spikes and leaves much smaller. d. Spikes mostly 3 cm. or more long when mature. c. Leaves pinnately nerved throughout; plants entirely glabrous; fruit tetragonous; bracts cupulate, glabrous 3. P. DARirNFNSE ee. Leaves pinnately nerved below the upper third (or sometimes throughout in P. phytolaccacfoliuni) . I. Young twigs and/or leaves hairy. g. Spikes 6-8 cm. long; peduncle 1.5-2 cm. long 4. P. lincolnense ^g. Spikes 3-4 cm. or less long; peduncle 0.5-1 cm. long, h. Leaves ovate, 7-12.5 cm. wide; twigs retrorsely hirtellous 5. P. plrvenosum hh. Leaves lancc-elliptic, 3.5-5 cm. wide; twigs villous 6. P. piruligerum hhh. Leaves elliptic-obovate or obovatc, 7-12 cm. wide 7. P. exiguicaule ff. Entirely glabrous. g. Leaves ovate, base rounded; spikes 5 cm. or more long 8. P. crassinervium var. tocotanum gg. Leaves not ovate, base acute to acuminate. h. Leaves lance-elllptic; spikes 2-4 cm. long 9. P. phytolaccae- FOLIUM hh. Leaves obovate, strongly revolutc 10. P. macropunctatum dd. Spikes mostly 2 cm. or less long when mature. e. Leaves with 6-8 main lateral nerves. f. Leaves 10 cm. or more wide, nerves beneath puberulent 11. P. latiuracteum ff. Leaves less than 5 cm. wide, nerves beneath glabrous 12. P. candelarianum ec. Leaves with 3-5 main lateral nerves. f. Leaf base 5-10 mm. shorter on owe side, glabrous 13. P. magnantherum ff. Leaf base essentially equilateral, or if unequal more or less pubescent. g. Young twigs and/or leaves hairy. h. Leaves more or less long-ciliate, base acute 14. P. garagaranum hh. Leaves not ciliatc, base more or less cordulate. 1. Twigs, petioles, etc. minutely hirtellous 5. P. pervenosum ii. Twigs, etc. densely villous. j. Leaves about three times longer than wide, oblong- lanceolatc- - 15. P. puuistipulum jj. Leaves about 2 times longer than wide, elliptic 16. P. colon-insulae (4) 1950] FLORA OF PANAMA (Piperaccae) 5 gg. Young growth glabrous. h. Leaves mostly 6 cm. or more wide. i. Leaves broadly elliptic, abruptly short-acuminate, subrugose 17. P. distigmatum ii. Leaves ovate, gradually acuminate, not rugose 18. P. Davidsonii hh. Leaves mostly narrower 9. P. phytolaccaiz- FOLIUM obviously an error. Jahrb. 40:247. 1908, as peta Piper fluvii-initii Trel. in Ann. Mo. Bot. Gard, 27:291. 1940. A shrub or small tree, 3-8 m. tall, nodose; upper internodes rather short, thick, subreflcxed-hirtellous with an intermixture of scattered long hairs; leaves elliptic- to round-ovate, 5-11 cm, wide X 13-17 cm. long, apex sharp-acuminate, base rounded, subinequilaterally obtuse, one side 1-2 mm. shorter, bullate-lacunose, pinnately nerved from the lower half, the nerves about 6 or 8 on each side, rusty- villous above, at least along the nerves, rather densely so beneath, especially along the nerves, with a mixture of long and short hairs, margin ciliate, drying firm, dark, opaque; petiole 10-20 plus 1-2 mm.^ long, villous, vaginate and narrowly winged toward the base; spikes 8-10 mm. thick X 9-13 cm. long; peduncle stout, 1—2 cm. long, villous; bracts oblong-subligulate, pilose to near the subacute, curved apex; fruit subcylindrical, glabratc, stigmas 3, elongated, recurved on a moderate or short, somewhat sparsely hispid or glabrate style. Western Colombia and Panama, commonly at rather high altitudes. CHiRiQuf: trail from Cerro Punta to headwaters of Rio Caldera, alt. 2250-2500 m., Allen 1445; Bajo Chorro, Boquete District, alt. 2100 m., Davidson 204. This species rather closely resembles P. lactinosnm of western South America but differs in the type of pubescence, shape of the bracts, and more elongated fruits. The Chiriqui specimens agree in all respects with those seen from South America, 2. Piper albopunctulatissimum Trel. in Ann. Mo. Bot, Card. 27:287. 1940. A somewhat nodose shrub, 4 m. tall; upper internodes glabrous, sparingly warty-papillate; leaves elliptic, apex acute, base obliquely cordate-auricled, one side about 5 mm. shorter at the petiole, the larger side about equaling the petiole, about 18 cm. wide X 28-30 cm. long, pinnately nerved from below the upper third, the nerves 6-8 on each side, submarginally loop-connected and with num- erous cross-connecting and anastomosing nervules, glabrous above, the nerves beneath puberulent or crisp-pubescent, marginal ciliation lacking, sparsely yellow- glandular-dotted on both sides, microscopically white-dotted above when dry, drying rather firm, subtranslucent; petiole 45 plus 5 mm. long, winged-vaginate to near the blade, sparsely warty; spikes 5 mm. thick X 25 cm. long; peduncle "The base of the blade in many species of ?iper Is unequal on the two sides with one side some- what shorter at the petiole. The pctiolar length is given plus the difference in length between the two sides. (5) [Vol. 37 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 4 cm. long; bracts triangular-subpeltate, clliatCj tbe pedicel broad, concave; stigmas 3, linear, on a short, stout style. Known only from the following locality. cocle: north rim of El Valle de Anton, alt. 600-1000 m., Allen 1652, 3. P1P1.R DARiENENSE CDC. in DC. Prodr. 16^374. 1869. Otloma gJauccscens Miq. !n Seem. Bot. Voy. Herald, 200. 18 54, not Viper glaiicescem Jacq. Viper Jaricnse CDC. in Candollca 1:168. 1923. Piper Uxispicum Trel. in Contr. U. S. Nat. Herb. 26:24. 1927. Piper laxispicum var. latijoliuvi Trel. loc. cit. 1927. Piper Permari Trel. in Ann. Mo. Bot. Card. 27:295. 1940. Piper fagopyricarpum Trel. loc. cit. 28:426. 1941. A glabrous slirub, mostly scarcely 0.5 m. tall; stems simple or occasionally branched, 2-3 mm. thick, somewhat finely striate, greenish; leaves oblong-elliptic, subovate or lance-elliptic, 3-7 or sometimes 8 cm. wide X 12-19 cm. long, apex attenuately sharp-acuminate, base narrowed, subcquilaterally acute or rounded and obtuse, minutely auriculatc at the petiole, pinnatcly nerved throughout, the midrib rather prominent beneath, the lateral nerves slender, 6-9 on each side, divergent and submarginally loop-connected, drying green above, paler and subglaucous be- neath, membranous, translucent; petiole mostly scarcely 5 mm. long, longitudinally striate, scarcely vaginatc at the very base; spikes 3-5 mm. thick X 3-5 cm. long when mature; peduncle about 5 mm. long; rachis smooth or obscurely papillate; bracts cupulate, glabrous; fruit mostly exserted, free, globose-tetragonous, smooth or obscurely papillate, stigmas short, 3-4, on a short, stout style. Panama and northern Colombia. BOCAS DEL TORO: uppcr Changuinoki River, Permar s. ;;.; vicinity of Chiriqui Lagoon, von Wedel IO42, cHiRiQuf: vicinity of Puerto Armuelles, alt. 0-75 m,, Wooihon & Scbery 8^2, CANAL ZONE: Barro Colorado Island, Gatun Lake, Standley 313T5I Mexia 6jjl; Bailey ^ Bailey 88. Panama: Indio, Madden Lake, Miller 206 j — leaves maximum size, as described, san blas: forests around Puerto Obaldia, nit. 0-50 m., Piitier 42S8, The type specimen did not have a sufficiently matured spike to show the char- acters of the fruit, which are significant in this species. The description of the vegetative characters, however, agrees in all respects with the other specimens in- cluded here. It was reported by Sccmann as being used by the Indians for stupeyf- ing fish. Mexia made the same report for the specimen she collected on Barro Colorado Island. 4. Piper lincolnense Trel. in Field Mus. Nat. Hist. Bot. Ser. 18:347. 1937. A shrub (?); flowering Internodes moderately slender and short, obscurely retrorscly hirteUous on one side, the other side papillate glabrescent; leaves broadly elliptic or ovate-elliptic, apex short-acuminate, base slightly incqullaterally cordu- late, 7.5-10.5 cm. wide X 16-20 cm. long, pinnately nerved from below the upper third, the nerves about 6 on each side, rather prominent beneath, glabrous above, glandular-dotted and crisp-pubescent beneath, especially along the nerves, not ciliate, drying dull, papery, translucent; petiole 1.5-2 cm. long, rather densely (Q 1950] FLORA OF PANAMA (Pij^evaceae) 7 reflexed-hirtellous, vaglnatc to the blade; spikes 3-4 mm. thick X 6-8 cm. long, peduncle 1,5-2 cm. long, thinly and obscurely hirtellous; bracts triangular, the pedicel broad and concave, glabrous; fruit depressed, slightly flattened laterally with the rachis, stigmas 3, slender, on a short style. Known only from Lincoln Creek, Bocas del Toro. BOCAS DEL TORO: Lincoln Creek, Changuinola Valley, Dunlap 408, The broadly elliptic leaves, pubescent beneath, and stylose fruits characterize this species. 5, Piper PERVENOSUM CDC. in Candolica 1:268. 1923, name only; Schroeder, in CandoUea 3:137. 1926. A shrub; flowering internodes short, moderately slender, retrorsely hirtellous; leaves ovate, apex gradually acute, base subequilateral, rounded, obtuse or sub- cordulate, 7-12.5 cm. wide X 13-18 cm. long, glabrous on both sides or with the nerves beneath obscurely and minutely hirtellous, pinnately nerved from the lower half or slightly above, the nerves 3 or 4 on each side, with shorter intermediates, and minor lateral nerves to near the apex, nervules very numerous and closely spaced as seen by transmitted light, glandular-dotted, the glands comparatively large, drying gray-green, firm, translucent; petiole 5-20 mm. long, stout, vaginate- winged to the blade, retrorsely hirtellous; spikes 3-4 mm. thick X 3 cm. long; peduncle 5-10 mm. long, hirtellous; bracts triangular-lunulate, papillate; ovary subglobose, stigmas small, on a short style. Known only from, the following locaHty. SAN BLAs: San Bias coast, high hills back of Puerto Obaldia, alt. 50-200 m., Vittkr 4302 The gray-green twigs and ovate, finely venulose leaves are distinctive features of this species. 6. Piper peruligerum Trel. in Ann. Mo. Bot, Gard. 27:295. 1940, Small and suflfruticose; twigs, petioles and leaves at first rusty subtomentose- villous; internodes short and rather slender; leaves lance-elliptic, apex long- attenuate, base obliquely cordulate, one side 2-3 mm. longer, 3.5-5 cm. wide X 11-14 cm. long, pinnately nerved from the lower half, the nerves 3-5 on each side, with strong cross-connecting-anastomosing nervules, becoming slightly pouched between veins, the midrib above moderately pubescent, subvillous beneath, the nerves rather strongly so, the hairs staring and of various lengths, the longest nearly 1 mm. long, drying papery, translucent; petiole scant 3 plus 2-3 mm. long, densely villous; spikes 3 mm. thick X 2.5 cm. long; peduncle 3 mm. long; stigmas on a short style. Known only from the following locality. BOCAS DEL toro: Isla de Colon, Woodson, Allen ^ Sclbert IQSQ. The above description is drawn largely from the type description by Trelease inasmuch as the fragment at Ilhnois consists of two leaves only. Trelease indicated that the type is at the Missouri Botanical Garden but it has not been seen by the writer. (7) [Vol. 2>7 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN 7. Piper exi^uicaule Yunckcr, sp. nov, Frutcx, 30 cm. altus; foliis obovatis vel elliptico-obovatis, apicc subacutis, basi subacquilatcralibus, ad petiolum abruptc acutis, a tertio supero deorsum pinnatim venosis, vcnis utrinque ca. 6, supra dlssitc villosis, subtus ad venas crispe vel adpresse pubcscentibus, sat obscure glanduloso-punctatis; petiolo 2 cm. vel ultra longo; spicis, ut adsunt, 3 mm. crassis, 3.5 cm. longis; pedunculo 2.5 cm. longo; bractcis triangulari-subpcltatis, conferte pallido-fimbriatis; ovarlo subgloboso, stylo brevi; stigmatibus 3, rotundatis. Slirub, 3 cm. tall; stem sparsely pubescent (?); leaves obovate or elUptic- obovate, apex acutish, base subequilatcral and abruptly acute at the petiole, 7—12 cm. wide X 12-19 cm. long, pinnately nerved below the upper third, the nerves about 6 on each side, with prominent cross-conncc ting-anastomosing nervules, sparsely villous above, the nerves beneath crisp- or appressed-pubescent, rather ob- scurely glandular-dotted, drying papery, translucent, finely pellucid-dotted; petiole up to 2 cm. or more long, vaginate toward the base (?) ; spikes as yet young, 3 mm. thick X 3.5 cm. long; peduncle 2.5 cm. long, sparsely pubescent; bracts triangular- subpeltatc, densely pale-fringed; ovary subglobose, stigmas 3, rounded, on a rather short, stout style; fruit not developed. Known only from the one collection. BOCAs DEL TORO: FisH Crcck, Apr. 9, 1941, von Wedrl 21S7 (Herb. Missouri Bot. Gard., type). The type specimen is rather poor and frag- mentary but it does not appear to agree with any other known species Th e very sm all Fig. 1. Piper crasdncrilum var. focofanum plants, obovate leaves, densely fringed bracts, and stylose ovaries are distinctive characters. 8. Piper crassinervium Fi.B.K. var. toco- tanum (CDC) Yuncker, comb. nov. Piper cxserrens (Miq.) CDC. var. tocotannm CDC in Ann. Conscrv. & Jard. Bot. Geneve 21:239. 1920. Viper papyracejim Trel. in Contr. U. S. Nat. Herb. 26:31. 1927. Piper noi'ae-helveticac Trel. in Ann. Mo, Bot Gard. 27:294. 1940. An essentially glabrous shrub or small tree, 2—5 m. tall; upper internodcs coniparatively short but soon lengthening, glabrous; leaves ovate, 5-14 cm. wide X 13-23 cm. long, apex acuminate, base rounded, obtuse or in large leaves sometimes shallowly cordate, equilateral or with one side slightly shorter, abruptly acute (^) 1950] FLORA OF PANAMA (Piperaccac) 9 at the petiole, pinnately nerved from below the middle, the nerves 4 or 5 on each side, prominent beneath, glabrous or at most only sparsely minutely puberulcnt along the nerves, drying firm, translucent; petiole about 15 mm. long, vaginate and narrowly winged to the middle or above; spikes 5-7 mm. thick X 12-15 cm. long when mature; peduncle stout, 5-15 mm. long, pubescent; bracts rounded- or triangular-subpeltate, marginally fringed, zonately arranged; fruit rounded, gla- brate, stigmas 3, recurved, on a rather short style. Venezuela, Colombia, and Panama; the species Colombia and Ecuador. CHiRiQUi: vicinity of "New Switzerland," central valley of Rio Chiriqui Viejo, alt. 1800-2000 m., Allcft IJjQ, 1416; forests around El Boquete, alt. 1000-1300 m., Pitlicr No significant difference has been found by means of which the Panamanian specimens can be distinguished from those from South America. 9. Piper phytolaccaefolium Opiz in Presl, Reliq. Haenk. 151. 1830. Viper brevhpicatiim Opiz, loc. cit. \5\, pU 28, 1830. Artanthc hcteropbylla Miq., Syst. Pip. 502. 1844. Artantbe brcvhpicata Miq., loc. cit. 508. 1844. Artanthc phytolaccacfolia Miq., loc. cit. 534. 1844. Peltobryart pbytolaccaefolium Presl, Epim, Bot. 224. 1849. Piper sirigidare CDC. in Jour. Bot. 4:161. 1866. Piper ampboricarpum Trel. in Contr. U. S. Nat. Herb. 26:29. 1927. Piper simulafis Trel. in Field Mus. Nat. Hist, Bot. Ser. 18:361. 1937. A glandular-granular shrub, 1-2 m. tall, glabrous or obscurely and transiently puberulent; twigs rather slender; leaves lance- or oblong-elliptic, 3-6 cm. wide X 10-13 cm. long, or occasionally up to 8 X 19 cm,, apex acuminate, base acute to cuneate, mostly equilateral, glandular-dotted, often strongly so, pinnately nerved throughout or some leaves from below the upper third, the nerves 4, or in large leaves 5-6 on each side, mostly scarcely ciliate; petiole 5-10 mm. long, vaginate at the base; spikes 5 mm. thick X 2-4 cm. long; peduncle 5-20 mm. long, glabrate or very sparsely hirtellous; bracts triangular- or rounded-subpeltatc, marginally fringed; fruit globose, stigmas 3, small, on a short subulate style. Colombia, Ecuador, and Panama. Panama: without definite locality, Nee jdj, J64, jd/. bocas del toro: Changuinola Valley, Diinlap 459. CHiRiQuf: Boquete District, Boquete, alt. 1140 m., Davidson 6ig; along Rio Caldera, south of El Boquete, alt. 1250 m., Killip 3611, No significant differences have been discovered between the Panamanian speci- mens cited here and those which have been studied from South America. Those from Panama, however, are, so far as seen, less conspicuously glandular-dotted th: are those of South America, A specimen collected by Nee without definite locality indicated has the leaves ciliate and with a few scattered long hairs on the upper surface. (^) IVoL. 37 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN 10. Piper macropunctatum Yuncker, sp. nov. Frutcx, ca. 0.5 m. altus, dissite hirtulus; intcrnodiis usque ad 6 cm. longis, obvie confertc glandulosis; foliis elliptico- obovatis, apice obtusis vel abrupte brcvi-acutis, basi inaequi- lateralitcr acutis, latere altero ca. 2 mm. breviore, utrinque glabratis vel subtus ad basem dissite pubesccntibus, a medio laminae deorsum pinnatlm venosis, vcnis utrinque 5, utrinque confertc glanduloso-punctatis, marginc revolutis; petiolo 2-4 cm. cum 2 mm. longo; basi vaginato; bractcis triangulari- lunato-subpeltatis; ovario ovoideo glanduloso, stylo brevi, crasso, subalato, stigmatibus parvis. A small, essentially glabrous sbrub, about 0.5 m. tall; stem unbranched, internodes up to 6 cm. long, strongly glandular-dotted; leaves clUptic-obovatc, apex obtuse or abruptly short-acute, base inequilaterally acute, one side about 2 mm. shorter, 6-8.5 cm. wide X 13-- 15 cm. long, glabrous on both sides or sparsely pubescent near the base beneath, pinnately nerved from the lower half, the nerves 5 on each side, with cross-connccting-anastomosing nervules, submarginally loop-connected, strongly glandular-dotted on both sides, drying rather thin, translucent, pellucid-dotted, revolute; petiole 2-4 cm. plus 2 mm. long, glabrous or sparse- ly pubescent, vaginate near the base; spikes 3 mm. thick X 2.5 cm. or more long; bracts triangular-lunate-subpeltate, glabrous (?); ovary glandular, ovoid, with a short, stout, subulate style, stigmas small, sessile. Known only from the type locality. BOCAS DEL TORO; Pumpkin River, July 28, 1941, von Wedel 2j6l (Herb. Missouri Bot. Fig. 2. Pil)er macropuncfatuw Gard., ). The small, simple, glabrous plants, strongly glandular-dotted with compara- tively large dots, obovatc, revolute leaves, and pointed ovary distinguish this species. The only spike present on the type specimen is fragmentary and young. 11. Piper latibracteum CDC. in Smiths. Misc. Coll. 71'':5. 1920. A shrub, about 1 m. tall; flowering internodes moderately stout and short, glabrous; leaves broad-elliptic, apex gradually acute, base narrowed, obtuse or slightly cordulate, one side 1-2 mm. shorter, 11-14 cm. wide X 20-24 cm. long, pinnntely nerved below the upper third, the nerves 7-8 on each side, with inter- mediates and slender cross-connecting and anastomosing nervules, glabrous above, the nerves beneath puberulent, somewhat rugose, glandular-dotted, drying subglossy and green above, paler beneath, papery, translucent; petiole 15-20 plus 1-2 mm. long, vaginate and narrowly winged to the blade; spikes 8 mm. thick X 2 cm, long, apiculate; peduncle stout, scarcely 10 mm. long; bracts triangular-lunate, (10) 1950] FLORA OF PANAMA (Piperaccac) 11 glabrous, or marginally papillate, pedicel broad, concave; ovary glabrous, stigmas 2, recurved, on a short, stout style. Southern Panama. darien: Boca de Pauarando, Sambu River, alt. 20 m., Pittier 5588; Cani-Cuasi Trail, Chepigana Dist., alt, 900 m., Terry 6 Terry 144I; alt. 1650 m., T^rr3; ^ Terry 1 581. The numerous nerves, puberulent beneath, short, thick, apiculate spikes, and stylosc ovaries distinguish this species. 12. Piper candelarianum CDC. in Linnaca 37:3 57. 1872. Piper globosum CDC. loc. cit. 340. 1872 not Poir. 1804. Piper psiloeladum CDC in Bull. Soc. Bot. Bclg. 30'^:211. 1891. Scandent shrub, glabrous; flowering internodes rather slender and short, red- granular; leaves lance-elliptic or the lower subovate, apex gradually acute, base sub- equally acute or one side sHghtly shorter than the other, 2,5-4 cm. wide X 8-12 cm. long, pinnately nerved throughout, the nerves mostly 6-7 on each side, with rather obscure intermediates and cross-connecting ncrvules, drying thin but firm, translucent, prominently yellow- or red-glandular-dotted on both sides; petiole scarcely 1 cm. long, vaginatc toward the base; spikes 4-8 mm. thick X 1.5-2.5 cm. long, mucronate; peduncle slender, about 1 cm. long, eventually recurving; bracts triangular-subpcltate, sparingly very short-fringed on the upper margin; fruit rather large, globose, attenuate into a short, stout style, conspicuously glandu- lar roughened, glabrous, stigmas 3. Costa Rica and adjacent Panama. BOCAS DEL TORO: Old Bank Island, vicinity of Chiriqui Lagoon, V07t Wedel 1994. The short, thick styles, short spikes, glandular twigs, and pinnately nerved leaves are distinctive characters of this species. It resembles P. ctanaralcuse of Colombia to some extent but differs because of its completely glabrous twigs and leaves and with the nerves of the leaves not submarginally loop-connected as in that species. 13. Piper magnantherum CDC in Smiths. Misc. Coll. 71^:16. 1920. A glabrous, strongly red-glandular shrub (?); flowering internodes short and slender; leaves lance-elliptic-oblong, apex acuminate, base obUquely acute, one side 5-10 mm. shorter, 3.5-5 cm. wide X 12-15 cm. long, pinnately nerved from the lower half, the nerves 4 or 5 on each side, or with additional lateral nerves to the upper fourth, with few cross-connecting nervules, rather loosely ciliate toward the apex, drying thin, translucent; petiole 5-10 plus 5-10 mm, long, or on lower leaves up to 2 cm., vaginate at the base; spikes as yet young, 2-3 mm. thick X 1.5 cm. long, mucronate, peduncle 5-8 mm. long; bracts triangular-subpeltate, marginally pale-ciliate; ovary ovoid, narrowing into a short, stout style, stigmas 3, small, short. Known only from the following locality. (U) [Vol, 37 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN darien: Ccrro dc Garagara, Sambu basin, southern Daricn, alt. 500-974 m., Piftier 3666, The densely red-glandular leaves, twigs, etc., strongly inequilateral base of the leaves, and stylose ovaries distinguish this species. It resembles P, pbytolaccacfolhim in many respects but differs because of the inequilaterally based leaves, and con- spicuous red-glands. 14. Piper garagaranum CDC. in Smiths. Misc. Coll. 71*': 15. 1920. A shrub; flowering Internodes moderately slender, rather short, moderately to sparsely loosely villous; leaves lance-elliptic, apex sharp-acuminate, base equilateral and obtuslsh, or more commonly inequilateral and acute with one side about 2 mm, shorter, 4.5-7 cm. wide X 13- 17 cm. long, pinnatcly nerved from below the middle, the nerves 3-4 or 5 on each side, with rather open, cross-connecting- anastomosing nervules, loosely long-villous, especially along the nerves beneath, somewhat long-ciliate, glandular-dotted beneath, drying rather thin and papery, translucent; petiole 20 plus 2 mm. long on lower leaves and vaginatc-winged to about the middle, reduced upy%^ard3 villous; spikes 3 mm. thick X 15-18 mm, long; peduncle about 5 mm. long, villous; bracts small, concave, short-ciliate along the concave margin; ovary ovoid, narrowing into a short, stout style, stigmas rather short, recurved. Darien Province. darien: foothills of Garagara, Sambu basin, alt. 30-500 m., Vtttler 5618; Chcpigana Dist., Cana-Cunsi Trail (Camp 2), alt. 600 m., Terry ^S Terry 1 432. The short spikes, and thinly long-villous and ciliate leaves distinguish this species. 15. Piper tubistipulum CDC. in Smiths. Misc. Coll. 71*':5. 1920. Piper pubisfipuliitfi var. ciidyhmim Trcl. in Contr. U. S. Nat. Herb. 26:28. 1927. A small shrub, 1 m, tall; upper internodes short, densely retrorscly hirsute; , leaves oblong-lanceolate, apex gradually rather long-acuminate, base somewhat narrowed, corduLite, one side 2-3 mm, shorter, glabrous above except for the mid- rib which is pubescent near the base, pubescent beneath with the nerves hirsute, 5—6 cm. wide X 15-19 cm, long, pinnatcly nerved from the lower half, the nerves 4 on each side, raised beneath, with cross-connecting and anastomosing nervules, sparsely glandular-dotted beneath, smooth or subrugose, drying firm, translucent; petiole stout, 6-8 plus 2 mm. long, densely hairy, deeply vaginate to the blade; spikes 3-4 mm. thick X 2 cm. long, mucronate; peduncle 5 mm. long, hirsute; bracts triangular-lunate, glabrous, pedicel broad, cupulate; ovary narrow^ed abruptly into a short, thick style, stigmas 2, recurved. Central Panama. CANAL zone: Barro Colorado Island, Gatun Lake, Standlcy JTJ64; hills around the Agua Clara Reservoir, near Gatun, alt. 20-3 m., Piftier 2661, Panama: Indio, Miller 20/ 8. (12) 1950] FLORA OF PANAMA (Piperaccae) 13 The densely pubescent stems, oblong-lanceolate leaves, pubescent beneath and with somewhat inequilateral, cordulate bases, short spikes, and stylose fruits dis- tinguish this species. The spike on the type of var. e7istylosnm is so young that It is not possible to determine the style character. The specimen agrees so well otherwise that it is believed that It is same as the species. 16. Piper colon-insulae Trel. in Ann. Mo. Bot. Gard. 27:290. 1940. A shrub, 2 m. tall; upper Internodes relatively slender, densely dingy-villous; leaves elliptic, apex acute or short-acuminate, base inequllaterally cordulate, mar- gins subequal at the petiole, 5-8,5 cm. wide X 13-16 cm. long, glabrous above or the midrib with a few hairs at the very base, villous beneath, at least along the nerves, pinnately nerved below the upper third, the nerves about 5 on each side, with numerous cross-connecting-anastomosing nervules, drying papery, translucent; petiole 1-1.5 cm. long, densely dingy-villous, vaginate to the blade; spikes 5 mm. thick X 2.5-3 cm. long; peduncle 8 mm. long, subhirsute; bracts narrowl) angular-lunate, glabrous; fruit subquadrangular-oblong, stigmas on a very short style. Known only from the following locality. BOCAS DEL TORO: Isla de Colon, ah. 25-75 m., Woodson, Allen & Seihert 1934, 17. Piper distigmatum Yuncker, sp. nov. Frutex (?) vel arbuscula glabratus 2-3 m. altus; internodils florifcris modice brevibus gracilibusque; foliis late ellipticis, apice abrupte breviter acuminatis, basi obtusis vel ad petiolum abrupte subacutis, ^ tn- rugosis, an fra medium pinnatim venosis, venis utrinque 5, subtus prominentlbus, rubro-glanduloso-punctatis; petiolo 1.5-2.5 cm. longo, ad laminam vaginato; spiels 5-8 mm. crassis, 1.5—2.2 cm. longis apiculatis; pedunculo ca. 1 cm. longo; bracteis tri^^ngu- lari-lunatis glabratis; drupa subglobosa, stig- matibus binis ligulatis recurvis, stylo sat longo crasso subulato fultis. A glabrous shrub (?) or small tree, 2-3 m. tall; flowering Internodes moderately short and slender; leaves broadly elliptic, apex abruptly short-acuminate, base rounded, obtuse or abruptly subacute at the petiole, 8-9.5 cm. wide X 14-17 cm, long, rugose, pinnately nerved from below the middle, the nerves 5 on each side, prominent beneath, not branched but with slender cross-connect- ing and anastomosing nervules, red-glandular- dotted, especially on the lower surface, drying firm, translucent; petiole 1.5-2.5 Fig. 3. Piper distigmatum (n) [Vol. 37 14 MISSOURI BOTANICAL GARDEN cm. long, vaginate to the blade; spikes 5-8 mm. thick X 1.5-2.2 cm. long, apiculate; peduncle about 1 cm. long; bracts triangular-lunate, glabrous, pedicel broad, curved; fruit subglobosc, stigmas 2, ligulatc, recurved, on a rather long, stoutish, subulate style. Known only from the type locality. CHiRiQUi: Boquctc Region, Cerro Horqucta, cloud forest, alt. 1950 m., April 17, 1940, von Hagcn ^ von Hagen 20og (Herb. Missouri Boc. Gard,, type); April 27, 1940, ton Hagen & von Hagcn 20§S. The broadly elliptic, somewhat rugose leaves, and bistigmatic, rather long- styled fruit distinguish this species. It differs from P. Daihhonii because of the shape of the larger, rugose, more abruptly acuminate leaves, 5-nerved on each side. 18. PiPiR Davidsonii Yuncker, sp. nov. Frutex parvus glabcr; internodlis supcris sat brevibus graclllbusque; foliis ovatis vel cUiptico-ovatis, apice acuminatis, basi rotundatis ad pctiolum abrupte acutis, infra medium pinnatim vcnosis, venis utrinquc 4, manifeste rubro-glanduloso- punctatis; pctiolo ca. 1,5 cm. longo, ad laminam vaginato; spicis, ut adsunt, 5 mm. crassis, 1 cm. longis; pedunculo 1 cm. longo; bractels triangulari-subpeltatis, glabratis; ovario globoso (?), stigmatibus 2, stylo longo fultis. A small, glabrous shrub, 1-2 m. tall; upper internodes rather short and slender; leaves ovate or clllptic-ovatc, apex acuminate, base rounded, abruptly short-acute at the petiole, 6-7.5 cm. wide X 14-16 cm. long, plnnatcly nerved from below the middle, the nerves 4 on each side or with a very slender fifth nerve at the base In large leaves, strongly red-glandular-dotted, especially beneath, green above, pale beneath, drying firm, translucent; petiole about 1.5 cm. long, vaglnate to the blade; spikes, as represented, 5 mm. thick X 1 cm. long; peduncle 1 cm. long; bracts triangular-subpcltate, glabrous, the pedicel broad, concave, papillate; ovary globose (?), stigmas 2, on an elongated style. Known only from the type locality. cHTRiQuf: Bajo Cliorro, Boquctc District, alt. 1800 m., Feb. 15, 193 8, M. £. Davidson 2Sj (Herb. Missouri Bot. Gard., typf). 11. KEY TO SPECIES WITH PALMATELY NERVED LEAVES (The nerves may coalesce within the lowermost 5 mm.) a. Fruit p.ipillate, with a prominent, glabrous, apical disc -- 19. P. ri-.ticulatum aa. Fruit lacking any evident apical disc. b. Leaf base cordate. c. Leaves 9- to 11-ncrved - 20. P. marginatum cc. Leaves 5-nerved - 21. P. aristolochiae- rOI IL'M bb. Leaf base rounded or narrowed, scarcely cordate, leaves more or less elliptic. C. Leaf base inequilateral with one side about 2 mm. shorter; flowers approximated. d. Petiole vagina margin glabrous; leaves lanceolate, 3-ncrved 22. P. humorigaudi ns dd. Petiole vagina margin ciliate; leaves lance-elliptic, 5-nervcd 23. P. coNVERSUM cc. Leaf base essentially equilateral, 5- to 7-ncrvcd. d. Flowers distant; bracts glabrous 24. P. pinoganense dd. Flowers more compact; bracts papillatc-pIlose .25. P. mpdium (14) 19 5 0] FLOPv^A OF PANAMA (Pipcraccae) 15 19. Piper reticulatum L. Sp. Pi. 1:29. 1753. Piper smilacifolium H.B.K. Nov. Gen. & Sp. 1:56. 1815. Piper latum H.B.K. loc. cit. 57. 1815. Enckea smilacifoUa Kunth, In Linnaca 13:605. 1839. Enckea lata Kunth, loc. cit. 606. 1839. Enckea reticulata Miq. Syst. Pip. 365. 1843. Macropipcr latum PresI, Epim. But. 223. 1849. Piper Duchassaiugii CDC. in DC. Prodr. 16^:251. 1869. A glabrous shrub or small tree up to 4-6 m. tall; branches slender, nodose; leaves elliptic, elliptic-ovntc, or broadly ovate, 8-15 cm. wide X 15-25 cm. long or occasionally up to 25 cm. wide X 30 cm. long, apex acuminate, base essentially equilateral, more or less acute, truncated, or in large leaves shallowly cordate, palmately 5- to 7-nerved, or larger leaves 9-ncrved, the nerves prominent beneath, reticulate-veiny with rather prominent cross-connecting nervules, drying firm and chartaceous; petiole 1—2 cm. long, glabrate or very obscurely puberulent-papillate, grooved above, vaginate near the base; spikes 5-6 mm. thick and up to 10 or 12 or occasionally 15 cm. long when mature; peduncle slender, 1-2 cm. long; rachis with membranous, fimbriated ridges; bracts pcltatcly dilated upward, pedicel and margin bearded; stamens 4; fruit obovoid, somewhat 4-sided, papillate, with an apical, smooth, disc-like area, stigmas 3 or 4, rounded. West Indies and on the mainland from Central America through northern South America. BOCAs DEL TORO: Lowcr Changuinola River, Stork JO. CHiRiQuf: Progreso, Cooper d Slater l8^. canal zone: along Cano Quebrado, Pittier 6g86; around Gamboa, alt. 40-80 m., Pittier 377^ > between Gorgona and Gatun, alt. 10—50 m., Pittier 227Q; near Gatun, Standlcy 27201; Mamci Hill, alt. 20-90 m., Pittier 3811; near Summit, Standley 25728; Cerro Gordo, near Culebra, Standley 26008; Rio Pedro Miquel, near East Paraiso, Standley 2gg6g; vicinity of Fort Sherman, Standley jogjg; Barro Colorado Island, Standley JI267; Miller 207 j; Matachin, Kuntze igi8; Obispo, Standley 3T726; vicinity of Gatuncillo, Piper 5603, 5620; Rio Agua Salud, near Frijolcs, Piper 58 sg, 5860; Empire, Miller 1867; Madden Dam Road, Miller 180O; along Rio Chagrcs between Gamboa and Alahucla, alt. 30-60 m., Allen gj6, colon: between France Field and Catival, Standley 30338; along Rio Fato, alt. 10-100 m., Pittier 3872, 3882; Fato, Pittier 3842. Panama: Rio Tapia, Standley 26160, 28035, 28241; near Tapia River, Juan Diaz region, Maxon & Harvey 66 jO, 6708; Juan Diaz, Standley 30548; Rio Tccumen, Standley 2g363, darien: along Sambu River, Pittier 5570, The large, glabrous, rather veiny leaves, comparatively short spikes, and especially the prominent glabrous disc at the top of the papillate fruit, characterize this species. There is a wide variation in the size of the leaves. 20. Piper marginatum Jacq. Icon. Pi. Ran 2:2, pL 215. 1786. Piper decnmanum Aubl. Pi. Guian, Fr. 1:21. 1775, not L. 1760, nor Miller 1763. Piper caialatum Vahl, Eclog. Am. 1:3. 1796. Piper alare Ham. Prodr. PI. Ind. Occ. 3. 1825. Schilleria caudata Kunth, in Linnaea 13:716, 1839. Schilleria marginata Kunth, loc. cit. 718. 1839. Artanthe caudata Miq. Syst. Pip. 3 80. 1843. Artanthe marginata Miq. loc. cit. 381. 1843. Artanthe alaris Miq. loc. cit. 406. 1843. (W [Vol. ^7 \6 ANNALS OF THE MISSOURI BOTANICAL GARDEN npcr regressum Anders, in Miq. loc. cit. 3 80. 1843, as synonym. Piper san-josearnim CDC. in Linnaea 37:351. 1872. Viper san-joseartnm var. chiriquiniim Trel. in Ann. Mo. Bot. Card. 25:826. 1938. Piper san-joseanuni var. kobcnse Trcl. loc. cit. 17\197, 1940. Piper san-joseuNUfn var. panaviauinu Trcl. loc. cit. 1940. piper saii-]oscanum var. tabogense Trcl. loc. cit. 1940. A shrub or small tree with moderate to rather short Internodes; branches slen- der, glabrous and blackish when dry; leaves round-ovate, 8-15 cm. wide X 10-20 cm. long, or rarely some leaves Fig. 4. P. fnarginatum somewhat larger, apex acuminate to long- attenuate-acuminate, base openly cordate or occasionally subtruncate, glabrous on both sides except for slight ciliation and submarginal silkincss, palmately 9- to 11- ncrvcd, nerves entirely free, or the Inner- most pair of laterals slightly coalescing with the midrib at the base, cross-con- necting and anastomosing ncrvules rather prominent, the leaf margins confluent across the top of the petiole, drying mem- branous, translucent, pcllucid-dotted; pet- iole up to 4 cm. long or on larger leaves somewhat longer, vaginate-winged to near the blade; spikes 2-4 mm. thick wben mature and up to 10-13 cm. or occas- II lonaiiy 20 cm. long; peduncle about 10 mm. long; bracts transversely triangul peltate, sessile. marginally fringed; fruit smooth, obpyramidal, angular; stigmas 3, linear, West Indies, on the mainland of Central America, and southward to Brazil and Ecuador. Panama: without Indication of locality, Nee jOO, j8/^. province (?): Santa Rita Trail, Cowell iT/'j T22. cHiRiQuf: between Rio Tabasara and Rio Tinta, WooJsou, Sc'ibcrf ^ Allen 416. VERAGUAs; Isla dc Uva, Contrcras group, Pitticr jII2, cocLE: El Vallc, Miller l8l2. canal Zone: Monkey Hill above Colon, Lehniann XLVJ; New Limon, Stevens TOlj; Culcbra, Stevens Q2/'; Fort Kobe road, Woodson, Allen ^ Seiberf 142^; western slope of Ancon Hill, vicinity of Balboa, alt. 20-75 m., Seiberf 3SS; Ancon Hill, Killip 3041, 12087; Standley 26326; Balboa, StanJlcy 2360O; between Frijolcs and Alonte Lirio, alt. 30 m., Killip 12116; Frijoles, Standley 2762/; Empire, Miller 1862; Juan Mina, Piper 36g6, j6gS; near old Fort Lorenzo, mouth of Rio Chagrcs, Piper ^SgS, SQiy, 5q62; Culebra, alt. 50-150 m., Pittier 2146; Ccrro Gordo, near Culebra, alt. 50-290 m., Piltier 3737; standley 2^g64; Corozal, Stevejis Tt8; Agua Clara Reservoir, Stevens 546; near Sum- mit, Standley 25731, 26g38; near Gatun, Standley 27247, 27288; Gamboa, Standley 28334, 28^36; near Fort Randolph, Standley 286$$; vicinity of Fort Sherman, Standley 30042; Stevens 267; Darien Station, Standley 31557; Barro Colorado Island, Standley 40832; Kenoyer 5gg. colon: between Fato and Playa de Damas, Pittier 3Q34; vicinity of Santa Isabel, Pittier 4176, Panama; Gorgona Beach, Woodson, Allen G? Seibert l6go; Taboga (16) 1950] FLORA OF PANAMA (Pipcvaccae) 17 Island, alt. 0-186 m., WooJson, Allen ^ Scibert Ijjl; KilUp 3165, 3195; Miller 1848; Pittier 3553; Standley 27044, 27057, 27847; Camino de la Granja, Bro. Heribcrto 166; Bella Vista, Killip I20I2; Staudley 25332; Madden Lake, Miller 2077; near Punta Paitilla, Standley 26274; near big swamp, cast of Rio Tecumcn, Staudley 265gg; along Corozal road, near Panama, Standley 26834; Ri^ Tapia, Standley 28207; between Fort Clayton and Corozal, Standley 2g2I4; Rio Tecumen, Standley 29482; Juan Diaz, Standley 3052I; between Las Sabanas and Matias Hernandez, Standley 31835. herrera: Pcse, alt. 50 m., Allen 801, darien: around Garachine, Pittier 5512. KEY TO THE VARIETIES OF P, MARGINATUM a. Sinus wide, open, lobes more or less divergent. b. Leaves not pilose above, the nerves glabrous or pubcrulent. c. Nerves scarcely pubcrulent 20. P. marginatum cc. Nerves pubcrulent, at least above 20a. var. anisatum bb. Leaves appresscd-pilose above, at least toward the apex; nerves beneath pubescent 20b. var. catalpae- FOLIUM aa. Sinus closed or nearly so, lobes approximated or overlapping 20c. var. clausum 20a. Piper marginatum var. anisatum (H.B.K.) CDC. in Urb. Symb. Ant. 3:172. 1902. Piper anisatum H.B.K. Nov. Gen. & Sp. 1:58. 1815. Piper san-]oseanum var. remediosense Trel. in Ann. Mo. Bot. Gard. 27:297. 1940. The nerves pubcrulent or hirtellous, at least on the upper surface. Occurs with the species. CHiRiQui: Rio ChiriquI to Rcmedios, alt. 15-50 m., Woodson, Allen ^ Seibert IIQI, PANAMA: Perlas Archipelago, San Jose Island, Johnston 319, 472; Saboga Island, Miller 1953; a^ong Charare River, near Chcpo, alt. 50-100 m., PHtier 4708, darien; trail be- tween Pinogana and Yavisa, alt. 15 m., Allen 260, 20b. Piper marginatum var. catalpaefolium (H.B.K.) CDC in DC Prodr. 16^:246. 1869, '/ ■/ 1815. 1839. Artantbe catalpaefolia Miq. Versl. en Medel. Kon. Nederl. Inst. 1842:197. 1843. Piper pseudomarginatuni CDC. in Bull. Herb. Boiss. 6:492. 1898. Leaves appresscd-pilose above, at least toward the apex, and pubescent along the nerves beneath. Occurs with the species. CANAL zone: Corozal, Stevens 63, Panama: Perlas Archipelago, Trapeche Island, Miller 1907, darien: vicinity of Boca de Cupe, alt. 40 m., Allen 869, 20c. Piper marginatum var. clausum Yuncker, var. nov, Foliis suborbicularibus, basi profunde cordatis, sinu angustato, lobis approximatis vel invicem obtegcntibus. Leaves suborbicular, base deeply cordate with narrow sinus, the lobes approxi- mated or overlapping. Central Panama. CANAL zone: Empire, March 5, 1937, Miller 1861 (U. S. Nat. Herb., type); vicinity of Salamanca Hydrographic Station, Rio Pequenl, alt. 80 m., July 28-29, 193 8, Woodson, Allen ^3 Seibert 1608 — this specimen shows somewhat more open sinuses than does the type. (17) [Vol. 37 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2L Piper aristolochiaefoliuni (Trel.) Yunckcr, comb. nov. Piper san-joscafinm var. arhtoloclnacfolunn TrcL in Contr. U. S. Nat. Herb. 26:22. 1927. A shrub, 1.5-2 m. tall, twigs somewhat zig-zag, granular-papillate, moderately slender; leaves bullate and dark green when growing, rather narrowly ovate, apex long-acuminate, base deeply cordate, sinus open, obtuse, with parallel margins, 8- 9 cm. wide X 15 cm. long, palmately 5 -nerved, the innermost laterals not continuing to the apex, the midrib strongly branched upward, the nerves and cross-connecting- anastomosing nervules prominent beneath, the innermost pair of laterals entirely free from the midrib, with scattered, soft, white hairs above, paler and with the nerves granulate-papillate or subhirtellous beneath, ciliate, the hairs restricted to a single marginal row, pale-ycllow-glandular-dotted beneath, drying thin, trans- lucent; petiole 3-4 cm. long, vaginate to about the middle, scarcely winged, granulate-subhirtcllous. No inflorescence present. Known only from the type locality. CANAL zone: wet forest, hills north of Frijolcs, "scarce," Dec. 19, 1923, Standley 2/J24 (U. S. Nat. Herb., typf). The specimen on which this species is based is sterile but the vegetative char- acters are believed to be sufficiently different from P. marginatnin to w^arrant specific rank. It differs from that species chiefly because of the 5 -nerved leaves, the midrib strongly branched upw^ard, the lateral nerves not continuing to the apex, the ciHation restricted to a single row^ of marginal hairs, and the shape of the basal sinus. 22. Piper humohigaudens Trel. in Ann. Mo. Bot. Gard. 27:293. 1940. A branching, nodose, glabrous shrub, 2 m. tall; upper internodes short and slender, finely ribbed; leaves lanceolate, apex rather gradually blunt-acuminate, base inequilaterally narrow^ed downward, one side about 2 mm, shorter at the petiole, the shorter side acute, the longer side rounded, subacute, 2.5-3 cm. wide X 8-11 cm. long, 3-nerved, the nerves unevenly coalescing w^ithin the lowermost 5 mm., salient beneath, cross-connecting and anastomosing nervules very slender, drying rather firm, subtranslucent; petiole scarcely 5 plus 2 mm. long, vaginate near the base; spikes 3 mm. thick X 4-5 cm. long; peduncle 1-1.5 cm. long; rachis ridges fimbriate; bracts small, subcucullate, marginally short-fringed; fruit somewhat granular, dcpressed-subglobose, laterally compressed; stigmas sessile. Known only from the following locality. cocle: wet north rim, El Valle, alt. 80-100 m., May 21, 1939, Allcu 1S33, 23. Ptpfr conveksum Trel. in Ann. Mo. Bot. Gard. 27:290. 1940. Subarborescent, 2 m. tall; young branches nodose, very slender, internodes short, finely striate, obscurely and minutely hirtellous glabrescent; leaves lance-elliptic, apex caudatcly acuminate, base about 2 mm. shorter on one side, acute, 3-4.5 cm. wide X 10-12 cm. long, palmately 5 -nerved with the outermost pair of nerves delicate and loop-connected with cross-connecting nerves upward, the innermost (18) 1950] FLORA OF PANAMA (Piperaceae) 19 pair of lateral nerves slightly coalcscent with the midrib at the base, prominent beneath, drying thin, translucent; petiole about 3 mm. plus 2 mm. long, obscurely hirtellous, vaginate toward the base, the margin of the vagina ciliate; spikes as yet young, 2 mm. wide X 2.5 cm. long; peduncle 5 mm. long, glabrous; bracts in- flcxcd-subpcltate, ciliate; ovary ovoid, stigmas apical. Known only from the following collection. CHiRiQuf: Llanos del Volcan, alt. about 1300 m., Alleri Tjjo, This species closely resembles P. /jumorigauJens but differs because of the shape of the 5-nerved, more attenuately acuminate leaves, subhirtellous twigs and petioles, and ciliate margins of the petiole vaginal groove. 24. Pn^ER PiNOGANENSE Trel. in Contr. U, S. Nat. Herb. 26:22. 1927. Piper tabasaramivi Trel. in Ann. Mo. Bot. Gard. 25:827. 1938. Viper salamauca}ium Trcl. loc. cit. 27:196. 1940. A shrub or small tree, 1.5-4 m. tall, nodose; intcrnodes slender, rather short, minutely and transiently pubcrulcnt, finely ribbed; leaves elliptic, apex caudatcly blunt-acuminatc, base rounded but subacute at the petiole, or narrowed and acute, 3-5.5 cm. wide X 8-12 cm. long, palmately 5-nerved, the nerves salient on both sides, the innermost pair of laterals coalescing with the midrib within the lowermost 5 mm., with numerous, fine, cross-connecting and anastomosing nervules, drying light green, firm, chartaceous, glossy, translucent; petiole scarcely 5 mm. long, granular-puberulcnt, channelled above, drying wrinkled, vaginate toward the base; spikes 3-4.5 cm. long, loosely flowered; peduncle scarcely 1 cm. long, obscurely puberulent; bracts very small, concave, glabrous; fruit ovoid, smooth or papillate; stigmas 3 or 4, sessile, Panama. CHiRjQui: banks of lower Rio Tabasara, WooJson, Alien & Seiberi 440, canal zone: vicinity of Salamanca Hydrographic Station, Rio Pequcni, alt. 80 m., Woochoti, Allen ^ Seibert IjQO. darien: around Pinogana, Pi f tier 66 j6, 25. Piper medium Jacq. Icon. Pi. Rar. 1:2. 1781. ■ Piper pliuifagineum Lam. Illustr. Encyc. Mcth. 1:80. 1791. Piper saururoides Ham. Prodr. Pi. Ind. Occ. 3. 1825. Piper reticiilatum Veil. Fl. Flum. 1: pi. 61. 1827, not L. 1753. Piper syringaefolium Balbis ex Kunth, in Linnaea 13:599. 1839. Piper orthostacbynm Kunth, loc. cit. 602. 1839. Enckea plaufagineu Kunth, loc. cit. 599. 1839. Enckea orthosfachya Kunth, loc. cit. 601. 1839. Enckea Sieberi Miq. Syst. Pip. 3 5 8. 1843. Piper victoriannm CDC. in Jour. Bot. 4:218. 1866. Piper figeriauum CDC. loc. cit. 3 00. 18 66. Piper Sieberi CDC in DC Prodr. 16^:248. 1869. A shrub; stems and branches glabrous or slightly puberulent when young; leaves lance- to round-elliptic or subrhombically obovate, 3.5-6 cm. wide X 8-11 cm. long, or rarely occasional leaves up to 10 X H cm., apex acuminate, base sometimes suboblique, acute or somewhat rounded, more or less subaunculate at (^9) [Vol. 37 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN the very base, or lowermost somewhat cordate, glabrous or pubcrulent along the nerves beneath, palmatcly 5- to 7-ncrved, reticulatc-ncrvulose, dr)Mng membranous and minutely pclIucid-dotted; petiole 5-10 mm. long, vaginate near tlie base, pubcrulent; spikes 4-5 mm. thick X 6-7 cm. long; peduncle 10-15 mm. long, pubcrulent or glabrous; rachis puberulent; bracts obovate-concavc, somewhat papillate-pilose; fruit ovoid, minutely papillate or glabrous; stigmas 3 or 4, rounded, scssi ile. West Indies, Central and South America. CANAL zone: forest along banks of Quebrada Fea, Q. Pura and Canon of Rio Chagres, alt. 70-100 ni., Dodge ^ Hvnfer 17404. chiriqui: Cerro Galera Chorcha, vicinity of Gualaca, alt. 3 00 m., Allen 5060, This species closely resembles P. pinogancfne but differs primarily because of its more membranous leaves and hairy rachis and floral bracts. ITT. KKY TO SPECIES WITH LEAVES MOSTLY 20 CM. OR MORE LONG AND MORE OR LESS LOBED AT THE BASE (The base may be only slightly lobed but none arc rounded or acute; the petiole mostly vaginate- wingcd to the middle or above.) a. Leaves rugose or bullate. b. Leaves scabrid 4L P. peracuminatum bb. Leaves not scabrid. C. Leaves loosely long-villous; peduncle 1-2 cm. long 26. P. rARTiTlCUSPE cc. Leaves hirtcllous to pubescent; peduncle 1-2.5 cm. or more long.... 27. P. Coopfri aa. Leaves scarcely rugose or bullate. b. Leaves and stems glabrous - 28. P. hastularum bb. Leaves and/or stems pubescent. c. Leaves scabrous 41. P. peracuminatum cc. Leaves not scabrous. d. Leaf margin densely clliolate, drying thin; spikes scarcely 5 mm. thick, yellow when dry 29. P. auritum dd. Leaf margin glabrous or essentially so. c. Young twigs, nodes and/or petioles and midrib beneath more or less fleshy-warty 30. P. imperiale ee. Not fleshy-warty. f. Stems matted brown-hairy, leaves 5- to 7-nerved. g. Leaves acute, basal sinus wide, essentially glabrous above.... 31. P. tardens g^. Leaves acuminate; basal sinus narrow, nerves pubescent above 32. P. campanum ff. Stems glabrous; leaves mostly 7- to 9-nerved. g. Leaves glabrous above; petiole vaginate below the middle; bracts glabrate 33. P. carrilloanum gg. Leaves more or less hairy above; petiole vaginate to the blade; bracts fringed 34. P. Pittieri 26. Piper partiticuspr TreL in Ann. Mo. Bot. Gard, 27:294. 1940. Suffruticosc, 3 m. tall; internodcs comparatively slender and elongate, for a time crisp-villous; leaves round-ovate, apex falcately acuminate, base inequilaterally cordate with one side surpassing the petiole, sinus open, 15-28 cm. wide X 20-3 5 cm. long, pinnately nerved from the lower half, the nerves 6-8 on each side, with prominent cross-connecting and anastomosing nervules, venosc-buUulate with 2-4 (20) 1950] FLORA OF PANAMA (Pipcraccac) 21 points to each cusp, the cusp points and the midrib near the base above long-villous, the nerves beneath loosely long-villous, drying firm, translucent; petiole 3-4 cm. long, densely villous; spikes 5 mm. thick X 15 cm. long; peduncle 1-2 cm. long, loosely villous; bracts triangular-subpeltate, slightly ciliate; fruit small, sub- trigonous, glabrous; stigmas small, sessile. Known only from the one locality. cocLE: vicinity of El Vallc, alt. 600-1000 m., Allen II95. The villous, round-ovate leaves with multipointed, villous, bullulate elevations characterize this species. 27. Piper Cooperi Yuncker, sp. nov. Frutex vel arbuscula, 5 m. altus; internodiis florifcris sat crassiusculis elongatis- que conferte crispe-pubescentibus; foliis elliptico-subovatis rugosis, apicc breviter subacuminatis, basi oblique cordatis, latere longiore petiolum excedente, infra medium pinnatim venosis, venis utrlnque 6-7, laminae facie supra hirtella Infra pubesccnte (ad vcnas quam maxime) ; petiolo 4-5 cm. longo, ad medium vel ultra vaginato-alato, conferte crlspe pubesccnte; spicis immaturis 3 mm. crassis, 6 cm. longis; pedunculo 2.5-4 cm. longo, pubescente; bractels rotundato-subpeltatis, atro-pubescentibus. A shrub or small tree, 5 m. tall; stem 8 cm. thick, with clustered basal shoots; flowering internodes moderately stout and elongated, matted-crisp-pubescent; leaves elliptic-subovate, apex shortly subacuminate, base obliquely cordate, the longer lobe surpassing the petiole, 15-20 cm. wide X 16-3 cm. long, pinnately nerved from the lower half, the nerves 6 or 7 on each side, with cross-connecting and anastomosing nervules, hirtellous above, at least along the nerves, pubescent beneath, the nerves densely so, bullate, drying dark, firm, opaque; petiole 4-5 cm, long, vaginate-winged to the middle or above, densely crisp-pubescent; spikes as yet ypung, 3 mm. thick X 6 cm. long; peduncle 2.5-4 cm. long, pubescent; bracts round-subpeltate, dingy-hairy. Known only from the type locality. BOCAS DEL TORO: Vicinity of Almirante, Daytonia Farm, Jan. -Mar. 1928, G. Proctor Cooper 374 (U. S. Nat. Herb., type). This species differs from P. partitictispc^ which it resembles somewhat, because of the character of the pubescence, longer peduncles, pubescent bracts, etc. 28. Piper hastularum Yuncker, sp. nov. Frutex primo intuitu glaber, ligno moHi, 1-3 m. altus; internodiis floriferis elongatis crassiusculisque; foliis lanceolato-ovatis vel late lanceolatis, 6-11 cm. latis, 22-29 cm. longis, apice sensim acutis acuminatisve, basi oblique inaequilateral- ibus, ad petiolum utrinque obtusis vel latere altero obtuso acutove ad petiolum 3-10 mm. breviore, caetero rotundato-obtuso subcordatove, a tertio supero pin- natim venosis, venis utrinque 4-5, venulisque submarginalibus gracilibus puberulis, subtus prominentibus trabibus anastomosantibus manifestis, arctius revolutis. (21) ♦ '. ' '^*j [Vol. 37 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN opacis; peciolo 20-25 cum 3-10 mm. longo, ad laminam vaginato-alato; spiels 3-4 mm. crassis, 13 cm, longis apiculatis; pcdunculo 10-20 mm. longo, gracili; bracteis triangulari-sublunatis margine tantum papillaris; drupa laevi, subglobosa, truncata; scigmatibus 3, linearibus, sessilibus. An essentially glabrous, soft-wooded shrub, 1 3 m. tall; flowering internodcs moderately slender to stoutish; leaves lance-ovate or broadly lanceolate, apex gradually sharp-acute to acuminate, base ob- liquely inequilateral, one side 3-10 mm. shorter at the petiole, both sides obtuse at the petiole or the shorter side subacute, the longer sometimes sub- cordate, 6-11 cm. wide X 22-29 cm. long, pin- natcly nerved below the upper third, the nerves 4-5 on each side, somewhat raised beneath, with atl rather strong cross-connectmg nervuies ul » th e s ub- marginal nervule m.ore or less papillate or pubcru- lent, drving chart aceous-subcoriaceous, opaque, somewhat glossy, narrowly revolutc, glandular- dotted and sometimes microscopically black-mottled beneath; petiole 20-25 plus 3-10 mm. long, strongly vaginate-winged to the blade; spikes 3-4 mm. thick X 13 cm. long; peduncle slender, about 1-2 cm. Fig. 5. Pi[)cr hasfidarum long; bracts narrowly triangular- or lunate-subpcltate, smooth or papillate, the pedicel broad, concave, glabrous or ciliate; fruit truncate subglobose, glabrous; stigmas sessile. Western Colombia and Panama. Colombia: choco: Bahia Solano, dense forest along Quebrada Jcllira, alt. 50-100 m., Feb. 22, 1939, Killip 6 Garcia 33368 (U. S. Nat. Herb., type). Panama: bocas del toro: Old Bank Island, von Wcdel 1934- cocle: El Valle de Ant6n, along Rio Indio Trail, alt. 500-700 m., llunfcr & Allen 316, oarien: Chcplgana District, Cana-Cuasi Trail (Camp 2), alt. 1200 m., March 10, 1940, Terry ^ Terry 1461. The lanceolate leaves and submarginal puberulent nervuies distinguish this species. 29. Piper auritum H.B.K. Nov. Gen. & Sp. 1:54. 1815. Schiller'ia aiiriia Kunth, in Linnaca 13:713. 1839. Arlauthe aur'ita Miq. Syst. Pip. 400, 1844. Art ant he Secmauuiana Miq. in Seem, Bot. Voy. Herald, 199, pL 39. 18 54, not Piper Seetnafinlanum CDC. Viper auritum var. amplifoliutn CDC. in DC. Prod. 16':321. 1869. Piper auritum var. Seemannianum (Miq.) Trel. in Contr. U. S. Nat. Herb. 26:40. 1927. Piper heraIJi Trcl. in Ann. Mo. Bot. Card. 27:292. 1940. Piper heraldi var. ampUus Trcl. loc. cit. 1940. (22) 19 5 0] FLORA OF PANAMA (Pipcraceac) 23 Piper heraldi var. coclcannm Trel. loc. cit. 1940. Fiper Alstoni Trel. loc. cit. 286. 1940. A small, soft-wooded, commonly more or less aromatic tree, 3-6 m, tall, often occurring in clumps; flowering intcrnodes moderately slender and long, striate, often drying black, glabrate or sometimes slightly pubescent; leaves ovate, oblong- or elliptic-ovate, 12-20 cm. wide X 20-30 cm. long, or occasionally up to 25 X 40 cm. or more, apex acute or short-acuminate, base deeply cordate with one side 1-2 cm. longer at the petiole and with the lower lobe longer, sinus rounded to the petiole, the midrib with 3 or 4 upcurved branches from below the upper third and 3 or 4 branches from the base, rather thinly short-hairy above at least along the nerves, more densely so beneath, densely white-cillolate throughout, drying thin, translucent; petiole 4-9 plus 1-2 cm. long, more or less pubescent or glabrate, vaginate-winged to the blade; spikes creamy or hght-yellow when dry, 3-5 mm. thick X 10-25 cm. long; peduncle slender, 2-8 cm, long, glabrate, often black when dry; bracts round- or triangular-subpeltate, marginally fringed; fruit small, glabrate, obpyramidal-trigonous; stigmas 3, sessile. West Indies and on the mainland from Mexico through Central America and northern South America. BOCAS DEL TORO: Changuitiola Valley, Dunlap 40J; Laguna dc Chiriqui, Harf 126; vicinity of Nicvcclta, alt. 0-50 m., ^^ooiUon, Allen & Seibert /5/J; vicinity of Chiriqui Lagoon, von Wcdcl J 194. chiriqui: Finca Lerida to Boquete, alt. 1300-1700 m., Wood- son, Allen & Seibert TOQj; Ccrro dc la Plata, near San Felix, alt. 120-150 m., Pittier 5159; vicinity of David, alt. 30-80 m., Piffier ^829. cocle: Penonomc, Williams 4^4; north rim of El Valle, Alston & Allen 1 842; vicinity of El Valle, alt. 600-1000 m., Allen 1 192, CANAL zone: Barro Colorado Island, Bailey & Bailey 2J; Bangham 59O; Monkey Fiill, Lebmann IOO4; Cowell 22; between Chagrcs Batteries and Fort San Lorenzo, Fort Sherman Military Reservation, Maxon 6 Valen/ine 698J; Rio Agua Salud, near Frijolcs, Piper 5844; between Gorgona and Gatiin, alt. 10-50 m., Pittier 226j; Gatiin, Sfandley 2/^244; Stevens 833; Riley loS; Fort Sherman, Standley 30934; Stevens 263; headwaters of the Rio Chinilla, above Nucvo LImon, Maxon 6898; forests of northerly arm of Qucbrada Sala- manca as far as falls, alt. 70 m., Steyermark 6 Allen 17140. colon: ''Aspinwall," Hayes S351 around Porto Bello, alt. 5-100 m., Pittier 2441 ; beach between Fato and Playa de Damas, Pittier 3933; along Rio Fato, alt. 10-100 m., Pittier 3909; vicinity of Santa Isabel, Pittier 4177; Puerto Obaldia, San Bias coast, alt. 0-50 m., Pittier 4380. Panama: Man- zanillo Island, Ihiyes 40, 706; Chararc River, near Chepo, alt. 100-200 m., Pittier 4709; vicinity of Arcnoso, lower Rio Trinidad, alt. 26-50 m., Seibert 623, daiuln: Boca de Cupe, Williams 785; headwaters Rio Chico, alt. 150-225 m., Allen 4603, The thin, deeply and incquilaterally cordate leaves more or less pubescent on both sides and densely ciliolate, long, blackening peduncles, and comparatively small, yellowish spikes distinguish this species. 30. Piper imperiale (Miq.) CDC. in DC. Prodr. 16^339. 1869. Artantbe imperialis Mlq. in Seem. Bot. Voy. Jierald, 198. 1854. Piper portobellense CDC. Smiths. Misc. Coll, 71*':6. 1920. Piper affectans Trel. In Ann. Mo. Bot. Card. 27:287. 1940. Piper frijolesanum Trel. var. grandifoliiim Trel. loc. cit. 291. 1940. Piper gigas Trel. loc. cit. 292. 1940. A shrub or small tree, commonly 2-5 m. tall; flowering intcrnodes rather elongate and stoutish, commonly somewhat fleshy-warty especially near the nodes, (23) [Vol. Z7 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN pubescent or glabrescent; leaves broadly ovate, apex acute or short-acuminate, base somewhat obUquely cordulate with a rather narrow sinus, both sides essentially equal at the petiole or one side a few mm. shorter, up to 29 cm. wide X 50 cm. long but usually somewhat smaller, pinnately nerved below the upper third or fourth, the nerves 7-8 on each side, the lowermost 3 or 4 approximated near the base, with cross-connecting and anastomosing ncrvules, submarginally loop-con- nected, glabrous above, pubescent beneath, at least the nerves puberulent, the midrib beneath with warty excrescences at least near the base, drying firm, brown, opaque; petiole 6-8 or 12 cm. long, puberulent, vaginate-winged nearly to the blade, with warty-excrescences; spikes 8-10 mm. thick X 30-50 cm. or more long; peduncle 4-6 cm. or more long, sparsely hairj' glabrescent; bracts rounded- or triangular-subpeltate, inflexed, yellow-puberulcnt or glabrate; fruit rather large, obovold, glabrous, stigmas linear, on a short, stout, deciduous style. Panama. CHiRiQUi: vicinity of Casita Alta, Volcan de Chiriqui, alt. 1500-2000 m., Woochof?, Allen & Seibcrt 846, 86 j; Boquete District, Bajo Chorro, alt. 1800 m., Davidson 374^ cocle: vicinity of E! Valle, alt. 600-1000 m., Allen //pj. canal zone: between Frijoles and Monte LIrio, alt. 30 m., Killil) T2ljj; hills north of Frijolcs, StanJley 2744T; Barro Colorado Island, StanJley 3 12/ J, 4^TI^, colon: forests around Porto Bcllo, alt. 5-200 m., Maxon S795' Panama: Taboga Island, Nee 350. dari^n; Chcpigana Distr., Cana-CuasI Trail, alt. 600 m., Terry ^ Terry T^OO. The types of ?. affccfans and ?. friplesamim var. grandijoliuvi are less warty and the stems arc more glabrous than with the other specimens cited, while the young growth of the type of P. gigd^ is somewhat more densely pubescent. The type specimens of both P. affccfans and P. gigas are stated as being trees and with a trunk 3 cm. in diameter. No other differences have been noted in the herbarium specimens, and it is believed that they all represent variations of the same species. So far as discovered, P. frijolcsauum is a novtcn nudum as used for var. grandifolhirn, 31. Piper tardens Trel. in Ann. Mo. Bot. Gard. 27:298. 1940. A shrub or small tree, 1-5 m. tall; flowering internodes rather slender and elongated, densely brown-tomentose-pubescent ; leaves pentagonally ovate, apex acute, base incquilatcrally oblique, rather shallowly and openly cordate, 10-16 cm. wide X 17-28 cm. long, pinnately nerved from the lower half, the nerves 5~6 on each side, salient beneath, with rather prominent cross-connecting and anastomosing ncrvules, glabrous above or the midrib sparsely pubescent near the base, hirsute beneath, the nerves densely so, drying coriaceous, opaque; petiole 3-5 cm. long, vaginate-winged to the blade, densely tomentose-pubescent; spikes 5-8 mm. thick X 25 cm. long; peduncle 2.5-4 cm. long, densely hairy; bracts rounded- or tri- angular-subpeltate; pedicel brown-villous; fruit globose, stigmas sessile or on a short style (?). Chiriqui, Panama. CHiRiQuf: vicinity of Casita Alta, Volcan de Chiriqui, alt, 1500-2000 m,, Woodson, Allen ^ Seibert 84S; Boquete Distr., Bajo Chorro, alt. 1800 m., Davidson j8^ 3/3, (24) 1950] FLORA OF PANAMA (Piperaccae) 25 The densely matted twigs, pentagonally-ovate leaves, glabrous above and hirsute beneath, are characteristic of this species. From the material at hand it has not been possible to determine with certainty regarding the presence of a style. Most of the fruit examined do not exhibit any but in a few there appeared to be a short, thick style. This condition, however, may have resulted from the manner In v/hich the fruit dried and may not represent a true style* 32. Piper campanum Yuncker, sp. nov. Frutex 2 m. altus, internodiis supcris gracilibus elongatis, conferte brunneo- pubcscentibus; foliis lanccolato-ovatis, apice sat attenuatim acuminatis, basi cordato- auriculatis, latere altero 5 mm. breviore, sinu obliquo, a tertio supero deorsum pinnatim venosis, venis utrinque 6-7, utrinque ad venas pubescentibus, glanduloso- punctatis; pctiolo 2.5-4.5 cm. cum 2-5 mm. longo, conferte brunneo-pubescente, ad laminam vaginato-alato; spicis immaturis, 1 mm. crassis, 5 cm. longis; pcdunculo gracili, conferte pubescente. A shrub, 2 m. tall; upper internodes slender and elongate, densely brown- matted-hairy; leaves lance-ovate, apex somewhat attenuately acuminate, base cordate-auriculate, one side up to 5 mm. shorter at the petiole, the sinus lateral, 11-12 cm. wide X 26 cm. long, or upper leaves smaller, pinnately nerved below the upper third, the nerves 6-7 on each side, branched upward to some extent, with cross-connecting and anastomosing nervules, the nerves pubescent on both sides or only toward the base on the upper surface, drying rather thin, subtranslucent, petiole 2,5-4.5 plus 2-5 mm. long, densely brown-hairy, nate-wmged to the blade; spikes as yet very young, 1 mm. thick X 5 cm. long; peduncle slender, 1 cm. long, densely hairy; bracts subpeltate, bristly. Known only from the type locality. PANAMA: Ccrro Campana, vicinity of Campana, alt. 1000 m., April 21, 1941, Allen 241 1 (U. S. Nat. Herb., type). This species resembles P. tardcns to some extent but differs because of the more acuminate leaf apex, cordate-auriculate base, and more pubescent nerves above. 33. Piper carrilloanum CDC. in Bull Soc. Roy. Bot. Belg. 30^:209. 1891. Piper vallicolum CDC. loc. cit. 222. 1891. Piper paiilowni folium CDC. In Anal. Inst. Fis. Geogr. Costa Rica 9:173. 1898. Piper tiliaefolium Cham. & Schlechtd. var. paulownifolium CDC in Candollea 1:154 1923. Piper Omega Trel. in Contr. U. S. Nat. Herb. 26:146. 1929. A shrub, 2-4 m. tall, glabrous except the puberulous nerves on the lower surface; upper internodes moderately slender and elongated; leaves subequilaterally ovate, apex rather long-acuminate, base subtruncate to moderately deeply cordate with open sinus, the blade margin commonly meeting over the petiole, rather small and elongate, 6-8 cm. wide X 15 cm. long or becoming more rounded and as much as 2 5 X 3 5 cm., pinnately nerved to about the middle, 7-9 nerves on each side, the lowermost 3-5 from near the base, salient beneath, with prominent cross- connecting and anastomosing nervules, very finely nervulose by transmitted light, (25) [Vol. ^7 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN drying light grcciij papery, translucent; petiole slender, 2-5 or even 10 cm. long, vaginate winged below the middle; spikes about 3 mm. thick X 15-2 5 cm. long; peduncle 1-2 cm. long; bracts small, triangular-subpcltate, glabratc; fruit small, obpyramidal-trigonous, gray-papillate, stigmas small, sessile. Costa Rica and Panama. cHiRiQuf: Cerro de la Plata, near San FcIIx, ale. 120-150 m., Pilficr 5^5^'^ '^^'^^ de Qucbrada Gata, between Hato Jobo and San Felix, Vliiier 54^7- canal zone: lillls north of Frljoles, StauJJcy 27612; hills west of the canal, near Gatun, Sfamilcy 2J236; Mamci Hill, alt. 20-90 m., Pitticr 3812; along Cano Quebrado, Pifticr 6666; Barro Colorado Island, Miller 2073; Maxon, Hancy & Vulcutiue 6Sog; Staudley J/2^^, 3^^93^ 40^7^^^ 40964; Bailey ^ Bailey 30; Sferefis 645; Pearson Trail, alt. 100 m., Wetmore ^ Abbe 123; between Gorgona and Gatun, alt. 10-50 m., P/7/;>r 226r. colon: Fato (Nombrc de Dios), P/7//Vr 3S44. dariln: Boca de Pauarando, on Sambu River, alt. 20 m., P/7/;Vr J^i?/; Chcpis^ana DIstr., Cana-Cuasi Trail, alt. 600 m., Terry 6 Terry 1475, This species closely resembles P. casimiriannm HcmsL of Nicaragua from Avbich It appears to differ because of the leaf shape and pubcrulence, as described. It is P T! :ie great variation in the size of the leaves often on the same specimen, leaf shape, puberulent nerves beneath, etc. are distinguishing characteristics. 34. PiPLK riTTTFRi CDC. in Bull. Soc. Bot. Bclg. 29':69. 1890; 3(V:204. 1891. A soft shrub, 1 m. or more tall; flowering internodcs rather thick and elongated, glabrous, glandular-dotted; leaves broadly ovate, apex abruptly acute or sbort- acumlnatc, base truncately shallow-cordate, 14-16 cm. wide X 15-22 cm. long, pinnately nerved from below the upper third, the nerves about 8 on each side, gradually approximated downward, submarglnally loop-connected, with rather prominent cross-connectlng-anastomosing nervules, thinly pubescent above, the nerves heneatb rather densely villous laterally, ciliate, strongly glandular-dotted on both sides, drying rather thin but firm, translucent; petiole about 6 cm. long, vaginate- winged to the blade, glabrous, glandular-dotted; spikes 5 mm. thick X 15 cm. long or in fruit up to 12 mm. thick X 18 cm. long; peduncle stout, 1.5-2.5 cm. long, puberulent glabrescent, glandular-dotted; bracts triangular-subpeltatc, narrowly fringed; fruit angularly oblong, with pointed, substylose apex, stigmas 3. Costa Rica and adjacent Panama. BOCAs DEL TORO: Robalo Trail, northern slopes of Cerro Horqucta, alt. 1800-2100 m., Allen 3012. The large, somewhat pubescent leaves, rather large spikes, and substylose fruits are distinctive characters of this species. IV. KEY TO SPECIES PINNATELY NERVED FROM SOME^X^F^AT BEI.OW THE MIDDLE TO THE UPPER THIRD OF THE BLADE AND >XTTH THE LEAVES MOSTLY LESS THAN 20 CM. LONG; IF LONGER, THE BASE IS NOT LOBED. a. Leaves scabrous. b. Twigs glabrous. c. Leaves glabrous on both sides, scabrous beneath, smooth abtjvc 35. P. davioanum cc. Leaves hairy beneath, at least along the nerves. d. Leaves mostly 3-4.5 cm. wide, strongly granular-dotted beneath.. 36. P. MiNUxr- SCABIOSUM (26) 19 SO] FLORA OF PANAMA (Pipcraceae) 17 dd. Leaves 5-6 cm. wide, scarcely granular -dotted beneath 52. P, Storkii bb. Twigs more or less hairy, at least when young. c. Spikes strongly and regularly curved; peduncle longer than the petiole.. - 37. P. aduncum cc. Spikes not strongly or regularly curved. d. Stem hairs stiff-hispid, comparatively short; leaves liarshly scabrous; fruit papillate-puberulcnt at apex,„. 38. P. HisriDUM dd. Stem hairs rctrorse; leaves finely scabrous. e. Plant scandent; leaf base obtuse or acutish 39. P. vitabundum ee. Plant shrubby; leaf base rounded, cordulate 78a, P. Wedelii var. RETROHTRSUTUM ddd. Stem more softly pubescent or villous. e. Hairs on the midrib beneath erect or at least not upcurved- strlgose. f. Hairs on midrib beneath mostly up to 1 mm. or more long, g. Leaves mostly 18-20 cm. or more long. h. Leaves 6-8 cm. wide, very slenderly acuminate, base obtuse or cordulate 40. P. persubulatum hh. Leaves mostly 1 cm. or more wide, not slenderly acuminate, base rounded-cordate 41. P. peracuminatum ^'^. Leaves scarcely up to 15 cm. long. h. Leaves broadly elliptic, less than two times longer than wide 73. P. cativalense hh. Leaves 2—3 times longer than wide. i. Leaves rhomboid-elliptic, strongly inequilateral; bracts glabrous 42. P. erubescenti- SPICUM ii. Leaves lance-elliptic, elliptic, or elliptic-obovate. j. Petioles scarcely 5 mm. long; stem hairs scarcely 1 mm. long; blades slightly scabrous.. 72. P. pseudo- garagaranum ]). Petioles up to 2 cm. long downward. k. Blades mostly 5—8 cm. wide, scarcely scabrous above _ 43. P. culebranum kk. Blades 3-4.5 cm. wide, harshly scabrous above.. 44. P. viridicaule ff. Hairs on midrib mostly scarcely 0.5 mm. long, dense. g. Leaves less than 5 cm. wide; veins above impressed In ago.. 45. P. pervelutinum gg. Leaves 6-8 cm. wide; veins not impressed 46. P. variitrichum ee. Hairs on the midrib beneath strongly upcurved and appressed or strigose. f. Leaves less than 4 cm. wide 47. P. changuinolanum ff. Leaves mostly wider, g. Blades moderately to densely hairy on the upper surface, h. Peduncle scarcely 5 mm. long; blades villous above and harshly scabrous; bracts of young spikes not conspic- uously bristly 48. P. vieliramui.um hh. Peduncle 5-10 mm. or more long; blades crisp-hirtellous or subhispid above, slightly scabrous; young spikes very short, conspicuously bristly from erect marginal hairs. i. Leaves mostly narrowed downward, usually only the longer side rounded at the base , 49. P. taboganum il. Leaves rounded-cordulate on both sides .. 50. P. pseudo- cativalense gg. Blades sparsely hairy or glabrous, hairy mostly only along the nerves above. h. Young spikes not conspicuously bristly because of the erect bract hairs. i. Bracts prominently densely yellow-fringed 70. P. leptocladum ii. Bracts very narrowly and sparsely fringed. j. Nerves beneath sprcadlng-hlrsute 51. P. euctgaudens jj. Nerves beneath obscurely strigose 52. P. Storkii iii. Bracts glabrous, with a dorsal callus 78. P. Wedelii hh. Young spikes about 1 cm. long, conspicuously bristly because of erect bract hairs 49. P. taboganum {27) [Vol. Z7 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN aa. Leaves not noticeably scabrous. b. Spikes short, scarcely exceeding 2 cm. in length. c. Stems and leaves glabrous. d. Spikes less than I cm. long; bracts glabrous 53. P. Wagnlri dd. Spikes about 2-3 cm. long; bracts densely ciliate 54. P. Dunlapi CO. Stems and/or leaves hairy. d. Stems glabrous or sparsely pubescent; nerves beneath appresscd- puberulcnt 5 5. P. palmasanum dd. Stems rather densely hairy, at least when young 16. P. colon-insulae bb. Spikes longer, mostly 3-5 cm. or more long (or unknown). c. Stems and leaves essentially glabrous (nerves hircellous or puberu- lous in P. carriUoamim and P. acqualc var. lanrifolium) . d. Peduncle 4-7 cm. long .56. P. reptabundum dd. Peduncle scarcely 2 cm. long at most. e. Leaves ovate. f. Base mostly acute at the petiole; petiole 1-2.5 cm. long 57. P. chirtquinum ff. Base obtuse to cordate. g. Leaves 10-14 cm. long; petiole 5 mm. long 65. P. tecumiznse gg. Leaves 15 cm. or more long; petiole 1-2 cm. or more long. h. Nerves beneath pubcrulous; peduncle 1-2 cm. long 3 3. P. carriixoanum hh. Nerves beneath glabrous; peduncle 0.5—1 cm. long 93. P. grande ce. Leaves elliptic, lance-elliptic, subobovate or obovatc. f. Leaves mostly 2.5—4.5 cm. wide (rarely wider); apex sharp- to strongly mucronatc-acuminate. g. Leaves scarcely more than 2-2/2 times longer than wide. h. Base acutely dccurrcnt, apex strongly mucronatc 58. P. tfnuimucro- NATUM hh. Base rounded, obtuse 59. P. septuplinervium gg. Leaves mostly 3—4 times longer than wide, or base not decurrent, h. Bracts smooth; leaves not glandular. L Leaf base rounded, obtuse; nerves 4-5 on each side..-. 59. P. septuplinervium ii. Leaf base narrowed, acute to cuneate; nerves 3—4 on each side 60. P. subquinquenerve hh. Bracts conspicuously yellow-fringed; leaves more or less glandular-dotted. i. Leaves drying dull, yellow 61. P. infraluteum ii. Leaves drying glossy, green 80b. P. utrteelipetioeum var. Harveyanum ff. Leaves larger, mostly 5 cm. or more wide. g. Leaves nerved from about the lower third, sharply ab- ruptly acuminate 62. P, cricamolense gg. Leaves nerved to about or above the middle. h. Leaves mostly 4- to 5-nervcd on each side of the mid- rib (or sometimes 6 in P. aequale). i. Leaves obovate, revolute, apex mostly obtuse 10. P. macropunctatum ii. Leaves oblong-elliptic-subobovate, apex abruptly blunt-acuminace 63. P. aequale iii. Leaves elliptic, apex gradually acuminate. j. Leaves, petioles, etc. strongly glandular-dotted 64. P. SAMBUANUM jj. Leaves, etc. not glandular-dotted 65. P. tecumense hh. Leaves 7-nervcd on each side of the midrib, oblong- ovate, 10-13 cm. wide... 66. P. crassispicatum cc. Stems and/or leaves more or less hairy (minutely and obscurely so in P. caUibracfmm) , d. Leaves nerved from the lowermost y^ or Y^ 67. P. bogotense dd. Leaves nerved mostly to about the middle or above. e. Leaf base with one side strongly auriculate and covering the petiole (auricle about 1 cm. or more long), f. Pubescence strongly dimorphic; leaves 6 cm. or more wide.... 68. P. sperdinum ff. Pubescence not dimorphic; leaves less than 5 cm. wide 69. P. pi rbrevicaui E ee. Leaf base may be cordulatc but scarcely auriculate. f. Leaves hairy on the upper surface, at least along the mid- rib above the very base. (2^) 1950] FLORA OF PANAMA (Piperaceae) 29 g. Leaves 10 X 25 cm. or more 85. P. colonense gg. Leaves mostly smaller. h. Pubescence on the upper leaf surface restricted to the nerves. i. Nerves beneath glabrous; spike 2-5 cm. long 59a. P. septuplinervium Var. SUBPUBINERVIUM ii. Nerves beneath obviously hairy 70. P. leptocladum hh. Pubescence not so restricted. i. Leaves 6—7 cm. wide, scarcely 3 times longer than wide; spikes straight. j. Crisp-hirtellous or subhispid above; young spikes short, bristly 49. P. taboganum jj. Sparsely long- villous above, glabrescent; young spikes not noticeably bristly, k. Peduncle up to 6 cm. long; stem and leaf hairs strongly dimorphic 71. P. dimorphotrichum kk. Peduncle much shorter; hairs not obviously dimorphic. 1. Apex gradually acuminate, blades 15—20 cm. long 72. P. fseudo- GARAGARANUM 11. Apex abruptly acuminate, blades 12-14 cm. long 73. P. CATIVALENSE ii. Leaves mostly narrower, 3—4 times or more longer than v/idc, rugcsccnt, appresscd-pubescent above; spikes commonly curved. J. Nerves mostly 7-8 on each side; leaves mostly 3-5 cm. or more wide 74. P. pseudo- lance aefolium jj. Nerves mostly 4—5 on each side; leaves mostly 2—3.5 cm. wide 75. P. linearifolium ff. Leaves glabrous above. g. Fruit and ovary papillate-pubescent, h. Petioles scarcely 10 mm. long, i. Spikes 2—3 mm. X 7—8 cm.; plant vine-like. 3 9. P. vitabundum ii. Spikes 3-5 mm. X 4-5 cm.; plant shrubby 76. P. panamense hh. Petioles up to 3 cm. long; spikes 8—10 cm. long 77, P. callibracteum gg. Fruit glabrous, or unknown. h. Stems glabrous or essentially so. i. Leaf nerves beneath thinly strigose; bracts glabrous.... 78. P. Wedelii ii. Leaf nerves beneath obscurely puberulent; bracts fringed. j. Leaves 7-10 cm. wide .79. P, verruculae- PETIOLUM jj. Leaves less than 6 cm. wide 65. P. TECUMENSE hh. Stems obviously pubescent, at least when young. i. Leaf base narrowed, acute or obtusish, the sides nearly equal. j. Nerves beneath glabrous; spikes about 4-5 cm, long 80. P. HIRTELLIPETIOLUM jj. Nerves beneath pubescent; spikes 5-6 cm. long.... 80a. P. iitrtelltpetiolum var. TAPIANUM ii. Leaf base with one or both sides rounded-obtuse, or the longer side mostly cordulate. j. Bracts conspicuou<;ly fringed; spikes 3-4 mm. thick. k. Lateral nerves loop-connected upward, not con- tinuing toward the apex 81. P. subcaudatum kk. Lateral nerves not strongly loop-connected up- ward, continuing unbroken toward the apex 43. P. culebranum jj. Bracts essentially glabrous; spikes 7—8 mm. thick.... 82. P. gonocarpum (29) [Vol. Z7 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN 35. Piper davidanum CDC. in Smiths. Misc. Coll. 7V:9. 1920. A shrub, 2 m. tall, nodose; flowering intcrnodcs moderately slender, glabrous, granular, with subnodal lenticels; leaves elliptic, apex rather short-acuminate, base inequilateral, one side 4—5 mm. shorter and acute, the longer side rounded, 6-9 cm. wide X 16-19 cm. long, glabrous on both sides, finely scabrous and glandular- dotted beneath, smooth above, pinnately nerved below the upper third, the nerves rather prominent beneath, 5—6 on each side, with slender cross-connecting and anastomosing nervules, drying pale green, membranous, translucent; petiole 6 plus 4-5 mm. long, vaglnate at the base, glabrous; spikes 3 mm. thick X 8-10 cm. long; peduncle 5-7 mm. long, glabrous; bracts triangular-lunate-subpeltate, sparsely papillate-fringed; fruit oblong-tctragonous, granulate-papillate at the top, stigmas sessile. Known only from the following locality. cHiRiQui: vicinity of David, along streamlet, alt. 30-80 m., Pifiicr 2Sj/, The glabrous stems and leaves, and scabrous lower leaf surface characterize this species. 36, Piper minute-scabtosum Trel. in Ann. Mo. Bot, Card. 27:294. 1940. Piper mifiuic-scubiosuni var. arborcscens Trel. loc. clt. 1940. A shrub or small tree, up to 3 m. tall, somewhat twiggy, nodose; flowering internodes slender, Icpidote-granular, glabrous; leaves narrowly elliptic or lance- elliptic, apex caudately acuminate, base inequilateral, acute or obtuse, one side 1-3 mm. shorter, 2.5—4 cm. wide X 9-15 cm. long, pinnately nerved from the lower half, the nerves 4 or 5 on each side, with inconspicuous cross-connecting nervules, glabrous and smooth above, scabrous and the nerves strigose-hairy beneath, gland- ular-dotted on both sides, drying rather firm, translucent; petiole 5-8 plus 1—3 mm. long, granular, vaginate at the base; spikes 2-3 mm. thick X 8 cm. long, cuspidate; peduncle 1 — 1.5 cm. long, glabrous; bracts triangular-lunate-subpeltate, rather sparsely short-fringed; ovary compressed-ovoid, papillate about the apex; stigmas sessile. Known only from the following locality, k: north rim of El Vallc de Anton, alt. 600-1000 m., Allen l6jO; vicinity of El Valle, alt. 600-1000 m., Allen llSS; along Rio Indio Trail, alt. 500-700 m., liuutcr G? Allen 333, The only difference which can be noted between the type of the variety and that of the species is the collectors' notations of the habit of growth, Because of the great variation in this character it is not believed to be valid in this case. The species was described as a shrub 3 m, tall, and the variety as a small tree 2.5 m. tall. 37. PirER ADUNCUM L. Sp. Pi. 1:29. 1753. Viper elongafum Vahl, Enum. Pi. 1:312. 1805, sensu Trelcasc in Contr. U. S. Nat. Herb. 26:36. 1927. Viper celtidifolium H.B.K. Nov. Gen. & Sp. 1:50. 1815. (}0) 19 5 0] FLORA OF PANAMA (Ptperaceac) 31 Arfantbe adiinca Miq. Comm. Phyt. 49. 1838. •# 1939. Ut lifolia 1848. 1920. 'i •i 1920. Nat. Herb. 26:37. 1927. Viper intersifum Trel. In H. Daniel, Una Ascension Al Cerro de '*La Vieja," Medellin, Colombia, 12. 1940, In part. Fipcr intersiium var. porcccitcnse Trel. loc. cit. 13. 1940. Shrub or small tree, mostly 3-9 m. tall, nodose; flowering internodes rath- er slender, short, rather sparsely pubes- cent glabrescent; leaves lanceolate to subelliptic, 4-8 or scarcely 10 cm. wide X 12—24 cm. long, apex sharp- acuminate, base incquilaterally rounded or cordulate, one side mostly 3-4 mm. shorter, pinnately nerved to the middle or upper third, the nerves 6-8 on each side, sharply ascending, the innermost meeting at the apex, gradually approxi- mated downward with usually 2 or 3 on each side very near the base, sca- brous and more or less pubescent above, at least along the nerves, commonly tardily somewhat rugulose, pubescent or glabrescent between the nerves and somewhat glandular-dotted beneath, the nerves commonly subappressed-hairy, dry- ing firm, translucent, pellucid-dotted; petiole 3-8 plus 3-4 mm. long, pubescent, vaginate at the base; spikes 3-4 mm. thick X 8-15 cm. long, short-cuspidate, gradually curved or hooked; peduncle slender, 8-20 mm. long, mostly about 10-12 mm., puberulent; bracts triangular- or rounded-subpeltate, marginally fringed; fruit small, obovoid, more or less tri- or tetragonous, glabrous, stigmas sessile. West Indies and on the mainland through Central America and northern South America. BOCAs DEL TORo: Cricamola Valley, vicinity of Almirante, Cooper jJT; Isla Colon, alt. 0-120 m., vOfi WeJel J/J. canal zone: vicinity of Fort Sherman, Staudley 30g4I ; near Rio Medio, Battery 46 Hill, Miller 1770; near old Fort Lorenzo, mouth of Rio Chagres, Piper jpSj; between Chagres Batteries and Fort San Lorenzo, Fort Sherman Military Reser- vation, Maxon ^ Valentifte 6gg2; hills west of the canal, near Gatun, Sfaudley 272^6; alluvial bottom near Bohio, alt. 10-20 m., Maxon 4766; Madden Dam Road, Miller l8oj, colon: along the beach between Fato and Playa de Damas, Pit tier 3935y 3930; forests around Porto Bello, a!t. 5-100 m., Pittier 2438. Panama: near Tapia River, Juan Diaz region, Maxon & Harvey 6671; Tumba Mucrto Road, near Panama, Staitdley 29803; forests, on dry limestone, around Alhajuela, Chagres Valley, alt. 30—100 m., Pittier 35 JO, Fig. 6. Piper aJuneum (31) [Vol. Z7 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN 37a. Piper adlxcum var. brachyarthrum (Trel.) Yuncker, comb. nov. ril)i'r cloffgafum var. brachyarthrum Trcl. in Contr. U. S. N.u. Herb. 26:37. 1927. Viper Cuafrecasasl Trel. in Trab. Mus. Nac. Hist. & Jard. Bot. Madrid, Sen Bot. 3 3:48, 1936. Piper cumbricola Trcl. loc. cit, 193 6. Ptpcr cumbricola var. mouteiuigum Trel. loc. cit. 1936. Viper iUndev% Trel. loc. cit. 50. 19 36. Stems densely hirsute-villous; leaves moderately to densely soft-villous beneath. Occurs with the species. Panama: without locality, Seemaun I ly, chiriqui: vicinity of El Boqucte, alt. 1000-1300 m., March 2-8, 1911, Maxoii jljQ (U.S. Nat. Herb., type), veraguas: vicinity of Santiago, alt. 50 m., Allen loi6, cocle: EI Vallc, Miller 1S15, 1S16, canal zone: Chiva-Chiva Trail, Red Tank to Pueblo Nucvo, Viper 3749. This is a widespread species which exhibits considerable variation in the size of the leaves and character and amount of pubescence. The rather slender, regularly curved spikes, short petioles, and somewhat longer peduncles are characteristic features. Variety lacvifoliiini has slightly less scabrous leaves but otherwise agrees in all respects. 3 8. Piper hispidum S\v. Prodr, Vcg. Ind. Occ, 15. 178 8. Viper scabruni Sw. Fl. Ind. Occ. 1:59. 1797. Viper hirsnlum Sw. loc. cit. 60. 1797. Stefferisia hirsufa Kunth, in LInnaca 13:640. 1839. Sfeffe/jsia scabra Kunth, loc. cit. 1839. Arfaftthe hirsufa Mlq. Syst. Pip. 446. 1844. Artaufhe scabra Miq. loc. cit. 447. 1844. Viper ereclamenium CDC. in Smiths. Misc. Coll. 71^:14. 1920. Viper scabriUmbnm CDC. in Candollca 1:121, 276. 1923. Viper Wriliamsii Trcl. in Contr. U. S. Nat. Herb. 26:3 2. 1927. Viper Killipi Trcl. loc. cit. 33. 1927. Viper Killipi var. calJeranum Trel. loc. cit. 1927. Viper sancti-felicis Trel. loc. cit. 3 5. 1927. Viper rivi-i'ctusli Trel. in Ann. Mo. Bot. Card. 24:186. 1937. Viper barbirostrc Trcl. loc. cit. 27:288. 1940. Viper casiteuse Trel. loc. cit. 2 89. 1940. Viper cerro-pHfiioense Trel. loc. cit. 1940. Viper Iw)iIii)itoense Trel. loc. cit. 293. 1940. Viper Margarctae Trel. loc. cit. 1940. Viper pervicax Trel. loc. cit. 296. 1940. Viper pseuJo-i'iriilicanle Trel. var. nierecifafunfj Trcl. loc. cit. 1940. A shrub, tip to 5-8 m. or more tall, rather nodose; flowering internodes mod- erately slender and rather short, hispid, sometimes glabrescent downward, hairs rigid, commonly more dense about the nodes, erect to reflexed, often of varying lengths; leaves elliptic or elliptic-ovate, 4-1 1 cm. wide X 1 1—20 or 2 5 cm. long, mostly 6-8 X 1 1^15 cm., apex acuminate, base obliquely inequilateral, one side 2—5 mm. shorter, mostly acute, the longer side usually rounded and obtuse, or in large leaves sometimes cordulate, pinna tely nerved from the (32) 1950] FLORA OF PANAMA (Piperaccae) 33 lower half, the nerves 5—6 on each side or 7 1 in some large leaves, wi 1 ith cross-connecting ncrvules, sometimes rugesccnt in age, Icpi- dotc-scabrous, more or less hirtellous or his- pid along the nerves above, the nerves beneath hispid-hirsute, commonly glandular- dotted, drying papery, translucent; petiole 5-10 plus 2-5 mm. long, hirtellous or hispid, vaginate at the base; spikes 3 mm. thick X 8-14 cm. long, mucronate; peduncle rather thick, scarcely 1 cm. long, hirtellous or hispid; bracts triangular-subpeltate, margin- ally fringed; fruit rather small, oblong- subpyramidal-trigonous, laterally compressed, papillate-pubcrulent at the truncate or de- pressed apex; stigmas small, sessile. In a number of scarcely distinguishable forms throughout the West Indies, Central America, and much of South America. Fig. 7. Piper hispiditm BOCAs DEL TORO: Rio Cricamola, between Finca St. Louis and Kontlntoe, alt. 10-50 m., WooJsofi, Alien fj Seibcrt j8g4; vicinity of Nievecita, alt. 0-50 m., Woodson, Allen & Seibcrt iSjQ; Daytona Farm, region of Almirante, Cooper l6j. chtriqut: vicinity of '*New Switzerland,'* central valley of Rio Chiriqui Viejo, alt. 1800-2000 m., Allen 1 365; valley of the upper Rio Chiriqui Viejo, vicinity of Monte Lirio, alt. 13 00-1900 m., Seibert 192; on trail to Gcrro Punta, White 52^ Ij8; Bajo Chorro, Boquctc Distr., alt. 1800 m., Davidson 58, 342; EI Boquctc, alt. 1400-1650 m., Killip 3549; around Las Siguas Camp, southern slope of Cerro de la Horqucta, alt. 1700 m., Pittier 3191 ; Rio Caldcra, above El Boquete, alt. 1600 m., Killip 3S44^ vicinity of San FeHx, alt. 0-120 m., Pittier 51 24; vicinity of Cnsita Alta, Volcan dc Chiriqui, alt. 1500-2000 m., Woodson, Allen & Seibert 843> 902, 978. CANAL zonk: Empire, Miller 1868; Barro Colorado Island, Gatun Lake, Standley 3 1 38/, Panama: Marraganrl and vicinity, alt. 3-60 m., Williams 986; Indio, Madden Lake, Miller 2064. darien: Boca de Pauarando, on Sambu River, alt. 20 m., Pittier 5584 — some of the leaves on this specimen are up to 10-11 X 22-24 cm. 3 8a. Piper hispidum var. trachydermum (Trel.) Yuncker, comb. nov. Piper trachydennuni Trcl. in Contr. U.S. Nat. Herb. 26:33. 1927, Piper non-retrorsuvj Trel. loc. cit. 36. 1927. Differs from the species in having the short, stout hairs on the upper internodcs upwardly curved and more or less appressed. Occurs with the species. Panama: without location, 1859-60, Hayes 791 (Herb. New York Bot. Card., type of P. trachydermum), cocle: El Valle, valley floor and lower slopes, along highway, Miller 18 1 7 — leaves on this specimen are larger than usual, canal zone: Barro Colorado Island, Gatiin Lake, alt. 120 m., Standley 40872; vicinity of Fort Sherman, Standley 30121, colon: around Porto Bello, alt. 5-100 m., Pittier 2429. Panama: Rio Tapia, Standley 28208. (^V I Vol. 37 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3 8b. Piper hispiduai var. ellipticifolium Yuncker, var, nov, Caule hispido (pilis rigidis soorsum recurvantibus) ; foliis ellipticisj deorsum ad basem scnsim angustatis, basi utrinque peracutis. Stem hispid, the hairs rigid, upcurved; leaves elliptic, gradually narrowed to the base which is scarcely rounded and strongly acute on both sides, 7-S cm. wide X 13-16 cm. long. Known only from the type locality. cocle: Bismarck, above Penonomc, alt. 600-900 m., March 5-19, 1908, Williams 34$ (Herb. New York Bot. Gard., type). There is a great amount of variation in the size and to a lesser degree also the shape of the leaves on the specimen included here. All agree in having harshly scabrid leaves, hispid stems, and papillate-puberulent fruit. A study of the type specimens of the numerous species proposed by DeCandollc and by Trelease has failed to show any consistent characters upon which it is believed specific distinc- tions can be justified. 39. Piper vitabundum Trel. in Contr. U. S. Nat. Herb. 26:38. 1927. Viper conscenJerJs Trel. in Field Mus. Nat. Hist. Bot. Ser. 18:338. 1937, Climbing; flow^ering internodcs rather slender and long, moderately rctrorse- strigose; leaves elliptic, apex acvmiinate, base inequilateral, one side 1-3 mm. shorter and acutish, the longer side acutish or obtuse, 3-4 cm. wide X 10-12 cm. long, pinnately nerved from the lower half, the nerves 4—5 on each side, Icpidote, finely scabrous, and glabrous above, the nerves beneath appressed-pubescent, drying dark, subopaque; petiole about 5 plus 1—3 mm. long, pubescent, vaginate at the base; spikes 2—3 mm. thick X 7-8 cm. long; peduncle scarcely 1 cm. long, sparsely crisp-pubescent; bracts triangular-subpeltate, narrowly fringed; fruit oblong-sub- tetragonous, laterally compressed, papillate-puberulent at the top, stigmas sessile. Known only from Changuinola Valley, Bocas del Toro. BOCAS DEL TORO: Changuinola Valley, Dnnlap JjS, The vine-like habit of this species is distinctive. P. consccudcin was based on the same collection as P. litabundiini. 40. Piper persubulatum CDC. in Smiths. Misc. Coll. 71*'*:4. 1920. Shrub (?) ; flowering Internodcs rather slender and short, densely brown- villous, the hairs up to 1 mm. or more long; leaves lance-elliptic-oblong, w^ith the midrib nearer one side, apex very narrowly long-acuminate, base subequally rounded, 5.5—8 cm. wide X 19-24 cm. long, pinnately nerved from the lower half, the nerves 6—7 on each side, with prominent cross-connecting nervules, the upper surface somewhat scabrous, loosely villous, the nerves rather densely so, marginally strongly long- ciliate, more densely dingy-hairy beneath, drying papery, subopaque; petiole scarcely 5 mm. long, densely villous, vaginate at the base; spikes 2 mm. thick X 10 cm. long; peduncle scarcely 5 mm. long, villous; bracts subpeltatc, densely dingy-hairy; ovary elongated with the rachis, glabrous, stigmas sessile. (34) 1950] FLORA OF PANAMA (Plperaccae) 35 Known only from the following locality. COLON: Loma dc la Gloria, near Fato (Nombre de Dios), Pit tier 4240, The densely villous stems and leaves, long, slenderly acuminate leaves, and densely hairy bracts distinguish this species. 41. Piper peracuminatum CDC. in Smiths. Misc. Coll. 71^^:9. 1920. A shrub or small tree, 2 od stout, hirsute, the hairs of various lengths; leaves elliptic-obovate, apex narrowly sharp-acuminate, base inequilateral, cordulate, the longer side 2—4 mm. longer at the petiole and covering the petiole, 8-13 cm. wide X 16-27 cm. long, scabrous above and loosely villous or at least pubescent along the nerves, thinly long-hairy beneath, ciliate, somewhat rugescent or nearly smooth, pinnatcly nerved from the lower half, the nerves 6 or 7 on each side, drying rather thin, subtranslucent; petiole 5-10 plus 2-4 mm. long, densely hirsute, vaginate-winged to the blade; spikes 5 mm. thick X 10-13 cm. long; peduncle rather stout, 2 cm. long, with intermixture of short, stiff, and long, crisp hairs; bracts rounded- or triangular- subpcltate, fringed, pedicel villous; fruit oblong, compressed, rusty-hirtellous, stigmas sessile. P anama. CANAL zone: Chagres, Fencller 26g. colon: around Dos Bocas, Rio Fato valley, alt. 40-80 m., Pittier 421O; Rio Indio de Fato, Pittier 4254, The rather large, scabrid, cordulate-based leaves, mixture of long and short stiff hairs, and hairy fruits distinguish this species. 42. Piper erueescentisptcum Trel. in Ann. Mo, Bot. Gard. 27:291. 1940, as ertibescojfispica, A shrub, 2 m. tall, nodose; flowering Internodes rather short and slender, evanescently villous; leaves rhombic-elliptic, apex sharp-acuminate, base inequi- lateral, cordulate, one side 2-4 mm. shorter, 4-6.5 cm. wide X 10-14 cm. long, pinnately nerved from the lower half, the nerves 4-5 on each side, with rather prominent cross-connecting nervules, harshly scabrous, sparsely villous above, the hair bases enlarged, whitish, loosely villous beneath, at least along the nerves, the hairs more or less erect, up to 1 mm. or more long, marginally ciliate, young leaves woolly- villous when unfolding, drying papery, translucent; petiole scarcely 5 plus 2-4 mm. long, at first villous, vaginate at the base; spikes red, as reported, 2 mm. thick X 6 cm. long; peduncle about 5 mm. long, glabrcscent; bracts glabrous, triangular-subpeltate, dorsally somewhat projected and callose, slightly ciliate beneath; fruit not developed, stigmas sessile. Bocas del Toro. BOCAs DEL toro: vicinity of Nievecita, alt. 0-50 m., Woodson, Allen ^J Scibert 181J; Sansan Swamp, region of Almirante, Cooper IQO, The villous, more or less rhombic-elliptic, scabrous leaves, red spikes, and irreg- ularly shaped bracts distinguish this species. r3 5; [Vol. 37 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN 43. Piper culebranum CDC. in Candollca 1:121, 243. 1923. A shrub or small tree, 2 m. tall; upper internodes comparatively short and slender, loosely to moderately crisp-vlHous; leaves elliptic, elliptic-oblanceolate or elliptic-subobovate, apex acutely long-acuminate, base 3-5 mm. shorter on one side, obtuse, or longer side cordulatc, 5-7 or sometimes 9 cm. wide X 15-20 cm. long, pinnately nerved from the lower half, the nerves 4-5 on each side, with cross- connecting and anastomosing nervules, sparsely wdiitc-villous glabresccnt above, mostly crisp-villous and slightly scabrous beneath, the nerves more densely so, drying rather thin, green, somewhat glossy, translucent; petiole niostly about 5 plus 3-5 mm. long, crisp-villous, vaginatc at the base; spikes 4 mm. thick X 7-9 cm. long; peduncle 1 cm. long, crisp-villous; bracts round-triangular-subpeltate, with a wide, dense yellow fringe; fruit subcylindrical, truncate; stigmas sessile. Central Panama. CANAL ZONr: F.mpire, Miller lS66; near Fort Randolph, StauJlcy 28622, 2866^, colon: along the Rio Culcbra, above Santa Isabel, near sea-level, Viificr 41 54; between France Field, Canal Zone and Catival, SiamUcy 30375. panamA: Juan Diaz, Sfiiudlcy J0611 ; along Rio Juan Diaz above Juan Diaz, alt. 30 m., Allen Q34. There is considerable variation in the size of the leaves on the specimens in- cluded here, especially Miller's No, 1866, but they agree very well otherwise, and it is believed they all represent the same species. 44. Piper viridicaule Trel. in Contr. U. S. Nat. Herb. 26:32. 1927. A shrub, 2 m. tall, branches subdichotomously forked; flowering internodes slender, elongating downward, villous, the hairs up to 1 mm. or more long; leaves lancc-clliptic, apex sharp-acuminate, base inequilateral, cordulate, one side 2-4 mm. longer and covering the petiole, 3-4.5 cm. wide X 9-12 cm. long, pinnately nerved from the lower half, the nerves 4 or occasionally 5 on each side, with slender cross- connecting nervules, sparsely villous and harshly scabrous above, paler beneath and apprcssed- villous, especially along the nerves, marginally long-cillate, pale-gland- ular-dotted beneath, drying thin, translucent; petiole 5-20 plus 2-4 mm. long, villous, vaginate at the base; spikes unknown. Known only fiom Barro Colorado Island, CANAL zone: Barro Colorado Island, Gatun Lake, Stavdley 3^4^^- The absence of inflorescence on the only specimen known of this species makes its position questionable. The vegetative characters, especially the villous twigs and leaves, distinguish it from any other known species. 45. Pn>i:R pekvelutinum Trek in Contr. U. S. Nat. Herb. 26:39. 1927. Shrub (?), nodose; flowering internodes short, rather stout, densely subvillous; leaves lance-oblong, apex subacuminate, base inequilateral, rounded, obtuse, one side 2-5 mm. shorter, the upper surface rugose, scabrous, hirsute, especially along the nerves, dull, densely dingy-villous and slightly scabrous beneath, not ciliate, pinnately nerved from the lower half, the nerves 6-7 on each side, impressed above, (^6) 1950] FLORA OF PANAMA (Pipcraceac) 37 gradually approximated downward, drying rather thick, opaque; petiole 8-10 plus 2-5 mm. long, densely dingy-villous, clasping and vaginate at the base; spikes as yet immature, 2 mm, thick X 1-5 cm. long; peduncle short, densely hairy; bracts round-subpeltate, fringed, the dorsal hairs much longer; ovary not formed. Known only from the one specimen. Panama: without locality, Hayes /g6. 46. Piper variitrichum Yunckcr, sp. nov. Frutex 1-2 m. altus, caulibus nodosis; internodiis florifcris gracilibus con- ferte atro-pubescentibus; foliis ovato-ellipticis, apice acute acuminatis, basi inacqui- lateraliter obliquis, latere angusto obtuso vol acuto, latere lato rotundato cordulato, supra conferte scabridis, a tcrtio supero deorsum pinnatim venosis, venis utrin- que 5-6, supra hispidis subtus pubes- centibus, ad venas sat conferte hirsutis. pilis erectis brcvioribus longioribusque intermixtis; petiolo 1-1,5 cm. longo, conferte hirsuto, deorsum vaginato ; spicis Immaturis 2 mm. crassis, 4 cm. longis apiculatis; pedunculo 5 mm. longo conferte pubescente; bracteis rotundato-subpeltatis fimbriatis. Shrub, 1—2 m. tall, nodose; flower- ing internodes slender, lengthening downward, densely dingy-pubescent ; leaves ovate-elliptic, apex narrowly at- tenuately acuminate, base inequilater- ally oblique, essentially equal length at the petiole, the narrower side narrow- ing downward, obtuse or acutish, the broader side rounded, cordulate, 6—8 cm. wide X 15-23 cm, long, harshly scabrous above, less so beneath, pin- nately nerved below the upper third, the nerves 5-6 on each side, with slender, cross-connecting ncrvules, glandular- dotted and hispid above, pubescent beneath w^ith the nerves rather densely staring- hair}^ with a mixture of various length hairs, scarcely ciliate, drying rather thin, translucent; petiole 1-1.5 cm. long, densely hairy, vaginate toward the base; spikes as yet young, 2 mm. thick X 4 cm. long, apiculate; peduncle 5 mm. long, densely pubescent; bracts round-subpeltate, fringed, with a few dorsal hairs longer; ovary and fruit not developed. Known only from the type locality. Fig. 8. P/per variitrichum 07) [Vol. Z7 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN OARIEN: Chepigana Distr., Cana-Cuasi Trail (Camp 2), alt. 600 m., March 9, 1940, Terry d Terry 1428 (Herb. Missouri Bot. Card., type). The comparatively large leaves, narrowly acuminate apex, the margins essen- tially equal at the petiole, and various length hairs on the nerves beneath distinguish this species. 47. Piper cttanguinolanum Trel. in Contr. U. S. Nat. Herb. 26:37. 1927. A nodose shrub with scabrous twigs and leaves; flowering internodes short and comparatively slender, finely hispid and granular; leaves inequilateral, narrowly elliptic-oblong, apex acuminate, base 2—4 mm. shorter on one side and acute, the longer side rounded, cordulate, 3-4 cm. wide X 12-15 cm. long, pinnately nerved from below the middle, the nerves 5-6 on each side, rather slender but conspicuous beneath, with slender cross-connecting nervulcs, slightly rugose in age, the nerves above hispidulous, paler and granular-dotted beneath and appressed-pubesccnt on the nerves, appressed-hispidulous ciliate, drying papery, translucent; petiole scarcely 5 plus 2-4 mm. long, hispid, vaginate at the base; spikes 3 mm. thick X 8-10 cm. long; peduncle scarcely 1 cm. long, subappressed-hispid; bracts roundcd-triangular- subpeltate, thinly stiff-ciliate, the anterior setae longer; fruit oblong, laterally- compressed, apex depressed- truncate, granulate or essentially glabrous, stigmas sessile. Known only from Changuinola Valley, Bocas del Toro. BOCAS DEL toro: Changuinola Valley, DtinJap 2JJ. 48. Piper villiramulum CDC. in Smiths. Misc. Coll. 71^:11. 1920. Pilfer bocascfisc Trcl. in Field Mus. Nat. Hist. Bot- Ser. 18:333. 1937. Piper yapearnim Trel. in Ann. Mo. Bot. Card. 25:827. 1938. A shrub or small tree, commonly 2-3 m. tall; flowering internodes moderately slender, moderately to rather densely villous; leaves ovate or elliptic-ovate, apex acuminate, base incquilaterally oblique, one side 3-5 mm. shorter and obtuse, the longer side rounded, cordulate, mostly 5-8 cm. wide X 13-18 cm. long, occasional leaves somewhat larger, scabrous on both sides but more harshly so above, mod- erately apprcsscd-hairy on both sides, becoming somewhat bullulate. pinnately nerved from the lower half, the nerves 6 or 7 on each side, drying rather thin, translucent; petiole 5-10 plus 3-5 mm. long, vaginate at the base, moderately to densely villous; spikes 3 mm. thick X ^~9 cm. long; peduncle rather stout, scarcely 5 mm. long, hairy; bracts rounded-triangular-subpeltatc, fringed, the hairs com- paratively short, coarse and somewhat rigid; fruit tetragonous, laterally compressed, glabrous or obscurely papillate at the apex, stigmas sessile. Panama. BOCAS DEL toro: region of Almirante, Cooper 4O/; Changuinola \'al!cy, Dnnhip 2J4; vicinity of Chiriqui Lai^oon, t'on WeJcl 121 y. Water Valley, von WcJcI 6()S^ ^777, 2660; Shepherd Island, ton WeJel 26jg — this specimen is more rui;ose than usual, canal 7on'E: Rio Pedro Miguel, near East Paraiso, Stiunllcy 2gg66; around El Paraiso, alt. 3 0-100 m., Piffier 25J4; Corozal, Stciefis 124; Rjlcy Ij8; alon.^ the old Las Cruces Trail, between Fort Clayton and Corozal, Sfandley 2gigg; between Miraflores and Corozal, alt. 20-30 m., (3S) 1950] FLORA OF PANAMA (Piperaceac) 39 Piftier 2igj; Las Cascadas Plantation, near Summit, Sfandley 2j6g2, 2§JOly 26012; Summit, Sfandlcy 26g^y; Cerro Gordo, near Culebra, SfanJIey 2^gg8; Balboa, Stavdley 26013, 29267; Gamboa, Standley 28^22; Empire, Miller l86g; Empire to Mandlnga, Piper 5514; vicinity of Ancon, Piper 6028; Bro. Celestine 32; Rio Agua Salud, near Frijolcs, Piper 586J; near Rio Medio, Madden Dam Road, Miller 1768, Ij6g, ^77^, l8o2; vicinity of Madden Dam, near Rio Cliagres, alt. 50-75 m., Seibert 54g; near old Fort Lorenzo, mouth of Rio Chagres, Piper 5971. colon: Loma dc la Gloria, near Fato (Nombre de Dios), alt. 10-104 m., Pittier 4^83. tanama: Sabana de Juan Corso, near Chepo, alt. 60-80 m., Piftier 4537; along the Corozal Road, near Panama, Standley 26843; Juan Diaz, Standley 30623; Rio Tapia, Standley 28134; 3 miles east of Juan Diaz, alt. 75 m.. Killip 3123; Bella Vista, Macbride 2732; Camino de La Granja, Bro. Heriberto 165. darien: near mouth of Rio Yape, alt. 20 m., Allen 351. 48a. Piper villiramulum var. gamboanum (CDC.) Trel. in Contr. U. S. Nat. Herb. 26:36. 1927. Piper gamboaniim CDC, in Candollea 1:249. 1923. Piper hispidiim Sw. var. gamboanum CDC in Smiths. Misc. Coll. 71^:12. 1920. Piper gamboanum var. yapense Trel. in Ann. Mo. Bot. Gard. 27:291. 1940. Leaves 2-4 or occasionally up to 4.5-5 cm. wide. Occurs with the species. CANAL zone: around Gamboa, alt. 20-100 m., Pittier 3410; Corozal, Stevens I16, 117; foot of Corozal Hill, Miller 1/88; along the old Las Cruces Trail, between Fort Clayton and Corozal, Standley 29048; Las Cascadas Plantation, near Summit, Standley 29639; Obispo, Standley 31794^ Panama: near the big swamp, east of Rio Tecumen, Standley ^659^; between Las Sabanas and Matias Hernandez, Standley 31818. darien: vicinity of Yape, Allen 854. The apprcsscd-villous, scabrous, buUulate leaves, and short, stout peduncles characterize this species and distinguish it from P. tabogamim in which the leaves are less scabrous, scarcely bullulate, more densely hairy, and with longer peduncles. The young spikes of P. faboganiim are also usually very short and bristly from the erect bract hairs. The type of P. yapcamim is reported as a tree 15 m. tall. The type specimen agrees in all other respects with those reported as being much smaller plants. 49. Piper taeoganum CDC. in Smiths. Misc. Coll. 71^:4. 1920. Piper obaldianum CDC. loc. cit. 7, 1920. Piper dnmeticola CDC var. pana?nense CDC. loc. cit. 1920. Piper pallidibracteum CDC loc. cit. 8. 1920. Piper saliuasanum CDC var. parvifolium CDC. in Ann. Conserv. & Jard. Bot. Geneve 21:234. 1920. Piper barbinerve Trel. in Contr. U. S. Nat. Herb. 26:35. 1927. Piper Bigelotm Trel. loc. cit. 2 3. 1927. Piper siibdilatatuvi Trel. loc. cit. 3 5. 1927. Piper chagresianum Trel. loc. cit. 37. 1927. Piper breve CDC. in Trel. loc. cit. 3 8. 1927. Piper breve var. pallidineunnn Trel. loc. cit. 1927. Piper breve var. pseudo-dicnieticola Trel. loc. cit. 1927. Piper breve var. bellae-vistae Trel. loc. cit. 39. 1927. Piper breve var. divaricatnm Trel. loc. cit. 1927. Piper breve var. pseudo-fafoannm Trel. loc. cit. 1927. Piper cayazasense Trel. in Ann. Mo. Bot. Gard. 25:826. 1938. (39) IVoL. 37 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN Piper amplnbium Trcl. loc. cit, 27:288. 1940. Viper in^ulicolum Trcl. loc. cit. 293. 1940. Piper fahcrnillaitum Trcl. var. ancouense Trcl. loc. cit. 298. 1940. Piper femblaJorense Trcl. in Ll. Williams, Expl. Bot. Guyana Vcncz. 177. 1942 (Scrv. But. MInist. Agr. y Cria). A shrub, mostly 1-3 m. tall; upper intcrnodcs rather short and slender, soft- villous, crisp-pubcsccnt, or subhispid, more or less cancsccnt; loaves rhombic- subobovate, elliptic, or less commonly lance-elliptic, mostly 4-6 cm. wide X 10-15 cm. long, or occasional leaves up to 9 X 22 cm., apex acuminate, narrowed down- ward to the inequilatcrally rounded base, one side commonly 2-5 nmi. shorter, the longer side obtuse or cordulate, pinnately nerved from below the middle, the nerves 5-6 or sometimes 7 on each side, often sUghtly rugesccnt near the margin or toward the base, crisp-hirtcllous or subhispid above with the midrib densely so, somewhat scabrous from the persistent hair bases, soft-pubescent beneath, the nerves usually densely so, the hairs appressed, drying rather thin, translucent; petiole 5-10 or occasionally up to 20 plus 2-5 mm. long, densely hairy, vaginate commonly to the middle or above, the groove smooth or hairy within; spikes scarcely 1 cm. Ion when young and more or less bristly from the suberect bract hairs, becoming 3-4 mm. thick X 4-10 cm. long, commonly apiculate; peduncle mostly 5-10 or oc- casionally up to 20-25 mm. long, pubescent; bracts rounded or triangular-sub- pcltate, marginally fringed, the hairs rather stiff, those on the dorsal margin usually longer and suberect in young spikes; fruit oblong-subtrigonous, glabrate or slightly papillate when young; stigmas small, sessile. Northern South America and Panama. Panama: without locality, Bigelow s. n.; Nee jjS, 765. chirh^ui: Boquctc District, Boquete, alt. 1140 m., Davidson diy. veraguas: trail between Canazas and the foot of the Cordillera Central, headwaters of Rio Canazas, alt. 300-600 m., Allen 185. cocle: vicinity of Ola, alt. 100-350 m.. Pfttier 3034; El Valle dc Anton and vicinity, alt. 500-700 m,y Seihert 436; El Vallc, Miller 1818, l8ig — small-Icaved specimens — 1820; between Aguadulce and Anton, alt. 15-50 m., Woodson, Allen ^ Seiheri 1222; vicinity of Santa Clara beach, Woodson, Allen 6 Seibert Tjoy, between Las Margaritas and El Valle, Wood- son, Allen Gf Seibert Jjo6, 172S. canal zone: Empire, Miller 1S64, l86g; Empire to Mandinga, Piper 5523; near old Fort Lorenzo, mouth of Rio Chagrcs, Piper 5944; between Chagrcs Batteries and Fort San Lorenzo, Fort Sherman Reservation, Maxon & Valentine 6q86; vicinity of Fort Sherman, Standley 30918, 30982, 3^1 4O; western slope of Ancon Hill, vicinity of Balboa, alt. 20-75 m., Seibert I18, IIQ, 399; Woodson, Allen 6 Seibert 1327; Standley 25203, 26382, Killip 120/8; Balboa, Standley 25420, 25603^ 25363; 2 miles w^est of Balboa, Correll 12261; Las Sabanas, Bro, Celestine HO; Riley I16; Cocli, Riley 12/; Matachin, Cowell 204; foot of Corozal Hill, Miller JjgJ; Corozal, Standley 2^341; near Fort Randolph, Standley 28639; Maxon ^ Harvey 6304, ^543i Fo^'t Kobe road, Woodson, Allen & Seibert 1412; Brazos Brook Reservoir, Stevens Y32; Culcbra, Stevens 993; alt. 50-150 m., Pittier 3445; around Gamboa, in thickets, alt. 40-80 m., Pittler 3709; Standley 283J4, 28481; between Gamboa .ind Cruccs, alt. 50-80 m., Pittier 378J; France Field, Stevens 981; Margarita Swamp, south of France Field, Maxon ^ Valentine 7^551 between France Field, Canal Zone and Catival, Colon, Standley 30164; Mount Hope Cemetery, Standley 28776; Obispo, Standley 31764; Juan Mina, Piper 5692; Cerro Gordo, near Culcbra, Standley 25975, 26018; Chlvi-Chlvi Trail, 2 miles above Red Tank, Maxoi ^ Harvey 6591, colon: Chagrcs, Pendler 268; Rio Culcbra, above Santa Isabel, Pittier 4148; Rio Indio de Fato, near sca-levcl, Pittier 4^7^ I ^long the beach between Fato and Playa de Damas, Pittier 3923, 3930, Panama: near Matias Hernandez, Standley 28957; (40) 1950] FLORA OF PANAMA (Pipcraccac) 41 near Las Sabanas, Standley 4()/'/g; between Las Sabanas and Matias Hernandez, SfunJley Jl8j2; Rio Tapia, SiafiJley 2810J; near Tapia River, Juan Diaz Region, Maxofj & Haricy 6620, 6626; Sabana dc Juan Corso, near Chepo, alt. 60-80 ni., Pittier 4539; Rio Tecumen, Standlcy 2g4jl ; near big swamp cast of Rio Tecumen, Standlcy 26jjl; near Punta Paitilla, Standley 26jOj; Stevens 34Q; Juan Diaz, Staudley jodj/; Tumba Mucrto Road, near Panama, Staudley 2g/24; Bella Vista, at sea-level, Killip 12045, 12004; Piper 5331; vicinity of Juan Franco Race Track, near Panama, Standley 2"// 13^ ^7734, ^7777\ Taboga Island, Woodson, Allen & Seibert 1 527; Staudley 27019, 27920; Miller 1S43, 1846 — a large-leaved specimen — , 1847; alt. 90 m., Collcuette 487; Killip 3194; Pittier 3529; Taboguilla Island, Miller 2003; Perbs Archipelago, San Jose Island, Johnston 687, I Gig, 1404; Erlan- son 376 — this specimen differs in having more elliptic-ovate leaves scarcely narrowed toward the base, but it agrees very well in other characters; Trapechc Island, Miller 1908, Allen 2618; Saboga Island, Miller TQjI — a large-leaved specimen — 1952, san blas: forests around Puerto Obaldia, alt. 0-5 m., Pittier 4366, 4396 — a large-leaved specimen. darien: Chcpigana District, Cana-Cuasi Trail, alt. 600 m., Terry & Terry 1428. This species is one of the most common in Panama where it occurs in a great variety of scarcely diflfcrentiablc forms. It appears to be rather closely related to P. niolliconiuni of Brazil, from which it differs, however, because of its mostly less venulose and more finely rugescent leaves which have the margins less rounded below the micidle, and with generally shorter peduncles. The type specimen of P. moUicomum has not been examined by the writer and the above contrast is based on notes and sketches made by the late Professor Trclease from specimens in European herbaria. There is a considerable range in size and amount of pubescence shown by the leaves, often on the same specimen. The contrast between large- leaved and small-leaved specimens is very great. Other specimens, however, show intergradations in size to such an extent that it does not seem advisable to establish or maintain varieties based on this character alone. The type of P. Bigelovii is a rather poor specimen which appears to agree well with this species although it is somewhat more tomentose than usual for most of the specimens included. 50, Piper pseudo-cativalense Trel. in Contr. U. S. Nat. Herb. 26:34. L927. Shrub, 2 m. tall; flowering internodcs rather slender and somewhat elongate, crisp-pubescent ; leaves elliptic or elliptic-ovate, apex bluntly short-acuminate, mucronate, base inequilaterally rounded, cordulate, larger leaves strongly so, equal at the petiole or one side about 2 mm. shorter, mostly 5—6 cm. wide X 10-13 cm. long, pinnately nerved from the lower half, the nerves 4-5 on each side^ with cross- connecting-anastomoslng ncrvules, scabrous, crisp-hairy on both sides, the nerves more densely so, the hairs on the nerves beneath upcurved, drying papery, trans- lucent; petiole about 10 plus 2 mm. long, crisp-pubescent, vaginate to the middle or above; spikes as yet very young, bristly with the erect bract hairs; peduncle pubescent; ovary or fruit not developed. Known only from the following locality. colon: France Field, Canal Zone, and Catival, Standley 30399. The elliptic or somewhat ovate leaves, hairy on both sides, and with the base rounded-cordulate on both sides, and bristly young spikes distinguish this species. It is apparently closely related to P. tabogannm but differs because of the shape of the leaf, and more rounded cordulate base. (41) [Vol. 37 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN 5L Piper lucigaudens CDC. In Smiths. Misc. Coll. 71*': 10. 1920. Piper [)seuJo-i'iriJicauIe Trel. in Ann. Mo. Bot. Gard. 27:296. 1940. A shrub or small tree, up to 3-4 m. tall, nodose; flowering intcrnodcs somewhat zigzag, slender and rather short, granular, sparingly crisp-pubescent; leaves elliptic or clUptic-oblanceolate, the midrib somewhat nearer one side, apex narrowly atten- uately acuminate, base Inequilateral, one side 2-5 mm. shorter, acute or the longer side more rounded and obtusish, 4-6 cm. wide X 13-17 cm. long, somewhat scabrous, pinnately nerved from the lower half, the nerves 4-5 on each side, with cross-connecting-anastomosing nervules, glabrous above, hirsute along the nerves beneath, the hairs subcrect, mostly 0.5 mm. or more long, drying papery, green above, paler and glandular-dotted beneath, translucent; petiole about 5 plus 2-5 mm. long, somewhat hirsute, vaginate at the base; spikes as yet young, 2-3 mm. thick X 8-10 cm. long, peduncle up to 1 cm. long, granular, glabrous; bracts rounded- or triangular-subpeltate, narrowly and sparsely fringed; ovary or fruit not developed. Panama. CANAL zone: forest along the Rio Indio de Gatun, near sea-level, Piftier 2/gi , 2ygia; near Rio Medio, Miller TJ'66; Barro Colorado Island, SfanJley 31288, 3^3^7' 51a. Piper lucigaudens var. Alleni (Trel.) Yuncker, comb. nov. Piper Alleni Trel. in Ann. Mo. Bot. Gard, 25:826. 1938, not CDC. 1910. Tree 7 m. tall; flowering internodes more densely subhispid; leaves somewhat larger, hairs beneath on midrib appressed, less than 0.5 mm. long. Known only from the type locality. darien: trail between Pinogana and Yavisa, alt. 15 m., Allen 2'JO (Herb. Univ. III., type) — "Roots used by Indians to deaden pnin, leaves used as snake bite remedy." The specimen upon which P, pseuJo-virklicauIc was based is sterile. It ag very well in its vegetative characters with the others Included here, and it is believed to be the same. 52. Piper Storkii Trel. in Contr. U. S. Nat. Herb. 26:32. 1926. Shrub (?) , nodose, nearly glabrous; flowering internodes slender and short, granular, sparsely pubescent when young; leaves elliptic or lance-cUiptic, the mid- rib nearer one side, apex attenuately acuminate, base strongly inequilateral, one side 2-3 mm. shorter, acute, the longer side rounded, obtuse, pinnately nerved from the lower half, the nerves 4—5 on each side, approximated downward, with cross- connecting-anastomosing nervules, 4-5 cm. wide X 14—16 cm. long, glabrous and somewhat scabrid above, very obscurely apprcssed-pubescent beneath, at least along the nerves, drying thin, green, translucent; petiole about 5 plus 2—3 mm. long, ranular, glabrous, vaginate at the base; spikes as yet young, 2 mm. thick X 6 cm. long; peduncle scarcely 5 mm. long, stout, glabrous; bracts triangular-subpeltate, glabrous or with a few, short, ventral hairs; ovary and fruit not matured. (42) 1950] FLORA OF PANAMA (Piperaceue) 43 Known only from Bocas del Toro. BOCAS DEL TORO: Rubber Tree Station, Stork 124. The thin, nearly glabrous leaves, and essentially glabrous bracts characterize this species. 53. Piper Wagneri CDC. in DC. Prodr. 16^:302. 1869. Piper minufispicum CDC. in Smiths. Misc. Coll. 71^:1. 1920. Piper Wagneri var. mluiitispicum Trel. in Contr. U. S. Nat. Herb. 27:29. 1927. A glabrous, nodose shrub; upper intcrnodes relatively short and slender; leaves lance- or ovatc-elliptic, apex moderately acuminate, base subequilateral, the margin equal at the petiole or with one side commonly 1-5 mm. shorter, acute or occasion- ally with the longer side subcordulate, 4-7 cm. wide X 10-14 cm. long, pinnately nerved from below the middle, the nerves 3-5 on each side, with fine, cross- connecting-anastomosing nervules, glandular-dotted beneath, drying rather papery, translucent; petiole 10-25 plus 1-5 mm. long, vaginate to the blade; spikes up to 4 mm. thick X 9 mm. long, mucronate; peduncle 5-8 mm. long; bracts concave- inflexed, glabrous; ovary glabrous, stigmas sessile. Southwestern Panama, CHiRiQUi: Wagner; oak-palm forest above Sabana de El Salto, above El Boquete, alt. 1 500-1750 m., Pittier 3116. DeCandoUe described P. minntispiciirn as dioecious. The spikes on the type are very young and, as in young spikes generally, the ovary is scarcely developed. It is believed that specimens with mature spikes will show that the flowers are perfect. Otherwise, the types of P. W^agneri and P. rninutispicum appear to be the same species. The glabrous stems and leaves and very short spikes are distinctive characters. 54. Piper Dunlapi Trel. in Contr. U. S. Nat. Herb. 26:23. 1927. (?), nodose, elabrous; flowerine internodcs rather short and slender. Shrub glandular-dotted; leaves ovate or lance-ovate, apex gradually sharp-acuminate, base inequilateral, acute, or equilaterally rounded and obtuse, 3.5-5.5 cm. wide X 10—13 cm. long, pinnately nerved from below the upper third, the nerves about 4-5 on each side, with faint cross-connecting-anastomosing nervules, strongly glandular- dotted on both sides, drying rather thick, narrowly revolute, subtranslucent, glossy above, dull beneath; petiole 1—1.5 cm. long, vaginate at the base, or on larger leaves to the blade; spikes as yet immature, somewhat curved, apiculate, 3 mm. thick X 2—3 cm. long; peduncle slender, scarcely 1 cm. long; bracts round- or subquadrangular-subpeltate, densely yellow-fringed, the fringe hairs filiform; ovary and fruit not developed. Known only from the following locality. BOCAS DEL TORo: Changuinola Valley, Dunlap 2JI, (43) [ VuL. 37 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN The glabrous stems and leaves, short spikes, and densely ycIlow-frlnged bracts distinguish this species. It resembles P. birtcllipctiohiin var. siibtriuerve to a con- sidcr.;ble extent but differs because of its more rounded floral bracts which arc scarcely umbonate when young, and with more filiform fringe hairs. The Gray Herbarium specimen, obviously the same collection as that of the type in the U. S. National Herbarium, has scarcely glandular-dotted leaves. 55. Piper palmasanum CDC. in Smiths. Misc. Coll. 71*^:3. 1920. A shrub; flowering intcrnodcs moderately slender, elongating downward, from very sparingly soft-pubescent quickly glabrescent; leaves ovate-lanceolate, apex gradually sharp-acuminate, base subcquilaterally rounded or with one side 1-2 mm. shorter at the petiole, obtuse or abruptly acutish at the petiole, 3-5.5 cm. wide X 10—15 cm. long, pinnatcly nerved from below the middle, the nerves about 4 on each side, gradually closer togetlier downward, impressed above, raised beneath, with cross-connecting-anastomosing nervules, glabrous above, the nerves beneath apprcsscd-hairy, obscurely glandular-dotted beneath, drying firm, subtranslucent; petiole 5—15 mm. long, sparsely pubescent glabrescent, vaginatc-wingcd to the middle; spikes yellow, as yet young, 2 mm. thick X 2 cm. long, mucronate; peduncle 5-10 mm. long, sparsely pubescent; bracts triangular-subpeltate, yellow- fringed; ovary and fruit not developed. Known only from Cucsta de Las Palmas, Chiriqui. CHiRiQUi: humid forest of Cucsta dc Las Pahiias, southern slope of Ccrro dc la Hor- qucta, alt. 1700-2100 m., Vittier 3225. The relatively small, yellow spikes, and the character of the pubescence char- acterize this species. 56. Piper reptabundum CDC, in Bot. Gaz. 70:169. 1920. "Repent or scandent," glabrous; flowering intcrnodes slender and short, granu- lar-scurfy; leaves strongly inequilaterally, somewhat falcately, oblong-elliptic, the margin on one side quite straight and subparallel with the midrib to near the apex, the other side strongly curved from the base to the tip, apex sharply acuminate, base inequilateral with one side about 2 mm. shorter, the longer side obtuse or cordulate- aurlculate and covering the petiole, 3.5—5.5 cm. wide X 13—20 cm. long, pinnately nerved from below the upper third, the nerves 4-6 on each side, strongly curved toward the apex, with strong cross-connecting veinlets and finely nervulcse Avirh anastomosing nervules, finely granular beneath, drying rather firm, translucent; petioles nearly obsolete, 1—2 plus 2 mm. long, scurfy; spikes 2 mm. thick X 4—8 cm. long, somewhat curved; peduncle slender, 4-7 cm. long; bracts small, tri- angular-subpeltate, smooth or granular-papillate; fruit obpyriform- trigonous, granular-papillate; stigmas sessile. Costa Rica and adjacent Panama. BOCAS DEL TORo: Sibub! Falls, Sixaola Valley, RotcJec ^ Roulcr J02 (Cited as J/'2 by Trclcasc in Contr. U. S. Nat. Herb. 26:38. 1927). The subarcuate leaf shape and very long peduncles strongly characterize this species (44) 1950] FLORA OF PANAMA (Pipcvaccae) 45 57. Piper chiriquinum CDC. in Smiths. Misc. Coll. 71^2. 1920. Piper paw-anchocnse Trcl. in Ann. Mo. Bot. Gard. 27;29'^. 1940. A shrub, 2-3 m. tall, glabrous, nodose; flowering internodes rather slender and short, smooth or with round lenticcls; leaves ovate or lance-ovate, apex bluntly acuminate, base mostly rounded, acute or obscurely and shallowly cordate, abruptly contracted onto the petiole, 3.5-8 cm. wide X 8-14 cm. long, pinnately nerved from below the upper third, the nerves 3-4 on each side in small leaves or 4-5 in larger leaves, with 2 or 3 pairs from near the base and 1 or 2 pairs above, with a few shorter intermediates and cross-connecting-anastomosing nervules, drying firm, pale green, translucent; petiole 10-25 mm. long, vaginate at the base or occasionally to about the middle; spikes 2-3 mm. thick X 5-8 cm. long, usually mucronate; peduncle 5-10 mm. long; bracts rhomboid-subpeltate, umbonate, submarginally ciliolatc; fruit obpyramidal-trigonous, glabrous, stigmas sessile. Panama. CHiRiQui: pastures around El Boquetc, alt. 1000-13 00 m., Vittier 2gjO; Maxon 4944; Davidson 725; forests along the Rio Ladrillo and vicinity, above El Boquete, alt. 1200-13 00 m., Maxofi 5557, 555^1 trail from Paso Ancho to Monte Lirio, upper valley of Rio ChiriquI Viejo, alt. 1500-2000 m., Allen 1 579. This species bears some resemblance to P. aeqnale from which it is to be dis- tinguished, however, because of the leaf shape and type of venation. The type of forma b has smaller leaves but the range in size of leaves on the different specimens is so great that it is not believed that it merits recognition. 58. Piper thnuimucronatum CDC. in Smiths. Misc. Coll. 71^:12. 1920. A glabrous shrub, 1-3 m. tall; flowering internodes slender and rather short, pale-granular or glandular-dotted downward; leaves lance-ovate or elliptic, apex attenuatcly rather long-acuminate, commonly bristle-pointed, base subequilaterally acute, 3-5.5 cm. wide X 8-12 cm. long, pinnately nerved from below the middle, the nerves 3-4 on each side, with cross-connecting-anastomosing nervules, finely granular- or glandular-dotted beneath, drying green above, paler beneath, rather thin, translucent; petiole 10-15 mm. long, vaginate at the base only; spikes 3 mm. thick X -^-7 cm. long, apiculate; peduncle 1-2 cm. long; bracts triangular- or lunulate-subpcltate, margin conspicuously white-fringed; fruit globose or ovoid, laterally somewhat compressed; stigmas 3, sessile. P anama. CHIRIQUI: humid forest around Los Siguas Camp, southern slope of Cerro de la Hor- queta, alt. about 1700 m., Piltier JlSj; Maxon 542I; Bajo Chorro, Boquete District, alt. 1800 m., Davidson J2I, Jig; forest along the Rio Ladrillo and vicinity, about El Boquete, alt. 1200-1300 m., Uaxon $395 y Monniche Trail, Chiriqui Volcano, alt. 1700 m., Killil} 3523^ The shape, rather prominent bristle tips, and nervation of the leaves characterize thi IS species (45) IVoL. 37 46 ANNALS OF THE MISSOURI BOTANICAL GARDEN 59. Piper SEPTUPLiNERViuM (Miq.) CDC. in DC. Prodr. 16^313. 1869. Artanthe septnplinervia Miq. in Seem. Bot. Voy. Herald, 199, pL 40. 1854. Shrub (?), glabrous, strongly nodose; flowering intcrnodcs short and slender, smooth or somewhat scurfy-granular; leaves lance- or oblong-ovate, apex rather long and sharply acuminate, base equilateral, rounded, obtuse, 3-4.5 cm. wide X 9-12 cm. long, pinnately nerved below the upper third, the nerves 4-5 on each side, prominent beneath, with fine cross-connecting-anastomosing ncrvulcs, drying pale green, firm, narrowly revolutc, scarcely glandular-dotted; petiole about 5 mm. long, channelled above, vaginate at the very base only; spikes 4 mm. thick X 3-4 cm. long when mature; peduncle 5-10 mm. long; bracts round-subpcltate, glabrous in the center, marginally somewhat ciliate; fruit obpyramidal-trigonous, puberulous toward the top, stigmas 3, sessile. Panama. Panama: without location, Scvuiium lo66, colon: forests around Porto Bello, alt. 5-100 m., Viitkr 2477. 59a. Piper septuplinervium var. subpubinerviuni Yuncker, var. nov. Foliis 4.5-5 cm. latis, 12 cm. longis, luteo-glandulosis, vcnls supra pubcrulcntis subtus glabratis; bracteis triangulari-subpcltatis, margine luteo-fimbriatis. Leaves 4.5-5 cm. wide X 12 cm. long, yellow-glandular-dottcd, nerves above puberulent, glabrous beneath; bracts triangular-subpcltate, marginally yellow - fringed. Known only from the type locality. Panama: without locality, 18 59-1860, Hayes 78] (no 67S also on the bottom of label) (Herb. New York Bot. Gard., type). The glabrous, strongly nodose twigs, and glabrous, round-based leaves char- acterize this species which appears to be rather closely related to P. acqunle. 60. Piper subquinquenerve Trcl. in Field Mus. Nat. Hist. Bot. Ser. 18:363. 1937. A shrub (?), nodose, essentially glabrous; flowering intcrnodes slender and short; leaves lance-elliptic, apex acuminate, base subeo^uilatcral, acute, 2.5 3 cm, wide X S-10 cm. long, pinnately nerved from the lower half, the nerves 3 on each side, long-ascending, connected with the midrib with cross-connecting ncrvules, commonly with two pair near the base and the third pair arising at about the middle, yellowish and microscopically puberulent glabrescent beneath, drying rather thin and dull, translucent; petiole 5—12 mm. long, grooved above but vaginate only at the very base; spikes as yet young, 2 mm, X 3.5 cm. long; peduncle about 1.5 cm. long; bracts round-subpeltate, smooth above; ovary and fruit not developed. Known only from the following locality. BOCAS DEL TORO: Changuinola Valley, Dunlap 4j2. (46) 1950] FLORA OF PANAMA (Pipcraccae) 47 61. Piper infraluteum Trel, in Contr. U. S. Nat. Herb. 26:33. 1927. A glabrous shrub; flowering internodes slender and short; leaves lance-elliptic or somewhat oblong, apex acuminate, base inequilateral with one side 2-3 mm. shorter, acute, 3-4 cm. wide X 10-12 cm. long, pinnately nerved from the lower half, the nerves slender, 4-5 on each side, with fine cross-connccting-anastomosing nervules, glandular-dotted on both sides, drying yellow beneath; petiole 5-7 plus 2-3 mm. long, vaginate at the base; spikes 2 mm, thick X 4-5 cm. long; peduncle about as long as the petioles; bracts round- or triangular-subpeltate, strongly yellow-fringed; ovary subglobose, smooth, stigmas slender, sessile. Panama. Panama: without locality, Hayes 7p5. 62. Piper cricamolense Trel. in Ann. Mo. Bot. Gard. 27:290. 1940. A brittle, somewhat nodose shrub, 2-3 m. tall; flowering internodes short and relatively slender, microscopically strigulosc especially about the nodes, quickly glabresccnt; leaves elliptic or elliptic-obovate, apex abruptly sharp-acuminate, base subcquilateral, obtuse, 4-7 cm. wide X 9-13 cm, long, pinnately nerved from below the middle, the nerves 4-5 on each side, strongly curved and unbranched upward, salient beneath, with conspicuous cross-connecting nervules, finely nervu- lose by transmitted light, glabrous on both sides or microscopically strigulosc along the nerves, obscurely glandular-dotted beneath, drying papery, translucent; petiole scarcely 5 mm. long, microscopically strigulosc, vaginate at the base only; spikes 2-3 mm. thick X 5-7 cm. long; peduncle 5 mm. long, rather stout, glabrous or microscopically strigulosc; bracts triangular-subpeltate, marginally densely yellow- fringed; ovary narrowly ovoid-trigonous, yellow-glandular; stigmas 3, slender, sessi ile. Known only from the following locality. BOCAs DEL TORO: Rio Cricamola, between Finca St. Louis and Konklntoe, alt, 10-50 m., Woodson, AUch & Scibcrt Jg2'^. The more or less microscopically strigulosc twigs, low branching of the midrib, densely yellow-fringed bracts, and ovoid ovary characterize this species. 63. Piper aequale Vahl, Eclog. Amer. 1:4, /?/. 5. 1796. Scbilleria aequalis Kunth, in Linnaea 13:687. 1839. Artauthe aequalis Miq. Syst. Pip. 511. 1844. ?Pipcr Fuuckii CDC. in DC. Prodr. 16^:3 10. 1869. Viper substenocarpum CDC. in Bull. Herb. Boiss. 6:492. 1898. Piper coticin7inm CDC var. triuttense CDC in Urb. Symb. Ant. 3:304. 1902. Piper Jolmsfoni CDC in Proc. Amer. Acad. 40:68 5. 1905. Piper caledoniunum CDC in Fcdde. Rep. Sp. Nov. 15:1. 1917. Piper aequale var. substenocarpum CDC in Ann. Conscrv. & Jard. Bot. Geneve 21:253. 1920. Piper amphioxys Trel. in Contr. U. S. Nat. Herb. 26:31. 1927. Piper iba^uense Trel. in Trab. Mus. Nac. Hist. & Jard. Bot. Madr. Scr. Bot. 33:50. 1936, Piper zediogoJiense Trel. ex Ll. Williams, Expl. Bot. Guyana Venezol. 177. 1942. (47) [Vol. 37 48 ANNALS OF THE MISSOURI BOTANICAL GARDEN A glabrous, nodose shrub, 1-3 m. tall; flowering internodes slender and com- paratively short; leaves elongated-elliptic, lance-elliptic or ovate, 3.5-9 X 12-17 cm. long, apex more or less attenuate, bluntly acuminate, base cquilaterally acute or obtusish, pinnately nerved to the upper third or nearly throughout, the nerves mostly 4-5 or 6 on each side, with fainter intermediates and cross-connccting- anastomosing ncrvulcs, commonly 2 or 3 pairs approximated near the base and 1 or 2 pairs above the middle of the blade, the nerves often golden in dry specimens, drying rather firm, somewhat glossy, narrowly revolute, translucent; petiole about 5-10 mm. long or sometimes up to 1.5 cm. on lower leaves, commonly vaginate near the base but occasional leaves vaginate to the middle or above; spikes 2-4 mm. thick and up to 10 cm. long; peduncle commonly about as long as the petiole; bracts rounded dome-shaped when young with submarginal fringe, becoming round- or triangular-subpeltatc and loosely fringed; fruit small, obpyramidal- trigonous, convex at the top, glabrous; stigmas slender, sessile. West Indies, northern South America, and Panama, CANAL zone: Barro Colorado Island, Bailey ^ Bailey 4gj. colon: Loma do la Gloria, ncnr Fato (Nombre dc Dios), alt. 10-104 m., Pifticr 423Q. 63a. Piper aequale var. laurifolium (Trel) Yuncker, comb. nov. Pilfer amphioxy^ Trcl. var. JaurljoliiDU Trcl. in Cor^tr. U. S. Nat. Herb. 26:31. 1927. The nerves beneath very minutely hirtellous; the leaves mostly with 3 strongly ascending nerves on each side with two from near the base and the third from near or above the middle. Known only from the type locality. DARiEN: foothills of Garagara, Sambu basin, alt. 300-500 m., Vittier 559^ (U. S. Nat. Herb., TvrE). This species appears to be widely distributed in tropical America. Th Panamanian specimens here cited agree in all respects with those from South America and the West Indies believed to represent this species. 64. Piper sambuanum CDC. in Smiths. Misc. Coll. 71^:13. 1920. A shrub (?), nodose, glabrous; flowering internodes slender and short, obscurely granular and glandular-dotted; leaves lance-elliptic to obliquely lanceolate, apex ■ acuminate, base inequilateral with one side about 2 mm. shorter, obtuse or the longer side subcordulate, 5—6.5 cm. wide X 13-17 cm. long, pinnately nerved from below the upper third, the nerves about 4 on each side, or 5 with the lower- most obscure, with rather strong cross-connecting-anastomosing ncrvules, gland- ular-dotted, drying papery, translucent, pellucid-dotted; petiole about 3 plus 2 mm. long, stout, vaginate-winged to the lower blade margin; spikes 2 mm. thick X 8—10 cm. long; peduncle scarcely 1 cm. long, stoutish; bracts small, round- or triangular- subpeltatc, marginally ciliolate; ovary glabrous; stigmas sessile; fruit not developed. Known only from Sambu basin, Daricn. DARIEN: foothills of Garagara, Sambu basin, alt. 30-500 m., Vittier 366^, 5726, (48) 1950] FLORA OF PANAMA (Piperaccae) 49 The glabrous, glandular-dotted, nodose twigs, short-petioled leaves, and slender spikes characterize this species. 65. Piper TECUMENSE Trel. in Contr. U. S. Nat. Herb. 26:29. 1927. A shrub, scarcely 2 m. tall, essentially glabrous; flowering internodcs slender and elongating downward, glabrous or very slightly evanescently crisp-pubescent, slightly pubescent at the nodes; leaves elliptic, elliptic-ovate, or lance-ovate, apex gradually sharp-acuminate, base nearly equilateral, rounded, one side commonly about 2 mm. shorter, pinnately nerved from the lower half, the nerves about 4 on each side, submarginally loop-connected, with cross-connecting-anastomosing nervulcs, not glandular-dotted, 4-5.5 cm, wide X 10-14 cm. long, glabrous above, the nerves beneath obscurely and minutely appressed-pubescent or glabrescent; petiole scarcely 5 plus 2 mm. long, vaginatc at the base; stipules and stipular line slightly pubescent; spikes as yet young, 1,5 mm. thick X 3.5 cm. long; peduncle slender, 1 cm. long, glabrous; bracts rounded-subpeltate, yellow-fringed; ovary and fruit not developed. Known only from the following locality. PANAMA: Rio Tecumcn, Standley 2pjj8. 66. Piper crassispicatum Opiz in Prcsl, Reliq. Haenk. 1:152. 183 0. Art ant he crassispicata Miq. Syst. Pip. 521. 1844. A shrub (?), nodose, glabrous; leaves elliptic- or oblong-ovate, apex rather abruptly acuminate, base equilatcrally subacute, 10-13 cm. wide X 20-22 cm. long, pinnately nerved from below the upper fourth, the nerves about 7 on each side, drying subcoriaceous; petiole 1.5-2 cm. long, vaginate-winged downward; spikes 4 mm. thick X 5-6.5 cm, long; peduncle about 5 mm. long; bracts tri- angular-subpeltate, ciUate; fruit said to be obpyramidal-triquetous and stigmas sessi lie. Known only from the type specimen in the Prague herbarium collected by Haenke. An obscure species apparently distinguished by the rather large leaves and number of nerves. The type has not been seen. The description is based on notes of the type made by the late Professor Trelease from the specimen at Prague. 67. Piper bogotense CDC. in Jour. Bot, 4:214. 1866. / fist 1866. scesens CDC. in DC. Prodr. 16^:291. 1869, not Artanthe } Viper durirameiim CDC in Bull. Herb. Boiss. 6:484. 1898. Conoc [1902.] Viper oblitnm Trel. in Contr. U. S. Nat. Herb. 26:22. 1927. Viper bogotense CDC var. nsaqiiemanense Trel. in Trab. Mus. Nac. Hist. & Jard. Bot Madr. Ser. Bot. 33:48. 1936. Viper nsaquemense (Trel) Perez Arbelaez, PL Utiles de Col. 207, f, 2d^. 1947. (49) [Vol. 37 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN A shrub or small tree, up to 8 or 10 m. tall; flowering intcrnodcs comparatively short, sparsely pubescent or glabrate; leaves ovate, mostly 6-8 or 9 cm. wide X 10-15 cm, long, or Infrequently up to 15-18 X 20 cm., apex acuminate, or some- times rather blunt, base more or less inequilateral with one side often shorter, rounded, obtuse, cordate, or acute, pinnately nerved from the lower third, the major nerves 3 or 4 or in larger leaves 5 on each side, with reduced nerves upward, more or less pubescent above at least along the nerves, often somewhat glossy when dry, from densely crisp-pubesccnt to subglabratc beneath, but with the nerves mostly persistently rather densely crisp-pubescent, the hairs more abundant on the sides of the nerves, mostly densely ciHolate, glandular-dotted, drying rather dark, firm, translucent; petiole 1-1.5 or 2 cm. long, or on lower leaves up to 5 cm., mostly densely crisp-pubescent-tomcntose, vaglnate at the base; spikes 5-7 mm. thick and up to 8 cm. long but mostly somewhat shorter; peduncle about 10 mm. long, thinly crisp-pubescent glabrescent; bracts round- or trlangular-subpeltate, marginally fringed; fruit obovoid, stigmas 3-4, lanceolate, recurved, sessile. Northern South America and Panama. Panama: without location, Hayes So J. It has been impossible to find any differences between this specimen from Panama and those believed to represent P. hogoicnisc from South America. 6^. PiPHR SPERDiNUAi CDC. in Smiths. Misc. Coll. 71*': 1. 1920. A low shrub; flowering intcrnodes relatively short, densely villous with a mix- ture of scattered hairs up to 2 mm. or more long; leaves inequilaterally elliptic or subovate, the midrib much nearer one side, apex sharp and long-acuminate, base strongly inequilateral with one side 1-3 mm. shorter at the petiole and rounded- cordulate, the longer side aurlculate, the lobe 1-2 cm. long and covering the petiole, 6-9 or 10 cm. wide X 15-24 cm. long, pinnately nerved from below the middle, the nerves mostly 4-5 on each side, prominent beneath and with rather prominent cross-connecting-anastomosing ncrvules, submarginally loop-connected, glabrous above except for a few hairs on the midrib near the base, pubescent beneath w^ith the nerves densely so with an Intermixture of spreading long hairs, margin not ciliate, obscurely glandular-dotted, drying rather papery, translucent; petiole about 3—5 plus 1—3 mm. long, densely hairy, vaglnate at the base; spikes about 3 mm. thick X 6-7 cm. long, with a slender, bearded, aristate tip up to 1 cm. long; peduncle up to 2 cm. long, densely pubescent; bracts truncately triangular-sub- peltate, with usually 2-4 rather prominent dorsal setae, or glabrous, the pedicel margins setose; rachis ridges fimbriate; fruit obpyrlform- trigonous, granular- papillate, stigmas slender, sessile. Near Puerto Obaldia^ San Bias Coast. SAN BLAs: hills back of Puerto Obaldia, alt. 50-200 m., ?ittier 4301^ 4348. (50) 1950] FLORA OF PANAMA (Pipcraceac) 51 A very well-marked species with its dimorphic pubescence, auriculate leaves, and papillate fruit. 69. Piper perbrevicaule Yuncker, sp. nov. Suffrutex, ca. 15 cm. altus, caudice conferte villoso, pills 1-2 mm, longis; foliis rhombico-elliptico-obovatis, apice abrupte brcviter acuminatis, basi inacqualiter subcordato-auriculatis, lobo longiore pctiolum obtegente, utrinque longe villosis, pilis 1 mm. vel ultra longis, subtus ad venas patentibus, a medio infcro pinnatim venosis, venis 4 ~\- 5—6; petiolo baud 1 cm. longo, dense longe villoso, sub base vaginato; spicis 1-2 mm. crassis, 3 cm. longis; pcdunc- ulo 3—5 cm, longo, dissitc villoso; bractcis triangulari- subpeltatis, glabra tis; drupa ovoideo-subglobosa sub- acuminata; stigmatibus gracilibus scssilibus. A very small subshrub, 15 cm. or less tall; stem drying somewhat ridged, densely villous, the hairs 1 mm. or more long; intcrnodcs mostly 1-2 cm. long; leaves somewhat pustular above, obliquely clliptic- obovate-rhomboidal, the midrib much nearer one side, 2.5-4 cm. wide X 7-9 cm. long, apex abruptly short- acuminate, base subequal at the petiole, the shorter side obtuse or cordulate, the longer side auriculate with the lobe, about 1 cm. long and covering the petiole, long- vlllous on both sides, the nerves beneath more strongly so with the hairs essentially erect and 1 mm. or more long, pinnately nerved from the lower half, the nerves 4 + 5-6 with cross-con- necting-anastomosing nervules, drying rather thin, green, translucent; petiole scant 1 cm. long, densely long-villous, vaginate toward the base; spikes red, purple, or olive-green when fresh, 1-2 mm. thick X 3 cm. long; peduncle 3-5 cm. long, slender, sparsely villous glabrescent; bracts triangular-subpcltate, glabrous; fruit very small, ovoid-subglobose with pointed apex, glabrous; stigmas slender, sessile. Known only from the type locality. BOCAs DEL TORo: vicinity of Chiriqui Lagoon, Water Valley, Oct. 1, 1940, ro/; Wald gyi (Herb. Missouri Bot. Gard., type); Nov. 1, 1940, von WcJcI 146Q, 69a. Piper PERiiRLVicAULE var, sub^labrilinibum Yuncker, var. nov. Foliis supra glabratis, subtus ad venas baud conferte villosis, pilis baud 1 mm. 1 Fig. 9. Viper pcrbrei imiile ongis. Leaves 4.5 cm. wide X 8-9 cm. long, upper surface of blade glabrous, the nerves beneath thinly villous, the hairs less than 1 mm. long. Known only from the type locality. BOCAS DEL TORO: Vicinity of Chiriqui Lagoon, Oct. 8, 1940, 1077 Wcclel IOJ4 (Herb. Missouri Bot. Gard., type). The very small plants, long-villous stems, and strongly auriculate leaves are distinctive characteristics of this species. rn; [Vol. 37 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN 70. Piper leptocladum CDC. in Anal. Inst. Fis.-Geogr. Costa Rica 9:164. 1897. Piper Piper Jilatatum Rich. var. leptocladum CDC. in Candollea 1:96, 1923. diazanum Trel. In Contr. U. S. Nat. Herb. 26:34. 1927. Piper diazanum var. liae-koheaiiae Trel. in Ann. Mo. Bot. Gard. 27:290. 1940. Piper tabcmillannm Trel. !oc. cit. 298. 1940. A slirubj up to about 3 m. tall; flowering internodes slender and rather short, 1 sparsely crisp-hairy; leaves subrhombically elliptic, lanceolate, or subovate, smootl or slightly rugose, occasionally slightly scabrous, apex gradually sharp-acuminate, base rounded, obtuse to cordulate, or subcordate, equal at the petiole or with one side sHghtly shorter, 4—5 or sometimes 7 cm. wide X 10-17 cm. long, pinnately nerved from the lower half, the nerves 4—5 on each side, puberulent above along the nerves, rather sparsely appresscd- or crisp-pubescent on the nerves beneath, glandular-dotted, drying rather thin, green, translucent; petiole 5—10 mm. long, crisp-villous, vaginate at the base; spikes 3 mm. thick X 6-9 cm. long; peduncle about 5 mm. long, pubescent or glabrate; bracts triangular-subpeltate, yellow- fringed; fruit obpyramidal-trigonous, glabrous, green to dark-colored, stigmas small, sessile. Panama and Costa Rica. CANAL zone: vicinity of Fort Sherman, Standley jopjjj 30Q74y 3^99^1 Monte Lirio, Maxon 6S43; Matachin to Las Cascadas, Cowell 33^> L^s Cascadai Plantation, near Sum- mit, Standley 2^/^q8, 2g62^; Fort San Lorenzo, Fort Shcrmnn Military Reservation, Maxofi ^ Valentine J012 — the leaves on this specimen are smaller and less pubescent than cus- tomary for this species but it appears to agree in all other characters; Barro Colorado Island, Bailey ^ Bailey jj, 104, 242; Kenoyer 307, 308; Standley 40862, 410JJ; Cerro Gordo, near Culebra, Standley 26006; around Tabernilla, along the railroad, alt. 20-25 m., Pittier 3828; Empire, Miller 176$; Mt. Hope Cemetery, Standley 2SS02; Obispo, Standley 31671; Fort Kobe road, Woodson, Allen ^ Seibert 1412. colon: forests around Porto BcUo, alt. 5-100 m., Pittier 242S, panama: Juan Diaz, deep woods, KtlUp 3114, Thi IS species ra the closely rcscmDics bl P, dilatatnm of South America but differs because of its glabrous fruit and other minor characters. 7 1 . Piper dimorphotrichuni Yuncker, sp nov. Fig. 10. Piper Jimorpbotrichum Frutex, 3 m. altus, Internodiis floriferis sat brevibus crassisque, conferte subrctrorse brunneo- hirsutis, pilis brevibus longisque intermixtis, pilis longis ad 3—4 mm. vel. ultra longis; foliis late ellipticis, apice attcnuatim acuminatis, basi ob- lique cordulatis vel subaunculatis, lobis sub- aequalibus vel lobo longiore petiolum fere acquantc, a medio infcro pinnatim venosis, venis utrinque 5—8, trabibus gracilibus anastomosanti- bus, supra sparse villosis, subtus pubescentibus (confertius ad vcnas) , pilis erectis, heteromorphis, brevioribus longioribusque intermixtis; petiolo vix 1 mm. longo, a medio deorsum vaginato; 9 ; 19 50] FLORA OF PANAMA (Pipcraccae) 53 spicis 5 mm. crassis, 5-6 cm. longis; pedunculo gracili, 4-6 cm. longo, villoso; bracteis triangulari-subpeltatis, margine fimbriatls; drupa oblonga lateraliter com- pressa, glabrata, glandulosa, stigmatlbus sessllibus. A slirub, up to 3 m. tall; flowering internodes ratber sbort and slender, densely subretrorsely brown-bairy, tbe hairs of dififerent lengths up to about 1 mm. with intermixed scattered long-spreading hairs up to 3-4 mm. or more; leaves broadly elliptic, apex attcnuately acuminate, base obliquely cordulate or subauriculate, sub- equal at the petiole, the lobes subequal or one larger and about as long as the petiole, 5-10 cm. wide X 15-20 cm. long, pinnately nerved from below the middle, the nerves 5-8 on each side, approximated dowriward, with prominent, slender, cross- connecting-anastomosing nervules, sparsely villous above, staring pubescent beneath, the nerves more densely so, interspersed with scattered long hairs, drying thin, brown, translucent; petiole scarcely 1 cm. long, densely villous with mixed long hairs, vaginate below the middle; spikes 5 mm. thick X 5—6 cm. long when matvirc, with a slender, seta-like apex up to 7 mm. long; peduncle 4-6 cm. long, villous with an intermixture of long hairs; bracts triangular-subpeltate, marginally fringed with rigid hairs, those on the dorsal margin somewhat longer; drupe compressed- oblong, glabrous, glandular, stigmas slender, sessile. Known only from the type locality, BOCAS DEL TORo: vicinity of Chiriqui Lagoon, vo7i Wedel lOIj; Water Valley, von Wedel 144^ — this specimen is not as densely pubescent as the others; Nov. 9, 1940, van Wedel IS6q (Herb. Missouri Bot. Gard., type). The wide difference in the length of the hairs, rather large, subauriculate leaves, setose spikes, and long slender peduncles are distinctive characters of this species. 72. Piper pseudo-garagaranum Trel. in Contr. U. S. Nat. Herb. 26:28. 1927. Shrub, 1—2 m. tall, nodose; flowering internodes short and slender, moderately to sparsely villous, glabrescent downward, hairs up to 1 mm. or more long, some- what granular and glandular-dotted; leaves eHiptic or elliptic-lanceolate, apex rather long-acuminate, base rounded, inequilateral, one side 2—4 mm, shorter, obtuse, the longer side cordulate, pinnately nerved from the lower half, the nerves mostly alternate, 4 or in larger leaves 5 on each side, with slender cross-connecting and anastomosing nervules, 4.5-7.5 cm. wide X 14—20 cm. long, smooth or very slightly scabrous, sparsely long-villous above, loosely long-villous beneath, at least along the nerves, the hairs mostly erect, drying papery, glandular-dotted, trans- lucent; petiole 2-4 plus 2 mm. long, villous, vaginate at the base; spikes 3 mm. thick X 7.5 cm. long; peduncle scarcely 1 cm. long, granular, glabrous; bracts round-subpcltate, moderately fringed, a few dorsal hairs longer; fruit obpyramidal- trigonous, glandular-dotted, apex truncate, glabrous, stigmas sessile. Panama. CANAL zone: Barro Colorado Island, Standlcy 3l40g, 41 164, darien: Cana-Cuasi Trail (Camp 2), Cheplgana Distr., alt. 600 m., Terry ^ Terry 14^/^. (^V [\'(.L. 11 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN The specimens cited differ considerably In the size of the leaves. Those on No. 4.1 164 are scarcely 5 cm. wide X 14-15 cm. long while those on No. 142^ are 7-8 cm. wide X 18-20 cm. long. No. 3^40g is intermediate in size. The description of the spike characters are from Terry & Terry 142J. The spikes on the other two specimens are very young and do not exhibit ovary or fruit characters. All speci- mens agree otherwise. 73. Piper cativallnse Trel. in Contr. U. S, Nat. Herb. 26:34. 1927. Shrub, 2-3 m. tall; flowering intcrnodes slender, rather elongate, villous; leaves elliptic to subobovate, apex rather abruptly acuminate, base inequilateral, cordulatc, one side slightly shorter, 6-7 cm. wide X 12-14 cm. long, pinnately nerved from the lower half or two-thirds, the nerves mostly 5 on each side, with cross-connect- ing and anastomosing ncrvules, somewhat glossy green, very slightly scabrous and sparsely villous above, the nerves scarcely hairy, villous beneath, especially along the nerves, marginally somewhat ciliate, glandular-dotted on both sides, drying thin, translucent; petiole 5-10 plus 2 mm, long, villous, vaginate toward the base; inflorescence not known. Known only from the following locality. COLON: between France Field, Canal Zone, and Catival, Stafidlcy JO206, The absence of inflorescence leaves the position of this species somewhat questionable. 74. Piper psEUDO-LAXCEAFroLiuM Trel. in Contr. U.S. Nat. Herb. 26:170. 1929. Pifjcr lirafificrve Trel. in Ann. Mo. Bot. Gard. 24:186. 1937. A shrub, 2 m. or more tall; upper intcrnodes slender, sparsely to densely apprcssed-villous; leaves sublnequllaterally lanceolate, apex gradually sharply long- acuminate, base ineciuilateral with one side about 2 mm. shorter, obtuse, cordulate, or sometimes acutish, mostly 4-5 cm. wide X 14—17 cm. long, pinnately nerved from the lower half, the nerves mostly 7-8 on each side, impressed above, salient beneath, veiny, rugcscent and subappresscd-pubesccnt above, scarcely lacunosc beneath, appressed-pubesccnt, the nerves densely so, obscurely pale glandular-dotted, drying firm, translucent; petiole about 5 plus 2 mm. long, vaglnatc-wingcd to near the blade, villous; spikes 2-3 mm. thick X 8-12 cm. or more long, aplculate, curved; j.x,'duncle up to 2—3 cm. long, pubescent; bracts triangular-subpeltate, densely yellow-fringed; fruit obpyramidal-trigonous, dark, apex truncate, glabrous, stigmas slender, sessile. Costa Rica and Panama. Panama: without locality, Ni'e JjQ, 400. cjiiriqui: valley of the upper Rio Chiriqui Vicjo, vicinity of Monte Lirio. alt. 13 00-1900 m., Scibert 1 58 ; vicinity of Casita Alta, Volcan de Chiriqui, alt. 1500-2000 m., Woodwu, Allen & Scibert Qio. (54) 1950] FLORA OF PANAMA (Pipcraceac) 55 This species resembles P. lanceaefoVnim In most characters and may be too close to itj but it differs because of its less but more coarsely rugose leaves which are scarcely lacunose beneath, and with the base much less cordulate-aurlculatc. 75. Piper linearifolium CDC. in Linnaea 37:355. 1872. Viper ahcafum Trel. in Ann. Mo. Bot. Gard. 27:287. 1940. Piper arctilimbiim Trel. loc. cit. 288. 1940. Piper arctilunbum var. Alleni Trel. loc. cit. 1940. A low, bushy shrub, scarcely 2 m. tall; flowering internodes moderately slender and short, gray-pubescent; leaves lanceolate, apex gradually long- and sharp- acuminate, base inequilateral with one side 2—3 mm. shorter, obtuse, cordulatc, or some leaves acute, mostly 2-3.5 cm. wide X 10-17 cm. long, subrugescent in age, pinnately nerved from the lower third, the nerves 4-5 or occasionally 6 on each side, prominent beneath, somewhat impressed above, subequally spaced, subparallel and sharply ascending from the midrib, appressed-silky-pubescent on both sides, the nerves more densely so beneath, glandular-dotted, drying rather firm, trans- lucent, pellucid-dotted; petiole 3-5 plus 2-3 mm. long, pubescent, vaginate to near the blade; spikes 2-3 mm, thick X 5-9 cm. long, generally curved; peduncle 5-15 mm. long, pubescent; bracts triangular-subpeltate, densely yclIow-fringcd; fruit obpyramidal, 3- or 4-sidcd, glabrous; stigmas small, sessile. Costa Rica and Panama. BOCAS DEL TORo: vicinity of Nicvecita, alt. 0-150 m., Woodson, Allen ^ Seibert iSj/; Chiriqui Trail, von WeJel 21/6. chiriqui: vicinity of El Boquetc, alt. 1000-13 00 m., Maxon 5^4^'^ Killip 3^3^ > Isla dc Coiba, Mendez 122; rocky plains about 5 miles south of Boquete, alt. 900 m., Allen 4705. cocle: El Vallc, Miller 1813; vicinity of El Vallc dc Anton, alt. 600 m., Allen 2003; Penonomc and vicinity, alt. 15-300 m., Williavis 54, colon: along Rio Fato, alt. 10-100 m., Pitiier 388 1, F The narrowly lanceolate, silky-pubescent leaves, with sharply ascending nerves, curved spikes, and yellow-fringed bracts distinguish this species. 76, Piper panamense CDC. in Jour. Bot. 4:216. 18 66. Piper citrifolium Lam. var. panamense CDC. in Candollea 1:114. 1923. A shrub; flowering branches sliort, nodose, rather slender, densely subretrorscly crisp-pubescent laterally below the petioles, glabrcscent downward, glandular- papillate; leaves elliptic-ovate, apex acute to short-acuminate, base with qne side slightly shorter at the petiole, rounded, obtuse, or one or both sides acutish, pin- nately nerved below the upper third, the nerves 4-5 on each side, with cross- connccting-anastoniosing nervules, 5-7 cm. wide X 1 0-1 5 cm. long, glabrous above, pviberulcnt beneath, at least along the nerves, glandular-dotted, drying firm and somewhat glossy, translucent; petiole scarcely 1 cm. long, rather densely sub- retrorscly ptibcscent, vaginate near the base; spikes 3 mm. thick X 4-5 cm. long; peduncle about 5 mm. long, pubcrulent; bracts triangular-subpeltate, ciliatc; fruit oblong, compressed, apex truncate, depressed about the stigmas, strongly papillate- pubcrulent; stigmas sessile. 0^) [Vol. 37 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama. BOCAS DEL TORO: Laguna de Chlriqui and vicinity, Hart 124. canal ZONE: Chagrcs, Fcndlcr 2yo. The densely papillate fruit and comparatively short, stout spikes distinguish this species, 77. Piper callibracteum CDC. in Smiths. Misc. Coll. 71^:13. 1920. Viper charnhsonis (Miq.) Stcud. var. rubcUibractcum CDC. loc. cit. 1920. Piper s24bfJ7idibracteHni CDC loc. cit. 14. 1920. Piper nitidifolium CDC loc. cit. 1920. A shrub, 1.5-3 m. tall; flowering internodes rather slender and short, glabrate, pale-dotted; leaves elliptic-lanceolate or subovate, apex acuminate, base inequilateral with one side mostly slightly shorter, acute on both sides or one side obtusish, 5-8 cm. wide X 13-22 cm. long, smooth or slightly rugose, pinnatcly nerved from the lower half, the nerves 4 or sometimes 5 on each side, with cross-connecting and anastomosing nervules, glabrous above, obscurely appressed-pubcscent beneath, at least on the smaller nerves, drying green, slightly paler and pale-glandular-dotted beneath; petioles up to 3 cm. long, glabrous, vaginate toward the base; spikes 3 mm. thick X 8-10 cm. long, short-inucronate; peduncles 1-1.5 cm. long, glabrous; bracts rather small, triangular-lunate-subpeltate, somewhat umbonate, sparsely short-fringed; fruit compressed, white-papillate-puberulent at the top, stigmas sessi lie. Western Panama. CHiRiQuf: forests around El Boquete, alt. 1000-1300 m., Pitiier 2040; Maxon 49431 patter 28gS, 28gg; between Ccrro Vaca and Hato del Loro, alt. 850-1100 m., PitHer SJQO, The petioles of the type of P. siibmidibractcum are scarcely 1 cm. long and the leaves are slightly more pubescent along the nerves beneath than are the types of the other species included here. All agree, however, in the pubescent ovary and fruit, small, scarcely fringed bracts, and other characters to such a degree that it is believed they all represent a single species which usually has petioles 1-3 cm. long, and nerves scarcely pubescent beneath. 78. Piper Wedelii Yuncker, sp. nov. Frutex, 1—2 m. altus; internodiis floriferis gracilibus brevibusque, delicate costulatis, dissitc hirtulis; foliis subrhombico-ellipticis, apice longius acute acumi- natis, basi oblique, latere altero obtuso, ad petiolum 3 mm. breviore, caetero rotundato cordulato, a medio deorsum pinnatim venosis, venis utrinquc 5—6, supra glabratis vcl dissitc scabridis, subtus ad venas sat dissite substrlgosis; petiolo ca. 5 cum 3 mm. longo, ad basem vaginato; spicis immaturis, 2 mm. crassis, 6 cm. longis, mucronatis; bracteis triangulari-subpeltatis, glabratis, dorso papilloso-callosis; drupa ignota. A tree-like shrub, 1-2 m. tall; flowering internodes short and slender, finely ridged when dry, granular-roughened, young growth obscurely hirtellous; leaves (^6) 1950] FLORA OF PANAMA (Piperaceac) 57 subrhomboidal-elliptic, apex sharply long-acuminate, base oblique, one side 3 mm. shorter at the petiole and obtuse, the longer side rounded, cordulate, 9-10 cm. wide X 2 0-2 2 cm. long, pinnately nerved from the lower half, the nerves 5-6 on each side with slender cross-connecting-anastomosing nervules, glabrous or shghtly scabrous above, the nerves beneath rather sparsely substrigose, glandular-dotted on both sides, drying thin, translucent; petiole about 5 plus 3 mm. long, vaginate at the base; spikes as yet young, red when fresh, 2 mm. thick X 6 cm. long, mucronate; bracts triangular-subpeltate, glabrous, with a dorsal papillate callus; ovary and fruit not present. Known only from the type locality. BOCAS DEL TORo: vicinity of Chiriqui Lagoon, Oct. 2, 1940, van Wedel I02I (Herb. Missouri Bot. Gard., type). 78a. Piper Wedelii var. retrohirsutum Yuncker, van nov. Tnternodiis floriferis retrorse hirsutis; foliis elHpticis, 4-5 cm. latis, 11-13 cm. longis, supra scabridis obscure hispiduHsque, subtus ad venas strigosis, glanduloso- punctatis. Flowering internodes retrorsely hirsute; leaves elliptic, 4-5 cm, wide X 11-13 cm. long, obscurely hispidulous and scrabous above, the nerves beneath strigose, glandular-dotted. Known only from the type locality. BOCAS DEL TORO: vicinity of Chiriqui Lagoon, Water Valley, Nov. 9, 1940, von Wedel 1375 (Herb. Missouri Bot. Card., type). This species resembles P. ertcbescentispicum In the appearance of the spikes but it differs because of its larger, non-vlllous, conspicuously glandular, and scarcely scabrous leaves. The dorsally calloused floral bracts are rather distinctive. The smaller leaves and strongly retrorsely hirsute twigs contrast the variety. 79. Piper verruculaepetiolum Trek in Field Mus. Pubk Bot. 18:368. 1937. Shrub, nodose; flowering Internodes comparatively short and slender, glabrous, pale granular-dotted; leaves obliquely ovate-elliptic, apex short-acuminate, base obliquely inequilateral, one side 2-3 mm. shorter, the longer side strongly cordulate, 7-10 cm. wide X 17-24 cm. long, pinnately nerved below the upper third, the nerves 6-8 on each side, with cross-connecting and anastomosing nervules, glabrous above, very finely puberulent on the nerves beneath, yellow glandular-dotted on both sides, drying thin, translucent; petiole 7 plus 2-3 mm. long, strongly granular, glabrous or sparingly puberulent, vaginate at the base; spikes 3 mm. thick X 7-8 cm. long; peduncle about 5 mm. long, glabrous, granular; bracts triangular-sub- peltate, marginally yellow-fringed; fruit trigonous, glabrous, glandular-dotted, truncate; stigmas sessile. Known only from the following locality. BOCAS DEL TORO: Farm Six, Changuinola Valley, Dunlap 494. 07) I Vol. i7 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN 80. Piper hirtellipetiolum CDC. in Smiths. Misc. Coll. 71*^:3. 1920. A conical, tree-like shrub, L5-2 m. tall, nodose; flowering internodcs very short and slender, glandular-dotted, glabrous or at first with a few scattered hairs; leaves lanceolate, apex gradually acuminate, base subinequllatcral, one side 1-2 mm. shorter, acute or obtusish, 2.5-3.5 cm. wide X 9-13 cm. long, pinnately nerved from below the middle, the nerves 3-4 on each side, loop-connected upward, with cross-connecting and anastomosing nervulcs, glabrous on both sides or the midrib sparsely hairy near the base, drying light green, glossy above, pale glandular-dotted beneath; petiole scarcely 5 plus 2 mm. long, loosely villous, vaginate at the base; spikes 3 mm. thick X 4-5 cm. long; peduncle 5 mm. long, puberulent above the middle; bracts triangular-subpcltate, copiously yellow-fringed, usually annular, alternating with the dark fruits; fruit rounded-obovoid (?) , glabrous, black; stigmas small, sessile. Known only from David, Chiriqui. cuiRiQuf: vicinity of David, alt. 30-80 m., Vitiicr 28j2, 80a. Piper hirtellipetiolum van tapianum (Trel.) Yunckcr, comb. nov. Piper tapiuNinu Trek in Contr. U.S. Nat. Herb. 26:30. 1927. Shrub, 3 m. tall; twigs moderately to rather densely crisp-villous; leaves 3.5-6.5 cm. wide X 10-16 cm. long, glabrous and glossy above, strongly glandular- dotted beneath with the nerves crisp-pubescent; petioles crisp-hairy; spikes 5-6 cm. long. Central Panama. PANAMA: near Tapia River, Juan Diaz region, June 1-3, 1923, Maxo/i ef Harvey 6jog (U. S. Nat. Herb., typl) ; Rio Tapia, StiiHillcy 26122; Pcrlas Archipelago, San Jose Island, Johnston 1360; Erlanwn Ij6, sob. Piper HiRTtLLiPETioLUM var. Harveyanuni (Trek) Yuncker, comb. nov. Viper subtrinerve var. Uarieycvunn Trek loc. cit. 30. 1927. Viper suhtriucrve Trek in Contr. U. S. Nat. Herb. 26:29. 1927. Viper perfugii Trcl. in Ann. Mo. Bot. Card. 27:295. 1940. An essentially glabrous, somewhat nodose shrub, 1,5-3 m. tall; leaves lance- elliptic, 2-4.5 cm. wide X 7-15 cm. long, glabrous or rarely with a few scat- tered short hairs on the nerves beneath; petiole glabrous; spikes 3-4 mm. thick X 4—6 cm. long; peduncle with a few short hairs near the spike. Fig. 11. Viper hirtcUipctiolum var. liuricyauMm (58) 1950] FLORA OF PANAMA (Pipcraccac) 59 Panama. BOCAS DEL TORo: Chinguinola Valley, Dinilap 2JI — the spikes of this specimen are apiculate. chiriqui: vicinity of Casita Alta, Volcan dc Chiriqui, alt. 1500-2000 m., Wooihon, Allen ^ Seibcrt Q2S — this specimen has leaves somewhat smaller than usual. cocle: El Valle, Miller 1814, canal zone: Las Sabanas, Bra, Celesfine HO; foot of Cor- ozal Hill, Miller 1790, Panama: near Tapla River, Juan Diaz region, Maxon & Harvey 6684, 6/'J4; Juan Diaz, Standley J0600; along the Corozal Road, near Panama, StauiUey 26854] Tumba Mucrto Road, near Panama, Standley 2gjlO; Punta Paitilla, Piper 5410; east of Rio Tecumcn, Standley 26j6j; between Las Sabanas and Matias Hernandez, Stand- ley 3l8j6; Taboga Island, Standley 2'/86g; Pittier 3618 (U.S. Nat. Herb., type); Perlas Archipelago, San Jose Island, Johnston 81, 754' The specimens included under this species, including the varieties, all agree to such a degree that it docs not appear advisable to maintain the varieties as separate species. The chief distinction appears to be in the character and amount of pubescence. 8L Piper subcaudatum Trel. in Contr. U. S. Nat. Herb. 26:3 L 1927. Shrub (?), nodose; flov/ering internodes slender and short, somewhat ridged and striate-pilose, especially below the petioles; leaves obliquely elliptic or elliptic- obovatc, apex abruptly subcaudately long-acuminate, tip about 2.5 cm. long, base narrowed downward, obtuse or subcordulate, one side about 3 mm. shorter, 6-8 cm, wide X 13-16 cm. long, pinnatcly nerved below the upper third, the nerves 5 or 6 on each side, with cross-connccting-anastomosing ncrvules, submarginally loop- connected upward and, excepting the innermost pair, not continuing to the apex, glabrous above, the nerves beneath appresscd-puberulent, botK sides glandular- dotted, drying somewhat glossy, thin, translucent; petiole about 5 plus 3 mm. long, hairy-lined beneath, vaginate near the base; spikes as yet very young, 1 mm. thick X 1 cm. long; peduncle sparsely hairy; bracts rounded-subpeltate, marginally ciliate; ovary and fruit not developed. Known only from Sambu basin, Daricn, darien: foothills of Garagara, Sambu basin, alt. 30-5 00 m., Pittier 56/2. 81a. Piper subcaudatum var. maternale (Trel.) Yuncker, comb, nov. Piper maternale Trel. in Field Mus. Publ. Bot. 18:349. 1937. Leaves somewhat oblique, elliptic or elliptic-subovate, apex tip about 1 cm. long, base more strongly cordulate. Panama. BOCAS DEL TORo: vicinity of Almirante, Cooper l8g^ 2IO (Herb. New York Bot. Gard. and Herb. Univ. 111., isOTYrLs) ; Changuinola Valley, Bar Mouth, Dufilap jj6; Old Bank Island, von Wedel 2047, The shape and venation of the mostly cordulate-based leaves characterize this species. It is to be distinguished from P, C7ilcbra)iiini chiefly because the lateral (^9) 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. n nerves do not continue toward the apex as in that species. There is some question whether var. viatcrnale differs sufficiently from the species to merit recognition. The leaf shape and shorter apex are the main differential features, 82. Piper gonocarpum Trel. in Contr. U. S. Nat, Herb. 26:28. 1927. A shrub, 2 m. tall; flowering intcrnodes moderately slender and elongate, densely rctrorsely dingy-pubescent ; leaves lance-elliptic or suboblong, apex gradually acuminate, base somewhat Inequilateral, rounded or cordulate, one side slightly shorter, 5-6.5 cm. wide X 15-17 cm. long, pinnately nerved from below the upper third, the nerves about 6 on each side, glabrous above, the midrib impressed below the middle, hirsute beneath, the nerves rather densely so, the hairs staring, drying rather thick, subtranslucent, glossy above, narrowly revolute; petiole about 10 plus 2 mm. long, densely pubescent, vaglnate to the blade; spikes 7-8 mm. thick X 4 cm. long, caudate at the tip; peduncle scarcely 1 cm. long, sparsely hairy; bracts broad, narrowly lunate-concave, glabrous; fruit depressed-quadrangular, 2-3 mm. in diameter, ridged, granular, glabrous; stigmas 2, broad, sessile. Canal Zone, CANAL ZONE: hills north of Frijoles, Standley ^7437- The short, thick spikes, and large, angular fruits are distinctive of this species. V. KEY TO SPECIES WITH LEAVES PINNATELY NERVED TO THE UPPER FOURTH OR THROUGHOUT. a. Midrib branches 10—15 on each side, subcqually spaced; leaves mostly 10 cm. or more wide; spikes mostly 8-10 mm. tlilck 83. P. AUGUSTUM aa. Midrib branches usually less than 10 and commonly variously spaced; leaves mostly narrower. b. Leaf base strongly inequilateral with one side mostly 5—10 mm. shorter at the petiole. c. Leaves somcwliat scabnd; stems villous; petiole vaginatc to the middle. 84. P. Steve NSi CC. Leaves not scabrid; stems glabrous or puberulcnt; petiole vaginate- winged to the blade, the wings often protracted at the blade. d. Leaves 25 cm. or more long; nerves 6—7 on each side 85. P. colonfnse dd. Leaves mostly less than 20 cm. long; nerves 8—10 on each side. e. Leaf apex very short and abruptly acute; leaves mostly less than 12 cm. long ...86. P. tubmrculatum ee. Leaf apex gradually acute to acuminate; leaves commonly 15 cm. or more long (shorter in var.) 87. P. arboreum bb. Leaf base mostly essentially equilateral or sometimes oblique or with one side slightly shorter. c. Leaves mostly 7-9 cm. long; spikes scant 2 cm. long 88. P. subrepens CC. Leaves mostly 10-15 cm. or more long; spikes longer (short In P. cafiilflarianjim) , d. Petiole strongly vaglnate-wiaged to the blade; leaves bronzed beneath, nerves prominently submarginally loop -connected; peduncle scarcely 5 mm. long S9. P. cordulatum dd. Petiole usually not vaginate-wingcd to the blade though some- times grooved above; leaves green or brown but not bronzed, e. Bracts cupulate, glabrous; fruit free on the rachis, exscrted, very short stylose 3. P. darienense cc. Bracts more or less peltate; fruit congested on the rachis. (60) 1950] FLORA OF PANAMA (Pipevaceae) f. Leaves mostly less than 5 cm. wide. g. Leaves obviously glandular-dotted beneath. h. Spikes 5 cm. or more long; stigmas sessile 90. hh. Spikes less than 3 cm. long; fruit short-stylose 12. gg. Leaves not obviously glandular-dotted. h. Stems glabrous 91. hh. Stems densely hairy.., 92. if. Leaves 6 cm. or more wide 95, 61 P. ACUTISSIMUM P. CANDELARIANUM P. PERLASENSE P. VIA-CHICOENSE P. GRANDE 83. Piper augustum Rudge, PL Gulan, Rar, 1:10, pL /. 1805. Piper anguslum Rudge ex Spreng. Syst. Veg. 1:108, 1825, sphalm. Arfanfbc augusta Miq. Syst. Pip. 398. 1844. Artanfbe calocoma Miq. In Seem. Bot. Voy. Lierald, 199. 18 54. Viper pediceUare CDC in Jour. Bot. 4:211. 1866. Piper calocama CDC. in DC Prodr. 16^-264. 1869. A glabrous sKrub, 2-3 m. tall, or small tree up to 8 m.^ branches somewhat nodose, with moderate intcrnodes; leaves elliptic-ovate, 9-15 or 18 cm. wide X 15-3 5 cm. long, apex acute, base acute or obtusish, usually subequilatcral, pinnately nerved throughout, the nerves 10-16 on each side, not strongly curved upward, densely ciliate, strongly glandular-dotted, drying membranous, translucent; petiole 5—10 mm. long or up to 4 cm. on lower leaves; spikes when young 3 mm. thick X 4-5 cm. long, becoming 1-1.5 cm. X 8-12 cm. when mature; peduncle 1-2 or 3 cm. long; bracts sublunate-triangular-subpeltate, marginally yellow-fringed; fruit oblong, compressed-angular, truncate; stigmas sessile. Northern South America and Panama. COcle: Bismarck, above Penonome, alt. 600-900 m., Williams 332, 611, canal zone: In forest along the Rio Indio de Gatiin, Pittier 27go^ 2jgi. 83a. Piper augustum var. cocleanum (Trel.) Yunckcr, comb. nov. Piper polyneurum CDC in CandoIIea 1:270. 1923, name only in index; Schroeder, loc. cit. 3:138. 1926. Piper gatunense Trel. in Contr. U. S. Nat. Herb. 26:27. 1927. Piper ladrillense Trel. loc. cit, 1927. Piper gatunense var. cocIea?iHm Trel. in Ann. Mo. Bot. Gard. 27:292. 1940. Piper gafiinensie var. latum Trel loc. cit. 1940. Upper internodcs sparsely to rather strongly puberulent or pubescent; leaves above from glabrous to rather strongly white-villous, puberulent to villous beneath, at least along the nerves. Panama. CHiRiQui: forests along the Rio Ladrillo and vicinity, above El Boqucte, alt. 1200-1300 m., Pittier 3164, cocle: north rim, vicinity of El Valle de Anton, alt. 600-1000 m., Allen 17S4, 180S, canal zone: Gatiin, Hayes 302; hills north of Frijoles, Standley 2/463, 2'/4gi^ 2/4^3, Fig. 12. Piper augustum var. coeleanuvi (61) [Vol. 37 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN darihn: Ccrro dc Garagara, Sambu Basin, alt. 500-974 m., Piificr $977 (U. S. Nat. Herb., type). This species Is to be identified because of its large, strongly pinnately nerved, densely ciliate, glandular-dotted leaves, and thick spikes. The original P. augnstum of Rudge, so far as can be determined, was entirely glabrous. Variety coclcanum has pubescent stems and more strongly pubescent leaves. Standley*s A^05. 27^6^ and 274gi arc conspicuously white-villous. No other differences have been noted, h owever. 84. Piper Stevensi Trel. in Contr. U. S. Nat. Herb. 26:27. 1927. A shrub (?) ; flowering internodes comparatively long and stout, rather densely subrctrorsely crisp-villous; leaves elliptic, apex abruptly short-acuminate, base inequilateral, one side 5—10 mm. shorter and acute, the longer side rounded to the petiole, 10-12 cm. wide X 20-23 cm. long, pinnately nerved to the upper fourth or nearly throughout, with intermediate cross-connecting nervulcs, the nerves 7-8 on each side, finely scabrous, sparsely to moderately crisp-pubescent on both sides, the nerves densely hirsute, sparingly glandular-dotted beneath, drying papery, trans- lucent, finely pellucid-dotted; petiole 20 plus 5-10 mm. long, vaginate to the middle or above, densely crisp-hairy; spikes 5 mm. thick X 7-8 cm. long; peduncle 1.5-2 cm. long, densely subrctrorsely crisp-hairy; bracts triangular-subpcltate, densely ciliate; frviit oblong, glabrous, stigmas sessile. Known only from Frijoles, Canal Zone. CANAL zone: Frijoles, Sfeieus 1247. The rather large, somewhat scabrous, pinnately nerved leaves, dense pubescence, and comparatively thick spikes characterize this species. 85. Piper colonense CDC. in Smiths. Misc. Coll. 71*^:11. 1920. Shrub, 2-5 m. tall, nodose; flowering internodes rather short and thick, puberu- lent; leaves oblong-elliptic, apex rather abruptly acuminate, base rounded, obtuse or cordulate, one side about 5 mm. shorter, 10-11.5 cm, wide X25-31 cm. long, pinnately nerved from below the upper fourth, the nerves 6 or 7 on each side, with * intermediates and cross-connecting and anastomosing nervules, the nerves on both sides puberulcnt but more conspicuously so above, drying green, paler and pale glandular-dotted beneath, translucent; petiole stout, 10-15 plus 5 mm. long, puberulent, vaginate to near the blade; spikes 3 mm. thick X 10 cm. long; peduncle 15 mm. long, rather stout, puberulent; bracts roundcd-trlangular-subpeltate, con- spicuously yellow-fringed; ovary glabrous; stigmas filiform, sessile. Central Panama. colon: around Dos Bocas, Rio Fnto valley, alt. 40-80 m., Pit tier 4^21 . The large leaves, puberulent on both sides, puberulent twigs, and vaginate petioles distinguish this species. (62) 1950J FLORA OF PANAMA (Piperaccac) 63 86. Piper tuberculatum Jacq. Collect. 2:2, /;/. 27/. 1788. Piper macronrnm H.B.K. Nov. Gen. & Sp. 1:54. 1815. Piper nutans Opiz in Prcs!, Reliq. Haenk. 1:156, pL 28, f. 2. 1830. Steffensia tuberculata Kuntli, in Linnaea 13:614. 1839, Piper scabrum Willd. in hb. p.p., fide KuntK, loc. cit. 1839. Piper obliquum Balbis in hb., fide Kunth, loc. cit. 1839. Arfanthe deciirrens Miq. Syst. Pip. 402. 1844. Artanthe tuberculata Miq. loc. cit. 497. 1844. Piper tuberculatum var. rigid o-membranaceum CDC. in DC. Prodr. 16^:266. 1869. A very nodose shrub or small tree, 2-4 or sometimes up to 6 or 7 m. tall; branches often warty, puberulent when young or less commonly glabrous; leaves subelliptic, subovatc, or mostly elliptic-oblong, 3-6 or occasionally up to 10 cm. wide X 7-14 cm. long, apex shortly and rather abruptly acute, base rounded, one side 5-10 mm, shorter at the petiole, the shorter side more elliptic and slightly wider than the longer side, ptnnatcly nerved to near the apex, tlie nerves 8—10 on each side, with intermediates, rather prominent beneath, glabrous or sparsely puberulent above, puberulent beneath, at least along the nerves, or rarely glabrate, drying membranous, somewhat translucent, obscurely minutely pellucid-dotted; petiole about 5-10 plus 5-10 mm. long, vaginate and narrov/ly winged throughout with the wings acutely protracted at the base of the blade; spikes 3-5 mm. thick X 5-10 or 12 cm, long; peduncle mostly 10-15 mm. long, rarely up to 2 or 3 cm.; bracts triangular-subpeltate, marginally fringed; fruit somewhat tctragonous, glabrous; stigmas 3, sessile. West Indies, Central America, and throughout northern South America. Panama: without location, Hayes 8o2; Bro, Heriberto 26^. canal zone: Balboa, SfafuIIey 3^40g; near Bojii Station, Panama R. R., Hayes lOl/'; Empire, Stevens II4U Miller l86j; near Gorgas Memorial Laboratory, White 84; western slope of Ancon Hill, vicinity of Balboa, alt. 20-7 5 m., Seibert IIQ; Las Cruces Trail, government forest, alt. 75 m., Hunter & Dodge 759. Panama: along the Rio Chilibrc, Piper 5681; vicinity of Bella Vista, Piper 5375; along Rio Juan Diaz, above Juan Diaz, alt. 30 m., Allen Q43 ; Taboga Island, Miller 2041 ; alt. 0-350 m., Allen 1287; alt. 0-186 m., Woodson, Allen & Seibert 1508; Perlas Archipelago, Saboga Island, Miller IQ50, 86a. Piper tuberculatum var, minus CDC. in DC. Prodr. 16^:266. 1869. Leaves smaller, mostly 3-7 or sometimes 8 cm. long. Occurs with the species. cocle: Penonome and vicinity, alt. 15-300 m., Willia?ns /J, ^^55- canal zone: near Gorgas Memorial Laboratory, White 84; Empire, Miller l86j; Sabanas north of Panama City, Bro. Paul J77. 87. Piper arcoreum Aubl. Hist. PL Guian. Fr. 1:23. 1775. Piper macrophyllum Sw. Prodr. Veg. Ind. Occ. 15. 1788. Piper geniculatum loc. cit. 178 8. Piper verrucosum loc. cit. 1788. Piper nitidurn loc. cit. 178 8. Piper nodidosum Link in Jahrb. d. Gewachsk. 1^:62. 18^0, name only; Link, Enum. Hort. Bcrol. 1:36. 1821. f 1839. (63) [Vol. 2^7 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN Piper xylopioliles Kunth, loc. cit. 18 39. nua? ntfl.la Kunth. loc. clt. 611. 1839, ■# 1839. Sfcffensia gcfuculafa Kunth, loc. cit. 612. Piper secufiJnm Poepp. in lib. fide Kunth, loc. cit. 1839. fi\ifi? ijprrnrnRa Knnrh. loc. cit. 613. 18 39. f 1839. Steffensia Ltischiiathiana Kunth, loc. cit. 616. Piper Luschnatljiuuum Kunth, loc. cit. 617. 1839. Artanfhc Lcsscrfiana Miq. Syst. Pip. 405. 1844. ArtiOitbe xcsfophylla Miq. loc. cit. 491. 1844. Arfanfhc xylopioidcs Miq. loc. cit. 492. 1844. Artantbc gcnicidata Miq. loc. cit. 493. 1844. Artantbe Lu^chtuithiana Miq. loc. cit. 494. 1844. Artantbc verrucosa Griscb, Fl. Br. W. Ind. 171. 1864. Artantbc viacropbylla Griscb. loc. cit. 1864. Piper arborcum f. gcnicidata Fawc. & Rend., Fl. Jam. 3:23. 1914. Piper laevibracfeum Trcl. in Contr. U. S. Nat. Herb. 26:26. 1927. Piper subnuilispicnm Trel. loc. cit. 1927. Piper iubcrcnlatum var. AUeni Trcl. in Ann. Mo. Bot. Gard. 27:298. 1940. A glabrous, somewhat nodose shrub, or small tree, mostly 2-7 m. tall; flowering internodes rather sliort but slender, more or less verrucose- warty; leaves elliptic- ovate, oblong, or lance-elliptic, mostly 6-11 cm. wide X 10-2 5 cm. long, apex gradually acute to short-acuminate, rounded at base, or narrow leaves acute, one side 5-10 mm. sborter at the petiole, or occasional leaves equilateral, pinnatcly nerved throughout, the nerves 8-10 on each side with some intermediates; petiole 5-10 plus 5-10 mm. long, vaginate- winged to the blade witb the wings protracted beyond the lower blade margin; spikes rather slender and elongated, 2-4 mm. thick and up to 15 cm. long but mostly 5-10 cm,; peduncle 5-10 mm. long and mod- erately stout; bracts triangular-subpcltate, marginally ciliolate; fruit elongated witb the rachis, glabrous; stigmas 3, sessile. West Indies, northern South America, and Panama. CHiRiQui: Ccrro Vaca, alt. 900-1136 m., Pittier 5324, cocle: north rim of El Vallc dc Anton, alt. 600-1000 m., Allen 1640, canal zone: Barro Colorado Island, Gatun Lake, StanJley 410SQ; Maxon, Harvey ^ Valentine 6S13; vicinity of Fort Sherman, Standley JIIlS- colon: Loma de la Gloria, near Fato (Nomhre de Dios), alt. 10-104 m., Pittier 4241. 87a. Piper aruoreum van hirtellum Yunckcr, var. nov. A speciei forma typica diffcrt innovationibus, petiolis, pcdunculis, venis subtus plus minusve puberulcntis. Similar to the species except that the young branches, petioles, peduncles, and nerves, especially on the lower surface, are more or less minutely pubcrulent or hirtellous with erect hairs. — British Guiana: northwest district: Waini River, April 3-18, 1923, de la Cruz 362J (U. S. Nat. Herb., type). Northern South America and Panama. CHiRiQUi: vicinity of San Felix, alt. 0-120 m., Pit tier 513I. canal zone: hills north of Frijoles, Standley 2/3IQ. Panama: Pcrlas Archipelago, San Jose Island, Jobnston 424, 594' (64) 1950] FLORA OF PANAMA (Vipevaccae) 65 87b. Piper arboreum van falcifolium (Trel) Yuncker, comb. nov. Piper falcifolium Trel. in Contr. U. S. Nat. Herb. 26:25. 1927. Leaves oblong-lanceolate, 3-5 cm. wide X 10-20 cm. long, apex acuminate, base inequilateral, rounded, obtuse or acute. West Indies, northern South America, and Panama. CHiRiQui: Boquete District, alt. 1140 m., Davidson 6l8; vicinity of El Boquete, alt. 1000-1300 m., Maxan 4959, 5381; Pittier 2958; Bro. Maurice 738; vicinity of San Felix, alt. 0-120 m., Dec. 1911, PUlitr 3137 ; Sabana de la Tortuga, between El Boquete and Caldera, alt. 3 00-700 m., Piifier 3327. Panama: Pcrlas Archipelago, San Jose Island, Johrntou 636, T373, This species appears to be extremely variable in the size of the leaves and to some extent also the shape. Aside from these variations and the presence of short, stiff hairs in one variety, no characters have been discovered by means of which the specific distinctions suggested by the rather long list of synonyms may be maintained. They are believed to be merely varying members of a rather poly- morphic species. 88. Piper subrepens Trel. in Ann. Mo. Bot. Gard. 24:187. 1937. A nodose, glabrous shrub; flowering internodes short and moderately slender, drying yellowish-glossy; leaves lance-ovate, apex acuminate, base rounded, obtuse, equilateral, or one side very slightly shorter, 2-3.5 cm, wide X 6.5-9 cm. long, pinnately nerved to the upper fourth or nearly throughout, the nerves 3-4 on each side, strongly curving toward the tip submarginally, with intermediates and cross- connecting and anastomosing nervules, the nerves beneath and the midrib above salient, drying firm, glossy, not glandular-dotted, subopaque, narrowly revolute, light green; petiole up to 1 cm. long, glossy, vaginate-winged to the middle or below; spikes 3 mm. thick X 2 cm. long; peduncle slender, up to 1 cm. long; bracts triangular- or rounded-subpeltate, submarginally fringed; fruit obpyramidal- trigonous with dome-shaped apex, glabrous, stigmas filiform, sessile. Known only from the following locality. cocle: very near the crest of Cerro Valle Chiquito, alt, 700-800 m., Seiberf 304. The short spikes and moderately small leaves with rounded, subcquilateral bases distinguish this species. 89. Piper cordulatum CDC In Jour. Bot. 4:217. 18 66. •J Artanthe trtjicrvia Miq. in Seem. Bat. Voy. Herald, 200, pi, 41. 18 54. Piper trinerve CDC in DC Pradr. 16^:292. 1869, not Vahl. 1804. ?Piper cordulatum van granulatum Trel. in Contr, U. S. Nat. Herb. 26:26. 1927. Piper sau-lorenzauum Trel. loc. cit. 29. 1927. A glabrous shrub or small tree, 1-3 m. tall; flowering internodes rather slender and short, obscurely finely striate; leaves lanceolate or lance-oblong, apex sharp- acumin:uc, base rounded or commonly somewhat narrowed downward, one side (65) I Vol. 37 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN usually 1-2 mn^. shorter, the longer side corduhue, 3-6 cm. wide X 14-21 cm. long, pinnatcly nerved throughout, the nerves 7-12 on each side, strongly sub- marginally loop-connected, the longer nerves prolonged, salient, with cross-con- necting and anastomosing ncrvulcs, drying rather coriaceous, narrowly revolute, somewhat glossy, usually bronzy-brown beneath, often with fine mottled markings, opaque; petiole 10-20 plus 1-2 mm. long, prominently vaginate-winged to the blade; spikes 3 4 mm. thick X 5-7 cm. long, apiculate; peduncle about 5 mm. long; bracts concave or Innately subpcltate, papillate or glabrous; fruit depressed- obovoid, slightly elongated with the rachis, glabrous, glandular-dotted; stigmas 3, sessile. Pannma. CANAL ZONT.: Gati'in Station, Ihiyes s. /?., 66, fjOJ, J^C); between Gori;ona and Gatun, alt. 10-50 m., Pit Her 2j6y; A^ua Clara Reservoir, Stevens jSg; around Las Graces, alt. 20-100 m., Piftier 2624; lulls north of Frijolcs, Standley 26450, 2^578; Rio Paraiso, above East Paraiso, Standley 2gr)0j; Barro Colorado Island, Wet more ^ Ahbe 2^; Standley 3^355y 4^^471 r*^3d to Madden Dam and Las Cruccs Trail, ah. 75 m., Hunter ^ Allen 460, colon: Chagres, Vendler 26"/. Panama: thickets and forests near Arraijan, alt. 15 m., Woodson, Allen & Seibert 782; Indlo, Madden Lake, Miller 20jO, 2065. The somewhat bronze-colored leaves with cordulate base, loop-connected nerves, and strongly va;nnate-winged petioles distinguish this species. 90. Piper acuhssimum Trcl. in Contr. U. S. Nat. Lierb. 26:25. 1927. Piper Incaeufnun Kunth var. magnifolinm CDC, in DC. Prodr. 16^:322. 1869. Viper sedncentifoliiim Trel. in Ann. Mo. Bot. Gard. 17:297. 1940. A shrub or small tree, 2 3 m. tall, more or less nodose, twigs straight or some- what /ig-zag, glabrous; flowering Internodes short and rather slender; leaves lance- elliptic, 2.5 6 cm. wide X 10-18 cm. long, apex acuminate, base subequilaterally acute, plnnately nerved throughout, the nerves 5-8 or 10 on each side, loop-con- nected upward, glabrous or rather sparsely pubescent along the nerves beneath, drying thin, translucent, glandular-dotted; petiole from 5 mm. long above to 10-15 mm. long bclov/; spikes 3-4 mm. thick X 5-9 cm. long; peduncle filiform, 1-2 cm. long; bracts triangular- subpcltate, sparsely marginally fringed; fruit trigonous, more or less gr.mular-papillate, scarcely longer than thick, truncate-concave at apex; stigmas 3, short, sessile. Panama and northern South America. CANAL zone: hills north of Frijolcs, Standley 27504, 27599. colon: Cliat;res, ¥end- ler 266. PANAMA: thickets and forests near Arraijan, alt. 15 m., Woodson, Allen & Seibert 781; Indio, Madden Lake, Mdler 2o6o, 2079. darien: Bawe 887. 90a. PiPLR ACUTLSSIMUM var. trichopus (Trel.) Yuncker, comb, nov. Viper trichopus Trcl. in Contr. U. S. Nat. Herb. 26:25. 1927. Youn^ grow th sparsely to densel)' pubescent; nerves beneath appresscd- pubescent, Panama. (66) 1950] FLORA OF PANAMA (Pipcraccae) 67 Panama: without location, 18 59-60, Hayes jgj (Herb. New York Bot. Gard., type). The glabrous twigs, and pinnately nerved, lanceolate, acuminate, and glandular- dotted leaves arc distinctive features of this species. It differs from P. hicacaniim of Brazil, according to Kunth's description of that species, because of its larger leaves, longer peduncles, and lack of styles. Variety trichopjts appears to differ only in the pubescence. 91. Piper perlasense Yuncker, sp. nov, Frutex glabcr 2-3 m. altus, caulibus nodosis; intcrnodiis floriferis brevibus gracilibusque; foliis lanceolatis vel sat oblongo-lanceolatis, apice sensim subfalcatim longe acuminatis, basi oblique subinequilateralibus latere altero 1-2 mm. brcviore, latere longiore rotundo obtuso, totis vel e quarto supero tantum venosis, vcnis utrinque 6-8; petiolo deorsum vaginato; spicis 2—3 mm. crassis, 5—6 cm. longis; pedunculo ca. 5 mm. longo; bracteis triangularibus vel rotundato-pcltatis marglne fimbriatis; drupa parva glabrata obpyramidali-trigona, stigmatibus parvis sessilibus. A glabrous, nodose shrub 2-3 m. tall; flowering internodes short and slender; leaves lanceolate or somewhat oblong-lanceolate, apex rather gradually subfalcately long-acuminate, base obliquely slightly inequilateral with one side 1-2 mm. shorter and acute, the longer side rounded and obtuse, 2-4.5 cm. wide X 9-15 cm. long, pinnately nerved to the upper one-fourth or throughout, the nerves 6-8 on each side, with intermediates, divergent at the midrib then strongly curved upward and submarginally loop-connected, drying papery, light green, somewhat glossy, trans- ■m lucent; petiole scarcely 5 plus 1-2 mm. or occasionally 1 cm. long, vaginate toward the base; spikes 2-3 mm. thick X 5—6 cm. long when mature; peduncle about 5 mm. long; bracts triangular- or rounded-subpeltate, fringed; fruit small, obpyra- midal-trlgonous, glabrous, stigmas small, sessile. Panama. CANAL zone: Barro Colorado Island, Steveits J, panama: Perlas Archipelago, San Jose Island, Johnston IJ4, 262, JIJ\ Jan. 22, 1946, Johnston 1 21 5 (Herb. Univ. 111., type), 1327. This species appears to be rather closely related to P. aeqnale of South America from which it differs, however, because of its mostly smaller, lanceolate leaves which are more definitely pinnately nerved, and rounded on one side at the base. 92. Piper via-chicoense Yuncker, sp. nov. Frutex (?); intcrnodiis floriferis gracilibus brevibusque, conferte villosis, pilis usque ad 1 mm. longis plus minusve heteromorphis, brevioribus longioribusque intermlxtis; foliis elliptlco-vel lanceolato-oblongls, apice acuminatis, basi obtusis vel pro more cordulatis, totis pinnatim venosis, venis utrinque 5 vel ultra, supra glabratis vel ad basem dissite pubcscentibus, subtus crispe pubescentibus, venis conferte indutis, pilis dissitis longioribus; petiolo vix 5 mm. longo, conferte hirsuto, basi vaginato; spicis immaturis, 2 mm, crassis, 5 cm. longis, mucronatis; pedunculo 5 mm. longo, pubescente; bracteis triangulari-subpcltatis, margine fimbriatis; ovario I ovoideo (?); stigmatibus sessilibus; drupa ignota. (67) 68 I Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Shrub (?); flowering internodes sliort and slender, densely hairy, hairs of various lengths, the longest up to 1 mm. long; leaves elliptic- or lance-oblong, apex acuminate, base with one side slightly shorter at the petiole, both sides obtuse or mostly cordulate, 3-4.5 cm. wide X 10-13 cm. long, pinnately nerved throughout, the nerves 5 or more on each side, more promi- nent downward, submarginally loop-connected, with cross-connecting-anastomosing nervules, glabrous above or sparsely pubescent at the very base, crisp-pubescent beneath, the nerves densely so and \v ith intersperse d 1 ong liai rs, Fig. 13. Pi per via-chicoensc drying firm, subopaque; petiole scant 5 mm. long, densely hairy, vaginate at the base; spikes as yet young, 2 mm. thick X 5 cm. long, mucronatc; peduncle 5 mm. long, pubescent; bracts trlangular-subpeltate, marginalK fringed; ovary ovoid (?); stigmas sessile; fruit not developed. Known only from the type locality. CANAL 70NE and adjacent Panama: forests along telephone cable trail between splice S16 and S49, Rio Indio, Chico Trail, Jan. 12, 1935, Stcycnnark & Allen 17431 (Herb. Missouri Bot. Gard., type). 93. Piper grande Vahl, Eclog. Amer. 2:3, /?/. //. 1798. Pspcr ^r^r audi folium H.B.K. Nov. Gen. & Sp. 1:46. 1815. Piper rrpariuff? H.B.K. loc. cit. 48. 1815. Scbilleria riparia Kunth, in Linnaca 13:707. 1839. Scbilleria ^ramVtjoUa Kunth, loc. cit. 709. 1839. ?Piper granatensc D. Dictr. Syn. Pi. 1:117. 1839. Artantbe grauJifolia Miq. Syst. Pip. 408. 1844. Artauthe riparia Mlq. loc. cit. 409. 1844. Piper pseudo-i'ariabile Trel. in Contr. U.S. Nat. Herb. 26:23. 1927. Piper pseiido-variabile var. pacbypus Trel. loc. cit. 24. 1927. Piper pseudo-t^ariabile var. coUinm Trel. loc. cit. 1927. Piper jnarnwreiiuj Trel. In Ann. Mo. Bot. Gard. 24:186. 1957. Piper spissifterviuni Trel. fide Bro, Tomas Alberto in Colcgio San Jose, McdcUin, Colombia. Publ. 53-70, p. 18. 1942, name only. A glabrous shrub or small tree, up to 5 m. tall; flowering internodes slender and moderately elongated; leaves ovate with the midrib commonly somewhat nearer one side, 8-23 cm. wide X 16-30 cm. long, or some leaves may be slightly smaller, apex abruptly acuminate, base nearly equilatcrally rounded, the upper acute or subacute, the larger lower leaves obtuse to subcordulate, or occasionally cordate, pinnately nerved to the upper one-third or essentially throughout, the nerves 6-8 or 10 on each side, with shorter intermediates, commonly yellow and prominent (68) 1950] FLORA OF PANAMA (Pipcraccae) 69 beneath, the lowermost 2 or 3 pairs more or less approximated at the base, drying chartaceous, somewhat glossy, translucent; petiole 10-3 mm. long, the longer vaginate to or beyond the middle; spikes 4-5 mm. thick and up to 15 cm. long; peduncle rather stout, 5-10 mm. long; bracts rounded-subpcltate, umbonate, smooth above, villous beneath; fruit small, obpyramidal-trigonous, apex somewhat dome-shaped; stigmas 3, sessile. Colombia and Panama. CHiRiQui: vicinity of San Felix, alt. 0-120 m., Pit tier jlSp, 5262, veraguas: IsLi de Coiba, Mendcz 121, cocle: vicinity of El Vallc, alt. 600-1000 m., Allen 180J; vicinity of Ola, alt. 100-3 50 m., Fit tier 5075; vicinity of El Valle dc Anton, alt. 600 m., Allen 2001, CANAL zone: along Cano Qucbrada, Viiiicr 6662; Barro Colorado Island, Miller 2074; Kenoyer J06; Chrysler 484O; CbarJon & Nolla Iq6; Standley 31308, 40S7J, 40930, 4^^54 — a narrow leaved form, 41076; Rio Paraiso, above East Paraiso, Standley 2g8g2; Las Cascadas Plantation, near Summit, Standley 23700, 29318, 29638; hills west of the canal, near Gatun, Standley 27191; hills north of Frijolcs, Standley 27333; between Frijolcs and Monte Lirio, alt. 30 m., Killip 121 43; Las Cascadas Plantation, near Summit, Standley ^3733; Battery 46 Hill, Madden Dam road, Miller 1772, 1773; vicinity of Fort Sherman, Standley 31022; Obispo, Standley 31702; Ancon Hill, alt. 180 m., Williams 19; Standley 23179. colon: Loma dc la Gloria, near Fato (Nombrc de Dios), alt. 10-104 m., Pittier 3^43'y along trail to triangulation station on top of Tumba Vieja, alt. 90-200 m., Dodge, Steyermark & Allen 16924, Panama: Pacora River, Killip 314I; Rio Tapia, Standley 41188; Indio, Madden Lake, Miller 2o6l, 2062, 2063, 2o66; Rio Tapia, Standley 26130 — the leaves are somewhat smaller and this may represent a large-leaved form of P. aequale; Juan Diaz, Standley 30610, 30377; Perlas Archipelago, San Jose Island, Erlanson 334; vicinity of Arenosa, lower Rio Trinidad, alt. 26-50 m., Seibert 622, The large, ovate, glabrous leaves, drying somewhat glossy, with prominent yellow nerves, and smooth, umbonate floral bracts are distinctive features of this species. The leaves show considerable variation in size often on the same specimen. The smaller-leaved specimens approach P. acqnale with -which this species is very closely related. These two species may, in fact, represent extremes in leaf size of a single strongly variable species. VL KEY TO SPECIF.S WITH PELTATE LEAVES a. Petiole 7 cm. or less long, vaginate to the middle or above; mostly with 3 nerves on each side of the midrib above the base. h. Leaves strongly bulla tc- rugose; young growth more or less rusty- tomcntose 94. P. Hartwegianum bb. Leaves not bullate-rugose, not hairy 95. P. Maxonii aa. Petiole 12 cm. long, vaginate at base only; with 2 nerves on each side of the midrib above the base 96. P. vhraguense 94. Piper LLartwegianum (Benth.) CDC. in DC. Prodr. 16^:369. 1869. Artanthe Hartivegiana Benth. Pi. Hartweg. 2 54. 1839. Piper insignifolium CDC. in Ann. Conserv. & Jard. Bot. Geneve 21:242. 1920. Piper Hayesii Trel. in Contr. U. S. Nat. Herb. 26:39. 1927. A shrub or small tree, 3-5 m. tall, rusty-villous except for the upper leaf surface which is glabrescent or pubescent along the nerves only; tv^igs stout; leaves ovate or cUiptic-ovatc, 16-25 cm. Avidc X 28-40 cm. long, apex acuminate, base incquilaterally rounded, obliquely retuse or shallowly cordate, peltate 3-6 cm. (69) I Vol. .\7 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN from tlic margin, with 7-9 nerves radiating from the petiole, and tlic midrib ^vilIl 3 or 4 branches on each side from the lower half, strongly rugose-bullatc, drying firm and subopaque; petiole 5-7 cm. long, vaginatc-wingcd to near the base of the blade; spikes up to 10 mm. thick and 35 cm. long when mature; pedimcle 3-4 cm. long; bracts clavatc, thickened and bristly at the top, pedicels bearded; fruit oblong-ovate, angular, glabrate; stigmas filiform, sessile. Suuilnvestcrn Colombia and Panama. Panama: without locality, in 1859-60, lluycs 76 j. No differences have been found between the above Panamanian specimen and those seen from South America which would justify maintaining P. JJaycsii as a separate species. 95. Pipr.R Maxonii CDC. in Smith< 15-25 or even 3 cm, long, apex acuminate, often abruptly so, base cuneate-decurrent, pinnately nerved, midrib prominent, lateral nerves inconspicuous; petiole 1—4 cm. long, margined by the dccurrent blade; spikes terminal, 5 or 6 mm. thick and up to 2 5 cm. or more long, densely flowered; peduncle 15—20 mm. long, commonly with a small leaf or bract near the middle; floral bracts round-peltate; fruit about 1 mm. long, ellipsoidal, yellow at the base, abruptly and truncately short-pointed; stigma apical. Northern South America and Central America, CHIRIQUI: vicinity of Casita Alta, Volcdn de Chiriqui, alt. 1500-2000 m., WooJsof?^ Allen ^ Seibert gj2; trail from Ccrro Punta to headwaters of Rio Caldcra, alt. 2250-2500 m., Allen 1452; Boqucte District, Bajo Chorro, alt. 2250 m., Davidson 2o6, The large, thick, pinnately nerved leaves, long, thick spikes, and fruit shape distinguish this species. 2, Peperomia acuminata Ruiz & Pav. Fl. Peruv. & Chil. 1:32, pL 5/, /. a, 1798, not P. ac7i7ninata (L.) Dahlst. 1900, Piper acjiminafiim Pers. Syn. 1:3 5. 180 5, not P. acuminatum L. 175 3. Piper nemorosum Vahl, Enum. PL 1:341. 1805. Peperomia pyrifolia H.B.K. Nov. Gen. & Sp. 1:61. 1815. Peperomia basellaefolia H.B.K. loc. cit. 66. 1815. (77) [Vol. ?>7 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN /#•**. Viper basellaefolium Poir. in Lam. Encycl. Mcth. 4:SuppI. 468. 1816. Piper murhium Roem. & Schult. Syst. Veg. 1:325. 1817. Viper pyrifoUum D. Dictr. Encycl. 122. 1831, not P. pyrifolium Vahl. 1805. Vcperomia nemorosa CDC. in DC. Prodr. 16^:415. 18 69. ?Peper(>mra dichroophylla Sodiro, Contr. Conoc. FI, Ecuator. Monogr. 1. cd. 2. 157. [1901]. Vepcromia Kniziana CDC. in Candollea 1:301. 1923. Veperomia Ekmaui Trel. in Fedde. Rep. Sp. Nov. 23:31. 1926. Vepcromia acuminatifolia Trel. in Contr. U. S. Nat. Herb. 26:48. 1927. Vepcromia oblanceaelimba Trel. in Badlllo in Cat. Fl. Venez. 1:245. 1945, name only. An assurgcnt, simple or sparingly branched, glabrous, succulent herb; stem 4-5 or sometimes 10 mm, thick when dry, ascending from the decumbent, rooting base to 60 cm. or more in large specimens; internodcs 1—3 or 4 cm. long, leaf scars large, nodes commonly more or less tumid; leaves alternate, elliptic, lanceolate or oblance- olate, 2-4 cm. wide X 6-13 cm. long, apex attenuate and sharply more or less arcuately acuminate, base cuneate, dccurrcnt, pinnately nerved, drying coriaceous and subopaque, commonly minutely dark-dotted and glandular beneath, minutely ciliolate at the apex; petiole 5-10 mm. long, or occasionally 15-20 mm. on lower leaves, more or less winged by the decurrent blade; spikes solitary, terminal, closely flowered, 3-4 mm. thick X 10-13 cm. long; peduncle 3-5 cm. long, sometimes with a small leaf or bract; floral bracts round-peltate; fruit about 1 mm. long, ellipsoidal, abruptly and truncately scutellate, sometimes yellowish at the base, stigma apical. West Indies, Panama, and northern South America, Panama: without locality, Hayes 817. 3. Peperomia ciliolibractea CDC. in Candollea 1:360, 383. 1923. A moderate-sized, assurgent herb; stem 3-4 mm. thick, ascending to 15 cm. or more from the decumbent, rooting base, velvety pubescent; internodes scarcely 1 cm. long, nodes somewhat tumid, leaf-scars large; leaves alternate, oblong-ovate or subobovate, apex shortly sharp-acuminate, base rounded, cordulate to acute, mostly 2-5 cm. wide X 5-8 cm. long, occasionally up to 7 X 14 cm., pubcrulent on both sides, rather densely so along the nerves beneath, ciliolate, 9- to 11- plinerved, the uppermost lateral nerves branching off the midrib near the middle, drying membranous, translucent, reddish-glandular-dotted beneath; petiole 2-5 or sometimes up to 8 cm. long, velvety; spikes 2 mm. thick X 10-14 cm. long, closely flowered; peduncle up to 5 cm. long, velvety; bracts round-peltate, rcddish- glandular-dotted, marginally more or less ciliate; fruit about 1 mm. long, sub- globose, apex pointed, slightly oblique, stigma essentially apical, basal part of the fruit yellowish, upper half reddish with reddish-glandular dots. Panama. CANAL ZONE: Las Cascadas Plantation, near Summit, StatiJley 2^Y^I ; Barro Colorado, Stevens 6j7; Wheeler Trail, Sbattuck 214, Panama: Charare River near Chepo, alt. 50-200 m., Vittier 4/12, The pllnerved leaves, velvety stems, leaves, etc, short internodes, and shape of fruit arc distinctive of this species. (7^) 195 0] FLORA OF PANAMA (Pipcraceac) 79 4. Peperomia brevipeduncula Trel. in Contr. U. S. Nat. Herb. 26:43. 1927, not P, Heydei var. brevipedtcncnla CDC. A short, rhizomatous, leafy, glabrous herb; stem ascending 6-8 cm, from a fibrous-rooted, somewhat contorted-nodulose (when dry) rhizome about 4 mm. thick, internodes up to 10 mm. long; leaves alternate, round, round-ovate, or elHptic-ovate, apex rounded or slightly pointed, base cordulate or shallowly cordate, 4.5-6 cm. wide and occasionally up to 9.5 cm. long, palmately 5-nerved, or large leaves 7-ncrved with the outermost laterals slender and indistinct, nerves sparingly branched upward, drying dull, membranous, translucent, pellucid-dotted; petiole 2-6 cm. long, clasping; spikes opposite the upper leaves, and axillar}% rather stout, 2-3 mm. thick X S-15 cm. long, rather loosely flowered; peduncle 5-15 mm. long; bracts round-peltate, glandular-dotted; fruit globose, about 1 mm. long, apex mammiform, stigma apical. Southern Panama, darien: Boca de Panarando, on Sambii River, alt. 20 m., Pitfier SjpO. When Trclease described this species he considered it as being the same as P. Heydei var. brevipednjicula, which is now believed to be P. pseudo-dependens. It differs from that species because of its fewer-nerved leaves with cordulate rather than cordate base. From P. Heydei it differs in the shape of the fewer-nerved leaves, shorter peduncles, etc. 5. Peperomia pseudo-dependens CDC. in Jour. Bot. 4:137- 1866. Peperomia Heydei CDC. var. brevipediinenJa CDC in Ann. Conserv. & Jard. Bot. Geneve 21:247. 1920. Peperomia brevipeduneida (CDC) Trel. var. major Trel. in Ann. Mo. Bot. Card. 27:^01. 1940. A short-stemmed, glabrous, succulent, few-leaved herb, perennial from a small corm; leaves alternate, round-ovate, up to 11 cm. in diameter, apex obtuse or sub- acute, base deeply cordate, palmately 9- to 11 -nerved, the nerves forked upward, drying very thin and translucent; petiole up to 10 cm. long, mostly about 5-7 cm.; spikes terminal and axillary, rather loosely flowered, 2-3 mm. thick and up to 14 cm. long; peduncle 7-10 or even 20 mm. long; bracts round-peltate, red-glandular- dotted; fruit about 1 mm. long, globose-ovoid, sulcate ventrally toward the base, apex mammiform, stigma apical. Venezuela, northern Colombia, and Panama. PANAMA: Rio Las Lajas, alt. 20 m., Allen 20J/. 6. Peperoml^ pellucida (L.) H.B.K. Nov. Gen. & Sp. 1:64, 1815, Piper pellncidum L. Sp. Pi. 30. 1753. Piper eoncinnum Haw. Rev. PI, Succ. 198. 1821. Peperomia eoncinna A. Dietr. Sp. Pi. 1:164. 1831. Micropiper pellucidum Miq. Comm. Phyt. 54. 1840. Peperomia pelhcida var. pygmaea Willd. Herb, ex Miq. Syst. Pip. 81. 1843. Peperomia pellucida var, minor Miq, in Mart. Fl. Bras. 4^:10. 18 52. (79) [Vol. 37 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN A rather delicate, erect, glabrous, pellucid, branched herb; stem up to 5 mm. thick near the base and 50 cm. tall; leaves alternate, round-deltoid-ovatc, mostly 15-2 5 mm. wide, apex somewhat attenuate, acute, base subtruncatc-cordatc, palm- atcly 5-nerved, drying very thin, semi-transparent; petiole mostly 1-2 cm. long, clasplng-dccurrcnt; spikes terminal, axillary, or leaf -opposed, slender, 2-5 cm. long, very loosely flowered; peduncle about 5 mm. long; bracts round-peltate; fruit about 0.8 mm. long, ellipsoidal, longitudinally more or less sulcately striate; apex pointed, stigma apical. A common weedy species occurring throughout the West Indies and tropical North and South America. It is also found in the Old World tropics and Pacific Islands. Panama: without locality, Dro. Heribcrto y8, bocas dll toro: vicinity of Nievecita, alt, 0-50 m., WooJson, Allen d Seibcrt lS66; vicinity of Chlriqui Lagoon, von WcJcl J20; lower Changuinola River, Stork 7. canal zone: Balboa, SianJlcy 2j8j/, 2/ljQ; Foster s. n.; Gatun, Courll 2Q2; Summit, Stevens 32j; Sfandley 26goy, Ancon, alt. 2080 m., Pit tier 3962; Ancon Hill, Stevens 794, colon: Chai;rcs, Femller 302; vicinity of Porto Bello, alt. 5-10 m., Pit tier 2469, Panama: Cliepo, alt. 60 m., Pit tier 4450; Las Sabnnas, StunJIey 2jgjO; Taboga Island, StantUey 2/063, 2/SSl; Pcrlas Archipelaj^o, San Jose Island, Johnston 126, lyS, san blas: vicinity of Puerto Obaldia, alt. 0-50 m., P/7/;Vr 4370. This species, together with the East Indian P. exignay constituted the subgenus Ogmocarimdtum of Dahlstedt. 7. Peperomia cordulatii ormis Trel. in Bot. Gaz. 73:143. 1922, An ascending, glabrous, rather large herb; stem 3-4 mm. thick when dry, ascending to 50 cm. or more, freely and widely branching, internodes up to 10 cm. long, rooting from the lower nodes; leaves alternate^ round-ovate, apex bluntly short-acute, not at all bristly, base rounded, cordate, scarcely peltate as originally described, 3-5 cm. wide X 4-6 cm, long, smaller upward in young growth, palm- ately 5- to 7-nerved, the midrib scarcely branched upward, drying membranous, translucent, glandular-dotted; petiole up to 5 cm. long on larger, lower leaves, reduced upward with smaller uppermost leaf petioles 5 mm. long, channelled above, clasping-dccurrent; spikes leaf -opposed, 1 mm. thick X 4-5 cm. long, rather loosely flowered; peduncle 5-10 mm. long; bracts round-peltate, glandular-dotted; fruit globose-ovoid, about 0.5 mm. long, apex scarcely oblique, stigma essentially apical. Canal Zone. CANAL zone: Mamei Hill, alt. 20-90 m., on rocks, Pittier jSo6; vicinity of Madden Dam, near Rio Chat;rcs, alt. 50-75 m., on banks of brook and in water, Seihevt 353^ Alajuela, rocks along Chagres River, Pittier 34^7^ Trelease described the leaves as "obscurely subpeltate." The margin, however, does not meet over the petiole in the type specimen. It differs from P. corJut.i because of its smaller, non-peltate leaves, and character of fruit. It rather closely resembles P. pdliicida in many respects but differs because of its usually larger leaves in which the midrib Is scarcely branched upward. (^0) 19 501 FLORA OF PANAMA (Pipcraceac) 81 8, Peperomia insueta Trel. in Ann. Mo. Bot. Gard. 27:303. 1940. A rather small, terrestrial, acaulcscent herb; leaves arising from a short, thick- ened, rhizome-like base, elliptic, apex obtuse, base obtuse, cordulate, or subacute, 2-4 cm. wide X 4-7.5 cm. long, moderately to sparsely crisp-pubescent above, moderately crisp-pubescent beneath, more densely so along the nerves, scarcely ciliate, palmately 5-ncrvcd, or large leaves with an additional pair of slender nerves at the base, the innermost pair of nerves appearing to coalesce with the midrib within the lowermost 5 mm., the midrib and lateral nerves branched upward, drying darker above, paler beneath, membranous, translucent, glandular-dotted beneath; petiole 3-8 cm. long, crisp-pubescent; spikes 1.5 mm. thick X 6-12 cm. long, rather closely flowered; peduncle about equilong, crisp-pubescent; rachis pubescent at the base and with a few, scattered hairs above; bracts round-peltate, glandular- dotted; fruit about 0.6 mm. long, globose, stigma apical or at most only slightly subapical. Pana ma. cocLE: Vicinity of El Valle, alt. 600-1000 m., Allen 1220. canal zone: Qucbrada Ancha, alt. 70 m., Dodge & Stcyermark l/06l; Steyermark 6 Allen IJo6la, The acaulcscent plants, long-pctiolate, elliptic leaves with pubescent blades and petioles, long spikes on long peduncles, and somewhat pubescent rachiscs are distinctive characters of this species. 9. Peperomia umbrigaudens Yuncker, sp. nov. Herba succulcnta rhizomatosa; foliis alternls, rotundato-ovatis usque ad 11 cm. latis, apice obtusis vel subacutis, basi profunde cordatis, sinu aperto, palmatim 11- nervils supra sat dissite villosis subtus ad venas plus conferte villosis; petiolo usque ad 15 cm. longo, sat gracih, proximo laminae villoso, deorsum glabrato; spicis solitariis; pedunculo ca. 6 cm. longo, dissite villoso; ovario subturbinato, apice sub- mammiformi, stigmatc apicaU interdum bifido. A rhizomatous, succulent, shade-loving herb; leaves alternate, few, clustered at the end of a fleshy, fibrous-rooted rhizome 5-10 mm. thick when dry, round- ovate, apex obtuse or obscurely acutish, base deeply cordate with open sinus, up to 11 cm. in diameter, palmately 11 -nerved, the nerves strongly forked upward, rather sparsely villous above, more densely so along the nerves beneath, drying thin, trans- lucent, finely pellucid-dottcd, paler and prominently glandular-dotted beneath; petiole up to 15 cm. long, rather slender, villous near the blade, glabresccnt down- ward; spikes solitary, terminal (?) or axillary (?), 2 mm. thick X 6 cm. long, closely flowered; peduncle equilong, thinly villous; bracts round-peltate; ovary subturbinate, apex submammiform, stigma apical, sometimes bifid. Known only from the type locality. PANAMA: Cerro Trinidad, on moist boulders in deep shade, alt. 800-1000 m., Oct. 20, 1946, Allen 3766 (Herb. Missouri Bot. Gard., type). This species resembles P. psciido-Jependens in some respects but differs because of the pubescence of the leaves, petioles, and peduncles, longer peduncles, etc. (U) [Vol. Z7 82 ANNALS OF THE MISSOUPvI BOTANICAL GARDEN 10. Peperomia PANAMENSis CDC. in Candollea 1:329. 1923, A small, spreading, stoloniferous epiphyte; stem branched, 1 mm. thick, fruiting branches ascending to 5 cm., internodes mostly 1-3 cm. long, minutely hirtcllous; leaves alternate, suborbicular, round-obovate, or occasionally cUiptic-subovate, 6 14 mm. wide X 8-20 mm. long, apex rounded, acute at the very base, palmatcly 3- to 5-nerved, both sides glabrous or sparsely minutely hirtcllous, ciliolate toward the apex; petiole 3~8 mm. long, mostly about 5 mm., sulcate above, clasping, glabrous or minutely hirtcllous; spikes terminal and axillary, 2 mm. thick X 10-15 mm. long, closely flowered; peduncles mostly 5—8 mm. long, sparsely minutely hirtcllous or glabrate; bracts round-peltate; fruit about 1 mm. long, basally immersed or eventually on pseudopedicels, subturbinate-cylindrical, apically mammiform, stigma apical. Panama and northern South America. cocl6: mountains beyond La Pintada, alt. 400-600 m., Hiuifcr & Allen 574- canal zone: in forest along the Rio Indio de Gatiin, near sea-level, Pitfier 2/8p. The minutely hirtcllous, repent stems, rounded leaves, short spikes, and shape of fruit characterize this species. II. KEY TO SPECIES MOSTLY WITH OPPOSITE OR VERTICILLATE LEAVES; FRUIT STYLOSE AND WITH A PSEUDOCUPULE. (Subgenus Micropiper (Miq.) Dahlst.). a. Leaves scarcely 10—12 mm. long, apex scarcely protracted. b. Raclus pubcrulcnt II. P. rhflexa var. AMERICANA bb. Rachis not pubcrulcnt. c. Stems and leaves glabrous; leaves longer tban wide, commonly whorlcd 12. P. QUAORiroMA cc. Stems and leaves more or less hairy. d. Leaves In whorls of 4, round or obovate-elliptic 13. P. quaternata dd. Leaves mostly opposite, orbicular 14. P. cvclophylla aa. Leaves mostly 10-20 mm. or more long, b. Leaves about 10-20 mm. long, opposite, apex scarcely narrowed 15. P. quadrangularis bb. Leaves mostly much lunger, or whorled and constricted toward the apex, c. Stem hirtcllous; leaves bluntly constrictcd-pulntcd; peduncle 4-5 cm. long 16. P. bocasensis cc. Stem essentially glabrous, or, if hirtcllous, leaves larger, narrowly pointed. d. Leaf apex minutely hirtcllous; leaves drying rather thin, 3 cen- tral nerves prominent - 17. P. martagonii olia dd. Leaf apex glabrous. e. Spikes densely flowered, commonly 6—8 cm. long; bracts con- tiguous or overlapping; leaves drying coriaceous opaque; spikes often multiple 18. P. Seemanniana ee. Spikes loosely flowered, commonly 10 cm. or more long; bracts mostly remote; leaves drying membranous, translucent, spikes mostly solitary IV. P. viridispica 11. Pkperomia reflexa (L. f.) A. Dictr. var. Americana (Miq.) Dahlst. in Kgl. Sv. Vet.-Akad. Handl. 33':175. 1900. Vcpcroniia reflexa f. amcrlcana Miq. Syst. Pip. 173. 1843, Peperotjua reflexa var. eneriulosa CDC. in DC. Prodr. 16^:452. 1869, in part (?). Peperomia americana Hertcr in Rev. Sudamer. Bot. 6:150. 1940. (U) 1950] FLORA OF PANAMA (Pipcraceae) 83 A somewhat caespitose or repent, branched, spreading, succulent, epiphytic herb; stems 1-2 mm. thick, fruiting branches ascending up to about 5 cm,, inter- nodes variable, commonly 1-4 cm. long, drying angular-grooved, hirtellous-sub- villous, the hairs often rather scattered, stiffish, and of varying lengths, up to 1 mm. long; leaves mostly in whorls of 4, rhombic-ovate, ovate, elliptic or sub- orbicular, commonly 5-10 mm. wide X 8-15 mm. long, apex rounded or slightly contracted, base acutish, glabrous above, loosely subvillous beneath, obscurely palmately 3 -nerved, drying coriaceous and subopaque; petiole mostly 1-2 mm. long, subvillous; spikes terminal, densely flowered, 2 mm, thick X 1-3 cm. long; peduncle up to 2-3 cm. long, loosely subvillous; rachis ridges hispid; bracts round- peltate; fruit immersed in pits, about 1 mm. long, subcylindrical, with basal pseudo- cupule, apex conical, stigma apical. West Indies and on the mainland from Mexico through Central America and northern South America. CHiRiQUi: vicinity of Casita Alta, Volcan de Chiriqiii, alt. 1500-2000 m., Woodson, Allen & Seibert S/Q; Chiriqui Volcano, Boquete District, alt. 2100 m., Dcnuhon 8jy, El Potrero Camp, alt. 2600 m., Killlp 3603; Rio Caldera, above El Boquete, alt. 1300-1350 m., Klllip 3512; valley of the upper Rio Chiriqui Viejo, vicinity of Monte Lirio, alt. 1300-1900 m., Seibert 147; near Los Siguas Camp, southern slope of Ccrro de la Horqueta, alt. 1700 m., Maxon 5539* Plants which are believed to represent the typical form of this species have stems minutely subreflcxed-hirtellous with the hairs only 0.1-0.2 mm. long. The contrast in pubescence appears to hold in all of the specimens examined and is the chief character upon which the variety is based. 12. Peperomia quadrifolia (L.) H.B.K. Nov. Gen. & Sp. 1:69. 1815. Pilier quadrifoUum L. Sp. PL ed. 2. 1:43. 1762. Troxinim quadrifoUum Raf. Sylva Tellur. 85. 1838. Vcperomia subquadrifoUa Trel. in Contr. U. S. Nat. Herb. 26:43. 1927. A small, glabrous, arboricolous herb; stem 1.5-2 mm. thick, repeatedly forking, rooting from the lower nodes, drying yellowish, somewhat scaly, angular; leaves in whorls of 3-7 but commonly 4 at a node, obovatc, oblong-obovate, or obovate- elliptic, emarginate, the notch bristly-hairy, base acute to subcuneate, 5-8 mm. wide X 8-14 mm. long, obscurely palmately 3 -nerved, drying coriaceous, opaque, narrowly subrevolute; petiole 1-2 mm. long, channelled above, occasionally micro- scopically hirtellous; spikes terminal, closely flowered, 1.5 mm. thick X 2-3 cm. long; peduncle 1-2 cm. long, comparatively slender; bracts round-peltate; rachis pitted; fruit obliquely ovoid, about 1 mm. long, acutely narrowed into a short, stout style, with basal pscudocupule, stigma apical. Mountains of Chiriqui province and elsewhere throughout Central America, the West Indies, and northern South America. cmiUQuf: Volcan de Chiriqui, vicinity of Casita Alta, alt. about 1500-2000 m., Wood- son, Allen ^ Seibert 8lj, gY2; eastern slope, Camp Aguacatal, alt. 2400-2 500 m., M^axon 52g8; vicinity of Cerro Punta, alt. 2000 m., Allen 1532; valley of the upper Rio Chiriqui rs3; [V(iL. Z7 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN of El Baquctc, alt. 1525-1600 m., Cornwau 3334; vicinity of El Boquete, alt. 1000-L^OO m., Maxo?j J013; Dexter Trail, El Boquete, Killip 3534- The glabrous stems and leaves, shape, size and arrangement of the leaves, and rather long terminal peduncles characterize this species. No consistent diflference has been noted by which P. snbquadrifolia can be maintained as a distinct species. 13. Pf.pi:romia quatf.unata Mlq. in Seem. Bot. Voy. Herald, 197. 1854. A small, low, compactly and divaricately branched, succulent herb; stcni sparingly hirsute; leaves In whorls of 4, round- or obovate-elliptic, rounded at both ends, scarcely 7 mm. wide X 1*^ nim. long, revolute, imprcssed-punctate, sparingly hirsute beneath, paimately 3-nerved; petiole 1-2 mm. long, retrorsely hispid; spikes terminal, closely flowered, scarcely 2 cm. long; peduncle 1 cm. long, hirsute; bracts round-peltate, golden- or red-glandular; rachis glabrous; fruit not known. Panama and (?) adjacent South America. CHiRiQUi: Volcano Chlriqui, Seemann Sgj. 14. Pepi-romta cyclophylla Miq. in Mart. FL Bras. 4^:219. 1852, name only; Seemann, Bot. Voy. Herald, 198. 1854. A small, repent, more or less crisp-pilose herb; stem about 1 mm. thick when dry, branching, internodes mostly 1-2 cm. long; leaves opposite, orbicular, mostly 5-10 mm. in diameter, occasional leaves 3-5 mm., pilose-pubescent on both sides, commonly sUghtly peltate, or the margin scarcely continuous over the petiole, loosely ciliatc, drying coriaceous, subopaque, nerves scarcely evident or faintly paimately 3-nerved; petiole m.ostly about 1 mm. long, sparsely crisp-pubcsccnt; spikes terminal or axillary, 1 mm. thick X 3-6 cm. long; peduncle 2-bracteate at or above the middle, crisp-pubescent, bracts linear or spatulate, pubescent; floral bracts round-peltate; fruit ovoid, pointed, stigma terminal. Costa Rica and Panama. Also, In closely related forms, In the West Indies and South America. Panama: without locality, Sceman>i 607, chiriqui: forests around Puerto Remedies, alt. 0-30 m., Vittier 3381. vera<;uas: trail between Caiiazas and the foot of the Cordillera Central, headwaters of Rio Cariazas, alt. 300-600 m., Allen 149. canal zone: western slope of Ancon Hill, vicinity of Balboa, alt. 20-7 5 m., Scihcrf 1 23; vicinity of Gatuncillo, ?ipcr 5652; Rio Pedro Miguel, near East Paraiso, StanJlcy 29987, Panama: second growth, near Pacora River, Wilier 1775; near big swamp east of Rio Tccumcn, Staudley 26342; Rio Tapia, SiaiiiUey 26193, 30670; between Matias Hernandez and Juan Diaz, StanJlcy 31938; Chorrcra, K/////? 340S; swamp between El Jagua Hunting Club on Rio Jagua and El Congor Hill, alt. 2 ni.. Hunter ^ Allen 487, darien: trail between Pinogana and Ynvisa, alt. about 15 m., Allcft 232, (84) 1950] FLORA OF PANAMA (Piperaccae) 85 15. Peperomia quadrangularts (Thomps.) A. Dietr. Sp. PL 1:169. 1831. Peperamia angulafa H.B.K. Nov. Gen. & Sp. 1:66. 1815 /. /. 1808. Piper angulatum Poir. in Lam. Encyc!. Meth. 4:Suppl. 468. 1816. Willd 1843. Peperomia ajigidata var. orbicans Dahlst. in Kgl. Sv. Vet.-Akad. Handl. 33": 157. 1900. Peperomia Carlo-Wrigbtil Trel. in Fcdde Rep. Sp. Nov. 23:23, 1926. Peperamia AUcui Trel. in Ann. Mo. Bot. Gard. 25:825. 1938. Peperomia saginans Trel. in Badillo in Cat. Fl. Venez. 1:246. 1945, nomcn nudum. A comparatively small, repent herb, epiphytic or on rocks; stem quadrangular, 1-2 mm. thick, minutely hirtellous to glabrate; leaves opposite, or occasionally roots replacing one of the leaves, rarely ternate, elliptic, obovate, or orbicular, 10—22 mm. wide X 12-3 mm. long, apex obtuse or very slightly pointed, base acute to rounded, palmately 3 -nerved, nerves impressed above and more or less salient beneath, drying coriaceous and subopaque, sparsely hirtellous or glabrate, more or less variegated when living; petiole 2-4 mm. long, hirtellous; spikes axillary or occasionally terminal, 2-3 cm. long; peduncles 2-4 cm. long, 2-bracteate near the middle, hirtellous; floral bracts round-peltate, glandular-dotted; fruit ovoid, stigma at the apex of a short, thick style. West Indies, Panama, and northern South America. darien: trail between Pinogana and Yavisa, alt. about 15 m., Allen 262, The specimen on which P. Allcni was based is rather fragmentary. It agrees in all respects, so far as can be determined, however, with those taken to represent this species from northern South America and Cuba. 16. Peperomia bocasensis Trel. in Ann. Mo. Bot. Gard. 27:300. 1940. A repent-assurgent, red-tinged, succulent, epiphytic herb; stems about 2 mm. thick and sulcate when dry, internodes 2-7 cm. long, crisp-hirtellous; leaves in whorls of 3-4, elliptic, apex constricted, obtuse, base subobtuse or acute, 1-2 cm. wide X 1.5-3.5 cm. long, mostly about 1.5 X 2.5 cm., palmately 3-ncrved, crisp- hirtellous above, especially along the nerves, glabrous beneath, ciliolatc toward the apex, punctate, drying coriaceous, glossy, opaque; petiole hirtellous, channeled; spikes axillary and (?) terminal, 1 mm, thick X 7-11 cm. long, closely flowered; peduncle 4-5 cm. long, hirtellous; rachis with inter-ovary, anastomosing ridges; bracts round-peltate; fruit subglobose, about 1 mm. long, obliquely pointed into a stoutish style, stigma terminal. Known only from Bocas del Toro. bocas del toro: vicinity of Nicvecita, stem and peduncles deep red, alt. about 0-50 m., WooihoHy Allen G? Seibert iSjQ. The hirtellous stems and upper surface of the blades, long spikes, and long peduncles distinguish this species. (S5) [Vol. 37 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN 17. Peperomia MARTAGOXii olia CDC. in Ann. Conserv. & Jard. Bot. Geneve 21:262. 1920, nnmc only; in Candollc:i 1:313. 1923; Sclirocder in CandoUca 3:128. 1926. An erect, somewhat branched or simple, essentially glabrous, fleshy, terrestrial, or epiphytic herb; stem up to 4 mm. thick below when dry, intcrnodcs commonly 3-4 cm, long above, the lowermost 15 cm., drying strongly ridged; leaves mostly }-A at a node, lance-elliptic, apex sharply long-acuminate, base acute-decurrent, 2-3.5 cm. wide X 5-9 cm. long, palmately 5-nervcd, the midrib and innermost pair of nerves prominent, the apex minutely hirtcllous, drying dull green, sub- membranous, subrcvolute, translucent; petiole 1-2 cm. long, glabrous, or sparsely microscopically hirtellous; spikes terminal and axillary, closely flowered, 2 mm, thick X 10-14 cm. long; peduncle mostly 8-15 mm, long, glabrous or micro- scopically hirtellous; rachis reticulately ridged between the flowers; bracts very small, round-peltate; fruit ovoid, obUquely pointed, about 1 mm. long, with basal pseudocupule, stigma apical. Mountains of Costa Rica and northern Panama. CHiiuQui: Cuesta de las Palmas, southern slope of Ccrro de la Horqueta, alt. 1700- 2100 m., Pifficr 3260; Bajo Cliorro, Boquete District, alt. 1800 m., Davidson IQQ, The conspicuous nerves, hirtellous leaf apices, lengths of petioles and peduncles, small bracts, etc. distinguish this species from P. Sccnianu'iana and other related fo rms. 18. Peperomia Seemanniana Miq. in Seem. Bot. Voy. Herald, 198, /;/. J/. 18 54. A glabrous, branched, suberect, terrestrial herb; stem 3-4 mm. thick when dry, 15-3 cm. or more tall, internodes 2-3 cm. long above, much longer downward, dr}ing ridgcd-sulcate; leaves commonly opposite, less frequently ternate, lance- elliptic, occasional leaves oblanccolate or suboblong, apex attenuately sharp- acuminate, base acute-decurrent, 2-3 cm. wide X 6-10 cm. long, palmately 5- to 7-nerved, drying coriaceous, glandular-dotted, opaque; petiole 5-10 mm. long, grooved above, stout; spikes terminal and axillary, commonly multiple, 2 mm. thick X 6-S cm. long, densely flowered; peduncle mostly 1-2 cm. long; bracts ovate-oblong-peltate, contiguous or overlapping; fruit globose-ovoid, pointed, stigma apical. Northern Panama. CHiRTQui: vicinity of El Boquctc, ait. 1000-1300 m., Vifitcr 2g2I; Maxon 3037; alt. 1400-1650 m., Killip 3542; Bajo Mono, Boquctc District, alt. 1350 m., Daiidsou 5^6; Chiriqui Volcano, Sccmanu S06, 19. Peperomia viridtspica Trel. in Contr. U. S. Nat. Herb. 26:44. 1927. Peperomia an^nslata H.B.K. var. saniamartac CDC. in Candollea 1:306. 1923. Peperomia viriJispica var. percjil Trel. Contr. U, S. Nat. Herb. 26:44. 1927. Peperomia sarcocarpa Trcl. loc. cit. 1927. A rather large, repent, pendent, glabrous, succulent, epiphytic herb; stem 3-5 mm. thick when dry X 30 cm. or more long, rooting at the lower nodes, drying (U) 1950] FLORA OF PANAMA (Pipcraccae) 87 angular-sulcate, Internodes mostly 3-6 cm. long, occasionally up to 10 cm.; leaves 2-4 at a node, more commonly opposite, subrliombically lance-elliptic, oblanceolatc, or rounded, 1.5-4 cm. v^ide X 3-7 cm. long, apex attcnuately acuminate, blunt, base acute or subcuncate, drying submembranous, translucent, palmately 3- to 5- nervedj if 5-nerved the outermost pair of laterals thin and obscure; petiole 5-10 mm. long; spikes terminal or occasionally axillary, commonly solitary, loosely flowered, 2-3 mm. thick X 8-15 cm. long or occasionally somewhat longer; peduncle 1-5 cm. long; bracts oblong-ovate-pcltate; fruit globose-ovoid, about 1 mm. long, obliquely pointed, stigma apical. Panama and Colombia. cocle: Penonome and vicinity, alt. 15-300 m., Williams j8j, canal zone: Balboa, Standley 2'JII'J, 28jjO, 28^68; vicinity of former town Empire, Culcbra Cut and vicinity, alt. 80 m., Hunter ^3 Allen jS^; on boulders in deep shade, Allen 2028. Panama: Taboga Island, Macbride 2S08; Standley 2/'8qJ\ Tumba Muerto Road, near Panama, Standley 2gS2J; between Las Sabanas and Matias Hernandez, Standley JlS/J, JlSgj; near Pacora River, Miller 1 7/6, Z///, 1 778; hills northeast of Hacienda La Joya, 50-5 00 m., Dodge, Hunter, Steyertnark & Allen i6q02. No significant difference has been found by means of which the Panamanian specimens can be differentiated from those seen from Colombia. The distantly spaced flowers and usually more membranous (when dry) leaves distinguish it from P. Seemauniana, I in. KEY TO SPECIES WITPI GLOBOSE-OVOID, SUBBASALLY ATTACHED FRUIT WITH OBLIQUE SOMEWHAT POINTED APEX AND SUBAPICAL STIGMA. (Subgenus Sphaerocarpidium Dahlsc). a. Leaves opposite or verticillate (lower stem leaves sometimes alternate). b. Stems minutely velvety hirtcllous. c. Leaves 5 mm. or less wide, 3 times or more longer than wide. d. Leaves scarcely 2 mm. wide, linear-oblong 20. P. apoda dd. Leaves 2—5 mm. wide. e. Leaves spatulatc, much longer than wide 21. P. galioides ee. Leaves elliptic, scarcely twice longer than wide 37. P. albescens cc. Leaves wider and proportionately shorter 22. P. palmana bb. Stem not hirtcllous. c. Stems glabrous; leaves commonly alternate, subplincrvcd 23. P. allagotacta cc. Stems more or less hairy. d. Stems villous, at least toward the base. e. Leaves 4-7 mm. wide 24. P. bifrons ee. Leaves 10-20 mm. wide 25. P. valliculae dd. Stems villous at the nodes only. e. Leaves mostly 3-4 at a node , 26. P. barbinodis ee. Leaves 2 or 1 at a node 27. P. Seiberth aa. Leaves alternate. b. Rachis conspicuously papillate, plants subcaulescenc 28. P. Killipi bb. Rachis not obviously papillate. c. Leaves scarcely 1 cm. long, or occasionally up to 2 cm., plants repent. d. Leaves 2-3 mm. wide, oblong-spatulate 29. P. nievecetana dd. Leaves mostly wider, rounded, elliptical, or lanceolate, c. Leaves or some of them 2-3 times longer than wide, f. Stem essentially glabrous or with only a few scattered hairs; leaves very variable in shape 3 0. P. chiriquiensis ff. Stem evidently hairy. g. Leaves elliptic-obovate, apex round-emarginate 31. P. HYGROPHiLomES gg. Leaves narrowly elliptic, apex acute 32. P. Chryslert (87) 88 l\()i.. },7 ANNALS OF THE MISSOURI BOTANICAL GARDEN ee. Leaves mostly scarcely twice as long as wide. f. Spikes scarcely 5 mm, long 33. P. coclkana ff. Spikes mostly 1-2 cm. or more long. g. Leaves, or some of them, elliptic or lance-elliptic, sub- P. CRUENTATA p. TENUICAULIS acute. h. Stems villous 34. hli. Stems minutely lilriellous 3 5. gg. Leaves mostly round, elliptic, or subobovatc, obtuse. h. Plants long-repent, leaves all alternate 36. P. rotundholia hh. Pkiots subccspitose, leaves congested 2-3 together above 37. P. ALliLSCENS cc. Leaves usually much larger. d. Leaves conspicuously black-dotted. c. Leaves elliptic-lanceolate, or ovate, mostly broader at or below the middle 38. P. GLABLLLA ee. Leaves ellipuc-oblanceolate, mostly wider above the middle. f. Leaves hairy on both sides; spikes about IS cm. long 39. ff. Leaves glabrous; spikes shorter 40. dd. Leaves not obviously black-dotted though sometimes dark- glandular. C. Stems glabrous (or somewhat pilose at the nodes only). f. Inner nerves coalescing within the lowermost 5—10 mm.; leaves more or less elliptic-oblanceolate. g. Spikes 3 cm. or more long 41. gg. Spikes 1 cm. long; peduncle equilong 23. ff. Nerves not obviously coalesccnt. g. Leaves 2.3—4.5 cm. wide. ' h. Plants ascending scarcely 20 cm.; leaves abruptly and bluntly short-acuminate, nervules by transmitted light numerous and anastomosing..... 42. hh. PUnts to 1 m, tall; leaves, at least above, narrowly attenuately sharp-acuminatc, nervules not conspicuous- ly numerous or strongly anastomosing 43. Leaves mostly 2 cm. or less wide. i. Leaves less than twice as long as wide 44. 11. Leaves 2—3 times longer than wide. j. Nodes more or less pilose; upper leaves opposite 27, jj. Nodes glabrous; upper leaves alternate 45. ce. Stems hairy. f. Leaves mostly less than 2 cm. wide. g. Plants ascending 10-15 cm.; stems villous upward 46. Plants ascending to 30 cm. or more; stems villous downward, glabrcscent above 25. ff. Leaves mostly 2-3.5 cm. wide, obtuse 47. P. flagi llispica P. rrRCUNfATA p. DIRUnORUM p. ALLAGOTACTA p. PAbCUICOLA P. ILATA ss P. LEUCOSTICTA P. Seibertii P. PARIETARIAEFOLIA P. OBSCURirOLIA Sg P. VALLICULAE P. DAVIDSliNIt 20. Peperomia apoda Trel. in Contr. U. S. Nat. Herb. 26:44. 1927. A small, stoloniferous herb; stem 2 mm. thick below when dry, rooting at the lower noJes; branches 1 mm. thick, ascending 5-8 cm., densely velvety-hirtellous; intcrnodcs mostly less than 1 cm. long; leaves mostly 4 at a node, oblong, apex rounded, sometimes emarginulate with bristle-like hairs, base acute, 2 mm. wide X 10 mm. long, glabrous on both sides or sparsely hirtellous near the base above, ciliolatc near the base, 1 -nerved, conspicuously ycllow-glandular-dotted, dryin firm, opaque; petiole about 0.5 mm. long, flat, hirtellous, marginally ciliolate; spikes terminal (?) and axillary, slender, 1 mm. thick X 4 cm. long, moderately flowered; peduncle 1—2 mm. long; bracts comparatively large, round-peltate, yellow-glandular-dotted; fruit ovoid, about 0,6 mm. long, apex rather acutely and obliquely pointed, stigma subapical. Rio Caldera valley, ChiriquJ. a (U) 1950] FLORA OF PANAMA (Pipcraceac) 89 cHiRiQuf: valley of the Rio Caldera, from El Boquete to the Cordillera, alt. 1400-1600 m., KilUp 3514, The densely hirtellous stems, whorled, narrowly oblong, glandular-dotted leaves, and obliquely, rather sharp-pointed fruits distinguish this species. 21. Peperomia galioides H.B.K. Nov. Gen. & Sp. 1:71, pi z/. 1815. ?iper galioides Poir. in Lam. Encycl. Mcth. 4:SuppL 470. 1816. Peperomia suaveolcns Ham. Prodr. Pi. Ind. Occ. 2: 1825, fide CDC. ?Peperomla MoIIiigo Willd. in Miq. Syst. Pip. 156. 1843, as synonym. Peperomia flagclliformis Hook. f. ex Miq. in Hook. Lond. Jour. Bot. 4:423. 1845. Peperomia Jamewni Regcl in Bull. Soc. Nat. Moscou 31^:544. 1858. Peperomia agapatensis CDC. in DC Prodr. 16^:455. 1869. Peperomia galioides var. lofigifolia CDC loc. cit. 464. 1869, Peperomia galioides var. nigro-punclulafa CDC loc. cit. 1869. Peperomia galioides var. umbrosa Henschen in Nov. Act. Soc. Sci. Upsal. ITT, 8:37. 1873. Peperomia galioides var. aprica Henschen, loc. cit. 1873. Peperomia sjibcorymbosa Sodiro, Contr. Conoc. Fl. Ecuator. Monogr. 1. ed. 2. 181. [1901]. Peperomia galioides var. minutifolia CDC. in Ann. Conserv. & Jard. Bot. Geneve 21:262. 1920. Peperomia granata Trel. in Fedde Rep. Sp. Nov. 23:24. 1926. A subcrect, branched, succulent, epipliytic or terrestrial herb; stem 2-3 or rarely 8 mm. thick, up to 100 cm, long but mostly much shorter, branching di- or trichotomous or verticillate, the branches often slender, wand-likc, and commonly rather strongly divergent, moderately to densely hirtellous; leaves 3-9 at a node, commonly 4-5, elliptic, elliptic-oblong, subspatulate, or oblanceolate, 2-5 mm. v^ide X 5-3 mm. long, apex obtuse, base acute, obscurely palmately 3 -nerved or subplincrved, the midrib faintly branched upward, commonly more or less puberu- Icnt along the m.idrib at least near the base above and sometimes slightly so beneath, ciHolatc toward the apex, often glandular-dotted, drying membranous; petiole very short, hirtellous; spikes terminal and axillary, solitary or in verticels of 3-6 spikes, 4-7 or rarely up to 15 cm. long; peduncle mostly about 5 or occasionally up to 10 fru long, globose-ovoid, apex oblique, stigma subapical. Wes CHiRiQUi: Chiriqui Volcano, alt. 2450 m., Killip 3j8^, This species does not appear to be common in Panama. The only specimen seen comes from the heights on Chiriqui Volcano. This specimen has leaves slightly more hirtellous than is common for the South American specimens and it appears more dingy when dry and with the leaves slightly more opaque. No other characters of any significance have been observed, however, by which it can de distinguished from this species, 22. Peperomia palmana CDC in Bull. Soc. Bot. Belg. 29^:71. 1890; 30^:233. 1891. Peperomia palmajia var. glabrior CDC in Candollea 1:304. 1923. Peperomia glabrior Trel. in Contr. U. S. Nat. Herb. 26:44. 1927. Peperomia laesa Trel. in Ann. Mo. Bot. Card. 27:402. 1940. (S9) [Vol. 37 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN An erect or subassurgcnt, freely branching, terrestrial or epiphytic herb; stem 2-3 mm. thick toward the base when dry, up to 20-30 cm. tall, rooting from the lowermost nodes, moderately to densely velvety-hirtellous, internodcs 3-5 cm. long below, reduced upward; leaves 2, 3, or 4 at a node, rather variable in size and shape, often on the same plant, elliptic, lance-elliptic, suborbicular or obovate, apex bluntly acuminate, base acute, 0.8-2.0 cm. wide X 1^5 cm. long, commonly 1-1.5 X 3-4 cm., very obscurely hirtellous on one or both sides, glabrescent, ciliolatc toward the apex, distinctly palmately 3 -nerved or the broader leaves 5- ncrvcd, drying thin and translucent, glandular-dotted, somewhat pellucid-dotted; petiole up to 15 mm. long, mostly 5-10 mm., velvcty-hirtellous; spikes axillary, 1-2 mm. thick X 2-6 cm. long, closely flowered; peduncle slender, hirtellous glabrescent; bracts round-peltate, rather conspicuously dark-glandular-dotted; fruit subglobose, about 0.5 mm. long, apex oblique, stigma subapical. Costa Rica and western Panama. CHiRiQuf: vicinity of Casita Alta, Volcdn dc Chlriqui, alt. about 1500-2000 m., Woodson, Allen GS" Scibcrt 876, gyi; vicinity of "New Switzerland," central valley of Rio Chiriqui Vicjo, alt. 1800-2000 m., Allen 1 422; valley of the Upper Rio Chiriqui Viejo, vicinity of Monte Lirio, alt. 1300-1900 m., S^cibert J 44, 149; Boquctc District, Bajo Chorro, alt. 1800 m., Davidson 178; above E! Boqucte, alt. 1400-1500 m., KiUip 3519; vicinity of El Boquete, alt. 1000-1500 m., Coruman 20 j2; between the Rio Ladrlllo and Los SiKu:is Camp, southern slope of Cerro de la Horqucta, alt. 1200-1700 m., Maxon J401, 5541 y 5553; Cuesta de la Palmas, southern slope of Cerro dc la Horqueta, alt. 1700-2100 m., Vittier 3217. In tbc original description of this species DeCandoUe said that the leaves and young stems were slightly puberulous, while he described variety glabrior in the Clavis as having glabrous stems. Trcleasc, later, in raising van glabrior to specific status, likewise claimed that the plants representing the variety were glabrous. He also described P, lacsa as glabrous. The type specimen of van glabrior has definitely hirtellous stems and some of the leaves arc likewise obscurely hirtellous. The type of P. Icicsa also shews the same characters. The leaves on the type of var. glabrior are somewhat larger than is common for other specimens cited here but this character is so variable that it does not appear advisable to use it for varietal distinction. I 23. Peperomia allagotacta CDC. in CandoUea 1:317, 378. 1923. A rather small, erect, fleshy, glabrous herb; stem 3-4 mm. thick when dry, 20 cm. or more tall, branched upward, the branches slender, divergent, internodes about 1 cm. long, those of the branches somewhat longer, drying wrinkled and subglossy; leaves alternate, those of the branches appearing opposite because of congestion, elliptic, elliptic-lanceolate, or suboblanceolate, apex somewhat narrowed but obtuse, base acute, 7 mm. wide X 2 cm. long, or those on the main stem 1.5 X 4.5 cm., 5-plinerved, the laterals coalescing with the midrib near the base, the midrib branched upward, ciliolate toward the apex, glandular-dotted, drying membranous, translucent, paler beneath; petioles 2-5 mm. long, grooved above; spikes terminal, closely flowered, 1 mm. thick X 1 cm. long; peduncle about as (90) 1950] FLORA OF PANAMA (Pipcraccac) 91 long as the spike; bracts round-peltate, about 0.5 mm. wide, glandular-dotted; ovary ovoid, apex oblique, stigma subapical; matured fruit not present, Chiriqui Volcano. CHiRiQui; around El Potrero Camp, Chiriqui Volcano, alt. 2800-3000 m., Pitiier 3T08. The plinerved leaves, congested and simulating an opposite or whorled arrange- ment on the branches, and short spikes characterize this species. 24. Peplromia bifrons Trel. in Ann. Mo. Bot. Gard. 27:3 00. 1940. A medium-Sized, assurgent herb, branching from the base, simple above; stem 3—4 mm. thick when dry, rather densely villous when young, becoming glabrous above in older stems, drying sulcate and somewhat glossy, internodcs mostly 1-3 cm. long; leaves mostly 3 or 4 at a node, oblong-elliptic or suboblanceolate, apex obtuse or subemarginulatc, base cuneate, 4-7 mm. wide X 1.5-3 cm. long, glabrous on both sides, ciliate toward the apex, obscurely palmately 3 -nerved, drying rather firm, opaque, glandular-dotted beneath; petioles 2-5 mm. long, glabrous, flattened above, clasping-decurrent; spikes terminal and axillary, 2 mm. thick X 3.5 cm. long, rather closely flowered; peduncle comparatively thick, 1-2 cm. long, glabrous; bracts round-peltate; ovary ovoid, apex oblique, stigma subapical; mature fruit not present. Known only from one locality in ChiriquJ. CHiRiQuf: Finca Lerida to Boqucte, alt. about 1300-1700 m., Wooclsan, Allen ^ Seibcrt II32. 25. Peperomia valltculai- Trel. in Ann. Mo. Bot. Gard. 24:186. 1937. A rather large, assurgent herb; stem 3-5 mm. thick below when dry, ascending to 3 cm. or more, villous toward the base, glabrcscent upward, internodcs 3-4 cm. long below, reduced upward; leaves alternate or sometimes 2 or 3 at a node, sub- rhombically oblanceolate, apex blunt, emarglnulate, base cuneate, 1-2 cm. wide X 2.5-5 cm. long, rather obscurely palmately 3 -nerved, glabrous on both sides, cillo- late toward the apex, dark-glandular-dotted, drying firm and opaque, paler beneath; petiole 5-10 mm. long, grooved above, glabrous, clasping-decurrent; spikes terminal and axillary, 1 mm. thick X 10-18 cm. long, closely flowered; peduncles 1-2 cm. long, glabrous; bracts round-peltate, glandular-dotted; fruit ovoid, about 0.5 mm. long, apex oblique, stigma subapical. Known only from Cerro Valle Chiquito, Code, cocle: western slope and summit of Cerro Valle Chiquito, alt. 700-800 m., Seibert 503. The stems villous downward, alternate-opposite-whorled, oblanceolate leaves, and long spikes are distinguishing characters of this species. 26. Peperomia barbinodis Trel. in Contr. U. S, Nat. Herb. 26:200. 1929. Veperoinia chiqucroana Trel. in Ann. Mo. Bot. Gard. 27:301. 1940. A rather large, erect, openly branching herb; stem 3-4 mm. thick below when dry, up to 30 cm. or more tall, glabrous except for the prominently villous nodes, (91) IVoi.. 37 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN intcniodcs 10 cm. long below, somewhat shorter upward; leaves 2, 3, or 4 at a node, elliptic, elliptic -lanceolate, or suboblanceolate, apex acuminate, base acute, 1-2 cm. wide X 3.5-6 cm. long, glabrous and paler beneath, the nerves above crisp- pubescent, densely ciliolate toward the apex, 5-nervcd, the midrib and innermost pair of lateral nerves prominent, the outermost pair inconspicuous, coalescing near the base, drying membranous, translucent, faintly glandular-dotted; petiole about 5 mm. long, glabrous, grooved above; spikes terminal and axillary, loosely flowered, very slender, 6 cm. long; peduncle about 1 cm. long, glabrous; bracts round- peltate; ovary ovoid, apex oblique, stigma subapical. Costa Rica and w^estern Panama. CHiRiQUi: Bajo Mona, mouth of Quebrada Chiqucro, along Rio Caldera, alt. about 1500-2000 m., Woodson, Allen 6 Scihcri 103 j. The type, Tondnz IO419, from Costa Rica, has leaves smaller than the Pana- manian collection cited above, but otherwise the specimens appear to be nearly idcnticah 27. PianiROMiA Seiberth Trcl. in Ann. Mo. Bot. Card. 24:185. 1937. A moderately large, openly branched, essentially glabrous herb, drying yellow; stem 3 mm. thick below when dry, up to 3 cm. or more tall, somewhat villous at the nodes, internodes up to 10 cm. long below, reduced upward; leaves both alternate and opposite, broadly lanceolate, apex sharply acuminate, base acute, 1-2.5 cm. wide X 2.5-6 cm. long, palmately 3- to 5-nerved, the midrib and inncrmos.t pair of lateral nerves salient bencatli, pale-glandular-dotted and glabrous beneath, pubescent at the base above, ciliatc toward the apex, drying membranous, trans- lucent; petiole scarcely 5 mm. long, grooved above, alately decurrent; spikes terminal and axillary, 1 mm. thick X 5-7 cm. long, loosely flowered; peduncle slender, 5-10 mm. long; bracts round-peltate; ovary ovoid, apex oblique, stigma subapical; fruit not present. Near Monte Lirio, Chiriqui. CHTRiQL'f: valley of the upper Rio Chiriqui Viejo, vicinity of Monte Lirio, alt. 1300- 1900 m., Sciberi 301. The alternate-opposite leaf arrangement, leaves pubescent at the base above, and pilose nodes distinguish this species. 28. Pepf.romia KiLLiPi Trel. in Bot. Gaz, 73:143. 1922. Pepcrowhi bytfiVfioJcs Trcl. in Contr. U. S. Nat. Herb. 26:43. 1927. A small, very short-stemmed herb; stem 3 mm. thick, subrhizomatous, ascend- ing 2-3 cm., glabrous, internodes very short, drying wrinkled and scaly; leaves alternate, round- to elliptic-ovate, apex acute or obtusish, base cordate or cordulate, 1.5-3.5 cm. wide X 3-6 cm. long, sparsely appressed-pubescent above, glabrous beneath, rather densely ciliolate throughout, palmately 7-nerved, drying very thin, translucent, pale glandular-dotted; petioles very slender, commonly 3-5 cm. long, glabrous or sparsely pubescent along the groove, abruptly enlarging into a thick- (92) 1950] FLORA OF PANAMA (Piperaccae) 93 cned base which remains as part of the stem when the leaf falls off j spikes apparently axillary, 1 mm. thick X 5-9 cm. long, closely flowered; peduncles up to 5.5 cm. lc>ng, glabrous; racliis ridges conspicuously papillate; bracts round-peltate, granu- late; fruit ovoid, scarcely 0.5 mm. long, apex oblique, stigma subapical. Panama. CHiRiQuf: eastern Chiriqui, vicinity of San Felix, alt. 0-120 m., Piftier 5266. e of which has not been examined, appears to be very closely related uo P. caitdiililhnba. As described, it appears to differ in its glabrous plants and 5-nerved leaves. However, some leaves of P. caudulilimba may be 5- (m) [Vol. 37 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN nerved with an obscure outer pair of lateral nerves, and the stems and leaves are sometimes nearly glabrous. It is possible that further study may show that this and P. cauiIuJiliiiiba arc diflFcrent forms of the same species. 49. Peperomia caudueilimba CDC. in Ann. Conscrv. & Jard. Bot. Geneve 21:269, 1920. Peperomia gafiincmis CDC. in Candollea 1:363. 1923. Vcperomia pipcroruni Trel. in Concr. U. S. Nat. Herb. 26:49. 1927. Peperomia catululdimba var. cryptopoJa Trel. loc. cit. 1927, ?Peperomia caiiJHlilimba var. Jongepotdiila CDC. loc. cit. 1920. A moderate-sized, often long-pendent, arboricolous herb; stem 2-3 mm. thick when dry, rooting from the lower nodes, branches often long and pendent, more or less villous, usually at least about the nodes, intcrnodcs up to 7-8 cm. long below, reduced upward; leaves alternate, lanceolate or lance-clKptic, apex narrowly sharp- acuminate, base somewhat rounded, obtuse or subacute, mostly 1-2.5 cm. wide X 4—8 cm. long, occasionally up to 4 X 1 1 cm., glabrous on both sides, or rarely sparsely hairy beneath, pinnatcly nerved from below the middle with two pair of nerves approximated near the base and a third pair toward the middle, usually more or less dilate, mostly somewhat yellow glandular-dotted beneath, drying coriaceous, opaque; petiole less than 5 mm. long, stout, commonly ciliate; spikes solitary or often paired on bracted stalks 5-10 mm. long, closely flowered, 1-2 mm. thick X 8-12 cm. long; peduncle glabrous or sparsely hairy, 1-1.5 cm. long; bracts rcund-pcltate; fruit reddish, cylindrical, about 2 mm. long, apex obliquely truncate, scutcUiform, stigma central on the scutcUum. Panama. BOCAS DEL TORO: Old Bank Island, vicinity of Chiriqui Lagoon, I'on WeJe! lS^2; Water Valley, rotj Wedel l6pO; Farm 6, Chant;uinoIa Valley, Duulap 4S6, cocle: above Pcnonomc, Williiims j66. canal zone: Couell 2JQig; railroad relocation between Gor- gona and Gatun, alt. 10-5 m., Pit tier 2280; Rio Agua Salud, near Frijoles, Piper 6ojj; vicinity of Fort Sherman, Sia/nlley jIllS; forest aloni; the Rio Indio de Garun, near sca- Icvcl, Piffier 2/88; Maxon 4818, 4864; near Rio Medio, Miller TJ'jj near Salamanca Hydro- graphic Station on the gorge of the Rio Pequcni, alt. 70-80 m., Dodge^ Steyermark d Allen 16082 ; Barro Colorado Island, Gatiui Lake, Sfaudley JIJ2I, J 1 336, 4116/. dariln: Boca de Panarando, on Sambu River, alt. 20 m., Piffier 35Ti . 49a. Peperoaua caudulilimba var. cyllndribacca (CDC.) Yuncker, comb, nov. Peperomia cylindribacca CDC. in Candollea 1:370. 1923. Leaves 4-5 cm. wide X 9-11 cm. long, densely ciliate; fruit 2.5 mm. long, scutellum acute. Known only from the type locality. CHIRIQUI: vicinity of San Felix, alt. 0-120 m., Dec. 1911, Piffier 527J (U. S. Nat. Herb., type). This species is one of a series, including P. uvicrostacbya of South America, which appears to be very closely related and may, indeed, represent only variations of a single polymorphic species. Careful field observations, including mass collec- (104) 195 0] FLORA OF PANAMA (Piperaccac) 105 tions, are needed to interpret adequately the various forms which have been described. It Is very closely related to P. oblongibacca from which it differs because of its mostly shorter petioles, more coriaceous leaves, and with villous rather than hirtellous type of pubescence. 50. Peperomta oblongibacca CDC. in Candollea 1:288. 1923. A modcrat:e-sized, epiphytic herb; stem 2-3 mm. thick, branches somewhat pendulous, glabrous or obscurely and minutely hirtellous, internodes mostly 2-3 cm, long; leaves alternate, elliptic, lance-elliptic or suboblanceolate, apex sharply long-acuminite, base acute to cuneatc, glabrous on both sides, loosely ciliate, 5- or obscurely 7-plinerved from below the middle, glandular-dotted beneath, drying thin, translucent; petiole slender, up to 2 cm. long below, reduced upward; spikes 1, 2, or sometimes 3 on short, bracted, sympodial branches, 1—1.5 mm. thick X 15-20 cm. or more long, closely flowered, peduncle 1 cm. long, obscurely hirtellous glabrescent; bracts comparatively large, round-peltate; fruit subcylindrical, 2-3 mm. long, apex obliquely scutelliform, stigma central on the scutcllum. Rio Fato, 'Colon. colon: along Rio Fato, in forests or thickets, alt. 10-100 m., Pit tier jSqi. KEY TO THE VARIETIES OE P. OBLONGIBACCA a. Mmutely pubcrulcnt on young growth, or glabrous; petioles not ciliate. b. Leaves 6—7 cm. long. c. Leaf base subcuneatc; stems obscurely hirtellous cc. Leaf base acute; stems puberulent bb. Leaves 10—12 cm. long aa. Nodes more or less sparsely villous; petioles ciliate P. oblongibacca Var. MARRACANTTNA var. MACROrilYLLA var. SUBVILLOSA Fig. 19. Peperomia obi au^^r folia var. viacropbylla 50a. Peperomia oblongibacca var. marragantina (CDC. ) Yuncker, comb. nov. Peperomta cauJuUVimba var. marragantina CDC. in Candollea 1:382. 1923, as viar- reganfinay name only. Young growth velvety-puberulent; leaves lance-ovate. Known only from the type locality. PANAMA: Marraganti, April 6, 1905, Williams 6q4 (U. S. Nat. Herb., type). 5 Ob. Peperomia oblongibacca var. macrophylla Yuncker, var. nov. Caule juvenili minute hirtello vel glabrato; foliis 7-plinerviis. Young growth obscurely and minutely hirtellous or glabrous; leaves 3.5-5 cm, wide X 12—14 cm. long, 7-plincrved. Known only from the type locaHty. (105) [Vol. 37 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONI-: drowned forest near Vigla and San Juan on Rio Pequeni, alt. 66 ni., Nov. 26, 1934, Dodge, Steycrmark ^ Allen r6^22 (Herb. Missouri Bot. Card., type). 50c. Pbperomia oiiLONGiiiAccA var, subvillosa Yunckcr, var, nov. Caulibus ad nodos dissitc villosis; foliis ovatis, 5-7-plinerviis; pctiolo ciliato. Nodes more or less sparsely villous; leaves ovate, 5- to 7-plinerved; petiole cotr- monly ciliate. Known only from the type locality. CANAL zone: near Rio Medio, Feb. 11, 1937, Milla 1740 (U. S. Nat. Herb., Tvpr). The plants Included here are all obviously rather closely related but differ in the characters as described. It is not believed that they represent individual species and there may be some question regarding their status even as varieties. 5L Peperomia serpens (Sw.) Loud. Hort. Brit. 13. 1830. Piper serpens Sw, Trodr. Vcg. Ind. Occ. 16-1788. Peperowia scanJcns Ruiz & Pav. Fl. Peruv. & Chil. 1:32, pL 5/, 5.!?. 1798. Piper scandens YM.Enum. l:}46, 1804, not Sw. 1788. Piper bracteatiun Thomps. in Trans. Linn. Soc. 9:203, /)/. 21, 1808. Peperowia repens H.B.K. Nov. Gen. & Sp. 1:65. 1815. Peperomia reniformis Hook. Exot. Fl. 3:]?/. 164. 1827. Piper Guildingianum Sprcng. Syst. Vcg. 4. Cur. Post. 21. 1827. Peperomia pulicaris Opiz in Presl, Reliq. Haenk. 1:162, /)/. 30. 1830. Peperomia myosnrus hb. Willd. in A. Dietr, Sp. Pi. 1:159. 1831. Peperomia Guddingiana A. Dletr. loc. cit. 162. 1831. Piper myosuriis hb. Willd. in D. Dietr. Syn. Pi. 1:110. 1839. Verbuellia serpens Miq. Syst. Pip. 49. 1843. Acrocarpidiiim pulicare Miq. loc. cit. 57 1843. Acrocarpidium repens Miq. loc. cit. 59. 1843. Acrocarpidium scandens Miq. loc. cit. 1843. Piper herbctcenm Miq. loc. cit, 1843. Acrocarpidium Guildingianum Miq. In Hook. Lond. Jour. Bot. 4:412. 1845. Bot. 4:143. 1866. / 1869. A comparatively small, creeping or scandent herb; stem slender, crisp-pubescent; leaves alternate, round-subdeltoid, mostly 1-2 cm. wide or occasionally up to 3 cm., apex acute or obtuse, base truncate-cordulate, palmately 3- to 5-nerved, drying rather membranous and thin, more or less crisp-pubescent on both sides; petiole 5-30 mm. long, crisp-pubescent; spikes axillary and sympodially terminal, up to 3 cm. long; peduncle 1- or rarely 2-bracted, usually about as long as the spike, crisp-pubescent; floral bracts round-peltate; fruit about 1 mm. long, ellipsoidal, stigma at the base of the slender, pointed beak. West Indies, and on the mainland southward from Panama to Brazil and Peru. BOCAS DEL TORO: Vicinity of Chlriqu! Lagoon, von Wedel TOJ4. canal 7one: Rio Indio dc Gatun, near sea-level, Maxon 4^57- 52. Peperontta urocarpoides CDC. in CandoUea 1:362. 1923. A rather small, repent, crisp-hairy herb; stem 1-2 mm. thick, rooting from the nodes, interncdcs mostly 2-3 cm. long; leaves alternatCj ovate, apex gradually bluntly pointed, base rounded, obtuse, cordulate, or acutish at the petiole, 1.5-3 (106) 1950] FLORA OF PANAMA (Piperaceac) 107 cm. wide X 2-3.5 cm. long, crisp-pubescent on both sides, cilia te, palmately 5 -nerved, the nerves branched upward, more or less glandular-dotted, drying sub- coriaceous, subopaque ; petiole 1—2 cm. long, crisp-pubescent ; spikes axillary or terminal, 1 mm, thick X 3 cm. long; peduncle crisp-pubescent, 1 cm. or more long, bracteate at about the middle; floral bracts round-peltate; fruit oblong, obliquely slender-mucronate, stigma near the base of the beak. Panama. cocle: mountains beyond La Plntada, alt. 400-600 m.. Hunter ^ Allen 5//. darien: Cerro de Garagara, Sambu basin, southern Darien, alt. 500-974 m., Piftier $^7^- This species differs from P. 7irocarpa in the shape of the 5 -nerved leaves, etc. 53. Peperomia digitinervia Trei. in Ann. Mo. Bot. Gard. 27:302. 1940. A repent, epiphytic herb; stem 1—2 mm. thick, rooting at the nodes, rather densely crisp-puberulent, internodes 2—4 cm. or more long; leaves alternate, ovate, apex acute, base subtruncate, cordulate or acute at the petiole, 2—3 cm. wide X 2.5—4 cm. long, subpalmately 5 -nerved, the innermost pair of lateral nerves coalescing with the midrib within the lowermost 2-3 mm., sparsely pubescent above, paler and crisp-puberulent beneath, at least along the nerves, ciliate, drying coriaceous, subopaque; petiole 1—3.5 cm. long, pubescent glabrescent, channelled above; spikes leaf-opposed, 2 mm. thick X 3.5 cm. long, densely flowered; peduncle 2 cm. long, crisp-puberulent, 1-bractcate near the middle; floral bracts round- peltate; fruit ellipsoidal, about 0.8 mm. long, rostrate, stigma anterior at the base of the beak. El Valle, Code. cocle: vicinity of El Valle, alt. 600-1000 m., Allen I22I, This species resembles P. ciibensis to some extent but differs because of the pubescence, smaller leaves, etc, 54. Peperoaita distachya (L.) A, Dietr. Sp. Pi 1:156. 1831. Piper dhtachyon L. Sp. Pi. 1 :30. 1753. Peperomia prod net a Griseb. Fl. Br. W. Ind. 166. 1864. Peperomia Bakerii CDC. in Urb. Symb. Ant. 5:296. 1907. ^ A sparsely hairy or glabrate, creeping or sometimes scandent herb; stems 2—3 mm. thick, rooting at the nodes; leaves alternate, elliptic-ovate, 4-5 cm. -u'ide X 7-8.5 cm. long, apex acuminate, typically rounded at the base and sometimes cordulate or barely subpeltate, drying rather thin and dull, sparsely hairy beneath when young, or glabrous on both sides, ciliolate towards the apex, 9-plinerved with the nerves coalescing below the middle; petiole 3-5 or sometimes even 9 cm. long, glabrous or somewhat ciliate; spikes 3—5 cm. or more long, mostly paired at the ends of terminal or axillary stalks 3-4 cm. long; peduncle scarcely 2 cm. long; bracts round-peltate; fruit ellipsoidal, rostrate, stigma at the base of the beak. Common in the West Indies. Apparently rare in Panama. Also reported from the Guianas and Ecuador. (107) [Vol. 37 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN liOCAS DLL TORo: Fish Creek, vicinity of Chiriqui Lai;oon, ivfi WeJcI 21 Q'^, cocLr : mountains bcyoad La Pintada, alt. 400-600 m., Hunter ^ Allen 5Jj, panama: summit of Ccrro Campana, alt. 800-1000 m., Allen 2224, 55, Peperomia machaerodonta Trel. in Contr. U. S. Nat. Herb. 26:47. 1927. A rather small, suberect, epiphytic herb; stem 3 mm. thick, simple, ascending to 10 cm. or more, glabrous, drying wrinkled, yellow-glossy, intcrnodcs 1~2 cm. long; leaves alternate, elliptic, round-elliptic, or subobovate, apex abruptly short- acuminate, base acute or subrounded, glabrcsccnt beneath, crisp-puberulent ahove, especially along the nerves, pinnately nerved to above the middle, the nerves about 4 on each side, 2—3 cm. wide X 3—5.5 cm. long, drying coriaceous, opaque, some- what glossy-wrinl'Jed, especially near the margin, with scattered glandular-dots; petiole 5—10 mm. long, glabrous, yellow-glossy, channelled above; spikes terminal, 2 mm. thick X 6~8 cm. long, rather closely flowered; peduncle 1.5-2.5 cm. long, glabrous; bracts round-peltate, pale-glandular-dotted; fruit ellipsoidal, about 1 mm. long, rostrate, stigma anterior near the base of the pale, subteretc, erect, slightly curved, equilong beak. Valley of the Rio Piarnasta, east of El Boqucte, Chiriqui. CHIRIQUI: valley of Rio Piarnasta, about 5 miles east of El Boquete, alt. 1525-1600 ni., KilUp 3538, The rather small plants, pubcrulent upper leaf surface, small, pinnately nerved leaves, and type of fruit characterize this species. 56. Peperomia obtusifolia (L.) A. Dietr. Sp. Pi. 1:154. 1831. ?iper obtusijolium L. Sp. Pi. 30. 1753. ^Viper cuneifolitim Jacq. Collect. 4:127, pi 214, 1788. Piper humile Mill. Card. Diet. no. 4, ex Polr. in Lam. Encycl. Meth. 5:473. 1804, fide CDC. in Urb. Symb. Ant. 3:254. 1902 Viper Milleri Roem. & Schult. Syst. Veg. 1:337. 1817, fide CDC, loc. cit. 1902 Ant. 3:254. 1902. 'folia 1825, fide CDC In Urb. Symb. ?repero?nia cuneifolia A. Dietr. Sp. PL 1:146. 1831. Peperomia coinmutata Trcl. in Fcdde Rep. Sp. Nov. 23:29. 1926, In part. Peperofnia hayafana Trel. loc. cit. 30. 1926. Peperomia daiquiriana Trcl. loc. cit. 1926. Peperomia Lunana Trel. loc. cit. 1926. Pe[)eromia Earlei Trel. loc. cit. 31. 1926. Peperomia iloJecafheontophylla Trcl. in Contr. U. S. Nat. Herb. 26:48. 1927. Peperomia Antoni var. reJucfa Trcl, in Ann. Mo. Bot. Card. 27:299. 1940. A fleshy, stoloniferous-assurgcnt, terrestrial or epiphytic herb, glabrous except for the peduncle which is minutely hirtellous as is also sometimes the lower leaf- surface; stem up to 5 mm. thick when dry, ascending to 15 cm. or more from the prostrate, rooting base, incernodes mostly 2—3 cm. long; leaves alternate, elllptic- obovate, 3—8 cm. wide X 5—20 cm. long, occasional specimens with smaller leaves, but commonly about 4-6 X 8-12 cm., apex rounded, obtuse, sometimes slightly emarginate, base cuneate, 7- to 11-plInerved from below the middle, the midrib branched upward, drying more or less coriaceous, opaque; petiole mostly 1 .5-4 (10^) 1950] FLORA OF PANAMA (Piperaceac) 109 cm, long, or occasionally somewhat longer, more or less winged by the decurrent blade; spikes 2-4 mm thick X 5-15 cm, long, densely flowered, solitary or paired on a 1-bracced, terminal, leaf-opposed, or occasionally axillary, usually minutely hirtellous stalk 3-5 cm. or more long; floral bracts 0.3-0.4 mm. wide, round- peltate; fruit ellipsoidal, about 1.2 5 mm. long including the slender, erect, terminally hooked beak, stigma anterior at the base of the beak. General in the West Indies and on the mainland from Mexico through Central America and northern South America. cocle; vicinity of El Valle, alt. 600-1000 m., Allcfi 1223, canal zone: along Rio Indio de Gatiin, Viltier 280O; Summit, Lindsay 402. 56a. Peperomia obtusifolia van emarginata (Ruiz & Pav.) Dahlst. in Kgl. Sv. Vet.-Akad, HandL 33^:66. 1900. Peperofnia emarghjata Ruiz & Pav. Fl. Peruv. & Chil. 1:33, pL 4g, f.a. 1798. Piper emarghiatum Vahl, Enum, Pi. 1:339. 1804. Stems short ; leaves oblong-elliptic-obovate, apex somewhat narrov/cd and rounded, emarginate, mostly 2 to 3 times longer than wide; spikes 2-3 mm. thick, up to 8—15 cm. long, single, or more commonly divergently paired on a common, terminal, or leaf-opposed stalk 1-4 cm. long, or occasionally subpaniculately 3- spiked; fruit about 1 mm, long. Found with the species. veraguas: vicinity of Santa Fe, forested slopes of Cerro Tute, alt. 900 m., Allen 4J34^ canal zone: near Salamanca Hydrographic Station on the gorge of the Rio Pequeni, 70-80 m., Dodge, Steyermark & Allen- l6j2j, l6j2jb; near Rio Medio, Miller 1736. 56b. Peperomia OBTusii'OLiA var. longi- bracteata Yuncker, var. nov. Foliis ellipticis; spicis 3 in panicula clongata laxa axillari bracteata dispositis, bracteis lineari- lanceolatis usque ad 3 cm. longis; pedunculo 2—4 cm. longo minute hirtello; ovario ellipsoideo f.J Fig. 20. Pepcro?rtia obtusifolia var. longihractcata rostrato. Leaves elliptic, 4-5 cm. wide X 10—12 cm. long; spikes 3 mm. thick X 5 cm. or more long, in a 3-spicatc, elongate, loosely paniculate, axillary inflorescence, the main stalk 8—10 cm. long, in- dividual peduncles 2—4 cm. long, minutely hirtel- lous, in the axils of linear-lanceolate bracts up to 3 cm. long; floral bracts subquadrangular-pcltate, strongly imbricate in young spikes; ovary ellip- soidal, rostrate, stigma at the base of the beak; fruit not matured. South-central Panama. COCLE: region north of El Valle de Anton, alt. 1000 m., Sept. 27, 1946, Allen 37 30 (Herb. Missouri Bot. Gard.,TYPE) ; La Mesa, Aug. 31, 1941, Allen 2728. (109) [Vol. 37 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species exhibits great variation in the size and shape of the leaves ant! length of the spikes, sometimes on the same plant. It has not been possible to main- tain as distinct the several species proposed by Trcleasc. The variability of the plants, readily observed in the field, leads one to conclude that only one species is represented. The hirtellous peduncle and commonly also the lower leaf surface, much smaller bracts, and more slender and strongly hooked beak of the fruit distinguish it from P. magnollacfoha. Variety hugibracfeata bears considerable resemblance to many specimens be- lieved unquestionably to represent P. oh fusi folia. The elongated 3-spicate, long- bracteate, paniculate inflorescence, and strongly imbricated floral bracts are rather strong differential characters. Additional collections may show that this variety merits specific status. In the meantime, however, it is given a rather questionable varietal status. The axillary inflorescence, etc. distinguishes it from P. viagnoJiae- folia, and the leaf shape and size, especially, from P. W 57. Peperomia magnoliaefolta (Jacq.) A. Dietr. Sp. PL 1:153, 1831. Tiller Ttta^uoliaefolium Jacq. Collect, 3:210. 1789. Viper subrofumlum Haw. Rev. PI. Succ. 2. 1819. Pepcrojuia stibrotunda A. Dictr. loc. cit. 154. 1831, Vcpcromia auiplexicauUs (Sw.) A. Dictr. var. magnoJiacjolia (A. Dietr.) Griseb. Fl. Br. W. Ind. 176. 1859. Teperomta euosma Trcl. in Fcdde Rep. Sp. Nov. 2 3:29. 1926. Peperofftra Antoni Trel. in Ann. Mo. Bat. Gard. 24:185. 1937. Pepcromia Afitoui f, fcrtilior Trcl. loc. cit. 17:299. 1940. Pepcromta Antoni f. lutca Trel. loc. cit. 300. 1940. A moderately large, assurgent, glabrous, succulent herb; stem 3-5 mm. thick, rooting at the lower nodes, ascending to 2 5 cm. or more, intcrnodcs mostly 1-5 or 6 cm. long; leaves alternate, round, obovate-elliptic, or subspatulate^ 3-8 cm. wide X 6-15 cm. long, apex rounded, often emarginate, base cuncate or sometimes rather abruptly contracted, drying dull, coriaceous, opaque, 7- to 9-plinerved from near the base, the rather prominent midrib pinnately branched upward, the lateral nerves slender and obscure; petiole 1-3 cm. long, more or less winged by the decurrent blade; spikes 3 mm. thick and up to 15-18 cm. long, closely flowered, 1-2 or sometimes 3 on a terminal or leaf-opposed, bracted stalk 5-10 cm. long; peduncle 1.5-3 cm. long, glabrous or rarely obscurely hirtellous; floral bracts round-peltate, mostly about 0,5 mm. wide; fruit about 1.25 mm. long, ellipsoidal, rostrate, the beak gradually narrowed from a broad base, from one-fourth to as long as the fruit, sharp and more or less curved but usually not hooked at the apex; stigma at the base of the beak. West Indies, Panama, and northern South America. BOCAS DEL TORO: Carccning Cay, Isla Colon, alt. 0-120 m., von Wcdcl j6^. cocle: El Valle de Anton and vicinity, alt. 500-700 m., Scibcrt 437; alt. 800-1000 m., Allen 760, Il60y collection labelled "lata" on the herbarium sheets; between Las Margaritas and El Valle, Woodson, Allen ^Jf Seibert I2g8, canal zone: Experimental Gardens, Stanivood 402. (110) 195 0] FLORA OF PANAMA (Piperaccac) 111 This species Is closely related to P. obtnsifol'ia from which it is to be distin- guished, however, because of its glabrous peduncles and lower leaf surface, wider floral bracts, and with the fruit beak mostly stouter and ordinarily not hooked at the end. 58. Peperomia HispiDULA (Sw.) A. Dictr. Sp. Pi. 1:165. 1831. Piper hispidulum S\<. Prodr. Veg. Ind. Occ. 15. 1788. Acrocarpidiuju hispidulum Miq. Syst. Pip. 54, 1843. Acrocarpidiuni Selloviannm Miq. loc, cit. 55. 1843. ?Peperamia tenera Miq. in Mart. Fl. Brasil. 4^:19, pL /, /. I. 1852. ?Peperomia pilinervia CDC. in Bull. Herb. Boiss. 6:509. 1898. ?Peperomia hispidula var. Sellowiana Dahlst. in Kgl. Sv. Vet.-Akad. Handl. 33-;14. 1900. Peperomia perhhpidula CDC. in Engl. Bat. Jahrb. 40:257. 1908. Peperomia hispidula var. perhispldula CDC in Candollea 1:335. 1923. Peperomia Woodsonii Trel. in Ann. Mo. Bot. Card. 27:305. 1940. A rather small, assurgcnt, branched, spreading herb; stem 2-3 mm. thick below when dry, branches ascending to 20 cm. or more from the decumbent, root- ing base, hispid at least at the nodes, the hairs rigid to rather flaccid, internodes variable in length; leaves alternate, ovate-orbicular or rhombic-ovate, 8—15 mm. in diameter, apex rounded, obtuse, base rounded or triangular, obtuse, loosely hispid above or glabrate, loosely hispid beneath at least along the nerves, palmately 5- nerved with the midrib strongly branched at about the middle, drying membranous and subpcllucid; petiole commonly 5-15 mm, long, slender, hispid to glabrate, but usually with at least a few hairs near the blade; spikes leaf-opposed or terminal, slender, mostly 1-2 cm. long, loosely and alternately flowered; peduncle slender, 10—15 mm, long, hispid to glabrate; bracts round-peltate; fruit globose-ovoid, about 1 mm. long, covered with white, hispid hairs and on a pedicel up to 0.5 mm. long when manure, stigma apical on a conic style. West Indies, Central America, and northern South America. Apparently in- frequent in Panama. CHiRiQuf: vicinity of Casita Alta, Volcan de Chiriqui, alt. 1500-2000 m., Woodson, Allen e? Seibert pSj;. Dahlstedt included this species in the subgenvis Acrocarpidtum (Miq.) Hcnsch. VI. KEY TO SPECIES WITH PELTATE LEAVES a. Fruit scarcely rostrate, apex acute, mammiform, or obliquely scutelli- form. b. Stems very short, scarcely I cm. long; rachis strongly hispidulous 59. P. hispidorhachts bb. Stem5 much longer. c. Leaves pubescent beneath; petioles more than half as long as the blade, slender 60. P. cerro-puntana cc. Leaves glabrous; petioles shorter, stout 61. P. cordulata aa. Fruit rostrate. b. Petiole attached 1—2 cm. or more from the leaf margin. c. Main lateral nerves conspicuous and strongly curved toward the apex - - 62. P. flexinervia cc. Main lateral nerves rather obscure and ascending more sharply. d. Leaves mostly 6—8 cm. wide 63. P. ciliifera dd. Leaves mostly 5 cm. or less wide 64. P. conserta (111) [Vol. 37 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN bb. Petiole attached within 1 cm. of the margin. c. Leaves 10 cm. or more long 65. P. maculosa cc. Leaves 3 cm, or less long 66. P. pfrglandulosa 59. Plperomia hispidorhachis Yunckcr, sp. nov. Hcrba paiva eplphytlca; caule pcrbrevc; foliis alternis ovato-lanceolatis, apice sensim acutis, basi rotundatls ca. 5 mm. ct basi peltatis utrinque cllssitc crispc-villosis (pilis usque ad 1 mm. longis) , ciliatis, palmatim 7— 9-nerviis lutco-glanduloso- punctatis; petiolo gracili 4-5 mm. longo dissite villoso, ad basim dilatato; spicis axillaribus; pcdunculo filiformi spicam ipsam aequantc, rhachi confertc hispido; drupa angusto-cUipsoidea, brevc-pedicellata, apice subobliquo, stigmatc apicall. A small, epiphytic plant; stem very short, scarcely 1 cm. long X 3 mm. thick, tumid with thickened leaf scars; leaves alternate, ovate-lanceolate, apex gradually acute, base rounded, peltate about 5 mm. above the base, 2.5-4 cm. wide X 6—8.5 cm. long, loosely crisp-villous on both sides, the hairs up to 1 mm. long, ciliate, palmately 7- or In large leaves faintly 9- nervcd, the innermost pair of lateral nerves coalescing with the midrib within the lowermost 5-10 mm., the lateral nerves simple or in large leaves more or less forked, drying very thin, pellucid, yellow-glandular-dottcd; petiole slender, 4—5 cm. or more long, loosely villous, base much enlarged; spikes axillary, scarcely 1 mm. thick, 7-9 cm. or more (?) long, loosely flowered; peduncle filiform, about as long as the spike, loosely villous; rachis densely and prominently hispid; bracts round-peltate; fruit narrowly ellipsoidal, apex slightly oblique, narrowed toward the base to form a short stalk, stigma essentially apical. Known only from the type locality. PANAMA: Rio La Maestra, alt. 0-25 m., Dec. 4, 1936, Allen J/ (Herb. Missouri But. Gard., type). This species bears some resemblance to P. tccticola of Costa Rica, Tt differs, however, according to DeCandolle's description, because of the leaf shape, shorter spikes, hispid rachis, and shape of the substalked fruit. It closely resembles P. KilUpi in general appearance. It differs, however, because of its villous, definitely peltate leaves, strongly hispid rachis, and fruit shape. 60. Peperomia cerro-puntana Trek in Ann. Mo. Bot, Gard. 27:301. 1940. Fig. 21. Pcpcratftia hispidorhachis A terrestrial or epiph)'tic mountain herb; stem 2-3 mm. thick, more or less decumbent and rooting from the lower nodes, glabrous, internodes 2—5 or 6 cm. long; leaves alternate, round-ovate or ovate, apex slender-acuminate, base round, obtuse, peltate about 1 cm. from the margin, 4-5 cm. wide X 6-8 cm. long, glabrous above, loosely pubescent beneath, the hairs about 0.5 mm. long, drying ratlier membranous, translucent; petiole slender, up to 7 cm. long, clasping, sparsely hairy glabrcscent; spikes leaf-opposed, 2-3 mm. thick X 7-12 cm. long, moderately flowered; peduncle 2-3 mm. long, pubescent; rachis somewhat papillate; bracts (112) 19 5 0] FLORA OF PANAMA (Piperaccac) 113 round -peltate; ovary obovoid-turbinate, stigma apical. Mountains of Chiriqui province. CHiRTQUi: trail from Cerro Punta to headwaters of Rio Caldera, alt. 2250-2500 ni., Allen 145^1 Bajo Chorro, Boquete District, in rain forest, alt. 1800 m., Daihhou 2ig^ The peltate leaves with villous lower surface, rather long spikes, and subturbi- nate ovary with apical stigma arc distinctive characters of this species. 1866. 61. Peperomia cordulata CDC. in Jour. Bot. 4:137. A pendent, glabrous, epiphytic herb; stem 5 mm. thick when dry, internodes 2-4 or 6 cm. long; leaves alternate, round-ovate or ovate, apex acute to subacumi- nate, base rounded, peltate within 1 cm, of the margin, 5-8 cm. wide X 7-11 cm. long, palmately about 11 -nerved, drying rather firm, subtranslucent; petioles stout, 2—3 mm. thick, 1-4 cm. long; spikes on short, bracteate, leaf-opposed branches, 2 mm. thick X 15 cm. long, densely flowered; peduncle 1.5-2 cm. long, bracts rather large, lanceolate; floral bracts round-peltate; fruit subglobosc, apex obliquely truncately scutelliform, stigma central. Panama. CANAL zone: near Madden Dann and along Azote Caballo Road near Alahuela, alt- 90-100 m., Dodge l6S9S> colon: Chagrcs, Fcndlcr 265. 62. Peperomia flexinervia Yuncker, sp. nov. Herba repens terrestris carnosula fere tota glabra; foliis altcrnis, rotundato-ovatis, apice sensim acute acuminatis, e basi rotundatis vcl cordatis, peltatis ad 15 mm. procul ex marginc, ex petiolo 7-9-nerviis, conferte ciliatis, glandu- loso-punctatis; petiolo usque ad 16 cm. longo; spicis axillaribus, usque ad 20 cm. longis, singulis e caudiculo 1-2-bracteato; ovario ovoldeo, ros- ^ trato. A repent, succulent, terrestrial, essentially glabrous herb; stem creeping, 3 mm. thick when dry, rooting at tlie nodes, "densely pink-spotted," internodes 6-10 cm. or more long; leaves alter- nate, round-ovate, apex attenuately sharp-acumi- nate, base rounded to cordate, peltate within 15 mm. of the margin, rather prominently 7- to 9-nerved from the petiole, the lateral nerves strongly curving toward the apex, the innermost pair slightly coalescent with the midrib which is faintly branched upward, 7-9 cm, wide X 12-15 cm. long, glabrous, yellow-glandular-dotted, densely ciliate, drying firm, subopaque, finely pellucid-dotted; petiole up to 16 cm. or more Fig. 22. Peperomia flexinervia (113) [Vol, 37 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN long; spikes axillary, up to 20 cm. or more long, solitary or sometimes multiple (?), on a 1- or 2-bractcd stalk up to 10 cm. or more long, bracts 2 cm, or more long, narrowly lanceolate; floral bracts round-peltate; ovary ovoid, rostrate, stigma at the base of the beak; fruit not developed. Known only from the type locality. cocLf;: Las Minas, Sept. 1, 1941, AUcjj 2/^j8 (Herb. Missouri Bot. Gard,, type). This species resembles P. parrjiafa ol Costa Rica but differs in the creeping habit of growth, and type of venation. From P. manilafa it differs especially in the shape of the leaves and type of venation, 63. Peperomia ctliifera Trel. In Contr. U. S. Nat. Herb. 26:212. 1929. Pcperomia hcruatuVifolta (Vahl) A. Dletr. var. ciliifcra Trel. In Bot. Gaz. 73:145. 1922. A repent, glabrous, vine-like herb; stem 3—4 mm. thick when dry, rooting at the nodes, internodes up to 12 cm. or more long; leaves alternate, round-ovate, apex abruptly sharp-acuminate, base round, obtuse, peltate 2—3 cm. from the margin, rather densely dilate, at least when young, 6-10 cm. wide X 9-15 cm. long, obscurely about 9-nerved from the petiole, drying dark, coriaceous, opaque; petioles up to 12 cm. long; spikes solitary, closely flowered, 2 mm. thick X 6-10 cm. long, on axillary, 1-bracteate stalks; peduncle about 1 cm. long as represented; bracts round-peltate; fruit ellipsoidal, rostrate, stigma anterior at the base of the recurved beak. Mountains of central Costa Rica and adjacent Panama, CHTRiQuf: Boquete Region, Ccrro Horquetn cloud forest, on fallen trees, alt. 19 50 m., V(n! Hagcn (3 ion Hagcri 21 jj. The specimen from Panama agrees in all essential characteristics with this species. The spikes arc not fully developed and are somewhat fragmentary, but so far as can be determined they agree well with Trelease's description of those of P. ciliifcra, 64. Peperomia conserta Yuncker, sp. nov. Herba glabra repcns carnosa epiphytica; foliis alternis, ovato-elllptlcis, apicc acuminatis, basl rotundatis ex lamina 1-2 cm. peltatls, obscure palmatlm 9-nervIiSj sat conferte clliatls, miiuite pellucido-punctatis; petiolo graclH; spiels terminallbus axlllarlbusque; drupa globoso-ovoidea rostrata. A repent, spreading, epiphytic, glabrous herb; stem 3 mm. thick when dry, Internodes up to 10 cm. or more long, rooting at the nodes; leaves alternate, ovate- elliptic, apex sharp-acuminate, base rounded, obtuse, peltate 1-2 cm. from the margin, 4-5 cm. wide X 6-8 cm. long, obscurely palmately 9-nerved, margin strongly clliate, drying coriaceous, narrowly revolute, opaque, finely pellucid - dotted; petiole slender, variable in length, 4-10 cm. long; spikes terminal and axillary, 2 mm. thick X 8 cm. long, closely flowered; peduncle 10-15 mm. long; bracts small, round-peltate; fruit globose-ovoid, about 0.8 mm. long, rostrate, stigma anterior, near the base of the equilong, rather thick, granulate, curved. (114) 195 0] FLORA OF PANAMA (Piperaceac) 115 blunt beak. — Colombia: caqueta: Cordillera Oriental, in forest between Sucre and La Portada, alt. 1200-1350 m., April 5, 1940, Cmtrecasas 9120 (Nat. Herb. Colcmbia, type; fragment m U. S. Nat. Herb.). Southern Colombia and Panama. Panama: chiriqui: Bajo Chorro, Boquete District, rain forest, alt. 1800 m., Jan. 10, 193 8, Davidson 85, So far as it lias been possible to determine, the Panamanian specimen agrees in all essential characters with the one from Colombia. 65. Pepekomia maculosa (L.) Hook. Exot. Fl. 2:/;/. 92. 1825. Piper vujcuhsnm L. Sp. Pi. 1:30. 1753. Peperomia variegata Ruiz & Pav. Fl. Peruv. & Chil. 1:33, pi 52, f,2. 1798. Peperomia septiipUnervia CDC. in Jour. Bat. 4:142. 1866. Pcpcrornia monderifolia Griseb. Cat. Pi. Cub. 64. 1866, Pepcromia leridana Trel. in Ann. Mo. Bot. Card. 27:303. 1940. Pepcromia ienebracgaudens Trel. loc. cit. 305. 1940. Pcperomia Scheryi Trel. loc. cit. 28:426. 1941. A rather large, sparingly hairy to glabratc, assurgent, succulent herb; stem spotted when growing, up to 10 mm. thick and ascending to 20 or 3 cm., rooting from the nodes of the decumbent basal part; leaves alternate, peltate within 1 cm. of the margin, cUiptic-ovate, 6-11 cm. wide X 10-18 cm. long, apex abruptly short-acuminare, base rounded or truncatcly subcordate, rather faintly 7- to 11- plinerved below, or large leaves with as many more branches above the middle, drying dull and opaque, somcv/hat pubescent or glabrate; petiole 6-15 cm. long, fleshy, pubescent or glabrate; spikes terminal, solitary or paired, up to 25 cm. long; peduncle up to 5 cm. long, pubescent or glabrate; bracts round-peltate; fruit ellipsoidal, rostrate, stigma at the base of the curved beak. West Indies, northern South America, and Panama. CHiRTQUi: Finca Lerida to Boquete, alt. 1300-1700 m., Woodson, Allen & Seibert 1 176; Bajo Mona, mouth of Quebrada Chiquero, along Rio Caldera, alt. 1500-2000 m., Woodwn, Allen 6 Seibert 993; Bajo Mona, Boquete District, alt. 13 50 m., Davidson 486; forests around El Boquete, alt. 1000-1300 m., Pitiier 2984; Quebrada Vclo, alt. 1800 m., Woodson ^ Scbery 247; Boquete, Boquete District, alt. 1140 m., Davidson 727, All the specimens seen from Panama are less pubescent than those which have been examined from the West Indies and South America. P. variegata was de- scribed as glabrous and the Panamanian specimens agree in this particular. There is so much variation in this character among individual plants, however, that it does not appear advisable to base specific or varietal distinctions on it. It has been impossible to recognize P. tencbracgaudcnSy P. Scheryi, and P. leridana as distinct species, although they differ somewhat in the texture of the dried plants, leaf size, length of spikes, etc. None of these characters, however, appear suf- ficiently distinctive to warrant specific or varietal recognition. Field studies and mass collections are required to provide evidence for proper segregation of thes questionable species, if such is indicated. All specimens seen from Panama are from an altitude of 1000-2 000 meters in the mountains of Chiriqui. n (115) 116 [Vol. 37 MISSOURI 66. Peperomia perglandulosa Yuncker, sp. nov. Hcrba cpiphytica repens carnosa; caule conferee rubro-villoso; foliis altcrnls dcltoldco-ovatis, apice rotundatis, basi truncatis, plcrumquc subpeltatis, palmatlm 5-nerviis5 snpra glabratis subtus pubescentibus saltern ad costam confertc ciliatis utrinciue conferte rubro-glanduloso-punctatis; petiolo 1.5-4 cm. longo confcrtt rubro-villoso, dissite rubro-glanduloso-punctato; spicis oppositifoliis; pcdunculo 2.5 cm. longo conferte rubro-villoso ad medium bracteato; ovario ellipsoidco acute rostrato. A repent, succulent, epiphytic herb; stem 2 mm. thick, densely red-villous, rooting at the nodes, internodes 2-5 cm. or more long; leaves alternate, deltoid- ovate, apex rounded, base truncate, mostly subpeltate with the petiole attached 1-2 mm. from the margin, palmately 5-nervcd, the laterals slender and obscure, glabrescent above, pubescent beneath, at least along the midrib, the margin densch ciliate with a zone of submarginal hairs on the upper surface^ L5-3 cm. wide X 1.5-3 cm. long, drying rather thick and opaque, densely and conspicuously rcd- glandular-dotted on both sides, the glands comparatively large; petiole 1.5-4 cm. long, densely red-villous, sparsely red-glandular; spikes leaf-opposed, 2 mm. thick X 7 cm. long, densely flowered; peduncle 2.5 cm. long, densely red-hairy, with scattered red glands, bracteate near the middle; floral bracts round-peltate, con- spicuously and densely red-glandular; ovary ellipsoidal, red-glandular, sharply rostrate, stigma anterior at the base of the beak; fruit not present. Known only from the type locality. cocLi: EI Valle de Anton, crest of Cerro Pajlta, alt. 1100 m., Sept. 28, 1946, Allen 375^ (Herb. Missouri Bot. GarJ., type). The densely red-villous plants, and red-glandular leaves, bracts, etc. are dis- tinctive features of this species. VII. KEY TO SPECIES WITH NUMEROUS SPIKES ARRANGED IN A COMPOUND. PANICULATE INFLORESCENCE. (Subgenus Panicularia Miq.). a. Spikes scarcely more than 1 cm. long; fruit subcylindrical with oblique apex; inflorescence stalk microscopically hirtellous 67. P. pernambucensis a. Spikes mostly 3 cm. or more long; fruit not as above. b. Leaves sessile or subsessile 68. P. duricaulis bb. Leaves obviously pctiolate. c. Spikes mostly less than 10 in each panicle. d. Leaves obovate, mostly 10 cm. or more wide 69. P. Williamsii dd. Leaves clliptlc-oblanceolatc, scarcely 5 cm. wide. e. Stems, leaves, etc. villous 70. P. W'udllii cc. Stems glabrous 5^,b. P. obtusifolia van longibractf.ata cc. Spikes mostly 12 or more in each panicle; leaves elliptic, elliptic- oblong, or oblanccolate, mostly 6-8 cm. wide. d. Leaves ghibrous; stem only a few cm. long, scarcely erect 71. P. mameiana dd. Leaves pubescent, at least beneath. e. Plants rhizomatous; leaves villous along the nerves beneath, glabrous above 70. P. Wldelii ec. Plants with erect stems up to 100 cm. tall; leaves silky-ap- pressed-villous on both sides 72. P. substriaTa (116) 1950] FLORA OF PANAMA (Piperaccac) 117 67. Peperomia pernambucensis Miq. in Hook. Lond. Jour. Bot. 4:420. 1845. Pcpcromia paniculata Rcgcl, in Bull. Soc. Nat. Moscou 31:542, pL J. 1858. Peperomia loug'tfoUa CDC. in DC. Prodr. 16^:405. 1869. Peperomia multiplex Leprleur ex CDC. loc. cit. 406. 1869, as synonym. Peperomia lopezensis Trel. in Ann. Mo. Bot. Card. 27:303. 1940. A simple, short-stemmed, succulent, terrestrial, or epiphytic herb; stem 3-5 mm. or more thick, ascending to 5-10 cm. from a briefly decumbent, rooting base, intcrnodcs very short, glabrous; leaves alternate, elliptic^ elliptic-oblong, or ob- lanceolate, 4—6 or sometimes up to 12 cm. wide X 15—3 cm, long, apex sharp- acuminate, base cuneate, decurrent, pinnately nerved with 5-8 branches on each side of the prominent midrib, glabrous on both sides, minutely hirtellous at the tip, usually drying rather chartaceous and subopaque; petiole 3-5 cm. long; spikes about 2 mm. thick X 10—12 mm. long, vcrtlcillate in a loose, terminal or axillary, spike-like paniculate inflorescence up to some 8 cm. long, on a common stalk 4-8 cm. long, the main stalk of the panicle somewhat hirtellous or puberulent; peduncles about 3 mm. long; bracts round-peltate; fruit about 1 mm. long, subcylindrical, slightly obliquely truncate at the apex, stigma central. Northern South America and extending into southern Central America. BOCAS DEL TORo: Fish Crcck Mountains, vicinity of Chiriqui Lagoon, von Wedel 22jjy 22S6, CANAL zone: Quebrada Lopez, Allen 2Tl8. The leaves on the specimens seen from Panama are somewhat more elongated than most of those seen from South America and consequently do not appear as abruptly acuminate. This character which, so far as has been noted, is not sup- ported by other differences, docs not appear to be sufficient to warrant the main- tenance of P. lopczensis as a distinct species. 68. Peperomia duricaulis Trel. in Ann. Mo. Bot. Gard. 27:3 02. 1940. An erect or assurgent, glabrous, leafy, terrestrial or epiphytic, alpine herb; stem 5 mm. thick downward when dry, ascending to 20—3 cm., internodes 1-3 cm. long, nodes rather prominent; leaves alternate, elliptic-oblanccolate, apex sharp- acuminate, minutely hirtellous, base cuneate-decurrent, 3—4.5 cm. wide X 10—13 cm. long, pinnately nerved^ the nerves 5—6 on each side of the prominent midrib, drying submembranous, translucent, faintly but densely pellucid-dotted; petiole very short, broadly winged to the base by the decurrent blade, the wings clasping the stem; spikes 1 mm. thick X 3-5 cm. long, in umbels of usually 3—5 spikes in the axil of a lanceolate bract, the umbels alternately arranged in an open, divaricate, paniculate, axillary Inflorescence eventually up to 15 cm. or more long, the basal part to the first umbel up to 7 cm. long in anthesis; peduncle of individual spike 5-10 mm. long; floral bracts round-peltate; fruit ellipsoidal, rostrate, stigma at the base of the elongate beak. Southwestern Panama. CHiRTQuf: trail from Cerro Punta to headwaters of Rio Caldera, alt. 2250—2500 m., Allen 1441; Bajo Chorro, Boquete District, alt. 1800 m., Daiidsoj? 2gg, (117) 118 [Vol, Z7 ANNALS OF THE MISSOURI BOTANICAL GARDEN 69. Peperomia Williamsii Trel. in Contr, U. S. Nat. Herb. 26:48. 1927. Veperomia flavhpica Trel. loc. cit. 1927. A glabrous, succulent, subsimple, arborlcolous herb; stem up to 1 cm. thick when dry, rather short, internodes 2-3 cm, long; leaves alternate, elliptic-obovate, apex rounded, obtuse, or abruptly acute, commonly emarginulate, base cuncatc, dccurrent, 9-11 cm. wide X 15-20 cm. or more long, pinnately nerved to above the middle, the lateral nerves very slender, about 6 on each side, drying thick, glossy, coriaceous, opaque, subrevolute; petioles stout, 5-12 cm. long, winged by the decurrent blade; spikes 2 mm. thick X 10-15 cm. long, closely flowered, several, solitary, or 2 or 3 from the nodes of a zig-zag, elongated stalk-Kkc con- tinuation of the stem, inflorescence up to 15 cm. or more long; peduncles of individual spikes 2-4 cm. long; rachis with anastomosing ridges; bracts round- peltate; fruit ellipsoidal, about 1 mm. long, rostrate, stigma at the base of an awl-like, equilong beak with hooked apex. Panama. COCLE: Bismarck, above Pcnonome, alt. 600-900 m., Williams 43$- canal zone: Las Cascadas Plantation, near Summit, Standlcy 25J2g; drowned forest near Vigia and San Juan on Rio Pequeni, alt. 66 m., Dodge, Steyermark ^ Allen l6j2j — leaves 5-7 X 12-15 cm. PANAMA: Rio Tapia, Standley 26ig6, The large, succulent, obovate leaves with stout petioles and paniculatcly clustered spikes are distinctive features of this species. No specific diflferences have been noted between this species and P. flavhpica. 70. Peperomia Wedelii Yuncker, sp. nov. Herba subrepens rhizomatosa epiphytica; caulc dissite villoso, indumento ad 1 mm. longo, intcrnodiis brcvibus; foliis alternis ob- lanceolato-cllipticis, apice breviter acumi- natis, basi cuneatis, plnnatim venosis, venis utrinque ca. 5, supra glabris subtus ad venas dissite villosis; petiolo 1-3,5 cm. longo, dissite villoso; spicis in panlculam tcrminalem dispositis; pedunculo ca. 1 cm. longo glabro; drupa angustata erccta oblongo-cylindrica rostrata. A subrcpenf , rhizomatous, e p I p h v 1 1 c herb; stem 2 mm. thick when dry, loosely villous, hairs up to 1 mm. long, internodes short; leaves alternate, oblanceolate-elliptic, apex short-acuniinate, base cuncate-dccur- rent, 3-5 cm. wide X 10-15 cm. long, pin- nately nerved, the hueral nerves about 5 on each side, glabrous above, loosely villous Fig. 23. Peperomia WcJclii al ong the nerves beneath, drying rather (118) 1950] FLORA OF PANAMA (Piperaccac) 119 papery, translucent; petiole 1-3.5 cm. long, loosely villous, shallowly grooved above; spikes 1 mm. thick X 3-6 cm. long, single or 2-3 in alternate clusters in an elongated terminal panicle, the lowermost spikes about 7 cm. above the base of the loosely villous stalk; peduncles of individual spikes about 1 cm. long, glabrous; floral bracts minute, round-peltate; fruit slender, erect, oblong-cyHndrical, rostrate, stigma anterior at the base of the short, flexible beak. Known only from the type locality. BOCAS DEL TORO: Fish Creek, vicinity of Chiriqui Lagoon, April 9, 1941, H. ton Wedel 200 j (Herb. Missouri Bot. Gard., type). This species appears to be rather closely related to P. cahifolia of Costa Rica. It differs, however, in the pubescence of the stem, leaves, etc., fewer lateral leaf nerves, longer petioles and spikes, etc. 71, Peperomia mameiana CDC. in Candollea 1:290, 396. 1923, as P. mawaiana CDC. in Candollea 2:188. 1925. A glabrous, rather succulent, rhizomatous herb, on rocks and trees; stem 4-8 mm. thick when dry, rooting at the nodes, internodes 1-2 cm. long; leaves alternate, elliptic, lance-eUiptic, or suboblanceolate, apex acute, base cuneate-decurrent, 4-10 cm. wide X 15-25 cm. long, obscurely pinnately nerved, the nerves 6-7 on each side of the stout midrib which continues to the very apex, drying rather glossy, coriaceous, opaque, narrowly revolute; petiole stout, up to 10 cm. long, channelled and winged upward by the decurrent blade; spikes 1-1.5 mm. thick X 6-12 cm. long, densely flowered, loosely paniculate on long, stout, axillary or terminal stalks, the entire inflorescence up to 30 cm. or more long; peduncles 1-5 cm. long; rachis with anastomosing ridges; bracts round-peltate; fruit scarcely 1 mm. long, ellip- soidal, rostrate, stigma anterior at the base of the curved beak. Central Panama. CANAL zone: Mamei Hill, alt. 20-90 m., Vlttier 380S; vicinity of Las Cruces, alt. 26-40 m., Seibert 571 ; Barro Colorado Island, Kenoyer 30J; near Salamanca Hydrographic Station on the gorge of the Rio Pcqueni, alt. 70-80 m., DoJge, Sieyermark ^ Allen l6j2ja; Experimental Gardens, Lindsay 402a, 72, Peperomia substriata CDC. in Candollea 1:291. 1923. A moderately large herb; stem somewhat silky-pubescent, erect or scandent, rooting at the nodes, with erect branches, up to 5 mm. thick when dry, internodes up to 7 cm, or more long; leaves alternate, elliptic, apex short-acuminate, base acute, 2-4 cm. wide X 4.5-8 cm. long or up to 6.5 X 16 cm., pinnately nerved nearly throughout, the lateral nerves 6-8 on each side, appressed silky-villous on both sides, drying rather thick, opaque; petiole up to 6 cm. long, moderately appressed silky-villous, channelled and narrowly winged; spikes 2 mm. thick X 6-10 cm. long, closely flowered, several on a bracted, silky-villous stalk 3 cm. or more long; bracts large, foliar, elliptic-obovate, sparingly silky; peduncle of indi- (119) [VuL. 37, 1950] 120 ANNALS OF THE MISSOURI BOTANICAL GARDEN vidual spikes 5-10 mm. long; floral bracts round-peltate, glandular-dotted; fruit ellipsoidal, about 1 mm. long, rostrate, stigma anterior near the base of the beak. Costa Rica and adjacent Panama. BOCAS DEL TORO: Robalo Trail, northern slopes of Cerro Horqueta, alt. 1800-2100 m., Allen 4950. CHiRiQui: Bajo Chorro, Boqucte District, rain forest, a!t. 1800 ni., Duiihon 217. (120) STAFF OF THE MISSC BOTANICAL GABDEI >( Director Georgc T. Mooke Herm^xn von Schrenk, _ „ : 3gist ROBE^.T W, SCHERY, Jesse M. Greenman, Curator £m«ntiis of the Herbarium Gc5TAV A> L. Mhhlquist, Research Horticulturist Carroli. W. Dodce, A ■-'^'-:.nt Curator of iierb-nriuin the Edgar Anderson, Geneticist George B. Vak Sckaacx, Haacrary Curat or of G ROBEF.T E. '^^. '^'^O^-, Jr, Curator of the Herbarium JixiAN A. Steyermasj:,^ Hctiorar J Rese^rcli Associate Henky N. Andrews, J'::.leo;:. ■ .. ^ J X, Librarian, and ,Edi of P:.:b!ic.atsons GE"RAtD UlRIO, TS. \.; , . - -iS ■\ !.- J"j V BOARD OF TRUSTE IL OF THE MISSOURI EOTA2>FICAL GARDEN Presideni Daniei- S* CAixm Second Vice EU<7EKH L. Ray Car-te^ n"r,v Hrr^- >». T EthaH a. H* Shef^ey E:X-0F.riCJ0 MEMBERS ■S^ieace of St, L^tis ■ basFH M. Daest *i« Cht of VlM-iAM Sc:Hi;J,iaO-^l^K. Pre^ideni m. tfee 'B^ 'ALt*. #\ Orii Eclnrstl'^ of St. t'crttm'y ^ ^ -^ i ^ ■.J ' ^■_ V Volume XXXVil Number 2 Anna of the Missouri MAY, 1950 Flora 0| Panama. Part V, Fas. 2 CResedaceae to Ee .osae,'lB 1 « Woodsoa, |r.,-Bobcft W: S assd- 1.21 ^- ^ , I 1 i^s'MJEftS-sii OF T^Mm^ r n^ .it>-£'f ^^ 'it^ ■*■'..%. ^'Wmmih ^.'i^rt. :-^«te »t #a^i^^Na^^: raam%: ^' .^ ^■ ~n ,\f^6^i.-m 'd^J-.Z^LL. "■ ,_, = ,- -. -^ ^ rr ^" Anna of the S Missouri Botanical Garden Qmn€s:ly Journal containinj? Scientific Co Missouri Botamcai Garden and the Henrv Shsw Sclioo: Uaiversitf otaay of affiliatsoa with tKe Missouri Botanical Informaikm Tfe Akhais OT -am Mmoixmi Botakical Garden appean four times dmnag the calendar year: February, May. September, and Movtmhtr. Four ssimibers ajnstatute a TCHimw; Sabtcriprioa Price Single NimslKJts . ■**-fr $10.00 per volume 2.50 eacH ■ ^ CjSBtents of pieriocs ,ia«es of tlie Asrw ats of tke Missooti OTAHICAI. Agrkulmral Indfcx* pufrliAsd by tiie K W. Wikoa I - ^^;- -T \' ^ F. ^ -ti-C. ■ -^J^.. -i ■ .-,"-/ -"^ . - rf . ' rf*wirj^"L.'^ . h ■- - L ^ ' -Tm ir^ _ FLORA OF PANAMA Part V. Fascicle 2 RESEDACEAE A small family of annual or perennial herbs with alternate or fasciculate leaves and spicate racemes of small, asymmetrical flowers with white or yellowish, cleft petals. Wo being native to the Mediterranean area. The genus Reseda (''mignonette", reseda) has been introduced into Panama and is cultivated in gardens in the Boquete region. MORINGACEAE pec 'i World resinous or gummy branches, and whitish bark. The leaves are large, alternate, and twice or thrice pinnate. The inflorescence is paniculate and axillary with rather mediocre but fragrant, white or reddish flowers. The fruit is a rather large and elongate, 3 -angled capsule dehiscent by three valves; the seeds are 3 -winged and oily. The vernacular name of M. oleifera is "horseradish tree" or marango, a name arising from the flavor and odor of the roots. The oil from the seeds is comestible or may be used as a substitute for ben oil which is produced as a lubricant for ■pf CRASSULACEAE Usually fleshy or succulent herbs or subshrubs, with thick stems and alternate (infrequently opposite), many-ranked leaves. Leaves usually thick and fleshy, mostly simple and entire or crenulate, infrequently pinnately compound, in many species living for a considerable time after being removed from the plant and in some cases capable of producing plantlets from the marginal indentations. In- florescence frequently cymose, less often racemose or paniculate. Flowers typically coarse and stiff, perfect, regular, polypetalous or sympetalous; calyx 4- to 5-parted, the sepals free or united; petals 4-5, free or united, mostly persistent; stamens in 1 or 2 cycles, each cycle isomerous with the petals; carpels as many as the petals or sepals, essentially free or united at the base, the ovules usually numerous from norm Fruit follicular from the individual carpels; seeds small, punctulate. Issued June 24, 1950. (121) (107) [Vol. 37 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN A family represented in almost all parts of the world, especially in Africa; best represented in warmer latitudes, not uncommonly under subalpine conditions. A. Berger (in Engl & Prantl, Nat, Pflanzcnfam, 18a:352-483. 1930) probably has given the most acceptable recent treatment of the family. a. Tall herbs; leaves opposite, pctiolatc, often pinnate; flowers 4-mcrous, the calyx tubular, inflated 1. Bryophyllum aa. Small herbs; leaves spiral, sessile, simple; flowers 5-mcrous, the sepals essentially free 2. EcHEVERiA 1. BRYOPHYLLUM Salisb. Bryophyllum SaHsb. Parad. Londim pi. J. 1805. Physocalycium (?) Vest, in Flora, III, 2:409. 1820. Crassoui'ia Commcrs. ex DC. Prodr. 3:395. 1828. Kalanchoe R. Hamct, in Bull. Herb. Boiss. II, 7:870. 1907, in part. Erect, succulent, perennial herbs or subsbrubs, usually glabrous. Leaves mostly simple but sometimes pinnatcly compound, opposite, thick and fleshy. Inflorescence usually a terminal panicle. Flowers perfect, regular, usually nodding, mostly 4-merous; sepals 4; usually united into an Inflated tube; petals 4, sympetalous, somewhat exceeding the calyx, usually constricted above the ovary, the lobes shorter than the tube; stamens inserted on the lower corolla tube, the filaments about equalling the corolla; carpels partly united or distinct; ovules numerous; disc bearing 4 glands or scales. Fruit normally of 4 follicles. The genus is distinguished from the New World genera of the family in pos- sessing the large, inflated calyx and opposite, sometimes compound leaves. It resembles the African and Asian genera Kalanchoe and Kitchingia^ differing from these in the insertion of the stamens toward the base of the corolla tube in the large, nodding flowers. A single species is naturalized widely in Panama, but is not frequently collected. 1. Bryophyllum pinnatum (Lam.) S. Kurz, in Jour. Asiatic Soc. Bengal 40^:52. 1871. Cotyledon plnnata Lam. Encycl. Meth. 2:141. 1786. Calatichoe pinnata Pers. Syn. PL 1:446. 1805. Bryophyllum calycirtum Salisb. Parad. Londin. pL J. 1805. Perennial, Httle-branched herb as much as 1 m. tall or taller, the stems fleshy, glabrous, lustrous, the roots diffuse. Leaves opposite, simple or less frequently 3- to 5 -pinnate (usually both simple and pinnate leaves on the same plant), in compound leaves the terminal leaflet the largest; blade of leaf or leaflets oblong, rounded at base and apex; coarsely crcnate, glabrous, as much as 13 cm. long and 7 cm. broad, fleshy and obscurely nerved, capable of producing plantlets from the indentations of the leaf margin; petiole conspicuous, commonly 3-4 cm. long on older leaves, fleshy, glabrous, widened toward the base, exstipulate but with de- nt intcrpetiolar lines on the subtending intcrnode. Inflorescence a glabrous, terminal, thrice-compound panicle as much as 5 dm. long, the bracts and bractlets (108) 1950] FLORA OF PANAMA (Crasstilaceae) 123 minute and caducous. Flowers large and conspicuous, with pedicels mostly about 1 cm. long; calyx broadly tubular, inflated, usually about 3 cm, long and 1 cm. broad, 4-lobed for one-fourtb or one-fifth its length, greenish often tinged reddish; corolla tubular, 4—5 cm. long, mostly enclosed by the calyx, markedly constricted just above the ovary and thence ventricose, 4-lobed, the lobes triangular and acuminate, glabrous, reddish; stamens 8, inserted at the base of the corolla tube, the filaments almost as long as the corolla. Native of tropical Africa and Madagascar; introduced and naturalized in the tropics throughout the world. CHiRiQuf: Boqucte, Woodson ^ Schery JSQ, The species Is quite common In the American tropics, both as a cultivated plant and as an escape. The rather bizarre flowers are not unattractive and the peculiar ability to produce plantlets at the notches of the leaves enhances its interest, earning the popular names "life plant" and ho]a del aire, Standley reports (Fl, Panama Canal Zone, p. 187. 1928) : "Among the West Indians of the zone [Canal Zone] a favorite method of testing the fidelity of a sweetheart is to place a leaf of this plant above one's door. If a new plant is produced at each notch, her faithfulness can not be questioned." 2. ECHEVERIA DC. EcHEVERiA DC. Prodr. 3:401. 1828. Caulescent or scarcely stemmed perennial herbs, the leaves frequently borne in a basal rosette, fleshy and succulent. Leaves spirally arranged, sessile, entire, usually glabrous and somewhat glaucous. Inflorescence usually a lax spike or raceme, the flowers arising singly or in pairs in the axils of the reduced stem-leaves. Flowers fleshy, variously colored; sepals 5, more or less free, equal or unequal, frequently erect, mostly greenish; petals 5, usually exceeding the sepals, free or somewhat united at the base, frequently erect and angled; stamens 10, the outer cycle mostly Inserted at the base of the petals; carpels 5, united at the extreme base, tapered into slender styles. Fruit follicular. An Interesting and attractive genus much used for ornamentals because of the rosettes of thick, handsome foliage. Distinctive characters frequently are lost in making herbarium sheets of such succulents, which may account for the widespread confusion of species limits. Fiortlculturallsts have been particularly concerned with the difficulty in naming specimens accurately, and such students of the group as Britton and Rose (N. Am. Fl. 22:13-26. 1905) and von Poellnitz (Fedde Rep. Spec. Nov. 39:193-270. 1936) apparently have overdone the splitting of species. The genus occurs in the southern United States to middle South America, cen- tering in Mexico. A single species is known to occur In Panama. 1. EcHEVERiA AUSTRALis Rosc, in BuU. N. Y. Bot. Gard. 3:6. 1903< Caulescent, little-branched, succulent herb to a few decimeters tall, glabrous and more or less glaucous. Leaves sessile, obovate or spatulate, up to 7 cm. long im) [Vol. 37 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN and almost 3 cm, broad in the larger, basal leaves, progressively smaller along the flowering stems, rounded or abruptly acute at the tip, broadly cuneate at the base, entire. Inflorescence terminal, appearing racemose by the insertion of flowers In the axils of reduced, frequently caducous leaves; pedicels 2-5 mm. long. Flowers moderate, pink; sepals 5, narrowly or broadly ovate, rather unequal, 5-10 mm. long; petals 5, lanceolate, about 1 cm. long, rather thick and rigid, mucronate- acute, essentially free; stamens 10, almost equalling the petals; filaments slender, glabrous; anthers oblong, about 1 mm. long. Costa Rica and Panama. CHiRiQuf: Rio Chirlqui Viejo valley, P. White 39- \ PODOSTEMONACEAE By p. van ROYEN Small, usually matted herbs growing in rapids and waterfalls, flowering when the water subsides seasonally, very variable in size, the smaller ones thallold and closely adhering to the supporting rocks, the larger ones with distinct stems often arising in pairs from long and thin roots, in other Instances rootless; sterile plants often larger and coarser than the fertile ones. Leaves alternate, di- or tristlchous, in the stcmlcss species sometimes united at the base and springing In 2 rows from an irregular base, entire or much divided, sometimes with bundles of minute threads on one or both sides; stipule 1, a sheath and a ligula sometimes present. Flowers solitary or fascicled between the leaf-bases or at the ends of short branches or In distinct Inflorescences, hermaphrodite, actlnomorphic or zvgomorphic, the younger ones enveloped by a thin spathe or by a few leaves. Tcpals 2 to many, n distinct or reduced to minute scales, free or united. Stamens 1 to many, alternate with the tepals In a complete or Incomplete v/horl, free or united to the tepals; anthers sagittate, 4-cellcd, dehiscing by 2 longitudinal slits, introrse or extrorse, dorsl- or baslfixed; pollen 1- or 2-celled. Ovary superior, sometimes oblique or nearly perpendicular to the (frequently) distally enlarged pedicel, 2- or 3-, or sometimes 1 -celled; carpels equal or unequal, ribbed or smooth without, the placenta axilc, fleshy, with 2 to numerous anatropous ovules; ovules with 2 in- teguments; styles 2-3, free or cohering, each with a distinct stigma. Fruit de- hiscing septlfragal with 2 or 3 valves, In fruits with unequal cells the smallest valve sometimes soon falling oflf; seeds 2 to many, w^ithout endosperm. About 200 species distributed in the tropics throughout the world, ascending the southern Appalachian Mountrins in the southeastern United States, a. Flower buds enclosed between a few leaves; perianth distinct; ovary 3-cellcd; leaves entire 1. Tristicha an. Flower buds enclosed in a membranaceous spathe; periantli reduced to minute scales; ovary 2-ce!led; leaves much divided...... 2. Maratjikum (110) 1950] FLORA OF PANAMA (Podosfcmonaceac) 125 1. TRISTICHA Du Petit Thouars Tristiciia Du Petit Thou. Gen. Nov. Madagasc. 2. 1806. Dufourea Bory, ex Willd. Sp. PI. 5:55. 1810. Philocrena Bong, in Mem. Acad. Sci. St. Petersb. VI, 3-:80-82. 1835. Potamobryon Liebm. in Forhandl. Skand. Naturf. Kjoeb. 5:512. 1847. Moss-like herbs widely diflfering in shape and size, much branched, forming dense mats on rocks; stems thin, terete. Leaves tristichous, the rows in the older plants sometimes indistinct, entire, membranaceous, nerved or unnerved, the apex obtuse or acute, often divided into 2 or 3 parts in older leaves. Flowers terminal or on short side-branches, enclosed between 2-3 distinctly larger leaves when young; perianth well-developed, membranaceous, 3 -partite, marcescent; stamen 1, the filament slender, the anther ovate, introrse, with elongate connective, the pollen globose; ovary 3 -celled, with 3 equal carpels, ovoid to subglobose, rounded or attenuate at the base, the placentas swollen, dehiscing with 3 equal valves each with 3 ribs; seeds numerous. Tropical and subtropical America, Africa, and Asia. Fig. 54. Tristicha trifaria 1. Tristicha trifaria (Willd.) Tul. in Arch. Mus. Hist. Nat. 6:179-189. 1852; Warming, Fam. Podost. 6:28-30, /. 24. 1901. Dufourea trifaria Willd. Sp. PI. 5:55. 1810; in Mag. Ges. Naturf. Fr. Berlin 5:63-64. 1811. Dufourea bypuoidcs St. Hil. in Mem. Mus. Paris 10:472. 1823. Tristicha hypnoiJcs (St. Hil.) Spreng. Syst. Veg. 4^:10. 1827; Cario, in Bot. Zeit. 39:25, tab. 1-24. 1881. Dufourea Boryi A. Rich, in Chev. Diet. Drogues 5:636. 1829. Philocrena pusilla Bong, in Mem. Acad. Sci. St. Pttcrsb. VI, 3^:80, tab. 6, 1835. (Ill) [Vol. 37 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN Potamohryon coucbnium Licbm. in Forhandl. Skand. Naturf. Kjocb. 5:513. 1847. Pofamobryon laxum Llebm. loc. cic, 514. 1847. Potamobryon patithim Liebm. loc. cit. 515. 1847. Trhticbta c(»icinna (Liebm.) I. M. Johnston, in Sargentia 8:130. 1949. Small herbs with shoots forming dense mats attached to much-branched roots; stems terete, 0.2-9.0 cm. long. Leaves tristichous, the rows at first regularly alternating with one another, the older leaves in irregular whorls of 3-6; one row always on the dorsal side of the stem, these ovate, 0.5 mm. wide to 1 mm. long; the otKer two rows lateral and at the same level, obliquely inserted, these spathulate, to 1 mm. wide, to 3 mm, long; all leaves definitely nerved, entire, obtuse, sessile, sometimes all alike or the older with 2-4 acute apices; fertile branches with 2-3 membranaceous leaves 2.0-2.5 mm. long, which enclose the young flower. Perianth 3 -fid or -partite, the lobes obtuse, nerved, 1-2 mm. long; stamen 1.5-2.5 mm. long, the filament spathulate, distinctly nerved, 1-2 mm. long, the anther sagittate or ovate, truncate, slightly cmarginate at the base, about 0.8 mm. long; pollen globose, 16-17 fi In diameter, yellowish; ovary ellipsoid, roundish or attenuate at the base, 0.5-1.0 mm. long, about 0.8 mm. In diameter; styles 3, linear, about 0.5 mm. long, often papillate; pedicel 3-15 mm. long, terete. Widespread in Central and South America, Cuba, and Africa. CANAL zone: Rio Pedro Miguel, StauJley 2Qpjj, ^9034; Rio Paraiso, Sfamllcy 2gS/S; Quebrada Ancha, Steyermark & Allen T^^IJO; Las Cascadas, near Summit, Dodge & Hunter 5. ».; Rio Boqucron, Steyermark & Allen //^i/; Rio Indio, Dodge & Allen 17400. 2. MARATHRUM Humb. & Bonpl. Maiiatiirum Humb. & Bonpl. PL Aequinox. 1:40-41, tab. II. 1808. Lack Mart. Nov. Gen. & Sp. 1:6. 1824. Small to medium-sized stcmless herbs known to be attached to roots in only a few species, the sterile plants often larger and coarser than the fertile, with an irregular, sometimes branched, adhesive base. Leaves either distichous or inserted along the margin of the thallold base, repeatedly pinnate with all the pinnae alter- nate or slightly opposite, or entire with a few lobes along the margin; petiole sometimes fleshy, often dilated at the base. Flowers at first enclosed in a persistent, membranaceous spathe which splits irregularly at the tip, 1 to many, solitary or fascicled between the leaf-bases; pedicel scarcely enlarged at the tip in some species, In others abruptly enlarged; tepals 3-25 in a complete or incomplete whorl, in- conspicuous, squamiform to filiform, Inserted on the margin of the receptacle in species with enlarged pedicels, but sometimes inserted at different levels In species with normal or scarcely enlarged pedicels; stamens 2-25, in a complete or incom- plete whorl (the latter unknown in Panamanian species), rarely united at tKe base, the filaments lanceolate, 3 -angled at the base, sometimes branched, nerved, the anthers sagittate, introrse; pollen ellipsoidal to subglobose, 3-sulcate; ovary 2- cclled, ellipsoidal, attenuate at the base, with 2 equal carpels, 8-ribbed; placenta (112) 1950] FLORA OF PANAMA (Podostemouaceae) 127 of the same shape as the ovary, with many ovules; styles 2, fiUform or cylindric, cohering at the base or very rarely free, often emarginate. Fruit with 2 equal, persistent, 5-ribbed valves. Central America, Antilles, northwestern South America, a. Pedicels not thickened at the tip; leaves 3-5 times pinnate. b. Stamens one-third to one-half as long as the ovary; pollen ellipsoidal, 17 y 14 fi 1. M. STENOCARPUM bb. Stamens about as long as the ovary. c. Tepals 0.5-1.0 mm. long; styles 2-3 mm. long; pollen ellipsoid, 16 X 13 H- ^' ^^* ^XYCARPUM cc. Tepals 0.5 mm. long or shorter; styles 0.5-2.0 mm, long. d. Leaves shorter than 3 cm., usually some dilated and cuneately fasciate-fiabelliform; pollen subglobosc, 15 X 14 ft -3. M. pusillum dd. Leaves 0.5-20 cm. long, of essentially similar form throughout. e. Ultimate leaf-divisions filiform or nearly so. f. Leaf rachis not winged, transparent when young; pollen narrowly ellipsoid, 20 X H M; styles 0.5-1.0 mm. long 4. M. LEPTOPHYLLUM ff. Leaf rachis distinctly winged, not transparent when young; pollen ovoid, 16 X 14 fx; styles 1.0-2.0 mm. long .5. M. indifferens ec. Ultimate leaf -divisions usually round to spathulate or tri- angular, not filiform, f. Leaf rachis narrowly winged; ovary 2.5-3.0 mm. long; pollen subglobosc, 15 X 14 M ^' M. cheiriferum ff. Leaf rachis not winged; ovary 3.5-4.0 mm. long; pollen ellipsoid, 17 X 14 M 7. M. Allenii aa. Pedicels thickened at the tip; leaves repeatedly bifurcate 8. M. foeniculaceum Fig. 5 5. Marathrum stcnocarpum 1. Marathrum stenocarpum (Wedd.) v, Royen, comb. nov. Marathrum Schiedeanuyn (Cham.) Tul. var. stenocarpum Weddell, in DC. Prodr. 17:54. 1873. Small to medium-sized heAs; base cuneiform to more or less discoid, 1-2 cm. wide, 1-3 cm. high. Leaves 3 times pinnate, 3-20 cm. long, all the pinnae ascend- ing at an angle of about 45°; primary pinnae 0.5-5.0 cm. long, 2-15 cm. apart; secondary pinnae 1-7 mm. long, 2-5 mm. apart; tertiary pinnae repeatedly forked, (113) 128 [Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 56. Muruthrum oxycarpum the ultimate divisions to 1 mm. long and 0.3 mm. wide, spathulate, acute, mostly nerveless; petiole 0.5-8.0 cm. long, 1-2 mm. wide, flattened, membranaceously dilated at the base and with a more or less distinct, obtuse, membranaceous Hgula LO-1.5 mm. long. Flowers few, solitary; pedicel 1-4 cm. long; young spathe clavate, acute, 1.0-L25 mm. long, the older infundibuliform, 3-5 mm. long; tepals 9, lanceolate, acute, nearly 1 mm. long; stamens 9, 1.5-2.0 mm. long, the filaments lanceolate, about 1.5 mm. long, the anthers obtuse, nearly 1 mm. long; pollen ellipsoidal, 17 X H /x, the lobes tapering toward one of the poles; ovary narrowly ellipsoid, acute, attenuate at the base, with 6 indistinct ai distinct ribs marking the sutures, 3.5-4.0 mm. long, the styles 2, cyHndricnl, 3- edgcd at the base, the narrow wings passing Into the midribs of the valves, papillate at the apex only, about L5 mm. long. Fruit with 2 distinctly 5-ribbed valves. Panama and Colombia. d 2 more (114) 1950] FLORA OF PANAMA (Podostemonaceae) 129 PANAMA: Juan Diaz River, Killip Jjpo. canal zone: Rio Boqueron, Steyermark & Allen 17255. 2. Marathrum oxycarpum Tul. in Ann. Sci. Nat. Ill, 11:94. 1849. Medium-sized herbs; base 2.5-30.0 mm. long, 2-8 mm. wide. Leaves 3 times pinnate, 3-50 cm. long, with all the pinnae alternate and ascending at an angle of 30-60°; primary pinnae 0.3-9.0 cm, long, 0.2-3.0 cm. apart; secondary pinnae 0.1-2.5 cm. long, 0.5-7.0 mm. apart, sometimes subopposlte; tertiary pinnae 3-10 mm, long, 1-2 mm. apart, alternately pinnate with forked lobes, the ultimate divisions spathulate, acute, nerveless, rarely distinctly nerved, to 3 mm. long and 0.3 mm. wide; petiole flattened, slightly ribbed, membranaceously widened, 0.3-12.0 cm. long, 0.5-4.0 mm. wide; rachis straight, with an obtuse membrana- ceous llgule about 2 mm. long. Flowers soHtary; pedicel 4-9 cm. long; young spathe clavate, acute, 2-5 mm. long, the older Infundibuliform, 8-10 mm. long; tepals 8-10, triangular, acute, 0.5-1.0 mm. long; stamens 8-10, 5-6 mm. long, the filaments lanceolate, membranaceous, somewhat scaphiform at the base, 4.0-4.5 mm. long, the anthers obtuse, 1-2 mm. long; pollen ellipsoidal, 16 X 13 /x, the lobes tapering toward one of the poles; ovary ellipsoid, acute, base attenuate, flat- tened, 3.5-5.0 mm. long, 1.5-2.0 mm. wide, ribs 8, distinctly raised, the styles filiform, with widened sometimes cmarglnate apex, 2-3 mm. long. Fruit with 2 equal, distinctly 5-ribbed valves. Honduras, Panama, and Colombia. cocLE; Penonome, Williams 371. Panama: Rio Sta. Maria, Scemann 34. um Herbae pusillae. Folia plnnata; pinnis primariis fere 8 pro folio, Interdum ala lata ad rhachem dccurrente; pinnis secundariis Interdum suboppositis; plnnis tertlariis repetito furcatis; laciniis ultlmls lanceolatis, acutis, enerviis, 0.5-2.0 mm. longis; incisionlbus obtusls. Flores 1-3, solitarii vel fasciculati; pedicello aplce non dilatato; spathella juvenile clavata, acuta, vetusta infundlbuUforme; tepalis 7, squamlformibus; staminibus 7, filamentls lanceolatis basi trlangulare; antheris obtusls, emarglnatis vel mucronatis; granulls poUlnis subglobosis; ovarlo ovoidco, acuto, basl attenuato, valde applanato, distlncte 8-costato, costis paulo promincnti- bus, duobus eorum Indistlnctlssimis; stylls cylindrlcis, aplce oblique truncatis, facie interne emarglnatis, aliquando papillatis. Fructus valvis 2 acquallbus, utraque valva 5-costata. Small species with base 1-4 mm. long and wide, 1-2 mm. thick. Leaves pinnate, 0.5-2.5 cm. long; primary pinnae about 8 per leaf, 1-7 mm. long, some- times with a broad wing running down the rachis; secondary pinnae 0.5-2.5 mm. long; tertiary pinnae repeatedly forked, 1.0-2.5 mm. long, the ultimate divisions lanceolate, acute, nerveless, to 2 mm. long, the Incisions obtuse. Flowers 1-3, solitary or in small groups; pedicel 0.7-2.0 cm. long; young spathe clavate, acute, splitting at the top or side, 1.5-3.0 mm. long, the older infundibuliform, 4-5 mm. lone:; tepals 7, squamiform, about 0.3 mm. long; stamens 7, about 3.5 mm. long, (115) 130 [Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig, 57. Marat/jrum pusillum cm. filaments lanceolate, 3-sided at the base, about 2.5 mm. long, anthers LO-1.5 mm. long, obtuse, cmarginatc or mucronate, the thecae unequal; pollen subglobose, about 15 X H ^; ovary ovoid, acute, base attenuate, strongly flattened, with 8 very slightly raised ribs (2 of them vcr}^ indistinct), 2.0-2.5 mm. long, about 1 mm. wide, the styles cylindrical, apex obliquely truncate, cmarginate at the inner side, slightly papillate, 1.5 mm. long. Fruit with 2 equal 5-ribbed valves. Leaves sometimes with a dilated and cuneately fasclatc-flabclliform rachis about 1 long and 2-3 mm. wide at the tip and there with 3-5 lobes with ultimate divisions as described above. PANAMA: Chagres River, Piftier 2303 (holotype in U. S. Nat. Herb.). This species is recognized easily by Its size, since It is the smallest Maratlnnm ever found. The cuncate leaves, which are often on the same specimen with the normal leaves, are characteristic, too, and by these leaves the plants are different from all other Marathrums, (116) 19 SO] FLORA OF PANAMA (Podosfe?nonaceac) 131 Fig, 58, Marathrufft Icptophyllum 4. Marathrum leptophyllum v- Royen, sp. nov. Herbae parvae, acaules e radicibus longis, tenuibus appalantisque binatlm nascentes. Folia tenuia, 3- vel 4-tlm pinnata; petiolo juventate byalino, intcrdum brevi tenui ligula 1 mm. alta instructo; pinnJs primariis pinnatis vcl bifurcatis; pinnis tcrtianis repetito bifurcatis, laciniis ultimis lanccolatis, subfiliformibus, membranaceis, acutis, encrviis, Flores pauci, juveniles soli noti: spatha 3-5 mm. longa, clavata, mucronata, subpapillosa, adulta ignota; tepalis 7, lanceolatis, acutis; staminibus 7, filamentis membranaceis, lanceolatis; antheris obtusis vcl emarginatis; granulis poUinis 20 X H /^j lobis angustis, ovoideis; ovario ovoideo, acuto, basi rotundato vel attenuato, applanato, costis 8, quarum 2 indistinctis; stylis filiform- Ibus, apice oblique truncatis, facie interne emarginatis. Fructus valvis 2 aequalibus, utraque valva 5-costata, costis prominentibus. (117) 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN |V',L. 37 Herb with small shoots arising in pairs from long, thin, flattened roots; base about 0.5 cm. long and wide, 2 mm. thick. Leaves delicate, repeatedly pinnate, 3-6 cm. long, with all the pinnae ascending at an angle of 45-90°; petiole 0.5-1.5 cm. long, about 1 mm. wide, membranaceous, transparent when young, sometimes with a distinct, membranaceous ligule about 1 mm. high at the base; primary pinnae 2.5-15.0 mm. long, pinnate or forked, alternate or slightly opposite; secondary pinnae 2-6 mm. long; tertiary pinnae repeatedly forked, about 2.5 mm. long, the ultimate divisions lanceolate, membranaceous, nearly filiform, acute, nerveless, to 2 mm. long. Flowers few, only young ones known: pedicel 1.5—2.0 cm. long; spathe clavate, mucronate, slightly papillate, 3-5 mm. long; tepals 7, lanceolate, acute, about 0.5 mm. long; stamens about 2.5-3,0 mm. long; filaments membranaceous, lanceolate, 1.5-2.0 mm. long, the anthers obtuse or cmarginate, 1,0-1.5 mm, long; pollen 20 X 1 1 /^j lobes narrow, tapering toward the 2 poles; ovary ovoid, acute, rounded or attenuate at the base, 2.5-3.0 mm. long, about 1.5 mm. wide, flattened, 8 -ribbed, all the ribs very slightly raised, 2 of them indistinct; styles 2, filiform with obliquely truncate apex, emarginate at the inner margin, 0.5-1.0 mm. long. Fruit with 2 equal valves, each distinctly 5-ribbcd, the ribs slightly raised, 2.5-3.0 mm. long. CHiRlQUi: Boqucte, Woodson ^ Scbery 8oj (holotype in Gray Herb.; isotype in Herb. Missouri Bot. Gard.). This species somewhat resembles young specimens of Marathrum hnliffercNs but differs from that species because the rachis and petiole are not winged. It differs from all the other Maratlynim species in its transparent petioles, its nearly filiform ultimate leaf-divisions which also are transparent, and in the long and very narrow but distinctly 3-sulcate pollen grains. It closely resembles M. tetmey but that species has only 2 or 3 stamens and its rachis and petiole are narrowly winged. 5. Marathrum indifferens v. Royen, sp. nov. Folia ea Marathri tcuui simulantia sed basi instructa dilatationcm membrana- ceam distlncte punctatam, 3- vel 4-tim pinnata; pinnis primariis ala lata rhachi adnatis, interdum fere opposltis, 60-90^" rhachi decllnatis; pinnis tertiariis repetito bifurcatis, laciniis ultimis subfiliformibus, acutis, erneviis. Flores pauci; spathellis demum 4-12 mm. longis; pedicello saepe 2-alato; tepalis 5-8, squamiformibus; staminibus 5—8, filamentis lanceolatis, membranaceis, antheris acutis vel emargin- atis; granulis pollinis 16 X 14 i^; ovario ovoideo, 8-costato; stylis juvenilibus O.S mm., veteribus 1.5-2.0 mm. longis, linearibus, aplce cmarginato vel acuto. Small herb; base branched or unbranched, 0.5-4.5 cm. long, 0.5-10.0 mm. high, 1—3 mm. thick. Leaves 0.5—15,0 cm. long, 3—4 times pinnate, with all the pinnae alternate or slightly subopposite and ascending at an angle of 60-90°; primary pinnae 1-3 cm. long, widened in the axil and with a distinct wing decur- rcnt upon the rachis; secondary pinnae 1-4 mm. long, the ultimate divisions 0,5-5,0 mm. long, nearly filiform, acute, nerveless; petiole terete or flattened, widened at (in) 1950] FLORA OF PANAMA (Podosfemonaccae) 133 Fig. 59. Maraihrum inJifferens the base, 0.5-3.5 cm. long, 0.5-1.5 mm. wide, dilated with 2 pointed wings at the base. Flowers few; pedicel terete or sometimes 2-winged, 1.5-5.0 cm. long; young spathe unknown, the older infundibuliform, 4-12 mm. long; tepals 5-8, squami- form, 0.5 mm. long or shorter; stamens 5-8, 3.0-6.5 mm. long, the filaments lanceolate, membranaceous, 2.5-5.0 mm. long, the anthers acute or emarginate, 1.0-2.2 mm. long; pollen ovoid, 16 X 14 fx, 3-Iobed, lobes tapering toward one pole; ovary ovoid, acute, base attenuate, 2.2-4.0 mm. long, 1.5-1.8 mm. broad, flattened, 8-ribbed; styles 1-2 mm. long, linear, emarginate or acute. Fruit with 2 equal valves, each 5-ribbed. Mexico, Guatemala, British Honduras, and Panama. CANAL zone: Rio Indio, Dodge & Allen iT39^y ^7399 (holotype in Herb. Utrecht; ISOTYPE in Herb. Missouri Bot. Gard.) ; Rio Boqueron, Steyermark & Allen 1 72 56; Quebrada Ancha, Steyermark & Allen 17TI/'. chiriqui: San Felix, Tittier 5430, tentie distinctly winged; while the latter species has 2 ndiff^ very 5-8 stamens. The distinctly pointed dilation at the base of the petioles characterizes ■fferens from all other species of Marathrnm. (119) 134 [Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fiy. 60. MarafhrN?7i chehiferum 6. Marathrum cheiriferum v. Royen, sp. nov. Herba pusilla vel mcdiocris, acaulis; basi irregulari. Folia bis- vcl ter-pinnata, 2-20 cm. longa; pinnis primarlis Icvitcr nervatis, 30-45° declinatls> lacinils ultimis ovatis, obtusis vel acutis, Icviter nervatis vel encrvatis, vcl interdum oblongis, rotundatis, margine crenulato-Iobatis, 0.3-LO mm. longis, 0.3-0.8 mm. latls; petiolo tereti vel applanato, rhachide striata, anguste alata. Flores pauci, solitarii; spathella claviformi, acuta, deinde infundibuliforml; tepalis 6-8, triangularibus, acutis, minus quam 0.5 mm. longis; stamimbus 6-8, filamcntis subulato-lanceolatis, nervatis, anthcris aplce obtuso vcl mucronato; granulis pollinis globosis, 15 X 14 /a; (120) 1950] FLORA OF PANAMA (Podoslcmojiaccae) 135 ovario ellipsoideo, apice acuto, basi attenuate, costis 8 distinctis, costis suturalibus juventute Indistinctis; stylls filiformibus vel spatbulatis apicc cmarginatis, obtusis, basi paullo connatis, leviter papillatis; capsula valvis 2 aequalibus, distincte 8- costatis. Small to medium-sized herbs; base irregular, 0.5-2.0 cm. long, 2—8 mm. wide, 1—2 mm. thick. Leaves 2-3 times pinnate, with all the pinnae ascending at an angle of 30-45°, 2-20 cm. long; primary pinnae 0.6-4.0 cm. long, 10-20 per leaf, the ultimate pinnae a few times forked; ultimate divisions to 1 mm. long and about 0.5 mm. wide, ovate, obtuse to acute, slightly nerved or nerveless, or the ultimate divisions rotundate and margins crenulate to lobcd; petiole terete to flattened, 0.5—5.0 cm. long, 0.5-2.0 mm. wide at the base, the rachis distinctly striate and narrowly winged. Flowers few, solitary; pedicel terete, 0.5-5.0 cm. long; young spathe clavate, acute, 2—5 mm. long, the older infundibuliform, 6—10 mm. long; tepals 6—8, triangular, acute, less than 0.5 mm. long; stamens 6-8, 3.0-5.5 mm. long; filaments at first subulate but later lanceolate, membranaceous, 3-sided at the base, 2.5-4.5 mm. long; anthers obtuse or mucronate, 1.0—1.5 mm. long; pollen globose, slightly 3-sulcate, 17 X 14 />i; ovary ovoid to ellipsoid, acute, base attenuate, slightly flattened, definitely 8-ribbed, 2 of the ribs indistinct, in young flowers 2.5-3.0 mm. long, 1.0-1.5 mm. wide; styles filiform or spathulatc, obtuse, emarginate, slightly papillate, 1.0—1.5 mm. long. Fruit with equal valves, each distinctly 5-ribbed. Costa Rica, Panama, and Colombia. cocle: Rio Mata Ahogada, Valle Chiquita, Martin SQ/Q; Rio Anton, Ilioitcr ^ Allen JjT. PANAMA: Rio de Jcsiis, Boiichc s. n. (holotype in Herb, N. Y. Bot. Gard.). Closely resembles M. wiruitiflornm but is easily distinguished by the shortly triangular ultimate leaf-divisions, those of the latter being spathulate. 7. Marathrum Allenti Woodson, in Ann, Mo. Bot. Gard. 25:827. 1938. Small herbs; base irregular, sometimes branched, 0.5-7.0 cm. wide, 0.5—4.0 cm. high. Leaves 1—20 cm. long, 3—4 times pinnate, with all the pinnae ascending at an angle of 30-60°; primary pinnae 0.3-3.0 cm. long, membranaccously widened in the axils; secondary pinnae 1—1.5 mm. long, the ultimate pinnae a few times forked, the ultimate divisions spathulate, mostly acute, nerveless or seldom indis- tinctly nerved, to 1.5 mm. long and 0.3 mm. wide; petiole dilated at the base, with a membranaceous, acute to obtuse or emarginate ligule 1-2 mm. long and 1 mm. wide, which sometimes is markedly shifted toward the ventral side, 0.3—7.0 cm. long, 0.5-2.0 mm. wide. Flowers several; pedicel 1.0-7.5 cm. long, terete or 2- winged; spathe clavate, the young 2-8 mm. long, obtuse or mucronate, smooth or slightly papillate, the older infundibuliform, 7-15 mm. long; tepals 6-9 ^ scale-like, 0.5 mm. long or shorter; stamens 5-9, 4.0-5.5 mm. long, the filaments lanceolate, membranaceous, sometimes split at the top, the anthers acute, obtuse or emarginate. (121) [Vol. 37 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 61. Marat brum AUcnil 1,0-2.5 mm. long; pollen ellipsoidal, 17 X 14 ^, the lobes tapering toward one pole; ovary ovoid or ellipsoid, 3.5-4.0 mm. long, terete, acute, sessile or attenuate at the base, 8-ribbcd, the ribs slightly raised; styles filiform, narrow, 3-sided at the base, obtuse or slightly cmarglnate, papillate, nerved, 0.5-2.0 mm. long. Fruit with 2 equal 5-rIbbed valves, the ribs but slightly raised. Panama. ciTTRiQui: Remedios, Pit tier 5443- cocle: El Valle dc Anton, Allen 82; same locality, Wooihon fi> Schcry 160, (122) 1950] FLORA OF PANAMA (Saxifvagaceae) 137 8. Marathrum foeniculaceum Humb. & Bonpl. PL Aequinox. 1:40, tab. II, 1808. Lach foeniciilacea (H, & B.) Mart. Nov, Gen. & Sp. 1:6, 1824, Medium-sizcd herb; base irregular, 0.5—2.0 cm. long, 1.0—1,5 cm. wide. Leaves 2.5—3 5.0 cm. long, repeatedly bifurcated or rarely 3—6 times pinnate; lateral lobes 3—6 times bifurcated or rarely pinnate, 0.5—10.0 cm. long, ascending at an angle of 3 0-60°, the ultimate divisions 0.5-12.0 mm. long, narrow, obtuse or acute, nerveless or very rarely nerved; petiole 1—8 cm, long, flattened, membranaceously widened at the base, with a distinct, sometimes very irregular ligule 2—3 mm. long and 5 mm. wide. Flowers solitary or in fascicles between the leaf bases; pedicel 1.0-1.5 cm. long, thickened into a disc at the tip; young spathe acute, 0.5-1.0 cm. long, the older tubular or infundibuliform, 0.5— LO cm. long; tepals 5—8, triangular, acute or acuminate, about 1 mm. long; stamens 5—8, 3.5—5.5 mm. long, the fila- ment lanceolate, scaphiform at the base, sometimes slightly united, 3—4 mm. long, the anthers obtuse, about 1 mm. long; pollen nearly globose, about 18 X 15 ft, the lobes very wide at one pole and tapering toward the other; ovary ellipsoid, acute, base attenuate, flattened, distinctly 8-ribbed, 3.0—4.5 mm. long, 1.0—1,5 mm. wide; styles flattened or subulate, acute or emarglnate, 1.0-2.5 mm. long. Fruit dehiscing with 2 equal, 5-ribbed valves. Mexico to Colombia. Panama: locality unknown, Hayes 824, SAXIFRAGACEAE By ELIZABETH McCLTNTOCK Deciduous or evergreen shrubs or small trees, or perennial, rarely annual, herbs. Leaves alternate or opposite, mostly simple but variable in form, usually without stipules. Inflorescence various; flowers mostly perfect and actinomorphic, medium- sized or small, rarely large and showy; receptacle various, flat or often concave and forming a hypanthium more or less adnate to the ovary; sepals 4 or 5, rarely more; petals as many as the sepals, imbricate or valvate, free; stamens as many as petals and alternate with them, or twice as many as petals, rarely more numerous; fila- ments subulate or filiform; anthers rotund or oblong, 2-celled, opening lengthwise; carpels usually united and fewer than petals, seldom partially free and equal in number to the petals; ovary superior or partly or wholly inferior, placentae parietal or axillary, rarely otherwise, ovules usually numerous, styles the same number as carpels, free or partially united. Fruit a dehiscent capsule or a berry; seeds usually numerous, small and occasionally winged. About 70 genera, widely distributed in both hemispheres, mostly in temperate to subarctic regions, rare In the tropics where only certain woody genera are found. Four genera in Panama. (n}) [VUL. 11 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN The Central and South American species of Hydrangea belong to the Section CoRNTDiA Engler (Nat. Pflanzenfam. 3^^:76. 1890), characterized by being ever- green climbers or shrubs with coriaceous leaves. Cornidia does not occur ex- clusively in the New World, several species having been described from the Phihppine Islands and Formosa. In addition to Cornidia there are two other sections in the genus which occur in the eastern United States and in China and Japan where the genus Is concentrated. There are two types of Inflorescence In Cornidia, The most common consists of a single cyme, with or without sterile flowers. The second type consists of several cymes one above the other, and does not have sterile flowers. The Central American species have only the first type, but both types are found in the South American species. Tlie characters distinguishing the species within each group are usually found in the flowers: in the presence or absence of sterile flowers, in the size, shape and number of styles, and in the length of the stamens. Because of the small size of the flowers these difi*erences are not readily seen. Sterile, juvenile plants, quite difi'erent from the adult flowering plants, have been collected in Central and South America In areas where the genus occurs. Juvenile plants have small, membranous, serrate or entire leaves borne on slender climbing stems. There are no distinctive generic or specific characters associated with these plants, and they passed unrecognized until they were finally associated with Hydrangea. a. Flowers cpiphyllous, borne in fcw-flowcrcd cymes alon^ the midvcin on the upper leaf surface, inconspicuous, greenish; petals less than 1 mm. long; fruit a few-seeded berry 4. Phyllonoma aa. Flowers not epiphyllous, conspicuous, white or colored; petals 2 mm. or more long; fruit a many-seeded capsule. b. Inflorescence a many-flowered cyme, often bearin)^ a few large sterile flowers In addition to the smaller, fertile ones; fertile flowers with petals about 2 mm. long, deciduous during anthesis; stamens twice as many as petals; capsule opening at apex between the styles; styles developing during maturity of fruit to about double their size in anthesis 1. Hydrangea bb. Inflorescence a few-flowered cyme, or flowers borne singly, flowers alike; petals 10 mm. or more long, persistent during anthesis; stamens same number as petals or numerous; capsule opening from base be- tween the valves or from the apex loculicidally; styles not developing further during maturity of fruit. c. Sepals and petals 4, stamens numerous; petals 15-20 mm, long, obovate, spreading; capsule dehiscent loculicidally from apex; leaves deciduous; leaves and branchlets hirsute-strigosc.,- 3. PniLADELrHUS cc. Sepals, petals and stamens 5; petals 10 mm. long, linear-spathulate, erect, appearing to cohere and to form a tubular corolla; capsule dehiscing from base between the valves; leaves evergreen; leaves and branchlets resinous and glandular 2. Escallonia 1. HYDRANGEA L. Hydrangea L. Sp. Pi. 397. 1753. Cornidia Ruiz & Pav. Fl. Pcruv. et Chil. Pradr. 53, /. JJ. 1794, %arco%ty\e% Presl, ex DC. Prodr. 4:15. 1830. {124) 1950] FLORA OF PANAMA (Saxifragaccac) 139 Woody climbers or shrubs, inflorescence and adjacent stem pubescent with stellate hairs, or tomcntose with long, curling silky hairs. Leaves opposite, simple, blades penninerved, margins remotely denticulate or entire, oval, broadest near middle, petiole stout. Inflorescence cymose (in Central America), terminal, or occasionally lateral, subtended by chartaceous, caducous involucral bracts; sterile flowers showy, pink or white, not always present; fertile flowers white or maroon, small; hypanthium hemispheric; calyx-lobes 4, inconspicuous, deltoid; petals 4, oblong, valvate, deciduous during anthesis; stamens 8, occasionally 10, filaments long and fiKform or short and inconspicuous, anthers oblong to somewhat rounded; ovary inferior, 2- to 4-cellcd, styles 2-4, erect and spreading, or occasionally co- herent in bud and early anthesis, continuing to elongate with the maturing capsule; stigmas terminal and more or less decurrent on inner surface; ovules numerous, borne on axial placentae; capsule 2- to 4-celled, truncate, thick-walled, dehiscent at apex between the spreading styles; seeds numerous, very small, linear, arcuate. a. Sterile flowers present; styles usually 2, slender or clavate. b. Stamens conspicuous, filamuits 1-3.5 mm. long, 2-7 times as long as styles; styles (during anthesis) less than 1 mm. long, slender, acute at apex..... 1, H. Ofrstedii bb. Stamens inconspicuous, filaments 0.2-0.9 mm. long, shorter than styles; styles (during anthesis) 1-2 mm. long 2. H. peruviana aa. Sterile flowers absent; styles usually 3, thickened at base 3. H. Prfslii 1. Hydrangea Oerstedii Briq. in Ann. Conserv. & Jard. Bot. Geneve 20:407. 1919. Conihlia nuViafa Oerst. in Naturhlst. Foren. Kjoeb. Vidensk. Meddcl. 42. 18 56, non Hydrangea radhtta Walt. Fl. Carol. 2 51. 1788. Hydrangea duri folia Briq. loc. cit. 406. 1919. Hydrangea Gotidotii Briq. loc. cit. 404. 1919. Hydrangea platypbylla Briq. loc. cit. 401. 1919. ?Hydrangea Secmannii Riley, in Kew Bull. Misc. Inf. 207. 1924. Woody climbers or shrubs, pubescence of stellate bairs, sparse to moderate en inflorescence and adjoining stem, occasionally densely pubescent with long curling hairs (Woodson & Schery jii); leaves oval, 10-15 cm. long, 4-7 cm. wide, their length 2 to 2/2 times their width (Central America), usually pubescent as the inflorescence on lower surface, upper surface glabrous, margin remotely denticulate to nearly entire, petiole stout, 1-2 cm. long; inflorescence open and spreading, longest branches 4-10 cm. long; sterile flowers present, pink, fertile flowers maroon; hypanthium 1-1.5 mm. long; calyx-lobes 0,5 mm. long; petals 1-2 mm. long, 1 mm. wide; stamens 8, 2-4.5 mm. long, fil?.mcnts 1-3.5 mm. long, 2 to 7 times as long as styles; styles 2, rarely 3, 0.3-1 mm. long (during anthesis), slender, acute at apex, stigma inconspicuous; capsule 2 mm. long, 2.5 mm. wide at apex, mature styles 2 mm. long. ? Mexico, Costa Rica to Peru. CHiRiQui: central valley of Rio Chlriqui Viejo, 18 00-2000 m., Allcfi 1400; Bajo Mono and Quebrada Chiquero, 1500 m., Woodson & Schery 511. cocle: region north of El Valle de Anton, 1000 m., Allen 26c) j, 37 1 2. (125) [Vol. 37 140 MISSOURI 2. Hydrangea peruviana Moric. in DC. Prodr. 4:14. 1S30. Coruhfia peruviana (Moric.) Small, in N. Am. Fl. 22:161. 1905. Hydrangea Trianac Briq. in Ann. Conscrv. & Jard. Bot. Geneve 20:403. 1919. Hydrangea panamensis Standi. In Jour. Wash. Acad. Sci. 17:10. 1927. Hydrangea Weberbaucri Englcr, In Eni;!. & Prantl, Nat. Pflanzcnfam. 18a:207. 1930. Hydrangea caucana Engl. loc. cit. 206. 193 0. Woody climbers or shrubs, pubescence of stellate hairs, sparse to moderate on inflorescence and adjoining stem; leaves oval, 10-23 cm. long, 5-14 cm. wide, tbcir length 2 to 3 times their width, both surfaces usually glabrous, margin entire to remotely denticulate; inflorescence open and spreading, longest branches 5-15 cm. long; sterile flowers present, pink, fertile flowers maroon, hypanthium 1.5 mm. long; calyx-lobes 0.4 mm. long; petals 1.5 mm. long, 1 mm. wide; stamens 8, 0.4-1.6 mm. long, filaments 0.2-0.9 mm. long, always shorter than styles; styles 2, rarely 3, 1-2 mm. long (during anthcsis), clavatc, stigma conspicuous, extending from apex along inner surface; capsule 2 mm. long, 2.5 wide at apex, mature styles 2 mm, long. Costa Rica to Peru. colon: along Rio Fato, 10-100 m., Pittkr JQIQ- chiriqui: Boqucte Distr., Bajo Chorro, 2000 m., Dav'idwn J4; valley of the upper Rio Chiriqui Viejo, ^'hiic QOa. Hydrangea penniaiia Moric. is very close to H. Oersicdii Briq. They are similar in habit, vegetative characters and fruits, and both have showy, pink, sterile flowers. Their only difference lies in the length of their stamens and styles. In H. Oersfedii the filaments arc several times longer than the styles. The styles arc slender, lack a prominent stigmatic surface and are less than 1 mm. in length. The prominent stamens can be seen with a lens or even the naked eye. The filaments of H. pcrniiana, on the other hand, are shorter than the styles which are clavate and from 1 to 2 mm. long. With the aid of a hand-lens it should be possible to distinguish between these two species in the field, provided that flowers are present. 3. Hydrangea Preslii Briq. in Ann. Conserv. & Jard. Bot. Geneve 20:409. 1919. Cornidia umbeUata Ruiz & Pav. Syst. Veg. 91. 1798. Sareostyles peruviana Prcsl, ex DC. Prodr. 4:15. 1830, non Hydrangea peruviana iMoric. in DC. Prodr. 4:14. 1830. Hydrangea umbeUata Briq. loc. cit. 411. 1919, non RchJcr. Hydrangea ecnadorensis Briq. loc. cit. 410. 1919. Hydrangea Bviqnefii Englcr, In Engl. & Prantl, Nat. Pflanzenfam. ]8a:207. 1930. Woody climbers or shrubs, pubescence of stellate hairs sparse to moderate on inflorescence and adjoining stem; leaves oval, 9-20 cm. long, 4-9 cm. wide, their length 2 to 3 times their w^dth, margin entire and somewhat undulate, petiole stout, 15-20 mm. long; Inflorescence more or less compact, longest branches 4-9 cm. long, sterile flowers absent, fertile flowers maroon; hypanthium 1.5 mm. long; calyx-lobes 0.5 mm. long; petals 1.5-2 mm. long, 1 mm. wide; stamens 8, 1.5-2 mm. long, filaments 1-1.5 mm. long, approximately as long as styles (in Central America); styles usually 3, occasionally 2 or 4, 0.8-1.5 mm. long (during anthcsis), (126) 1950] FLORA OF PANAMA (Saxtfragaccae) 141 Fig. 62. Hydrangea pcritviana (127) [Vol. 37 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN thickened at base, stigma usually extending from apex along inner surface; capsule 2 mm. long, 2.5 mm. wide at apex, mature styles 1.5-2 mm, long. Costa Rica, Panama, Ecuador, and Peru. cocle: vicinity La Mesa, hills north of El Vallc de Anton, 1000 m., Allen 2^^2. Hydrangea Preslii differs from the other two species in the lack of sterile flowers and in the number and shape of the styles. The styles are usually 3, although oc- casional flowers may have 2 or 4. They are thickened at the base and attenuate toward the apex, while in the others they are slender or clavate. 2. ESCALLONIA Mutis EscALLONiA Mutis ex L. f. SuppL 2L 1781. Stcreoxylon Ruiz & Pav. Fl. Pcruv. et Chil. Prodr. 38, /. 6. 1794. Small trees or shrubs, evergreen, rarely deciduous. Leaves alternate, shortly petiolate, lanceolate, oblong-elliptic or lance-obovate, mostly leathery and glandular- serrate. Inflorescence racemose or paniculate, few- to many-flowered, or flowers borne singly; hypanthium nearly globose or turbinate, completely or partially united to ovary, cal)'x-lobes 5, ovate, subulate at apex; petals 5, imbricate in bud, linear-spathulate, with long, erect claws which appear to be coherent and to form a tubular corolla, bearing at their apices spreading blades; stamens 5, inserted on margin of hypanthium, erect; filaments filiform, anthers oblong-ovoid; ovary with 2—3 complete or incomplete chambers, with 4-6 placentae which hang from an apical, central axis and bear numerous ovules; style mostly simple, rarely 2 -parted at apex, with capitate or lobed stigma; capsule with 2-3 chambers, scpticidally de- hiscent from the base into 2 or 3 valves which hang together by the bundles of the hypanthium and the partitions, after the partial or complete shedding of the epi- carp (hypanthial wall and style) ; seeds many, usually small, often curved, with longitudinal grooves in the covering, fimbriate at base. A large genus, mostly of the Andes and southern Brazil in South America, with two species in the mountains of Central America. One species in Panama. 1. EscALLONiA POASANA Donn. SiTi, in Bot. Gaz. 23:243-244. 1897. Small tree up to 2 5 feet tall, irregular in shape; branchlets short, closely leafy, somewhat resinous and glandular; leaves 10--20 mm. long, 6-9 mm. wide, obovate, rounded at apex, cuneate at base and narrowed to a short petiole 2—3 mm. long, margins finely denticulate, at least in upper half, somewliat resinous, subcoriaceous, lower surface whitened and reticulate, upper surface green; flowers white, borne singly, apically or laterally, on the short branchlets, peduncles 5-15 mm. long; hypanthium united to ovary in lower half, 7 mm. long, diameter at upper margin 6 mm., calyx-lobes 2 mm. long, deltoid, acuminate toward apex, margins fimbriate; petals 10 mm. long, blade 3 mm. long, scarcely broader than claw, rounded at apex with a short mucro at tip, margins fimbriate; stamens 7.5 mm. long, filaments 5 (m) 1950] FLORA OF PANAMA (Saxifvagaceae) 143 mm. long, anthers oblong; style entire, 5 mm. long, stigma depressed-globose, crenate on margin; capsule 7-8 mm. long, at upper margin 7 mm. in diameter, bilocular, dehiscent at base, Costa Rica and Panama. CHiRiQui: Boquete Distr., Volcan de Chiriqui, 10,400 ft., Davidson T0J4, 1306; Chiriqui Volcano, 2800-3000 m., Piltier JO/Q; Rio Chiriqui VIejo, White 6. 3. PHILADELPHUS L. Philadelphus L. Sp. Pi 470. 1753. Syrifiga Adans., Fam. Pi. 2:244. 1763. Deciduous shrubs with slender, arching and occasionally scandent stems with close or sometimes flaking bark. Leaves opposite, entire or dentate, sessile or shortly petiolate, lower surface often pubescent with simple hairs, upper surface green and glabrous or with few scattered pili, usually with 3-5 prominent veins from base. Flowers white, showy, often fragrant, soUtary or in few-flowered terminal cymes; hypanthium turbinate, united with the ovary, calyx-lobes 4, rarely 5-6; petals 4, rarely 5-6, obovate, rounded or retuse at apex, convolute in bud; stamens 20 or more, filaments flat, subulate, anthers small, oblong; ovary inferior to half superior, 4 or rarely 3 or 5 chambers, placentae axial, bearing numerous ovules; styles 3-5, fihform, more or less united, each bearing an elongate stigma; capsule turbinate, ligneous or coriaceous, 3-5 chambers, dehiscing locuHcidally from the top; seeds numerous, oblong. About 40 species in North and Central America, eastern Asia and southern Europe. 1. Philadelphus myrtoides Bertol. Fh Guat. 21, fab, 7 (in Nov. Comm. Acad. Sci. Inst. Bonon. 4:421, pL 43). 1840. rhiJaJcIphiis irichopetalus Kocrn. in Kegel's Gartenfl. 16:73. 1867. Shrub with arching, strigose and often scandent stems; leaves 3-9 cm. long, ovate, acuminate at apex, rounded at base, remotely denticulate, 3- to 5-nerved at base, pale and hirsute-strigose beneath, green with few pili above, short-petiolate; flowers pedicellate, 1-3, on short branchlets; hypanthium and calyx grayish- strigose, calyx-lobes 5-10 mm. long, deltoid-acuminate; petals 15-20 mm. long, broadly obovate or suborbicular, glabrous or finely villous on both surfaces; stam.cns and style about one-half as long as petals; capsule 10 mm. long, Guatemala, also in Costa Rica and Panama, but perhaps indigenous only In Guatemala. Close to Philadelphus mexicamis Schlecht. of southern Mexico. chiriqui: Bajo Mona and Quebrada Chiquero, 1500 m., Woodson ^ Schery 588— "Side of a footpath, possibly an escape." (129) 144 [Vol. 17 MISSOURI 4. PHYLLONOMA Willd. Phyllonoma Willd. ex Rocm. and Schult. Syst, Vcg. 6:210. 1820. Dulongia HBK. Nov. Gen. & Sp. 7:76. 1825. Shrubs or small trees, completely glabrous; leaves alternate, membranous or coriaceous, lanceolate to ovate-lanceolate, long-acuminate toward apex, entire or denticulate, petiolate. Inflorescence few- to many-flowered, cymose or racemose, arising from the midvein on the upper surface of the leaves near the base of the acuminate tip; flowers small, greenish-white; hypanthium turbinate, completely attached to ovary, disc eplgynous, thick, covering base of petals and stamens; calyx-lobes 5, deltoid, minute; petals 5, valvate in bud, spreading in an thesis, deltoid-ovate, acute at apex; stamens 5, alternating with petals, spreading, fila- Fig. 63. Phyllouonid ruscifolia F ments short, subulate; ovary inferior, 1 -celled with 2 rows of erect ovules on 2 parietal placentae, very short, styles 2, scarcely protruding from disc, spreading, with stigmatic surface at apex; fruit a berry, with scars of sepals and petals at apex, imperfectly 2-celled; seeds 3-6, small, oblong. A small genus occurring from the high mountains of southern Mexico through the Andes to Bolivia, with one species in Panama. 1. Phyllonoma ruscifolia Willd. ex Roem. and Schult. Syst. Veg. 6:210. 1820. Dulongia acii7fUfiala HBK. Nov. Gen. & Sp. 7:78, pi. 623, 1825. Small tree to 20 feet; leaves ovate-lanceolate, coriaceous, blade 40-8 5 mm. long, -30 mm. wide, gradually narrowed to the slender pointed acumen 10-2 5 mm. 10 (130) 1950] FLORA OF PANAMA (Ctinoniaceae) 145 long, remotely denticulate below acumen nearly to base; inflorescence immediately below acumen or on its lower half, several-flowered; hypanthium broadly turbinate, disc 0.7 mm. across; calyx-lobes 0.2 mm. long; petals 0.5 mm. long; stamens less than half as long as petals; styles and stigmas 0,1 mm. long; berry 4 mm. In diameter, white turning black on drying. Costa Rica to Bolivia. CHiRiQui: Boquete Distr., Cerro Horqueta, i07t Hagcn 2043. CUNONIACEAE 1. WEINMANNIA L. WiENMANNiA L. Syst. ed. 10, 2:1005. 1759; Engl, in Engl. & PrantI, Nat. Pflan- zenfam. 18a:250. 1930. ArnoUia Bl. Bijdr. 868. 1826. Leiospcrtuunt D. Don, in Edinb. New Phil. Jour. 9:91. 1830. Ptcropbylla D. Don, loc. cit. 9 3. 1830. Ornithropus Boj. ex Engl, in Linnaca 36:636. 1870. Trees or shrubs, the younger branches usually somewhat flattened and rotatmg to about 45 "" between the nodes. Leaves opposite, usually imparipinnate, often with the rachis winged between the opposite leaflets, rarely simple; stipules interpctiolar, caducous. Inflorescence frequently appearing terminal but axillary and usually opposite in the upper leaf-axils, a racemiform or spiciform thyrse with the pedicels clustered and nearly sessile upon the short side branches. Flowers numerous, small, perfect or polygamo-dioccious; sepals 4-5; petals 4-5, white or somewhat pinkish; stamens 8-10; ovary superior, 2- or rarely 3-cellcd, each carpel with a gradually produced, persistent style with simple stigma; ovules numerous, in 2 rows upon the axile placentae; disc fleshy, hypogynous, surrounding the ovary but free from the stamens. Fruit a small, thin, septicldally 2- or rarely 3-valved capsule; seeds small, usually pilose. Wciumannia is a genus of upward of 100 species of the tropics and subtroplcs of both hemispheres. The species are notoriously variable, and much confusion has resulted from the failure of certain students to appreciate this fact. a. Leaves imparipinnate, the rachis broadly winged between the small leaflets; flowering branches relatively distant, the inflorescences not greatly aggregated "- ^- ^- tinnata aa Leaves simple; flowering branches conspicuously shortened, the in- florescences appearing greatly aggregated - 2. V. PSEUDOLAURINA 1. Weinmannia piNNATA L. Syst. ed. 10, 2:1005. 1759. Wcinwannta glabra L. f. Suppl. 228. 1781 (based on W. pwnafa L.)- Wcinmannia intermedia Cham. & Schlecht. in Linnaea 5:555. 1830. Weinjvanuia burseraefolia Standi, in Jour. Wash. Acad. Sci. 17:310. 1927. Weirimanfiia Liebmaunii Engl, in Engl. & Prantl, Nat. Pflanzcnfam. 18a:252. 1930, nom. subnud. (131) 146 I Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 64. Weinmamtla pinnaia Large shrub or small tree sometimes as much as 20 m. tali, the younger branches ferruginous-pilose to glabrate. Leaves imparipinnate, petiolate, 3-9 cm. long, the rachis broadly winged in rhombic sectors between the pairs of leaflets; lateral leaflets in 2-6 pairs, oblong-elliptic to obovate, unequally cuneate at the base, acute or obtuse at the tip, rather closely crenulatc, about 0.7-3.0 cm. long and 0.5-LO cm. wide, wholly glabrous or more or less pubescent beneath; terminal leaflet usually somewhat larger and more symmetrical than the lateral, rhombic-elliptic to obovate, obtuse to acuminate; petiole 0.7-2.0 cm. long. Inflorescences usually distinctly paired in the axils of the uppermost pair of leaves, narrowly cylindrical, 2-9 cm. long, the pedicels 2-3 mm. long, slender, usually more or less ferruginous- pilose like the peduncle; sepals triangular, persistent in fruit, about 1 mm, long; petals white or rarely pinkish, slightly longer than the sepals; stamens with white filaments 2-3 mm. long, the small anthers orange. Capsule glabrous, 3-4 mm. long; seeds pilose. Southern Mexico to Brazil and Peru; Cuba, Jamaica, and throughout the Antilles, growing in forests as a rule, at temperate altitudes. CHlRiQUi: valley of the upper Rio Chiriqu! Viejo, vicinity of Monte Lirio, alt. 13 00- 1900 m., Scibert 288; summit of Cerro Copcte, alt. 9000 ft., Allen 48q2; Finca Lcrida to Pcna Rlanca, WooJsou & Scbcry JJO; Volcan dc Chiriqui, Daihhon 977; Cerro Hor- quL'ta, cloud forest, alt. 6500 ft., von Hagen ^ von Hagcn 2o68; vicinity of Finca Lcrida, upper forested slopes of Quebrada Velo, alt. 5000 ft., Allen 4736; vicinity of Casita Alta! Volcan de Chiriqui, Woodson, Allen G? Scibert 821. (132) 1950] FLORA OF PANAMA (Rosaccac) 147 Wcinmannia pifmafa is a very frequent tree in Chiriqui at altitudes between approximately 1200 and 2500 m. The number, size, and shape of the leaflets, as well as their indument, is exceedingly variable; this has given rise to a formidable array of "species", only the more important of which, from Central America alone, are enumerated above. It is very difficult to understand why Engler (loc. cit. 1930) chose the name W. glabra L. f. for the nucleus of this complex, when that name is so clearly based upon the earlier W> phniata L. 2. Weinmannia pseudolaurina Woodson, Ann, Mo. Bot. Card. 28:429. 1941. Trees at least 8 m. tall; branches strongly compressed and densely ferruginous- pilosulose when young, the flowering internodcs greatly contracted. Leaves simple, shortly petiolate, broadly elliptic or eUiptlc-ovate, rather distantly crenulate, apex obtuse or broadly acute, base obtuse or broadly rounded, 5-9 cm. long, 3.5~5.0 cm. broad, subcoriaccous, glabrous above, inconspicuously pilosulose to glabrate beneath; petiole 2—5 mm. long. Inflorescences densely aggregated in pairs at the tips of the shortened flowering internodes, racemiform, narrowly cylindrical, 6-8 cm. long, the pedicels 3—4 mm. long, ferruginous-pilosulose like the peduncle; sepals per- sistent in fruit, triangular, about 1 mm, long, very minutely pilosulose; petals unknown; stamens 10, the filaments glabrous, about 2 mm. long. Capsules ovoid, about 3 mm. long, densely ferruginous-pilosulose; seeds densely plumose at both en ds. Endemic tree 'Very common along edges of grasslands" at altitudes of 600 800 m. PANAMA: Cerro Campana, trail from Campana to Chica, Allen 26 j6. This species is particularly conspicuous, and unique amongst Weinmannias of its immediate alliance, because of the densely aggregated inflorescences. ROSACEAE By ROGERS McVAUGH Herbs, shrubs or trees with leaves alternate, rarely opposite, simple or com- pound, usually with conspicuous stipules. Flowers usually perfect and regular, the perianth perigynous, the axis sometimes enlarged to form a flat or concave recep- tacle or hypanthium bearing the calyx-lobes, petals, and stamens on its margin, and usually lined inside with a glandular disk; calyx-lobes 5 (sometimes 4), imbri- cate; petals usually 5 and imbricate, sometimes wanting; stamens few to many, often about 20; carpels 1 to many, distinct or united, often connate with the receptacle; styles as many as the carpels, sometimes connate. Fruit various; seeds usually without appreciable endosperm, the cotyledons often fleshy and convex, rarely folded or convolute. The Rosaceae comprise approximately 100 genera, well represented in almost all parts of the world. They include many plants of economic importance, especially in temperate regions where they are much cultivated for food and orna- ^35; \ k 148 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 mcnt (e.g., the genera TragariUy KuhiSy Rosa, Spiraea, VrnmiSy Pyrus, Mains, Chacrwmelcs, Rbodotypus, etc.). Some workers have advocated the division of the family into several segregate families, on the basis of characters of the fruit; thus genera with the fruit an achcne or a follicle have been made to constitute the Rosaceae proper, genera with drupaceous fruit the Amygdalaccac, and genera with pomaceous fruit the Malaceae. These segregations appear unjustified because of the very large number of characters of the flowers, inflorescences and vegetative parts which are repeated in genus after genus without regard to fruit type, and which together make of the inclusive Rosaceae an easily comprehensible and evi- dently natural group of related genera. Precise determination of specific limits is difficult in many genera, related groups often seeming to merge morphologically or to differ by minor characters only. Apparently much of the difficulty arises from the frequent occurrence in the family of reproductive methods other than a sexual fusion of haploid gametes followed by regular reduction-division. Several genera are known to include species In which polyploidy, ancuploidy, apomlxis and other, irregularities have been demonstrated, and it may be supposed that these phenomena have contributed largely to the formation of the numerous minor geographic races which are known to exist, in nature Isolated in small disjunct areas or if growing with other related species then barely distinguishable from th cm. a. Leaves pinnatcly or palmately compound, with 3 or more leaflets, b. Upright or trailing shrubs with prickly canes and leaves (brambles) ; petals present and conspicuous; fruit of few or numerous drupelets borne on an enlarged receptacle 3. RuBUS bb. Herbaceous or low suffrutescent plants with unarmed leaves and stems; corolla wanting. C. Hypanthium covered with barbed spines, in fruit forming a bur; erect plants up to 1 m. tall, the inflorescence splcate or racemose.... 5. Acaena cc. Hypanthium unarmed; flowers very small (rarely as much as 3 mm. long) ; plants mostly prostrate or procumbent, the inflorescence cymose, often corymbiform , 4. Alchemilla aa. Leaves simple. b. Herbaceous, usually prostrate plants with lobcd leaves; flowers very small (rarely as much as 3 mm. long); corolla wanting; calyx-lobes alternating with bracts, the lobes apparently 8-10 in number; hypanthium at maturity containing a few achencs 4. Alchemilla bb. Shrubs or trees with simple unlobed leaves; flowers larger; corolla usually present; calyx-lobes usually 5, without alternating bractlets; fruit dry or fleshy. c. Styles 5, distinct; fruit of 5 thin-walled achcnes, or of 5 bony nutlets in a fleshy hypanthium (pome); leaves dentate, d. Fruit of 5 achcnes about 2 mm. long; unarmed shrubs with leaves white- tomentose beneath, and flowers in ample panicles 5-15 cm, long 1. Holodiscus dd. Fruit a pome up to 1 cm. long, red or black; splnesccnt shrubs, nearly glabrous or the branchlets rufous-pubescent, the flowers in corymbose cymes 2-3 cm. long 2. Hcsperomeles cc. Style 1; fruit a dry or fleshy drupe with a single large stone; leaves entire. d. Style terminal on the ovary or essentially so; stamens usually 20; flowers (in Panamanian species) in glabrous simple axillary racemes which are ebracteate at flowering time 6. Prunus / (134) 1950] FLORA OF PANAMA (Rosaceuc) 149 dd. Style arising from the base of the ovary; stamens usually 15 or fewer; flowers in cymes or panicles, if in simple racemes these bracteate and pubescent. e. Filaments 10-16 mm. long; stamens 3-7 9. Hirtella ee. Filaments 1—6 mm. long; stamens 15—20 (—3 0?) or if only about 3 anther-bearing, then less than 1 mm. long, f. Hypanthium 5-6 mm. long (excluding the calyx-lobes) in bud, elongate, somewhat gibbous, the ovary attached lateral- ly somewhat above the base of the hypanthium-cavity 10. Couepia ff. Hypanthium 3 mm. long or less in bud and in anthcsis, cup-shaped or campanulate, symmetrical, the ovary attached at the base of the internal cavity, g. Flowers 8-20, in nearly sessile axillary cymes (2-4 cm. long) shorter tlian the leaves; low shrub, or medium- sized tree up to 5-6 m. tall 7. Chrysobalanus gg. Flowers very numerous, in axillary and terminal panicles 10—40 cm. long, with divaricate leafless brandies; trees, often 10-15 m. tall or more 8. Licania A weedy immigrant from the United States, Votentilla norvegica L., was re- ported from San Jose Island by Johnston (Sargcntia 8:31, 132, 1949). It is not to be expected that this species will maintain itself permanently in competition with the native flora. It is a hirsute herbaceous annual or biennial with 3-foliolate leaves, leafy cymes, and conspicuous yellow flowers. 1. HOLODISCUS (Koch) Maxim. HoLODiscus (Koch) Maxim, in Acta Hort. Petrop. 6:253. 1879; Focke in Engl. & Prantl, Nat. Pflanzenfam. 3^:18. 1894; Ley in Bull. Torrey Bot. Club 70:275-288. 1943, nom. conscrv. Sericotheca Raf. Sylva Tellur. 152. 1838; Rydb. in N. Am. Fl. 22:261-266. 1908. Spiraea sect. HoloJiscus Koch, Dendr. 1:309. 1869. Deciduous shrubs or small trees up to 7 m. tall, with simple, short-petiolate, estipulatc, dentate leaves, the blades mostly with a long cuncate entire base, the veins ascending and parallel to the cuneate base, impressed on the upper surface, the lower surface often tomentose. Flowers white, 5-merous, in large terminal panicle-like clusters; pistils distinct, 2-ovulate, alternate with the sepals, inserted on center of disk, each forming at maturity an indehiscent 1 -seeded dry fruit. This genus has experienced several systematic revisions, and those who have worked on it have usually recognized two principal species-groups, one ranging from British Columbia to northern Mexico, the other from southern Mexico to Colombia. The southern group has been distinguished by having long-mucronate teeth on the leaf-blades, by having the stamens not longer than the sepals, and by the rather strongly pubescent outer surface of the petals. The two most recent monographers of Holodisc7is, Rydberg (in the North American Flora) and Ley (in the Bulletin of the Torrey Botanical Club) each recognized three species in this "southern group" of the genus, although the two authors differed in their ideas of how the species were to be delimited. (135) s [Vol. 37 150 ANNALS OF THE MISSOURI BOTANICAL GARDEN Examination of large series of specimens from western United States indicates that neither author is conservative in delimitation of species, each upholding as species several groups which are morphologically scarcely distinguishable, and in addition are connected by numerous intermediates. The differences between the supposed species are chiefly those in the leaves, which vary in size, shape and vesti- turc. Study of the meager amount of Central American and South American material which Is available Indicates that all the supposed species of the above ''southern group" are in actuality but representatives of a single species. The leaves are usually oblanceolate or obovate, with the lower surface tomentose and bearing in addition few or many straight silky hairs. Rydbcrg and Ley have at- tempted to separate species here on the basis of the pubescence of the upper and lower surfaces of the leaves, respectively, and Ley has also used the shape of the leaf-tip as a criterion. These characters do not seem to be correlated with other morphological ones, however, nor with any geographical area, and in any case both characters are notably untrustworthy in the Rosaceae. The single Panamanian specimen which has been available for study has some leaves rounded at tips and others acute (the usual condition in this genus and other rosaceous ones, the rounded ones being produced first at the initiation of the growing season), so that according to Ley's key it could be either Holodisnis ari^cnicu^ or f/. fissus; many of the leaves agree precisely in shape with her illustration (Bull. Torrey Bot. Club 70:287, /I^s;. J. 1943) of H, argentcus, but the pubescence of the lower surface is less abundantly provided with silky hairs than most specimens she has called H, argcntcus, and agrees well with those she has named H. fissns. The latter, how- ever, she does not report from south of Guatemala, and it is moreover excluded by having the upper leaf-surface glabrous, not strongly pubcrulcnt as In the Pana- manian material. The jumble of characters used to separate H. argcnfcus and //. jinus Is so inconclusive that the latter is here reduced to synonymy, following Standley and Steyermark In the Tlora of Guatemala* (Fieldiana: Bot. 24:453. 1946). 1. HoLODlscus ARGENTEUS (L. f.) Maxim. In Acta Hort. Petrop. 6:254. 1879. Spiraea argent ca L. f. Suppl. 261. 1781. Spiraea fissa Lindl, in Bot. Reg. 26:Misc. 73. 1840. HohJisms fissus (Lindl.) C. K. Schncid. 111. Handb. Laubh. 1:495. 1905. Sericotheca fissa (Lindl.) Rydb. in N. Am. FI. 22:265. 1908. Sericotbeca argeufea (L. f.) Rydb. loc. cit. 266. 1908. Shrub usually 1-3 m. high, with gray-tomentulose branchlets strongly but obtusely angled below the decurrcnt leaf-bases, the bark on older branchlets reddish- brown, exfoliating in long strips; leaves oblanceolate or the youngest obovate (on vigorous shoots becoming elliptic to ovate, larger than noted here, and with acute or attenuate tips), the upper surface appressed-pllose with hairs up to about 0.5 mm. long, or distally pubcrulcnt with much shorter hairs, the lower surface densely floccose-tomentose between the veins and in addition appressed-pilose on the veins with hairs up to 0.7 mm. long, and with similar hairs amid the tomentum; blades I \ X (136) 19S0] FLORA OF PANAMA (RoSaCCac) 151 Fig. 65. Holo(lfsc2is argcntciis mostly 1 cm. or less wide (up to 3 cm.), 2-3 (-6) cm. long, rounded to acute nt tip, the basal half entire, cuneate, attenuate to a margined petiole 1—3 mm. long, the margin dentate distally with 4-6 (-8) teeth on each edge, each tooth terminat- ing one of the primary veins of the blade, and tipped by a blunt mucro; veins prominent on both surfaces, strongly impressed above, the primary ones parallel to the sides of the cuneate leaf -base; flowers creamy white, in relatively large paniculi- form oblong to pyramidal leafless clusters terminating leafy twigs, the inflorescence mostly 5-15 cm. long, bracteate, often with its lower branches distant and elongate; petals 2-3 mm. long, with an ovate to orbicular entire blade and a short broad claw, pilose with white straight hairs on the outer surface at least near base on midvcin; stamens usually 20 (1 opposite each petal and 3 opposite each sepal), the filaments subulate, 1.5-2 mm. long, 0.2-0.3 mm. wide at base, the anthers nearly globose in dried material, about 0.3 mm. long; receptacle shallowly concave, tomentose within but the margin thickened Into a fleshy glabrous ring with the stamens inserted on its outer edge, the ring strongly 5-Iobcd In age; styles with capitate stigmas, in flower ascending and parallel, about 1 mm. long, pilose near base; ovaries flattened, heavily white-pilose with hairs 0.5-0.7 mm. long, especially on the sutures, the sides somewhat hairy or glabrous, but beset with numerous short-stlpitate yellowish capitate glands; hypanthium saucer-shaped or hemispheric, 1- to 3 -bracteate at base, at maturity about 3 mm. across, the persistent entire triangular sepals 1.5-2 mm. wide at base, 2-3 mm. long; fruit with membranous wall, laterally (i. e., radially) compressed, biconvex, stipitate, the body about 2 mm. long and 1 mm. deep, the Inner angle nearly straight from base of stipe to tip (137) [Vol. 37 152 ANNALS OF THE MISSOURI BOTANICAL GARDEN of the persistent style, the outer one abruptly outcurved at base of style, then gradually attenuate Into the stipe; total length, including stipe and style, about 3,5-5 mm. Southern Mexico and Guatemala; Costa Rica and western Panama; Cordillera Oriental, Colombia; highlands in forested areas. CHiRiQUi: valley of the upper Rio Chiriqui Viejo, P. White ^S, 2. HESPEROMELES LindL HespiiROMELEs LindL in Bot. Reg. sub pL IQjS. 1837; Macbr. in Field Mus. Publ. Bot. 13-:1065-1069. 1938. Osieomeles auct., quoad pi. Amor. Elcntberocarpnm Schlecht, in Lechl. Berb. Am, Austr. 59. 18 57. Evergreen shrubs or small trees, sometimes depressed and gnarled, often much branched and the young branchlets often spinose-tipped. Leaves simple, variable in size and shape, mostly coriaceous, 3—5 cm. long or less, crenate or dentate. Flowers 5-merous, pink or white, in small terminal corymbose cymes usually ex- ceeded by the leaves; ovary inferior at anthcsis, the 5 carpels distinct from each other from the flowering stage on but attached parietally, at maturity with bony endocarp, enlarged and often somewhat exserted from the fleshy hypanthium; ovule I (occasionally 2) in each locule; cotyledons accumbent. Fruit a small red, purple or black pome 1 cm. long or less, tipped by the persistent sepals. Hespcromeles is a genus of somewhat doubtful standing, scarcely morpholog- ically distinct from Ostcomcles, of which the (Hawaiian and Chinese) species have pinnate leaves but in technical characters agree precisely with the American Hespcromeles. The Old World genus Vyracantlja is evidently very closely akin to HcsperomeleSy as evidenced by similarities between species of the two groups, but differs in having the carpels regularly bi-ovulate, and the cotyledons incumbent. A hybrid between Osteoruclcs siibrotunda and Pyracantha crcnato-scrrata is known (X Pyracomeles Yilmoritiii Rchd. ex Guillaumin in Bull. Soc. Nat. Hort. France, VIj 4:191, 198. 1937). Possibly all three of the above genera should be united under the name Osteorneles LindL (Trans. Linn. Soc. Lond. 13:98. 1821), the oldest name. About 20 species of Hespcromeles have been proposed. These occur in nature at relatively high elevations (mostly above 2000 meters) either in the South American Andes or in the mountains of Panama or Costa Rica. Almost without exception the supposed species have been founded upon leaf characters alone (as witness the names corJata, cnveafa, cscallojilacfolla, ferrughica, lyctcropbylla, lati- foliiiy etc.), A study of the available herbarium material indicates that no general correlation is possible between leaf characters and those of the inflorescences. The flowers and fruits, inflorescences, types and distribution of pubescence, venation- patterns, and occurrence of spinescent branching are very nearly identical in all I < (in) 195 0] FLORA OF PANAMA (Rosaccac) 153 the supposed species within reasonable limits of variation — as nearly identical as would be expected in any genus which had long evolved not in a continuous geographical range but on the series of isolated areas which are the paramos and suhpdramos of the Andes. The differences between the leaves of the supposed species of Hcsperomeles are correlated in a very general way with habitat and with vigor of the plant itself; in other words the large-leaved plants, which are usually those with most vigorous apical growth of the branches, occur mostly at lower elevations under relatively mild climatic conditions. These differences in leaf characters are doubtless now transmitted genetically, even though they may reflect the effect of environments of past geological periods. Apparently many local races have developed, but as these vary so much as to defy precise definition, as they differ in such trivial ways and are all fundamentally so much alike morphologically, it would seem unwise to dignify them by calling them species, especially since such a course would entail the naming of many more than have already been so treated. A treatment that seems to be more in accord with the biology of the group is to consider most of the plants as belonging to a single wide-ranging species with numerous local races (subspecies, varieties, forms, ecotypes, apomictic strains, or whatever other categories further field study may suggest). These local races are of unknown phylogenetic or systematic position and should not be given formal Latin names until their interrelationships and exact distributions have been worked out. The Panamanian member of this group is a small-leaved race indistinguishable morphologically from races of the higher Andes in Colombia, Ecuador, Peru, and Bolivia. It appears to be conspecific with the slightly more vigorous and larger- it is treated here as belonging to that species. Mespiln I. Hesperomeles heterophylla (R. & p.) Hook. Ic. PI. 9:pL 846. 1852. MespHus heterophylla Ruiz & Pav. Fl. Peruv. 4:pL 42 jb. 1802. Mespilus lanuginosa Ruiz & Pav. loc. cit. 4:pL 42 ja, 1802. Osteameles obovata Pittier in Contr. U. S. Nat. Herb. 20:108. 1918. Hcsperomeles obovata (Pittier) Standi, in Field Mus. Publ. Bot. 18:480. 1937. Hesperomeles chiriqnensis Woodson in Ann. Mo. Bot. Gard. 26:288. 1939. Evergreen shrub or small tree up to 5-6 m. high, the smaller forms at higher elevations usually intricately branched, the branches often spinescent; bark on the older branches gray; branchlets reddish-brown with prominent pale lenticels, when young covered with pale or (usually) lustrous reddish-brown hairs about 0,5 mm. long, the hairs contorted at base and curved into a hook or a closed loop, but with their distal ends appressed toward the tips of the branchlets; bark of the branchlets ultimately glabrescent but much roughened by the persistent raised bases of the deciduous hairs; leaves coriaceous, obovate or oblanceolate to elliptic, the blades lustrous and rugose-reticulate above, paler beneath, with a prominulous network of veins, rounded to acute at tips, usually cuneate at base, mostly 1.5—2.5 (—4.5) cm. long and 0,5-1.5 (-2.5) cm. wide, varying from about 2-3 times as long as (139) LVoL. 37 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN wide (on vigorous shoots and especially near tips of branches) to about 1-L4 times as long as wide (on depressed and much-branched plants wuth little apical growth, and especially near the bases of branchlcts) ; petioles 1-4 mm. long; stipules subulate, 0.5-0.7 mm. long, early deciduous; blades glabrous or nearly so beneath, often glabrate above but usually with some pale or reddish hairs per- sistent at least near base In the groove formed by the sunken midvein; margins dentate except near base, with 8-11 low rounded teeth on each edge, the teeth at first tipped with minute glandular processes; flowers in small corymbose cymes terminating the leafy branches, the cymes pubescent and later roughened like the branches, with about 20 flowers or fewer, up to about 3 cm. across, the lower branches up to about 3 cm. long, all the branches bracteate, the bracts like reduced leaves but very narrow, mostly 3-6 mm. long, 0.3-1 mm. wide, very acute at both ends; flowers with a *'strong, sweet scent" (E. K. Balls), on pedicel-like cyme-branches 1-3 mm. long, or essentially sessile; petals white or pale pink, or white with pink margins, obovate-oblong, concave, erose-dentlculatc, glabrous, or pilose on the midline without, rounded at apex (sometimes cuspidate), mostly 3.5-5 mm. long and 3-4 mm. wide, narrowed at base to a stout claw 1 mm. wide; stamens 20, the filaments narrowly triangular, flat, about 1.5-3 mm. long, the anthers pink or rose-colored, about as broad as long, 0.7-1.3 mm. long; disk white- or rufous-tomentose, varying to almost glabrous; styles 5, filiform, 4-5.5 mm. long, distinct to the base but approximate, arising from the inner angles of the carpels, white-pilose for 1-2 mm. at base; stigmas terminal, capitate, about 0.3 mm. across, often slightly oblique; hypanthium in anthesis campanulate or turbi- nate, 3-5 mm. long, the apex truncate and surmounted by the 5 broad-based calyx- lobes which are 2-3 mm. wide at base, 2.5-4 mm. long, deltoid, or with a subulate tip; hypanthium and calyx-lobes vestite like the branches of the cymes, or the lobes glabrous at least toward tips; fruit red, or nearly black when ripe, globose to oblong, fleshy (in some races said to be edible), glabresccnt, up to 1 cm. long, the styles and calyx persistent; pyrenes 4-4.5 mm. long. Costa Rica and Panama; Colombia and Venezuela to Bolivia; thickets, open hillsides, and paramos, mostly at elevations greater than 2000 meters. CHlRiQuf: Potrcro Muleto, Volcan dc Chiriqui, Datidson 1055; Cerro Pando, valley of the upper Rio Chiriqui Viejo, P. White 4J; Loma Lari^a to summit, Volcan dc Chiriqui, Woodson, Allen 6 Scibcrt 10j8, 3. RURUS L. By L. H. bailey RuBUS L. Gen. Pi. ed. 5, no. 557. 1754. Woody plants, erect or trailing or somewhat climbing, bearing flowers and fruits on canes of the second year; in the first year the growths from the root are called primocanes, and in the second year arc known as floricanes when they bear fruit and perish; axes usually bear sharp prickles, and often hairs and stalked (HO) 1950] FLORA OF PANAMA (Rosaccac) 155 glands; leaves (In the known Panama species) 3-folioIate or 5-folioIate except sometimes a few of them simple in the inflorescence, frequently more or less per- sisting until the following year; inflorescence racemiform or paniculiform, axillary and usually terminating the canes, pedicels often lengthening in fruit; flowers perfect and complete (in our species); calyx 5-lobed, sometimes glandiferous; petals 5, distinct; stamens numerous, surpassing the head of many small closely packed pistils; fruit a syncarp composed of small simple coherent drupelets that are either attached as a body to the receptacle (blackberries) or forming a cap (raspberries) that falls intact from the receptacle. Hundreds of species in many parts of the world, particularly in the North Temperate Zone, in the tropics usually on higher lands and mountains. The brambles of Panama are not yet fully collected or well understood, and some of the identifications are subject to modification. Good complete specimens are needed to verify the determinations, with both primocane and floricane parts from the same plant, together with notes on stature and habit. I. Blackberries. Fruit (syncarp) composed of drupelets that adhere to receptacle and the aggregation falling as one body or "berry"; drupe- lets rather large, mostly loosely aggregated; main stems or axes usually angled or furrowed, neither glossy or pruinose. a. Axes of canes and inflorescence not hairy or hispid or setose, more or less closely pubescent; fruits pendent 1. R. panamanus aa. Axes prominently hairy or hispid or setose; fruits erect. b. Canes beset with glandiferous hairs 2. R. praecipuus bb. Canes not glandiferous. c. Leaflets abruptly contracted into very narrow apex; canes pubes- cent but not densely long-hairy; prickles many; clusters very ^<^"se 3. R. tantus CO. Leaflets gradually acuminate, somewhat pubescent; canes thickly shaggy; prickles few; clusters open 4. R. trichomallus IL Raspberries. Fruit a cup of drupelets separating from the receptacle (which remains on the plant); drupelets small, much compacted; main stems or axes usually evenly terete, poHshed and mostly P^"^"OSe 5. R. GLAUCUS 1. RuBUS PANAMANUS Bailey, in Gent. Herb. 6:355. 1944, Diflfuse and scandent; canes whitish-pubescent, with few and scattered hooked prickles; primocane foliage not recorded; floricane leaves minutely pubescent on upper surface, pubescent-tonientulose on veins underneath, finely and sharply serrate, 3-foliolatc or perhaps simple in the inflorescence; leaflets narrow-elliptic, terminal or central one 8-10 cm. long, 4-5 cm. broad, narrowed to long point; flowers a dozen or fewer in axillary and one terminal racemiform open cluster; corolla 1 cm. or less across, white; pistils dark-colored; fruit pendent, about 1 cm. long; drupelets many and closely packed. Panama. CHiRiQuf: Baja Chorro. Boquete Distr., 2000 m., Davidson 72. known in Panama. J (141) [Vol. 37 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. RuBUS PRAEciPUUS Bailey, in Gent. Herb. 6:3 56. 1944. Large bramble, canes sometimes 5 m. long, densely and finely pubescent and covered with brownish glandular hairs, and bearing scattered broad-based hooked prickles; primocanes not recorded; leaves of floricanes large, minutely hairy on upper surface, softly pubescent underneath, sharply minutely serrate, 3- to 5- foliolate; leaflets oblong-acuminate to elliptic-acuminate, 8-11 cm. long, 4-6 cm. broad; petiole glandular-hairy; inflorescence terminal, panicle-like, 20-3 cm. long, central rachis continuous, glandular-hairy; flowers white to pink, 2-2.5 cm, across; fruit oblong, 2 cm. or less long, very seedy. Panama. CHiRiQuf: Qucbrada Velo, 1800 m. alt., WooJson ^ Schery 2^^^ vicinity of Casita Aha, Volcan de Chiriqui, 1500-2000 m. alt., Wooitwiiy Allen (^ Scibcri Soi, S6g. 3. RuBUs TANTus Bailey, in Gent. Herb. 6:361. 1944. Upright and stout, canes pubescent but not long-hairy or shaggy, prickles many and recurved, not glandular; leaves somewhat pubescent on upper surface, gray-tomcntosc on under surface, closely sharply serrate, 3- to 5-folIolatc; leaflets ovate to ovate-elliptic, sometimes subcordate, 7—9 cm, long, about 4 cm. broad; petiole pubescent and prickly; inflorescence terminal and short, panicle-like, lateral clusters in upper axils; flowers many, crov/ded, small, pinkish-white; fruit small, with few drupelets. Panama. CHIRIQUI: vicinity of EI Boquctc, 1000-1300 m., Maxon 51OJ; pastures, Vifticr 312/. 4. Rur.us TiucHOMALLUs Schlccht. in Llnnaea 13:268. 1839. Vigorous, large, 2 m. high, the canes shaggy brown-hairy but not glandular, closely pubescent on surface, prickles remote and hooked; primocanc leaves many and large, pubcrulent on upper surface, grayish-tomcntosc on under surface, finely and unevenly sharp-serrate, 3- to 5-foliolate; leaflets narrowly ovatc-cUiptic- acuminate, 8—10 cm. long, 3-5 cm. broad, sometimes subcordate; petiole stout, hairy-pubescent, aculeate; inflorescence panicle-like and terminal with a few smaller clusters in upper axils, the peduncles brown-hairy and commonly prickly; flowers white, small, about 1 cm. across; fruit small, short-oblong, about 1 cm. long. Southern Mexico to Panama, but the identity of the Panama plant not yet clear. CHiRiQuf: Cerro Vaca, in forest or thickets, 900-1136 m. alt., Pitticr 572J; pastures around El Boquctc, 1000-13 00 m., Piffier 31 39; vicinity of El Boquctc, Maxon 4Q2/', COCLE: vicinity of El Valle, Allm uSg, 1647, 1776, 5. RuBUs GLAUCUS Bcnth. in Plant. Hartweg. 173. 1839. "Black Raspberry." Diffuse or erect, to 1.5 m. tall; primocanes smooth, glaucous, with uncinate scattered prickles 2-3 mm. long; leaves dull green, glabrous on upper surface, gray- tomcntose underneath, minutely serrate, 3-foliolate; leaflets elliptic-ovate to ovate- lanceolate, 6—10 cm. long, long-acuminate, petiole with hooked prickles; flowers mostly solitary, sometimes 2 or 3, In upper axils; corolla white sometimes tinged (142) t 1950] FLORA OF PANAMA (Rosaccae) 157 ? Fig. 66. Kubus pyaccipims (143) [Vol. 37 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN I •- I /* ^ Fig. 67. Rubi4s frichoffiallus (144) 1950] FLORA OF PANAMA (Kosaccae) 159 green, about 1.5 cm. broad; fruit oblong or thimble-sbaped, 1.5 cm. and more long, becoming black, of good quality, Panama, Colombia, Ecuador. CHiRiQui: Volcan de Chiriqui, Loma Larga to summit, 2500-3880 m. a!t., Woodson, Allen 6 Seibert io8i; vicinity of Cerro Punta, 1500-2000 m., Seihert 264; llanos dc volcan, about 1300 m., Allen 1546; El Boquete, 1000-3000 m., ?ittier 3133, 4. ALCHEMILLA L. Alchemilla L. Gen. PL cd. 5, 58. 1754; L. M. Perry in Contr. Gray Herb. 84:1-57. 1929. Alchemilla, sect. Lachcmilla Focke, in Engl. & Prantl, Nat. Pflanzenfam. 3^:43. 1894. Lachemilla (Focke) Rydb. in N. Am. Fl. 22:380. 1908. Perennial herbs, often prostrate or procumbent, often stolonifcrous. Leaves alternate, pctiolatc, usually palmatcly lobate or cleft, the upper smaller, with fewer lobes or divisions, sessile; stipules sheathing, usually foliaceous, lobcd or cleft, adnate to the petiole, the whole often appearing as a multifid perfoliate leaf or a many- lobcd sheath. Flowers minute, usually crowded into small axillary or terminal glomerulose cymes, less often in loose racemoid inflorescences; hypanthium per- sistent, urceolate, the 8—10 lobes 2-seriate, the outer ones smaller; petals none; stamens 1-4, free, opposite the calyx-lobes, the filaments short, usually not more than half the length of the calyx-lobes; disk with thickened margin, nearly closing the mouth of the hypanthium. Fruit of 1—6 achcnes, these sessile or stipitate, in- cluded in the hypanthium, with basal, ventral, ascending and persistent styles, and capitate or clavate stigmas. This group of species, whether it be treated as a genus, Lachcrnillay as done by Rydberg and more lately by Rothmaler (Fedde Rep. Sp. Nov. 42:164-173. 1937), or as a section or subgenus of Alchemilla, following Focke, Miss Perry, and m.ore recent American authors including Standley (Field Mus. Publ. Bot. 1^:477—47^. 1937), and Standley & Stcyermark (Fieldiana: Bot. 24:436-440. 1946), is easily recognized in the field, but the individual species arc difficult of interpretation. The flowers are very small (3—4 mm. long or less), and in consequence the species have been distinguished chiefly by the use of characters of leaf-shape and lobing, distribution and quality of pubescence, and other vegetative characters which are not always convincing. Miss Perry divided the group as a whole into six easily recognizable series, or species-groups, of which two, the Orbtculatae and the Aphanoides, are represented in Panama. a. Basal leaves 5- to 11-lobed or -cleft, the blades relatively broad and merely toothed, even those of the upper leaves much broader than and distinct from the stipules, b. Leaf-blades shallowly 5- to 11-lobed 1. A. orbiculata bb. Leaf-blades palmately 5- to 9-cleft to near their bases 2. A. pascuorum aa. Basal leaves 3- to 5-cleft or -parted, the blades with narrow finger-like lobes, those of the upper leaves often similar to and little exceeding the stipules 5. A. APHANomES (145) [Vol. 37 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Alchemilla orbiculata Ruiz & Pav. FI. Peruv. 1:68. 1798. Alchcmilla pcctinata HBK. Nov. Gen. & Sp. 6:226. 1824; Perry in Contr. Gray Herb. 84:14. 1929. LacbemiUa orbiculata (R. & P.) Rydb. in N. Am. Fl. 22:381. 1908. The stolons usually with well-developed leaves bearing offsets or floral shoots in their axils; basal and lower stem-leaves orbicular-reniform, 1.5—4 cm. broad, stiff, deeply cordate, the lobes serrate; blades green and glabrate above, appresscd- silky beneath, the surface beneath the silky hairs glaucous with minute waxy particles; petioles sericeous, 3-6 (-20) cm. long; stipules of the upper leaves fohaccous, forming a somewhat tubular sheath, deeply incised, the lobes not spread- ing nor recurving; inflorescence racemose, often terminal and 4-6 cm. long in Panamanian material seen; flowers relatively large, at maturity 3.5—4 mm. long, densely silky; achenes 2-4. Since the publication of Miss Perry's revision, American botanists generally have treated Alchcmilla pcctinata HBK. as a species distinct from the South American A. orbiculata^ w^hich differs in having greatly reduced or scale-like leaves on the stolons, sheathing (not merely amplexicaul) bracts subtending the flowers, and 1—2 (not 4-6) carpels. These differences do not seem to be greater than might be expected in geographically separated races of the same species, and in this case I prefer to accept the judgment of Rydberg, w^ho could usually be depended upon to find differences between species whenever such difl*crcnccs exi isted. Central Mexico to Bolivia, at the higher elevations, usually above 2000 meters, in clearings and on the paramos of the Andes. CHiRiQUi: Loma Larga to summit, Volcan dc Cliiriqui, Woodson, Allen & Scibcrf 1042; Potrero Mulcto to summit, Woodson ^ Scbcry 401, 4j8, 2. Alchemilla pascuorum Standi. Jour. Wash. Acad. 17:310. 1927. Lachcmilla pascjwnnn (Standi.) Rothm. in Fcdde Rep. Sp. Nov. 42:171. 1937. Basal and lower stem-leaves orbicular-rcniform, 2-4 cm. broad, the lobes cuneatc-obovate, rounded at apex, incised-dentate above the middle, green and sparingly pilose above, densely to sparsely appressed-silky beneath, the surface beneath the silky hairs glaucous with minute waxy particles; petioles 2 cm, long or less, apprcsscd-pilose; stipules of the upper leaves foliaceous, forming a short sheath, deeply Incised, the lobes spreading or recurving; inflorescences compact, in short axillary clusters or racemes 2-4 cm, long; flowers 2-2.5 (-3) mm. long, densely silky; achenes 2-4. Possibly not to be distinguished specifically from Alchcmilla vcnusta Schlecht. & Cham, (of Mexico), A. guatcmaJcnsis Rothm. (Guatemala), or Lachcmilla fnlicsccus (Perry) Rothm. (Colombia). Perry separates A, icnusta from A. pascuorum on the basis of its ^Villous, turbinate-urccolate" (not "sericeous, globosc- urceolate'*) hypanthium, and its erect to spreading (not erect to subconnivent as (146) 1950] FLORA OF PANAMA (Rosaceac) 161 Fig. 68, Alckcfnilla pascnornm in A. pasciiorinn) calyx-lobes. I find tKese characters difficult of interpretation and think the species rather too close together. As pointed out by Perry, Alchemilla pascnornm is also closely related to A. pecfinata (orhiculata) . All the Panamanian material I have been able to examine falls Into two rather well-defined groups, one with relatively large basal leaves often 2.5—3 cm. broad, and a distinct erect terminal racemose inflorescence 4-6 cm. long. The other group has the basal leaves somewhat more deeply divided (but not always to near the base), mostly 1.5—1.8 cm. broad, and the flowers In shorter clusters, 2-4 cm, long, from the axils of the well-developed stolons. The first group, with the larger leaves, undoubtedly represents AlcbcmiUa orhiculata. The second I have here called A, pascnornm^ although at least one of the specimens cited below as this species has been seen by Miss Perry and cited, with some doubt, as A. pcctinata {Woodson et al. 8g2, cited in Ann. Mo. Bot. Gard. 26:287. 1939). CHiRiQUi: Volcan de Chlriqui, Cerro Copcte, Allen 4858; Chlquero, Boqucte Distr., Davidson $46; Cerro Pando, valley of the upper Rio Chiriqui Vic jo, P. White 8; Volcan de Chiriqui, Casita Alta, Woodson^ Allen ^ Seibert 8g2. 3. Alchemilla aphanofdes L. f. Suppl. 129. 1781; Perry in Contr. Gray Herb. 84:36. 1929. AlcbcmiUa hirsuta var. alpestris Cham. & Schlecht. in Linnaea 5:572. 1830. Alchemilla stibalpcstris Rose, in Contr. U. S. Nat. Herb. 10:96. 1906. Lachemilla tripartita (R. & P.) Rydb. in N. Am. Fl. 22:383, 1908. Lachcmilla stibalpestris (Rose) Rydb. loc. cit. 384, 1908. Alchemilla aphanoides var. subalpcsfris (Rose) Perry in Contr. Gray Herb. 84:39. 1929, Lachemilla aphauoides (L. f.) Rothm. in Fedde Rep. Sp. Nov. 42:170. 1937. Stems erect to decumbent, subsimplc or branched, densely to sparsely pilose; leaves tripartite, the lower more densely pubescent than the upper, the segments oblanceolate to cuneate-obovate, multifid or incised-dentate; stipules foliaceous, 2- to 4-lobed or cleft, often with the sessile upper leaves forming many-lobcd or (147) 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 perfoliate "leaves"; flowers aggregated in small axillary and terminal cymes, usually very short-pcdicclhte; hypanthium glabrous or nearly so at least at maturity, the outer lobes about equalling the inner; flowers in Panamanian material less than 2 mm. long; achenes 1-3. Northwestern Mexico to Bolivia; mountains, often at elevations between 2000 and 3000 meters, in clearings and on paramos. The North American representatives of this species have lately been referred to var. siibalpi'stris, which has somewhat shorter flowers than the typical variety and IS somewhat less pubescent except for the hypanthium, which may be sparsely hairy in the Panamanian plant. Alchemilla apbafwiJes is very closely related to A. sihhaUIiacfolia HBK., but the Central American representatives of the latter are marked by rather strongly pubescent hypanthia, and I have not seen any such from Panama. ciiiRiQuf: Volcan de Chiriqui, Potrero Mulcto, Davidson 1040; Loma Lari;a to summit, Woodson, Allen 6 Scibert 1041; Potrero Muleto to summit, Woodson ^ Sc/jcry 41S. 5. ACAENA L. Acm:na Mutis, ex L. Mant. PL 145. 1771; Bitter, in BIbl. Bot. 74:1-336. 1910- 1911. Herbaceous or low suffrutescent plants usually with odd-pinnate leaves, toothed leaflets, and stipules more or less adnate to the petioles. Flowers in spikes or racemes, apetalous, the sepals and stamens variable in number (usually 4) ; pistil usually 1 with short terminal style and multifid stigma, at maturity forming an achene wholly inclosed in the prickly hypanthium. 1. AcAENA ELONGATA L. Mant. Pl. 200. 177L Acaena agriwontoides HBK. Nov. Gen. & Sp. 6:231, 1824. Plants shrubby at base, often with upright herbaceous branches, about 1 m. high or less, tlic bark on the older branches exfoliating in very thin reddish-brown sheets; plant ultimately glabrate, but juvenile stems and leaves vcstlte with loosely appressed silky straight yellowish-white hairs 0.7-1.5 mm, long. Leaflets often 13 (9-19 according to Bitter), the proximal ones much reduced In size, the 7 distal ones all alike (or the terminal one slightly larger), elliptic or oblong to obovate, often 5-8 mm. w^ide, 10-20 mm, long, acute at each end or obtuse at tips, the two edges very unequal at base, the proximal edge of at least the terminal pair dccurrent on the rachis, the petiolules mostly 1-2 mm. long; margins of leaflets rather coarsely crenate-serrate with 6-8 teeth on each edge, the teeth often pilose-tufted; stipules adherent to the rachis, scarious, often reddish-brown, con- nate at base and forming a sheath 1-2 mm. long about the stem, thence tapering gradually toward tips, the body 5-15 mm. long, the free herbaceous tips linear, entire, 2-6 mm. long; margins of stipules, rachis and sometimes axis of inflorescence with scattered stipitate glands; pubescence of leaf at maturity often reduced to (14^) ^-' 19 5 0] FLORA OF PANAMA (Rosaccae) 163 Fig. 69, Acacna elongafa tufts at bases of leaflets, on upper side. Flowers racemosely arranged on leafless axillary branches mostly 20-30 cm. long, relatively few (often 8-10) to a branch, each borne on a short lateral bracteate branchlet (up to 1.5 cm. long, or most often 1 mm. long or less) which is bibracteolate at summit and terminated by a pedicel 1-4 mm. long; calyx-lobes 4, green (contrasting with the reddish hypan- thium), ovate, acute, about 1 mm, wide and 1.5 mm. long, at anthesis crowning the flower and much exceeding the spines of the hypanthium, but at maturity Im- mersed in the spines; stamens 3-4, purplish, 1-2 mm. long, the anthers reniform, 0.5 (—0.8) mm, long and about 1.2 mm. wide after dehiscence; style very short, tipped by the expanded flattened peltate stigma which is about 1.5 mm. broad, bilateral, the divisions deeply lacerate; hypanthium in anthesis cyhndric, reddish, at maturity pendent, broadly ovoid or ellipsoid to globose, 3-5 mm. in diameter and 5-8 mm. long (exclusive of spines), the outer layer inflated, membranaceous, glabrous or puberulent; spines uniformly distributed, spreading, arising from the 10-12 ribs of the hypanthium (about 6 to each rib) which are concealed by the outer inflated layer; visible part of the spines 2-3 mm. long, purpUsh-red or brown, commonly with 3 retrorsely pointed barbs at apex. (149) [Vol. 37 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN Central Mexico to Guatemala; Costa Rica and Panama; Colombia and Ecuador; open meadows, thickets or wet forests, In highlands, mostly at elevations of 2000 meters or above, sometimes becoming a noxious weed. CH'RiQuf: valley of the upper Rio Chiriqul Viejo, P. White 62. 6. PRUNUS L. Prunus L. Gen. Pi. ed. 5, 213. 1754; Koehne in Engl. Bot. Jahrb. 52:279-333. 1915; Macbride in Tield Mus. Publ. Bot. 13-:1083-1090. 1938. Trees with alternate simple leaves. Flowers (in Panamanian species) white, racemose; calyx 5-lobed, the tube perigynous, cup-like, forming with the receptac- ular disk the hypanthium bearing the 15-20 stamens and 5 petals at its ma'gin; filaments free, filiform or somewhat dilated at base; carpel 1, with terminal style and peltate or truncate stigma; ovules 2, collateral Fruit a drupe, one-seeded, often with juicy pulp. A large genus, best known for the numerous species which are cultivated for food and for ornament. Segregate genera, e. g. Paclus and Laurocerasiis, are main- tained by some botanists but are based chiefly on characters of the Inflorescence and are probably best included in Pnnms. The native Panamanian species belongs to the Section Laurocerasus as defined by Koehne In 1915. This author included 47 species In the section, 42 of them American. At the time of his revisionary studies Koehne was unable, because of lack of material, to Integrate the results of his studies in the several distinct floristic areas in tropical and subtropical America, and it is probable that the actual number of species will prove smaller than his estimate. In the area including Mexico and Central America Koehne recognized 7 species, of which he described 5 as new. The characters used by him to dis- tinguish species in the group to which the Panamanian plant apparently belongs (I. e. the species having entire leaves, glabrous petals, solitary racemes, and super- ficial glands near the base of the leaf blade), appear to be of little value for this purpose. The number of glands on the leaf-blade, which was assumed by Koehne to be constant for a species, varies from to 3 on either side of the midrib of the same leaf, and from to 4 (rarely 6) on different leaves on the same plant. The unusual development of lentlcels and the transverse cracks in the branchlets, on which he based Prunns tnbcrciilata and P. annnJari^, respectively, do not appear to be constant characters. After study of somewhat more material than was available to Koehne, I doubt that more than one species exists In this group, at least in Cen- tral America. The oldest name which is applicable to the species Is Primus hracby- hotrya Zucc, (1837), from Mexico, but pending a revision of the whole group It seems wisest at present to refer the Panamanian plant to a species orlglnallv de- scribed from nearby Costa Rica, Prufuis annidar/s. (1^0) 1930J FLORA OF PANAMA (Rosaceac) 165 Fig. 70. Prntnis annularis 1, Prunus annularis Kochne in EngL Bot. Jahrb. 52:308. 1915. Large evergreen tree, up to 20 m. tall with trunk up to 50 cm. In diameter, entirely glabrous. Leaves coriaceous, entire, elliptic to lanceolate or ovate, 2.5-5.5 cm. wide, 6-13 cm. long, usually 2-2,5 times as long as wide, rounded at base, narrowed toward the apex from about the middle, with a short blunt acumen; petiole stout, 1-1.5 mm. in diameter, 1-1.5 cm. long; lateral veins often 5-7 pairs, inconspicuous, slender, arcuate-ascending, anastomosing well inside the mar- gin; glands usually present on the lower surface of the blade near base, round or oval, often about 0.5 mm. wide and 0.7 mm. long, 1—2 (rarely none, or 3) on each side of the midrib, mostly 5—10 mm, from the base of the blade and about 5 mm. from the margin, but sometimes as little as 1-2 mm. from base or margin; stipules very soon deciduous, not seen, probably narrowly linear, 2-4 mm. long. Flowers in leafless racemes 4—5.5 (-7) cm. long, these from the axils of the sub- persistent leaves of the preceding season; flowers 20-40, subtended by very small chartaceous bracts which are deciduous before the flowers open; pedicels naked, 3-7 mm. long (those at the base of the racemes the longest), 0.5-0.7 mm. in (lU) [Vol. 37 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN diameter at antKcsis, up to 1.5 mm. in diameter in fruit, mostly at right angles to the axis; petals white, glabrous, entire, rounded or reniform, about 2 mm. long including the short claw, 2-2.5 mm. wide; stamens about 20, glabrous, the fila- ments 2,5—4 mm. long, subulate; anthers (0.6?—) 0.7—1 mm. long, elliptic; hypanthium in anthesis campanulate, about 3 mm. broad and high, the calyx-lobes broadly triangular, blunt or acute, about L5 mm. wide at base, 1 mm. long, en- tire; hypanthium and calyx deciduous Immediately after anthesis; ovary glabrous, about L5 mm. long, tipped by the straight style 2-3 mm. long and 0.3 mm. in diameter; stigma peltate, 0.7-0.9 mm. across. Fruit probably a purple ovoid drupe with thin flesh, about 1.5 cm. in diameter, 2 cm. long; stone ovoid, pointed, about 10 mm. In diameter, 13 mm. long; seed globose, about 9 mm. In diameter. Costa Rica and Panama; related species or perhaps the same species from Vera Cruz and Chiapas through Central America into northern South America; chiefly in rain forests, at elevations from 1000 to 2000 meters. CHiRTQUi: Bajo Chorro, Boquete DIstr., DariJsov 2Jj; between El Volcan and Ccrro Punta, G. Wbife 14; vicinity of Casita Alta, Volcan dc Chirlqui, WooJson, Allen ^ Scibert 907. Pniniis occidcntalh Sw., reported from the Volcan de Chiriqui by Seemann (Bot. Voy. Herald, 119, 1852-53), is a West Indian species not known to occur in Panama; the report was doubtless based on what is here called Pruuns amnilarh, VrufiHs coYfiifolia Koehne, a Costa Rican species recognized by having a persistent calyx and tufts of hairs in the axils of the main foliar veins (Sect. Neocalycinia Koehne), was erroneously reported from Panama (Ann. Mo. Bot. Card. 27:312. 1940), but its known range is from southern Mexico to Costa Rica only. The common half-wild cherry of the highlands of Guatemala and southern Mexico, the capidin, 'Prnuns capuli Cav., has apparently not been reported from Panama, although it is widely naturalized from Venezuela to Bolivia. It may be recognized by the racemose inflorescence which bears leaves near Its base, the crenate-dentate leaves bearing reddish hairs beneath along the midvein, the persistent calyx and the sweet nearly black juicy fruit 1-1.5 cm. in diameter. 7. CHRYSOBALANUS L. CiiRYSOBALANUs L. Gen. PL ed. 5, 229. 1754. A small (possibly monotypic) genus of shrubby plants with entire coriaceous leaves. Flowers white, in axillary cymes; calyx (hypanthium) perigynous, the lobes 5; petals 5; stamens about 20, hirsute, coherent; ovary sessile at the bottom of the hypanthium; ovules 2; style basal. Fruit a drupe with one bony stone having 5-6 angles. 1. CuRYsoBALANUS ICACO L. Sp. Pi. 513. 1753; Hook. f. in Mart. Fl. Bras. 14-:7. 1867. Low shrub 1-2 m. high, or prostrate, or a medium-sized tree up to 5-6 m. high, glabrous or essentially so except in the inflorescence and on very young (1^2) 1950] FLORA OF PANAMA (KoSUCeac) 167 Fig. 71. Chrysobalanus Icaco growth, where strigose; bark of the branchlets reddish-brown, smooth except for the very numerous and conspicuous pale lenticels. Leaves varying considerably from one plant to another, elliptic to obovate or suborbicular, rounded or emarginate at apex or when elliptic narrowed to an obtuse apex, cuneatc to acute or rounded at base, up to about 8 cm. long and 6 cm. wide, (1-) 1.5-2 times as long as wide, on stout petioles 2-4 mm. long; blades dark green and lustrous above, dull beneath and usually with a depressed elliptic glandular area about 0.5 mm. long on each side of the midrib at extreme base; small veins prominently reticulate en both sides of the blade, the primary lateral veins mostly 4-6 pairs, straight at base but curving and anastomosing before reaching the margins. Flowers white, in short-pcduncled axillary cymes shorter than the leaves (mostly 2-4 cm. long, with 8-20 flowers), the cyme-branches often strongly flattened and 1-1.5 mm. wide, often glabrcscent near the b ase, the tips densely pale-strigosc like the hypanthium, the flowers and the tips of the cyme-branches appearing silvery white to the unaided eye; cymes usually naked In anthesis, the ovate, acute, finely glandular-ciliate bracts 1.5-2 mm. long but very soon deciduous; petals 5, glabrous, oblanceolate or spatulate, 3-5 mm. long, 2-3 times as long as wide, the tips rounded, often erose, the bases cuneate; stamens about 20 (up to 30, according to Hooker) in one series, the filaments pale (Pwhite), about 5 mm. long or less, hirsute on the inner surface except on the distal third, the flattened bases coherent a third or half the length of the filaments into a cyHnder; anthers about 0.5 mm. long, Ppink or Ppurplish; hypanthium campanulate or turbinate, at anthesis about 3 mm, long and wide, the inner surface hairy, the long-hirsute ovary sessile at the bottom of the tube, the fiUform style 6-7 mm. long, hirsute except at tip, erect, with a minute terminal stigma; calyx-lobes triangular-ovate, acute or blunt- tipped, heavily pubescent, the margins glandular-ciliate but otherwise entire. Fruit globose or oval, 2-5 cm. long, white to pink or purple, edible, with white juicy insipid flesh, the stone 1-2 cm. long with 5-6 acute longitudinal ribs. (153) [Vol. 37 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN Tropical America, from Florida and Tamaulipas southward through the West Indies and Central America to northern Brazil and to Ecuador; tropical West Africa; near sea level, in beach thickets and coastal swamps; often planted inland. BOCAs DEL TORO: Nances Cay, vicinity of Chiriqui Lagoon, von WeJel 2gj6. canal zone: Aspinwall, Hayes 6j6; Chagres, Fcndler lOJ, cocLE: Penonome, WHIictms 4^0, colon: Manzanilla Island, Hayes yog. paxNAMA: San Jose Island, [Johnston, Erlanson], 8. LICANIA Aublet LiCANiA Aubl. Hist. Pi. Guian. Fr. 119, pL 45. 1775; Hook, f, in Mart. Fl. Bras. 14-:8-19. 1867; Fritsch, in Ann. K. K. Naturhist. Hofmus. Wien 4:33-60. 1889. Moquilea Aubl. loc. cit. 521, pi 20S, 1775; Hook. f. In Mart. Fl. Bras. 14-:19-26. 1867; Fockc in Engl. & Prantl, Nat. Pflanzcnfam. 3^:58. 1894. Trees with alternate simple entire leaves. Flowers usually paniculate, the ultimate branchlets cymoscly branched; hypanthium globose or campanulate, the lobes 5; petals 5, minute or none; stamens 3 to many, inserted at the margin of the hypanthial disk, often connate at base, forming a complete ring or unilaterally disposed; ovary unilocular, sessile at the bottom of the hypanthium, hairy; style basal; fruit drupaceous, various, one-seeded, the pericarp usually leathery or woody. a. Leaves tomcntose beneath with fine white or nearly wlilte appressed tomentum. b. Leaves broadly rounded at apex, obtuse or (usually) subcordate at base, mostly 1.4-1.7 times as lon>; as wide; branchlets stout, often 3-6 mm. thick at base of Inflorescence; stamens about 20 1. L. arborea bb. Leaves acute or acuminate at apex, acute to rounded at base, mostly 2—2.7 times as long as wide; branchlets at base of inflorescence slender, 1—2 mm. thick; stamens 3 (sometimes in addition to a few vestigial rudiments) 2. L. hypolhuca aa. Leaves glabrous or essentially so 3. L. platypus 1. LicANiA ARiiOREA Seem. Bot. Voy. Herald, 118, pL 2^. 1852-53, Ucania Selerrana Loes. Verb. Bot. Ver. Brnndcnb. 53:5 5. 1911. Tree to 10 m. tall (or sometuiies up to 3 m., according to Standley) with broad stiff blunt, oval or oblong leaves; juvenile foliage, branchlets and inflorescence conspicuously yellowed, very densely hlrsutulous with short erect or somewhat matted tawny hairs; branches and upper surface of mature leaves glabrate, the lower leaf-surfaces strongly whitened with a closely felted tomentum; branchlets stout, often 3-6 mm, in diameter at the base of the inflorescence, the bark reddish- brown with inconspicuous pale lenticels. Leaf -blades coriaceous, ovate to elliptic or oblong, broadly rounded at apex, obtuse or (usually) subcordate at base, 5-12 (—15) cm. wide, 6-18 (-25) cm. long, usually 1.4-1.7 times as long as wide, lustrous and glabrous above; lateral veins about 12-15 pairs, widely divergent, the 3—5 proximal pairs much closer together than the others, all raised and forming conspicuous ribs on the lower surface; petioles stout, 5-10 mm. long, 2-3 mm. in diameter; stipules 1-1.5 cm. long, 1—2 cm. wide, linear, acute, appressed, on shoots (154) 1950] FLORA OF PANAMA (Kosaceac) 169 only. Flowers essentially sessile, very many together in axillary and terminal panicles at the tips of branches, forming an inflorescence often up to 40-50 cm. long and nearly as broad, with widely divergent branches; petals white, hirsute, oblanccolate or obovate, often about 1.5-2 mm, long and half as wide, obtusely pointed at apex, cuneate at base; stamens about 20 (the anther-bearing ones 10 or fewer), the filaments up to about 3 mm. long, hirsute, their bases coherent into a cyHnder and inserted with the calyx, the sterile ones often short and inconspic- uous; anthers about 0.5 mm. long; hypanthium deeply cup-shaped, about 1.5-2 mm. wide and high in anthesis, tomentose within, usually contracted toward apex and surmounted by the 5 erect or converging calyx-lobes which are triangular, acute, about 1-1.5 mm, wide and long; ovary sessile at the bottom of the hypan- thium, nearly globose, about 1 mm. long, hirsute; style about 2.5 mm, long, subulate, hirsute except for the distal 0.5 mm. which is slender, terete, and glabrous; stigma terminal, slightly flattened. Fruit (said by Seemann to be black) obovoid or oblong-obovoid, 2-3 cm. long or larger, rounded at tip, substipitate, glabrate. Western Mexico (Guerrero) to Panama, lowland, often in dry 1 Colombia. ru CANAL zone: Chiva-Chiva trail, Red Tank to Pueblo Nucvo, Piper 57I3y 5738; old Las Cruces trail between Fort Clayton and Corozal, Standley 2go8g. cocle: north rim of El Valle de Anton, Allen 174I; Pcnonome and vicinity, Williams 3j8. chiriqui: David, Pittier 2841. Panama: Sabanas, Bro, Paul 2g0; Corozal road near Panama, Standley 26847; Matias Hernandez, Standley 28gj2; between Matias Hernandez and Juan Diaz, Standley 32007. The large seed of this species is said to contain about 30 percent of oil, and to burn readily. The oil is similar to that produced commercially from the seeds of a closely related species, Licania rigida Benth., of Brazil. The type specimen of Licania arborea was collected by Seemann between Tole and David, in what is now the Province of Chiriqui. 2. Licania hypoleuca Benth. Bot. Voy. Sulphur, 91, pi. 32, 1844. Tree 10-15 m. tall, with a trunk-diameter of 10-40 cm., the ovate or lanceolate pointed leaves whitened beneath, the branchlets slender (1-2 mm. thick at the base of the inflorescences), nearly black, glabrate (in the juvenile state densely sordid- puberulent). Leaves ovate to lanceolate or elliptic, the upper surface lustrous and glabrous but the veins and vcinlets in fully mature leaves marked with lines of tiny closely set papillae, the lower surface (appearing glaucous to the unaided eye because of the fineness of the tomentose layer) whitened with a dense appressed tomcntum, the main lateral veins densely puberulent or thinly tomentose, arcuate, 6-S pairs, prominent beneath, inconspicuous above (in dried material) ; blades acute to acumi- nate at tips, acute to rounded at base, 5-12 cm. long, 2-5 cm. wide, usually about 2-2.7 times as long as wide; petioles stout, about 1 mm. in diameter and 4-7 mm. long, densely short-hairy, the hairs on the upper side extending 2-4 mm. in a nar- row inverted V onto the base of the blade; stipules subulate, 1-3 mm. long, subpcr- (155) [Vol. 37 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN sistcnt, erect and parallel to the branchlets. Flowers (said to be white) very small, numerous in leafy, terminal, mostly pyramidal panicles 10-20 cm. long; lower (axillary) branches of the panicle widely divergent, often 6-15 cm. long, most of the flowers borne in racemold arrangement along secondary branches in 2- to 5-fIowcred cymules on peduncles 2-5 mm. long; terminal flowers of the cymules often pedicellate 1-3 mm., the lateral ones usually nearly sessile; corolla none; anther-bearing stamens usually 3 (opposite 3 adjoining calyx-lobes), less than 1 mm. long, the filaments subulate, dilated at base and hirsute about the zone of insertion, the anthers about half as long as the filaments; hypanthium campanulate or turbinate, about 1.5-2 mm. long and wide in anthesis, often contracted slightly at mouth, surmounted by the 5 ovate-acute, entire calyx-lobes which are spreading at tips, about 1 mm. long and wide; receptacle thickly hairy within, with a prom- inent fringe of hairs at the somewhat thickened margin, which in addition often bears short sterile filaments alternating with the calyx-lobes, and less often bears very short vestigial filaments elsewhere; ovary sessile at the bottom of the hypan- thium, ovoid-globose, less than 1 mm. long, white-hirsute; style erect (in the flower opposite the stamens and so alternating with the stamenless calyx-lobes), about 2 mm. long, subulate, hirsute except for the slender tip 0.2-0.3 mm. long; developing ovary elongating at base and becoming stipitate at maturity. Fruit pyriform, red, thinly pubescent at least at base, about 1 cm. thick and 2 cm. long (including the attenuate base). Tabasco and British Honduras to Panama, lowland, in wet or moist forests; Colombia? The TYPE of this species came from "Veragua", according to Bentham's original description; in Seemann's "Flora of the Isthmus of Panama", published a decade after Bentham's description, the locality was given as "Province of Vera- guas", the implication being that no more definite locality could be given. At this period Veraguas Included all of what is now the Province of Chiriqui, so the type-locality of this species may have been almost anywhere along the Pacific Coast of western Panama. The species is reported by Standley (Contr. U. S. Nat. Herb. 27:189. 1928) from Barro Colorado Island, with the notation that this is the only locality at which it is known to occur in the Canal Zone. CANAL zone: Barro Colorado Island, Standley 31323, 31416, 4106^. darien: La Palma, Vifticr 549I. 3. LicANiA PLATYPUS (Hemsl.) Fritsch In Ann. K. K. Naturhist. Hofmus. Wien 4:53. 1889; Pittler in Contr. U. S. Nat. Herb. 13:443, pi S6. 1912; Standi. & Steyerm. in Fieldlana Bot. 24^:456. 1946. Moquiha platypus HemsL Diagn. Pi. 1:9. 1878; Biol. Centr.-Am. Bot. 1:366. 1880. Tree up to 50 m. high with large, narrow, glabrous, somewhat distichous leaves; branchlets stout, glabrous, purplish or red, often 2-3 mm. in diameter at the base of the inflorescence, the lenticels numerous, narrowly elliptic, 0.5 mm. long; leaf-blades coriaceous, elliptic-lanceolate or narrowly oblong, rounded grad- (156) 1950] FLORA OF PANAMA (KoSUCCae) 171 Fig. 72, Licania platypus ually to a short-acuminate tip, rounded or acute at base, 3-6 (-8) cm. wide, 10-30 cm. long, usually 3 times as long as wide, green and somewhat lustrous above, paler and somewhat glaucous beneath; midvein raised and rather prominent beneath; lateral veins 12—20 (-25) pairs, arcuate, anastomosing near the margin; lower surface of the blade bearing small oval or circular glands 0.2-0.3 mm. across, these 0.2—1.5 mm. from the margin, slightly sunken, rimmed, one near the end of each main lateral vein; petioles dark red (when dry), 9-15 mm. long, 2 mm. in diameter; stipules dark, erect, stiff and persistent, 2.5 mm. long, adnate to the petiole three-fourths their length, the tip free, rounded. Flowers white, fragrant, rnz; [Vol. 37 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN subsessilc or on pedicels 1-2 mm. long, very many together in axillary and terminal cano-tomentose panicles at the tips of branches, these forming a divaricately branching inflorescence 10-3 5 cm, long; panicle branches strongly flattened near base, bractlcss at flowering time; petals obovatc, 2-3 mm. long, finely white- tomcntulosc distally (often said to be ciliate) ; stamens 15 (-18), glabrous, sub- equal, the filaments up to 3 mm. long, inserted separately on the margin of the thickened disk, which is heavily retrorsely bearded; anthers about 0.5 mm, long; hypanthium cup-shaped, about 1.5 mm. high and 2 mm. wide in anthcsis, tomcntu- losc without, silky within^ the erect or spreading calyx-lobes obtusely triangular with rounded sides, about 1.3 mm. long and wide; ovary sessile at the bottom of the hypanthium, nearly globose, about 1 mm. long, hirsute; style 5-6 mm. long, subulate, hirsute except for the distal one-third which is terete, filiform. Fruit (according to Pittier) very large, 1-3 to each panicle, 15-20 cm. long, 10-14 cm. in diameter, with dark brown, vcrrucose skin covered with white Icnticels, and the inside flesh yellow, juicy, and sweet, somewhat fibrous; seed usually one, ovate- oblong, flattened, 6-8 cm. long, 4-4.5 cm. in largest diameter. Southern Mexico to Panama; reported from Colombia. In lowland forests, often at elevations less than 400 meters, often cultivated for ornament, PANAMA: along Rio Juan Diaz above Juan Diaz, Allen p^i". This species is reported from San Jose Island by Johnston (Sargcntia 8:132. 1949), who gives at the same time a good descriptive account of the tree as it occurs in the forest on the island. The native country of this tree is somewhat in doubt; Pittier remarks that he has never seen it in a truly wild state. Hemslcy's original material included a cultivated specimen from Nicaragua (designated as the type-specimen in Contr. U. S. Nat. Herb. 23:344. 1922), and Cuming's No. 12^2, originally supposed to have come from Panama (i. e. probably western Panama) . As Hemsley later pointed out (Biol. Centr.-Am. Bot. 1:3 66. 1880), "Cuming's specimens may have been collected further south, in Western Colombia." (i.e. probably what is now eastern Panama) . 9. HIRTELLA L. HiRTELLA L. Gen. PL ed. 5, 20. 1754; Hook. f. In Mart. Fl. Bras. 14-:27-40. 1867; StandL & Steyerm., in Ficldiana, Bot. 24^:450-452. 1946. Shrubs or trees with alternate, simple, entire leaves. Flowers in axillary and terminal panicles or racemes; hypanthium short and symmetrical or elongated and then usually gibbous, the lobes 5, rcflexcd; petals 5; stamens 3-7 (rarely more), inserted at the summit of the hypanthium disk, usually unilaterally inserted (i. e, on the side where the ovary Is attached, opposite the style and the enlarged side of the hjpanthium) ; uvary unilocular, hirsute, attached to the side of the hypan- thium-tube above the base (i. e. below the base of the middle stamen) ; style basal. Fruit drupaceous, often dry. (1>8) 1950] FLORA OF PANAMA (Rosaccac) 173 a. Inflorescence racemold, the 3 0-40 flowers on unbranched filiform bracteolate pedicels 5-8 (-12) mm. long; stamens 5-7 2. H. RACEMOSA aa. Inflorescence a leafless spike-like panicle, the lateral branches (cymes) often 1.5 cm. long, the spikes sometimes appearing racemoid when most of the cymes are reduced to 2 or even 1 flower each; stamens 3. b. Leaves glabrate or the principal veins with persistent apprcsscd yel- lowish hairs; bracts of the cymes lanceolate, attenuate, minutely glandular-denticulate 3. H. triandra bb. Leaves sparsely or densely short-hirsute, at least on the veins, with erect rufous or tawny hairs; bracts of the cymes round or oval, heavily beset with depressed or stalked and peltate glands 0.3-1 mm. ^^""OSS 1. H. AMERICANA 1. HiRTELLA AMERICANA L. Sp. Pi. 34. 1753; Standi. & Steyerm. in Fieldiana, Bot. 24:450. 1946, Hirtclla molUcoma HBK. Nov. Gen. & Sp. 7:263. 1825; Hemsl. In Biol. Ccntr.-Am. Bot. 1:366. 1880, as mollhsima; Standi, in Contr. U. S. Nat. Herb. 27:188. 1928. ?Hirtella glaiulidosa Spreng. Anlcit., />/. /, /. 1-4; Neuc. Entd. 1:303. 1820; DC. Prodr. 2:528. 1825; Seem. Bot. Voy. Herald, 118. 1852-53. Hirtclla guatcmalcnsh Standi, in Trop. Woods 11:19. 1927. Small or medium-sized tree, the branchlcts and inflorescences conspicuously rufous, densely short-hirsute with straight, slender, multicellular tawny hairs 0.5-0.7 mm. long; branchlcts 2-4 mm, in diameter below the inflorescence, the hairs persistent. Leaves hairy Kke the stems but less densely, the upper surface glabrate in age except the midrib, the lower surface densely hairy on the principal veins and uniformly but sparsely so on the anastomosing vclnlets, where the hairs are often about 3 times as long as the distance between them; hairs of the leaves with enlarged papillose bases; leaf-blades thick and subcoriaceous, oblong or elliptic, acute and gradually short-acuminate at apex, rather abruptly rounded or obtuse at base, 3-6 cm. wide, 6-15 cm. long, often 2-2.7 times as long as wide; lateral veins 8-12 pairs, arcuate, slightly impressed above, raised beneath, the small veinlcts finely reticulate and often raised on the upper surface, making it rough to the touch; glands abundant on the lower surface, usually concentrated near base and apex of the blade, elliptic or circular, 0.2-0.5 mm. across, with raised rim and depressed center; petioles very stout, 2-4 mm. long and about 2 mm. thick, their surfaces concealed by the hairs; stipules subpersistent, hirsute, subulate, apprcssed, 6-9 mm. long. Flowers many, white or purphsh-white, in very narrow spike-like leafless panicles 10-20 cm. long and 2-4 cm, thick, the lateral branches of the spike often about 1.5 cm. long, ascending, cymosely 1- or few-flowered; bracts of the cymes round or oval, 1-1.5 mm. long, heavily beset with large glands, these varying from depressed spots 0.3 mm. across to peltate glands 1 mm, across on stalks 1 mm. long, the larger glands often very numerous and conspicuous in the spikes after the abortion of most of the flowers; bracts subtending the lateral cymes (i. e. those of the main axis) eglandular or nearly so, hirsute, subulate, about 5 mm. long; petals white, broadly elliptic, bluntly rounded at base and apex, glabrous, about 4 mm. long and 3 mm. wide; stamens 3, opposite adjoining sepals, much exserted and curled into the flower, purplish or deep red, glabrous, the fila- (159) l\'oL. ^7 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN merits 1-L2 cm. long, flcsKy at base and tapering to tKe very slender tips, arising separately from the hypanthium-rim which is fleshy and projects about 1 mm. or a little less above the base of the calyx; anthers about 0.6 mm. long; hypanthium turbinate, the cavity funnelform, about 2 mm. deep measured from the summit of the fleshy rim, retrorsely hairy within the rim but glabrous at bottom; calyx- lobes broadly oblong, obtuse to rounded at tips, reflexed at and after anthesis, entire, 1.5-2.2 mm. wide, 2-2.7 mm. long, the outer surface thinly yellow-hirsute like the inflorescence, the inner surface pale, sordid-tomentulose; ovary hairy, sessile, attached above the middle of the hypanthium cavity (at the base of the middle stamen), the style about as long as the stamens and opposite them in the flower, filiform, long-pilose below the middle, with tiny capitate stigma, in bud bent abruptly away from the ovary and then coiled toward it. Fruit oval^ thinly yellow-hairy, rounded at apex, probably 1-1.5 cm. long or longer, black. Cuba; British Honduras, Costa Rica, and Panama; Colombia and northern Venezuela; thickets and savannas, lowlands. A distinct entity, but probably the same species, occurs in Guatemala and southern Mexico; the pubescence in general is toward yellowish rather than reddish, the calyx-lobes are sordid-tomentulose and pale on the outer surface as well as the inner, and the glands are usually absent from the lower leaf-surfaces but arc usually present on the upper. CANAL zonh: Gatuncillo, Piper 36541 between Rio Grande and Pedro Vidal, road to Arraijnn, Pitticr 2/oS; [Cruccs, Seerfiafm]; southwest of Pt. Salud, Barro Colorado Island, Wooduvrfh d Vestal 728, cocle: Penonome, Willia?rts 24/. Panama: Chorrcra, KiHip 34OQ; Pacora, Bro. Paul 2S4; Sabanas, north of Panama, Bro, Paul 412; San Jose Island, Erlanwn loS, 2. HiRTULLA racemosa Lam. Encycl. Meth. 3:133. 1789; DC. Prodr. 2:529. 1825; Seem, in Bot. Voy. Herald, 119. 1852-53; Standi. & Steyerm. in Field- iana, Bot. 24:450-452. 1946. Uhfclla amcricana Aubl. Hist. Pi. Guian. Fr. 247, /;/. 98. 1775; Hook. f. in Mart. Fl. Bras. 14-:33. 1867; Bcnth. in Bot. Voy. Sulphur, 91. 1844; Standi, in Contr. U. S. Nat. Herb. 27:188. 1928; auct. Amcr. plur., non L. llirtclla ohlofis^ifolia DC. Prodr. 2:529. 1825. Shrub or small tree, 2-3 (—6) m. tall, with slender pubescent branchlets 1-1.5 mm. In diameter below the inflorescence; hairs of two types, those of the young foliage and branchlets yellowish, appressed or erect, straight and slender, pointed^ multicellular, 0.5-1.5 mm. long, these also abundant In the inflorescence but there mixed with tiny erect white hairs mostly 0.2—0.3 mm. long. Leaves elliptic to oblong, lanceolate, or ovate, glabrate, only a few long yellow hairs persistent on the veins (at least of the lower surface), a few of the hairs with papillose bases; blades subcoriaceous, lustrous, 1.5-4 cm. wide, 3-8 cm. long, usually 2-3 times as long as wide, narrowed from the middle or above to a broad blunt or pointed acumen 1 cm. long or less, narrowed and rounded at base (or obtusely narrowed, with the extreme base rounded abruptly into the petiole) ; petiole short, stout, about 0.7 (160) 1950] FLORA OF PANAMA (Rosaceuc) 175 mm. in diameter, 2 mm. long; lateral veins 6-8 pairs, arcuate, slightly raised on both surfaces but not forming conspicuous ribs on either, often hardly visible on the upper; glands occurring on the lower surface of the blade near base, mostly nearly circular, about 0.2 mm. across, with raised rim and depressed center; stipules subpersistent, filiform-subulate, appressed-hairy, 2.5—5 mm. long, usually black in dried material, apprcssed to the branchlets. Flowers in axillary and terminal racemes 12—15 cm. long (or up to 20—25 cm. including the basal leafy part of the growing branchlet terminated by the raceme); "racemes" 3 0- to 40-flowered, flowering from base to apex but apparently always determinate, the terminal (bractless) flower abortive; lateral flowers on filiform ''pedicels" 5—8 (-12) mm. long, spreading at right angles or somewhat ascending, each with two tiny bracte- oles near base or about one-third the distance from base to apex, the bracteoles often with sessile or stalked glands on the margins, and borne at the summit of a thickened peduncular portion of the flower-stalk (only the part beyond them bein the pedicel proper) ; bracts subtending the flower-stalks lanceolate, usually re- curved, 1—2 mm. long, with basal marginal glands like those of the leaves (and sometimes a similar gland at apex) ; petals pink, purple or ''rosy mauve", glabrous, broadly elliptic, rounded at both ends, 2—3 mm. wide, 3—5 mm. long; stamens 5—7, much exserted, purple, glabrous, the filaments 1.2-1.6 cm. long, fleshy at base and tapering to the very slender tips, arising separately from the fleshy hypanthium-rim which projects 1 mm, or less above the base of the calyx, absent from the segment of the rim opposite the insertion of the ovary or there represented by vestigial staminodia only; anthers about 0.6 mm. long; hypanthium In anthcsis elongate- tubular, with hollow cylindric or oblong one-sided base up to about 1 mm. thick and 2 mm. long, and funnclform throat about 1 mm. long, the interior hairy about the summit of the throat, glabrous below; calyx-lobes oblong or elliptic, usually reflexed at anthesis, entire, rounded or subacute at apex, about 1-1,5 mm. wide, 2.5 mm. long, the outer surface with mixed short white and long yellow hairs, the inner surface glabrous proximally, distally whitc-tomentulose; ovary hirsute, sessile, attached to one side of the funnelform portion of the hypanthium above the bottom; style as in H. aviericanay but up to about 1.5 cm. long, arising from the flower toward the side where functional stamens are lacking (i. e. the side toward which the inflated base of the hypanthium is expanded). Fruit oblong- obovoid, sparsely hairy, dark red or purple, up to about 1.5 cm. long and 0.6 cm. in diameter, rounded at apex, the base substipltate. Guerrero and British Honduras to Panama, northern South America, and south in the Amazon Basin to central Brazil, eastern Peru and eastern Bolivia; lowlands, in forests and thickets. CANAL zone: Summit Road, D. G. Jories 2'^^; Ancon Hill, Piper 5575, 5596; Pittier 20/5; between Miraflores and Corozal, Pittier 2102; between Corozal and Ancon, Pittier 264I; road to Arraijan, between Rio Grande and Pedro Vidal, Pittier 2702; Ancon Hill, Staudley 26^66; Corozal, Standley 2/^/1; Gamboa, Staynlley 28^70; Summit, Standley JOIO/; Obispo, Standley JT742; Rio Chagres, Steyermark & Allen IJ500; Barro Colorado Island, Wetmore & Abbe 105; Woodxvorth ^ Vestal 5J2. cocle: Penonome, il. S. (161) 176 [Vol. Z7 ANNALS OF THE MISSOURI BOTANICAL GARDEN PKOVINCH UNKNOWN: Williams gj. colon: vicinity of Santa Rosa, Rio Gatuncillo, Allen 41 j6, darien: La Palma, Vittier S486. Panama: Rio Las Lajas, Alloi 160J; Las Sabanas, Killip 3361 ; MacbriJe 264S; La Chorrcra, Bro, Maurice 816; Isia Taboga, Miller 20jj; SianiUey 27036, 2S0IJ; Tumba Mucrto road near Panama, Standley 2gY36, "Panama", Duchassaing in 1850. According to Johnston (Sargentia 8:131-132. 1949), the true Hirtclla race- niosa is a South American species with larger flowers than the Panamanian and Central American species, which he refers to H. oblongifolia DC. Under the name of H, oblougifoUay he reports the species as a very common shrub on San Jose Island, 3. HiRTELLA TRiANDRA Sw. Prodr. Veg. Ind. Occ. 51. 178 8. Large shrub or a tree, often 5-6 m. high (sometimes to 20 m. with a trunk 15-20 cm. in diameter), with slender pubescent branchlets 1-2 mm. in diameter below the inflorescences; inflorescences and juvenile foliage and branchlets more or less appressed-pubesccnt with two types of hairs intermingled: straight, slender, pointed, yellow or tawny, usually appresscd hairs 0.5-1 mm. long, and straight or crisped often erect white hairs 0.3 mm. long or less; the longer, yellow hairs pre- dominating on the leaves, while in the inflorescence the two types arc often equally Intermingled except on the inner surfaces of the calyx-lobes, where the shorter, white hairs alone occur. Leaves elliptic or sometimes oblong, glabrate or the principal veins persistently hairy, the hairs of the leaves often with enlarged papillose bases; blades rather thin, 2-5 cm. wide, 6-12 (-15) cm. long, 2-3 times Fig. 73. Hirtclla trlanJra (162) > \ h t \ \ 1950] FLORA OF PANAMA (Rosaceac) 177 as long as wide, rather abruptly acute at apex and with a definite narrow sharp- pointed acumen 0.6-2 cm. long; base of blades acutely narrowed from below the middle, the extreme base not decurrent but often abruptly rounded and so truncate or even subcordate, and sharply delimited from the stout petiole 1-2 mm. thick and 2-4 mm. long; lateral veins 5-8 pairs, arcuate, not very prominent, sHghtly elevated on the lower surface; glands occurring on the lower surface of the blade (occasionally on the upper surface), mostly near base, often very small, 0.2 mm. across, but sometimes reaching 0.5 mm., elliptic or circular, with raised rim and depressed center; stipules subpersistent, hirsute, subulate, apprcssed, 3-5 mm. long. Flowers in axillary and terminal raceme-like panicles 3-7 cm. long, the "racemes" compound but the lateral cymes most often reduced each to 1 or 2 flowers and the inflorescence giving the appearance of a simple raceme with bracteolate pedicels 8-15 mm. long (actual pedicels 2-3 mm, long); bracts of the cymes lanceolate, attenuate, minutely glandular-denticulate, 1.5-3 mm.; petals white, broadly ellip- tic, rounded at both ends, about 5 mm. long and 3-3.5 mm. wide, glabrous; stamens 3, opposite adjoining sepals, much exserted, pink or purple, glabrous, the filaments 1.2-1.6 cm. long, fleshy and about 1 mm. wide at base and tapering to the very slender tips, arising separately from the hypanthium-rim which is fleshy and projects about 0.5 mm. above the base of the calyx; anthers about 0.6 mm. long; hypanthium campanulate, the cavity funnelform, about 1.8-2 mm. deep, retrorsely hairy within the fleshy rim but glabrous at bottom; calyx-lobes oblong to suborbicular, rounded at tips, reflcxed after anthesis, entire, 2-2.5 mm. wide, 3-3,5 mm. long, the outer surface with long yellow appressed hairs and scattered small white hairs also, the Inner surface white-tomentulosc; ovary hirsute, sessile; style as in H. amcricana, but up to 1.5 cm. long. Fruit dark red or purple, oblong- obovoid, up to 2.3 cm. long and 15 mm. thick, with rounded apex and thick, short, stipe-like base, rather densely yellow-hairy. West Indies; British Honduras and Guatemala to Panama; northern South America and south in the Amazon basin to southern Brazil, eastern Peru and eastern Bolivia; low and middle elevations, in moist or wet forests. BOCAs DEL TORo: Flsh Creek hills, von Wedel 24jy. canal zone: Barro Colorado Island, Bangham 398; Salvoza 865; Wetmore 6 Abbe TqS; WooJworth & Vestal 647; Frijoles, ?ittier 26S0; along Rio Indio de Gatiin, Vttfkr 2784; Fort Sherman, Staiidley 31 1 35. colon: along Rio Fato, Vittier 3869; Santa Isabel, Fitfier 4178. darien: El Real, Allen gdj; Garachine, Vittier 5523; Pinogana, ?ittier 6S77\ Cituro, K, S, Williams 667. PANAMA: hills above Campana, Allen 1689; Pacora, Allen 34531 near Vigia and San Juan on Rio Pcqueni, Dodge, Steyermark & Allen 16524; near Chepo, Kluge 48; Rio Tatare, Woodson ^ Schery IO08, ?san blas: Perme, Cooper 286; Marraganti [PMazar- gandi], R. S. Williams lOOO, 10. COUEPIA Aublet CouEPiA AubL Hist, PL Guian, Fr. 519, pi 207. 1775; Hook. f. in Mart. Fl Bras. 14^:40-49. 1867,. Trees or shrubs, with alternate, simple, entire leaves. Flowers in racemes or panicles; hypanthium elongate and somewhat tubular, often gibbous at the base, (163) [Vol. 37 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN the lobes 5; petals usually 5; stamens 10-15 or numerous, inserted at the summit of the hypanthium tube, the filaments more or less united at base in a complete or incomplete ring; ovary unilocular, hairy, attached to the side of the hypanthium- tubc above the base. Fruit a dry or fleshy drupe, the stone mostly woody, 1- sce ded. 1. CouEPiA PANAMENSis Standi, in Trop. Woods 42:22. 1935. Tree up to 20 m. tall, nearly glabrous, the petioles, young growth, and in- florescence sparsely strigose with appressed sordid hairs 0.5 mm. long or less, the inner calyx-lobes canescent-puberulent near tips; branchlets dark reddish brown, glabrcsccnt, about 2 mm. thick below the inflorescence, the Icnticels inconspicuous. Leaves ovate or elliptic, the tip acute and terminated by a blunt acumen about 1-1.3 cm. long and 2-4 mm. wide, the base of the blade acute and attenuate into a stout petiole 1.5-2 mm. thick and 4-6 mm. long; petiolar glands elliptic, near summit of petiole, about 1.5-1.8 mm. long, with raised margin and a flat sunken central area; similar but much smaller glands (about 0.5 mm. long) occur at inter- vals along the margins of the blades, especially near tips; blades 3-5 cm. wide, 8-12 cm, long, 2—2.5 times as long as wide, lustrous on both surfaces when dry, the veins inconspicuous, arcuate, about 5-6 pairs, those near apex usually anasto- mosing and not forming well-marked ribs; stipules not seen. Flowers *'grecni:jh to pale white", in small terminal panlcle-Hke clusters 6-8 cm. long; petals fleshy, glabrous, strongly concave (at least in bud; open flowers not seen), broadly elliptic, rounded at apex and base, about 4 mm. wide and 5 mm. long; stamens about 20, mostly anthcriferous, the filaments glabrous, about 6 mm. long in bud, their bases coherent into an incomplete ring, the anthers about 1 mm. wide and long; h}-pan- thium long-campanulate, in bud about 3 mm. in diameter and 5-6 mm. long, at- tenuate to a pedicel 5-8 mm. long, enlarged at apex into the 5 fleshy and strongly imbricate suborbicular entire calyx-lobes which are about 4 mm. long; receptacle funnelform and densely woolly within, the cavity 2-3 mm. deep, the basal half of the hypanthium lignified, the hirsute ovary 1 mm. or less above the bottom of the cavity; style about 3 mm. long in bud, the stigma white, peltate, flattened, about 1 mm. across. Fruit not seen. SAN BLAS: Perme, Cooper 2/g. CONNARACEAE Trees, shrubs, or woody lianas. Leaves alternate, imparlpinnate, rarely unl- foliolate, exstlpulate; leaflets entire. Inflorescence paniculate, terminal or lateral. Flowers relatively small, regular, perfect or rarely dioecious; sepals 5, Imbricate to valvate, free or rarely connate at the base, sometimes accrescent In fruit (Rourca) ; petals 5, free or connate at the base; stamens 10, the filaments united toward the base and in alternate cycles of two lengths, the longer opposite the sepals, the .shorter (sometimes staminodial) opposite the petals; carpels free, 5-1, but usually (164) 1950] FLORA OF PANAMA (Comiaraccae) 179 producing a single fruit, the ovules 2 in each carpel, collateral, erect, anatropous, basal or sub-basal. Fruit a follicle, sometimes indehiscent, or a legume, containing usually a single arillate seed. a. Carpels 5; follicles sessile. b. Sepals valvate, not enlargi^d in fruit; follicles densely fcrruginous- tomentose; leaves densely and persistently pubescent 1. Cnestidium bb. Sepals imbricate, enlarged in fruit; follicles and leaves glabrous or glabrate - 2. RouREA aa. Carpels 1; follicles stipitate, essentially glabrous; leaves glabrous or glabrate ^' Connarus I 1. CNESTIDIUM Planch, Cnestidium Planch, in Linnaea 23:439. 1850; Schellenb. in Engl. Pflanzenreich IV. 127 (Heft 103):191. 1938. Woody lianas. Leaves imparipinnate. Inflorescence paniculate, pscudoterminal in the axils of the upper leaves. Flowers small, eglandular, subsessile; sepals valvate, not enlarged in fruit; carpels 5. FolUcles almost always soUtary, sessile, very densely ferruginous-tomentose without, containing a single arillate seed. In addition to the following species, one other occurs in the Guianas. 1. Cnestidium rufescens Planch, in Linnaea 23:440. 1850. Kourca frutesccfis Aubl. ex Griseb. in Bonplandia 6:6, 1858. Rourca hondurcnsh Donn. Sm. in Bot. Gaz. 40:2. 1905. Woody lianas as much as 12 m. high, sometimes attaining the proportions of a slender tree. Leaves large, with 7-9 oblong to obovate-oblong leaflets 3-9 cm. long which arc persistently ferruginous-pubescent, particularly beneath. Flowers subsessile, numerous, in densely ferruginous-pubescent, pscudoterminal panicles 10-15 cm. long; sepals about 3 mm. long; petals white, very slightly longer than the sepals. FoUicles sessile, densely ferruginous-tomentose, 1.0-1.5 cm. long, 5-7 mm. broad. Moist or swampy forests and thickets from sea-level to about 120 m.; common in most of Panama; southern Mexico to Colombia; Cuba. TQWn CANAL , 137; Woodworth 6 Vestal SQQ; Van Tyne s.v,; Standley 41000; Bangham 403; drowned forest near Vigia and San Juan, on Rio Pequeni, alt. 66 m., Steyermark & Allen 16572; vicinity of Miraflores Lake, G. White 167; near Graces, Hayes 93^; Frijoles, Piper 5834; Allen 925; near Fort Randolph, Standley 28679; ^l^ng old Las Cruces Trail between Fort Clayton and Corozal, Standley 29028, 29139; near Summit, Standley 29592, 30090; vicinity of Fort Sherman, Standley 31087; Obispo, Standley 31/44; Gamboa, Stevens nil; between France Field, C. Z., and Catival, Province of Colon, Standley 30172, cHiRiQuf: vicinity of San Felix, alt. 0-120 m,, Vittier 5740. cocLfe: between Aguadulce and the Chico River, ?ittier 5104; Penonome and vicinity, Williams 186. bocas del toro: Water Valley, von Wedel 895. Panama: Punta Paitllla, Piper 34-9; between Pacora and Chepo, Woodsan, Allen ^ Seibert 164 j; Camino del Botlcario, near Chepo, Pit tier 4553; "ear Matias Hernandez, Standley 28953; ^io Tecumen, Standley 29432,29443; Cerro Campana, Allen 268 1; Tumba Muerto Road near Panama, Standley 297 41; San Jose Island, Erlanson 560; Taboga Island, Barclay s, n.; G. S. Miller 1842; Standley 27985. (165) [Vol. Z7 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. ROUREA Aublct RouREA Aubl. Hist. PL Guian. Fr. 1:467. 1775; Schellenb. In Engl. Pflanzenrelch IV. 127 (Heft 103):194. 1938. Rohcrgia Schrebcr. Gen. 1:309, 1789. Malbrancia Necker, Elem. Bot, 2:366. 1790. Eicblcriu Progel, in Mart. FL Bras. 12^:518. 1877. Woody Leaves imparipinnate. rarely unifoliolate, when pinnate tbe lateral leaflets usually approximate or alternate rather than strictly opposite. Inflorescence terminal, pseudoterminal, or lateral, paniculate, usually more slender and with fewer flowers than In the two other genera. Flowers small, usually distinctly pedicellate; sepals imbricate, conspicu- ously enlarged In fruit, sometimes glandular; carpels 5. Follicles almost always sohtary, sessile, glabrous or essentially so, containing a single arillate seed. A genus of the American tropics containing about 32 spec Schellenberg. The plants normally are woody lianas, but in the absence of suitable support may become erect or nearly so. a. Calyx glandular, tomentose. b. Flowers yellowish green; sepals 3-4 mm. long; leaflets 6-17 cm. long. densely tomentulose beneath __ 1, R. Pittieri bb. Flowers white; sepals 1.5-2.0 mm. long; leaflets 3-10 cm. long, very Inconspicuously pubcrulcnt to glabratc beneath 2. R. adenophora aa. Calyx cglandular, glabrous or inconspicuously ciliolate; leaflets 4-12 . long, glabrous 5^ r, glabra cm 1. RouREA Pittieri S. F. Blake, in Bull Torrey Bot. Club 50:274. 1923. Woody lianas. Leaves impariplnnate, leaflets 5-7, broadly elliptic, apex sub- caudate-acuminate, rarely obtuse, 6-17 cm. long, 3.5-8.5 cm. broad, glabrous above, densely tomentulose beneath. Sepals 3.5-4.0 mm. long in flower, 4.5-5.0 mm. in fruit, glandular and densely tomentulose without; petals yellowish green, 4-5 mm. long. FoUicles about 1.5 cm. long, glabrous. Panama, in moist thickets and forests, ascending to about 700 m. altitude. colon: Loma de la Gloria, near Fato (Nombre de Dios), Vittier 424J, darien: foot- hills of Garagara, Sambii basin, ?ittier 5613; Cana-Cuasi Trail, Chepi^ana Distr., Terry & Terry Ijj6. ^ 2. RouREA ADENOPHORA S. F. Blake, in Bull. Torrey Bot. Club 50:273. 1923. Woody lianas. Leaves imparipinnate, leaflets 3-7, rarely 1, broadly oval to rather narrowly oblong-elliptic, apex narrowly subcaudate-acuminate, rarely ob- tuse, 3-10 cm. long, 1.5-5.0 cm. broad, glabrous above, very inconspicuously pubcrulent to glabrate beneath. Sepals 1.5-2.0 mm. long in flower, 3.5-4.0 mm. in fruit, glandular and finely pllosulose without; petals white, about as long as the sepals. Follicles about 1.5 cm. long, glabrous or glabratc. Panama, in moist thickets and forests at altitudes near sea-level. ^ CANAL zone: vicinity of Frijoles, Viper 5812; near mouth of Rio Chagrcs, Allen 887; vicinity of Summit, Standley 30102; between Gatun and Lion Hill, Pf/fier 2j66; near Old (166) 19S0] FLORA OF PANAMA (Connaruceae) 181 Fort Lorenzo, moutli of Rio Chagres, Piper 39/6; Santa Rita Trail, Cowcll III; Barro Colorado Island, Standlcy 4.OQYI ; Bangbam 48/'; hills west of Canal, near Gatun, Standley 2^216, cHiRiQui; forests around Puerto Remedios, Pittier JjSo, 3. RouREA GLABRA HBK, Nov. Gcn. &Sp. 7:41. 1825. Connanis glaber DC. Mem. Soc. Hist. Nat. Paris 2:385. 1825. Robergia glabra (HBK.) Spreng. Syst. 4-:188. 1827. Kourea oblongifolia Hook. & Arn. Bot. Bccchey Voy. 283. 1836. Sanfalodes glabrum (HBK.) O. Ktze. Rev. Gen. Pi. 1:155. 1891. Fig. 74. Kourea glabra (167) 182 ANNALS OF THE MISSOURI [Vol. ^7 Woody lianas. Leaves imparlpinnate, leaflets 3-5, elliptic to oval, apex sub- caudate-acuminatc to obtuse, 4-12 cm. long, 2-6 cm. broad, glabrous. Sepals 2-5 mm. long in flower, 4-5 mm. in fruit, eglandular, glabrous or inconspicuously cilio- late; petals white, 5-6 mm. long. Follicles about 1.5 cm. long, glabrous. Southern Mexico to southern Brazil; Cuba and Jamaica. In Panama a frequent liana in moist thickets and forests at lower elevations. CANAL zone: near Fort Randolph, Standlcy 286S6; Gamboa, Stevens I0g4; Ancon Hill, StauJley 26j8l; Chagres, FendJer 5/; vicinity of Salamanca Hydrographic Station, Rio Pequcni, Woodson, Allen (3 Seibert 1556; between France Field, C. Z., and Cativa], Province of Colon, Sfandlcy JOlSo, coci^t: Penonome and vicinity, Williams 149, ama: between Matias Hernandez and Juan Diaz, Sfandley 32016; near the big swamp east of the Rio Tecumen, Sfandley 26/'OJ; Pnitilla, Bra. liertberto 2o8; Isla Taboga, Wood- son, Allen & Seibert I^JJ. 3. CONNARUS L. CoNNARUS L. Sp. PL 2:675. 1753; Schellenb. in Engl. Pflanzenreich IV. 127. (Heft 103):216. 1938. Oniphalobium Gaertn. Fruct. 1:217. 1788. Thysanns Lour. Fl. Cochinch. 1:284. 1790. Canicidia Veil. Fl. Flum. 4:pl. 139. 1827. Eryihrostlgnia Hassk. in Flora 25:Beibl. 2:45. 1842. Anisostewonl'uTcz. in Bull. Soc. Nat. Mosc. 2:152. 1842. Tricholobus Bl. Mus. Dot. Lugd. Bat. 1:256. 1850. Woody lianas, shrubs, or trees. Leaves imparipinnate, rarely unlfoliolate. In- florescence paniculate, terminal and lateral in the upper leaf axils. Flowers small, glandular; sepals imbricate to subvalvate, more or less conspicuously punctate- glandular, not enlarged in fruit; petals glandular or eglandular; stamens 10, or the inner whorl reduced to staminodia, the filament and connective bearing small, sessile or stlpitate glands; carpels 1. Fruit a stipitate, gibbous follicle containing a single arillate seed. About 121 species of the tropics of both hemispheres, according to Schellcn- berg. In Panama the species are woody lianas, occasionally erect or suberect, in- habiting moist forests and thickets at low elevations. a. Inflorescence densely and persistently ferruginous-tomentulose, the flow- ers sessile or subscssile; leaflets usually 3, 6-20 cm. long. 1. C. panamensis aa. Inflorescence Inconspicuously pilosulosc to glabratc, the flowers shortly but definitely pedicellate. b. Leaflets usually 5, broadly oval to oblong-elliptic, broadly obtuse to rounded at the base, 3-12 cm. long 2. C. TuRczANlNOWii bb. Leaflets usually 3, oblanccolate, rather narrowly cuncatc at the base, 10-25 cm. long 3. C. Williamsii 1. CoNNARUs PANAMENSis Griseb. in Bonplandia 6:6. 1858, non Turcz. Connarus haemorrboeus Karst. Fl. Columb. 2:73. 1866-69. Connarns Cooksii Pittier, ex Schellenb. in Engl. Pflanzenreich IV. 127. (Heft 103):223. 1938, nom. nud. In synon. Woody lianas, sometimes erect or suberect. Leaflets usually 3, rarely 5, eHiptic to oval, broadly obtuse at the base, 6-20 cm. Ion", 2-10 cm, broad on flowering? or fl68) 1950] FLORA OF PANAMA (Connaraccac) 183 fruiting branches, sometimes larger on vegetative stems, ferruginous-tomentulose to glabrate. Inflorescence densely and persistently ferruginous-tomentulose, spicate- paniculate, the small yellowish flowers sessile or subsessile. Sepals ovate, about L5 mm. long; petals about 2.5 mm. long. Follicles obliquely compressed-obovoid, about 1.5 cm. long and 1 cm. broad, with a stipe about 3—4 mm. long. Costa Rica to Colombia. CANAL Zone: 314^51 "^00 Starry 20'/'; Bailey 224; Bangham jpg, 41S; Aviles 10; Cerro Gordo, near Culebra, Pittier 2314; Rio Agua Saliid, near Frijoles, Piper jS62; vicinity of Pacora, Allen IO08; along Rio Teciimen, north of Chepo Road, Hunter ^J Allen 2J0; Rio Tapia, Standley 28212; Isla Taboga, Woodson, Allen ^ Seihert 1449; San Jose Island, Erlansoft 8^, chiriqui: Rcmedios and vicinity, Pittier 547^* cocle: Penonomc and vicinity, Williams 3/6; between Paso del Arado and Ola, Pittier 5010. 2, CoNNARUs TuRCZANiNOwii Tdana, in Ann. Sci. Nat. V. 16:364. 1872. Connarus panamensis Turcz. in Bull. Soc. Nat. Mosc. 22:277, 1859, non Griscb. Woody lianasj sometimes becoming erect or suberect. Leaflets usually 5, broadly oval to oblong-elliptic, broadly obtuse or rounded at tbe base, 3—12 cm. long, 1.5—5.0 cm. broad on flowering or fruiting branches, glabrate or glabrous. Inflorescence inconspicuously pilosulose to glabrate, the small greenish white or yellow^ish flowers shortly but definitely pedicellate; sepals narrowly lanceolate, 2.0-2.5 mm. long; petals about 5 mm. long. FoUicles obliquely compressed- obovoid, about 1.5 cm. long and 1 cm, broad, with a stipe about 2—3 mm. long. Apparently endemic to Panama. CANAL zone: Chagres, FenJler 128; Gatun, Hayes 166, 54§; vicinity of Frijoles, ^iper 5797, 5832; near Fort Randolph, Standley 28603; Barro Colorado Island, Bailey Sf Bailey 322; between France Field, C. Z., and Catival, Province of Colon, Standley 302^3, colon: Aspinwall [Colon], Hayes 633, 639, It is noteworthy, perhaps, that the cited specimens of C. Turczaninowii are all from the Atlantic slopes of the isthmus, while those of C. panamensis are from the Pacific slope, with the exception of the plants from Barro Colorado Island. 3. Connarus Williamsh Britton, in N. Am. Fl. 22^:560. 1918. Connarus Allenii Steyermark, in Ann. Mo. Bot. Card. 28:430. 1941. Woody lianas, sometimes erect or suberect. Leaflets usually 3, sometimes 5, oblanceolate, rather narrowly cuneate at the base, 10—25 cm. long, 2-8 cm. broad on fruiting branches, glabrous or glabrate. Inflorescence rather inconspicuously pilosulose to glabrate, the small flowers shortly but distinctly pedicellate; sepals 2—3 mm. long; petals 4.0—4.5 mm. long. Follicles obliquely compressed-obovoid, about 1.5 cm. long and 1,3 cm. broad, with a stipe 3—4 mm. long. Apparently endemic to Panama. darien: Marraganti and vicinity, Williams JO06; trail between Pinogana and Yavisa, Allen 24g. (169) [Vol. 37 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN LEGUMINOSAE Herbs, vines, shrubs or trees of exceedingly diverse habit and habitat, frequently armed (most frequently in the subfamily Miaiosoideae) , Leaves alternate, tri- foliolate, pinnate or bipinnate or rarely simple, usually large, sometimes sensitive (folding at night or upon disturbance) ; petiole and rachis prominent or infrequent- ly nearly obsolete, typically pulvlnate basally, usually sulcate on the upper side and often (MiMOSOiDEAE particularly) bearing one or more glands; leaflets many or few, as a rule small, almost invariably with the margins entire, commonly oblique basally, pubescent or glabrous, the venation prominent or obscure; stipules and usually stipels present. Inflorescence axillary or terminal, indeterminate, the flowers solitary or more commonly in racemes, panicles, spikes or heads; floral bracts usually present but inconspicuous. Flowers typically, but not always, perfect, complete and 5-merous, regular, sympetalous and valvate (the great majority of the Mimosoideae) or more commonly Irregular, polypctalous and imbricate (Caesat - PINOIDEAE and Papilionoideae) ; calyx usually synsepalous and campanulate, sometimes of separate sepals, frequently pubescent; corolla funnclform (most Mimosoideae), slightly irregular and with the upper petal to the inside (Caesal- pinoideae), or papilionaceous (strongly irregular with the upper standard outside the wings and keel: Papilionoideae) ; stamens usually 10 (most Papilionoideae) to many (most Mimosoideae), sometimes few (some Caesalpinoideae), except in the Caesalpinoideae and a few genera of other subfamilies the filaments united below; anthers bilocular, small or large, sometimes gland-bearing, in a few genera dehiscing by terminal pores; ovary unicarpellatc and 1-celled, bearing 1 to many ovules from the dorsal suture, usually oblong and compressed, stipitate or sessile, surmounted by a prominent, simple style and usually a small stigma. Fruit a legume, but this extremely variable in size and shape, dehiscent or in a number of ■ cases Indehisccnt, the seeds typically exalbuminous and bean-like. The Lcguminosae is an exceedingly large and diverse family, well represented in both the tropics and temperate regions (Mimosoideae and Caesalpinoideae largely tropical). In spite of its inordinate size and perhaps consequent diversity, the family is quite "natural'* and is seldom confused with other families. Some exceptions can be found to almost any defining characteristic, but most useful are the usually compound, alternate, stipulate leaves, flower form, and unicarpellatc ovary with dorsally affixed ovules becoming in fruit a legume. The floral difi'erences In tlie three subfamilies arc striking and clear-cut for the large majority of genera, so that ample justification can be had for division of the Leguminosae into three families, the Mimosaccac, Caesalpiniaceae, and the Papilionaceae or Fabaceac. Many of the world's most important and useful plants hail from the Lcguminosae. In the tropics their importance perhaps exceeds that of even the Gramineae (cereal grains, etc.). Aside from supplying timber, food, ornamentals, drugs, poisons, insecticides, resins, shade and ground cover, the Leguminosae are noted for the (170) 195 01 FLORA OF PANAMA (Lcgumhtosae) 185 peculiar symbiotic relationship of nitrifying bacteria on the roots which makes them very useful for soil improvement. In many tribes generic differences are exceedingly obscure and undependable, while in others they are sharply marked. Parallel evolution among genera and a multitude of exceptions to the characters on which they are based make It nearly impossible to form simple yet infallible keys. In the keys to the genera of Leguminosae in Panama an effort has been made to avoid obscure characters and those not likely discernible in herbarium material. It is fully realized that some of the distinctions will not always hold when species and genera outside the range of the flora are considered, nor can use of legume characteristics as the sole distinction be avoided in every case. KEY TO THE SUBEAMILIES a. Flowers regular, valvate (rarely imbricate) in bud, usually sympetalous; stamens frequently many and long-exsertcd, giving a delicate "fluffy" appearance to the bead or spike; leaves twicc-pinnate except in Inga and a few species of Pithecolohium Mimosoideae aa. Flowers irregular, imbricate In bud, polypetalous, petals unequal; stamens typically 10 or fewer, mostly briefly exserted or not exserted, rarely in heads or "fluffy" spikes; leaves twice-pinnate only in a few genera. b. Upper petal within the others in bud; flowers not papilionaceous; stamens usually free; leaves sometimes twIce-pInnate (Cafsal- pinieaf) Caesalpinoideae bb. Upper petal exterior in bud; flowers typically papilionaceous; stamens except In one tribe united below; leaves never twice-pinnate Papilionoideae Subfamily Mimosoideae (Mimosaceae of many authors) Trees, slirubs, vines or herbs, variously armed or unarmed. Leaves twice-pin- nate (except for Inga and a few species of Pithecohhmm in which the leaves are once-pinnate), usually large, in most genera bearing a gland or glands on the petiole and/or rachis, in some genera sensitive (folding at night or when disturbed) , the pinnae normally opposite; leaflets large and few or more frequently small and numerous; petiole usually conspicuously pulvinate. Inflorescence commonly of axillary, pedunculate heads, umbels or spikes, these often borne subterminally from non-foliate nodes and hence constituting a terminal raceme or panicle, usually colorful and ornamental. Flowers usually small, densely clustered, commonly 5- parted, regular, usually complete, rarely imperfect, sessile or less frequently pedicel- late, often fragrant; calyx synsepalous, valvate in bud or rarely imbricate, usually small, sometimes minute; corolla sympetalous or polypetalous, usually exceeding the calyx, ordinarily valvate; stamens few or many, free or united, long-exserted, usually the most consplcvious and colorful part of the flower; anthers minute to moderate, gland-tipped or cglandular; ovary 1-carpellate, mostly symmetrical, free, stipitate or sessile, glabrous or pubescent, the style simple and about equalling the stamens. Legume very variable, flat and thin to fleshy and torulose, variously de- hiscent or rarely even indehiscent. A distinctive subfamily well represented in the tropics, especially in the New World. Alany genera are nearly pantropical and only a few are confined to either (171) 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 the Old or New Worlds. A number of species are large trees of ornamental and economic importance. Other species are pestiferous armed weeds. The MiMosoi- DEAE as a whole are usually distinguishable w^ithout difficulty by the "fluffy" heads or spikes of clustered, regular flowers with long-exserted stamens. Most genera of the Mimosoideae are extremely difficult taxonomically. Parallel evolution, omnipresent variability, and repeated intcrgradation arc the rule rather than the exception. As a result, numerous segregate genera have at various times been proposed, which can scarcely serve any practical purpose. It is difficult enough to recognize the traditional genera of Willdenow and Bcntham without in- cluding ad infinitum the many possible segregable ''genera" usually based upon a single difference (this typically in the legume and uncorrected with distinctive vegetative or floral characters). Such segregates cannot be adequately separated in a key to the genera. It Is perhaps significant that the best-versed students of the group, preeminently Bentham, have refrained from inordinate segregation and dependence upon legume characteristics. Willdenow had established legume char- acters as primarily definitive of most genera, and indubitably such characters are valuable taxonomically. Yet they would seem to have been overstressed. Ample collections of specimens in fruit (seldom sufficient from the tropics, as most speci- mens of the usually meagre collections consist of flowering material alone) seem to present the Intergradatlon and variability commonly met with in this subfamily. The outstanding work on the Mimosoideae is Bcntham's classical ''Revision of the Suborder Mimoseac" (Trans. Linn. Soc. 30:335-664. 1875), and his treatment of the subfamily for Martius' 'Flora Brasiliensis.' More recent work in the group has been carried on by Ducke, Pittier, Taubert, and others, in a number of publica- tions, while floristically the Mimosoideae have been treated for North America by Britton and Rose (N. Am. Fl. 23:1-194. 1928), for Colombia by Britton and Killip (Ann. N. Y. Acad. Sci. 35:101-157. 1936), for Peru by Macbride (Field Mus. Publ. Bot. 13:1-113. 1943), and for Guatemala by Standley & Steyermark (Ficldiana: Botany 24:1-88. 1946). In spite of this generous attention a great deal of uncertainty still exists and a need for monographic study of the genera is evident. a. Stamens united into a short or long tube, usually many, rarely (viz. Entcrolohium) as few as 10; flowers generally rather large, the sym- petalous corolla frequently 4—5 mm. long or longer. b. Leaves oncc-pinnatc, 2-ranked; flowers whitish 1. Inga bb. Leaves twice-pinnate (except few species of Pitbccolohium which then have the leaves few-ranked), mostly few-ranked; flowers some- times white or cream but often pink. c. Either the petiole or rachis or both gland-bearing (the gland some- times obscure in Alhizzia); legume various but not clastically de- hiscent from above. d. Petiolar glands lacking or borne only at the apex of the petiole at or just below insertion of the pinnae; legume relatively slen- der and thick, often torulose, rarely septate, usually dehiscent and pulpy, the seeds usually longitudinal; corolla usually more or less tubular and greatly exceeding the calyx 2. Pithecolobium dd. Petiole bearing a gland or glands (except obscure in Alhizzia (172) 1950] FLORA OF PANAMA (Leguminosae) 187 carbonaria) , these usually near or below the middle; legume flat and thin but moderately broad, indehisccnt or tardily de- hiscent, scarcely pulpy, the seeds transverse; corolla (except few species of Pifhccofohhim) funnelform, usually only about twice as long as the calyx. e. Legume circinate to arcuate, somewhat fleshy, septate or falsely septate; pinnae and leaflets many (leaflets 1 0—60 pairs per pinna) ; leaflets shorter than 4 mm. or longer than 8 mm., or it 5—7 mm. long the midvein nearly central. f. Legume about 1 cm. wide, arcuate or somewhat coiled and twisting in dehiscence, falsely septate; leaflets 5—10 mm. long, the cost a nearly central; flowers, where known, es- sentially glabrous; 2 or 3 species only 2. Pithecolobium ff. Legume 2 or more cm. wide, circinate or rcniform, coiled into a complete or nearly complete circle, septate; leaflets shorter than 4 or longer than 8 mm,, the costa cxcentric; flowers tomcntulosc 3. Enterolobium ee. Legume straight or nearly so, thin and dry, not septate; pinnae and leaflets usually fewer (leaflets less than 10 pair per pinna except in 2 species, these with leaflets 5-7 mm. long, their mid veins excentric) 4. Albizzia cc. Petiole and rachis both eglandular; legume elastically dehiscent from above. d. Inflorescence capitate; stamens many 5. Calliandra dd. Inflorescence spicate; stamens 5 plus 5 staminodia 16. Pentacletiira ca. Stamens free or united only at the extreme base, except in Acacia 10 or fewer; flowers generally rather small, the corolla sympetalous or polypetalous and usually considerably less than 5 mm. long. b. Herbs, vines, shrubs or trees, seldom aquatic or of swamp habitat; all flowers visibly similar; stems generally lenticellate or armed; legume various. c. In Panama Inflorescence capitate or very short-splcate (except few species of Acacia which then have large, hollow, stipular spines) ; anthers eglandular (rarely glandular in Acacia), d. Petals more or less united (divided almost to base in species of Acacia with pedicellate flowers); anthers minute (fraction of a mm. long); flowers usually very small; plants frequently armed. e. Unarmed or armed, the armament frequently of stipular spines; stamens usually more than 10; petiole and rachis fre- quently glandular, without nodular apicules; legume unarmed.. 6. Acacia ee. In Panama stems armed with scattered, recurved thorns (ex- cept sometimes M. puailla); stamens few, usually 10 or fewer, petiole and rachis eglandular but commonly with nodular apicules; legume usually aculeate. f. Legume flat, oblong or broad, transversely articulate (in Panama) ; lower flowers of head rarely unisexual; pinnae and leaflets various but often not as below 7. Mimosa ff. Legume terete or turgid, usually narrowly linear, continu- ous; lower flowers of head unisexual; pinnae few to several rather remote pairs, with many leaflets 8. Schrankia dd. Petals free; anthers comparatively large (usually Yz—l mm. long); flowers small; plants unarmed. e. In Panama small shrubs or subherbs; stipules setiform, sub- persistent; heads with fewer, erect flowers, the peduncular torus little expanded; legume scarcely 5 mm. wide 10. Desmanthus ee. Large shrubs or trees; stipules ovate or caducous; heads orbic- ular, from an expanded peduncular torus; legume almost 20 mm. wide 11. Leucaena cc. In Panama inflorescence elongate-splcate; anthers gland-bearing, at least in bud. d. Leaves and inflorescences on condensed short-shoots; petals pilose within; stems with "nodular" spines - 12. Prosopis (173) 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. Z7 dd. Lacking short-shoots; petals glabrous; armament, if any, of scattered thorns. e. Trees or shrubs; leaflets less than 15 mm. long, numerous; legume not breaking transversely. f. Leaflets mostly alternate; legume indchlsccnt, unarmed 13. Stryphnodendron ff. Leaflets mostly opposite; legume dehiscent; unarmed or aculeate. g. Usually aculeate; petiole usually glandular; calyx-lobes valvate; flowers with 10 stamens, lacking staminodia; legume smaller, thinner , 14. Piptadrnia ^g. Unarmed; petiole eglandular, calyx-lobes Imbricate; flow- ers with elongate staminodia and 5 stamens; legume large, ligneous _ 16. Pentaclethra ee. Large vine-like plants, often tendrilcd; leaflets longer than 15 mm., few; legume valves eventually breaking into seg- ments 15. Entada bb. Herbs or subshrubs, of swampy habitats or even free-floating; lower flowers of head with large, petaloid staminodia, visibly quite different from upper perfect flowers; stems unarmed, not lenticcllate; legume short, broad, flat, unarmed, glabrous 9. Neptunia L INGA Scop. Inga Scop. Introd. Hist. Nat. 298, 1777. Amosa Neck. Elem. 2:459. 1790, fide Dalla Torre & Harms. Torcalia Nor. in Verh. Batav. Gen. 5, Art. IV. 4. 1790, fide Dalla Torre & Harms. Ingaria Raf. Sylva Tellur. 119. 1838. Fcuillcca O. Ktze. Rev. Gen. PL 1:182. 1891, in part. Unarmed trees or shrubs, the branchlets usually pubescent when young and noticeably Icnticellate. Leaves of moderate size to very large, 2 -ranked, simply compound, the leaflets 2-many pairs; petiole usually short, only rarely alate; rachis shore or elongate, unwingcd or alatc, almost invariably bearing subcupuhform, sessile or less frequently stipitatc glands between insertion of the petlolulcs; leaflets entire, rather small to very large, frequently pubescent, the veins normally promi- nent below; stipules mostly caducous; petiolules short. Inflorescence of 1 to feu pedunculate spikes from the axils, or paniculate by insertion of spikes from terminal or subterminal defoliate nodes; peduncular portion of spikes elongate to almost ob- solete, the florifcrous portion elongate and lax to condensed and clavate, capitate or umbellate; pedicels prominent in only a few species; bracts usually inconspicuous, caducous or subpersistcnt. Flowers white or whitish, regular, perfect, typically Mimosaccous; calyx cupular or tubular, very small to quite large, frequently pubescent, 5 -toothed; corolla sympetalous, tubular or funnelform, short and small to very elongate, usually pilose without, 5-lobed; stamens many, exceeding the corolla, the filaments united below into a staminal tube; ovary glabrous or pubes- cent; style elongate, usually equalling or slightly exceeding the stamens, sometimes capitellate. Legume from short and flat to elongate and subtercte, glabrous or pubescent, the margins frequently prominent. Distribution: New World tropics and subtropics; center of distribution ap- parently northern South America. Few genera exhibit the variability found in Inga^ and the difficulty in organ- izing the species into groups or workable keys. Sections originally proposed bv ' (174) 1950] FLORA OF PANAMA (Lcguminosac) 189 Be additional species and especially more material of many problematical or only par- tially known species. It seems rather futile to try to retain clear-cut sections within the genus at the present time, and a ''natural" key must be sacrificed to a large extent so that certain "artificial" distinctions may make it workable. Thus winging of the leaf rachis may not be a fundamental distinction, being possessed by or lacked by closely related species; but yet no character seems as usable for primary breakdown of the genus. Likewise absolute flower size and leaflet size, although undoubtedly somewhat variable with changing environmental conditions, are often more useful characters than are such extremely variable structural features as comparative length of stamlnal tube, shape and size of rachial glands or (in many species) number of leaflets per leaf. Legume characters are probably useful, but the lack of mature fruiting material of most species makes them impractical as a sole basis of segregation. Doubtless when fruiting material becomes as frequent as is flowering material today, variability in legume will become as noticeable as is variability in flower. Many species grade into others, and there is every reason to suppose that hybridization occurs in a number of species. Several rather well- defined complexes are discernible In the genus; and it is my belief that within these, species should be regarded in as broad a sense as possible and that considerable condensation is in order. Little correlation or condensation outside of Panama is attempted here because of the lack of many types in this hemisphere. Frequently in discussions at the end of the species descriptions, suggestions of similarity or relationship with extra-Panamanian species are given. Future collections may link new species or varieties described to previously known species, but at present it would scarcely seem advisable to lump all such divergent specimens into known categories without intermediate specimens bridging the gap. Bentham's "Revision of the Suborder Mimoseae" (loc. cit. 1875) is a landmark In the classification of the genus, but at that date, of course, Bcntham lacked the quantitv of specimens needed to show intergradation of related "species." Pittier has worked with the genus to a considerable extent, Introducing a number of new species and discussion of many others. His revision of the genus (Contr. U. S. Nat, Herb. 18:173-224. 1916) seems scarcely conservative enough, and many of his species are probably not valid segregates. This treatment was without keys, and is quite difficult to use for general determinative work. Pittier's "The Middle Ameri- can Species of the Genus Inga" (Jour. Dept. Agr. Porto Rico 13:117-177. 1929) is a more usable appraisal, correcting many of the deficiencies of the earlier work. Unfortunately, Bentham's sections are more or less rigidly adhered to, and the keys are not easily workable. Pittier can be excused for having difficultly workable keys In a genus such as h^gt^y but it might have been wiser to seek a substitute for the lesrume characters needed to determine section and series. The treatment acccroed higa by Britton and Rose (N. Am. Fl. 23:2-16. 1928) is from the practical standpoint almost useless. Seldom will the key work, and many segrega- tions appear untenable. Britton and Killip follow somewhat the same system in ^75; 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN [VuL, Z7 i their "Mimos.iceae :kn^ Caesalpiniaceae of Colombia" (Ann. N. Y. Acad. Sci. 35:110-124. 1936). Macbridc's "Flora of Peru— Lcgumlnosae" (Field Mus. Publ. Bot. Scr. l}:6-47. 1943) and Standley and Steyermark's "Flora of Guatemala" (Fieldiana: Botany 24:34-46. 1946) seem to give a more conservative treatment for their areas. Ducke (Arch. Jard. Bot. Rio de Janeiro 4:19. 1925; 5:119. 1930; 6:13. 193 3) has treated the species of Jnga in the lower Amazon valley. a. Leaf raclils wIngcJ, at least In part. b. Wing of rachis nearly obsolete; leaflets mostly less than 6 cm. long, subglabrous, blunt; legume flat, oblong, less than 4 times as long as broad; known only from the type (in fruit) T. L Daviosoniae bb. Rachis wing usually prominent; leaflets in most species much larger and frequently heavily pubescent; legume except in one or two species several times longer than broad. C. Flowers unusually large or gross; calyx funnelform or campanulate, mostly 10-15 mm. broad apically; legume flat. d. Calyx and leaflets subglabrous; leaflets 2 pairs 4. I. portobellensis dd. Calyx and leaflets usually heavily pubescent; leaflets mostly 4 P^*''S 6. I. GOLDMANH cc. Flowers thinner or more delicate; calyx rarely as much as 6 mm. wide, subtubular or else very small; legume various. d. Calyx glabrous or sparingly pubescent. €. Leaflets glabrous; inflorescence umbellate or capitate, or very long and laxly spicate. f. Inflorescence umbellate or capitate; calyx at least 2 mm. long. g. Calyx 2-3 mm. long; flowers long-pedicellate; terminal leaflets usually about 10 cm. long 2. I. myriantha gg. Calyx at least 15 mm. long; flowers short-pedicellate or subsessile; terminal leaflets usually 20-30 cm. long 4. I. portobizllensts ff. Inflorescence elongate, lax; calyx about 1 mm. long 28. 1. marginata ee. Leaflets pubescent at least on the veins; inflorescence a dense, short spike. f. Floral bracts 1 0-20 mm. long, persistent; stipules sub- persistent; calyx and lower leaf surface with darker "gland- ular** dots 1 1. I, panamlnsis ff. Floral bracts minute or caducous; stipules caducous, at least in age; plant not noticeably "glandular" dotted. g. Spikes conspicuously pedunculate; leaflets mostly acumi- nate; corolla shorter than 8 mm., or longer than 3 5 mm. h. Corolla never longer than 8 mm.; branchlets cinereous- pubescent; leaflets lightly pubescent (except on veins); legume subglabrous, blackish 5. I. dfnsitlora hh. Corolla longer than 35 mm.; branchlets dark-tomen- tose; leaflets rather heavily pubescent; legume densely ferruginous-strigose 8. I. mucuna gg. Spikes subsessile, the condensed floriferous portion ob- scuring the peduncular portion; leaflets acute to rounded, rarely acuminate; corolla 13—16 mm. long 9. I. Hayesii dd. Calyx noticeably pubescent, usually tomentose. C. Inflorescence umbellate, the flowers slenderly long-pedicellate.... 2. I. myriantha ce. Liflorescence spicate, the flowers sessile or rarely short- pedicellate. f. Leaflets glabrous or very sparingly appressed-hirsute; young branchlets markedly angled, quadrangular; floral bracts ovate, subpersistent, about 1 cm. long; legume flat 10. I. sphctadilis ff. Leaflets pubescent at least on the veins; young branchlets less markedly angled, subterete; floral bracts either nearly linear, or else small or caducous; legume tetragonal or sub- terete. V 4 (176) 1950] FLORA OF PANAMA (Lcgiiminosac) 191 g. Floral bracts 10-20 mm. long, subpersistent; legume broader, tetragonal, glabrous; leaflets minutely "glandular*' below. h. Leaflets mostly 3 (or fewer) pairs; petiole scarcely winged; terminal rachial internode more prominently winged than penultimate. i. Corolla about 1.5 cm. long; staminal tube barely exsertcd; leaflet sparingly pubescent or subglabrous, even on the veins 11. I. panamensis ii. Corolla about 2 cm. lung, the constricted tube ob- scured in bud by the calyx; staminal tube scarcely exserted; leaflet usually lightly hirsute or pilose over entire lower surface 11a. T. panamensis var. PiTTiERi iii. Corolla about 3 cm. long, the bud noticeably clavate; staminal tube well exserted; leaflet usually not prom- inently pubescent except on the veins lib. I. panamensis var. CLAVATA hh. Leaflets 4 pairs; petiole prominently winged; rachis about equally winged on all rachial internodes 12. L alatopetiola gg. Floral bracts inconspicuous, small or caducous; legume slender, subterete or angled, tomentose; leaflets eglandular on lower surface. h. Tomcntum of branchlets, petioles and young legume subhirsute, rubiginous or dark-ferruginous; buds and flowers dark, usually ferruginous or ferruginous-golden. i. Tertiary veins of leaflets conspicuously reticulate; calyx usually 3—6 mm. long; mature stamens 20—25 mm. long; rachis frequently scarcely winged except on terminal rachial internode, or this markedly larger than wing of lower rachial internodes 13. L Oerstediana ii. Tertiary veins of leaflets inconspicuous; calyx 8—10 mm. long; mature stamens almost 45 mm, long; rachis winged about equally on all or most rachial internodes 1 6. I. pauciflora hh. Tomentum of branchlets, petioles, and young legume of shorter hairs, cinereous-ferruginous; buds and flow- ers light, cinereous or cinereous-golden. i. Leaflets ovate to elliptic or obovate, 5-8 cm. wide; flowers less robust, the calyx normally less than 8 mm. long. j. Flowers rather large, the calyx usually 7—8 mm. long and the corolla almost 2 cm. long; branchlets less prominently lenticellate 14. L edulis jj. Flowers small, the calyx about 4 mm. long, the corolla about 8 mm. long; branchlets densely raised-lenticellate 14a. I. edulis var. MINUTULA ii. Leaflets narrowly ovate-lanceolate to obovate-lance- olate, 2-5 cm. wide; flower stouter, the calyx 10—20 mm. long, or if only 8 mm. long the branchlets sub- hirsute. J. Leaflets mostly more than 4 pairs; calyx 10-20 mm. long; branchlets tomentose 15. I, spuria jj. Leaflets 3 or 4 pairs; calyx 8-10 mm. long; branchlets subhirsute 16. L pauciflora aa. Leaf rachis unwinged or merely marginate, b. Leaflets 5 or more pairs. c. Pubescence setose or villous, of very long hairs; flowers long- pedicellate 17. L Saffordiana cc. Pubescence, if any, tomentose, of short hairs; flowers sessile. d. Leaflets moderately appressed-pubesccnt to tomentose below; leaflets mostly broadest at or below the middle; flowers larger, the cslyx 5-12 mm. long. (177) 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 e. Leaflets moderately pubescent; branclilets ferruglnous-tomcn- tose; legume somewhat flattened? 18. I. multijuga ee. Leaflets heavily pubescent; branchlcts dark-rublglnous-tomen- tosc; legume terete?; known only from the incomplete type.... 19. I. cocleensis dd. Leaflets subglabrous or scattcringly pubcrulcnt; leaflets broadest above or below the middle; flowers small, the calyx 3-4 mm. long. e. At least the terminal leaflets broadest above the middle; leaflets usually 6 pairs; flower rather large (calyx 3-4 mm., corolla 7-8 mm. long), densely pubcrulent .20. I. Ruiziana ee. Leaflets broadest at or below the middle; leaflets 4-5 pairs; flower smaller (calyx about 2 mm., corolla 4-6 mm. long), lightly pubcrulent 21. I. microstachya bb. Leaflets 3 (rarely 4) or fewer pairs. c. Flowers large and stout, the corolla 15-25 mm. long, the calyx cither about 9 mm. long or else wcxilly and nearly 5 mm. wide. d. Leaflets 4 pairs; calyx cupular, lanose-tomentose; floral bracts small, caducous 7. J. Standleyana dd. Leaflets 1-3 pairs; calyx somewhat tubular, not woolly; floral bracts 10 mm. long or longer, subpcrsistcnt. e. Leaflets glabrous or very sparingly appresscd-hirsutc; young branchlcts markedly angled; floral bracts ovate; legume flat.... 10. I. splctabilis ee. Leaflets pubescent at least on the veins; branchlcts subtercte; floral bracts linear; legume tetragonal lla. L Panamensis Var. CLAVATA cc. Flowers small, the corolla seldom more than 10 mm. long, the calyx 1-7 mm. long and up to 2 mm. wide. d. Floriferous portion of spike short, condensed, usually less than 1 cm. long. e. Inflorescence capitate or umbellate, bearing flowers from the terminal few mm. f. Flowers pubescent, somewhat pedicellate; leaflets rarely as much as 18 cm. long, the terminal ones usually \2-\6 cm, long. g. Pubescence of twigs and inflorescence setose or villous, of very long hairs; pedicels almost 2 cm. long; legume dark villous 17. I. Safeordiana gg. Pubescence of youne twigs and inflorescence tomentose, of short hairs; pedicels scarcely 0.5 cm. long; legume tomentulose, h. Pedicels as long as the calyx; calyx somewhat expand- ing basally and subcampanulate 22. I. Roussoviana hh. Pedicels conspicuously shorter than the calyx; calyx funnelform, narrow basally. 23. I. Williamsii ff. Flowers glabrous or subglabrous, prominently pedicellate or sessile; leaflets eltlier very large, 13-40 cm. long, or very rmall, 2-10 cm. long. g. Flowers prominently pedicellate; leaflets small, 2-10 cm. long, usually 1-2 pairs 3. L HETEROPHYLLa gg. Flowers sessile or subsessile; leaflets large, 13-40 cm. long, usually 3 pairs 24. L GIGANTUOLIOLa ee. Inflorescence short-splcate, bearing flowers for about 1 cm. of its length. f. Leaflets 2-3 pairs; twigs and leaflets strigose-puberulcnt; calyx 3-5 mm. long 25. L punctata ff. Leaflets 4—5 pairs; twigs and leaflets subglabrous; calyx about 2 mm. long .21. I. microstachva dd. Floriferous portion of spike elongate, lax, usually about 8 cm. long. e. R.ichis unwingcd, scarcely marginatc; leaflets usually broadly elliptic, bluntly short-acuminate. f. Leaflets 4-12 cm. long and 1.5-5.5 cm, wide, mostly 2 pairs; stipules and stipule-like bracts not apparent, or at most 6 mm. long 26. L laurina (178) 1950] FLORA OF PANAMA (Legumhiosac) 193 ff. Leaflets larger, the terminal pair about 15 cm. long and 6-7 cm. wide, mostly 3 pairs; stipules and stipule-like bracts of new growth and buds conspicuous, mostly 8—12 mm. long 27. I. Caldasiana ee. Rachis briefly winged or decidedly marginate terminally; leaflets usually narrowly elliptic or elliptic-lanceolate, sharply long-acuminate _ 2 8. I. margin ata 1. Inga Davidsoniae Standi, in Field Mus. Publ. Bot. 22:79. 1940. Small tree, the branchlets ferruginous-tomentose becoming glabrate and notice- ably lenticellate with age. Leaves comparatively small, about lO-foHolate; petiole 5-15 mm. long, terete, lightly tomentose; rachis 3-7 cm. long, very narrowly alate (wing most conspicuous on uppermost foliolar internode), pubescent although lightly so on the wings, bearing small, sessile, cupuliform glands between insertion of the peiiolules; leaflets unusually small, oblong or elliptic to ovate-lanceolate, about 7 cm. long and 3.5 cm. wide in largest (terminal) leaflets, scarcely 2 cm. long and 1 cm. wide in smallest (basal) leaflets, subtruncate to bluntly acute apically, rounded or obtuse and somewhat inequilateral basally, coriaceous, sub- glabrous except on the midvein and glossy above, noticeably pubescent en the veins and dull below; stipules caducous. Inflorescence an axillary, pedunculate spike nearly 5 cm., long, the floriferous portion little more than 1 cm. long. Flowers not known. Legume oblong, about 12 cm. long and 5 cm. wide, flat, more or less truncate apically and basally, transversely striate, tomentose becoming subglabrate in age, the valvular margins sulcate. Known only from the type. CHiRiQuf: Volcan dc Chiriqui, Datidson 943- The leaflets of this species have a ^^'Pithecolobium appearance," but the legume scarcely resembles any species of that genus with once-pinnate leaves. Lack of flowers prevents accurate comparison with other species of h^gci, but the small, coriaceous, glossy, rather variably shaped leaflets are distinctive. The legume much resembles that of 7, V7onficola Pittier. 2. IxGA MYRiAXTHA Pocpp. & Endl. Nov. Gcn. & Sp. 3:77, pi, 28g. 1845. Jn^a gracilipcs Standi, in Jour. Wash. Acad. 15:101. 1925. hi^^a Laivranccana Britt. & Killip, in Phytologia 1:23. 1933. Small tree, the branchlets subglabrous and very densely white-lenticellatc. Leaves of moderate size, 4- (rarely 6- or 2-) foliolatc; petiole 1-3 cm. long, terete basally, prominently marginate or alate apically, glabrous or somewhat pubcrulent above; raclns 2-5 cm. long, essentially glabrous, alate between leaflet pairs (win broadest at apex of rachial internode, narrowing baseward), bearing prominent patelliform or subcupulate glands above at the rachial nodes; leaflets usually pairs, mostly elliptic, the terminal pair about 10 cm. long and 4 cm. wide, the basal pair half or less this size, obtuse or broadly cuneate basally, short-acuminate apical- ly, completely glabrous, lustrous, the prominent lateral veins about 5 pairs; stipules linear, about 5 mm. long, caducous. Inflorescence of pedunculate umbels, axillary (179) 194 ANNALS OF THE MISSOURI [Vol. 37 or subtcrminal from young defoliate nodes; peduncles 2-5 cm. long, almost gla- brous; pedicels slender, almost 1 cm. long, lightly puberulent; floral bracts llnear- clavate, 2-3 mm, long, subpersistent. Flowers delicate, greenisli-white; calyx cupuliform-tubular, about 3 mm. long, subglabrous or lightly puberulent, th teeth triangular and about 0.5 mm. long; corolla tubular-funnclform, about 8 mm. long, slightly broadened apically, glabrous except puberulent on the lobes, sub- striate; stamens 2-3 cm. long, the stamina! tube cxserted. Young legume flat, subglabrous, margined. Panama. CANAL zone: Barro Colorado Island, Bangham 48S, ^22; Sahoza g2g; Wooduvrt/j & Vestal 587; France Field, Sfandley JOJjJ. An interesting species apparently without close affinities in Panama, distin- guished by the graceful, prominently pedicellate flowers. The species has not previously been reported from Central America as 7. myriantha. South American material of the species varies In flower size, and some Peruvian specimens have flowers almost \^^ again as large as the specimens here cited. Other Amazonian material seems identical with the Panamanian specimens. In deciding that 7. gracilipcs is not distinct from 7. viyriantha, It has been necessary for comparison to rely on the original description and illustration, plus limited herbarium material of 7. myrianfbay the type specimen not being available, 3, Inga tieterophylla Willd. Sp. Pi. 1020. 1806. Mimosa Varae Poir, in Lam. Encycl. Mcth. Suppl. 1:44. 1810. 7;/^^ iimbellata G. Don, Gen. Hist. Dichl. Pi. 2:391. 1832. Inga proiracla Steud. in Flora 26:758. 1843, fide Benth. Shrub or small tree, the branchlets puberulent when very young, soon becoming glabrous, Icnticellate. Leaves moderate, the leaflets 1-2 (sometimes more?) pairs; petiole short, usually less than 1 cm, long, glabrous in age, subsulcate above, cgland- ular; rachis frequently exceeding the petiole, unwinged, glabrous, bearing cylindrlc glands at insertion of the pairs of petiolules, at least in the larger leaves; leaflets elliptic, 2-10 cm. long and 1-4 cm. wide, in quadrifoliolate leaves the basal pair of leaflets the smaller, acuminate and markedly attenuate apically, cuneate basally, glabrous, the venation reticulate and prominent below; stipules linear, about 2 long, caducous. Inflorescence of axillary pedunculate umbels, slender and delicate; peduncles up to 2 cm. long; pedicels 4-12 mm. long, glabrous. Flowers moderate; calyx cupulate, scarcely 1 mm. long, glabrous; corolla tubular-funnclform, 5-6 mm. long, glabrous, the tube about twice as long as the lobes; stamlnal tube ex- serted. Legume Kncar, up to 13 or 14 cm. long and about 17 mm. wide, flat, glabrous, nearly straight, short-stlpitate. Panama to middle South America. cocle: Bismarck, Williams 600. The cited specimen, examined for the North American Flora, was found in- mm. (180) 1950] • FLORA OF PANAMA (Legumhiosac) 195 correctly determined as L laurina, and constitutes a new record for /. heterophylla north of South America. The specimen is in fruit, but is apparently referable to this species; certainly it is as much so as is South American material of various phylla. There is some doubt that I. viap J ^phylla. Other species related to I. hctcroph 7netrifolia Harms, 7. sertulifera DC, 7. tarapot and 7. coriacca (Pers.) Desv. 4, Inga portobellensis Beurl. in Svensk, Vet. Akad, Handl. 1854:122. 1856. Inga macrophylla Billb. ex Beurl. loc. cit. 123. 1856, fide Ind. Kew. Tree more or less glabrous throughout, the young branchlets with elongate • lenticels. Leaves large, bijugate; petiole usually 2-3 cm. long, winged (the wing narrowing baseward and the petiole subterete basally) ; rachis usually 6-8 cm. long, spatulate-winged like the petiole, bearing at insertion of the pairs of leaflets an elevated, cupuhform gland, bearing apically in young leaves a (deciduous) mucro- nate appendage as much as 15 mm. long; leaflets 2 pairs, obovate-oblong, 20-3 cm. long, 10-13 cm. wide in apical pair (basal pair only half as long), obtuse basally, abruptly acuminate apically, darker above than below, glabrous except occasionally puberulent on the costa; stipules ovate-subfalcate, about 16 mm. long, subpersistcnt. Inflorescence of solitary, axillary, pedunculate heads (or very condensed spikes); peduncle 3-8? cm. long, glabrous; bracts of head ovate or lanceolate, 7-8 mm. long, late-caducous. Flowers short to moderately pedicellate, large and showy; calyx tubular-funnelform, about 25 mm. long, striate, glabrous except pubescent on tips of the lobes; corolla tubular, almost 4 cm. long, glabrous except cancsccnt- pubcscent on the lobes; stamens as much as 7-8 cm. long, the stamlnal tube well- exserted; ovar)^ glabrous. Legume flat, about 16 cm. long and 2.5 cm. wide, arcuate, ligneous, thickened on the margins. Panama and Costa Rica. We have examined no Panamanian material (unless Allen 17282 be this species: see "Specimens of Uncertain Status"). The (European) type, however, is from Porto Bello, province of Colon, Panama. Costa Rican material does not match the original description in some respects, but we accept Pittier's and Britton and Rose's citations as correct for this name. The species seems to be unusual and striking. 5. Inga densiflora Benth. in Trans. Linn. Soc. 30:617. 1875. Inga mofjticolt Pittler, in Contr. U. S. Nat. Herb. 18:190, pL 96, 1916. Inga Java Pittier, in Trab. Mus. Com. Venez. 5:261. 1929. Large, spreading tree about 16 m. tall, the branchlets lightly or moderately cinereous-tomentose. Leaves moderately large, 8-foliolate; petiole scarcely 1 cm. long, terete, tomentose, swollen basally; rachis up to 10 cm. long, narrowly alate (Ul) 196 [Vol. 37 MISSOURI (sometimes obscurely so on lowermost rachial internodc), tomcntose along th axis, bearing above at the rachial nodes a sessile, pateUiform gland; leaflets normally 4 pairs, elliptic to elliptic-lanceolate, 10-20 cm. long and 5-7 cm. wide in the terminal pair, usually short-acuminate apically, obtuse to blunt and somewhat in- equilateral basally, heavily pubescent on costa and chief veins, loosely appressed- strlgose or subglabrous and "glandular" elsewhere; stipules linear, about 5 mm. long, caducous. Inflorescence of 1-2 pedunculate spikes borne from subterminal axils; peduncle 3-6 cm. long, pubescent; floriferous portion of spike usually 1-3 cm. long, the flowers very congested; bracts small, caducous. Flowers small, white, sessile; calyx tubular-funnelform, about 4 mm. long, subglabrous or sparsely stri- gose, striate; corolla tubular, 7-8 mm. long, hirsute-strigose; stamens 15 or more mm. long, the staminal tube scarcely exserted. Legume broad, flat, up to 16 cm. long and 5 cm, wide, slightly margined, transversely striate, subglabrous in age, blackish, several-seeded. Northern South America and Panama. cocLE: Bismarck, Williams jl6. san blas: Perme, Cooper 6jT. The San Bias specimen difl'ers from the Code one (type of /. vwnticoU) in degree of pubescence, size of leaflet, and in other minor ways. The latter probably reflects intermixture with other species towards the northern extremity of its range. The San Bias specimen is nearly identical with the type of L Java (from tree planted for coffee shade in Venezuela). 6. Inga Goldmanh Pitticr, in Contr. U. S. Nat. Herb. 18:198. 1916. Moderate-sized tree, the branchlets ferruginous-hirsute. Leaves large, mostly 8-foliolate; petiole usually about 1 cm. long, aiate, hirsute except lightly pubescent on the wings; rachis up to 20 cm. long, similar to the petiole, the wing about 5 mm. wide from midrachis to margin, bearing short-stipitate, cupuliform glands between insertion of the petiolules; leaflets mostly 4 pairs, ovate to elliptic, up to 25 cm. long and 15 cm. wide, the terminal pair about twice as long as the basal pair, acute and short-acuminate apically, rounded and inequilateral basally, sub- glabrous above except on the midvein (midvein heavily pubescent and usually bear- ing 1-2 small glands basally), laxly pubescent below especially on the veins; stipules broadly ovate, caducous in age. Inflorescence a stout terminal or axillary spike, the peduncular portion about 8 cm. long and hirsute, the floriferous portion up to 6 or 7 cm, long; bracts broadly ovate, scarcely 4 mm. long, hirsute, deciduous In age. Flowers very broad and stout; calyx subcampanulate, about 13 mm. long and almost as wide (apically), densely hirsute-tomentose; corolla tubular-campan- ulate, 2 cm. long or longer, densely villous without; stamens about 6 cm. long, the staminal tube included. Legume flat, about 18 cm, long and 4 cm, wide?, some- what arcuate, stipitate, densely ferruginous-hirsute, the margins not unduly thickened. Panama and Costa Rica, (182) 19 50] FLORA OF PANAMA (Leguminosae) 197 CANAL zone: Frijoles, Allen g22; Gatun, Goldman 1 866; Lion Hill Station, S. Hayes 5g8; Monney? Hill, Lehmann lOOI; near Salamanca, Steyermark ^ Allen 16754. This species has been segregated from Bentham*s original concept of J. Linden- lanuy and without reference to the (European) type of /. Limieniana it is impossible here to pass judgment on the validity of the segregation. Perhaps the most defi- nitive difference given by Pittier between 7. Goldmanii and J. Ltndeniana is the considerably lesser pubescence of the former. Certain herbarium specimens (in fruit) labeled /. mncuna are evidently referable here. 7. Inga Standleyana Pittier, in Contr. U. S. Nat. Herb. 18:204. 1916. Low tree, the branchlets densely dark ferruglnous-tomentose. Leaves large, 8-foliolate; petiole 2-4 cm. long, terete, tomentose like the branchlets, swollen basally; rachis about 11 cm. long, unwinged, similar to the petiole, bearing sessile, urceolate glands above between insertions of the petiolules; leaflets 4 pairs, broadly- ovate or obovate, about 12 cm. long and 7-8 cm. wide in terminal pair (lowermost pair about Yi as long), rounded or obtuse or briefly acute, and very briefly mucro- nate apically, obtuse or rounded and somewhat inequilateral basally, pilose below, subglabrous above except tomentose on the costa and chief veins, reticulate, darker above than below; stipules caducous. Inflorescence of 1-2 pedunculate spikes axillary and subtermlnal from the upper branchlets; peduncular portion 2-4 cm. long, tomentose like the branchlets; floriferous portion equally long, the flowers somewhat distant; floral bracts ovate, 1 mm, long, caducous. Flowers large, sessile; calyx cupular, about 5 mm. long and equally as wide, lanose-tomentose, the teeth scarcely discernible because of the pubescence; corolla funnelform, 17-20 mm, long, gross, lanose-tomentose; stamens about 5 cm. long, the staminal tube barely exscrted; ovary white-pubescent along sutural line; style 5 mm. long; stigma capitate. Legume reported flat and densely rufous-pubescent. Panama. darien: La Palma, Pittier 549^' This species is readily distinguished from others In Panama by the gross, woolly flower. It is said to be quite similar to 7. dysantha of South America, a species of which no specimens have been available for comparison. 8. Inga mucuna Walp. & Duchass. in Walp. Ann. 2:459. 1851-52. Moderate-sized tree, the branchlets dark hirsute-tomentose. Leaves moderate, the leaflets 6- or 8-foliolate; petiole about 2 cm. long, heavily tomentose, terete and unwinged; rachis up to 15 cm. long, with prominent interfoliolar wings about 5 mm. wide from rib to margin, the rib tomentose, the wings more lightly pubescent, with small, sessile, subcupuliform glands between insertion of the leaflets; leaflets apparently 3 or 4 pairs, mostly ovate, up to 15 cm. long and 7 cm. wide terminally, progressively smaller baseward, apically acute and short-acuminate, basally in- equilaterally rounded or obtuse, rather heavily pubescent with curved hairs below and above; stipules triangular-ovate, about 6 mm. long, soon deciduous. In- (U}) 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN I Vol. Z7 florescence of 1 to few? pedunculate spikes axillary from the upper leaves, the peduncular portion hirsute-tomentose, about 4 cm. long, the floriferous portion less heavily pubescent, 3-5 cm. long; bracts ovate-elliptic, about 6 mm. long, caducous. Flowers white, very elongate; calyx tubular, 13-17 mm. long, longitudinally striate or sulcate, subglabrous except at the apex of the teeth, the teeth 1-2 mm. long; corolla 4-5 cm. long, pilose without (sparingly so in age), the hairs 3 mm. long or longer, the lobes somewhat flaring, about 5 mm. long; stamens up to 10 cm. long, the staminal tube cxscrted about 2 cm. beyond the corolla; ovary tetra- gonal and indented on opposite faces. Legume up to 30 cm. long and 5 cm. wide, usually spirally twisted, densely ferruginous-strigose, the margins deeply sulcate along the line of dehiscence. Panama; northern South America? CANAL zone: Barro Colorado Island, Wood worth & Vestal 66 j. darien: Boca de Cupc, Allen SSj; Sambu River, Pit tier 552§, province unknown: ^'Panama," Duchas- saing 8l. The cited specimens certainly seem to belong to section Pseudinga, series LoNCiFLORAE as Considered by Pittier in his revision of Inga. Of this series only one species, /. mncuna, is reported north of South America: thus the species' aflfini- chiefly South American (Brazil especially). Without fruit as a criterion, a specimen of this species might key to (Pitticr's) /. oopbylla, I, RoJngiivz/ana, I. Rcfisoni, 7. Preussii or J. HoUofiii, but it differs from all of these in having the extremely elongate corolla. Brazihan specimens in the herbarium labeled 7. caJantha Ducke (Krukoff 6204, fruit only) appear very similar to the Panamanian speci- mens, and possibly are identical. 9. Inga Hayesii Bcnth. in Trans. Linn. Soc. 30:617. 1875. Feuillcea Hayesii Ktze. Rev. Gen. PL 1:188. 1891. Small tree, the branchlets rather heavily pubescent when young becoming glabrous. Leaves moderate, 4- to 8-foliolatc; petiole 1-2 cm. long, pubescent especially medianly, narrowly winged except at the extreme base; rachis 2-10 cm. long, similar to the petiole except the rachial internodes prominently alate (wing up to 5 mm. wide measured from midrachis), bearing sessile, patelliform or cupull- form glands between insertion of the pctiolules; leaflets 2-4 pairs, ovate, oblong or elliptic, the larger (terminal) ones as much as 15 cm. long and 7 cm. wide, the smaller (basal) ones about 5 cm. long and 2.5 cm. wide, acute, obtuse or even rounded apically, obtuse or rounded basally and not markedly inequilateral, with scattered curved-appressed hairs above and below, the veins prominent below; stipules broadly ovate, 2-3 mm. broad, soon caducous. Inflorescence very con- densed, of 1 to few? axillary spikes from the tips of branches, the peduncular portion usually less than 1 cm. long and obscured by the compact floriferous por- tion 1-3 cm. long; bracts lanceolate, inconspicuous, at most a few mm. long. Flowers whitish, densely compacted; calyx tubular, 6-9 mm. long, subglabrous except for small tufts of hair at the tip of calyx lobes and sometimes the base of (n4) 1950] FLORA OF PANAMA (Lc gummosac ) 199 Fig. 75. higil Haycsii the calyx, longitudinally striate, rather deeply cleft on one or two sides (the teeth thus unequal); corolla tubular-funnelform, 13-16 mm. long, silvery-pilose with- out, the lobes about 3 mm. long; stamens up to 4 cm, long, the stamina! tube included; ovary subglabrous, 4-angled due to lateral Indentations; stigma capitel- late. Legume linear-oblong, up to 17 cm. long and 2 cm. wide, flattened, ycUow- tomentose. Panama. CANAL zone: Ancon Hill, Sciberf II j; Fort Kobe road, Allen iSgO; Mlraflores; P. White 79. PANAMA: Matias Hernandez, Pitticr 6/I4; Pacora, WooJsony Allen ^ Seibcrt 759; Pearl Islands, Allen 2jg;^. The cited specimens comprise an entity in many ways imperfectly matching various descriptions of /. HaycsiL Nevertheless, knowing the extreme variation and intergradation common in In^^a, it would be presumptuous to regard them as a different species. The short-pedunculate spike is perhaps the species' outstanding characteristic, and this, along with the subglabrovis, striate calyx and relatively light leaf pubescence, tends to distinguish /, Hayesii from most (otherwise) similar species. Some specimens of /. spur/a much resemble the cited specimens except for larger flower size and generally greater leaflet number. 5) 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 Fig. 76, Inga specfahilis 10. Inga spectabit.is (Vahl) Willd. Sp. Pi 4:1017. 1806. Mimosa spectabilis Vahl, Skr. Natur. Sclsk. Kjob. 2:219, pi TO. 1792. Irtga fulgent Kunth, Mimos. PI. Legum. 36, pL IT. 1819. Ifiga hicida HBK. Nov. Gen, & Sp. 6:287. 1824. Feiiillcea spectabilh Ktze. Rev. Gen. Pi. 1:184. 1891. Moderate-sized tree, the branchlets lightly pubescent to glabrous and markedly angled, lenticellate. Leaves large, mostly 4-foliolatc; petioles scarcely 1 cm. long, angled or short-winged, sometimes pubcrulent; rachis up to 7 or more cm. long, somewhat puberulent, alate towards the upper part of each rachial internode (the (lU) 1950] FLORA OF PANAMA (Legummosae) 201 wing broadest just below Insertion of tbe leaflets), the wing from subobsolete to 7 or 8 mm. wide measured from midrachis, bearing a large, cupuHform or patelliform gland between insertion of the petiolules, usually puberulent at least medianly above; leaflets 1-3 (usually 2) pairs, elliptic or nearly so, up to 25 cm. long and 15 cm. wide in terminal pair, the basal pair smaller (often lacking in herbarium material), rounded to acute apically, more or less obtuse and inequilateral basally, coriaceous, glabrous above in maturity, subglabrous below, the veins impressed above and prominently elevated below, the alternate lateral veins much shorter than the others; stipules linear to linear-lanceolate or linear-oblanceolate, about 1 cm. long, subpersistent. Inflorescence mostly a terminal panicle of pedunculate spikes (the spikes sometimes soHtary from an axil), the peduncular portion gen- erally 4-5 cm. long and hghtly puberulent, the floriferous portion about 5 cm. long and tomentulose, the flowers more or less congested; bracts nearly elHptic, up to 2 cm. long and almost 1 cm. wide, more or less persistent, tomentulose. Flowers white; calyx tubular-campanulate, about 9 mm. long, pubescent, somewhat cleft to one side, the teeth about 3 mm. long; corolla tubular, about 2 cm. long, tomen- tose, the lobes somewhat flaring; stamens about 4 cm, long, the staminal tube about equalling the corolla. Legume flat, reported up to 60 cm. long and 7 cm. wide, the margins thick but not elevated, glabrous. Costa Rica, Panama, northern South America. BOCAS DEL TORO: Chinqui Lagoon, H. von Wedel 2Sg2, CANAL 70NE: Culebra, PHtier 242J; Las Cascadas, Pitfier 3746, cHiRiQuf: trail from San Felix, Allen 1944. cocle: Bismarck, Williams 383, 584. Panama: Taboga Island, Maxon 6q22. without locality: Cooler ^ Slater 203. 11. Inga panamensts Seem. Bot. Voy. Herald, 117. 18 53. Small tree, the branchlets slender, essentially terete, pubescent when young becoming glabrous In age, conspicuously lenticellate- Leaves large, 2- to 6-folio- late; petiole 4-5 cm. long, terete or somewhat angled above apically, lightly pubescent or nearly glabrous, swollen basally; rachis as long as (or longer than?) the petiole, similar to the petiole except narrowly winged, bearing at Insertion of the petiolules small, concave, sessile glands; leaflets mostly 2 pairs in isotype ex- amined, broadly oblong to elliptic-obovate, terminal ones up to 15 or more cm. long and almost 10 cm. wide (basal leaflets % or less this size), obtuse or rounded and somewhat Inequilateral basally, rounded and briefly mucronate apically, gla- brous above, scantily pubescent (especially on veins) and "glandular'' below, the veins prominently elevated below, the lateral ones about 12 pairs; stipules ovate, 6 or 7 mm, long, persistent. Inflorescence of axillary, pedunculate spikes 3-7 cm. long; peduncular portion 2-4,5 cm. long, lightly pubescent, striate; floral bracts Hnear, about 1 cm. long, persistent, lightly pubescent. Flowers moderate, whitish?; calyx cupular-tubular, 5-8 mm. long, lightly pubescent, striate, the teeth prominent and Irregular or unequal; corolla tubular-funnelform, about 14 mm. long, golden appressed-pilosc; stamens about 4 cm. long, the staminal tube some- bout (n7) 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 what exserted; ovary nearly glabrous; style exceeding the stamens; stigma capitcl- late. Legume (reported) up to 15 or more cm. long and about 4 cm. broad, thick, glabrous, the margins very prominent. Panama. CANAL ZONF?: Cruccs, Scemann 40^. Apparently this species has been collected but once. We have examined the Gray Herbarium isotype of the original collection which appears to be an aberrant form of a large Central American complex. The fact that no undoubted second collection has been made, although the type is from a comparatively well-known area, would support the belief in its aberrancy. Yet the name must take precedence over its more common varieties as here interpreted, in spite of being less "typical'* of the complex as known in lower Central America than are its varieties. Two pairs of leaflets to the leaf is unusual for the group in Panama, although Mexican material frequently shows this characteristic. A number of specimens have leaves with both 2 and 3 or 3 and 4 pairs of leaflets. Many taxonomists will doubtless take exception to my condensation and reduc- tion to varietal status (under /. panamcusis) of numerous "species'* of this com- plex. It is conceded that the types of such varieties scarcely resemble the species, and that certain extremes among them more resemble other accepted species than /. panafficJisis. Yet when all available material is examined, no clear-cut specific demarcation can be found within the complex. After days of fruitless effort at- tempting such, it was deemed wiser to regard as of varietal status some fairly con- stant but difficultly distinguishable entities. These embraced most of the species of Pittier's series Tetragonae. One of the entities (L panamensis var. clavata) appears fairly clear-cut. It is found throughout Panama (especially in Bocas del Toro) and in Costa Rica. It differs from the species in having a much longer flower, but vegetative differences (except leaflet number) arc scarcely apparent. Another less clear entity (I. pana- mensis var. Pittieri) is found in central and western (Chiriqui) Panama and ad- jacent Costa Rica. It has a flower only a little longer than in the species but the leaflet shape is different and its lower surface more heavily pubescent. Like J. panamensis var. clavata, it normally has 3 pairs of leaflets per leaf. A third variety (L panamensis var. Rodriguez/an^) , including a very variable group of "species" and specimens from Costa Rica and Nicaragua to Mexico all grading into one another, is proposed as a convenient catch-all within the complex for northern Central America. The type specimen (for the species from which the varietal name is taken) is quite unlike 7. panamensis — 4 pairs of leaflets, mod- erately long flowers, heavy pubescence — yet It seems to grade quite imperceptibly into iorms scarcely distinguishable from J. panamensis var. Pittieri. In flower size var. Kodrigueziana is intermediate between var. cJavafa and var. Pittieri. Some of the synonyms listed for it (viz. 7. Purptisii) may with further collections merit varietal standing of their own. (lU) 1950] FLORA OF PANAMA (Lcgiiminosac) 203 All in all, the species and its varieties comprise a difficult complex, on which it Is perhaps profitless to devote inordinate time in the attempt to make detailed taxonomic distinctions. Ha. Inga panamensis Seem. var. Pittieri (MicheH) Schery, comb. nov. Irtga Pittieri Micheli, in Bull. Herb. Boiss. 2:446, pL 1 3. 1894. Inga Jimeneziana Pittier, in Contr. U. S. Nat. Herb. 18:208. 1916. This variety differs from the species In having a slightly longer flower (corolla 20 mm. long vs. 14 mm. in species) and in having normally 6 large, tapered, more pubescent leaflets. The legume is linear-oblong, up to 25 cm. long and 2.0—2.5 cm. wide and thick, tetragonal, the valves deeply sulcate medianly (dry material), the Juncture of valves slightly protruding from the otherwise flat face presented by the valve margins, essentially glabrous, scarcely stipitate basally. Fig. 77, Inga patjamcnsis var. Pittieri t t _^*- *, (1S9) [Vol. 37 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama and Costa Rica. CANAL ZONE: Gatun, Wctmorc d Ahbe S4'> Quebrada La Palma, Dodge d Allen 7340 CHiRTQui: San Felix, Pit tier 5452. cocle: Bismarck 48g. Fig. 78. Tnga panavicns'n var. cJavafa lib. Inga panamensis Seem. var. cla\ata, Schery, var. nov. Arbor ab specie differt foliolis subtus frequenter minus pubescentibus, floribus longioribus, ca. 3 cm. longis, staminibus 7—8 cm. longis, tubo plerumque ex- sertissimo. 1?50] FLORA OF PANAMA (Lcgumtnosae) 205 On vegetative characters this variety is scarcely distinguishable from the species and the preceding variety. The flowers, however, are considerably longer than in the species, the corolla-tube extending well beyond the calyx and making the corolla in bud appear clavate. This same difference in degree is seen in the stamens and staminal tube, which are almost twice as long as in the species. Costa Rica and Panama, BOCAS DEL TORO: Changuinola Valley, Dunlap 525; Cooper ^ Slater 1 08; Water Valley, H. von Wcdd 6^2 (Herb. Missouri Bot. Card., type), 849, 2/4Q (Herb. Missouri Bot. Gard., cotypf). canal zoinje: Barro Colorado Island, Kenoyer 368; Darlen Station, Standley 31610 (sterile); Empire, Piper 31 12, cocle: Bismarck, Williams ^Sp (sterile)?. darien: Cituro, Williams 6J3. Fig. 7^. Inga pa7jamctisis var. Rodrif^ueziana lie. Inga panamensis Seem. var. Rodrigueziana (Pittier) Schery, comb. nov. Jnga Preussii Harms, in Fcdde Rep. Sp. Nov. 13:420. 1914, ex char. Inga Rodrigucziana Pitrier, in Contr. U. S. Nat. Herb. 18:209. 1916. h7ga bioUeyana Pittier, loc. cit. 207. 1916. Inga Purpusii Pittier, loc. cit. 199. 1916. This variety is distinguished from the species in having a more robust flower approximately 2.5 cm. long, and a more heavily pubescent leaflet (frequently sub- tomentose below). In keeping with a very broad interpretation of /• pafiamensis, the synonyms listed above should constitute a variety of that species. This variety is not known to occur in Panama, but Is found In southern Mexico, northern Central America, and perhaps aberrantly in Costa Rica (the type of /. bioUcyana of Costa Rica ap- pears to be but an extreme of this variety). Besides the type specimens of /. Rod- r'lgneziana and /. bioUeyana , specimens examined include: (191) 206 ANNALS OF THE MISSOURI IVoL. 37 Nicaragua: Baker ^j, Salvador: Caldcnht lyi; Pittier 1974; Standley IQlOg, Guatemala: Record 47; Tuerckbeim (J. D. Smith) 7S55. Mexico: Purpus 6811. Inga Prcnssiiy in spite of priority, is not chosen as the name-bringing synonym for the variety for the reason that the type has not been available for examination. 12. Inga alatopetiola Schcry, sp. nov. Arbor?, ramulis juvenalibus ferrugineo-pubesccntibus; foliis magnis, petlolis ca. 2 cm. longis, prominenter alatis, costa pubcscente, rhachibus ca. 15 cm. longis, glandulam substipitatam concavam ad insertioncm parium foliolorum ornatis, folio- lis 4-jugatis, ovato-cUipticis, ad 20 cm. longis, 8 cm. latis, basalibus parvioribus, brevc-acuminatis, basi inacqualitalitcr obtusis vel rotundatis, subglabris costa supra excepta, subtus leviter pilosis, ncrvis subtus prominenter elevatis supra indcntatis; inflorescentia axillari, ca. 7 cm. longa, pedunculo ca. 3 cm. longo, ferrugineo- tomcntoso; bractcis lincaribus, ca, 1 cm. longis persistentibus tomentosis; calyce tubulato, ca. 15 mm. longo, striato, adprcsso-piloso, dcntibus brevibus; corolla tubulata, ca. 3 mm. longa, dense pilosa; staminibus ca. 5 cm. longis, tubo vix exserto; Icgununc ignoto. Tree?, the young branchlets ferruginous-pubescent. Leaves large, once-pinnate, with 4 pairs of leaflets; petiole about 2 cm. long, alate, the wings subglabrous, the axis pubescent; rachis about 15 cm. long, similar to the petiole, prominently winged between all pairs of leaflets, bearing short-stipitate, concave glands at insertion of each pair of leaflets; leaflets ovnte-elliptic, terminal pair up to 20 cm. long and 8 cm. wide, basal pair scarcely half this si/e, short-acuminate aplcally, inequilaterally obtuse or rounded basally, subglabrous above except on the costa, sparingly pilose below, the veins Indented above, elevated below, coarsely reticulate. Inflorescence an axillary pedunculate spike, peduncular portion 3-4 cm. long, ferruginous- tomcntose, florifcrous portion similar, comparatively few-flowered, bracts essentially linear, densely pubescent, persistent. Flowers large, sessile; calyx tubular, about 15 mm. long, striate, appressed-pilose, the teeth short and somewhat unequal; corolla nearly tubular, about 30 mm. long, densely villous or pilose without, the lobes relatively short and slender; stamens approximately 5 cm. long, the staminal tube barely exsertcd; style elongate, exceeding the stamens, the stigma capitate. Legume not known. Panama. BOCAS DEL TORO: Almlrantc ret^Ion, Cooper & Slater 65 (U. S. Nat. Herb., type). The type specimen is somewhat fragmentary, but is sufficient to indicate dis- tinctiveness from known Panamanian species. Floral characters indicate affinities with f. pa?7awcf?sis (and Pittier's series Tetragonae), and future collections may link it to 7. panamerjsis var. RoJrigticzia7ia, The winged petiole is unusual in higa, and serves, along with the persistent, linear bracts, the large "panamensls-type" flowTrs, and various leaf characters, to distinguish the species rather easily. (192) 19S0] FLORA OF PANAMA (Lcgnmhosac) 207 Fig. 80. Inga Oerstcdiana 13. Inga Oerstediana Benth. in Seem. Bat. Voy. Herald, 117. 18 52-57. A Feuillcea Oerstcdiana Ktze. Rev. Gen. Pi. 1:188. 1891. higa criorhachis Harms, in Fedde Rep, Sp. Nov. 13:525. 1915. Inga cobanensis Pittier, in Contr. U. S. Nat. Herb. 18:188. 1916. Ifiga Tjierckheimii Pittier, loc. cit. 192. 1916, based on same type as J. cobancush, Inga chiriqiiemh Standi, in Field Mus. Publ. Bot. 22:78. 1940. Small or moderate tree, tKe branchkts dark ferruginous-tomcntose, becoming less densely so in age. Leaves large, normally 8-foliolate; petioles up to 5 cm. long, terete, densely dark ferruginous-tomcntose; rachis up to 15 cm. long, tomentose like the petiole except very lightly so on the wings, conspicuously alate for the uppermost rachial internode, progressively less alate in penultimate and succeeding (lower) rachial internode (s), bearing between Insertion of the petlolules large or small sessile glands (or these sometimes lacking), the glands laterally extended and markedly concave; leaflets 4 (sometimes 3 or 5) pairs, asymmetrically elliptic, ob- long or ovate, 7-16 cm. long and 3-8 cm. wide, rounded or subtruncate basally, inequilaterally acute apically, lightly pubescent above except on the veins where (193) 208 [Vol. 37 MISSOURI heavily so, moderately pubescent below except densely tomentose on the elevated veins, reticulate; stipules caducous. Inflorescence of 1 to few pedunculate spikes from foliate or young defoliate nodes, the spikes 3-7 cm. long, florifcrous for the upper 1-3 cm., densely tomentose Hke the petiole; bracts ovate, small, promptly caducous. Flowers quite congested, white; calyx campanulate-cupuliform, 3-6 mm. long, ferruginous-hirsute or pilose, the teeth somewhat unequal; corolla sub- tubular, about 1 cm. long, somewhat flaring apically although constricted for the lower half its length; densely ferruginous-hirsute or pilose, the lobes about 2 mm, long; stamens 2.0-2.5 cm. long, the staminal tube included within the corolla; ovary linear, the style exceeding the stamens. Legume apparently Hnear, 12 or more cm. long and less than 1 cm. wide, subterctc and longitudinally sulcate, sessile, ferruginous-tomentose. h So CHiRiQuf: Bajo Mona, Daihlson Jjd; Boquete, Pitticr JIJO; Cerro Punta, Allen 1 573; Vole an dc Chiriqui, Daihhon g28. Again It IS impossible without reference to the type to establish with certainty the correct name to the specimens cited. If 7. OersfeJiana should prove not to apply, 7. criorhachis could be used. Standley and Steycrmark (Fieldiana, Bot. 24:40. 1946) consider 7. cobavcnus and 7. Tuerckbefmii as synonyms of 7. Micheliava; yet It seems hardly possible to regard 7. Micheliana a synonym of our 7. Oersfcdiaria, although 7. cobancnsh and 7. Tuerckhciwii likely are this (L OersfeJiana^). Pittier, among others, separated some of the possible synonyms given above into separate sections of the genus on the basis of fruit characters. Only one of the cited speci- mens is with fruit, and on vegetative appearance the others seem to be the same species. It would seem that fruit characters have been overstressed in separating the genus Into often unworkable sections, especially in that fruiting specimens are so rarely available. Almost certainly when extensive fruiting specimens are gathered, variation and degrees of intermediacy will be found, perhaps indicating fruit characters to be little more rehable and certainly less practical taxonomically than floral and vegetative characters. 14. Inga EDULis Mart, in Flora 20:BeibL 113. 1837. Mimosa Ynga Veil. Fl. Flum. Ic. 11:/. J. 1827, fide Benth. Ifjga vera HRK. Nov. Gen. & Sp. 6:289. 1824, non Willd., fide Benth. Fcuillcca CiluJh (Mart.) Ktze. Rev. Gen. PI. 1:187. 1891. Small or medium tree, the branchlets ferruglnous-tomentulose, lenticellatc and angled or ridged. Leaves large, usually 10- or 12-foliolate; petioles mostly 2-4 cm. long, terete, rufous-tomcntose; rachis almost 20 cm. long, tomentose like th petiole, conspicuously alate, the wing usually 5 or more mm. wide on each side of the racbis, bearing prominent, oval, cupullform or patelliform glands be- tween insertion of the pctlolules, the glands broader than long; leaflets 3-6 pairs, apparently most commonly 5 or 6 pairs in Panama, ovate to elliptic, up to 15 cm. long and 5-7 cm. wide in terminal leaflets, usually about Yz as large in basal leaflets, (194) 1950] FLORA OF PANAMA (Legummosae) 209 acute and somewhat acuminate apically, mostly rounded basally, lightly tomentose below especially on the veins, sparsely appressed-pubescent above, the veins elevated and conspicuously reticulate below, conspicuous but not elevated above; stipules caducous, reported about 5 mm, long. ped spikes from the leaf axils, or often paniculate by insertion of such spikes at sub- terminal defoliate nodes; spikes mostly 6-8 cm. long, floriferous for the upper 2-3 cm., the flowers moderately congested; bracts ovate, about 5 mm. long, caducous in age. Flowers white, moderate; calyx turbinate, apparently 7-8 mm long in Panama, appressed short-pilose, the teeth regular and 1-2 mm. long; corolla tubular, almost 2 cm. Icng, relatively slender and Httle expanded apically, densely ascending- pilose, the lobes subequal and 2-4 mm. long; stamens mostly 4 cm. (or up to 5 cm.) long, the staminal tube slightly exserted; ovary glabrous, the style slightly exceeding the stamens. Legume as much as 1 m. long and up to 2 cm. wide or wider, ferrugineous-tomentose, somewhat tetragonous or subterete, the margins almost entirely covering the faces. Mexico to Panama; South America to middle Brazil. BOCAS DEL TORO: Almirante, Cooper & Slater 38; Changulnola Valley, DunJap 220; Chiriqui Lagoon, H. von Wedel WOO; Water Valley, H. von Wedel IO96. This species is very variable, and the description given here is mostly drawn from the Panamanian material. Probably synonymous with Z. edulis are L Endlichii Harms and /. latibracteata Harms (ex char.), and possibly a number of other "species" which intergrade with this. The Panamanian specimens cited resemble * more closely collections from South America (Amazon Valley) than most collec- tions labeled 7. cdidh from northern Central America and Mexico, Evidently no well-established distinction between J, edtdh and /. sjnnia has been apparent to most authors. Examination of all available specimens from Mexico to Brazil shows specimens, practically indistinguishable from each other, determined rather indis- criminately as either 7. edulh, L spiiriay or even other species. I. ediilis is here in- terpreted as having a more slender, less stocky corolla and a generally smaller calyx than 7. spuria. The leaflets hkewise appear significantly larger (especially wider) in 7. edulis than in 7. spuria (in Panama). The H. von Wedel specimens cited appear somewhat atypical (more leaflets; corolla more elongate). 14a. Inga edulis Mart. var. minutula Schery, var. nov. Arbor foliis speciei similibus; inflorescentiis prominentibus, floribus specie multo brevloribus, bracteis obovato-oblanccolatis, 3-4 mm. longis. Ferruginous-tomentulose tree with large 6- to 10-foliolate leaves. Inflorescence of 1 to few pedunculate spikes 3-8 cm. long (floriferous portion 1-5 cm.) from either foUatc or defoliate axils; bracts 3-4 mm. long; flowers small; calyx cinereous, appressed-pubescent, 4 mm. long; corolla cinereous, short-pilose, about 8 mm. long; stamens about 3 cm. long. Legume tetragonal-subterete, 15-20 cm. long, 10-15 mm. wide, brownish-tomentose, somewhat constricted between the seeds, the valves sulcate. Panama; Honduras? (195) 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 Fig. 81. Ifjga cJnlis var. minutula BocAs DEL TORO: Changuinola Valley, Dunlap jS2. canal zone: Barro Colorado Island, Zctek 357S; Kcnoyer 3 /'I; Gamboa, Alh-N igY2; Gorgona, VUiicr 26g6. El Real, Allen g6S (Herb. Missouri Bot. Card., type). DARIEN: This variety is well marked from the species by the considerably smaller flow- lat into crs, and Is perhaps worthy of specific rank except that it grades somewl both /. edulis and /. OcrstcJtaua. It resembles /. edulis var. grcnaJcnsh Urb., but differs from that variety in the longer inflorescence and the obovate or oblanceolate bracts. The cinereous flower ordinarily serves to distinguish /. eJnlh vunntnlu from 7. Ocrstcdiana. 15. Inga spuria H. & B. ex Willd. Sp. PL 4:1011. 1806. Mimosa spuria Poir. in Lam. Encycl. Meth. Suppl. 1:40. 1810. Feuilh'ea spuria Ktze. Rev. Gen. Pi. 1:189. 1891 7/;^^ fissicalyx Pittier, In Contr. U. S. Nat. Herb. 18:213, 1916. Small- to moderate-sized tree, the branchlets ferruginous-tomcntose, b glabrous in age. Leaves moderate to large, with few to several pairs of leaflets; ecoming (196) 1950] FLORA OF PANAMA (Legummosac) 211 petiole sKort, scarcely exceeding 1 cm., tomentose like the young branchlets, terete; rachis up to 15 cm. long, tomentose, conspicuously alate, bearing a small sessile, cupuliform or patelliform gland between insertion of the petiolules; leaflets 4-9 pairs (usually 6 or 7 pairs), ovate-lanceolate to narrowly obovate, the terminal ones 7-15 cm. long and 2—5 cm. wide, becoming progressively smaller downward, the basal pair usually only J/4 the size of the terminal pair, usually acute or slightly acuminate ;?pically, commonly obtuse or narrowly rounded basally, slightly pubes- cent above, somewhat more heavily so below, the veins prominent below but not above; stipules broadly linear, a few mm, long. Inflorescence of 1—3 pedunculate spikes from each of several leaf axils, the spikes 4-12 cm, long, floriferous in tlie upper few cm., tomentose, the flowers subscssile to somewhat pedicellate, more or less congested, white; calyx tubular, 10—20 mm. long, 3—6 mm. broad, cinereous- or golden-tomentose, the lobes 3—5 mm. long; corolla usually close to 2 cm. long, pilose without, the lobes 4-5 mm. long and slightly flaring; stamens many, fre- quently 5-6 cm. long, the staminal tube invested by the corolla. Legume linear, reported up to 30 cm. long and 1,5 cm. wide, subterete or angled, fulvous-tomen- tose, the margins thickened and expanded over the faces. Mexico to northern and eastern South America. CANAL zone: Miraflores, P. White /p; Sabana of Panama, Pit tier 2^j8; Trinidad River, Pitticr 3973; Victoria Fill, Allen 1704. chiriqui: Boqucte, Davidson 823, cocle: La Pintada, Hunter & Allen jl8; Penonome, Williams 1 37, 334, herrtra: Ocu, Allen 4o6g, PANAMA: Chcpo, Hunter ^ Allen 8g; Juan Diaz, Allen g3g; R. Las Lajas, Allen 160S; Las Sabanas, Stafulley 5/(^25. veraguas: Canazas, Allen 160, A very variable species, forming a vast complex from Mexico through northern South America. It intergrades with other (both doubtful or recognized) species and is difficult to delimit precisely. Single specimens of two extremes of this species may appear quite distinct; yet examination of many specimens will show all degrees of intermediacy and no clear-cut differences. Characters such as ab- solute calyx size, number and width of leaflets, presence or absence of pedicellar portion of calyx, etc., normally somewhat constant in most species, show little constancy or correlation in 7. spuria. Three Mexican "species," 7. eriocarpa Benth., 7. oop/jylla Riley, and J, sciadoclendron Harms, should probably be reduced to synonymy with 7. spvria and likely combined to form a variety of the species. 7. xaJapensis Benth., of Mexico and Central America as far south as Costa Rica (fide Pittier), is also in all probability 7. spuria or a variety of it, but It is not listed among the synonyms because the type has not been examined. Two centers of divergence in this complex seem discernible. One apparently occurs in south-central Mexico, and may be typified as the '"7, eriocarpa influence." Here the flowers are generally stockier, the leaflets broader, the corolla pubescence less rigid, the legume stout., This would constitute the "variety" above suggested, to which 7, eriocarpa would bring the oldest name. All degrees of gradation occur to the "typical" 7. spnria. The '7. eriocarpa influence" is particularly noticeable in Mexico and transitionally in^ Guatemala, but is little apparent in Panama. A 097) [Vol. 37 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN second less clear center of divergence seems to occur near Honduras or British Honduras. Here slender flowers, narrow* and peculiarly veined leaflets, and slender legumes are the rule. This influence is probably felt in Panama as well as northern Central America and Mexico. In addition, throughout its range, other species, 7. ednlis particularly, tend to Intergrade with J. spuria. As with a number of species of Jnga, L spuria is sometimes planted for coffee shade. 16. Inga pauciflora Walp. & Duchass. in Linnaea 23:746. 1850. Small tree, the branchlets fcrruginous-tomentose-hirsute. Leaves moderate, mostly 6- or 8-foliolate; petiole short, up to 1.5 cm. long, heavily pubescent like the branchlets, unwinged; rachis up to 8 cm. long, ferruginous-pubescent, alate (the wings, margin to margin, 4-10 mm. broad), bearing a small, sessile, cupuli- form gland between insertion of the petiolules; leaflets mostly 3 or 4 pairs, elliptic or nearly so, the terminal pair up to 13 cm. long and 5 cm. wide, the basal pair up to 5 cm. long and 2 cm. wide, acute and often somewhat acuminate apically, sub- cuncate to rounded basally, appressed-pubescent above, villous-tomentose below especially on the prominent veins; stipules ovate, about 4 mm. long, pubescent, early caducous. Inflorescence 1-2 axillary or terminal pedunculate spikes, the peduncular portion up to 3 cm. long and ferruginous-pubescent, the floriferous portion about as long or slightly longer; bracts very small, short and broad, caducous. Flowers several, white; calyx broadly tubular, 8—10 mm. long, tomentu- lose, striate; corolla tubular, about 15 mm. long, pilose; stamens almost 4.5 cm. long, the staminal tube included; stigma capitellate. Legume (young) densely ferruginous hirsute-tomentose, terete, becoming sulcate? or possibly flattened? in age. Panama. CANAL ZONE: Ancon Hill, Williams J2; Barro Colorado Island, Woodson 6 Schery g6g; Chiva-Chlva trail. Viper 572§, veraguas: Santa Fe, Allen 4417. The Woodson & Schery g6g specimen Is in very young fruit. Judging from this immature fruit the specimen is likely not 7. eriocarpa, L sptiria or similar species, although certainly the species grades into the /. sptiria complex. The fruit of /. pauciflora has not been known. Bentham regarded 7. pauciflora as a synonym of /. verUy and indeed a number of difficultly separable species might profitably be combined under that older name. 17. Inga Saffordiana Pittier, in Contr. U. S. Nat. Herb. 18:176. 1916. Small, slender tree, the branchlets dark setose-villous. Leaves large, 8- or 10- foliolate; petiole about 2.5 cm. long, terete, setose or sparsely villous; rachis up to 20 cm. long, unwinged, pubescent like the petiole, bearing long-stipitate glands above at the rachial nodes; leaflets 4 or 5 pairs, oblong to obovate-lanceolate, about 14 cm. long and 4 cm. wide in terminal pair (lowermost pair only ^^-% this size), tapered but blunt basally and inequilateral, acuminate apically, glabrous above, sparsely setose on the margins and veins below, the veins moderately prominent below, the lateral ones arcuate and subconfluent; stipules subulate, up to 12 mm. (198) 195 0] FLORA OF PANAMA (Leguminosac) 213 long, setose-villous. Inflorescence axillary or borne on older defoliate wood, umbellate, pedunculate; peduncles up to 13 cm. long, villous-setose; pedicels sim- ilarly pubescent, almost 2 cm, long; floral bracts similar to the stipules, subpcr- sistent. Flowers incompletely known; calyx apparently 7-8 mm. long, setose- villous, tbe teeth elongate and narrow; corolla tubular, about 15 mm. long, setose-villous, especially apically. Legume broadly linear, about 30 cm. long and almost 3 cm. wide, cuneate basally, flat and thin, lightly margined, densely dark villous, about 18-seeded, Panama. darien: Sambu basin, Pitlier j6/6. A distinctive species with unusual pubescence, long-stipitate rachial glands, and long-pediccllate, umbellate flowers. 18. Inga multijuga Benth. in Trans. Linn. Soc. 30:615. 1875. FeuHlcea mult:}uga Ktze. Rev. Gen. PL 1:188. 1891. Tree to several m., the branchlets densely ferruginous-tomentulose. Leaves large, with a variable number of leaflets (5-10 pairs, fide N. Am. Fl.), 6-9 pairs in Panamanian specimens seen; petiole less than 2 cm. long, essentially terete, tomentulose like the branchlets; rachis up to 2 dm. long, similar to the petiole, not winged, bearing a sessile, cupuliform gland 1-2 mm. in diameter between insertion of the petiolules; leaflets few to several pairs, oblong or elliptic to occasionally ovate-lanceolate, mostly about 10 cm. long and 3—4 cm. broad in the terminal leaflets, Yz or less this size in the lowermost leaflets, acute or short-acuminate apically, rounded or obtuse and somewhat Inequilateral basally, lightly strigose- puberulent or subglabratc above, rather densely pubescent and paler below; stipules apparently ovate, about 2 mm. long, caducous. Inflorescence of pedunculate spikes up to 6 cm. long, solitary, geminate or fasciculate from the axils of the leaves, the floriferous portion scarcely 2 cm. long; bracts ovate, very small, ferruginous- tomentose like the axis. Flowers congested, splcate on the axis, whitish; calyx tubular, up to 12 mm. long (reported as short as 5 mm. In Guatemalan and Costa Rican specimens), usually cleft on one side for as much as % its length, puberulent or tomentulose without, the teeth minute; corolla elongate, tubular, 20—25 mm. long, appressed-villous without, the lobes about 2 mm. long; stamens many, almost 4 cm. long, united below into a tube about equalling the corolla; style somewhat exceeding the stamens, the stigma expanded. Legume variously reported glabrous or tomentulose?, flat or subterete?, up to 25 cm. long and 1 cm. wide?, the valves expanded and costate at the edges. Honduras to Panama. CANAL ZONE: Barro Colorado Island, Bangham 528; **Chagres," Fendler jl; Margarita Swamp, Maxon ^ Valentine 7058. veraguas: Canazas, Allen 1^4- The usual concept of this species seems to include a remarkably variable group of specimens. Certainly various authors have been at great divergence In writing (199) 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 descriptions of it, and specimens seen show great differences in leaflet number, calyx size, etc. Dimensions given by Pittier (Contr. U. S. Nat. Herb. 18:187) for the FtucUer 5/ specimen at the Gray Herbarium in no way approximate those for Fcfti/lcr 51 at the Missouri Botanical Garden. Yet in spite of such lack of constancy the species seems to form an unit distinguishable from most other Ingas within its range (especially by the large, elongate corolla, the several pairs of leaf- lets, the wingless rachis, and the spicatc inflorescence). 19. Inga cocLEENSis Pittier, in Contr. U. S. Nat. Herb. 18:211. 1916. Tree, the branchlets densely dark rubiginous-tomcntosc. Leaves large, 12- foliolate; petiole about 2.5 cm. long, terete, tomcntose like the branchlets; rachis about 16 cm. long, unwinged, pubescent like the petiole, bearing above at insertion of the petiolules a large, sessile, oval, concave gland- leaflets 6 pairs, mostly elliptic- lanceolate, the terminal ones about 13 cm. long and 4 cm. wide, the basal pair somewhat smaller, rounded or obtuse basally, acute and somewhat acuminate apically, rather coarsely short-pubescent above except tomcntose on the costa, tomcntose below, the veins elevated below and slightly impressed above, reticulate; stipules not known. Inflorescence of axillary pedunculate spikes; peduncle 4-5 cm. long, tomcntose like the branchlets; floriferous portion thick, about 4 cm. long, the bracts caducous. Flowers sessile; calyx tubular, 6—7 mm. long, rubiginous- pubescent; corolla not known. Legume (immature) slender, terete by thickening of the margins, longitudinally sulcate, ferruglnous-tomentose, twisted. Panama. COCl4: Bismarck, Williams 40J, The species is known only from a broken fragment found on the ground, in- cluding one complete leaf and two or three Immature fruits. A photograph of the type shows resemblance to the rather hazily defined J. midtiju^a, of which further collections may prove 7. cocleense to be a synonym. 20. Inca RuiziANA G. Don, Gen. Hist. Dichl. PI. 2:391. 1832. Inga fagifoliu G. Don, loc. cit. 391. 1832, non Willd., fide Bcnth. Inga foliosa Bcntli. in Hook. Jour. Bot. 4:597. 184 5. Fcuillcca Kuiziana Ktze. Rev. Gen. PI. 1:189. 1891. Inga confusa Britt. & Rose, in N. Am. Fl. 23:5. 1928. Tree to 20 or more m., the branchlets conspicuously lenticellate and fcr- ruginous-pubcrulent towards the tip. Leaves large, even-pinnate, generally 12- foliolate; petioles 1-2 cm, long, terete or nearly so, ferruginous-pubcrulent, swollen basally; rachis about 15 cm. long, like the petiole except margined laterally above, bearing disc-like, sessile, generally depressed glands 1-2 mm. in diameter between insertion of the petiolules; leaflets usually 6 pairs, mostly elliptic, the uppermost often obovate-elliptic and up to 25 cm. long and 8 cm. broad, the lowermost usually only \(^ or 5/4 this size and somewhat ovate, short-acuminate apically, obtuse and somewhat inequilateral basally, the veins prominently reticulate and (200) 1950] FLORA OF PANAMA (Leguminosae) 215 Fig. 82. Ifiga Ruiziana puberulent below, lightly indented above, the leaflet subglabrous and drying darker above; stipules caducous. Inflorescence terminal and subterminal, tomcntulose, of 2-3 pedunculate spikes fasciculate from each of several nodes or axils, the spikes 3-4 cm. long, the floriferous portion ovate and about 1 cm. long. Flowers con- gested, white, subsessile; calyx subturbinate, 3-4 mm. long, contracted gradually baseward, densely puberulent; corolla 7-8 mm. long, strigose-pubcrulcnt; stamens many; filaments almost 2 cm. long, united below for at least half their length, the tube slightly exceeding the corolla; anthers minute. Legume reported 10-15 cm, long and 2-2.5 cm. wide, flat, rounded apically and basally, somewhat pubescent, the margins elevated. (201) [Vol. 37 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN Nicaragua to Peru and Brazil. CANAL zone: Gamboa, Pittier 6j20; Peluca Hydro^raphic Station, Sieyermark Gf Allen I/2/I; Qucbrada Salamanca, Dodge, Steyermark ^ Allen s. n, (1934). colon: R. Fato, Pittier 391 7> darien: Marraganti, Williams 768; Sambu River, Pittier 333 3» bocas del TORO: Chiriqui Lagoon, H, von Wedel 2012^ 2677; Water Valley, H. von Wedel 844. Although the type of the species is unavailable for examination, various descrip- tions and keys are precise enough to indicate /. confiisa Britt. & Rose as synonymous with L Ruiziana G. Don. Certainly herbarium material of the two species appears identical. 2L Inga microstachya Britt. & Killip, In Ann. N. Y, Acad. Sci. 3 5:115. 1936. Unarmed tree, the branchlets fulvous-pubcrulent when very young, soon be- coming glabrous or subglabrous, very pronouncedly raised white-lenticellate, some- what flcxuous, subangulate below the nodes because of dccurrcnt ridges from the axils. Leaves large, the leaflets 4 or 5 pairs opposite on the rachis; petioles mostly 1-2 cm. long, terete except briefly ridged laterally, subglabrous or lightly pubcru- lent; rachis up to 12 or more cm. long, unwlnged but ridged laterally especially below the foliolar nodes, sulcate above, puberulent to subglabrous, bearing a promi- nent, sessile, cupulatc gland at insertion of each pair of leaflets; leaflets mostly elliptic-lanceolate, up to 18 cm. long and 7 cm. wide in the larger (terminal) leaflets, the basal leaflets perhaps one-third as long and wide, acute- acuminate apically, Incquilaterally acute or obtuse basally, glabrous and lustrous above, the venation visible but not as prominent as below, glabrous below except often puberu- lent on the main veins, the venation prominently reticulate; stipules linear, early caducous. Inflorescence of fasciculate clusters of pedunculate spikes from terminal (Immaturely foliate) and subterminal axils; peduncles 2-5 cm. long, puberulent or tomentulose; spikes short, 1-2 cm. long, the floral bracts minute. Flowers sessile, moderately congested; calyx cupulate, slightly less than 2 mm. long, puberulent, the lobes irregular; corolla funnelform-cylindrlc, about 4 mm. long (in Panama), puberulent at least apically; staminal tube about equalling the corolla; ovary ob- long, short, compressed, glabrous except for a few setae marginally. Legume not known, but presumably flat. Panama and Colombia. CANAL zone: Fort Sherman Military Reservation, Maxon & Valentine 6g7g. This species, a new record for Panama, somewhat resembles both 7. Ktiiziana and 7. Jnmctata of that country, but it lacks the obovatc terminal leaflets of the former and possesses more (4—5 pairs) leaflets than the latter (2—3 pairs). Its exact relationships with South American species such as 7. uobiVn Willd., 7. /r?///- ccllata Bcnth., 7. Caldasiana Britt. & Killip, and others which it resembles, awaits work of the monographer. (202) 19 5 0] FLORA OF PANAMA (Legumiuosae) 217 22. Inga Roussoviana Pittier, in Contr, U. S. Nat. Herb. 18:175. 1916. Inga Schippii Standi, in Field Mus, Publ. Bot. 11:132. 1932. Small tree, the branchlets ferruginous-lenticellate and puberulent towards the tips. Leaves moderately large, mostly 6-foliolate; petioles usually 1-2 cm. long although reported up to 4 cm. long, subterete, puberulent or subglabrous; rachis mostly about 7 cm. long but often longer or shorter, flattened and margined above, lightly ferruginous-puberulent to glabrous, bearing small, knob-like, concave glands between insertions of the petiolules (the glands variable in shape and size, in- frequently suppressed); leaflets 2-4 (usually 3) pairs, generally oblong or elliptic, the uppermost somewhat obovate, up to 18 cm. long and 3-7 cm. wide, the lower- most only about % as large, broadly or narrowly cuneate basally, blunt to short- acuminate apically, glabrous except occasionally on veins below, the veins very prominent below, less so above; stipules reported ovate, about 5 mm. long, fer- ruginous-tomentose, caducous. Inflorescence a terminal or subterminal paniculate shoot, the pedunclcd umbels 1 to few from the defoliate nodes; the bracts and axis ferruginous-tomentose; peduncles 1-2 cm. long; flowers whitish, umbellate from the tip of the peduncle on prominent pedicels usually 4-5 mm. long. Flowers several in each umbel, whitish; calyx turbinate-subcampanulate, 3-6 mm. long, expanded upward, hirsute-tomentose; corolla about 9 mm. long, hirsute-villous without, the teeth about 2 mm. long; stamens about 18 mm. long, united below into a tube about 1 cm. long. Legume linear-oblong, up to 17 cm, long and 2.2 cm. wide, apiculate, rounded basally and stipitate, flat, ferruginous-tomentulose, prominently margined. British Honduras and Guatemala to Panama; Colombia? BOCAS DEI TORo: CHanguinoIa Valley, Stork 2J2; Cooper ^ Slater /J. canal zone: Earro Colorado Island, Bailey ef Bailey 2^0; R. Chagres, Sieyermark H Allen 167S1; France Field, Standlcy 30J2j. cHiRiQuf; San Felix, Pittier 52yo. p The Canal Zone specimens have smaller flowers and shorter pedicels than is usual for the species, and probably represent intergradation towards /. Williamsii Pittier, a species of doubtful specific distinction. /. santanderensis Britt. & Killip (in Ann. N. Y. Acad. Sci. 35:114), of Colombia, is probably nothing more than a robust form of /. Roussoviana^ but having seen no specimens of the former other than the type I cannot pass judgment on the constancy of this difference. The type of L Schippii, also with a more robust calyx than is typical of /. Rottssoiiaria, is nevertheless linked by a series of intergrading specimens. 23. Inga Williamsii Pittier, in Contr. U. S. Nat. Flerb. 18:176. 1916. Small tree, the branchlets ferruginous-lenticellate and puberulent towards the tips. Leaves moderate, mostly 6-foliolate; petioles about 2 cm. long, subterete, puberulent; rachis mostly 6 or 7 cm. long, puberulent like the petiole, flattened and margined above, bearing small glands between insertions of the leaflets or the glands partly or wholly suppressed; leaflets usually 3 pair, more or less elliptic, the upper- (203) [Vol. Z7 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN most up to ;ibout 12 cm. long and 4 or 5 cm. wide, the lowermost only Y^ as large, subcuneate basally, acute to emarginate apically, glabrous except on the veins below, the veins very prominent below, not elevated above; stipules reported lanceo- late, variable in size, caducous. Inflorescence a terminal or subterminal paniculate shoot or reduced to only 1 or a few pedunculate umbels from the defoliate nodes, ferruglnous-tomentose; peduncles 1-2 cm. long; pedicels 1-2 mm. long, arising more or less in umbellate or subcapitatc fashion from the tip of the peduncle. Flowers whitish; calyx about 4 mm. long, densely pubescent without; corolla short, typically 6 or 7 mm. long, villous-tomcntose without; staminal tube scarcely ex- sertcd, the filaments about 15 mm. long. Legume linear-oblong, up to 10 cm. long, about 2 cm. wide, arcuate, flat but thick, densely ferruginous-tomentosc when young. Panama ; Colombia ? CANAL zone: Barro Colorado Island, Wcimorc tf Abbe TOT; Bangham jT2; Tctck 392Q; R. Chagres, Dodge GS" Allen l74og, COLON: Bismarck, Williams 285, This species seems to differ from /. Rottswiiana almost solely in having notably shorter pedicels, and it probably should constitute but a variety of that species. We hesitate, however, to make this reduction until accumulation of more material can prove the frequency of intcrgradation. 7. Mnthi'i Britt. & Killip (Ann. N. Y. Acad. Sci. 35:114) of Colombia, with leaflets up to 22 cm, long, is apparently just a more robust form of 7. William siL 24. Inga gi^antifoHola Schery, sp. nov. Arbor ramulis glabris lenticellatis; foliis permagnis plerumque 3-jugatis, pctiolis aliquanto elongatis ca. 8 cm. longis teretibus exalatis glabris, rhachibus elongatis ad 20 cm. longis, exalatis, glandulam interfoliolarem disciformem sessilem, ornatis; petiolulis ca. 5 mm. longis; foliolis ovato-eHiptlcis ellipticisvc, 13-40 cm. longis, 7-14 cm. latis, abrupte angustatis acuminatisquc, basi late subcuncatis, stipulis non visis; inflorescentia spicibus tenuibus 1 vel pauclbus ex axi, parte peduncular! ca 5 mm. longo, parte florifero ca. 5 mm. longo, bracteis minutis; floribus subsessillbus albis; calyce tubulatl-cupuliformi, ad 2.5 mm. longo, glabro, estriato apice un- dulato; corolla tubulati-infundibuliformi, ad 9 mm, longa, subglabra; staminibus ad 2 cm. longis, tubo aliquanto exserto; ovario glabro, multiovulato; leguminibus risimiliter planis glabris. Small or moderate tree, more or less glabrous throughout, the branchlcts Icnti- cellate. Leaves very large, 6- (sometimes 4?-) foliolate; petiole elongate, almost 8 cm, long, terete and unwingcd, lenticellate, glabrous, callous at the extreme base; rachis up to 20 cm. long, terete and unwingcd like the petiole, bearing an incon- spicuous, flat, sessile gland between insertion of the pctiolules; petiolules about 5 mm. long, callous, glabrous; leaflets mostly 3 pairs, elliptic or ovate-elliptic, 13-40 cm. long and 7-14 cm. wide, rather abruptly narrowed apically into an acumen 1-2 cm. long, similarly contracted basally into a short subcuneatc base, completely glabrous, the veins somewhat impressed above and elevated below, chartaceous; (204) 19 5 0] FLORA OF PANAMA (Leguminosac) 219 Fig. 8 3. Inga gtganti folia Stipules not present. Inflorescence 1 to few, of very sKort, axillary, pedunculate spikes (sometimes borne on older, defoliate branches?), the peduncular portion about 5 mm. long and lightly puberulent, the floriferous portion about 5 mm. long also; bracts triangular-lanceolate, minute. Flowers small, white, scarcely pedicellate; calyx tubular-cupular, about 2.0-2.5 mm. long, subglabrous without, not striate, sometimes cleft to one side but otherwise the teeth broader than tall and the rim (205) [Vol. 37 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN appearing more or less undulate; corolla funnclform-tubular, 8-9 mm. long, sub- glabrous, the lobes 1-2 mm. long; stamens up to 2 cm. long, the staminal tube briefly exsertcd; ovary turgid-quadrangular, glabrous, multiovulate. Legume not known in type specimen, probably linear-oblong, flat, glabrous, the margins little thickened. Panama; South America? EOCAS ULi TORO: Fish Crcck Mts., H. von WeJcI 2J4g (Herb, Missouri Bot. Gard., type). The species would fall into the section Bourgonia of the genus as interpreted by Pittier following Bentham. It is quite distinctive from most species in the extremely large leaflet size. The type specimen appears identical in many respects with the Amazonian (Rio Solimoes) 7th Krukoflf Expedition specimens, Krukoff S2I§ and S22S. The Krukoflf material in flower (S2IJ) and fruit (822S) was found undetermined in the her- barium; the flowers were of slightly smaller dimensions, pronouncedly more pedi- cellate, and the calyx slightly more elongate than in the Panamanian material. Nevertheless, similarity in leaflet, inflorescence and floral structure would seem to indicate that the Krukoff material has close affinities with the Panamanian speci- men. In any event the Amazonian material supports the belief that, in spite of the separation of the inflorescence from the leaves in the H. von Wedel specimen, flower and branch are both from the same plant; and that similarity in ovary struc- # f 25- Inga PUNCTATA WiUd. Sp. Pi. 4:1016. 1806. Mimosa sericea Poir. in Lam. Encycl. Meth. Suppl. 1:42. 1810. In^a Icptoloba Schlecht. In LInnaea 12:560. 1838. Inga puucfafa var. panamcnsh Bcnth. in Trans. Linn. Soc. 30:613. 1875. FeuiHrca punctata Ktzc. Rev. Gen. Pi. 1:188. 1891. Inga punctata cbagronh Pittier, in Jour. Dept. Agr. Porto Rica 13:135. 1929. Small or moderate tree to 15 m., the branchlets strigose-puberulent and mark- edly lentlcellate. Leaves moderately large, pinnate, 4- to 6-foliolatc; petioles 2-4 cm. long, terete or nearly so, puberulcnt like the branchlets, swollen-callous basally; rachis similar to and (the intcrnodcs) about as long as or slightly longer than the petiole, wingless, bearing disc-like or subconic glands between insertions of the pctiolules (or these glands in part or entirely suppressed in some specimens) ; leaf- lets 2 or 3 pairs, elliptic to ovate, acuminate, those of the apical pair larger, up to 20 cm. long and 9 cm. wide, those of the lowermost pair usually about half as long and broad, strlgose- puberulcnt on both surfaces; stipules linear-lanceolate, 3-4 mm. long, caducous. Inflorescence an axillary, subtermlnal, paniculate or corymbi- form branch, the secondary spicate peduncles umbellate from tlie nodes, or some- times the spikes fasciculate or simple-umbellate directly from the axil; spikes short, the florlfcrous portion scarcely more than 1 cm. long, the many flowers densely f20G) 1950] FLORA OF PANAMA (LegMmiftosae) 221 congested. Flowers sessile, whitish; calyx subturbinate, 3-5 mm. long, the 5 teeth conspicuous but only about 0.5 mm. long, puberulent without; corolla tubular, somewhat expanded above the calyx, 7-9 mm. long, the lobes about 1 mm. long, pubescent without; stamens many, the filaments almost 2 cm. long in age, united into a tube for about Yz their length, glabrous, the anthers minute, bilocular, only a fraction of a mm. long; ovary and style glabrous. Legume linear-oblong, about 13 cm, long and 2.5 cm. wide, slightly arcuate, lightly pubescent, flattened, some- what margined, apparently indehiscent. Mexico and Central America to the Amazon Valley and Peru; Tobago. von Wedel lo6g, 1330, 2357, 2387, CANAL zone: Barro Colorado Island, Zetek 3837, 3847, 38671 "Chagres," tendler 8g; R. Chagres, Steyermark Gf Allen 16778; Gatun, Goldman 186^; Hayes jj, CHiRiguf: Boquete, Davidson 814. cocle: El Valle de Anton, Allen 3700, colon: R. Culebra, Pittier 4161, darien: El Real, Allen g66; Pinogana, Allen 4281. PANAMA: Chepo, Hunter ^ Allen pj. veraguas: near Sona, Woodson, Seibert ^ Allen 486, Inga leptoloba has been retained as a species distinct from /. pnnctata by most authors (Bentham, Britton & Rose, etc.), and Standley (Flora Guatemala) lists it as doubtfully distinct. Bentham's distinction between 7, leptoloba and 7. punctata is not clear, while Britton & Rose and Standley separate the two species on the basis of bifoliolate leaves in the latter vs. trifoliolate leaves in the former. However, many specimens occur with both bi- and tri-foliolate leaves on the same plant, and no supporting or correlating characters are apparent to strengthen this demarcation. Both bl- and tri-foliolate specimens exhibit the same range of variability In rachial gland structure, flower size, and type of inflorescence, and none of the vegetative distinctions mentioned by Pittier (Contr. U. S. Nat. Herb. 18:187) exhibit con- stancy when large numbers of specimens are examined. Thus it seems most prac- tical to unite these two names as a single species, or at least reduce 7. leptoloba to varietal status under 7, punctata, 26. Inga laurina (Sw.) Willd. Sp. Pi. 4:1018. 1806. Mhnosa fagtfolia Jncq. Stirp. Amer. Hist. 1:264, /. 164. 1763, non L., fide Benth. Mimosa laurina Sw. Prodr. Vcg. Ind, Occ. 85. 1788. Feuilleea laurina Ktze. Rev. Gen. PI. 1:184. 1891. Moderate-sized tree, the branchlets glabrous and markedly lenticellate. Leaves moderate-sized, usually 4- for sometimes 6-, or reported occasionally 2-) follolate; petiole 1-2 cm. long, terete except angled laterally above, glabrous; rachis 2-5 cm. long, glabrous, wingless but ridged and somewhat marginate laterally above, bear- ing a smallj sessile, disc-like or subcupuliform gland between insertion of the petiolules; leaflets usually 2 pairs, sometimes 3 or 1? pair, mostly oblong or elliptic, 4-12 cm. long and 1.5-5.5 cm. wide, usually acute and bluntly acuminate apically, cuneate and slightly inequilateral basally, glabrous above and below; stipules linear- oblong, up to 5 or 6 mm. long, pubescent, subpersistent or caducous. Inflorescence of 1 or 2 elongate spikes from each of a number of foliate or defoliate axils; spikes (207) [Vol. 37 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN subglabrous, about 10 cm. long, tbc peduncular portion only about 2 cm. long, tne many flowers scattered over the remaining floriferous portion. Flowers whitish, glabrous to puberulcnt; calyx cupuliform-tubular, 1-2 mm. long, the teeth minute but distinct; corolla usually 4-5 mm. long, narrowed basally, expanded apically, subglabrous; stamens 12-15 mm. long, fused basally into a tube greatly and notice- ably exceeding the corolla (Panama?) or only slightly exceeding the corolla. Legume reported flat and strongly compressed, 7—15 cm. long and 2—3.5 cm. wide, subglabrous, the margins thickened. Mexico to Panama; West Indies. CANAL zone: Barro Colorado Island, Bangham S47^ cuiriqui: Boca Chica, Vittier 511Q. cocle: Bismarck, Williams 3j2. herrera: Ocu, Allen 4037, Panama: Capira, Alien 1688; R. Pacora, Allen Si/, As with most species of the genus, I. lanrhm exhibits variability, noticeable here especially In length of the staminal tube relative to the corolla. In fact, certain specimens arc almost identical with respect to the elongate staminal tube to South American material generally accepted as 7. fagifolia Willd. Bentham (Trans. Linn. Soc, 3 0:608) considered the elongate staminal tube, along with a more pronouncedly bordered rachis and a fruit difference, as distinguishing 7. fa^^ifolia from 7. hnirifia, Panamanian (and Mexican) material could sometimes be con- sidered either of these two species, leading us to believe that combining these names (and probably that of 7. cylindrica (Veil.) Mart, also) might eventually prove in order. In Panama, 7. hiurina (regarding 7. fagifolia as a separate species not occur- ring there) is fairly readily delimited from other species except 7. marghiata by distinctive leaflet appearance and the elongate, relatively lax spikes. 7. coruscans H. & B. ex Willd., of northern South America, resembles 7. laurina except for the leaflet which is like that of 7. viarginata. 27. Inga Caldasiana Britt. & Killlp, in Ann. N. Y. Acad. Sci. 35:115. 1936. Forest tree to 20 m., the branchlcts glabrous or subglabrous, prominently lentlcellate, the Icntlcels and bark reddish-brown. Leaves large, normally 6- foliolatc, the basal pair of leaflets sometimes caducous, occasionally only one mem- ber of the terminal pair present; petiole short, at most 1.5 cm. long, the lower portion swollen and callous, glabrous or sparsely puberulcnt above; rachis usually 6—8 cm. long, unwinged but frequently slightly marglnate below the apical leaflets and somewhat angled above, subglabrous, bearing between insertion of the petiolulcs a sessile, patelliform gland about 1 mm. in diameter; leaflets elliptic or nearly so, about 15 cm. long and 6-7 cm. wide, the basal leaflets somewhat smaller, abruptly narrowed and short-acuminate apically, cuneate or subcuneate basally, glabrous except perhaps minutely puberulous on the costa; pctiolules 2-3 mm. long, callous, puberulous; stipules linear-oblong, usually 8—12 mm. long, puberulous. Inflores- cence of 1 to few axillary, short-pedunculate spikes, the peduncular portion less than 2 cm. long, the floriferous portion puberulcnt and as much as 10 cm. long but usually about 6 cm. long; bracts subtending the flowers minute, sublinear and ascending-expanded apically. Flowers moderately lax, sessile, white; calyx cupular, (20S) 1950] FLORA OF PANAMA (Legummosue) 223 less than 1 mm. long, pubcrulcnt; corolla tubular-funnclform, about 4 mm. long, subglabrous except canescent-pubcrulent at tips of the lobes; stamens 1 cm. long or slightly longer, the staminal tube exserted 2-3 mm. beyond the corolla; ovary oblong, compressed, glabrous; style truncate, exserted beyond the stamens. Legume not known. Colombia and eastern Panama. DARIEN: Yavisa, Alleti 45S5, The Panamanian specimen cited matches fairly well the type of /. Caldasiana, and as this species it constitutes a new record for Panama. Lawraucc ^84 from Boyaca, Colombia, is probably likewise 7. Caldasiana and not /• marginafa as found labeled in the herbarium. The inflorescence and flowers of /. Caldasiana might well be J. marginafa or 7. laurina, but it is hard to reconcile the leaf with either of these species. The unwinged, scarcely marginate leaf rachis and large leaflet size are unlike 7. marginafa, and the much larger leaflet and large stipule-like bracts are unlike 7. laurina. Also similar to 7. Caldasiana are the South American species 7. Klugii Standi, ex Macbride and 7. Bourgoni (Aubl.) DC. 28. Inga MARGINATA Willd. Sp. Pi. 4:1015. 1806. ?higa sapida HBK. Nov. Gen. & Sp. 6:286. 1824. Mimosa semialata Veil. Fl. Flum. Ic. 11:/. 5* 1827. Inga gtiayaqiiilensis G. Don, Gen. Hist. Dichl. Pi. 2:391. 1832, fide Benth. Inga odorata G. Don, loc. cit. 388. 1832, ex char, fide Benth. Inga cxceha Pocpp. & Endl. Nov. Gen. & Sp. 3:78. 1845, fide Benth. Inga piiberula Benth. in Hook. Lond. Jour. Bot. 4:589. 1845. Inga pycnostachya Benth. loc. cit. 1845. Feuillcca marginata Ktze. Rev. Gen. Pi. 1:188. 1891. Moderate-sized tree to 10 or 15 m., the branchlets glabrous, lenticcUate. Leaves moderate, 4-foliolate; petiole short, usually about 1 cm. long, marginate or alate apically, terete and callous basally; rachis 2—4 cm. long, usually prominently alatCj the wing broadest apically, bearing a sessile, patplliform gland between Insertion of the petiolules; leaflets 2 pairs, narrowly elliptic or elliptic-lanceolate, sharply acuminate apically, cuncate and somewhat inequilateral basally, the terminal pair up to 15 cm. long and 2—5 cm. wide, the basal pair about % as large, essentially glabrous above and below; stipules linear, about 5 mm. long, caducous. Inflores- cence of 1 to few spikes from the leaf axils, the spikes up to 10 cm. long and laxly floriferous for all except the lower several mm., glabrous or ligbtly puberulent. Flowers sessile, whitish; calyx campanulate or short-tubular, scarcely more than 1 mm. long, somewhat puberulent, the teeth prominent; corolla subfunnelform, 3—4 mm. long, gradually constricted basally, essentially glabrous; stamens about 12 mm. long, the staminal tube exserted 1—2 mm. beyond the corolla. Legume up to 12 cm. long and 1.5 cm. wide, flat, thickened about the seeds, lightly margined. Costa Rica, Panama, northern South America. BOCAS DEL lORO: Changuinola Valley, Dunlap 282; Chiriqui Lagoon, //. von Wedel 1378, 1549, j8oy, CANAL zone: Barro Colorado Island, Zetek Z4322; Gatun Valley, Vitficr 6512, cocLE: El Valle, Allen 22 31, darien: Sambii River, Pittier 55/9^ (209) [Vol. 37 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 84. In^a jnarginata SPECIMENS OF UNCERTAIN STATUS The Allen 1^282 (Canal Zone: R. Pequeni), in very young bud, lacking both fruit and corolla, had been filed in the Missouri Botanical Garden herbarium as Jnga panafncjjsis Seem. Such determination would seem decidedly incorrect. In Pittler's treatment of Inga (Jour. Dept. Agr. Porto Rico 13. 1929) the plant would likely fall into section Leptinga, with affinities near J. portohcllcmh Beurling, and may be that species. However, accurate comparisons cannot be made because of the incompleteness of the specimen, and its exact status must remain in doubt. Two sterile specimens labeled higa cdulis^ collected by James Zetek (No, jQoy from Barro Colorado Island, and No, S9^7 i^om near Chorrera, Panama), appear different, as judged by leaf alone, from any species listed from Panama. It is im- possible to give an exact determination to such sterile material, and unless dupli- (210) 19 5 0] FLORA OF PANAMA (Lcgiiminosae) 225 cates occur showing floral characters the specimens mentioned will have to remain indecisively located as to species. NAMES OF UNCERTAIN APPLICATION, POSSIBLY EMBRACING PANAMANIAN SPECIES Inga Lindeniana Benth. in Hook. Lond. Jour. Bot. 4:608. 1845. This species has been restricted by Pittier and Britton and Rose to the type, from Teapa, Mexico. Bentham, however, had considered 7. mucunu a synonym of this name and the species to be represented in Panama by Fendler and Duchas- saing specimens. We have not had opportunity to examine either the type or these specimens. Inga vera Willd. Sp. PL 4:1010. 1806. Inga Inga Britton, FL Burmuda, 170. 1918. This species has apparently been thoroughly confused with 7. spuria and related species. Bentham regarded 7. panciflora as synonymous with 7. vera, with a dis- tribution of the species in the West Indies, Central America and northern South America. Pittier later considered 7. vera confined to the West Indies, and reestab- lished 7. panciflora as a continental species. Recent floras of Colombia, Costa Rica, and Guatemala fail to list 7. vera: hence we include 7. panciflora but not 7. vera as occurring in Panama (see discussion of 7. panciflora) , 2. PITHECOLOBIUM^ Marrius PiTHEcoLOBiuM Mart. Hort. Reg. Monac. 188. 1829, as Pithcccllobium nom. conserv. /. ?. 1756. Spiroloba Raf. Sylva Tellur. 119. 1838. Cathormion Hassk. Retzia, 231. 1855. Fe7(iUeca Ktze. Rev. Gen. 1:182. 1891, in part. Siderocarpus Small, in Bull. N. Y. Bot. Gard. 2:91. 1901, homonym. Havardia Small, loc. cit. 1901. Samanea Merrill, in Jour. Wash. Acad. 6:46. 1916. Abarcma Pittier, in Trab. Mus. Com. Venezuela 2:86. 1927, Jutmnba Britt. & Rose, in N. Am. Fl. 23:24. 1928. Pimjnba Britt. & Rose, loc. cit. 28. 1928. Cojoba Britt. & Rose, loc. cit. 29. 1928. Ebenopsis Britt. & Rose, loc. cit. 33. 1928. Paiutcria Britt. & Rose, loc. cit. 1928. Chlorolcucon Britt. & Rose, loc. cit. 3 6. 1928. PseuJosamanea Harms, In Bot. Gart. Berlin Notizbl. 11:54. 1930. Arthrosamanca Britt. & Rose, In Ann. N. Y. Acad. Sci. 35:128. 1936. Macrosamauca Britt. & Rose, loc. cit. 131. 1936. ^The genus is written VithcccUobinm, PithccoUobium or Pithecolohium, The spelling Pitbcco- lohiumy although lacking prioricy, has been followed by most authors, including Bentham, Englcr & Gilg, Index Kew., Srandley, Ducke, Macbride and many others. Pittier and a few others have followed Martius' later alteration of the original spelling, PiihecoUobhim, Merrill and Britton Zc Rose have used the original spelling Pitbecellobhint, and the genus has been so listed in the con- scived name list of the International Rules of Botanical Nomenclature. On the basis of general acceptance and simplicity we follow here the spelling Pithccolobrum, (211) 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. Zl Trees or shrubs, unarmed or, less frequently, armed with stipuLir spines or thorns, the branchlets usually puberulent or tomentulose and Icnticellate. Leaves small or large, except in very few species (''Cojoba'') bipinnatc, the pinnae 1 to several pairs opposite on the rachls; leaflets 1 to many pairs opposite on the pinnular rachls (except frequently basal pair alternate or one leaflet of pair aborted) ; petiole unwinged, usually short, rarely bearing a gland on the upper side; rachis similar to the petiole, generally bearing a gland at insertion of each pair of pinnae; pinnular rachis usually similarly glandular, the glands commonly cupulatc; leaflets 2 to many per pinna, small or large, entire, usually Inequilateral, seldom heavily pubes- cent above; stipules minute or of prominent spines or thorns. Inflorescence of 1 to several axillary or supra-axillary, lateral or subterminal, pedunculate (rarely sessile) heads or spikes; floral bracts minute or conspicuous. Flow^ers pentamerous, sessile or rarely pedicellate, almost invariably whitish; calyx campanulate to tubular, pubescent or subglabrous, valvate, shallowly dentate; corolla tubular to funnelform, generally elongate, valvate, frequently tomentulose on the 5 lobes; stamens many, united below into a staminal tube included in or exceeding the corolla; ovary glabrous or pubescent, sessile or stipitate. Legume monlliform, compressed or flat, subclrcinate, arcuate or almost straight, normally dehiscent, the valves often twisting after dehiscence; seeds often imbedded in pulp. nd subtropics of New World; one section (Clypearta of Benth.) Far Eastern. Wo a temptation to authors favoring segregation. A few of the segregate genera, for example, Samanca Merrill (Jour. Wash. Acad. 6:46. 1916), have little in common ■with Pitbecolohhtm in general, and offer a case for establishment of separate (usually monotypic) genera. In the case of Samanca, Merrill argues his point well and convincingly, but fails to consider the results were each equally divergent species or grouping of species so accorded separate recognition. It is difficult enough to attempt to distinguish Pithccolobium from long-established genera of the MiMosoiDEAE, such as Calliaudra, Alhizzia, etc., without adding confusion In the form of numerous segregates such as Britton and Rose have proposed. It is often nearly impossible to find into which of such segregate genera a species falls, and the key to these genera is impossible to w^ork. Unfortunately, segregations of even universally recognized genera In the MiMOsoiDEAE have been made chiefly on the basis of legume characters. Many specimens, of course, are not in fruit, and such specimens often resemble one genus as much as another. Thus determination often becomes a hit-or-miss matchiniz of specimens. I am led to wonder whether legume characters would not have failed in many generic dehmitations had sufficient fruiting specimens been on hand to exhibit variability and degrees of intergradatlon. Certainly, in Panama, con- sideration of Fithecolob'ntyn in a broad sense brings together In one ''pigeon-hole" many diverse species which can then be comparatively easily keyed out on vegeta- tive and floral characters. (212) 1950] FLORA OF PANAMA (Legiiminosac) 227 a. Leaves once-pinnate; legume moniliform C^Cojoha^\ In part). b. Floral bracts minute, not cxserted nor readily visible in young head; costa and rachis glabrous or becoming glabrous in age; rachial glands mostly longitudinal - 1. P. membranaceum bb. Floral bracts 2-6 mm. long, cxserted beyond or visible among buds of young bead; costa and rachis pubescent except subglabrous in P. rufcscens var, vallcftse; rachial glands essentially isodiametric. c. Leaflets 1-5 pairs, only slightly asymmetrical, 2—15 cm. long (if small usually subglabrous on costa). d. Terminal pair of leaflets usually 10 cm. long or longer; costa of leaflet markedly pubescent below; floral bracts about 4 mm. long, exscrtcd, the young head appearing burr-like 2. P. rufescens dd. Terminal pair of leaflets seldom as much as 7 cm. long; costa of leaflet subglabrous below; floral bracts less than 3 mm. long, comparatively inconspicuous in young head 2a. P. rufescens var. VALLENSE cc. Leaflets 4—8 pairs, markedly oblique or inequilateral basally, 2—6 cm. long, tomentose on the costa 3. P. tubuliferum aa. Leaves bipinnate; legume various, seldom moniliform. b. Pinnae only 1 pair. c. Leaflets 3—12 per pinna; stems unarmed. d. Leaflets in pairs (even-pinnate} ; peduncle conspicuous, 2-24 cm. long; corolla elongate, 12—16 mm. long; legume moniliform. e. Leaflets 2—5 (usually 3 ) pairs per pinna; peduncle usually 2-3 but never more than 6 or 7 cm. long; corolla about 12 mm. long 4. P. Valerioi ec. Leaflets 4—6 pairs per pinna; peduncle 8—24 cm. long; corolla about 16 mm. long ....—. 5. P. caTenatum dd. Leaflets usually 3 or 5 (odd-pinnate) per pinna; heads or spikes scarcely pedunculate; corolla mostly 6-7 mm. long; legume flattened, of essentially uniform width ("Zygla'^) . e. Leaflets 3 or 5 per pinna, broadly elliptic (normally less than 3 times as long as broad); flowers in nearly sessile, very con- densed spikes or heads 6. P. laTIfolium ee. Leaflets 3 per pinna, narrowly elliptic (3 or more times as long as broad) ; flowers in short, lax spikes 7. P. longifolium cc. Pinnae bifoliolate; stems normally armed with stipular spines. d. Flowers in pronounced spikes, the floriferous portion mostly 3—1 2 cm. long; corolla 4—1 1 mm. long; legume arcuate or twisting. e. Corolla 7-11 mm. long, stamens 4-6 cm. long 8. P. hymeneaefolium ee. Corolla 4—6 mm. long, stamens no more than 2.5 cm. loni; 9. P. lanceolatum dd. Flowers capitate or short-spicate, the floriferous portion never more than 2 cm. long; corolla 2—4 mm. long; legume coiled or subcircinate. e. Flowers In short spikes; inflorescence an expanded, branched panicle, the peduncles mostly 1—3 cm. long 10. P. oblongum ee. Flowers capitate; inflorescence a contracted panicle, the heads subsessile or on short peduncles less than 2 cm. long 11. P. OULCE bb. Pinnae 2 to several pairs. c. Petiole bearing 1—3 sessile glands well below insertion of and not associated with the lowest pair of pinnae (or their scars) ; leaflets less than 10 mm. long. d. Commonly armed at some or most nodes with stipular spines; buds large, ovoid, with prominent bud scales; legume arcuate, the valves not coiling after dehiscence: (^'Chlorolcucon'') 12. P. mangense dd. Unarmed; buds slender, naked; legume circlnate, twisting In dehiscence, or unknown, e. Leaflets about 40 pairs j>er pinna, glabrous, narrower (up to 2 mm. wide) 13. P. pseudo-Tamarindus cc. Leaflets up to 16 pairs per pinna, pubescent below, broader (3-6 mm. wide) 14. P. Barbourianum cc. Petiole glandular at extreme apex only (if apparently glandular lower dowq, gland bordered by scars of aborted pinnae); leaflets 10 mm. long or longer. (213) [Vol. 37 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN I d. Leaflets 10-25 mm. long; flowers sessile; legume moniliform- subterete, the valves twisting after dehiscence ("Cojoba** ) , or unknown. c. Leaflets pubescent below, obtuse or rounded apically; flowers short (about 4 mm.? long), hirsutulous 14. P. Barbourianum ee. Leaflets essentially glabrous, more or less acute apically; flow- ers elongate (about 1 cm. long), glabrous or nearly so 15. P. COSTaricense dd. Leaflets 20-50 mm. long; flowers pedicellate; legume thickish but flat, uniform, the valves never twisting ("Samanca") , e. Cupular gland between insertions of basal pinnae large, as much as 1 cm. long; leaflets and flowers lightly pubescent..,.. 16. P. macradenium ee. Gland at insertion of basal pinnae small or nearly obsolete; leaflets and flowers densely pubescent 17. P. Saman 1. PiTHEcoLOBiUM membranaceum (Benth.) Schery, comb. nov. Ingj mcmhrauacea Bcntli. in Trans. Linn. Soc. 30:606. 1875. Cojoba glabra Britt. & Rose, in N. Am. Fl. 23:33. 1928. ritbecolobsum glabrum Standi, in Trop. Woods 34:40. 1933. Vithecolobium Darhhoniae Standi, in Field Mus. Publ. Bot. 22:82. 1940. Fig. 85, Pit hccolobium membranaceum Small or moderate tree, the branchlcts rufous-tomentellous when very young, soon glabrate, lenticellate. Leaves moderate, once-pinnate, the leaflets 2-4 pairs; petiole 1-3 cm. long, puberulent becoming glabrate, terete, bearing a sessile, con- cave, longitudinally extended gland shortly below insertion of basal pair of leaflets; rachis similar, 2-7 cm. long, bearing similar glands just below insertion of each pair of leaflets; leaflets elliptic, 3-11 (mostly 6-7) cm. long and usually about 3 cm. wide, briefly contracted apically into a bluntly acute tip, cuneate or sub- cuneatc basally, glabrous above and below, finely and noticeably reticulate below; (214) 1950] FLORA OF PANAMA (Legumhtosae) 229 stipules caducous or lacking in specimens examined. Inflorescence an axillary, pedunculate head; peduncle about 3 cm. long, puberulent, bearing a small bract (often caducous) on the upper half; head dense, subumbellate, the subtending bracts minute, rufous. Flowers sessile, greenish-white or pale greenish-yellow; calyx campanulate or campanulate-subtubular, 1-2 mm. long, glabrous except rufous-tomentose on the teeth; corolla tubular-funnelform, about 6 mm. long, glabrous except minutely puberulent on the teeth; stamens about 1 cm. long, the 5tamlnal tube included; ovary glabrous, sessile. Legume 2—10 cm. long, terete and moniliform, stipitate, red?, the valves twisting after dehiscence. Costa Rica and western Panama, cHTRiQUi: Boquete, Seemann Iig2; Callejon beco, Woodson t5 bcbery 4go; Davtdson 7/0; Casita Alta, Woodson, Allen ^ Seibert 864; Camiseca, M. £. (Davidson) Terry I3S2; R. Chiriqui Viejo, G. White 61; P. White 335; Volcan de Chiriqui, Davidson QOO; Quebrado Velo, Woodson & Schery 2/2. The species somewhat resembles P. mfescens, and grades into it, but as a rule has a more delicate, membranous leaflet and lacks the elongate subtending floral bracts so conspicuous in the (immature) head of P. ntfescens. It belongs to the ^^Cojoha^' section of Vitbccolobiuvt^ and is distinguished from all except 2 species of that genus (in Panama) by the once-pinnate leaves and the peculiar moniliform legume. The type of this species (^nga membranacca^ Seemann II92) has not been examined, but judging from a photograph of it in the U. S. Nat. Herbarium there seems little doubt but that it belongs with the other once-pinnate Pithecolobiums which Pittier transferred from IngUy and that the newer Pithecolohium species listed above can be referred to it. The species has rested as Inga since original publication without a second collection having been recognized. 2, PiTHECOLOBiUM RUFESCENS (Benth.) Pittler, in Contr. U, S. Nat. Herb, 18:181. 1916. Inga rufescens Benth. in Hook. Lend. Jour. Bot. 4:58 5. 1845. Jnga globiilifera Benth. loc. cit. 184 5, fide Pittier. Inga Billbergiana Benth, loc. cit. 1845, fide Pittier. Teuilleea rufescens Ktze. Rev, Gen. Pi. 1:189. 1891. Cojoba rnfescem Brltt. & Rose, in N, Am. Fl. 23:32. 1928. Tree, up to several meters tall, the branchlets ferruginous-tomentulose and usually prominently lenticellate. Leaves moderate or moderately large, once-pin- nate, the leaflets 1-5 (mostly 3-4) pairs; petiole short, normally 1 cm. or less long, pubescent; rachis up to 10 cm. long, pubescent, subsulcate above, bearing a sessile, concave gland at insertion of leaflet pairs; leaflets mostly oblong, 4-15 cm. long and 2-6 cm. wide, rounded to bluntly acute-acuminate aplcally, subcuneate to rounded basally, very slightly Inequilateral, usually glabrous above, lightly pubescent below except the costa rather heavily so, the lateral veins prominent and elevated below; stipules linear, early caducous. Inflorescence of solitary or geminate pedunculate heads from subterminal axils; peduncle 1—5 (usually about 2) cm. (215) [Vol. 37 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 8 6. Pithvcolobhim rnfcsccns long, rufous-pubescent, generally bearing a solitary bract above the middle; bead condensed; floral bracts linear, about 4 mm. long, clliate, giving the head prior to nnthcsls a burr-like appearance. Flowers sessile, white; calyx short-tubular, about 2 mm. long, rufous-pubescent apically; corolla tubular-funnelform, 6 or 7 mm. long, rufous-pubescent on the lobes; stamens about 15 mm. long, the staminal tube included; ovary sessile, puberulcnt apically. Legume linear-moniliform, up to 15 cm. long, reddish with black seeds, puberulent, the valves very twisted after dehi isce Panama and Colombia, CANAL zone: Ancon, Vitticr 2/4^; Barro Colorado Island, WooJuorfb ^ Vestal 6jy; Kcnoycr 36p; Miraflorcs, Allen 174Q, IJSQ; G. White Ij2; Pueblo Nucvo, P. White 30J; R. Tndio Hydrographic Station, Steyermark 174^9. cocle: Santa Clara, Muenscher 1624T, 16203, colon: Porto Bcllo, Vittier 24QO. Panama: Chepo, Dod^e^ Hunter, Steyermark d Alien 26626; Vittier 4493; San Jose Island, Erlanson 39, 368; Taboga Island, Standley 27943- (216) 195 0] FLORA OF PANAMA (Legumhtosac) 231 This species exhibits considerable variabiHty, and observation of material at hand supports belief that Pittier was entirely correct in reducing to synonymy Bentham*s 7. globiilifera and /. Billbergkna. Likewise, Pittier's transfer of this and related species from J^iga to Pithecolobhim seems justified for the reasons he gives, although from the standpoint of practical identification the species could be more easily located in Jnga^ by means of the once-pinnate leaf criterion established by Bentham. As now recognized by most authors these few species of Pithccojolmtm {'^Cojoba'' of Britt. & Rose) are the only species of the Mimosoideae aside from higa with once-pinnate leaves. Unfortunately rejection from Inga and inclusion in the "catch-all*' Pithccolobinin involves chiefly legume characters (moniliform fruit not found in Inga) not to be had in the majority of specimens. The legume resembles that of most species of PUbecoloblum little more than the legume in Inga, and on this basis there may be some justification for Britton and Rose's genus Cojoba. 2a. PiTHEcoLOBiuM RUFESCENS (Benth.) Pittier var, vallense Schcry, var. nov. Arbor ramulis inconspicue lenticellatis, foliolis specie minoribus subglabris; bracteis capitulcrum spcciei similibus scd minoribus. Tree of medium height, the branchlets rufous-tomentulose when very young, glabrate in age, inconspicuously lenticellate. Leaves small, once-pinnate, the leaflets 3-5 pairs; petioles very short, usually less than 5 mm. long, terete, pubcrulent; rachis up to 5 cm. long, like the petiole, bearing a sessile, concave gland just below insertion of the pairs of leaflets; leaflets oblong to narrowly elliptic, usually not longer than 6 cm., nearly glabrous, mostly obtuse apically and basally. Inflores- cence an axillary, pedunculate head; floral bracts similar to those of the species but somewhat smaller and perhaps intermediate to condition found in P. membra- nacc7(m. Flowers white, sessile; calyx campanulate, little over 1 mm. long, pubes- cent apically; corolla tubular-funnelform, about 6 mm. long, pubescent on the lobes. Legume moniliform, red, the seeds black. Apparently found only in vicinity of El Valle de Anton, Code, Panama. cocle: El Valle, Allen 7/0 (Herb. Missouri Bot. Gard., type), 1834 (cotype), 24C)y, Hunfcr ^ Allen ^§$. This entity has been described as a variety rather than as a species in that degrees of intermediacy {Hunter ^ Allen ^55 especially) occur towards P. rufes- ce^u. Its geographic distribution, as far as known, fits nicely the concept of a localized variety. 3. PiTHECOLOBiUM TUBULIFERUM Pittier, in Contr, U. S. Nat. Herb. 18:181. 1916. Inga fubulifera Benth. in Hook. Lend. Jour. Bot. 4:5 84. 1845. Cojoba fubuUfera Britt. & Rose, in N. Am. Fl. 23:32. 1928. Shrub or small tree, the young branchlets brownish tomentose, becoming sub- glabrous In age and inconspicuously lenticellate. Leaves moderately small, once- (217) [Vol. 37 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN pinnate, tlic leaflets 4-8 pairs; petiole usually less than 1 cm. long, terete, tomcntose like the young branchlets, often appearing to bear a gland because of abortion of lowermost pair of leaflets; rachis up to 10 cm. long, similar to the petiole, bearing sessile, concave glands at insertion of leaflet pairs; leaflets somewhat variable but mostly oblong, 2-6 cm. long and 1-2.5 cm. wide, bluntly acute to rounded or subtruncate-retuse (malformation? ) apically, obliquely rounded basally, sub- glabrous above, lightly pubescent below except heavily so on the costa; stipules linear, caducous. Inflorescence an axillary, pedunculate head; peduncle 3-(7?) cm. long, tomcntose, bearing a linear-lanceolate bract (this often caducous) above the middle; head condensed; floral bracts linear, about 2 mm. long, pubescent and ciliate. Flow^crs sessile, whitish; calyx campanulate-funnelform, about 1 mm, long, rufous-pubescent except at extreme base; corolla tubular or nearly so, about 7 mm. long, glabrous except on the lobes; ovary sessile, glabrous. Legume not known, / Panama and Colombia. cMiRTQUi: Boquctc, Davidson 62S, cocle: Penonomc, Wrlliaws 4Q. The description is chiefly from the Chiriqui specimen (in immature flower) . As far as can be determined from keys and rather incomplete descriptions, this speci- men is P. tuhulifcrnniy although no authentic specimens are available for com- parison. It is similar to P. rnfescens but differs in having more and generally smaller leaflets, and has the floral bracts less pronounced. 4. PiTHEcOLOBiuM Valertoi (Britt. & Rose) Standi, in Field Mus. Publ. Bot. 18:509. 1937. Cojoba Yalerioi Britt. & Rose, in N. Am. Fl. 23:32. 1928. Cojoba StanJlcyi Britt. & Rose, loc. cit. 1928. Vithecolobhim Statnlleyi Standi, loc. cit. 1937. Shrub or small tree, the branchlets rufous-puberulent when young becoming glabratc, lenticcllate. Leaves large, bipinnate, the pinnae 1 pair, the leaflets 2—5 (usually 3) pairs per pinna; petiole quite short, less than 1 cm. long, terete, puberulent, bearing an apicule about 4 mm. long terminally between insertion of the pinnae; pinnular rachis almost 10 cm. long, rufous-tomentulose, terete, bearing sessile, subcupular glands at insertion of leaflet pairs; leaflets ovate to elliptic, the terminal pair as much as 15 cm. long and 6 cm. wide, acute to short-acuminate apically, very obliquely blunt or rounded basally (the basal pair as small as 2.5 cm. long and 1 cm. wide, scarcely oblique basally, not strictly opposite), sub- glabrous above and below except on the costa, the veins prominent below; stipules linear, scarcely 2 mm. long. Inflorescence a pedunculate head arising slightly above the axil; peduncle 2— (6?) cm. long, puberulent, bearing a squamiform bract above the middle; head condensed, about 4 cm. in diameter; floral bracts linear-lanceolate, 2—3 mm. long, rufous-pubescent and ciliate. Flowers sessile, white; calyx cylindric, 2-3 mm. long, glabrous except tomentulose on the teeth; corolla tubular, about 12 mm. long, glabrous except tomentulose on the lobes; stamens about 18 mm. (2n) 1,9.5 0] FLORA OF PANAMA (Legummosae) 233 Fig. 87. nthecolob'tiim catcnatum valves twisting after long, the stamlnal-tube Included; ovary sessile, subglabrous. Legume apparently monillform, about 10 cm. long, red, the seeds black, the dehiscence. Costa Rica and Panama. BOCAs DEL TORo: Chlrlqui Lagoon, H, von WeJel 2246, 22J4; Water Valley, H, t^n We J el 1723. (219) [Vol. 37 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN The Water Valley specimen is rather poor, and had originally been determined as Jnga. The species Is poorly known, but seemingly is quite variable. The cited specimens are probably better regarded as conspecific with P. Valcrioi and P. Standlcyi than as distinct and coordinate with these species, although certain dif- ferences exist between collections of P. Valcrioi, P. Standlcyi and the cited speci- mens. 5. PiTHECOLOBiUM CATENATUM Donn. Smith, in Bot. Gaz. 48:294. 1909. Cojoha catenata Britt. & Rose, in N. Am. Fl. 23:32. 1928. Tree, the branchlets ferruginous-tomcntulose when young, longitudinally striate or rugose when dry, prominently lenticcllate. Leaves large, twice-pinnate, the pinnae 1 pair, the leaflets 4-6 pairs per pinna; petiole 2-6 cm. long, terete, ferruginous-pubescent, subsulcate above, usually bearing a small, sessile gland apically; pinnular rachis 10-20 cm. long, similar to the petiole, bearing a small, cratcriform gland at insertion of the pairs of leaflets; leaflets elliptic or obovate, the terminal pair quite large, 10-20 cm. long and 5-7.5 cm. wide, the basal pair 3-8 cm. long and 1.5-4.5 cm. wide, acute apically, inequilaterally acute or obtuse basally, subglabrous above except on the costa, Hghtly pubescent below or sub- glabrous except on the veins; stipules caducous. Inflorescence of very long pedunc- ulate heads lateral from the axils of the older wood; peduncle 8-24 cm. long, usually with a squamiform bract above the middle, tomentulose; head condensed, about 4 cm, in diameter. Flowers sessile, white; calyx tubular, 2.5-4.5 mm, long, glabrous except puberulent on the teeth; corolla tubular, about 16 mm. long and 1.0-1.5 mm. wide, glabrous, longitudinally substriate, the lobes thick and brown- pubescent apically; stamens about 2 5 mm. long, the staminal tube included; ovary terete, lightly pubescent, 12- to 14-ovulate; style glabrous. Legume moniliform, terete, up to 18 cm. long, red, tomentose, short-stipitate, the valves twisting after dehiscence. Costa Rica and Panama. BOCAS DEL TORO: Bastimcntos, H. von Wcdcl 2gog; Careening Cay, H. von Wcdcl 2813; Isia Colon, H. von Wcdcl 576; Old Bank Island, H. von Wcdcl /p77, 2066; without locality, H. von Wcdcl IQQ. This species is especially marked by the unusually long peduncles. 6. PiTHECOLOBiUM LATH OLiUM (L.) Bcnth. in Hook. Lond. Jour. Eot. 3:214. 1844. Mimosa latifoUa L. Syst. cd. 10, 1310. 1759. Zygia arhorcsccfjs St. Hil. Expos. Fam. Nat. 2:246. 1805, fide Britt. & Rose. Inga latifolia Willd. Sp. Pi. 4:1020. 1806. Calliandra latifoUa Griscb. Fl. Brit. W. Ind. 225. 1860. Tcnillcca latifoUa Ktze. Rev. Gen. PL 1:188. 1891. Zygia latifoUa Fawc. & Rcndlc, Fl. Jam. 4^:150. 1920. Pithccolobium chagrcnsc Pittier, in Contr. U. S. Nat. Herb. 20:465. 1922, Shrub or small tree, the branchlets glabrous, inconspicuously lenticcllate. Leaves moderately large, bipinnate, the pinnae one pair, the leaflets 3-5 per pinna; (220) 1950] FLORA OF PANAMA (Lcgiiminosae) 235 petiole very sKort, mostly 2-3 mm. long, callous, rugose, bearing a patelliform or subpeltate gland apically; rachis of pinna up to 10 cm. long, sulcate above, puberu- lent or subglabrous, bearing a small gland apically at insertion of terminal leaflets; leaflets elliptic, 5—15 cm. long and 1.5—6 cm. wide, the terminal pair opposite on the pinna, the penultimate pair alternate or lacking, tbe basal leaflet solitary on the outer side (sometimes basal leaf caducous), acute and sbort-acuminate apically, cuneate or subcuneate basally, slightly asymmetrical, glabrous above and below, with usually 3 prominent, ascending, arcuate lateral veins; stipules ovate. Inflores- cence of fasciculate sessile heads (or very condensed spikes) lateral on the older branchlets, the axis scarcely 5 mm. long, pubescent; bracts ovate, about 1 mm. long. Flowers sessile, pinkish; calyx cupular, about 1 mm. long, pubescent apically on the minute teeth; corolla tubular, about 7 mm. long, only slightly expanded above, subglabrous except puberulent on the lobes, striate; stamens usually about 17 mm, long, the stamlnal tube slightly exserted; ovary subsessile, lightly pubes- cent; style exceeding the stamens, the stigma capitellate. Legume linear, about 12 cm. long and 2 cm, wide (immature?), flat, minutely puberulent, arcuate. West Indies; Honduras to Panama; Colombia to Brazil and Bolivia. BOCAS DEL TORO: Garay Creek, H. von WeJel 26jJ; Old Bank Island, H. van WeJel 2014; Water Valley, H, von WeJel 86S. canal tone: "Chagres," Fendler QO; France Field, Standley 30368; Gatiin, Hayes 439; Pittier 280S, As with P. longifol'mm, P, latifolinm appears to be a fairly constant species. It is quite similar to P. glomcratum and P. cauJifhrnm, two South American species doubtfully distinct from each other, P. latifoUitm is distinguished from P. longi- fol}u7n In having a much shorter inflorescence, wider and often more numerous (5 vs. 3) leaflets. + 7. PiTHECOLOBiuM LONGii OLiUM (H. & B.) Standi, in Field. Mus. Publ. Bot. 4:212. 1929. Inga lon^ifoUa Humb, & Bonpl. ex WiUd. Sp. PL 4:1010. 1806. Mimosa ligustrina Vahl, Eclog. 3:34, f, 2/. 1807, not Jacq. 1801, fide Benth Jnga Yahliamim DC. Prodr. 2:43 8. 1825. Inga falciformis DC. loc. cit. 1825, fide Benth, Pithccolobhifji ligmfrinum Benth. in Hook. Lond. Jour. Bot. 3:213. 1844. Pitbecolobhim glomeratuDi van spicafnm Seem. Bot, Voy, Herald, 116. 1852-53, fide Britt. & Rose. Pithecolobium Yahlianum Benth. In Trans. Linn. Soc. 30:594. 1875. Pemlleea lou^tfoUa Ktze. Rev. Gen. Pi. 1:185. 1891. 2y^ia longifolia Britt. & Rose, in N. Am. Fl. 23:40. 1928. Moderate-sized tree, the branchlets glabrous or nearly so, lenticellate but not prominently so. Leaves moderate, bipinnate, the pinnae 1 pair, the leaflets 3 to the pinna (terminal pair and outer member of reduced basal pair?) ; petiole short and condensed, 3-10 mm. long, rugose, callous basally, bearing between Insertion of the pinnae a sessile patelliform or subpeltate gland; rachis of pinna usually about 2 cm. long, sulcate and puberulent above, commonly but not always bearing a gland between insertion of terminal pair of leaflets; leaflets narrowly ovate-lanceo- (221) [Vol. 37 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 8 8. Viihccolobium lofjgifolium latCj lanceolate or elliptiCj 4-12 cm. long and 1.5—3.5 cm. wide, cuncatc or sub- cuncatc basally, usually bluntly acute or acuminate apically, glabrous on botb sides, inconspicuously reticulate; stipules lanceolate, about 2 mm. long. Inflores- cence lateral on older wood, of short, lax, fasciculate spikes; spikes about 2 cm. lon^, scarcely pedunculate, subglabrous; bracts lanceolate, about 1 mm. long, sub- persistent. Flowers small, sessile, whitish or pinkish, fragrant; calyx cupular, about 1 mm. long, glabrous except the minute teeth puberulent or ciliatc; corolla tubular- funnelform, 6—7 mm. long, glabrous; stamens many, about 16 mm, long, the staminal tube well exserted; ovary pubescent. Legume linear-arcuate, up to 30 cm. long and about 13 mm. wide, glabrous, flat and thin, short-stipitate, the seeds longitudinal. (222) 1950] FLORA OF PANAMA (Lcguminosae) 237 Honduras to Panama; northern South America. BOCAS DEL TORo: Water Valley, H. von WeJel 655; Cooper ^ Slater 54b, canal zone: R. Chagres, Steyermark & Allen 16/8/ ; R. Indio Hydrographic Station, Steyermark 17418; Tumba Vieja, Steyermark ^ Allen 16/4^. cocle: Bismarck, Williams 24J; El Valle, Allen loSy 2'j6^. colon: Santa Isabel, Pittier 462. darien: Boca de Cupc, Allen Sjl , PANAMA; R. Las Lajas, Allen 161J; R. La Macstra, Allen 2^, The species in Panama seems to exhibit stability unexpected in the genus. Similar to P. latifolinm and other species, the differences, although minor, appear surprisingly constant. Fig. 89. Tifhccoloblum hymcncacfolittm 8. PiTHECOLOBiuM HYMENEAEFOLIUM (H. & B.) Benth. in Hook. Lond. Jour. Bot. 3:198. 1844. Jnga hymenaeaefoUa Humb. & Bonpl. ex Willd, Sp. PL 4:1008. 1806. Mimosa bymeneacfolia Poir. in Lam. Encycl. Mcth. Suppl. 1:3 8. 1810. Pithecolobium panamense Walp. & Duchass. in Linnaea 23:746. 1850. Shrub or small tree to a few meters tall; branchlets subglabrous, minutely lenticellate, with a few longitudinal lines or thin ridges and usually paired nodal thorns. Leaves moderate, bipinnate, the pinna bifoliolate; petiole mostly 2—4 cm. long, canaliculate above, glabrous, bearing apically a sessile, cupular gland; axis of the pinna up to 2 cm. long, like the petiole; mature leaflets asymmetrically ovate (223) [Vol. Z7 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN port or ovatc-lanccolatc, 6-13 cm, long and 2-5 cm. wide, bluntly acute apically, ob- liquely rounded basally, glabrous above and below except often barbate basally below on the narrow side; stipules modified as small to prominent recurved thorns or spines to either side of the petiolar insertion. Inflorescence of elongate, axillary, subtcrmJnal and terminal pedunculate spikes; peduncle L5-6 cm. long, sub- on of the spike mostly 3—6 cm. long, the axis tomentulose and few-sulcate, the flowers condensed; bractlcts minute, ovate-subulate. Flowers large, sessile, whitish; calyx cupular, about 2 mm, long, tomentulose, the teeth short and somewhat irregular; corolla funnelform, usually about 1 cm, long, tomentulose without, valvate; stamens 5-6 cm, long, united basally into a promi- nently exsertcd stamlnal tube 2-4 cm. long; ovary sessile, pubescent; style (plus ovary) about 6 cm. long, the stigma capitcllate. Legume 7-10 cm. long and about 15 mm. wide, falcate, compressed, tomentose, rugose, few-seeded, the seeds black. Panama, Colombia, Venezuela. CANAL zone: "north end of island" (Miraflores?) , P. White 124; R- Chagres, Steyer- mark ^ Allen ^6jgO; Salamanca Hydrographic Station, Woodson, Allen S? Seibcrt Js6j, PANAMA: Taboga Island, Woodson, Allen ^ Seibert J 539; Chepo, Dodi^e, Hunter, Steyer- mark ^ Allen r666j. The corolla of the cited specimens is slightly shorter than dimensions given for the species by most authors. There is some uncertainty as to application of the Willdcnow name and its Interpretation by later authors. 9. PiTHECOLOBTUM LANCEOLATUM (H. & B.) Bcnth. in Hook. Lond, Jour. Bot. 5:105. 1846. Mimosa U^ustrina Jacq. Fraj^m. Bot. 29, />/. J2, /. 5. 1801. In^^a lauceolata Humb. & Bonpl. ex Willd. Sp. Pi. 4:1005. 1806. Mimosa lanceolata Polr. in Lam. Encycl. Meth. Suppl. 1:37. 1810. Vitbecolobinni macrostacbyum Benth. in Hook. Lond. Jour. Bot. 5:198. 1844, excl. syn, Vahl., fide Benth. Vitbecolobium ligustriniim Kitz. ex Benth. in Trans. Linn. Soc, 30:571. 1875, not B^ntli. 1844. Vitbecolobium macrosipbon Standi, in Contr. U. S. Nat. Herb. 20:19L 1919, fide Stand!. & Steycrm. Vifhccolobium Winzerlingii Britt. & Rose, in N. Am. Fl. 23:193. 1928, fide Standi. & Steyerm. This species, to the best of my knowledge, has not yet been collected in Panama, but is known from Mexico, northern Central America and Costa Rica, and northern South America. It thus can be expected in Panama also. It is a small, armed tree similar vegetatively to P. ohlotigiim except that the leaflets average larger. The floriferous portion of the spike is considerably more elongate (4-12 cm. long) than in P. obloiignm^ and the corolla is 5-6 mm. long. The ovary is subscssile, the legume similar to P. oblongiim although perhaps somewhat larger. As with P. Juice, the aril of the seeds is said to be edible and used for making a beverage suggestive of lemonade. P. lauccolatuvi can scarcely be distinguished from P. pachyfnis Pitticr except in fruit. P. pacbypus, a species of Salvador, (224) 195 0] FLORA OF PANAMA (Legjimtuosae) 239 Guatemala, and Mexico, has a significantly broader, stockier fruit than does P. lanceolatum, but insufficient fruiting material is on hand to appraise accurately the delimitation and distribution of these species. 10. PiTHECOLOETUM OBLONGUM Benth. in Hook. Lond. Joun Bot. 3:158. 1844. Feiiilleea oblonga Ktze. Rev. Gen. Pi. 1:188. 1891. ?ithecolobiiim paniculatiim Pittier, in Contr. U. S. Nat. Herb. 20:462. 1922. Pithecolobium pulcbellum Pittier, loc. cit. 1922. Vithecolobium microsfachyicm Standi, in Jour. Wash. Acad. 13:439. 1923. Pithecolobium Pittieri Britt. & Killip, in Ann. N. Y. Acad. Sci. 35:125. 1936. Fig. 90. PithecolobtU7n oblongiim Small or bushy tree to several meters; branchlets essentially glabrous, with a few longitudinal lines or thin ridges and many prominent lenticels. Leaves mod- erately small, bipinnate, each pinna bifoHolate; petioles mostly 1-3 cm. long, slen- der, subglabrous, somewhat flattened, canaHculate above, bearing a short-columnar, concave gland apically above; axis of the pinna mostly 1 cm. or less long, similar to the petiole except the apical gland sometimes obsolete; leaflets asymmetrically ovate, oblong or obovate, up to almost 6 cm. long and 3.5 cm. wide, obtuse to rounded or subtruncate apically, obliquely obtuse basally, glabrous above and below except barbate basally on the narrow side below; stipules modified as small or moderate thorns or spines to either side of the petiole^ a few mm. to 1 cm. long or longer. Inflorescence an axillary panicle, usually subterminal, consisting of a (225) [Vol. 37 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN number of pedunculate spikes; peduncles slender, usually 1—2 cm. long at maturity, lightly pubescent or subglabrous; floriferous portion of spike 5-20 mm. long, condensed apically and somewhat more lax bascward; bractlets ovate-lanceolate, scarcely 1 mm. long, persistent, pllosulous. Flowers small, sessile, whitish; calyx cupular, little over 1 mm, long, shallowly lobed, puberulent, 5 -angled; corolla funnclform, 3-4 mm, long, valvate, puberulent at least apically; stamens about 8-9 mm. long, united below into a short staminal tube; ovary stipitate, glabrous, the stipe almost 1 mm. long. Legume lincar-circinate, up to 10 cm. long and about 1 cm. wide, frequently coiled upon itself, dehiscent, compressed but turgid at and indented between the seeds, nearly glabrous, reddish; seeds ovoid, lustrous-black. Mexico to Panama. CAN AT. zone: Farfan Beach, Huutcr & Allen 446; without locality, Alien l6j2; Hayes JIJ, /6l; Seemann 40J, Panama: beach, Panama Vieja, Allen 82/; Bro. Paul jSS; beach, San Carlos, Allen IIJJ; Bella Vista, Killip I20ip; Punta Paitilla, Piper 5120. No authentic specimens of P. oblongum have been available for comparison with the cited Panamanian material. Bentham's description fits the cited specimens as far as it goes, but it is scarcely complete and comprehensive in a modern sense. Yet, it seems very likely that our specimens are P. oblongum^ a species described from a Panamanian type, especially In that they key to or close to P. oblonguw as interpreted by various authors. Specimens of P. microstacbyiim Standi, seem to differ from our material only within the allowable limits of specific variation. P. pulchcllum Pittier and P. Vittcri Britt. & Killip differ from the Panamanian material inconstantly and in a very minor fashion. There is also apparent inter- gradation to P. uuguis-catiy restricted (for convenience?) to the West Indies. The species is evidently a sea beach plant. 11. P1THECOLO131UM uuLCE (Roxb.) Benth. in Hook. Lond. Jour. Bot. 3:199. 1844. Mimosa dulcis Roxb. PL Coromand. 1:67. 1795. higa dulcis Willd. Sp. PL 4:1005. 1806. Inga fyuugctjs H. & B. ex Willd. loc. cit. 1004. 1806, fide Benth. Minufsa pun^ens Poir. in Lam. EncycL Meth. SuppL 1:36. 1810. Inga jarana DC. Prodr. 2:436. 1825, fide Benth. MimoSii intguis-cati Blanco, Fl. Filip. 731. 1837, non L., fide Benth. Acacia obliquifolia Mart. & Gal. in Bull. Acad. Brux. 10^': 3 17. 1843, fide Benth. Jn^a leucantha Presl, Bot. Bcm. 65. 1844, fide Benth. Fciiilleea dulcis Ktze. Rev. Gen. PL 1:184. 1891. nthccolobium littovulc Brltt. & Rose ex Record, In Trop. Woods 11:15. 1927, fide Standi. & Stcycrm. Small to medium tree, often flowering when small and shrub-llkc, the branch- lets puberulent or subglabrous becoming glabrous In age. Leaves small, bipinnatc, each pinna blfioliolate; petiole mostly 1-2 cm. long, subglabrous or lightly puberu- lent, margined and sulcate above especially apically, callous basally, bearing a con- cave gland at Insertion of the pinnae; axis of the pinna like the petiole, up to 1 cm. long, the gland frequently smaller than the petlolar gland; leaflets 2 per pinna, mostly inequllaterally oblons;, 2-6 cm. long and 1-3 cm. wide, obliquely obtuse or (226) 1950] FLORA OF PANAMA (Leguminosac) 241 rounded bnsally, blunt or rounded apically, glabrous except barbate basally below; stipules modified as scarcely noticeable to moderate spines or thorns on either side of the petiole. Inflorescence a terminal or subterminal panicle of pedunculate heads; peduncles about 1 cm. long or longer at anthesis, puberulent; head globose, about 15 mm. in diameter, several-flowered, compact; bracts scarcely 1 mm. long. Flowers small, whitish; calyx cupular, about 1 mm. long, canescent-tomcntulose; corolla funnelform, 2-3 mm. long, tomcntulose, the lobes about 1 mm. long; stamens many, about 8 mm. long, united below into a staminal tube for about half their length, glabrous; ovary pubescent, stipitate. Legume torulose or subtorulose, up to 12 cm. long and 8-15 mm. wide, coiled, glabrous or puberulent, dehiscent, the valves twisting after dehiscence. Mexico to northern South America, CANAL zone: Balboa, Standley J0842. This species is very similar to P. tinguh-cati^ a species "not known to occur in Panama," from which it differs in having pubescent rather than glabrous flowers. 12. PiTHEcoLOBiuM MANGENSE (Jacq.) Macbr. in Contr. Gray Herb. 59:3. 1919. Mi?fiosa mangcnsh Jacq. Enum. PI. Carib. 34. 1760. Mimosa parvifolla Sw. Fl. Ind. Occ. 984. 1800. Acada pariifoUa Willd. Sp. Pi, 4:1086. 1806. Mimosa antillarum Lam. ex Poir. in Lam. Encycl. Meth. Suppl. 1:80. 1810. Acacia micranfha Desv. ex Ham. Prodr. FL Ind. Occ. 60. 182 5, fide Benth. Inga martbac Spreng. ex DC. Prodr. 2:441. 1825, fide Benth, Pithecolobium parvifolium Benth. in Hook. Lond. Jour. Bot. 3:22 3. 1844. Feuillcea wafigensis Ktze. Rev. Gen. Pi. 1:186. 1891. Entcrolobiitm mangensc Fawc. & Rendle, Fl. Jam. 4^:151. 1920. Chlorolcucon maugcusc Britt. & Rose, in N. Am. Fl. 23:38. 1928. Catbormium vjangcnsis Dugand, in Apunt. Hist. Nat. Dept. Atl. 32. 1933, fide Britt. & Killip. Shrub or small tree, armed at most (older) nodes with paired spines, the branch- lets glabrate and moderately lenticellate, frequently flexuous in active growth, bearing large, ovoid, axillary buds with prominent bud-scales. Leaves small, twice- pinnate, the pinnae 4-10 pairs, the leaflets 8-3 pairs; petiole slender, usually 1-2 cm. long, more or less puberulous, flattened on the upper side and bearing a small clavatc gland, usually below the middle; primary rachis similar, sulcate above, usually bearing a sessile, concave gland between some or at least the terminal pair of pinnae; rachis of pinna mostly 2-4 cm. long, pubescent, more or less terete; leaflets small, linear, usually about 6 mm. long and little over 1 mm. wide, some- what falcate, asymmetrically acute apically, obliquely rounded or truncate basally, lightly pubescent or subglabrous, the veins few and prominent below; stipules at most nodes becoming modified as spines, those straight or slightly curved, as much as 15 mm. long. Inflorescence an axillary, pedunculate head; peduncle slender, about 2 cm. long, somewhat puberulent; bracts linear, minute, caducous. Flowers white, sessile; calyx cupuliform-campanulate, about 2 mm. long, glabrous except (227) 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 perhaps on the lobes; corolla tubular-funnelform, 4-8 mm. long, glabrous except the lobes pubcrulcnt-tippcd; stamens 12 or more mm. long, the staminal tube about equalling the corolla. Legume linear, 6-14 cm. long and 10-12 mm. wide, compressed or nearly flat but thick, fleshy-coriaceous, glabrous, retlculate-vcined, curved or nearly straight. Panama and northern South America; Jamaica. VERAGUAs: Santiago, Allen I0S2, This species Is poorly represented in the herbarium, Its variability and exact * 4 taxonomic position uncertain. 13. PiTHEcoLOiiiuM pseudo-Tam ARiNDUs (Brltt.) Standi, in Field Mus. Publ. Bot. 4:212. 1929. Jupiitiba pscuJo-TamariuJus Britt., In N. Am. Fl. 23:193. 1928. Unarmed, buttressed tree up to 18 m. tall, the branchlets angulate and fer- ruginous-tomcntulose. Leaves twice-pinnate, the pinnae mostly 12-16 pairs, each pinna with about 30-40 pairs of leaflets; petiole 1-4 cm. long, terete, bearing 1-3 sessile glands towards or above the middle; rachis up to 20 cm. long, tomentulosc, bearing sessile glands between all pairs of pinnae; pinnae mostly opposite, 6-9 cm. long, the rachis tomcntulose, gland-bearing between the upper leaflets; leaflets narrowly oblong, 5-7 mm. long and almost 2 mm. wide, apically obtuse or rounded, basally sessile and obliquely rounded, glabrous, lighter below, the costa central, the secondary nerves only moderately prominent. Inflorescence subumbellate, axillary, the peduncle 2-12 cm. long, ebractcate, the pedicels 4-6 mm. long. Flowers (type specimen in fruit) white, the center (terminal) one of the umbel somewhat larger (as in ''Samaned") ; calyx campanulate, about 2 mm. long, pubcrulcnt, the teeth 5, triangular, about 0.5 mm. long; corolla funnelform, about 7 mm. long, lightly pubescent without, the lobes 5, nearly as long as the tube; stamens many, about 2 5 mm. long, united only for the basal 3 mm., glabrous; anthers minute; ovary cyllndrlc, about 3 mm. long, tomentulose, subtruncate apically, several- (about 14-) ovulate in 2 rows; style glabrous. Legume circlnate, 10-12 mm. wide, lightly constricted and falsely septate between the 3-5 seeds, the valves thin, reddish- brown, sparsely strigillose or almost glabrous. BOCAS DEL TORo: Bocas Island, Cooper 461 (fruit); Chiriqui Lagoon, H. von Wedd IIJ2 (flower). The species is similar to some of the West Indian ''Jupunbas" and "Abarcmas." It had been known only from the type (Cooper 461 ; in fruitj, but the von Wedel specimen (flowers only) appears to be the same species although the peduncle of the latter is markedly more elongate than in the type. The above description is pieced from both specimens. 14. PiTHEcoLOBiUM Barbourtanum Standi, in Contr. Arn. Arb. 5:74, pL IT. 1933. Unarmed tree, the branchlets fulvous hirsute-tomentose, slender. Leaves (228) 1950] FLORA OF PANAMA (Lcgummosae) 243 moderately large, twice-pinnate, the pinnae 8-11 pairs, the leaflets mostly 9-16 pairs per pinna; petiole 1-2 cm, long, somewhat thick, densely tawny hirsute, bearing a sessile, irregular, shallowly crateriform gland near the apex; primary rachis 5-12 cm. long, subterete, pubescent Hke the petiole, bearing similar solitary glands below insertion of each pair of pinnae; pinnular rachis 3-6 cm. long, pilose, glandular at most upper nodes; leaflets oblong or oblong-trapezoidal, 5-10 mm. long and 3-6 mm. wide, apically obtuse or rounded, basally oblique, glabrous (except sometimes on the depressed costa) above, lighter and pilose below, especially on the costa, the secondary veins prominent, the margins subrevolute and adpressed- ciliate. Inflorescence of axillary, pedunculate heads; peduncles usually geminate, slender, 4-6 cm. long, pilose; head about 1 cm. In diameter, multiflorate, very condensed; floral bracts obovatc-spatulate, about 2 mm. long. Flowers sessile, apparently inconspicuously dimorphic (terminal flower of head usually grosser, its ovary evidently not functional, the stamina! column enlarged) ; calyx funnelform, about 3 mm. long, golden-brown hirsutulous, briefly and obtusely 5- to 6-dentate; corolla apparently funnelform, about 4 mm. long (mature bud), densely golden- hirsute without; stamens about 13, the filaments briefly united below, glabrous, the anthers small, eglandular; ovary pilose, subsessile. Legume not known. CANAL zone: Barro Colorado Island, Shattuck ^J/'. The species Is known only from the type collection. Without fruit or mature flower its affinities are difficult to establish. It would seem, however, to fall near the well-represented P. Saman. 15. PiTHECOLOBiuM cosTARicENSE (Britt. & Rosc) Standi, in Trop. Woods 34:40. 1933. Cojoba costaricensh Britt. & Rose, In N. Am. FL 23:31. 1928. Moderate-sized, spreading tree, the branchlets fcrruginous-tomentosc becoming more or less glabrate in age, lentlcellate. Leaves moderately large, twice-pinnate, the pinnae 4-9 pairs, the leaflets mostly 10-14 pairs per pinna; petioles mostly 2-4 cm. long, terete, short-tomcntose, eglandular or inconspicuously glandular, scarcely callous basally; primary rachis (petiolar extension) like the petiole, bearing small, erect, truncate or concave glands between insertion of pairs of pinnae; pinnular rachis similar, but the glands frequently obsolete between the leaflets; leaflets ovate- lanceolate, 10-25 mm. long and 5-10 mm. wide, bluntly acute apically, obliquely rounded or subtruncate basally, subglabrous except puberulent on the costa, dull and inconspicuously reticulate above, lighter and more obviously reticulate below. Inflorescence of solitary pedunculate heads from older axils, the peduncle 3-8 cm. long, tom.cntose, eglandular; head 3-4 cm. In diameter, the flowers crowded; bracts minute or caducous. Flowers white, sessile; calyx cupular-tubular, about 2 mm. long, subglabrous except rufous-tomentulose on the teeth; corolla tubular, approxi- mately 1 cm. long, glabrous, the lobes scarcely expanded; stamens about 16 mm. (229) 244 I Vol. 37 MISSOURI BOTANICAL GARDEN Fig. 91. Pithecolobium costariccnsc long, glabrous, the stamina! tube included; ovary subglabrous, very sKort-stipitate. Legume monillform-subterete, reddish, the valves twisting after dehiscence. Costa Rica and Panama. CHiRTQui: R. Chiriqui Viejo, P, White 23. The P. White 22 specimen is sterile, but seems on vegetative characters to be P. cosfiiriccnsCy a species to be expected in Panama. P. chazufcnc Standi., of Peru, and several Central American species are quite similar to P. cosfaricense. 16. Pithecolobium macradenium Pittier, in Contr. U. S. Nat. Herb. 20:465. 1922. A moderate-sized tree, the branchlets more or less pubescent. Leaves moder- (230) 1950] FLORA OF PANAMA (LegJiminosac) 245 ately large, twice-pinnate, the pinnae usually 3 pair, each bearing 5-8 pairs of leaflets; petioles 3.5-7 cm. long, minutely pubescent; primary rachis (petiolar ex- tension) similar, bearing between insertion of the basal pair of pinnae a large, subcupular gland as much as 1 cm, long (possibly a disease condition), bearing between upper pairs of leaflets smaller glands or these more or less obsolete; rachis of pinna similar, bearing small glands between insertion of the pairs of leaflets; leaflets ovate or rhombic-ovate, 2-5 cm. long and 1-2.2 cm. broad, sessile, rounded apically, inequilateral basally, glabrous above except pubescent on the costa, paler and more or less pilosulous beneath; stipules linear, caducous. Inflorescence of axillary, pedunculate, condensed umbels or very short spikes; peduncles 5-6 cm. long, ferruginous-pubescent; flowers short-pedicellate, the pedicels 2.5 mm. long and sparsely puberulous. Flowers white, small; calyx campanulate, about 3.5 mm. long, puberulous, the teeth short, subacute; corolla funnelform, 6-6.5 mm. long, the tube glabrous, the lobes lanceolate, acute, and minutely brownish-pubescent; staminal tube included; pistil about 15 mm. long; ovary short-stipitate, glabrous. Legume about 10 cm. long and 2 cm. broad, thick, strongly arcuate, compressed, transversely sulcate between the seeds, short-stipitate. Panama. CANAL zone: Monte Lirio, Cbrhfopherson ig6. The species Is known only from the type (U. S. Nat. Herb.). The above description Is taken primarily from Pittier's original description. The most re- markable character of the species seems to be the very large gland between the basal pair of pinnae. I am familiar with no other species of the Mimosoideae bearing a gland of such proportions, and if this is a constant character not merely confined to an aberrant specimen, it may serve for ready identification of the species. 17. PiTHECOLOBiuM Saman (Jacq.) Benth, in Hook, Lond. Jour, Bot. 3:216. 1844. Mimosa Saman Jacq. Fragm. Bot. 15, pL p. 1800. hi^i^a clnerea H. & B. ex Willd. Sp. Pi. 4:1024. 1806, fide Benth. Mimosa pnbifera Poir. in Lam. Encycl. Meth. Suppl. 1:47. 1810. higa salutaris HBK. Nov. Gen. & Sp. 6:304. 1824, fide Benth. Calliandra tubulosa Benth. In Hook. Lond. Jour. Bot. 3:101. 1844, fide Benth. Vitbecolobium cincreiim Benth. loc. cit. 216. 1844. ? Acacia propinqua A. Rich. Ess. Fl. Cub. 466. 1845, fide Benth. Calliandra Samayi Griseb. Fl. Brit. W. Ind. 225, 1861, Albizzia Sawan F. MuelL Scl. Extra-Trop. Pt. 27. 1891, fide Rock, Legum. PL Hawaii. Feiiillcea Saman Ktze. Rev. Gen. PL 1:189. 1891, Enfcrolobiu?n Sarnan Prain ex King, In Jour. Asiat. Soc. Bengal 66^:252. 1897. Samanea Saman Merrill, in Jour. Wash. Acad. 6:46. 1916. A large, spreading, unarmed tree, the branchlets aureo-pubescent becoming glabrate, sparingly lenticellate. Leaves large, twice-compound, the pinnae 2-6 pairs, each pinna with 2-8 pairs of leaflets (terminal pinnae usually with the more (231) 246 [Vol. 37 MISSOURI BOTANICAL GARDEN Fig. 92. Pithi'colobium banian numerous leaflets) ; petioles mostly 4-6 cm. long, canescent- or yellowish-tomen- tose, somewhat sulcate above; primary rachis (petiolar extension) 4-6 cm. long, similar to the petiole, bearing at or just below insertions of each pair of pinnae a small, sessile, subcupuliform, laterally extended gland; axis of the pinna like the primary rachis except the glands between (slightly below) insertions of the leaflet pairs smaller; leaflets asymmetrically ovate, oblong or obovate, up to about 4 cm. long and 2 cm. wide, obtuse or rounded apically, obliquely obtuse or rounded basally, pubcrulent or subglabrous and lustrous above, yellow-tomentulose below; (232) 1950] FLORA OF PANAMA (Leguminosac) 247 stipules caducous. Inflorescence of 1 to few pedunculate umbels axillary and sub- terminal on tlie branchlets; peduncles at anthesis 4-8 cm. long, tomentose, sulcate; umbels congested, the pedicels mostly 1-3 mm. long (except central flower sessile) ; bracts (except central flower) lanceolate, about 3 mm. long. Flowers whitish or pinkish, dimorphic; central (sessile) flower of umbel larger, persistent, its calyx campanulate, with 10 teeth, about 5 mm. long and 3-4 mm. wide, its staminal column gross and thick but the many stamens relatively short, its ovary ovulate but comparatively underdeveloped, this flower surrounded by 5 oblanccolate bracts about 5 mm. long; remaining (pedicellate) flowers first to fall, their calyx funnel- form, about 5 mm. long and only 1-2 mm. wide, 5 -toothed, aureo-tomentose; their corolla funnelform, about 1 cm. long, valvate, aureo-pubescent apically only; their stamens about 3.5 cm. long, the stamlnal-tube included; their ovaries sessile, subglabrous except pubescent at base of style, the style becoming more or less glabrous apically and about equalling the stamens. Legume linear or linear-oblong, reported as much as 20 cm. long and 2 cm. wide, straight or slightly curved, r glabrate, the valves thick, the margins prominent, indented and septate between the seeds, the seeds transverse. Mexico, Central America, South America: naturalized in "West Indies and Old World tropics. BOCAS DEL TORO: Almirante, Cooper 442, canal zone: several localities, sterile speci- mens, herrera: Divisa, Allen 4433- veraguas: Santa Fe, Allen 4437. A well-known and frequently abundant species in Central America, the leaves and pods of which are often relished by stock. The wood is of local importance and the pulp of the pods is said to be edible. The most common English name for this species is ''rain tree." SPECIES OF PITHECOLOBIUM OF DOUBTFUL OCCURRENCE IN PANAMA PiTHECOLOEIUM COGNATUM Benth. Mamei railway station (S. Hayes 166; Seemann s.7t.), fide Hemsl. Biol. Centr.-Am. Bot. 1:359. 1888. The cited specimens, which I have not seen, are possibly P. latifolhim, P. cognatum (as 2ygia cognata), fide Britt. & Rose (N. Am. Fl. 23:39. 1928), is restricted to Mexico. PiTHECOLOBIUM FRAGRANS BcUth. Hacienda de San Juan, "Veraguas" (Seemann Iigo), fide Benth. In Trans. Linn. See. 30:592, 1875; and Hemsl. Biol. Centr.-Am. Bot. 1:360. 1888. This species is regarded as a new genus, Psendalbizzia by Britton & Rose (N. Am, Fl. 23:48. 1928), and is according to them restricted to the West Indies, It has been variously interpreted as Acacia Berteriana, Inga fragransy Pitbecolobium Berterianay Acacia Balbistiy Acacia littoraliSy Albizzia Berterianay etc. Without seeing the Seemann specimen it is impossible to know to what entity Britton & (2} 3) [Vol. 2,7 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN West Indies. It is likely close to Pitbccolohium costariccnse if really a Pifhccolobium (in the broad sense). 3. ENTEROLOBIUM Mart, Enterolobtum Mart, in Flora 20:Bcibl. 117. 1837. Feuilh-ea Ktze. Rev. Gen. Pi. 1:182. 1891, in part. Unarmed trees, usually large, the branchlets pubescent or glabrous. Leaves bipinnatc, moderate or large, 2- or few-ranked; petiole prominent, bearing a gland usually near the middle; pinnae few to many, opposite, the rachis bearing glands especially terminally; leaflets usually numerous and small, inequilateral; stipules inconspicuous, caducous. Inflorescence of solitary to fasciculate pedunculate heads, axillary, or paniculate by insertion at defoliate nodes; peduncles slender; heads small, globular; flowers condensed, sessile, small; bracts small and inconspicuous. Flowers whitish; calyx campanulatc to tubular-funnelform, valvate, briefly 5- dentate; corolla more or less funnelform, 5-lobed, valvate, about double the calyx; stamens 10 to many, exceeding the corolla, united below into a short, included stamlnal tube; anthers small, eglandular, quadrangular; ovary sessile, several- ovulate; style slender, equalling or exceeding the stamens, the stigma small. Legume diagnostic, circinnate or rcniform, coiled into a complete or nearly complete circle, compressed or almost flattened but nevertheless somewhat fleshy, usually glabrous and lustrous, fairly broad, ligneous, indehiscent, septate between the seeds, the seeds transverse. Mexico to Argentina. Introduced into Pacific Islands and elsewhere. This small American genus (8 or less species, mostly South American) is re- tained separate from PithecoJobium (and Albizzia) following traditional practice. Nearly all students of the group have realized that Eufcrolobiiini probably consti- tutes no more than a section of Pifhccolobium (in the broader sense), but for con- venience have retained the genera separate. Morphologically the distinctiveness of Entcrolobhini rests mostly with the unusual, coiled legume. a. Flower uniformly and densely ferrui;inoiis-tomentulosc; petiole, rachis, and peduncle ferruginous-tomentulosc; leaflets up to 60 or more pairs per pinna, only 2-4 mm. long 1. E. ScHONfuURCKii aa. Cal)'X and corolla cancsccnt-tomentulose at tips of lobes; petiole, rachis, and peduncle usually subglabrous, not ferruginous; leaflets 15—30 pairs per pinna, S-1 5 mm. long 2. E. cyclocarpum 1. ENTr.ROLOBiUM ScuoMBURGKii Bcnth, in Trans. Linn. Soc. 30:599. 1875. I Vithecolobinm Schomburgkii Benth. in Hook. Lond. Jour. BoC. 3:219. 1844. Feuillcea SchomburgkJi Ktze. Rev. Gen. Pi. 1:189. 1891. Slender, erect tree, the branchlets ferruginous- tomentulose or puberulent. Leaves moderate, twice-compound, 2 -ranked, the leaflets very small; petioles mostly 2—3 cm. long, terete, ferruginous-tomentulose, bearing m\ ovoid, sessile gland on the upper surface more or less at the middle; rachis usually 6-20 cm. long, fer- (234) 1950] FLORA OF PANAMA (Legtiminosae) 249 %. Fig. 93. Entcrolobhirn SchomhurgkU ruginous-tomentulosc, ridged above along the center, bearing a small prominence or gland between or just below insertion of all or most pairs of pinnae; pinnae generally about 20 pairs, opposite or subopposite, the pinnular rachis mostly 4-7 cm. long, bearing glands (hidden by leaflets) between Insertion of the terminal 1 to few pairs of leaflets; leaflets many (about 60?) pairs per pinna, jinear or linear-oblong, more or less falcate, only 2-4 mm. long, obliquely rounded basally, inequilatcrally obtuse or acute apically, the costa excentric, glabrous above, pubes- cent below; stipules early caducous. Inflorescence of 1 to few pedunculate heads from several foliate or defoliate nodes of the terminal branchlets; peduncles about 2 cm. long, ferruginous-tomcntulose; heads globular, about 1 cm, in diameter (in bud), many-flowered; bracts minute. Flowers small, sessile, whitish; calyx funnelform, about 2 mm. long, ferruginous-tomentulose without, valvate; corolla tubular-funnelform, about 4 mm. long, ferruginous-tomentulose without, valvate; stamens 10 (or more?), about 1 cm. long, the filaments united below for less than half their length into a short, included staminal tube; ovary pubescent, terete. (2} 5} [Vol. 37 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN Legume completely coiled upon itself into a circle about 5-7 cm. in diameter, compressed or almost flattened, 2—3 cm. wide, glabrous. Mexico and Guatemala to Brai^iL CHiRiQuf: San Felix, Pit tier 52 J^- Panama: Cerro Jefe, Allen 3434; Rio Indio drain- age, Barbour I0j8. By reason of the distinctive fruit, this species is graced with many local, often picturesque, common names (viz. Brazil — orelha de macaco y "monkey*s ear"; orclha dc prctOj "nigger's ear"; etc., etc.). This species seems to have little com- mercial use in South America, although it is reported of value for lumber and ties in northern Central America. Except when In fruit this and the following species might be mistaken for Albizzia carlbaca and ^4. inaJacocarpa or A. idiopoda. m 2. Enterolobium cyclocarpum (Jacq.) Griseb, Fl. Brit. W. Ind. 226. 186L Mimosa cyclocarpa Jacq. Fragm, Bot. 30, /. 34, f, i. 1801, Inga cyclocarpa Willd. Sp. PI. 4:1026. 1806. ?Prosopis Juhia HBK. Nov. Gen. & Sp. 6:309. 1824, fide Benth. Vithccolobium cyclocarpum Mart. In Flora 20^:Beibl. 115. 1837. Mimosa Parota Sessc & Mog. Pi. Nov. Flisp. 257. 1887. Teuillcca cyclocarpa Ktze. Rev. Gen. Pi. 1:184. 1891. Spreading tree, often very large and buttressed on the trunk, the branchlets puberulent to glabrous, smooth. Leaves moderate, twice compound, few-ranked; petiole up to about S cm. long, subterete, swollen basally, glabrous or somewhat puberulent, bearing an ovoid gland on the upper side usually near the middle; rachls up to 15 or more cm. long, shallowly sulcate and submarginate above, glabrous or somewhat puberulent, bearing subcupular glands between the terminal and usually penultimate insertions of the pinnae; pinnae 4-15 pairs, opposite on the rachis, the pinnular rachis with sessile or partly sunken, oblong, concave glands near insertion of the terminal 1—4 pairs of leaflets; leaflets 15-30 pairs, opposite, small, inequilaterally oblong or oblong-subf alcate, 8-1 5 mm. long, obliquely rounded basally, obliquely acute-mucronate aplcally, subglabrous and dark above, appresscd-puberulent and lighter below, the costa excentric; stipules not evident. Inflorescence of 1-3 pedunculate heads from several foliate or (more frequently) defoliate nodes of the young branchlets; peduncle 2—4 cm. long, puberulent or subglabrous; head globular, 10—15 mm. In diameter, multiflorate; bracts minute. Flowers small, sessile, whitish; calyx short tubular-campanulate, about 3 mm. long, puberulent or subglabrous on the tube, canescent-tomentulose on tips of the lobes, valvate in bud; corolla tubular-funnelform, 5-6 mm. long, subglabrous except canescent-tomentulose toward the tip> valvate In bud; stamens many, up to 12 mm. long, filaments united into a staminal tube for about half their length; anthers quadrangular, unappendaged; ovary glabrous, subsulcate laterally; style about equalling or slightly exceeding the stamens. Legume rcniform, about 10 cm. across and 3-6 cm, wide, compressed, glabrous, lustrous, curved Into a nearly complete circle. Mexico to northern South America. (236) 1950] FLORA OF PANAMA (Legumtnosae) 251 BOCAS DEL TORO: Changuinola Valley, Cooper ^ Slater g4> canal zone: Balboa, Stanclley 30S4 (sterile), chiriqui: Progreso, Cooper & Slater 283 (sterile), cocle: La Venta, Muenscher t6j26; Penonome, Williams 288, Panama: Chepo, Kluge Sj; Punta Paitilla, Piper 5/J9. The species apparently grades into £. tbnbotiva Mart, of eastern and southern South America. £. cyclocarjnun is said to have a number of local uses, such as the pulp of the fruit for food and as a soap substitute, a gum from the trunk as a remedy, and the wood of the harder specimens, easily worked and polished and seasoning well, as a decorative or cabinet wood. Except in fruit the species may easily be confused with Albizzia malacocarpa and A. idiopoda (northern Central America) : the leaflets in these species of AlbizziUy however, are not as markedly mucronate apically as with E. cyclocarpiim. 4. ALBIZZIA Durazzlm Alsizzia Durazz. Mag. Tosc. 3:11. 1772. Sericandra Raf. Sylva Tellur. 119. 1838. Serianthes Benth, in Hook. Lend. Jour. Bot. 3:84. 1844. Besenna A. Rich. Tent. Fl. Abyss. 1:253. 1847. Trees or shrubs, mostly unarmed, pubescent or glabrous. Leaves few-ranked, usually large, bipinnate, the pinnae and leaflets few to many pairs; petiole gland- ular, the gland usually borne near or below the middle, less frequently above the middle, but always below insertion of the lowermost pair of pinnae; petiolar gland concave, longitudinally oblong and sunken in the petiole to orbicular and erect; rachis and pinnular rachis usually also bearing glands, especially towards the tips; leaflets small and linear to moderate and ovate, variously pubescent; stipules usually caducous or lacking. Inflorescence of usually subterminal, axillary, pedunculate heads, spikes, or umbels. Flowers pentamerous, synsepalous and sympetalous, regu- lar, almost invariably perfect, white to pink; calyx campanulate to tubular, toothed apically, usually pubescent; corolla mostly funnelform, valvate; stamens numerous, united below into a staminal tube which is usually included; anthers small, egland- ular. Legume hnear to oblong, relatively broad but very flat and thin, straight, dry, not septate, Indehisccnt or tardily dehiscent, the valves never contorted nor elastically dehiscent, the seeds transverse. Pantropical but relatively rare in the New World. Albizzia is primarily an Old World genus, scarcely distinguishable from Entero- lobitim and 'Pithccolobitim (primarily New World). The traditional means of dis- tinguishing the genus (on basis of the legume) is highly inconvenient. New supplementary characters are thus utilized here (especially position of glands: whether on petiole or rachis only) which seem definitely more practical in sep- arating Albizzia from Pithecolobinm. Separation of Albizzia from Enterolobinvi is more difficult in the absence of fruit, but since few species are then involved individual specific characters usually serve to make determination possible. Certain (237) [Vol. 37 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN species of Acacia so resemble a few species of Albizzta that detailed examination of the stamens is necessary for determination. a. Leaflets small and narrow (5—1 1 mm. long and about 2 mm. wide) , numerous (9-50 pairs per pinna), b. Leaflets less tlian 30 pair per pinna, pubescent; mature peduncles 2—3 cm. long; flowers relatively large (calyx about 2.5 mm. long) L A. carbonaria bb. Leaflets usually 3 or more pairs per pinna, glabrous; peduncles less than 1.5 cm. long; flowers small (calyx about 1 mm. long) 2. A. CARiBAtA aa. Leaflets larger and broader (mostly 15-40 mm. long and 10-20 mm. wide), few or several (2—9 pairs per pinna). b. Leaflets tapered and bluntly acute apically; flowers sessile 3. A. AnixoCEPHALA bb. Leaflets obtuse or rounded to emarginate apically; flowers pedicellate. c. Flowers all alike, short-pedicellate (pedicels mostly about 2 mm. long) ; mature leaflets, young twigs, and legume sparingly pubes- cent or glabrous 4. A. Lcbbeck cc. Flowers dimorphic, all except center one long-pcdicellate (pedicels 8—15 mm. long); mature leaflets, yt)ung twigs, and legume tomcn- tose to moderately pubescent 5. A. LONGEPFnATA 1, Albizzia CARBONARIA Britton ex Britt. & Wils., Scientif. Surv. Porto Rico & Virgin Isl. 6:348. 1926. Alhizzia >nahtcocarpa Standi, ex Standi. & Cald. Fl. Salv. 96. 1925 (hyponym); Britt. & Rose, in N. Am. Fl. 23:44. 1928, fide Britt. & Killip. Moder.ite or large tree, the branchlets tomentose when young becoming gla- brous, moderately ienticeUate, unarmed. Leaves moderately large, bipinnate, the pinnae 7—15 pairs opposite on the rachis, the leaflets 10—2 5 pairs per pinna; petioles mostly 2—3 cm. long, tomentulosc, nearly terete, usually eglandular; primary rachis up to 10 cm. long, similar to the petiole, bearing at the apex at insertion of terminal pair of pinnae a sessile, subcupular gland; pinnular rachis 2-5 cm. long, pubcrulcnt, usually bearing a sessile gland at insertion of the terminal pair of leaflets; leaflets opposite, oblong-subfalcate, 4-6 mm. long and 1-2 mm. wide, asymmetrically acute apically, obliquely rounded or subtruncate basally, pubescent above and more heavily so below, somewhat paler below, the costa exccntric, the venation obscure; stipules early caducous. Inflorescence of several pedunculate umbels from terminal and subterminal defoliate or minutely foliate nodes; peduncles slender, up to 3 cm. long, tomentulose; bractlets linear-spatulate, caducous. Flowers briefly pedicellate, dense; calyx narrowly obconic, about 2.5 mm. long, tomentulose; corolla narrowly funnclform, about 5 mm. long, tomentose, the lobes Yi the length of the tube; stamens about 14 mm. long, the staminal tube well Included; ovary glabrous. Legume broadly linear, 8-11 cm. long and almost 2 cm. wide, stipitate, pubescent, falsely septate, the seeds transverse. Salvador, Panama, and Colombia. UAKiJiN: Cana, Williams Qj2, A, vialacocarpa was described in part (flowers) from a Panama specimen (WiUidws OS-)' On the basis of certain characteristics A. carhonayiu nnght well be considered Pilhecolol^ium (or Samanca^ if this be recognized as distinct from Pithccolobinm) instead of Alhizzia, Yet the legume better fits Alhizzia, (2} 8) 19 50] FLORA OF PANAMA (Lcgtiminosae) 253 2. Albizzia caribaea (Urban) Britt. & Rose, in N. Am. Fl. 23:44. 1928. Pithecolobium caribacum Urban, Symb. Ant. 2:260. 1900. Moderate to tall tree, the branchlets puberulent or almost glabrous, inconspicu- ously Icnticellate. Leaves large, bipinnate, the pinnae 5—10 pairs, opposite, the leaflets 30-50 pairs; petiole 1-5 cm. long, terete except sulcate above, more or less puberulent, bearing above near the middle an orbicular gland concave above; rachis up to 18 cm, long, similar to the petiole, frequently glandular at the upper 1 to few rachial nodes; pinnular rachis about 6 cm. long, similar, eglandular; leaflets inequilatcrally linear, 5-11 (usually 7-8) mm. long and 1-2 mm. wide, subtruncate and oblique basally, asymmetrically obtuse apically, glabrous, the costa excentric; stipules caducous or lacking. Inflorescence of subterminal, axillary, pedunculate heads; peduncles short, 6—15 mm. long, pubescent or subglabrous; heads globose, the bracts minute. Flowers small, pink, sessile; calyx turbinate, about 1 mm. long, puberulent-pilosulous; corolla funnelform, about 4 mm. long, subglabrous; stamens about 1 cm. long, the staminal tube included; ovary glabrous or shortly pilose apically, several ovulate. Legume linear, usually about 10 cm. long and 1.5—2.5 cm. wide, straight, flat and thin, the margins elevated, glabrous or subglabrous, transversely striate, the seeds transverse. West Indies; north coast of South America, and lower Central America. SAN BLAs: Puerto Obaldia, Pitticr 4326. This species is distinguished from most species of the genus in Panama by the fine ''fern-like" foliage and short -pedunculate heads. Unless in fruit it Is easily mistaken for Enferolobium: however, the leaflets arc somewhat larger than is the case with £. SchGinbiirgkii and somewhat smaller than is usual for £. cyclocarpttm, the only two species of EnteroJob'ium occurring in Panama. 3. Albizzia adinocephala (Donn. Smith) Britt. & Rose ex Record in Trop. Woods 10:22. 1927. Pithecolobium adhioccphalum Donn. Smith, in Bot. Gaz. 57:419. 1914. 'Pithecolobium discolor Pittier, In Contr. U. S. Nat. Herb. 20:464. 1922, not Brittan. Large forest tree, the branchlets glabrous except at the extreme tips and some- what corky or Irregular. Leaves large, bipinnatCj the pinnae (1) 2—3 pairs op- posite on the rachis, the leaflets (2) 3—4 (5) pairs per pinna; peuolc slender, 2—8 cm. long, terete, glabrous or lightly puberulent, swollen basally, bearing a longi- tudinally elongate, concave gland sunken in the upper surface of the petiole below the middle; primary rachis similar to the petiole, eglandular, longitudinally sulcate above; pinnular rachis similar, except bearing glands like that of the petiole at the upper pinnular nodes; leaflets opposite, ovate-lanceolate to ovate-elliptic, mostly 2—4 cm. long and 1-2 cm. v/ide, bluntly acute apically, mostly obtuse basally, dull and lightly puberulent above, lighter and puberulent below, the veins very promi- nent and reticulate above; stipules apparently lacking. Inflorescence a terminal or subterminal panicle, the ultimate divisions capitate from slender, puberulent, (239) [Vol. 37 254 ANNALS OF THE MISSOURI BOTANICAL GARDEN ebractcolatc peduncles; heads several-flowered, dense, minutely bracteolate. Flowers small, white, fragrant; calyx funnelform, little more than 1 mm. long, silvery- pubescent above, the teeth small; corolla funnelform, about 3 mm. long, pubcru- lent or subglabrous, the teeth as long as the tube; stamens many, about 1 cm. long, glabrous, the staminal tube about equalling the corolla; ovary glabrous. Legume broadly linear, 10-17 cm. long and L5-2.0 cm. wide, flat and thin, glabrous, trans- versely striate, dehiscent, 10- to 13-seeded. British Honduras and Salvador to Panama. CANAI. ZONE: Ancon, Vitticr 5737\ Balboa, Standley 2604J. panama: Culebra Island, Hayes 2S; sterile specimens from Standlcy afford many other records from the Canal Zone and nearby Panama. A fairly common tree of Central America, much resembling certain species of Lejicacfut. 4. Albizzia Lebdeck (L.) Benth. In Hook. Lond. Jour. Bot. 3:87, 1844. Mimosa Lebbcck L. Sp. PI. 516. 1753. Mimosa spcciosa Jacq. Ic. Pi. Rar. 1:19, /»/. igS. 1786. Acacia Lcbbcck Willd. Sp. Pi. 4:1066. 1806. Teuillcca Lcbbcck Ktze. Rev. Gen. Pi. 1:184, 1891. Many additional synonyms are given by Bentham (Trans. Linn. Soc. 50:562. 1875) for this Old "World species. Small or moderate, spreading tree, the branchlets pubcrulent when young soon becoming glabrous, conspicuously white-lenticellate. Leaves moderately large, mostly blph-inate, the pinnae 2-4 pairs opposite on the rachis, the leaflets 4-9 pairs; petiole 2-10 cm. long, glabrous, bearing on the upper side belovv' the middle a sessile, oblong, concave gland; primary rachis similar to the petiole, sulcate above, often bearing a sessile, scutelliform gland at insertion of the upper pair of pinnae; pinnular rachis similar, bearing at or just below the upper 2-3 rachial nodes glands like the gland of the petiole; leaflets opposite, asymmetrically oblong, 2-4 cm. long and mostly 1-2 cm. wide, roundcd-subtruncatc to obtuse apically, obtuse and very inequilateral basally, essentially glabrous above and below. Inflorescence of many long-pedunculate umbels axillary from the subterminal nodes (2-3 per node); peduncles 3-10 cm, long, sparingly puberulent; umbels dense, globose, the pedicels mostly about 2 mm. long, the bractlcts minute. Flowers cream-colored, fragrant; calyx funnelform, about 4 mm. long, unequally toothed or cleft, golden- tomentulose apically; corolla funnelform, 6-7 mm. long, puberulent, the teeth prominent; stamens many, as much as 3 cm. long, glabrous, the staminal tube well included; ovary glabrous. Legume broadly linear or narrowly oblong, usually 15 or more cm. long and 3-5 cm. wide, very thin and dry, glabrous, thinly mar- gined, transversely striate; seeds few, transverse. Native to Africa and Asia, Introduced Into various parts of the New World tropics. CANAL zone: Colon, Wittier 66^2, (240) 195 0] 1 FLORA OF PANAMA (Legummosae) 255 The popular name "woman's tongue" is given this species. It is quite common in the West Indies, more so than on the Central American mainland. In fruit it may be confused with species of Leiicacna. Fig. 94. Albfzz'ia longepedata 5. Albtzzia longepedata (Pittier) Britt. & Rose, ex Record In Trop. Woods 11:14. 1927. Fithecolobium longepedatum Pittier, in Contr. U. S. Nat. Herb. 20:464. 1922. Samanea Samanigua Pittier, In Boh Cient. y Teen. Mus. Com, Venez. 1:54. 192 5. Large tree, the young branchlets brownish-tomentose. Leaves large, bipinnatc, the pinnae 2—6 (usually 3—4) pairs opposite on the rachls, the leaflets 3—7 pairs per pinna; petioles usually 3—4 cm. long, brownish-tomentose, bearing a sessile, con- cave, patelliform gland on the upper side near or slightly above the middle; rachls 3—15 cm. long, similar to the petiole, often with sessile, concave glands at or below insertion of the pairs of pinnae; pinnular rachls similar, bearing small glands below the rachial nodes; leaflets ovate to obovate, 15-40 mm. long and 10-25 mm. wide. rou nded to subcuneate and somewhat obhque basally, rounded or emarginate aplcally, opposite, softly pubescent above and more densely so below, the venation (241) 256 ANNALS OF THE MISSOUFvI BOTANICAL GARDEN [Vol. 17 prominent above; stipules linear, caducous. Inflorescence of pedunculate umbels axillary from the subterminal nodes (1-3 per node); peduncle 3-9 cm. long, brownish-tomentosc; pedicels slender, 8-15 mm. long, tomentose; bractlcts linear, slender, early caducous. Flowers whitish, in many-flowered umbels, dimorphic, the central flower sessile and gross; calyx funnclform, 6-7 mm. long (tubular and 10-12 mm. long in central, sessile flower), tomentose; corolla funnelform, about 10 mm. long (tubular and 20-25 mm. long in central, sessile flower), tomentose, the teeth prominent and acute; stamens about 3 5 mm. long, the staminal tube included; ovary pubescent. Legume broadly linear, 15-20 cm. long and 2.5-3.5 cm. wide, pubescent, tardily dehiscent, the seeds several, transverse. Salvador and Guatemala to northern South America. CANAL zone: R. Pequeni, Steyermark & Allen 17155. This species is quite distinct from other species of Alb'izzia in Panama, difl'ering primarily by its pronouncedly umbellate, long-pediccllate flowers. 5. CALLIANDRA Benth. Calliandra Benth. in Hook. Jour. Bot. 2:13 8. 1840, nom. conserv. AftncsJia Salisb. Parad. Lond., pL 64. 1807. Juga Anncslcya G. Don, Gen. Hist. Dichl. Pi. 2:396. 1832, in part. Clclia Casar. Nov. Stirp. Brasil. Dec. 83. 1S45. CoJonafhlra Karst. Fl. Columb. 2:43, /. 122. 1862. Usually shrubs or small trees, unarmed except In one or two (non-Panamanian) species, mostly not heavily pubescent. Leaves bipinnate, the pinnae 1 to several pairs; leaflets 1 to many pairs per pinna; petiole and rachis cglandular; leaflets large and broad (ovate) or more frequently small and narrow (linear or Uncar-oblong) , glabrous In most species, frequcMitly ciliate marginally, almost invariably very oblique basally, the costa prominent, but the secondary veins obscure in many species; stipules mostly striate and subpersistent. Inflorescence commonly of axillary, pedunculate spikes, sometimes a terminal raceme or panicle. Flowers in dense, globose heads, sometimes dimorphic (center flower of head larger or more gross, sometimes sterile), normally 5-partcd and perfect, regular, sympetalous; calyx relatively broad, mostly cupulate or campanulate; corolla usually funnclform, never narrow and tubular like species of 7;/^^ and Pif/j('coJohii(ni, usually lobed for about half its length, the lobes valvate; stamens many, elongate, conspicuous and showy, obscuring the corolla, united below into an included or exscrteJ staminal lube; anthers small, usually eglandular; ovary usually sessile, glabrous, scveral- ovulate, the style filiform, the stigma terminal. Legume broadly linear, straight or nearly so, gradually constricted bascward and thus cuneate-substipitate, flat and thin, the margins elevated, bivalvate, tlie valves elastically dehiscent from the apex but not twisting, not septate, the seeds (oblique!}) transverse or nearly longi- tudinal. Primarily a New World genus, extending from southwestern United States and Mexico to southern South America: one or two Oriental or African species have (242) 19 50] FLORA OF PANAMA (Lcgtiminosae) 257 at times been recognized. The genus is fairly distinctive and seemingly quite natural. It is especially characterized by the glandiess petiole and rachis, the dense heads of "cxserted stamens" and the distinctive, elastically dehiscent legume. Specific boundaries within the genus are for the most part indistinct and often artificial. Several readily identifiable complexes can be distinguished, and within these a great many of the species can be united with profit. Probably not more than 20 or 25 "good" species occur in all Central America, most of these in the northern part. Eight species are here recognized from Panama, one of which is questionably recorded. a. Leaflets normally 3 per pinna, comparatively large and broad (about 1 cm. wide) ; pinnae 1 pair. b. Flowers large (corolla about 13 mm. long, stamens 6-7 cm. long); staminal tube well cxserted (about 4 cm. long); peduncles thick 1. C. glaberrima bb. Flowers smaller (corolla about 6 mm. long, stamens 2-2.5 cm. long); staminal tube included; peduncles slender 2. C. Seemanni aa. Leaflets few to many per pinna, comparatively small (never more than a few mm. wide) ; pinnae 1 to several pairs, b. Inflorescence of axillary, pedunculate heads; pinnae 1-1 1 pairs. c. Pinnae a single pair 3. C. Magdalenae cc. Pinnae 2—11 pairs. d. Corolla essentially glabrous; leaflets membranaceous, dull, the secondary veins obscure. e. Branchlcts terete; flowers rather small 4. C. caracasana ee. Branchlcts quadran!;ular, markedly angled; flowers larger 5. C. tetragona dd. Corolla pubescent; leaflets thicker, often subcoriaccous, darker or more lustrous above, venation more or less evident as far as known. e. Pinnae 2-6 pairs; leaflets usually 10-25 pairs, 4-7 mm. long; corolla about 8 mm. long.- 6. C. Cumingii cc. Pinnae mostly 7— 11 pairs; leaflets normally 20-65 pairs, mostly 3-5 mm. long; corolla 5-6 mm. long -7. C. Pittieri bb. Inflorescence a terminal raceme or panicle; pinnae several to many (usually about 15) pairs 8. C. CONFUSA 1. Calliandra glaberrima (Bentb,) Britt. & Klllipj in Ann. N. Y. Acad. Sci. 35:134. 1936. CcilliiniJra glypboxylon var. glaberrima Benth. in Trans. Linn. Soc. 30:539. 1875. Small tree or slirub?, the branchlets essentially glabrous, striate, callous but hardly lenticellate. Leaves moderately small, bipinnate, the pinnae only 1 pair, the leaflets 3 (2-4?) per pinna; petioles mostly 1-2 cm. long, subglabrous, sulcate above, eglandular, aplculate apically from the lower side; pinnular rachis similar to the petiole; leaflets oblong-falcate, the terminal pair mostly 2-4 cm. long and about 1 cm. wide, the lower leaflet usually half or less this size, blunt or bluntly acute apically and minutely mucronulatc, very obliquely obtuse or rounded basally, glabrous and lustrous above and below, sparingly ciliate on the margins, the 2-3 main veins palmate, smaller veins conspicuously reticulate, coriaceous; stipules QY^te, 1-2 mm. long, longitudinally striate, subperslstcnt. Inflorescence of axillary, pedunculate heads; peduncles about 1 cm. long; heads dense; bracts more or less ovate, little more than 1 mm. long, persistent. Flowers large, red; calyx cam- panulate, 2-3 mm. long, striate, glabrous, the teeth short and broad; corolla funnel- (243 ) I Vol. Z7 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN form, about 13 mm. long, glabrous; stamens as much as 6-7 cm. long, united below into a glabrous stamina! tube often about 4 cm. long, thus strongly exserted; ovary glabrous, tapered basally. Legume typical of the genus. Colombia and Panama. CANAL zone: R. InJio, DoJgc ^ Allen 173/0, cocLE: Bismarck, Williams 14T, PANAMA; north of Panama City, Bro. Paid 612, A few South American species, including C. carbonar'ia, are similar to our Panamanian specimens, and with additional collections may possibly prove to grade into this species and/or Into one another. Knowing how variable species in this group may be, wc have refrained from describing as new the DoJge £3" Allen ^737^ specimen cited, even though its flowers are considerably larger than is typical for C. glaherrima or other related species examined. The description here given is taken mostly from the DoJi^e d Allen 1737^ specimen and is not necessarily "typical" of C. glabcrrima, C. glaberrima is allied to the well-known C. emarginata of northern Central America and Mexico, and to other species of Britton & Rose's Ti^kgi-mixae. The Tergeminae as constituted by Britton and Rose is a highly unworkable complex of doubtful entities. Seemingly it should be condensed Into a half-dozen or less species. In spite of marked variability in the species, none of the Mexican or Cen- tral American ones possess flowers nearly so large as those of our Panamanian cita- tion. C. glaberrhua, along with C. Secnianui and Its extra-Panamanian allied species, are easily distinguished from all other species of Calliandra in Panama by the bi- pinnular, few-foliolate leaves with relatively large leaflets. Larger flowers and extremely exserted stamlnal tube serve readily to distinguish C. glabcrrinta from C. Scemafini. 2, Calliandra Seemanni Benth. In Seem. Bot, Voy. Herald, 116, pL 22, 13 53. Feuillcea Seemanni Ktze. Rev. Gen. Pi. 1:189. 1891. Annesha Seemanni Britt. & Rose, In N. Am. Fl. 23:54. 1928. Large shrub or small tree?, the branchlets glabrous or lightly canescent-puberu- lent. Leaves moderately small, biplnnate, the pinnae a single pair, the leaflets 1 pair (plus a third smaller leaflet below to the outside) per pinna; petiole slender, up to 2 cm. long, essentially glabrous, sulcate above, eglandular, apiculatc from below terminally; pinnular rachis similar to the petiole; leaflets obliquely oblong, mostly 2-4 cm. long and about 1—1.5 cm. wide, obtuse to acute aplcally, rounded and very inequilateral basally, with one chief nerve and two smaller nerves palmate from the base, the secondary venation reticulate, glabrous except usually ciliolate on the margin; stipules lanceolate, small. Inflorescence of axillary, pedunculate heads; peduncles slender, mostly 2-3 cm. long; heads several-flowered, globular; floral bracts lanceolate, subpersistent. Flowers dense; calyx narrowly cupulate, about 2 mm. long, glabrous, striate, the teeth prominent; corolla funnelform. (244) 1950] FLORA OF PANAMA (Legumifiosae) 259 about 6 mm. long, glabrous; stamens 2-2.5 cm. long, united below into an in- cluded staminal tube. Legume broadly linear, up to 10 cm. long and about 1 cm. wide, glabrous, the few brown seeds reported white-dotted. Reported from northern South America (erroneously from Colombia, fide Britton & Killip) ; Costa Rica?. The species is based upon a Seemann collection from Veraguas, Panama. The type (the only Panamanian record) has not been available for examination, but is well illustrated with the original description: it is strongly suspected that the species is merely a variant of the older C. emarginata (which in turn is probably scarcely separable from C. tcrgemhia), and incorporation under one of these names may eventually be in order. As mentioned under discussion of C. glaberrima, it seems impossible to accept Britton & Rose's inordinate splitting of the Tergeminae, into which section the doubtfully distinct C. Seemanni falls. 3. Calliandra Magdalenae (Bert.) Benth. in Lond. Jour. Bot. 5:102. 1846. Acacia Magdalenae Bert, ex DC. Prodr. 2:455. 1825. Feuillcca Magdalenae Ktze. Rev. Gen. Pi. 1:188. 1891. Calliandra riparia Pittier, Arbol. y Arbust. Legum. Dec. 6-8:80. 1927. Anneslia Magdalenae Britt. & Rose, in N. Am. Fl. 23:60. 1928. Auneslia Tonduzii Britt. & Rose, loc. cit. 61. 1928. Anneslia cbiapensJs Britt. & Rose, loc. cit. 1928. Calliandra Schultzei Harms, in Feddc Rep. Sp. Nov. 24:209. 1928. Calliandra Tonduzii Standi, in Field Mus. PubL Bot. 4:309. 1929. Calliandra angusiidens Britt. & Killip, in Ann. N. Y. Acad. Scl. 35:134. 1936. Tree to 10 m. tall, the branchlets somewhat ascending-pubescent when young, becoming glabrous in age. Leaves moderate, bipinnate, the pinnae 1 pair, the leaflets several (usually 10-20) per pinna; petiole short, about 1 cm. long, terete, tomentulose, eglandular, bearing terminally an apicule from the lower side; pin- nular rachis 5-9 cm. long, ridged and tomentose above; leaflets linear-ovate or linear-lanceolate to linear-oblong, mostly about 1 cm. long and 3 mm. wide, broadly acute or sometimes obtuse apically and minutely mucronulate, obliquely truncate basally, glabrous above and below, lustrous above, often ciliate, the costa slightly excentric, 1-2 minor veins palmate from the base, secondary veins promi- nently reticulate; stipules lanceolate, 3-6 mm. long, striate, subpersistent. In- florescence of axillary, pedunculate heads; peduncles about equalling petioles, puberulent; floral bracts lanceolate, quite small, persistent. Flowers rose-red, several In dense globular h about glabrous, striate; corolla funnelform, 5-7 mm. long, glabrous, the lobes prominent; stamens In age 2.5 or more cm. long, the staminal tube about equalling the corolla. Legume linear-oblong, reported up to 14 cm. long and about 1 cm. wide, pubescent when young, said to be (erroneously?) glabrous at maturity. As here interpreted (broadly), southern Mexico, Central America to northern So CHiRiQui: Progreso, Cooper 6 Slater 318; Remedies, Allen 3666; San Felix, Allen 1958; R. Tinta, Woodson, Allen ^ Seibert 401. veraguas: Sona, Allen 1040, (245) 260 [Vol. Z7 MISSOURI While the species assumes a fairly constant form in Panama, it grades into a number of variants in northern Central America and Mexico, and in northern South America. As a result numerous bipinnular "species" have been described which are nearly impossible to delimit precisely. South American (Colombian) specimens taken by Britton & KiUip to be C. MagJalenac may appear quite dif- ferent from those of northern Central America and even Panama. On the other hand, there seems little doubt that the Colombian C. Schulfzci is the same species as our Panamanian citations. Yet Bentham cites a Seemann collection (as being C. MagJalcfiac) from the same province as the Allen IO40 collection. With un- certainty as to the exact boundaries of C. Magclalcnae, a few possible and probable synonyms have been combined for practical reasons under the oldest name. Should C. Sc/jul/zei in reality be distinct from C, MagJalcnae, both it (on the basis of recent collections) and C. Magdalenae (solely on the basis of Bentham's citation) would have to be considered as occurring in Panama. 4. Calliandra caracasana (Jacq.) Bcnth. in Trans. Linn. Soc. 30:543. 1875. Mimosa caracasana Jncq. Coll. 4:142. 1790. Acacia caracasana Willd. Sp. PI. 4:1068. 1806. ?Calliafu[ra portoriccusis Bcnth. in Hook. Lond. Jour. Bot. 3:99. 1844. (For numerous synonyms to this species see Bcnth. in Trans. Linn. Soc. 30:543. 1875.) Fcuillrca caracasana Ktze. Rev. Gen. Pi. 1:187. 1891, Anncslia caracasana Britt. & Rose, in N. Am. Fl. 23:65. 1928. CallianJra Toroana Britt. & Rose, in Ann. N. Y. Acad. Sci. 35:137. 1936. Shrub or weak slender tree, the branchlets puberulcnt to glabrous. Leaves moderately large, blplnnate, the pinnae 2-6 (mostly 4-5) pairs, the leaflets 10-3 5 pairs; petiole slender, 1-3 cm. long, glabrous or nearly so, eglandular; rachis mostly 3-4 cm. long, similar to the petiole, bearing a terminal apicule from the lower side; pinnular rachis up to 7 cm. long, glabrous except frequently puberulcnt above; leaflets linear or linear-oblong and more or less straight, mostly 6-8 mm. long and about 1 mm. wide, obtuse or somewhat acute apically, asymmetrically rounded or subtruncatc basally, glabrous (in age) or pubescent, dull above and below, the costa somewhat excentric, other veins obscure; stipules lanceolate, 3-6 mm. long. Inflorescence of axillary, pedunculate heads; peduncles slender, 2-5 cm. long or even longer, glabrous or puberulcnt; floral bracts minute, usually ciliate; head small, several-flowered, globular. Flowers whitish or pinkish; calyx funnelform-cam- panulate, scarcely more than 1 mm. long, subglabrous; corolla broadly funnclform, little more than 2 mm. long, glabrous or subglabrous; stamens about 16 mm. long, the staminal tube included. Legume broadly linear, about 8 cm. long and almost 1 cm. wide, constricted basally, glabrous or lightly puberulcnt, Venezuela to Ecuador; Panama. "Species" probably conspeciflc with C. cara- casana if broadly regarded, occur in Mexico, northern and middle Central America and the West Indies. CHiRigui: R. Chiriqui Vicjo, P. White 342. (246) 1950J FLORA OF PANAMA (Legumhiosae) 261 The White J42 specimen cited (fruit only) is in rather poor condition. It seems identical with C. tetragona and logically should be cited as C. tetragona^ except for the absence of the characteristic stem angulations, a character possibly of httle genetic weight. Even if this specimen be regarded as /. tetragona, the entity represented by C caracasana is to be expected in Panama. C. portoricensis has been distinguished from the usually slighter C, caracasana in reportedly having white stamens while the latter has pink, red, or purple stamens. This character- istic, in view of morphological intcrgradation, seems of dubious value for specific distinction. Bcntham himself states (Trans, Linn. Soc. 30:543. 1875) concerning C. caracasana: "The six preceding species [C. portoricensis is the first preceding one and C. tetragona the first subsequent one], however, are very closely alHed to each other, and might almost be classed as varieties of a single species." Mono- graphic work is sorely in need here and will likely show a widespread, variable complex centered in these tropical American species, in which speclation has not as yet segregated clear-cut entities. 5. Calliandra tetragona (Willd.) Benth. in Hook. Lond. Jour. Bot. 2:139. 1840. Acacia tetragona Willd. Sp. Pi. 4:1069. 1806, Mimosa quadrangularis Poir. in Lam. Encycl. Mcth. Suppl. 1:72. 1810. Acacia qnaJrangularis Link, Enum. Hort. Berol. 2:445. 1822. Anncslia tetragona Donn. -Smith, Enum. PI. Guat. 1:10. 18 89. Fetdllcea tetragona Ktzc. Rev. Gen. PI. 1:189. 1891. Calliandra portoricensis var. miiltijuga Micheli, in Bot. Gaz. 20:285. 1895, fide Britt, & Rose. Shrub or small tree, the branchlets puberulent when young becoming glabrate, usually markedly quadrangular, longitudinally striate, scarcely lenticellate. Leaves moderately large, bipinnate, the pinnae 4—8 pairs opposite on the rachis, the leaflets 15-3 5 pairs per pinna; petiole mostly 2-3 cm. long, puberulent or subglabrous, angled, somewhat sulcate or ridged above, eglandular; rachis similar to the petiole, often bearing tufts of hair at insertion of the pinnae; pinnular rachis 3-8 cm. long; leaflets linear to linear-oblong, nearly straight, mostly about 8 mm. long and little more than 1 mm. wide, obtuse or somewhat acute apically, asymmetrically rounded to truncate basally, glabrous or cillate on the margin, dull above and below, the costa somewhat excentric, the other veins obscure; stipules narrowly lanceolate, about 6 mm. long, striate, subpersistent. Inflorescence of 1 to few pedunculate heads from the upper axils, the terminal and subterminal nodes fre- quently defoliate; peduncles elongate, not uncommonly 7—8 cm. long in age, sim- ilar to the petiole; heads dense, several-flowered, globular; floral bracts minute. Flowers white; calyx funnelform-campanulate, about 2 mm. long, subglabrous, the teeth acute; corolla funnelform, about 6 mm. long, glabrous; stamens 3 cm. long or longer, the staminal tube included. Legume up to 12 cm. long and about 1 cm, wide, glabrous, otherwise typical of the genus. Southern Mexico to northern South America. (247) 262 [Vol. 37 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fig. 95. Calliaudra ictragana CHiRiQuf: Boquetc, ?itticr JI40; Hato del Jobo, Pitficr 5423. COOLt: El Valle, Allen 381Q; Hufiler (^ Allen 357. This species IS distinguished, perhaps somewhat "unnaturally," by the tetra- gonal branchlets. Certainly some species normally identified as C portoricensis or C caracasana seem identical T\uth C. tctragona except for the angled stem. It may be that a single or few-gene characteristic (angled stem) has appeared throughout the range of what might be called the C caracasana~C. porforicer^sis complex. Yet this character is sufficient to distinguish (somewhat arbitrarily) this species from others of the genus, 6. Calliandra Cumingii Benth. in Hook. Jour. Bot. 2:140. 1840. lni;j speclosa Mart. & Gal. In Bull. Acad. Brux. 10-:320. 1843?, fide Ind. Kew. Feuilleea Cumingii Ktzc. Rev. Gen. PI. 1:187. 1891. AnnesUa Cumingii Britt. & Rose, in N. Am. Fl. 23:57. 1928. Shrub, the branchlets puberulent or lightly pubescent, becoming glabrous in age. Leaves moderately small, bipinnate, the pinnae usually 2-3 pairs opposite on the rachls, the leaflets several to many pairs per pinna; petiole usually less than 1 cm. (24S) 195 0] FLORA OF PANAMA (Leguminosac) 263 longj lightly pubescent, eglandular; racKis 5-20 mm, long similar to the petiole, eglandular, bearing a terminal apicule from the lower side; pinnular rachis slender, up to 2 cm. long, puberulent, eglandular; leaflets linear-oblong or narrowly oblong, usually 4-7 mm. long and 1-2 mm. wide, blunt or obtuse and often miniitely mucronate apically, obUquely rounded or subtruncate basally, sparsely pubescent or subglabrous, the costa slightly excentric, the venation fairly prominent below, somewhat lighter In color and duller below than above; stipules small, lanceolate- subulate. Inflorescence of pedunculate heads solitary or rarely geminate in the upper axils; peduncles 2-3 cm, long, lightly pubescent; floral bracts small, puberu- lent. Flowers in compact, several-flowered heads, pubescent or lightly pilose apically, purplish; calyx campanulate, 1-2 mm. long, pubescent on the teeth; corolla funnelform, about 8 mm. long, lightly pubescent with long hairs, the lobes about Vs—Va the length of the tube; stamens about 2.5 cm. long, the staminal tube mostly included; ovary glabrous. Legume linear, up to 8 cm. long and about 8 mm. wide, puberulent?, dehiscent as typical of the genus. Panama, Mexico?, northern South America? PROVINCE unkno^-'N: without locality, Cuming 1 248, The species was described from a Panamanian collection by Cuming (1248)^ which we have not had opportunity to examine. No later collections from Panama have come to our attention. Bentnam regarded Mexican and Venezuelan material as conspecific with C CnmingU, but Britton and Rose have regarded the species as confined to Panama. The above description is pieced together from various in- complete descriptions of the species and certain herbarium sheets that presumably are (or are very similar to) C CuniingiL 7. Calliandra Pittieri Standi, in Contr. U. S. Nat. Herb. 18:104. 1916. Calltajulra bcUa van Triauac Benth. in Trans. Linn. Soc. 30:556. 1875. Ccilliaficlni Lehnianii Harms, in Feddc Rep. Sp. Nov. 17:90. 1921. Anfjcslia Pittieri Britt. & Rose, in N. Am. Fl. 23:63. 1928. Small tree with flat crown, the branchlets sparingly appressed-pubcscent or puberulent when young, becoming glabrous. Leaves moderate, bipinnate, the pinnae mostly 7-11 pairs opposite on the rachis, the leaflets several or many (20-65 pairs) ; petiole short, normally less than 1 cm. long, terete, puberulent, eglandular; rachis up to 9 cm. long, similar to the pctialc; pinnular rachis 2—5 cm. long; leaf- lets small, linear or linear-oblong, 3-5 mm. long and less than 1 mm. wide, acute to obtuse apically, inequilaterally rounded basally, usually minutely ciliolate but otherwise glabrous, the costa conspicuous and somewhat excentric, more or less lustrous on the upper surface; stipules linear-lanceolate, about 3 mm. long, per- sistent. Inflorescence of axillary, pedunculate heads; peduncles 1.5-4.5 cm. long, puberulent; heads few- to several-flowered, dense, spherical, the bracts small and subpersistent. Flowers red; calyx campanulate, little more than 1 mm. long, glabrous or puberulent, the teeth short and broad; corolla funnelform, 5-6 mm. long, pubescent; stamens about 3 cm. long, the staminal tube barely exserted. (249) 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 Legume lincar-spatulate, 5-8 cm. long and scarcely 1 cm, wide, pubescent, the margins thick. Colombia and Panama. darien: Cana, Williams 707. This species, from a Colombian type {Vitticr QSl), was recorded in the original description as occurring also in Panama (Cana, Darien, Williams 7^7). Britton and Rose, in the N. Am. Fl., so accepted its range. However, Britton and Killip (Ann. N. Y. Acad. Sci. 35:136, 1936) indicate that the Panamanian identifica- tion was in error, and that the species is in reality confined to Colombia. Morpho- logically, there seems little reason why the Williams 707 specimen cannot be considered as C. Piitieri, and it is so listed here. 8. Calliandra coniusa Spraguc & Riley in Kew Bull. Misc. Inf. 1923:371- 1923. Calliandra similis Sprague & Riley, loc. cit. 372. 1923. Anneslia confusa Brltt. & Rose, in N. Am. Fl. 23:70. 1928. AnncsUa similis Britt. & Rose, loc. cit. 71. 1928. Small tree or shrub, the branchlets lightly hirsute to subglabrous, longitudinally striate. Leaves large, bipinnate, the pinnae 10-20 or more (usually about 15) pairs, opposite, the leaflets numerous (30-60, usually 40-50, pairs) ; petiole usually about 2 cm. long, pubescent to subglabrous, sulcate above, eglandular; rachls sim- ilar, 8-15 cm. long; pinnular rachis about 4 cm, long, ridged above, lightly pubes- cent* leaflets linear, usually about 5-6 mm. long and 1 mm. wide, acute or narrowly obtuse apically, obliquely truncate basally, glabrous above and below except ap- pressed-ciliate on the margins, only the midvein prominent, this more or less central; stipules linear-lanceolate, 4-5 mm. long, glabrous, striate, caducous. In- florescence a terminal raceme or panicle; peduncles mostly 5-8 mm. long, geminate or few-fasciculate, subglabrous; pedicels 2-4 mm. long; bracts lanceolate-caducous. Flowers in few-flowered umbels, purplish-red; calyx broadly campanulate, scarcely 2 mm. long, glabrous, 5-toothed, the teeth broad; corolla 5-lobed, cleft almost to the base, the lobes elliptic or oblong, 5-6 mm. long, glabrous, the tube about 1 mm. long; stamens as much as 6 cm. long, united below into a short staminal tube 2-3 mm. long; ovary glabrous. Legume almost 8 cm. long and about 1 cm. wide, strigose-pubescent, dehiscing as typical of the genus. Panama to northern Central America and southern Mexico; northern South America? culRlQuf: Boquete, Terry & Terry l6j7; Woodson Sf Schery 7^7; R- Chiriqui Viejo, G. & P. White 112; Allen 4651. This species is a member of the "Houstoniana complex" (Houstonianae of Britton and Rose) very common in Mexico and to a lesser extent in northern Cen- tral America. There arc probably 3 or 4 tenable species in that area, of which C. Houstoniana dating from 1768 and C. f^ramli flora dating from 1788 are the best (250) I ysoi FLORA OF TANAMA (Lcguminosae) 265 Fig. 96, Calliandra confusa known (C. anoinaja and C. strigillosa of the early 18 00's may also be "good" species, but several additional "species" in this group, proposed by Britton and Rose and others, seem anything but clear-cut). The southern extension of this com- plex in Central America (Honduras to Panama) seems to be confined to the single species here listed. It is impossible to say without examining the type whether or not the earlier name, C. calothyrstis Meissn., for a South American specimen (col- lected in 1846 by Kegel in Surinam), can apply to the entity found in Panama. In describing C. confusa and C, similis, Sprague and Riley seem to have hit upon two extremes of the same entity. Our cited specimens and others from Costa Rica to Guatemala appear to be quite intermediate between characteristics described for these names. Hence it has been necessary to regard C. s'wiilis as a synonym of C. confusa. (251) [Vol. 37 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN C. confusa is similar to C. granJiflora, but lacks the floral pubescence of tbat species. It is quite distinct from other species of Calliauclra occurring in Panama, being the only species there having a terminal, racemose or paniculate type of in- florescence. The plant is very attractive because of the showy purplish-red flowers and fine foliage, and deserves attention as an ornamental. 6. ACACIA Willd. Acacia Willd. Sp. Pi. 4:1049. 1806. Numerous synonyms occur for the genus. Segregate genera proposed by Rafincsquc (Sylva TeUur. 118-120. 1838) include: Cuparilln, Drepapbyla, Ehurnax, Esclcrofui, Gumi- fcra, Hccatandra, Popofrax, ScfiegaUdy and Zignmloha, Britton and Rose segregates (N. Am. Fl. 23:85-96. 1928) include: Araciclla, Acaciopsisy Babumiu, Fishlockia, Lucaya, Myrme- coJi'fiilrofjy Tauroceras, and Fcruracin Britt. & Bro. Leon. Dalla Torre £c Harms furtlicr list the following synonyms: Arthospriofi Hassk.; ChifojiuNt/jus Lehm.; Farncsia Gasparr.; Hoopcsiu Buk^kl.; PbyUodocc Link; Sassu Bruce ex Gmelin; Tctrachcilos Lchm.;V a cbdlia Wight & Arn. Woody plants, usually armed with scattered thorns or stipular spines. Leaves (Central American species) bipinnate, the pinnae few or many, opposite, the leaf- lets mostly small and numerous; petiole in the majority of species bearing some sort of a gland or glandular field; rachis short or elongate, glandular or less often eglandular; stipules usually small and caducous but sometimes modified as promi- nent spines, in the "bull-horn" species these very large and myrmecophilous. In- florescence basically of axillary, solitary or fasciculate, pedunculate heads, spikes, umbels or racemes, often appearing paniculate because of suppression of leaves at terminal and subterminal, floriferous nodes. Flowers small, usually numerous and dense, predominately whitish or yellowish, normally exceeding the bractlets; calyx usually campanulatc, sympetalous, 4- or 5-toothed; corolla typically funnclform, sympetalous for most of its length or in some species 4- to 5-cleft nearly to the base; stamens diagnostic, numerous, free or rarely somewhat united at the extreme base; anthers small, rarely glandular; ovary small, the style and terminal stigma scarcely visible among the stamens. Legume usually oblong to linear, terete or more frequently compressed or flattened, usually straight, dehiscent and with slightly elevated margins, the seeds transverse. Tropics and subtropics of New and Old Worlds, very prominent in Australia. The genus is highly variable, with many groups of species simulating diverse other genera (parallel evolution) and reflecting environmental adaptation (xero- phytism, etc.). It is thus extremely difficult to give a generalized yet definitive description for the genus (the numerous, free or nearly free stamens being the only binding characteristic), but it is even more diflficult to establish clear-cut sub- divisions of or within the genus. Most of the numerous segregates of Acarhj such as those recognized by Britton and Rose (N. Am. Fl. 23:84. 1928), depend upon subjective evaluation (viz.: coriaceous vs. cartilaginous; linear vs. oblong; subtercte vs. turgid; promptly vs. tardily dehiscent; etc.). It becomes highly in- convenient with these segregates to determine the '*genus," let alone the species, (252) 19S0] FLORA OF PANAMA (Lcgiimhiosae) 267 except where perfect material (with fruit) and at least modest familiarity with the group is to be had. Even then many marginal cases exist, where a species may well fall into either of two '^genera." Thus, for convenience, if nothing else, I prefer to recognize Acacia in the traditional broad sense, In spite of certain groups of species forming islands within the genus markedly different from other group- ings of species. The species of Acacia usually present considerable variability and intergrada- tion. It would seem that formulation of species has been unusually inordinate in this genus. When material from Mexico to middle South America Is examined and compared, or in areas from which abundant material is available, many specific distinctions fade. This appears especially true in the segregate genera ^'Acaciclla^^ and ^^Senegalia^* as considered by Britton and Rose. On the other hand, many of the "bull-horn" species, differing but difficult of separation, may prove constant with further collections. Collections of all species in Panama are rather meagre, with the possible exception of species occurring on Barro Colorado Island. It is as yet not possible to study degree of variation of these species within the country, nor to confidently correlate single or few collections with better known Mexican and northern Central American expressions of (presumably) the same entity. a. Flowers pedicellate; petiole eglanjular; twigs unarmed C'Acaciella''). h. Leaflets nearly linear (about 6 mm. long and 1 mm. wide), numerous (frequently 60 or more pairs per pinna), subglabrous; twigs com- monly subglabrous 1. A. axgustissima bb. Leaflets oblong or ovate (usually 8-9 mm. long and at least 2 mm. wide), fewer (9-25 pairs per pinna) , pubescent; twigs often villous.... 2. A. villosa aa. Flowers sessile or subsessile; petiole with 1 or more glands; twigs rarely unarmed, b. Armament of scattered thorns or lacking; flowers whitish c. Flowers in short, oblong spikes, comparatively large 3. A. Hayesii cc. Flowers capitate, small. d. Leaflets relatively large (usually 10 mm. long), more or less oblong, pubescent at least below; thorns frequently lacking 4. A. glomerosa dd. Leaflets small (usually A— 6 mm. long), linear or sublinear, glabrous except often ciliate marginally; thorns usually present. e. Costa of leaflet exccntric, submarginal; petlolar gland usually solitary; legume glabrous or subglabrous in age 5. A. ti-nuifolia ee. Costa of leaflet subcentral, at least apically; petiole frequently bearing 2 glands; legume tomentulose 6. A, rip a hi a bb. Armament of paired stipular spines; flowers yellowish. C. Stipular spines slender, usually whitish, solid, not ant-infested; peduncle not involucrate; in A. farncsiava older branch lets with axillary "cushions" or short-shoots, and the leaves small (scarcely 6 cm. long), with few pinnae. d. Leaves less than 6 cm. long, with 2—6 pairs of pinnae CVachellia") 7. A. farnesiana dd. Leaves usually 10 cm. long or longer, with 8—60 pairs of pinnae (^'^Popoiia.x"), Not known from Panama outside of cultivation, but to be expected there 8. A. macracantha cc. Stipular spines mostly gross, dark, hollow, myrmecophilous; pedun- cle involucrate; leaves large (usually 1 dm. long or longer), with several pinnae {"MyrmecoJcndron*^ except no. 12). d. Inflorescence capitate; leaflets with a single prominent vein; leaf rachis glandular. e. Heads axillary; petiole eglandular..... ,. 9. A. Cookii (253) [Vol. 37 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN ec. Inflorescence paniculate, the nodes not foliate; petiole bearing few to several glands 10. A. melanoceras dd. Inflorescence spicate; leaflets with 2—5 prominent veins; leaf rachis usually eglandular. c. Legume turgid-compressed, dehiscent (often tardily), short- beaked, the pericarp thick; spike thin (about 5 mm. thick), without protruding bract lets; petiolar gland of 2—3 crateri- form ncct.iries 11. A. costaricensis ee. Legume terete, indehiscent, long-beaked, the pericarp thin; spike stout (almost 1 cm, thick), with protruding bractlcts; petiolar gland with a solitary, crateriform nectary (''Tauro- ccras*'). To be expected In Panama 12. A. spadicigera L Acacia angustissima (Mill.) Kcze. Rev. Gen. PL 3^:47. 1898. Minwsa angustissimn Mill. Gard. Diet. cd. 8, no. 19, 1768. filicioiJes Cav. Icon. Pi. 1:55, pi. jS filiciTUi Willd. Sp. PI. 4:1072. 1806. 1791. Mimosa picriciua Poir. in Lam. Encycl. Mcth. Suppl. 1:74. 1810. Acacia glabrafa Schlecht. in Linnaea 12:569. 1838. Acacia anp^tdosa Bcrtol. Fl. Guat. 42. 1840. Acacia insignis Mart. & Gal. in Bull. Acad. Brux. 10-:315. 1843. Acacia jjlicioidcs Trclcasc, Rcpt, Ark. Gcol. Surv. 1888^:178. 1891. Acacia boliviaua Rusby, in Bull. N. Y. Bot. Gard. 4:348. 1907. Acaciclla angusUssiifia 'Rrkt. Sc Rose, in N. Am. FL 23:100. 1928. Acaciclla augulosa Britt. & Rose, loc. cit. 100. 1928. Acaciclla costariccfuis Britt. & Rose, loc. cit. 101. 1928. Acaciclla Retnonii Britt. & Rose, loc. cit. 101. 192 8. ?AcacielIa Holtonii Britt. & Killip, in Ann. N. Y. Acad. Sci. 35:140. 1936. Acaciclla martensis Britt. & Killip, loc. cit. 1936. Acaciclla saiifafidcrerisis Britt. & Killip, loc. cit. 1936. Acacia Vifticriana Standi. In Field Mus. Publ. Bot. 18:489. 1937. Bentliam (Trans. Linn. Soc. 30:532. 1875) further lists tbe following species as synonyms of A. angtisfissiuia: A, chlorafifba Zucc, A. cuspiJata Schlecht., A. cicgans Mart. & Gal., A. elegans Schlecht., A. Harfwcgi Benth., A. hirsufa Schlecht., A. birfa Nutt., A. pcuicillijcra L.ig., A. stipcllata Schlecht., A. tcxcusis Torr., A. utnhcllijcra Kunth. Wiggins (Contr. Dudley Herb. 3:229—233. 1942) hsts the following names as synony- mous with several subspecies of A. afrgusfissiwa: Acacia Lcnn?mnii Rose, A. birfa var. Sbrevci jfrutcscan Kearney & Peebles, A. $ujf sa Britt. & Rose, A. cilia/a Britt. & Rose, A. biria Brirt. ?: Rose, A. Lonnionii Britt. & Rose, A. sali'aJorcnsis Britt. & Rose, A. Shrciei Britt. & Rose, A. Siuitbii Britt. & Rose, A, 5;//- frutcscais Britt. & Rose. Shrub or tree, the branchJets pubescent or pubcrulent to subglabrous, not noticeably lenticellate, longitudinally striate or angled. Leaves moderately large, bipinnatc, the pinnae few to several pairs (about 15 pairs in Panamanian material cited), opposite or subopposite on the rachis, the leaflets numerous (up to 60 or more pairs per pinna); petiole up to a few cm. long, puberulcnt, cglandular, sul- cate above; rachis up to 12 or more cm. long, like the petiole; pinnular rachis 3-9 cm. long, somewhat margined, seemingly eglandular, bearing basally a pair of stipule-like leaflets; leaflets nearly linear, usually about 6 mm. long and 1 mm. wide, acute (mostly bluntly so) apically, obliquely truncate basally, glabrous except ciliate-margincd, dull above and below but darker above, the costa visible but secondary venation obscure; stipules linear-subulate, mostly 4—6 mm. long. Inflorescence a terminal or subterminal panicle of pedunculate umbels (heads). (2 -1 4) \ 19 50] FLORA OF PANAMA (Legummosae) 269 Fig. 97. Acada an^usiissima these solitary to few-fasciculate from the (usually) defoUate nodes; peduncles usually about 1 cm. long; floral bracts obovate-spatulate, little more than 1 mm. long; pedicels mostly 1-2 mm. long, borne from the terminus of the peduncle in umbellate or very short-spicate fashion. Flowers few to several per umbel, whitish; calyx campanulate or turbinate, about 1 mm. long, glabrous or sometimes sparingly ciliate, the teeth short and broad; corolla funnelform, about 2 mm. long, glabrous, the lobes divided almost to the base; stamens many, about 4 mm. long, free except at the extreme base; anthers eglandular; ovary scarcely stipltate, glabrous or minutely pubcrulent, the style slightly exceeding the stamens. Legume linear- oblong, 4-8 cm. long and about 1 cm. wide, flat and thin, stipitate, glabrous in age; the seeds few, transverse. Mexico to middle South America. CHiRiQuf: R. Chiriqui VIejo, P. Wbiie 3$, IQO. After examining Panamanian and Central American material of this species, and further noting its complicated synonymy, I must agree with Standlcy and Steyermark (Ficldiana: Bot, 24':5. 1946) that a not-unusually-variable species has been made difficult by extreme splitting. Exact specific delimitation must await (255) 270 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 the monographer, but again I concur with Standley and Stcycrmark's findings for Guatemala, In that Panamanian material won't "work" in Wiggins' separation of A. afi^itstissima from A, filkioiclcs. 2. Acacia villosa (Sw.) Willd. Sp. Pi. 4:1067. 1806. Mimosa villosa Sw. Fl, Ind. Dec 2:982. 1800. Acacia lophauthoides DC. Prodr. 2:457. 1825. Acacia carhonaria Schlecht. in Linnaca 12:571. 18 38, fide Benth. Acacia Cumingii Benth. in Hook. Lond. Jour. Bot. 1:525. 1842, fide Benth. Acacia YalcuzucUna A. Rich. Ess. Fl. Cub. 1:462. 1845. Acaciclla villosa Britt. & Rose, in N. Am. Fl. 23:104. 1928. Acaciclla Cumirigii Britt. & Rose, loc. cit. 1928. ?AcacicUa Oerstcdii Britt. & Rose, loc. cit. ex char. 1928. Shrub up to a few m. tall, unarmed, the branchlets heavily pubescent to almost glabrous. Leaves moderately large, bipinnate, the pinnae 4-10 pairs opposite on the rachis, the leaflets 9-2 5 pairs per pinna; petiole mostly about 2 cm. long, pubescent, normally cglandular, sulcate above; rachis similar to the petiole, 2-8 cm. long; pinnular rachis about 4 cm. long, pubescent, usually eglandular; leaflets broadly oblong or ovate, mostly about 8 or 9 mm. long, more or less inequilaterally obtuse apically, obliquely rounded basally, subglabrous or lightly pubescent above, appressed-pilose below, obscurely pinnately veined; stipules linear-lanceolate, 2-3 mm. long. Inflorescence of axillary, pedunculate heads, usually appearing more or less paniculate towards the tips of twigs because of insertion of peduncles at de- foliate or imniaturely foliate nodes; peduncles about 2 cm. long, pubescent; head compact, subglobose or very short-spicatc, about 3 0-flowered. Flowers small, briefly pedicellate; calyx broadly cupulate, about 1 mm. long, shallcwly 5-toothed, glabrous, sometimes ciliate; corolla funnelform-campanulate, 2-3 mm. long, gla- brous except sometimes pubescent apically, deeply 5-parted; stamens numerous, about 5 mm. long. Legume oblong, 5-6 cm. long and almost 1 cm. wide, flat and thin, tapered basally, stipitate, lightly pubescent, the seeds transverse. Southern Central America and West Indies. According to Rrltton and Rose, this species Is confined to Jamaica, while A. Cufuingii is recognized as a distinct species known only from the Panamanian type. We have been unable to examine type material of either of these species, or, for that matter, of most of the synonyms listed. However, Bentham, author of A, Ctuniugii, with the accumulation of more material, later reduced his species to synonymy with A. villosa (Trans. Linn. Soc. 30:532. 1875). There seems to be no reason for not accepting Bcntham's judgment, at least without reference to type material. Bentham's type sheet of A. Cumuigii (Cnming 1242) thus becomes the record on which we base inclusion of A. villosa in Panama. No additional collections from there have come to our attention. Necessarily, the description above given is a composite from several sources and does not reflect any given specimen. Acacia villosa somewhat resembles A. angnstissiway but can be distinguished from It in having significantly broader and usually larger, more pubescent leaflets. (256) 1950] FLORA OF PANAMA (Leguminosac) 271 3. Acacia Hayesii Benth. in Trans. Linn. Soc. 30:524. 1875. Sencgalia Hayesii Britt. & Rose, in N. Am. Fl. 23:114. 1928. Sencgalia acanthophylla Britt. & Rose, loc. cit. 117. 1928. Acacia tclonis Standi, in Field Mus. Pub!. Bot. 4:308. 1929, Acacia acanthophylla Standi, loc. cit. 18:488. 1937. Liana or scandcnt shrub, armed with small, scattered, recurved thorns, the branchlets puberulcnt when young becoming glabrous. Leaves large, bipinnate, the pinnae usually about 10 pairs opposite on the rachis, the leaflets nearly 20 pairs per pinna; petiole 2 or more cm. long, pubcrulent above, bearing 1 or 2 cupulate, sessile glands; rachis up to 20 or more cm. long, similar to the petiole, bearing cupulate glands just below insertion of all or most pairs of pinnae; pinnular rachis 4-5 cm. long, presumably puberulcnt and with occasional small glands at the distal rachial nodes; leaflets linear-oblong, 8-10 mm. long, somewhat falcate, briefly mucronatc apically, obliquely subtruncate basally, reportedly glabrous in a frag- ment of the type but apprcssed-pubcscent below, the costa markedly excentric; stipules obovate? Inflorescence of 1 to few pedunculate spikes from subterminal, defoliate or minutely foliate nodes, thus appearing paniculate early; peduncles 1-2 cm. long, puberulcnt; spikes 10—12 mm. long, dense. Flowers subsessile, presum- ably whitish; calyx cupulate, about 2 mm. long and equally as broad, glabrous, moderately lobed; corolla cylindric-campanulate, about 4 mm. long, glabrous, rather briefly lobed; stamens many, 7—8 mm, long. Legume linear-oblong, flat, thin, stipitate, glabrous. Panama, Costa Rica, Honduras. canal zone: Mammee railroad station, S. Hayes s, n. Examination of a photograph and a fragment of the type shows that specimens referable to the Fionduran A. felcnsis and the Costa Rican Sencgalia acanthopbylla constitute a second record for A. Hayesii^ previously known only from Panama. 4. Acacia glomercsa Benth. in Flook. Lond. Jour. Bot, 1:521. 1842. Sencgalia glomcrosa Britt. & Rose, in N. Am. Fl. 23:116. 1928. Usually a moderate to large tree, often buttressed, unarmed or sparingly armed witli a scattering of recurved prickles on twigs and rachis, the branchlets pubescent when young becoming glabrous with age. Leaves quite large, bipinnate, the pinnae several to many pairs, the leaflets many (about 30) pairs per pinna; petiole 2-4 cm. long, tomentulose to subglabrous, flattened or sulcata above, bearing 1 (—2?) sessile gland usually below the middle; rachis up to 20 cm. long, similar to the petiole, sometimes aculeate below, usually bearing sessile glands above just below the distal few rachial nodes; pinnular rachis up to 10 cm. long, pubescent, gland- ular at the distal foliolar nodes; leaflets narrowly oblong or linear-oblong, mostly about 10 mm. long and 2-3 mm, wide, asymmetrically obtuse or acutish apically and thus subfalcate, obliquely truncate basally, puberulent or subglabrous above, appressed-pubescent below, the costa markedly excentric but not marginal, other (257) [Vol. 37 272 ANNALS OF THE MISSOURI BOTANICAL GARDEN venation visible but obscure; stipules linear, caducous. Inflorescence a terminal panicle of pedunculate heads; peduncles 5-15 mm. long, tomentulose; heads com- pact, globose, 12- to 20-flowered, the bracts minute. Flowers small, cream or white; calyx cupulate, about 1 mm. long, puberulent; corolla funnelform, 2—3 mm. long, pubescent, the lobes about "^^ as long as the tube; stamens many, mostly 5-6 mm. long. Legume linear-oblong, up to 20 cm. long and 3 cm. wide, roundcd- subcuncate apically and basally, flat, puberulent becoming subglabrous In age, the margins slightly elevated, the seeds few, transverse. Mexico to middle South America. CANAL zone: Barro Colorado Island, Aiilcs lob. This species is quite variable, and in its extended range exhibits a number of diverse forms. Apparently it is commonly found as an erect tree, but sometimes It assumes somewhat the aspect of a liana. Most specimens are unarmed, but not a few bear scattered, recurved prickles, nor Is this characteristic always associated with the subscandent habit. The Panamanian specimen cited possesses a longer pubescence than Is typical for the species either In South America or north of Panama. It is also "atypical" in being armed and having more than the usual num- ber of pinnae. Yet It can probably be considered within the specific boundaries of A. glovicrosa despite uncertainties and perhaps intermcdiacy towards A. tenui- folr.r. The plant is evidently uncommon in Panama. Our specimen (and others passing as the species) could almost as well be called A. polyp/jyila DC. as A, glotfiewsa. In fact, considered over their entire range. It is doubtful that A. glonicrosa can be maintained as a species distinct from the older A, polyphylla. 5. Acacia tenuifolia (L.) Willd. Sp. Pi. 4:1091. 1806. Mimosa tcniujolia L. Sp. PI. 523. 1753. Acacia pauiculata Willd. Sp. Pi. 4:1074. 1806. Mimosa grafiJ'isHiqua Veil. Fl. Flum. Ic. 11:/. J/. 1827, fide Benth. Acacia graudisiliqua Bcnth. in Hook. Lend. Jour. Bot. 1:518. 1842. Acacia Claimeni Benth., loc. cit. 1842, fide Benth. Acacia martiniccnm Presl, Bot. Bcmerk. 65. 1844, fide Britt. & Rose. Acacia microccphala A. Rich. Ess. Fl. Cub. 1:469. 1S45, fide Brirt. & Rose. Scncgalia tenuifolia Britt. & Rose, in N. Am. Fl. 23:118. 1928. Tall liana, climbing through tops of trees, the branchlets puberulent or minute- ly tomentose, armed with broad-based, recurved thorns. Leaves large, bipinnate, the pinnae 20 or more pairs opposite on the rachis, leaflets numerous (about 70 pairs) ; petiole up to a few cm. long, terete, lightly tomentulose or puberulent, armed, bearing about 1 cm. from the base a sessile, longitudinally oblong gland; rachis about 20 cm. long, flattened or subsulcate and puberulent or tomentose above, armed with thorns, bearing at the uppermost several rachial nodes prominent orbicular glands; pinnular rachis about 7 cm. long, glabrous below, tomentose above, eglandular; leaflets asymmetrically hnear, about 4 mm. long and less than 1 mm. wide, broadly and inequilatcrally acute apically, obliquely truncate basally, (258) 1950] FLORA OF PANAMA (Lcguminosae) 273 glabrous except ciliate-margincd, the costa excentric, secondary veins prominent below but not above, dull; stipules caducous. Inflorescence paniculate, up to 3 or more dm. long, the primary axis armed, tomentulose; ultimate divisions of 1 to few pedunculate heads from nodes on which leaves have not yet developed; peduncles mostly 10—15 mm. long, tomentulose; heads several-flowered, globose, the floral bracts minute. Flowers dense, white; calyx funnclform-cupulate, about 1 mm. long, puberulent ; corolla f unnclf orm, scarcely 2 mm. long, minutely puberulent, deeply lobed; stamens many, 3—4 mm. long, glabrous, free except at the extreme base; anthers mostly eglandular (some apparently abortively glandular in bud); ovary pubescent, long-stipitate. Legume presumably linear-oblong, flat, thin, non-septate, glabrous in age. Mexico to middle South America; West Indies. dari^n: R. Chico, Allcfi 4646; El Real, Allen 961. san blas: Puerto Obaldia, Piitier 4326. Britton and Rose (N. Am. Fl. 23) and various other authors have not recog- nized the synonymy of the South American A. paniculata and several Central American and Mexican "species" which probably represent the Linnean name A. tcnuifoUa, Yet a comparison of a series of Mexican, Central American, and West Indian specimens with South American material shows remarkably little difference. I have thus accepted Bentham's judgment that A. pantcnlata and A. tcnuifoUa arc the same, and constitute merely one, not overly variable, species. 6. Acacia riparia HBK. Nov, Gen. & Sp. 6:276. 1824. }Mhi70Sa rctusa Jacq. Enum. PL Carib. 54. 1762, fide Ind. Kew. Mimosa pauicidata West, Bidr. til Beskr. Ste.-Croix, 312. 1793; non authors, fide Ind. Kew. Acacia Westiana DC. Prodr. 2:464. 1825, fide Benth. Acacia guatialupensis DC. loc. cit. 1825. }Mimosa plana Veil. Fl. Flum. Ic. 11:/. 2c?. 1827, fide Benth. Acacia quadnglandiilosa Mart, in Flora 20, Beibl. 2:110. 183 8, fide Ind. Kew. Acacia tubuUfcra Benth. in Hook, Lond. Jour. Bot. 1:520. 1842, fide Macbride. Acacia sarmcntosa Griseb. Fl. Brit. W. Ind. 221. 1861; non Desf., fide Ind. Kew. Senegalia riparia Britt. & Rose, In Ann, N. Y. Acad. Sci. 35:144. 1936. Shrub or tree (or often subscandent) , sparingly aculeate on leaf rachis and twigs, with small, scattered, recurved thorns, the branchlets puberulent soon be- coming glabrous, striate. Leaves moderately large, bipinnate, the pinnae 6-20 pairs opposite on the rachis, the leaflets many (mostly 20-40) pairs per pinna; petiole about 2 cm. long, sulcate above, subglabrous, bearing on the upper side 1 or more (usually 2) sessile, patcUiform, longitudinally oblong glands; rachis up to 15 cm. long, sulcate and puberulent above, bearing glands like those of the petiole just below the several distal rachial nodes; pinnae usually 4-5 cm. long, eglandular; leaflets lincar-oblong, 3-7 (usually 4-5) mm. long and about 1 mm. wide, more or less rounded to broadly acute apically, obliquely rounded or subtruncate basally, glabrous above and below, usually ciliate marginally and barbate at the base on the inner margin below, the costa subccntral except at the inequilateral base, it and a (259) 274 [Vol. Z7 MISSOURI few remote lateral veins prominent below, dull; stipules caducous or lacking. In- florescence seemingly a raceme or panicle of pedunculate heads, these 1- to few- fasciculate from subterminal defoliate or tardily foliate nodes (nodes foliate in ngc or with fruit) ; peduncle 1-2 cm. long at anthesis, tomentulose; head dense, sub- globose, about 30-flowered, Flowers sessile, whitish; calyx cupular, 1 mm. long, pubcrulcnt; corolla funnelform, 3-4 mm. long, puberulent apically, the lobes brief; stamens many, free, about 6 mm. long. Legume linear-oblong, usually 10-15 cm. long and 20-25 mm. wide, stipitate, tomentulose, usually sharp-beaked apically, flat and thin, the margins slightly elevated, the seeds transverse. Middle South America to southern Central America; perhaps northern Central America and southern Mexico? CANAL zone: Barro Colorado Island, Allies 14; Brown 14S; Shatfnck 436; Wctmorc 6 Abbe 205; Woodwortb ^ Vestal 512. CHiRiQuf: San Felix, ?itticr 5141, Possibly the name Acacia retusa should apply to this species, but inasmuch as none of the types are available and Bentham himself had only ''ex char." acquaint- anceship with Mimosa retusa Jacq,, which he listed as synonymous, we do not sug- gest any change in the familiar name, A. riparia. The species seems to grade into what commonly passes for A, riparioiJes in northern Central America and Mexico, differing from it primarily in having a somewhat longer corolla. The Chiriqui specimen is somewhat atypical, having larger flowers approaching those of A. Haycsii, 7. Acacia farnesiana (L.) Willd. Sp. Pi. 4:1083. 1806. Mimosa farnesiana L. Sp. Pi. 521. 1753. Acacia acicularis W'illd. Enum. 1056. 1809. YacbclUa farnesiana Wi^ht & Arn. Prodr. Fl. Penins. Tnd. Or. 272. 18 34. Listed by Bentham (Trans. Linn. Soc. 30:502. 1875) as additional synonyms arc: Acacia eJulis H. & B., A. lenticeUata F. Mucll., A. leptopbylla DC, A. peduuculata \v'i!lJ., Farnesia oJora Caspar., Mimosa peduuculata Poir., Al. scorpioides Forsk. Shrubs or occasionally small trees, the branchlets frequently somewhat pilose when young becoming glabrous, prominently lenticellate, armed with slender, solid (but having a soft pith), whitish, stipular spines up to about 3 cm. long, these not ant-infested. Leaves moderately small, bipinnate, the pinnae 2-6 pairs opposite on the rachis, the leaflets mostly 10-20 pairs per pinna; petiole up to 1 cm. long, terete except somewhat flattened above, usually tomentulose-subhirsute, bear- ing a sessile, oblong gland above the middle; rachis 1-5 cm. long, similar to the petiole, usually bearing a gland at insertion of the terminal pair of pinnae; pinnular rachis 1-3 cm. long, flattened above, often bearing a minute gland just below insertion of the terminal pair of leaflets; leaflets narrowly oblong, about 4 mm. long and 1 mm. wide, rounded apically, obhquely rounded or truncate basally, glabrous except frequently ciliate marginally, the costa prominent but other venation usually obscure. Inflorescence of axillary, pedunculate heads, these 1 to few from foliate nodes; peduncles mostly 1-2.5 cm. long, pubescent; heads dense, (260) 1950] FLORA OF PANAMA (Legumhiosae) 275 globular; floral bracts narrowly oblanceolate, tipped white-pilose. Flowers small, yellow, sessile, fragrant; calyx funnclform-cupulate, a'bout 1 mm. long, glabrous except on the shallow teeth; corolla narrowly funnelform, a'bout 2 mm. long, gla- brous except on tips of the lobes, the lobes relatively short; stamens many, little more than 3 mm. long, free except at the extreme base. Legume linear-oblongoid, up to 6 or 7 cm. long and almost 1 cm. wide, turgid-subtcrete, glabrous, longi- tudinally conflucnt-striatc, tardily dehiscent, coriaceous, with a sweet pulp, pithy in age, septate, the seeds transverse. Southern United States and Mexico to Argentina; West Indies; introduced into many parts of the Old World. CANAL zone: Balboa, Sfandley ^0841; Las Cruces Trail, Hunter G? Allen 6S2; Pitiier 2621. cocLE: Penonome, Williams 1 36. Panama: R, Jagua, Hunter ^ Allen 4/(); Juan Diaz, Standley 3^533^ Nucvo San Francisco, StanJley 3^7^3' This abundant species is of some economic importance. Leaves and pods serve as food for stock, while the heavy wood is used for fuel. The bark and legume are a source of tannin. An exudate from the trunk is reported useful as a mucilage. In southern France especially the species has been extensively cultivated for the perfume industry. Acacia farncsiana is not clearly distinct from what commonly passes as another Linnean species in Central and northern South America, A, tortuosa, "Without access to authentic material of these two species, no attempt is made to pass judg- ment on them. The pod of A. tortnosa is typically pubescent, while that of A, farucsiana is glabrous and shorter. Yet intermediacy in this character is found, as well as In other characteristics which are quite different in their extremes. The unsnarling of these few related species will require more critical study than Is possible In this Flora. 8. Willd, Sd. PL 4:1080. 1806 Acacia macracantboides Bert. In DC. Prodr. 2:463. 1825. Poponax macracantboides Brltt. & Rose, in N. Am. Fl. 23:89. 1928. For multitudinous additional synonyms see Benth. Trans. Linn. Soc. 30:500. 1875. Small tree, the branchlets armed with small to large, solid, paired stipular spines, in youth pubescent to subglabrous, later subglabrous and prominently lenticellate. Leaves moderately large, bipinnate, the pinnae 8— (reportedly) 60 pairs opposite on the rachis, the leaflets many. Petiole about 1 cm. long, pubescent, canaliculate above, bearing a longitudinally extended, crateriform gland slightly above the middle; rachis similar, gland-bearing only at terminal 1 to few rachial nodes; pinnae 2—3 cm. long, eglandular; leaflets linear or linear-oblong, mostly about 3 mm. long and scarcely 1 mm. wide, rounded or obtuse apically, obliquely rounded to subtruncate basally, glabrous or lightly pubescent, only the costa prominent; stipules modified as spines. Inflorescence of axillary, pedunculate heads, solitary or few-fasciculate; peduncles about 1 cm. long, tomentose, ebracteate (except at head) ; head orbicular, dense, less than 1 cm. in diameter; floral bracts linear- (261) [Vol. Z7 276 ANNALS OF THE MISSOURI BOTANICAL GARDEN spatulatc, about 1 mm, long. Flowers small; calyx funnclform, about 1 mm. long, tomcntulose aplcally; corolla tubular-funnelform, about 2 mm. long, glabrous except on rbc prominent lobes; stamens many, free, about 3 mm. long. Legume variable, Hncai, reported up to 10 cm. long and 12 mm. wide, subterete, straight or somewhat curved, usually puberulent and glandular. Northern South America and West Indies; in a broad sense, possibly also northern Central America. CANAL zone: Summit, Lindsay 24g (presumably cultivated: at Canal Zone Experi- ment Gardens) . This very variable species is evidently not known in tbe wild from Panama, but occurs in Colombia and other portions of northern South America and the West Indies. It somewhat resembles A. farnesiana, these two species alone (in Panama) possessing solid stipular spines, A. ntacracafitha is readily distinguished from A. farucsiana by the larger, multi-plnnular leaves. 9, Acacia Cookii Safford, in Science n. s. 31:677. 1910, Acacia hnccrophera B. L. Robins., in Proc. Am. Acad. 49:502. 1913. Myrmccodcndron Cookii Britt. & Rose, in N. Am. Fl. 23:93. 1928. Shrub or tree, the branchlcts puberulent soon becoming glabrous and heavily lenticcllate, armed with paired stipular spines, many of these becoming very large, blackish, hollow, ant-infested. Leaves large, bipinnate, the pinnae about 15 pairs, the leaflets about 30 pairs per pinna; petiole short, puberulent at least above, eglandular, sulcate above; rachis 8-15 cm. long, similar to the petiole, bearing cupulate glands just below Insertion of all or most pairs of pinnae; pinnular rachis about 3 cm. long, tomcntulose, eglandular; leaflets essentially linear, in the Panamanian specimen 3-4 mm. long and less than 1 mm. wide (usually about 6 mm. long for the species), asymmetric and bluntly acute apically, obliquely trun- cate basally, glabrous except ciliate-marglned, the venation obscure; stipules modi- fied as spines, these when enlarged (ant-infested) briefly connate at the base and resembling bull horns. Inflorescence of axillary pedunculate heads; peduncles short, clustered, bearing a 3 -parted involucre near the middle; head very compact, globose. Flowers small, yellowish; calyx cupular, about 1 mm. long, glabrous; corolla funnelform, barely exceeding the calyx, glabrous except puberulent on tips of the lobes; stamens numerous, about 2 mm. long. Legume reported linear, 10-30 cm. long, somewhat compressed, usually curved, subglabrous, sessile, beaked aplcally, 2-valvate. Guatemala, British Honduras, Honduras; on basis of cited specimen, Panama. BOCAS DEL TORo: Changuinola Valley, DunJap 462, The Panama specimen cited is very poor, and its determination cannot be en- tirely certain. It appears to match (descriptions and specimens of) A. CooJui better than A. welanoceras^ tbe only other described species it might likely be. Our description is of the Panama plant insofar as possible (vegetative characters). The species is a member of the distinctive "bull-horn" group of Acacias. (262) 1950] FLORA OF PANAMA (Legiimlnosae) 277 10. Acacia melanoceras Beurl. in Sv. Vet. Akad. Handl. 1854:123. 1856. Acacia multiglandulosa Schenck, in Fedde Rep. Sp. Nov. 12:362. 1913, fide Britt. & Rose. Myrmecodendron melanoceras Britt. & Rose, in N. Am. Fl. 23:93. 1928. Shrub or sometimes small tree, tKe branchlets pubcrulent when young, armed with large, terete, hollow, myrmecophilous, stipular spines, these briefly connate at the base and resembling bull horns. Leaves large, bipinnate, the pinnae normally at least 15 pairs, the leaflets 15—30 pairs per pinna; petiole 1-2 cm. long, sulcatc above, usually puberulent, bearing on the upper side a number of conic glands in one or more rows; rachis similar, as much as 4 dm. long, bearing a conic gland just below insertion of most pairs of pinnae; pinnular rachis mostly 4-5 cm, long, if glandular obscurely so; leaflets linear, about 5—6 mm. long and scarcely more than 1 mm. wide, Inequilaterally obtuse aplcally, obliquely rounded basally, glabrous, dull, with a single prominent vein; stipules modified as large, lustrous, brown or black spines as much as 4 or 5 cm. long. Inflorescence a terminal, raceme-like grouping of pedunculate heads, these 1 to few from the axils of defoliate or minutely foliate nodes; peduncles short, 2—12 mm. long, bearing near the base a 3- to 4-partIte Involucre; heads globular, densely flowered, the floral bracts peltate, about equalling the flowers. Flowers minute, yellow; calyx cupular, about 1 mm. long, glabrous or puberulent aplcally, obscurely lobed; corolla funnelform, little more than 1 mm. long, puberulent aplcally, more prominently lobed; stamens many, 2-3 mm long. Legume linear-oblong, up to 10 cm. long, slightly com- presscd-subtcrete, short-beaked, longitudinally striate, glabrous, tardily dehiscent. Panama. CANAL zone: Barro Colorado Island, Aviles 2ob; Kenoyer 3/ J; France Field, Fairchild Aug, 1924; Shropshire, Feb, ip4S' The species Is known only from Panama (types = Billhcrg 2(?9, 18 2§, from Portobello, Colon). Possibly one or two species of northern Central America, such as A. Donnelliana, A. Cookii and A. glohulifera, may prove to be the same entity; but Insufficient collections are available to pass judgment on this question. 11. Acacia costaricensis Schenck, In Fedde Rep. Sp. Nov. 12:361. 1913. Acacia panamefjsis Schenck, loc. cit. 362. 1913, fide Britt. & Rose. Acacia penonomensis Safford, In Jour. Wash. Acad. 4:363. 1914, fide Britt. & Rose. Acacia Nelsonii Safford, loc. cit. 1914, fide Britt. & Rose. Myrmecodendron costariccnse Britt. & Rose, in N. Am. Fl. 23:93. 1928. Shrub or small tree, the branchlets glabrous and laxly lenticellatc, armed with large, hollow, myrmecophilous stipular spines shaped like the horns of a bull. Leaves large, bipinnate, the pinnae 4-12 pairs opposite on the rachis, the leaflets 13—24 pairs; petiole almost 2 cm. long, glabrous, flattened or sulcate above, bearing on the upper side below the middle usually 2—3 conic glands; rachis up to 15 cm. long, similar to the petiole, usually eglandular; pinnae somewhat lax, glabrous, eglandular, the axis margined; leaflets linear, usually 8-9 mm. long and only slight- ly more than 1 mm. wide, Inequilaterally rounded or obtuse aplcally, obliquely (263) [Vol, 37 278 ANNALS OF THE AIISSOURI BOTANICAL GARDEN Fig. 98. Acaciii cosfariccftsis rounded basally, glabrous and dull above and below, with 2-3 prominent palmate veins from the base; stipules large, terete spines frequently 3-4 cm. long, those of a pair fused at the base, lustrous, brownish. Inflorescence of condensed, peduncu- late, axillary spikes about 3 or 4 cm. long; peduncle usually about 1 cm. long, bearing a tripartite involucre near the base; floriferous portion elongate, ament- likc, becoming thicker in fruit; bractlets peltate, equalling the flowers. Flowers minute, very dense on the spike, yellowish; calyx cupulatc, less than 1 mm. long, glabrous (or minutely puberulent marginally), obscurely lobed; corolla funnel- form, little more than 1 mm. long; stamens many, about 2 mm. long. Legume narrowly oblong, mostly 4-5 cm. long, bluntly and briefly beaked apically, sessile, slightly compressed-subterete, glabrous but more or less scurfy, longitudinally striate, pulpy, reportedly 2-valvate but apparently tardily dehiscent, the seeds obliquely longitudinal. Mexico to Colombia. CANAL zone: Chivi-Chivi trail, Maxou ^ CHiRiQuf: Puerto Rcmcdios, Piffier 3378, PANAMA: R. Abajo, Bro. Paul 267; Bella Vista, Piiticr 6785; Staudlcy 25374; Macbridc ^758 y Nuevo San Francisco, Sfandley 3^737^ This species, a member of the interesting "bull-horn" group very prominent in northern Central America and Mexico, runs into A, Collhisli (^ A. yncata^icinis) and possibly A. hirf/pcs and A. Himlsi/ of that region. Three or four species of Harvey 657Q; Miraflores, Allen 1743. Penonomc, Williams II 3 (photo) . (264) 1950] FLORA OF PANAMA (Legiunlnosac) 279 Acacia in Panama have the large, hollow, ant-infested stipular thorns the basis for such popular names as "cockspur," casa de hormigas^ cormzucloy etc. Of these A. cosfariceusis can be readily distinguished by the spicate inflorescence. 12. Acacia spadicigera Schlecht. & Cham, in Linnaea 5:594. 183 0. Acacia nicoyensis Schenck, in Fedde Rep. Sp. Nov. 12:360. 1913. Acacia ciibensis Schenck, loc. cit. 1913, fide Britt. & Rose. Acacia furcella Safford, in Jour. Wash. Acad. 4:359. 1914, fide Britt. & Rose. Acacia Hcrnauclezii Safford, loc. clt. 3 5 8. 1914, fide Britt. & Rose. Tanroceras spaJicigerum Britt. & Rose, in N. Am. Fl. 23:85. 1928. Shrub or small tree, the branchlets glabrous, armed with large, hollow, grayish, paired, myrmecophilous stipular spines connate at the base and resembling bull horns. Leaves moderate, bipinnate, the pinnae mostly 2-8 pairs, the leaflets many (12-3 or more) pairs per pinna; petiole about 1 cm, long, glabrous, expanded above into a large, longitudinally oblong, glandular area bearing a solitary, ex- tended conic ^'nectary"; rachis a few cm. long, glabrous, sulcate above, sometimes bearing an orbicular gland at insertion of the lowermost pair of pinnae; pinnae up to 7 cm. long, glabrous; leaflets linear, 5-10 mm. long and almost 2 mm. wide, rounded-mucronulate apically, obliquely truncate basally, glabrous, both costa and secondary veins noticeable; stipules modified as large, myrmecophilous spines as much as 8 cm. long. Inflorescence of axillary, pedunculate spikes; peduncles very short, usually about 3 mm. long, glabrous, bearing below the middle a 4- partite involucre; spikes oblong, mostly about 3 cm, long and almost 1 cm. wide, gross, spadix-likc, the tips of the bractlets protruding. Flowers very small, very condensed, presumably yellow; calyx tubular-truncate, about 1 mm. long, glabrous; corolla included within calyx; stamens scarcely 2 mm. long. Legume oblong, about 8 cm, long (including beak) and about 13 mm. in diam., stipitate, long- beaked (the beak about 3 cm. long), terete, indehiscent, pulpy, the pericarp thin, the seeds transverse. Mexico, northern Central America, Costa Rica; Panama? This species is reported by modern authors from Costa Rica but not from Panama. Bentham incorrectly listed Cuming 12^0 from Panama as A. ^[mdicigeYa (Trans. Linn. Soc. 30:514), the specimen actually being A, costaricensis. Never- theless, there remains a distinct possibility that the species docs occur and will eventually be collected in Panama. As with a number of pulpy legumes, the fruit of this species is commonly eaten by local peoples in Central America. SPECIES DOUBTFULLY RECORDED FROM PANAMA Acacia Hindsii Bcnth. (Fiook. Lond. Jour. Bot. 1:504. 1842) has been reported from Panam,a, apparently in error. A specimen so determined in the herbarium of the Chicago Museum of Natural History proved to be A. cosfaricensiSy and no additional collections from Panama of A. Hiudsii have come to my attention in (265) [Vol. 37 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN any oi the larger United States herbaria. This species is characterized by having the legume reputedly dehiscent only along the lower suture, being the only species among the '*bull-horn" Acacias having this characteristic. 7. MIMOSA L. Mimosa L. Sp. Pi. 516. 1753. ?Panthocarpa Raf. Neogen. 2. 1825, fide Dalla Torre & Harms. Eburnax Raf, New Fl. N. Amer. 1:42. 1836, in part- Lomoplis Raf. Sylva Tell. 118. 1838. Scnsitiva Raf. loc. cit. 119. 183 8. Pteromimosa Britt., in N. Am. Fl. 23:171. 1928. Neomimosa Britt. & Rose, loc. cIt. 172. 1928. Mimosopsis Britt. & Rose, loc. cit. 174. 1928. Acanthopteron Britt., loc. cit. 179. 1928. llaitimimosa Britt., loc. cit. 179. 192 8. Coarse herbs or shrubs (in Panama), usually sprawling, decumbent to sub- scnndcnt, heavily or lightly armed (in all Panamanian species except sometimes M. pusilla) with recurved thorns, variously pubescent. Leaves often sensitive (re- sponding visibly to touch or other stimuli), bipinnate, the pinnae 1 to many pairs, opposite, the leaflets 2 to many pairs per pinna; petioles pulvinate, eglandular (in Panama), usually aculeate; rachis eglandular but often spinose- or subulate- appendaged; pinnae usually short, pulvinate, the pulvinus commonly setose; leaf- lets large and few or more commonly small and many, inequilateral basally, if pubescent usually strigosc; stipules narrow, subpcrslstcnt. Inflorescence of pedunc- ulate heads (in all Panamanian species) or spikes, these axillary from lower foliate and/or subterminal non-foliate nodes; floral bractlets usually shorter than the corolla, subpersistent. Flowers pinkish or whitish, normally sessile, small, numerous, usually perfect; calyx commonly very minute; corolla several times exceeding the calyx, usually funnelform, 3-6 (mostly 4) -lobed, the lobes equalling or shorter than the tube, valvate; stamens as many as or twice as many as the corolla lobes, free, exserted, the small anthers eglandular; ovary generally short, few- to many- ovulate, the style about equalling the stamens, the stigma not expanded. Legume linear to broadly oblong, flat, thin, with persistent margins, usually (as with all Panamanian species) transversely articulate, commonly setose-spinose. Primarily western hemisphere from United States to Argentina, especially abundant in tropics and subtropics; a few species are known from Africa and Asia, On the whole, the genus presents a rather natural grouping, grading, however, into certain (generally recognized) segregates such as Schrankia, Many of the species of Mimosa seem constantly distinctive, while others have evolved into a number of indistinct forms which are usually graced with (doubtfully tenable) specific names. As a result considerable condensation is in order, a common con- dition among most genera of the MiMOSomEAE. In addition to Bentham's original classical work on the Leguminosae, B. L. Robinson presented a "Revision of the North American and Mexican species of (266) 1950] FLORA OF PANAMA (Leguminosac) 281 Mimosa'* (Proc. Am. Acad. 33:305. 1898). Britton and Rose in the 'North American Flora' list about 150 species of Mi?nosa north of South America, of which 9 have been collected in Panama. A number of additional species are known from neighboring regions, some of which well may be collected in Panama eventually. Listed from Costa Rica are: M. alb'ula^ M, Calderom, M. dormiens^ M. flavcscertSy and M. Watsonii. Listed from Colombia by Britton and Killip (Ann. N. Y. Acad, Sci. 3 5:148-154. 193 6) are: M. albida, M. AnJreana, M. colombiana, M. debilis, M. dormienSy M. esmeraldae, M. flavescens, M. floribtifiJa, M. pramenensis^ M. Killifyiiy M. Lawranceana, M. leiocarpa, M. Martensis, M. oligacatitha, M. polycarpa, M. cpiitensisy M. santanderensisy M. spiciflora, M. temiifloruy and M. TrianaCy a number of them recognized species of long standing. a. Pinnae a single pair per leaf. b. Leaflets 10—40 mm. long, 4—15 mm. wide, 2—6 pairs per pinna; legume 4—12 mm. wide, c. Leaflets only 2 pairs per pinna, mostly 3-4 cm. long. d. Leaflets 10—15 mm. wide, ovate to elliptic (broadest below the middle); legume mostly 6—10 mm. wide; head more robust. e. Twigs glabrous or nearly so, copiously armed; leaflets spar- ingly if at all strigose; floral bractlcts prominent, exceeding and obscuring the subglabrous buds 1. M. Velloziana ee. Twigs usually pubescent, sparingly aculeate; leaflets densely strigose; floral bractlets no longer than and scarcely visible among the puberulent buds 2. M. albida dd. Leaflets mostly 4—10 mm. wide, more or less obovate (broadest above the middle); legume about 4 mm. wide; head less robust.. 3. M. panamensis cc. Leaflets 3 or more pairs per pinna, mostly 1—3 cm. long 4. M. casta bb. Leaflets 3—12 mm, long, 1-2 mm, wide, about 16 pairs per pinna; legume about 4 mm. wide 5. M. pudica aa. Pinnae 2 or more pairs per leaf, at least on some or most leaves. b. Rachls very condensed or obsolete, the pinnae thus subdigitate; sta- mens as many as the corolla lobes. c. Pinnae no more than 2 pairs; peduncle about 2 cm. long in maturity 5. M. PUOICA cc. Pinnae 3-5 pairs; peduncles short, about 1 cm. long 6. M. polydactyla bb. Rachis approximating or exceeding the petiole, the pinnae compara- tively remote and quite obviously not subdigitate; stamens twice as many as the corolla lobes. c. Slender plants, with shorter thorns and softer pubescence; leaflets with 1 main vein or costa; pinnae usually 2-5 (1-8) pairs; rachis approximating the petiole, setiform-appendaged. d. Corolla membranous, not striate; pubescence not glandular; twigs strongly angled or terete. e. Twigs terete or nearly so, scarcely if at all aculeate; pinnae 1—3 pairs; secondary veins of leaflet conspicuous; heads some- what obconic, conspicuously bracteate 7. M. pusilla ee. Twigs strongly angled or tetragonal, heavily aculeate; pinnae usually 4—5 pairs; secondary veins of leaflet obscure; heads nearly orbicular, inconspicuously bracteate 8. M. invisa dd. Corolla striate; pubescence almost always glandular-tipped; twigs subtcrcte : , 9. M. sOMNiANs cc. Coarse plant, formidably armed and setose-hispid; leaflets longi- tudinally few-veined; pinnae usually at least 10 pairs; rachis much exceeding the petiole, spinose at insertion of the pinnae 10. M. pigra (267) [Vol. Ti"? 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1. Mimosa Velloziana Mart, Herb, FL Bras, 18 5. 18 37. Mimosa iha Veil. FL Flum. Ic. 11:/. JJ. 1827, not L. Woody, vine-like shrub, scandcnt or decumbent, large or small, tbc branchlets subteretc, glabrous or subglabrous, copiously armed with recurved thorns. Leaves reportedly sensitive, moderate, bipinnate, the pinnae a single pair, the leaflets only 2 pairs per pinna; petiole up to 5 or 6 cm. long, subglabrous, eglandular, armed with recurved thorns, conspicuously pulvinate basally, apiculate terminally; pin- nular axis 1-2 cm. long, appendagcd above the pulvinus, scarcely if at all aculeate; leaflets asymmetrically ovate-lanceolate, mostly 3-4 cm. long and 10-15 mm. wide, the inner member of the lower pair very reduced, acute-subacuminatc apical- ly, obliquely rounded basally, glabrous and darker above, sparingly setose-strigose or subglabrous and lighter below, setose-ciliate on the margins; stipules ovate, about 3 mm. long, ciliate. Inflorescence of axillary, pedunculate heads, often few- fasciculate; peduncles mostly 1-2 cm. long, unarmed, subglabrous; heads ovoid, compact, multiflorate; floral bractlets linear-lanceolate or hncar-oblanceolatc, setose-fimbriate, about 2 mm. long. Flowers small, sessile, reportedly pinkish; calyx exceedingly minute, fimbriate-lobed; corolla funnelform, about 2 mm. long, subglabrous, the 4 lobes less than Yz the length of the tube; stamens 4, 7-8 mm. long, the anthers orbicular; ovary setose. Legume oblong, 1-3 cm. long and 8-10 mm. wide, flat, segmented, densely sctose-spinose on the persistent margins but little if at all so on the segments, otherwise glabrous. Southern Mexico through Central and South America to Paraguay. BOCAs DEL TORO: Changuinola Valley, Dunlap 2gj, canal zone: "Chagrcs", Fcndlcr pi; 'Tanama", Sccmann lOO, The species usually occurs in clearings, and fast becomes a pest, forming nearly impenetrable tangles which tear clothing and flesh. It is reported sensitive to touch, contracting the leaves when disturbed. Mimosa Velloziana has consistently been regarded as a distinct and tenable species, in spite of the fact that it docs not always exhibit hard and fast demarca- tion from certain species of greater antiquity. Remarkably like it, but usually without prominent, appressed, pectinate floral bractlets are M. scnsifiia L. of Brazil (usually exhibiting markedly greater pubescence of twig, leaf and legume); M. albiJa H. & B. ex Willd. of the same general range (also more pubescent) ; M. flori- humla Willd. = M. sfrigosa Willd. (which, according to Britton and Rose, are synonyms of M. alhiJa) ; and five or six more recent species. Bentham states (Trans. Linn. Soc. 30:390. 1875): "[M. sefisifiva, M. Velloziana] and the three or four following species [which include M. alhiJa] pass so nearly one into the other that they would probably all have been included by Linnaeus under his M. scnsifiia" Bentham likewise included AL ']iramcncn$is Karst. of Colombia as but a variety of M. Velloziana, but Britton and Killip (Ann. N. Y. Acad. Sci. 3 5:150. 1936) recognize it as a distinct species. (26^) 1950] FLORA OF PANAMA (Lcgiimhiosae) 283 2. Mimosa albida Humb. & Bonpl. ex Willd. Sp. PL 4:1030. 1806. Mimosa strigosa Willd. Sp. PI. 4:1030. 1806, fide Brltt. & Rose. Mimosa jloribiinJa Willd. loc. ck. 1031. 1806, not Vent., fide Britt. & Rose. Mimosa Willdenoivii Poir. in Lam. Encycl. Meth. Suppl. 1:50. 1810. Mimosa racemosa Schlecht. in Linnaea 12:557. 1838, fide Britt. & Rose. Mimosa albida van floribunda B. L. Robins, in Proc. Am. Acad. 33:311. 1898, fide Britt. & Rose. Mimosa albida var. strigosa B. L. Robins, loc. cit., fide Britt. & Rose. Mimosa albida van euryphylla B. L. Robins, loc. cit., fide Britt. & Rose. Mimosa albida var. glabrior B. L. Robins, loc. cit., fide Britt. & Rose. Mimosa Williamsii Standi, in Contr. U. S. Nat. Herb. 18:105. 1916, not Rusby. Mimosa Standleyi Macbride, in Contr. Gray Herb. 59:12. 1919. Usually a slender vine with trailing or clambering stems; branchlets terete, glabrous or more commonly (Panama) strigose or lightly tomentose, unarmed or ^ /o* Fig. 99. Mimosa albida (269) [Vol. n 284 ANNALS OF THE MISSOURI sparingly aculeate (Panama). Leaves moderate, bipinnate, the pinnae a single pair, the leaflets only 2 pairs per pinna; petiole up to 5 or 6 cm. long, eglandular, pubescent like the branchlcts, sometimes aculeate, apiculatc terminally; pinnular axis 1-2 cm. long, appcndagcd above the pulvinus, usually pubescent; leaflets usually asymmetrically oblong or elliptic, broadly acute or obtuse and briefly mucronatc apically, obliquely rounded basally, the costa strongly cxccntric, stri- gosc-pubescent above and more heavily so below; stipules linear to linear-lanceolate, 3-4 mm. long, rigid, erect, pcctinatc-ciliate. Inflorescence of axillary, pedunculate heads, sometimes few-fasciculate; peduncles mostly 1-2 cm. long; heads ovoid, compact, multifloratc; floral bractlets sublinear, with subulate tips, scarcely equal- ling the corolla. Flowers small, sessile, whitish; calyx campanulate, minute; corolla funnelform, about 2 mm. long, usually 4-lobcd, not striate, usually tomentulose apically in bud; stamens 4 (5?), mostly 4-6 mm. long, glabrous; ovary glabrous or subglabrous. Legume more or less oblong, 1-3 cm. long and usually d-l mm. wide, flat, 1- to few-segmented, short-stipitatc or subsessile, acute and subulate apically, setosc-strlgosc or sctose-splnose marginally. Mexico to northern South America. cocle: Pcnonome, Williams 101, The cited specimen is the type collection for M. Williamsii Standi. (= M. SfanJlry/), It appears to be another of the aberrant forms commonly found in M. alhida, a species known to vary extravagantly. Standley himself, with the original description of M. Willianisii, stated his species to be closely related to M. alhiJa. Bcntham (Trans. Linn. Soc. 30:390. 1875) lists Panama as included in the range of M. alhiJa as he interpreted the species. Without any attempt to evaluate B. L. Robinson's subdivision of M. albiJa Into varieties, I have here fol- lowed the 'North American Flora' in pigeon-holing a number of different variants, including M. Williannii Standley, under the early name of AL albida. Even then it Is difficult to find any hard-and-fast difference between this species and M. scfisitiia L. of South America. M. albida is of a weedy nature, invading cleared and pastured lands where it often becomes a serious pest In forming nearly impenetrable tangles. Like other members of the group it is at least moderately "sensitive'* to contact. 3. Mimosa panamunsis (Benth.) Standi, in Contr. U. S. Nat. Herb. 18:104. 1916. MimoSii ifchilis H. & B. ex Willd. var. pauamcush Benth. in Trans. Linn. Soc. 30:391. 1875. A sprawling subherbaccous or suffruticose annual, the branchlcts slender, spar- inglv to heavily hirsute, sparingly to moderately recurved-aculeate. Leaves mod- ratcly small, bipinnate, the pinnae a single pair, the leaflets only 2 pairs per pinna; petiole slender, 15-40 mm. long, sparingly armed or unarmed, eglandular; pinnular axis very short, seldom 5 mm. long; leaflets asymmetrically obovatc to oblong, usually broadest above the middle, up to 2 cm. long and 4-7 mm. v^'Ide, the inner (270) 1950] FLORA OF PANAMA (Lcguminosae) 285 member of the basal pair very reduced, usually obtuse or rounded and briefly mucronate apically, obliquely rounded basally, strigose-pubescent on both upper and (more heavily) lower surface, the venation prominent below; stipules linear, about 4 mm. long, pectinate-ciliate. Inflorescence of axillary, pedunculate heads, racemose terminally because of non-foUate nodes; peduncles up to 3 cm, long, sub- glabrous; heads dense, globular; floral bractlets linear-lanceolate, 1-2 mm. long, pectinate-ciliate- Flowers small, sessile, pinkish-white?; calyx campanulate, minute; corolla funnelform, about 2 mm, long, membranous, usually puberulent apically, 4- (5?) -lobed, the lobes less than half as long as the tube; stamens 4 (5?), about 6 mm. long, glabrous, the anthers orbicular; ovary subglabrous. Legume oblong, mostly 1-1.5 cm. long and 3-5 mm. wide, flat, few-segmented, the persistent margins heavily setose-spinose, the valves less heavily so. Costa Rica, Panama, and Colombia. CANAL zone: Ancon Hill, Bro. Celestine 66; Standley 2jigi, 26321 ; Kiltip 12082. cocLE: Aguadulce, Wittier 4Q^2; Nata, See?nanrt g8. Panama: Corozal Road, Standley 26784, 2gi72; Ktllip J146; northeast of Panama City, Bro, Paid 206. veraguas: Sona, Allen 1 06 1. It is possible that the species should be M. senutiva L, (considered broadly) or but a form of M. albiJa (which in turn does not seem to be clearly distinct from M. sensitiva). The point is debatable as to whether it is worthy of better than varietal rank, but as a variety it would seem as much akin to M. alhida or M. Velloziana as to M. debilis. 4. Mimosa casta L. Sp. Pi. 518, 1753. Mimosa dmniniciana Desv. In Ann. Sci. Nat. Bot. 9:424, 1826, fide Benth. Scandent, suffruticosc plants, the branchlets subterete but angled or ridged, glabrous or somewhat pubescent, densely armed with recurved, broad-based thorns. Leaves moderately large, biplnnate, the pinnae 1 pair, the leaflets mostly 3-4 pairs; petiole elongate, as much as 10 cm. long, eglandular, callous basally, aplculatc terminally, armed like the twigs; pinnae 3-7 cm. long, the rachis pulvinate and usually aculeate; leaflets Inequilaterally elliptic to oblong-Ianccolate, mostly 10-30 mm. long and 5-15 mm. wide, acute apically, obliquely rounded basally, callous basally and with a few conspicuous palmate veins, setose-ciliate on the margins, sometimes with a few appressed setae on the lower surface; stipules narrowly linear- lanceolate, falcate, commonly about 5 mm. long, striate. Inflorescence of axillary, pedunculate heads; peduncles short, 5-15 mm. long; heads globular, several- to many-flowered, the floral bractlets Hnear-lanceolate, 1-2 mm. long. Flowers small, sessile, dense, whitish; calyx very minute; corolla funnelform, 1-2 mm, long, glabrous, 4-lobed, the lobes half as long as the tube; stamens 4, about 1 cm. long, free, glabrous; ovary short, setose, bilateral, the style slightly exceeding the stamens. Legume oblong, usually 3 0-40 mm. long and about 12 mm. wide, flat, glabrous, 4- to 5-segmented, the margins sctose-spinose and persistent. Panama and north coast of South America to Brazil; Lesser Antilles. (271) [Vol. 37 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONE: Miraflorcs, Pit tier 2ig8; Pedro Miguel, Allen 2, 5. Mimosa pudtca L. Sp. PL 518. 1753. Bentham (Trans. Linn. Soc. 30:397. 1875) lists tlie following synonyms for M. puJica: M. enjymlonh Mart.; M. birsuta Mo^^. & Sess.; M. bispiiliila HBK.; M. irriiab'iUs Presl; M. p7tdibufu{a Willd.; M. strialo-stipula Stcud.; M. tctranJra H. & B.; M. nuijuga Duchass. & Walp. Prostrate herb to somewhat suflfruticose, the branchlcts terete, glabrous or pubescent (often both glabrous and pubescent branchlcts on same plant), heavily or lightly armed with recurved thorns, the thorns broad-based and black-tipped. Leaves moderately small, sensitive, bipinnate, the pinnae 1 or (more commonly) 2 pairs, the leaflets mostly about 16 pairs; petiole elongate, up to 4 cm. long, sub- terete, glabrous or pubescent and sometimes even aculeate, eglandular, with a con- spicuous basal pulvinus; rachis short (2-3 mm.) or obsolete; pinnae congested, the pinnular rachis and pulvinus usually setose, with a pair of squami just above the pulvinus; leaflets linear or linear-oblong, 3-12 mm. long and mostly 1-2 mm. wide, abruptly acute or subobtusc apically, inequilaterally truncate or rounded basallv, with 1 or 2 smaller basal veins in addition to the costa, venation remote but visible below, glabrous except the margins (and sometimes the lower surface) yellow-setose; stipules linear-lanceolate, falcate, 4-8 mm. long, setose. Inflorescence of axillary, pedunculate, head-Uke spikes; peduncle about 2 cm. long In maturity; spikes short, very condensed, many-flowered; floral bractlcts linear-lanceolate or lincar-oblanceolate, setose. Flowers pinkish, sessile; calyx extremely minute, only a fraction of a mm. long; corolla funnclform, 1-2 mm. long, glabrous, 4-lobed, the lobes ribout half as long as the tube; stamens 4, about 7 mm. long, free, gla- brous; anthers minute, nearly orbicular; ovary small, glabrous, bilateral, the style about equalling the stamens. Legumes typically in a dense cluster, usually 10-15 mm. long and about 4 mm. wide, flat, glabrous, segmented, somewhat constricted between the articulated, 1 -seeded segments, heavily setose-aculeate on the margins, the margins persisting after dispersal of the segments. Southern Mexico to middle South America; West Indies. Introduced into Old World tropics; cultivated under glass in temperate regions. bocas dfx toro: Chiriqui Lagoon, IL von Wcdcl 1492, l6ig^ 2S'0. canal zone: "Chagres", Fendler g6; Gatun, Hayes ijQ; Las Cruces trail. Hunter & Allen /'26; Quebrada La Palma, Dodge ^ Allen 1733S. chiriqui: Boquete, Woodson 6 Scbery /'2I; Llanos del Volcan, Seibert 324, 55/. cocle: Aguadulcc, Pittier 4S61; Pcnonome, Williams lOO, PANAMA: Chepo, Pittier 4/6o; Taboga Island, Standley 2JOJ2, This interesting "plant oddity" is well known for its ability to quickly fold and droop its leaves with contact or other stimuli. Walking through a field of "sensitive plant" in the tropics one can look back and easily discern the path taken because of the altered position of the leaves brushed against. The plant shows a similar response to darkness or disturbance by rainfall. It is frequently kept as a curiosity, both in the tropics and In greenhouses of temperate climates. The species is abundant in open lowland areas in Panama, but also occurs at higher (272) 1950] FLORA OF PANAMA (Lcguminosac) 287 elevations. It is similar to M. polyJactyla, and is distinguishable from the Mexican M. affhiis only by a difference in legume setae. 6. Mimosa polydactyla Humb. & Bonpl. ex Willd. Sp. PL 4:1033. 1806. An herbaceous to sufFruticose plant, the branchlcts terete, pubescent or glabrate, sparingly or densely armed with broad-based, black-tipped thorns. Leaves mod- erate or moderately large, sensitive, bipinnate, the pinnae 3-5 (usually 4) pairs clustered at the apex of the petiole (hence subdigitate) , the leaflets usually about 30 or more pairs; petiole elongate, up to 8 cm. long, eglandular, usually setose; rachis very condensed, scarcely 3 mm, long; pinnae as much as 12 cm. long, the basal pinnular pulvinus setose; leaflets linear, 4-9 mm. long and usually 1-2 mm. wide, usually acute apically, inequilaterally rounded basally, the costa and remote lateral veins subpromincnt, glabrous except setose marginally and occasionally below; stipules linear-lanceolate, falcate, about 5 mm. long, setose. Inflorescence of axillary, pedunculate heads or very short spikes; peduncles short, usually little more than 1 cm. long; heads dense, many-flowered; floral bractlcts sublinear, setose- laciniatc. Flowers pinkish or purplish, sessile; calyx extremely minute; corolla funnelform, about 2 mm. long, glabrous, 4-lobed; stamens 4, several mm. long, free; ovary glabrous, bilateral, the style equalling the stamens. Legumes typically in a dense cluster, usually 10-15 mm. long and about 4 mm. wide, flat, glabrous, segmented, somewhat constricted between the articulated, 1 -seeded segments, heavily setose-aculeate on the margins, the margins persisting after dispersal of the segments. Panama to Brazil. "WESTLRN PANAMA": Siork, July-Aug. 1923. Mnuosa polydactyla is apparently less frequent In its range than Is M. [nulica. It is very similar to M. pnJica, but possesses more numerous pinnae and leaflets and has shorter peduncles than is characteristic of the latter species. 7. Mimosa pustlla Bcnth. In Bot. Voy. Sulphur, 90. 1844. Low, prostrate or semi-pros t rate herb, the branchlets glabrous to setose or lightly hirsute, scarcely armed (thorns, if present, few, nearly straight, weak), terete or subterete, Inconspicuously lentlcellate. Leaves small, bipinnate, the pinnae 1-3 pairs opposite on the rachis, the leaflets 5-20 pairs per pinna; petiole less than 1 cm. long, glabrous or pubescent like the twigs, terete, eglandular; rachis similar to the petiole, eglandular but with setiform appendages, the rachial internodes about equalling the petiole; pinnae remote (i.e. not subdigitate), 5-35 mm. long; leaflets linear-oblcng, 3-8 mm. long and 1-1.5 mm. wide, broadly acute to obtuse and briefly mucronulate apically, obliquely rounded or subtruncate basally, glabrous except often ciliate-margined, the costa and lateral veins quite conspicuous (the lateral veins curving upward to subparallel the costa); stipules ovate, 2-4 mm. long, acuminate, longitudinally striate. Inflorescence of solitary, axillary, peduncu- late heads; peduncles short, rarely even as much as 1 cm. long, pubescent as the (273) [Vol. 37 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN twigs, ebracteate except at tKc head; head small, more or less obconic, few- to several-flowered, nearly cnsheathed (at least in youth) by the prominent floral bracts; floral bracts ovate-lanceolate, usually 2-3 mm. long, subscarious and striate, usually clliate but otherwise glabrous. Flowers small, pinkish; calyx campanulate, minute, glabrous; corolla funnelform, 1—2 mm. long, membranous, glabrous, 4- lobed; stamens 8 (sometimes fewer?), 3-4 mm, long, glabrous; ovary glabrous. Legume oblong, typically less than 1 cm. long and about 4 mm. wide, flat, setose, 1- to 3 -jointed, the joints separating from the margins in age. Mexico to Panama. Panama: Pearl Islands, San Jose Island, 7. M. Jobtjsfon ///j ^^^> 8l8A, The Johnston collections constitute a new record from Panama for this inter- esting species, concerning which I. M. Johnston is publishing notes elsewhere (in mss. as this is written). The species is the smallest, least offensive of any of the genus In Panama, and is not apt to be confused with other species except perhaps M. puJica. I have not had opportunity to examine the type of the species, but there seems little doubt that the name applies to the Panama collections. The Colombian M. martcusis Britt. & Rose may be a more pubescent variety of Af. pusilla. 8. Mimosa invisa Mart, in Flora 20:Bcibh 2:121. 1837. Schnwkia brachycarpa Bench. In Hook. Jour. Bot. 2:130. 1840, fide Benth. Mimosa diplotricha C. Wright in Sauvalle, An. Acad. Habana 5:405. 1896, fide Britt. & Rose. Morongia pilosa Standi, in Contr. U. S. Nat. Herb. 18:105. 1916, fide Britt. & Rose. Schrankia pilosa Macbridc, in Contr. Gray Herb. 59:11. 1919. Herb or small shrub, prostrate, decumbent or scandent, the branchlets strongly angled, subtetragonal, usually lightly pilose (sparingly so on older twigs especially), copiously armed with recurved thorns. Leaves reportedly sensitive, moderate, bipin- nate, the pinnae mostly 4-7 pairs opposite on the rachis, the leaflets several to many (usually about 15—20) pairs per pinna; petiole elongate, 2—5 cm. long, characteristically exceeding the rachis, eglandular, pulvinate, aculeate; rachis equal- ling or almost equalling and similar to the petiole, bearing setiform appendages at insertion of the pinnae; pinnae comparatively remote, mostly 2-3 cm. long, bearing just above the pulvinus usually paired, setiform appendages; leaflets linear or linear- oblong, 2-5 mm. long and about 1 mm. wide, bluntly acute or subobtuse apically, incquilatcrally rounded or subtruncate basally, lightly pubescent or subglabrous, the costa visible but secondary venation obscure; stipules setiform, about 4 mm. long. Inflorescence of pedunculate heads solitary to few-fasciculate from older foliate and subterminal non-foliate nodes; peduncles normally no more than 1 cm. long, aculeate and usually sparingly pubescent; heads compact, globular, several- to many-flowered; floral bractlcts oblanccolate, about 1 mm. long, glabrous. Flow- ers sessile, pink; calyx cupular, much less than 1 mm. long, glabrous; corolla funnelform, 2—3 mm, long, glabrous, 4-lobcd, the lobes about equalling the tube; (274) 19 SO] FLORA OF PANAMA (Le gum'tnosae ) 289 ► Fig. 100. Mimosa sovtnrans (275) [Vol. 37 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN stamens 8, about 6 mm. long, the filaments thickish, the anthers about Yz mm. long; ovary short, vcrrucose laterally, the style equaUing the stamens, the stigma narrow. Legume oblong, usually less than 2 cm. long and about 5 mm. wide, flat, segmented, norn'ially pilose, the persistent margins sctose-splnose, Mexico to Argentina; West Indies. BOCAS DEL TORO: Almirantc, Rati'Ice d Stark 4gS. canal zone: Corozal, Sfandlcy 2/'j68; Summit, StanJlcy 26g§l, Panama: Chcpo, Pittkr 4544; Juan Franco Race Track, Standley 2Jjg8, province unknown: Las Delicias, Carlelort 41 , Mimosa ini/sa is on the ''continental divide" between Mimosa and Schrankia; vegetative and floral appearance is that of Scbrankia, but the comparatively broader legume necessitates inclusion in Mimosa. The species is another "sensitive plant" or Jormilona^ rather easily distinguishable from M. pudtca by the comparative remoteness of the pinnae. The plant frequently invades cleared areas, where it becomes an objectionable weed, potentially a source of severe lacerations to the unwary. Pittlcr reports various parts or infusions of the plant to be both mildly toxic and medicinal. M, iniisa has a great many expressive local names, such as vialici'i ilc mulhcr in eastern Brazil and raspacanilla in lower Central America. 9. Mimosa somnians Humb. & Bonpl. ex Willd. Sp. Pi. 4:103 6. IS 06. Mimosa palpitans H. & B. ex Willd. loc. cit. 1806, hdc Renth. Mimosa somnicidosa HBK. Nov. Gen. & Sp. 6:257. 1824, fide Benth. Mimosa acittijlora Bentli. In Hook. Jour. Bot. 4:^97, 1842, fide Benth. Mimosa poJocarpa Benth. loc. cit. 1842, fide Benth. Mimosa quaJrijuga Salzm. ex Benth. loc. cit. 398. 1842, fide Benth, Decumbent or nearly erect or even subscandcnt herb or subshrub, the branchlets subtcrete, longitudinally striate, usually pilose with gland-tipped hairs, sparingly or moderately aculeate with broad-based, recurved thorns. Leaves moderate, bi- pinnate, the pinnae usually about 4 (2-8) pairs opposite on the rachis, the leaflets many (20 more or less) pairs per pinna; petioles moderate, 1-3 cm. long, normally shorter than the rachis, armed and pubescent like the branchlets, otherwise egland- ular; rnchis 1.5-8 cm. long, like the petiole but less pubescent, bearing a subulate appendage at insertion of each pair of pinnae; pinnae 1—4 cm. long, setose-splncse particularly on the pulvinus; leaflets broadly linear, 2-5 mm. long and usually less than 1 mm. wide, rounded or obtuse apically, obliquely subtruncate basally, gla- brous or sparingly puberulent, the venation obscure except for the costa; stipules lanceolate-subulate, about 5 mm. long, pubescent. Inflorescence of pedunculate heads solitary or few-fasciculate from both non-foliate subterminal nodes and lower foliate nodes, thus appearing racemose distally; peduncles up to 3 cm. long in age, most frequently glabrous; heads globular, compact; floral bractlets ovate-lanceolate, about 1 mm. long, ciliate. Flowers sessile, pink; calyx minute, much less than 1 mm. long; corolla funnelform, about 3 mm. long, glabrous, markedly striate, the 4 lobes about equalling the tube, subrigid and pointed in bud; stamens 8, about 7 mm. long, glabrous, the filan"icnts somewhat thickish. Legume narrowly linear, (276) 1950] FLORA OF PANAMA (Legtimiuosac) 291 4-7 cm. long and about 4 mm. wide, flat, several-segmented, stipitate, usually glandular-pubescent and setose when young, less prominently so in age, the per- sistent margins short aculeate-setose. Mexico to Argentina. CANAL ZONE: Balboa, Standley 26063; Fort Kobe Road, Woodson^ Allen & Scibert 1410. CHiRiQUi: Boquete, Woodson ^ Schery /'g6; Pittier 3321. cocle: El Valle de Anton, Allen 4/0, 28iy. Panama: without data, M.B.G. I0p02gj; "Sabanas", Bro, Paid SYl Tumba Muerto Road, Standley 2g//'0, province unknown: without locality, Seemann 10^, This species is rather well distinguished by the glandular pubescence (usually) and striate corolla. Along with certain other species it seems to bear the common name dormilona in Central America, indicating that it too is a "sensitive plant" responding visibly to contact or other stimuli. It seems to prefer open, moist, often sandy habitats at moderate elevations, 10. Mimosa pigra L. Cent. Pi. in Amoen, Acad. 4:274. 1755. Mimosa aspcrafa L. Syst. Nat. ed. 10, 1312. 1759. Mimosa polyacantha Willd. Sp. PI. 4:1034. 1806, fide Benth. Mimosa bispida Willd. loc. cit. 1037. 1806, fide Benth. Mimosa caucscens Willd. loc. cit. 103 8, 1806, fide Benth. Mimosa pcllita H. & B. ex Willd. loc. cit. 1037. 1806, fide Benth. Mimosa dliata Willd. Enum. Hort. Berol. 1048. 1809, fide Benth. Bcntham also lists other African and Far Eastern synonyms. Shrub to a few meters tall, sometimes scandent, the branchlets usually con- spicuously setose-hispid and armed with formidable recurved thorns. Leaves re- ported sensitive, moderate to large, bipinnate, the pinnae up to 15 or even more pairs opposite on the rachis, the leaflets usually about 25 or 30 pairs per pinna; petiole comparatively short, seldom more than 1 or 2 cm. long, eglandular, hispid like the twigs, usually aculeate; rachis 4—2 5 cm. long, similar to the petiole, egland- ular, the thorns frequently becoming acicular; pinnae 2-7 cm. long, hispid; leaflets linear, 2-8 mm. long and approximating 1 mm. in width, acute apically, inequi- laterally rounded or subtruncate basally, appressed-pubescent (especially below) and setose-cilia te margined, longitudinally few- veined, these veins prominent below; stipules ovate-lanceolate, as much as 8 mm. long, pubescent. Inflorescence of solitary or few-fasciculate, pedunculate heads, axillary from foliate and sub- terminal non-foliate nodes; peduncles up to 5 cm. long in age, hispid; head ovoid, dense; floral bractlets linear, 2-3 mm. long, pectinate above the middle. Flowers sessile, whitish; calyx tubular, about 2 mm. long, cleft to one side, laciniate apically, glabrous; corolla funnelform, about 4 mm. long, short-hispid apically, the 4 lobes less than Yz the length of the tube; stamens 8, free, the filaments scabrous and expanded distally; ovary setose-hispid. Legume linear-oblong, mostly 4-7 cm. long and little more than 1 cm. wide, short-beaked, stipitate, flat, 7- to 20-seg- mented, densely hispid-setose both on the persistent margins and on the segments. Mexico to Brazil and Peru; West Indies; Africa and Far East (introduced?). (277) [Vol. 2,7 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN CANAL ZONE: *'Chagrcs", FcnJler qS; Frijoles, Bro, Heriberto 84. cocle: Penonome, Williams 111. PANAMA: between Panama and Chepo, Dodge, Hurrter, Stcycrmark ^ Allen 16684; Bro. Maurice j6l; Tumba Muerto Road, Statnlley 2g8j2, The species is distinctive except in Mexico where a smaller, less pubescent form has been regarded as a separate i>pecies (AL BerlanJieri A. Gray) or variety (var. Berlandieri (Gray) Robins.). The heavy, coarse pubescence and multipinnular leaf serve to distinguish it from other species through most of its range. M. pigra quickly when disturbed. It bears, as does M. pud. p pica- p. ope 8. SCHRANKIA Willd. ScHRANKiA Willd. Sp. Pi. 4:1041. 1806, not Medic. 1792; nom. conserv. Leptoglottis DC. Mem. Legum. 451. 1825. Morongia Britt. in Mem. Torrcy Bot. Club 5:191. 1894. Perennial herbs or subshrubs, often decumbent or clambering, armed with re- curved thorns, the branchlets usually markedly angulate. Leaves moderate or moderately small, bipinnate, usually sensitive, the pinnae few or several remote pairs opposite on the rachis, the leaflets several to many pairs per pinna; petiole comparatively elongate, in many species exceeding the rachis, aculeate, eglandular; rachis similar to the petiole, eglandular but with subulate appendages at insertion of pairs of pinnae; pinnae usually bearing a pair of bract-like mucrons just above the basal callus; leaflets small, usually inequilateral basally, the venation obscure or less frequently prominent ; stipules linear-subulate, arcuate, subpersistent. In- florescence of axillary, pedunculate heads; peduncles short (most South American species) or moderately elongate (many Mexican and North American species) ; heads dense, globular; bractlets small, concave but not peltate. Flowers small, essentially sessile, mostly 5-merous, pinkish, the upper ones perfect, the lower ones usually staminate; calyx small, nearly campanulate, prominently toothed, usually glabrous; corolla funnelform-campanulate, sympetalous, the teeth or lobes promi- nent, usually glabrous; stamens 8-13 (usually 10), free (except perhaps at base), exserted; anthers small, ovate-orbicular, eglandular; ovary nearly symmetrical, usually glabrous, several- to many-ovulate, briefly stipitate; style about equalling the stamens; stigma terminal, not expanded. Legume narrowly linear, terete or at least turgid, usually aculeate, not jointed, the face of the valves separating from the margins in dehiscence (thus at length longitudinally 4-valvate), usually with a brief or elongate acuminate beak; seeds longitudinal, oblong, obliquely truncate, blackish. Temperate and tropical North America to middle South America; West Indies; introduced into Africa, Pacific Islands, and Far East. (27S) 1950] FLORA OF PANAMA (Legumlnosac) 293 Schrankia is a small genus, distinguisha:ble from Mimosa only with difficulty, and reliably only in mature fruit. Except that Schrankia has been traditionally separated from Mimosa, there seems little more justification in recognizing it than a great many Britton & Rose segregate genera not commonly accepted. As is typical of most genera of the Mimosoideae, species within the genus are not clear- cut. Of the 27 species listed by Britton & Rose (N. Am. Fl, 23:138-144) only a fraction are likely tenable. Two species are reported to occur in Panama, prob- ably both of them ''good" but possibly grading into "species'* of Mexico and the United States. a. Pinnae mostly 2—3 pairs; venation of leaflet except for the costa obso- lete; legume about 4 mm. wide, long-beaked I, S. leptocarpa aa. Pinnae mostly 4—5 pairs; leaflet venation delicately reticulate below; legume 5-6 mm. wide, short-beaked 2. S. hamata 1. Schrankia leptocarpa DC. Prodr. 2:443. 1825, Leptoglottis leptocarpa (DC.) Standi, in Jour. Wash. Acad. 15:458. 1925. ?Schrankia trijuga Goyena, Fl. Nic. 1:379. 1909. Perennial, subherbaceous plant, decumbent, straggling or somewhat scandent; branchlets lightly to heavily armed with recurved thorns, glabrous, angled or sub- tetragonal. Leaves moderate but slender and lax, bipinnate, the pinnae mostly 2—3 pairs (rarely 1 or 4-5 pairs), the leaflets ordinarily about 10-15 pairs per pinna; petiole elongate, 2—8 cm. long, subglabrous, generally armed like the branchlets, eglandular; rachis 1—4 cm, long, similar to the petiole, bearing above at insertion of the pairs of pinnae 1—2 subulate, stipule-like appendages (these sometimes cadu- cous) ; pinnae 1—4 cm. long, callous basally and usually bearing there a pair of minute, bract- like mucrons; leaflets linear-oblong to narrowly obovate-elliptlc, 3-10 mm. long and 1-2 mm. wide, rounded-submucronate or obtuse apically, inequilaterally rounded basally, glabrous except ciliate-margined, dull, the venation obscure; stipules linear-subulate, curved, few to several mm. long. Inflorescence of axillary, pedunculate heads; peduncles short, mostly about 1 cm. long, eglandular; heads several-flov/cred, orbicular, the bractlets lanceolate or oblanceolate. Flowers subsessile, pinkish; calyx campanulate, considerably less than 1 mm. long, glabrous, irregularly 5-toothed; corolla funnelform-campanulate, about 2 mm. long, gla- brous, the 5 teeth almost equalling the tube; stamens 10, about 6 mm. long, free, glabrous, the anthers ovate-orbicular; ovary substipitate, verrucose. Legume linear, up to 12 cm. long, acuminate-subulate, moderately aculeate, the thorns in 3 rows on each margin with an additional row on each valve, subglabrous, the valves separating from the margins in dehiscence. Central America and West Indies to middle South America; introduced into Old World tropics. CANAL zone: Ancon, Pit tier 6/6p. cocle: Aguadulce, Pit tier 48 3S; Pcnonomd, Williams log. Panama: Las Sabanas, Bro, Heriberto 26 J; Standley ^JPJJ. (279) [Vol. Z7 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN Certain species of Mexico and tlic soutKern United States are remarkably similar to S. Icpfocarpa, For example, S. RonTicriana, a number of specimens commonly passing as S. vucvopbylla (^^ S. borriJula, = S. uncinafa, = S. augustata), and several Small and Britton & Rose species are difficultly separable from it. Yet the name S. leptocarba has priority over most possible synonyms, and will likely endure. Fig. 101. Scbrafjk ia ha mafa 2. ScHRANKiA HAMATA Humb. & Bonpl. cx Willd. Sp. PL 4:1042. 1806. Mimosa tetragoiia Poir. in Lam. Encyck Mcth, Suppl. 1:56. 1810, fide Bench. Lepioglottis hamafa Standk in Jour. Wash. Acad. 15:458. 1925. Slender, clambering, mostly glabrous, herbaceous plant, the branchlets penta- gonal and copiously armed on the ridges with recurved thorns. Leaves bipinnate, the pinnae usually 4—5 pairs, the leaflets mostly 10—20 pairs; petiole moderate, about 5 cm. long, it and the rachis aculeate; rachis usually exceeding the petiole, eglandular but with interpinnular mucrons; leaflets linear, up to 7 mm. long, acute (2W) 19 5 0] FLORA OF PANAMA (Legtiminosae) 295 or obtuse apically. inequllatcrally truncate basally, glabrous except sparingly ciliatc on the margin:*, delicately reticulate-velned below. Peduncles axillary, about 1 cm. long, pubescent, often aculeate; bead globose. Flowers small, dense, sessile; calyx campanulate, about Yz mm. long, glabrous, briefly 5-dentate; corolla funnclform, about 3 mm. long, glabrous, 5-lobate, united below for about % its length; stamens 10, about 6 mm. long, glabrous, free; antbers ovate, about Yz mm. long; ovary glabrous, briefly stipitate, multi-ovulate, angled, with the style 4-5 mm. long. Legume 6-10 cm. long and about 5 mm. wide, terete-tetragonal, heavily setose- aculeate, the beak being about 1 cm. long. Panama and Colombia. CANAL ZONE: AncoH, PittscT 686j; Balboa, StanJley 321IJ; Bella Vista, Piper 51 14. darien: Pinogana, Allen 42JJ. VK^sM.k\ Corozal road, Standley 26/61; Panama, Mac- bride 2609; Standley 2//'00; Tumba Muerto road, Standley 29702. Scbranha kamata is reported quite common in Colombia and Panama. It ap- parently is distinct from S. Icpfocarpa (in having a thicker, more spiny legume, larger leaves with more pinnae, and leaflets with lateral veins distinct), but may not be separable from certain Mexican species. \ 9. NEPTUNIA Lour. Neptunia Lour. Fl. Cochinch. 653. 1790. Hemidesmas Raf. Sylva Tellur. 119. 1858, in part. Generally herbs but sometimes somewhat woody, of moist or swampy ground and sometimes free-floating on ponds or quiet water, the branchlets unarmed, not lenticellate, in miost species glabrous, occasionally spongy-inflated in submerged stems. Leaves usually moderate, bipinnate, the pinnae opposite, few, the pairs moderately remote, the leaflets several to many pairs, small; petiole prominent, glandular or eglandular; rachis similar to and usually approximating the petiole, eglandular; leaflets nearly linear, inequilateral basally, usually glabrous but often ciliate, the venation prominent or obscure; stipules conspicuous, often cordatc- subclasping, sometimes striate-nerved. Inflorescence of axillary, pedunculate heads or very short spikes, usually soUtary; peduncles elongate, commonly bearing 2 small or large bracts; heads dense, mostly obovoid, the basal flowers staminate or sterile and smaller in bud, the upper flowers perfect, larger in bud. Flowers sessile, usually yellowish, the basal ones with prominent petaloid staminodia, the perfect ones relatively inconspicuous; calyx synscpalous, more or less cupulate, 5-partcd; corolla of 5 free petals or these in some cases coherent near the middle (but free below and above), usually glabrous; stamens or staminodia mostly 10, free; ovary generally glabrous, the stout style elongate and bearing an expanded stigma. Legume short and broad, flat, unarmed, 2-valved, not septate nor the valves sep- arating from tho margins, the seeds transverse. World tropics and subtropics; in the Western Hemisphere from middle United States to Brazil, (281) [Vol. Z7 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ncptunia is a small genus, containing 6 or fewer species In the Americas- De- CandoUe and others included it as a section of Desmanthus, but more recently it has been generally accepted as a distinct genus. It is unusual and distinctive In possessing basal flowers of the head with conspicuous petaloid staminodia, and in preferring aquatic or swamp habitats. a. Stipules strlate-ncrved; leaflets ciliate, the venation remote but promi- nent below; petals coherent about at the middle; peduncular bracts borne above the middle - '■ N. pubf.scens aa. Stipules scarcely nerved; leaflets glabrous, the venation obscure; petals free; peduncular bracts borne near and below the middle^ or absent. b. Not floating, the stem seldom developing a spongy outer tissue; petiole glandular; flowers yellow 2. N. plena bb. Frcc-floatini;, with a thick spongy outer tissue on the submerged stem; petiole cglandular or merely callous apically; flowers whitish?., 3. N. prostrata 1. Neptunta I'UBFSCENS Benth. in Hook. Jour. Bot. 4:3 56. 1842. Neptunia floridaua Small, in Bull. Torrey Bot. Club 25:138. 1898. Neptunta Ltfulheimeri B. L. Robins., in Proc. Am. Acad. 33:333. 1898. Neptunta microcarpa Rose, in Cantr. U. S. Nat. Herb. 8:300, 1905. Slender herb or subshrub, prostrate or ascending from a somewhat woody base, the branchlets terete, glabrous or pubescent. Leaves moderate or moderately small, bipinnate, the pinnae 2-4 (normally 3) pairs opposite on the rachis, the leaflets many (12-40) pairs per pinna; petiole slender, 1-2 cm. long, pubescent or gla- brous, eglandulur; rachis about as long as the petiole, eglandular, terminally apicu- latc; pinnae mostly 2-4 cm. long, moderately remote, the axis somewhat alate; leaflets lincar-oblong, 3-5 mm. long and 1-2 mm. wide, obtuse apically, obliquely rounded basally, usually glabrous except for a ciliate margin, the venation promi- nent below but remote; stipules ovate or lanceolate, about 4-5 mm. long, promi- nently longitudinally striate-nerved, subpersistent. Inflorescence of solitary, axillary, pedunculate heads; peduncles elongate, up to 6 cm, long in age, usually minutely 1- to few-bracteate near or above the middle; heads globose to briefly oblong, bearing dimorphic flowers; floral bractlcts lanceolate or oblanceolate, about 2 mm. long, ciliate or glabrous. Flowers of 2 (3?) types, the few lower ones of head unisexual and with conspicuous petaloid stamens or staminodia; upper flowers without petaloid stamens or staminodia, usually perfect (occasionally with pistil aborted), sessile, yellow; calyx cupulate-funnelform, about 2 mm. long in perfect flowers, somewhat shorter in basal flowers, prominently 5-lobed, the lobes usually ciliate; corolla funnelform, about 3 mm. long in perfect flowers, somewhat shorter in basal flowers, glabrous, 5-lobate, the lobes about equalling the imperfectly closed tube; stamens (or staminodia) usually 10, glabrous, 5-6 mm. long; ovary glabrous, subfusiform, the style stout and about cqualUng the stamens. Legume broadly oblong, mostly 2-3 cm. long and 1 cm. wide or wider, flat, rounded apical- ly and briefly mucronate, substipitate basally, 2-valved, few-seeded, glabrous, the seeds transverse and sHghtly oblique. (in) 1950] FLORA OF PANAMA (Legumifiosae) 297 Florida to Texas and Mexico; West Indies; southern Central America; Peru; southern South America. cocLE: Aguadulce, Pittier 4g62. This species is readily distinguished from others of the genus in Panama by the striate stipules, adherent petals, and ciHate leaflets with prominent venation. The species much resembles N. lutea of the United States. 2. Neptunia plena (L.) Benth. in Hook. Jour. Bot. 4:355. 1842. Mimosa plena L. Sp. Pi. 519. 175 3. Mimosa punctata L. Sp. PI. ed. 2, 1502. 1763. Desmanthus plenus Willd. Sp. Pi. 4:1045. 1806. Desmanfbus punctatus "Willd. lac. cit. 1047, 1806. Mimosa lycopodioides Desf. in Pers. Syn. PI. 2:263. 1807, fide Benth. Acacia lycopodioides Desv. in Jour. Bot. 3:68. 1814. Desmanthus polypbylhis DC. Prodr. 2:444. 1825, fide Benth. Acacia punctata Desf. Cat. Hort. Par. 300. 1829. Mimosa adenanthera Roxb. Fl. Ind. 2:5 54. 18 32, fide Benth. Neptunia polyphylla Benth. in Hook. Jour. Bot. 2:129. 1840, Neptunia surinamensis Steud. in Flora 26:759. 1843, fide Benth. Desmanthus comosus A. Rich. Ess. PL Cub. 475. 1845, fide Griseb. Subprostrate or ascending subshrub of swampy habitats, the branchlets gla- brous, not Icnticellate. Leaves moderately small, bipinnate, the pinnae 2-4 (mostly 3) pairs opposite on the rachis, the leaflets 10-25 pairs per pinna; petiole 1-2 cm. long, glabrous, somewhat flattened above, bearing a sessile, orbicular gland apically just below insertion of lowermost pair of pinnae; rachis 1-4 cm. long, similar to the petiole^ cglandular (except for gland already noted at apex of petiole), termi- nally apiculate; pinnae 1-5 cm. long, the pairs moderately remote from one an- other; leaflets linear or linear-spatulate, 3-10 mm. long and 1-2 mm. wide, rounded or obtuse and usually very briefly mucronate apically, obliquely subtruncate basally, glabrous, all venation obscure; stipules obliquely ovate-lanceolate and subclasping, usually about 5 mm. long. Inflorescence of axillary, pedunculate heads or very short spikes; peduncles up to 10 cm, long in age, glabrous, conspicuously bibract- eate below the middle; heads dense, the floral bractlets obovate-lanceolate, little more than 1 mm. long. Flowers sessile, yellow, the upper ones perfect, the lower ones staminate or infertile and with 10 conspicuous petaloid staminodia 7—8 mm. long; calyx cupulate, about 1 mm. long, 5-lobed, glabrous; corolla of 5, free, spatulate petals about 3 mm, long, glabrous; stamens 10?, 4-5 mm. long, glabrous, the anthers almost 1 mm. long; ovary glabrous, dark; style stout, exceeding the stamens; stigma expanded, truncate. Legume oblong, 2—4 cm. long and about 1 cm- wide, apiculate, flat, glabrous, conspicuously stipitate, 2-valved, the seeds several, transverse. Mexico and Guatemala, West Indies, Panama and Colombia to Brazil. COCLE: Penonome, Williams p7- (2^3) [Vol. 37 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species characteristically grows in marshy, open areas such as the shallow ponds of the savannas. N. plena can be distinguished rather readily from other species of Nepttinia, except N, prostratay by a variety of characters. However, occasionally plants appearing identical with N". plena (or other species) will develop the thick, fleshy, floating stems characteristic of N. prostrata. This stem char- acteristic has been the chief means of distinguishing N. prosfrata, 3, Neptunia prostrata (Lam.) Baillon, in Bull. Soc. Linn. Paris 1:356. 1883. Mimosa prostrata Lam. Encycl. Meth. 1:10. 1783. Neptunia oleraeea Lour. Fl. Cochinch. 654. 1790. Mimosa nutans Vahl, Symb. 3:102. 1794 (L.f. Suppl. 439. 1781. ~ nomen nudum), Miftiosa lacustris H. & B. Pi. Aequin. 1:55. 1806. Desmanthus lacustris Willd. Sp. PI. 4:1044. 1806. Desmantbus nutans Willd. loc. cit. 1806. Mimosa aquatica Pers. Syn. Pi. 2:263. 1807, fide Benth. fer DC. Prodr. 2:444. 1825, fide Bcnth. 1830-33. Acacia lacustris Dcsf. Cat. Hort. Paris, 301. 1829. Neptunia stoloniferu Guill., Perr. & Rich. Fl. Seneg. 2 39. Prostrate, floating herb or ''subshrub," the submerged stems spongy-inflated with a pithy outer surface, bearing nodular roots, the above-water stems glabrous, flexuous, not lenticellate. Leaves moderate, 2-ranked, bipinnate, the pinnae mostly 3 pairs opposite on the rachis, the leaflets usually about 15 pairs per pinna; petioles mostly 2-3 cm. long, glabrous, somewhat flattened above, cglandular or with an apical callus just below insertion of the basal pair of pinnae; rachis similar to and approximating the petiole; pinnae usually 2-4 cm. long, the pairs moderately re- mote from one r.nothcr, glabrous; leaflets linear or linear-spatulatc, 3-12 mm. long and 1-3 mm. wide, rounded or obtuse and often briefly mucronulate apically, obliquely rounded or subtruncate basnlly, glabrous, the venation obscure; stipules inequilaterally ovate-lanceolate, generally about 5 mm. long, glabrous, not striate. Inflorescence of axillary, pedunculate heads or very short spikes; peduncles elongate, as much as 15 cm. long, glabrous, usually bearing near or below the middle 2 rather small bracts; heads obovoid in bud, dense, bearing staminate or infertile flowers basally and perfect flowers apically. Flowers sessile, bi-(tri?) -morphic, reportedly whitish fading yellow; calyx of perfect flowers, cupulate-funnelform, about 2 mm. long (smaller in basal flowers), glabrous, 5-lobed; corolla of 5 lincar-spatulate petals, about 4 mm. long, glabrous; stamens of perfect flowers about 10, 7-8 mm. long; staminodia of basal flowers normally 10, petaloid, as much as 15 mm. long, glabrous; ovary glabrous; style exceeding the stamens; stigma expanded, truncate. Legume oblong, 10—2 5 mm. long and about 8 mm. wide, flat, somewhat curved, stipitate, briefly beaked, glabrous, few-seeded, the seeds transverse. World tropics. Tn Western Hemisphere: Mexico and Guatemala, West Indies, Panama and Colombia to Peru and Brazil. cocle: Aguadulce, Woodson, Allen & Seibert l2op. There is some question as to whether the cited specimen may more properly (284) 1950] FLORA OF PANAMA (Lcgumhtosae) 299 belong to this species or to iV, plena. The floating nature of the spongy, inflated stem, however, warrants its inclusion as N. prostrata. Probably N, prostrata is not a valid species, but merely a form of N. plena growing floating in water and assuming a prostrate position. 10. DESMANTHUS Willd. Desmanthus Willd. Sp. PL 4:1044. 1806, nom, conserv. Acuati Medik. Theodora, 62. 1786. Darliugtouia DC. in Ann. Sci. Nat. Bot. 4:97. 1825. Aeuania Ktzc. Rev. Gen. PL 1:158. 1891. Perennial, small, usually diffuse or decumbent shrubs, or subherbaceous. Leaves moderately small, bipinnate, the pinnae one to many pairs opposite on the rachis, the leaflets few or many pairs, opposite; petiole short, either it and/or the rachis bearing one or more subcupular glands, very rarely eglandular; leaflets small, ob- scurely veined In most species, glabrous or infrequently pubescent; stipules seti- form. Inflorescence of axillary, pedunculate heads, the peduncle usually slender; head dense, usually few-flowered, the flowers more or less erect and the head then less than hemispherical; floral bracts nearly lanceolate, apparently never peltate as In most species of Leticaena. Flowers small, sessile, whitish or greenish; calyx synsepalous, more or less campanulate, 5-dentate; corolla essentially polypetalous, 5-parted, valvale, the petals narrowed basally; stamens 5 or 10 (staminodes 5 or 0), free, normally exceeding the perianth; anthers comparatively larger as in Lencaena, not smaller as in Acac/ay eglandular; ovary subsessile, bilateral, several-ovulate, usually glabrous; style thickened above Into a truncate-concave stigma. Legume small, linear, flat, 2-valved, straight or curved, sometimes somewhat constricted between the seeds, dehiscent from the apex (the rather thin, dehisced valves sub- perslstcnt), subscptate or continuous; seeds usually oblique, compressed. World World Desmanthus is obviously close to Lencaena, the basic floral and fruit characters being almost identical. However, whereas Laicaena consists chiefly of arborescent species, species of Desmantfms are small shrubs or subherbaceous. In Central America additional characters serve to rather readily separate these genera, the more obvious of which are used in the key to the genera. Desmanthus is a small genus, but exceedingly troublesome from the standpoint of specific delimitation. As collections have accumulated it has become apparent that many of the characters on which species had been formulated do not hold constant, and that intergradatlon Is rampant between almost all "species." Two specimens may appear distinct, but usually "connecting" specimens exhibiting all degrees of intergradatlon can be found to link them. Thus, for practical reasons, if no other, not more than a handful of species appear worthy of recog- (2S5) [Vol. 37 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN nition, and even these may not be readily distinguishable. On the basis of general observation of the genus the three "species" reportedly occurring in Panama are here combined under the oldest name, which thus considered is the only species known from Central America. L Uhsmantiius virgatus (L.) Willd. Sp. Pi. 4:1047. 1806. Mimoh! lirgata L. Sp. Pi. 519. 175 3. Wvnna pcniawhuratia Mill. Card. Diet. cd. 8, no. 3. 1768, not L., fide Bntt. & Rose. Mimosa angusfhiliqua Lam. Encycl. Meth. 1:10. 1783. Acuan firgafinii Mcdik. Theodora, 62. 1786. Acacia I'ir^ata Gacrtn. Fr. & Scm. 2:317. 1791. Dcsmanlhus dcpressus H. & B. ex Willd. Sp. PI. 4:1046. 1806. Desmanihus diffiisiis Willd. loc. cit. 1806. Mivuna deprcssa Poir. in Lam. Encycl. Meth. Suppl. 1:58. 1810. Desnunif/uis sf rictus Bertol. Giorn. Arcad. 21:190. 1824. Desmaitthus IcptophyUus HBK. Nov. Gen. & Sp. 6:264. 1824, fide Bcnth. Dcsmanthus tcncUus DC. Prodr. 2:445. 1825. Acacia angustisiliqita Desf. Cat. Mort. Paris, ed. 3, 300. 1829. Desman thus pratorum Macf. Fl. Jam. 1:311. 18 37. Acacia depaupcrata Mart. In Steud. Norn. Bot., ed. 2, 4. 1841, fide Bcnth. Desmanfbus virgatus strict us Griscb. Fl. Br. W. Lid. 218. 18 60. Acacia Icptospcrma Bcllo, In Anal. Soc. Esp. Hist. Nat. 10:265. 1881. Acuatt deprcssum Ktze. Rev. Gen. Pi. 1:158. 1891. Acuan latum Britt. & Rose, in N. Am. Fl. 23:132. 1928. Acuan bahamcuse Britt. & Rose, loc. cIt. 1928. Acuan insularc Britt. & Rose, loc. cit. 133. 1928. Small shrub or subherbaceous, nearly erect or more commonly diffuse or de- cumbent, the branchlcts essentially glabrous, angled or subtetragonal, scarcely Icnticellate. Leaves moderately small, bipinnate, the pinnae 1-7 (mostly 2-4) pairs opposite on the rachis, the leaflets mostly 10-20 pairs per pinna; petiole usually no more than 5 mm. long, subtercte, perhaps ciliate above, bearing apically at insertion of the lowermost pair of pinnae a large or small, orbicular to obovate, cupulate or patcUiform, sessile gland; rachis usually flattened and lightly pubescent or ciUate above, cglandular (except for gland already described at juncture of rachis and petiole), apiculate from the lower side apically; pinnae 1-4 cm. long; leaflets linear or linear-oblong, 2-8 mm. long and mostly 1-2 mm. wide, obtuse or rounded apically, obliquely truncate basally, glabrous except usually ciliate, th costa but not secondary veins visible; stipules jetiform, mostly 3-5 mm. long. Inflorescence of axillary, pedunculate heads towards the tips of the twigs; pedun- cles up to 6 cm. long (in age), usually scatteringly pubescent; head small, dense, few-flowered, the flowers all erect; floral bractlets lanceolate, about 2 mm. long, subrigid. Flowers whitish, sessile; calyx cupular-tubular, 2-3 mm. long, glabrous or subglabrous, prominently dentate; corolla of 5 essentially free, obovate, glabrous petals 3-4 mm. long; stamens 10, free, glabrous, the filaments 6-7 mm. long; anthers ovate, almost 1 mm. long, glabrous; ovary linear-oblong, about 3 mm. long, curved, glabrous, bilateral; style about 5 mm. long, expanded apically Into a truncate stigma. Legume linear, mostly 4-6 cm. long and 3-4 mm. wide, flat, glabrous, short-beaked, dehiscent on both valves, the seeds oblique. (286) 195 0] FLORA OF PANAMA (Legtiminosac) 301 Florida and Texas to Argentina; West Indies; Galapagos; probably introduced Isewb eisewnere. CANAL zone: Balboa, Standley 2Jl^Yy S^H^^ cocle: Aguadulce, Vitticr 4970, PANAMA: Matias Hernandez, Pit tier 6go6; Las Sabanas, Standley JoSqj; Bro, Paul 598. A fairly common but inconspicuous weedy plant in Central America, generally flowering during the rainy season. Standley and Steyermark, following Fawcett and Rendle, bave recently recog- nized the synonymity of D. Jepressus and D, virgatus. My observations similarly indicate tbat no reasonably distinct boundary exsits between these two species. Perhaps D. Jcprcssics could be profitably reduced to varietal status under D. vir- gafusy for the name well reflects a tendency within the species-complex towards more diffuse habit, smaller (fewer pinnae) leaves, and less prominent rachial gland. However, I cannot say into which category Panamanian specimens would likely fall. 11. LEUCAENA Benth. Leucaena Benth. in Hook. Jour. Bot. 4:416. 1842. Kyncholeiicaena Britt. & Rose, in N. Am. Fl. 23:130, 1928. Caudoleucaena Britt. & Rose, loc. cit. 1928. Unarm.ed shrubs or trees, glabrous or pubescent. Leaves bipinnate, the pinnae few or several pairs opposite on the rachis, the leaflets few and large or many and small; petiole evident, usually glandular; rachis pronounced, glandular or egland- ular; pinnular rachis often gland-bearing; leaflets linear to ovate, pubescent or glabrous, usually dull. Inflorescence of axillary or racemose pedunculate heads; peduncles usually stout, solitary or fasciculate, ebracteate except at the base of the head; head globular, dense, the floral bractlcts evident, usually peltate. Flowers valvate, white; calyx subcampanulate, markedly sympetalous but 5 -toothed, usually pubescent aplcrlly; corolla of 5 narrow, essentially free petals; stamens 10, free, exserted, the anthers relatively large, pubescent or glabrous; ovary stipitate, gla- brous or pubescent, inequilateral (bilaterally symmetrical) , multlovulate; style thicker than the filaments, gradually expanded into an obliquely truncate stigma. Legume linear to oblong, straight, flat, thin, dehiscent from both margins, the seeds usually nearly transverse. Chiefly New World tropics and subtropics; a few species described from the Far East, and some introduced into various parts of the world. This comparatively small genus, a well recognized segregate of Acacia, super- ficially resembles Albizzia. The flowers, however are very diflferent from those of AlbizziUy and afford the diagnostic criteria for the genus. Only two species arc known from Panama. a. Leaflets small, sublinear, 8—15 mnl. long, about 15 pairs per pinna 1. L. glauca aa. Leaflets larger, nearly ovate, 15—40 mm. long, about 4 pairs per pinna.... 2. L. multicapitula (2U) [Vol. 37 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN L Leucaena glauca (L.) Bcnth. in Hook. Jour. Bot, 4:416. 1842. Mimosa glauca L. Sp. PI. 520. 1753. Mimosa Icucoccphala Lam. Encycl. Meth. 1:12. 1783, Acacia glauca Willd. Sp. PL 4:1075. 1806. Acacia Icucoccphala Link, Enum. 2:444. 1822. Shrub or tree, the branchlets tomentulose becoming less pubescent in age, mod- erately Icnticellate, Leaves large, bipinnate, the pinnae 5-10 pairs opposite on the rachls, the leaflets about 15 pairs per pinna; petioles mostly 3-4 cm. long, pubcru- Icnt, sulcatc or flattened above, usually bearing a sessile, conic gland aplcally just below insertion of the basal pair of pinnae, but this frequently absent; rachis up to 15 cm. long or longer, similar to the petiole, eglandular; pinnae up to 10 or 12 cm. long, the pinnular rachis bearing a small, pateUiform gland aplcally at insertion of the terminal pair of leaflets; leaflets Hncar-elllptic, 8-15 mm. long and 2-3 mm. wide, inequilatcrally acute or subobtuse basally, acute aplcally, subglabrous, dull, usually clliate-rnarglned, the costa (and perhaps 1-2 smaller basal veins) promi- nent, the secondary venation obscure; stipules small, ovate, pubescent. Inflores- cence of large, pedunculate heads, 1-3 axillary from subtermlnal nodes; peduncles about 4 cm. long in age, stout, tomentulose; heads orbicular, dense, almost 2 cm. in diameter, the receptacle expanded. Flowers white; calyx tubular-campanulate, 1-3 mm. long, quite briefly dentate, pubescent aplcally; corolla of 5 free petals, these linear-spatulate, about 5 mm. long, puberulent without; stamens 10, 7-8 mm. lone, <:Iabrous, the anthers ovate-oblong, about 1 mm. long; ovary Inequi- lateral, pubescent aplcally, many-ovulate; style thick, expanding gradually into a broad, obliquely truncate stigma. Legume linear, about 15 cm. long and nearly 2 cm. wide, minutely puberulent, stipltate, briefly apiculate, flat and thin, de- hiscent on both margins, the seeds somewhat obliquely transverse. Tropical America; introduced Into various parts of the world. CANAL zone: Balboa, Allen 2^^6; Staiulley 30QI2. The species Is frequently cultivated. Standley reports a widespread popular belief in Central America that consumption of this species by horses, mules, and pigs will lead to loss of their hair. 2. Leucaena multicapitula Schery, sp. nov. Arbor vcl frutex ramis teretibus primo tomentulosis dcnique subglabrls; foliis magnis distichis bipinnatls, pinnis plcrumque 6 vel 8, follolis 6-10, petlolis 2-3 cm. Icngis tomentulosis subglabrlsve 1-glanduliferis, rhachibus ca. 6 cm, longis, cglanduliferis, foliolis ovatis vel ellipticis, 15-50 mm. longis, 7-25 mm. latls, basi Inequilnteralltcr late acutis apice plcrumque acutis, supra leviter puberulentis subter dense cnnescenti-puberulcntis; Inflorcscentia 3-composita, capitulis ultimis pcdunc- ulatis fasciculatis, pcdunculis tomentosis bracteatis solum subtus capitulos, capitulis orbicularibus ad 1 cm. diamctro crebrls, bracteis peltatis; floribus parvis albis scssili- bus; calycc inl undibuUfcrmi-campanulato ca. 1 mm. longo; petalls 5 lineari- oblongis ca. 2 mm. longis glabris; staminibus 10, libcris 3-4 mm. longis glabris, (2S8) 19.5 0] FLORA OF PANAMA (Legumtnosae) 303 Fig. 102. Leucacna 7nuUicapi/ula anth ens magnis pubescentibus giaonsve, ovano labri inaequilatcrali glabro brcvi- stipitato, stigmatc oblique truncate; legumlne late linear! ca. 12 cm. longo et 3 cm. lato tcnui glabro. Unarmed tree or shrub, the branchlets tomcntulose when young becoming sub- glabrous, moderately lenticellatc, terete. Leaves 2-ranked, large, bipinnate, the pinnae normally 3-4 pairs opposite on the rachis, the leaflets 3-5 (mostly 4) pairs per pinna; petiole about 2-3 cm. long, terete, tomentulose, becoming subglabrous, bearing a sessile, conic gland slightly below insertion of the lower pair of pinnae; rachis about 6 cm. long, similar to the petiole, eglandular or with a sohtary gland at insertion of terminal pinnae; pinnular rachis 2-8 cm. long, puberulent or sub- glabrous, bearing a conic gland apically just below insertion of the terminal pair (289) [Vol. 37 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN of leaflets; leaflets ovate or elliptic, 15-50 mm. long and 7-25 mm. wide, broadly and somewhat inequlLiterally acute basally, acute apically (occasionally rounded on same leaf), dull, densely canesccnt-pubcrulent below (at least In young leaflets) and to a lesser extent above, lighter below than above; stipules caducous. Inflores- cence thrice compound, consisting of several terminal and subtcrminal racemes of few-fasciculate pedunculate heads; peduncles about 1 cm. long, tomentosc, ebracteate except just below the head; head orbicular, less than 1 cm. in diameter, dense, beset (in bud) with pubescent, peltate bracts. Flowers w^hitish, sessile, small; calyx funnelform-campanulate, little more than 1 mm. long, puberulent on the teeth, the 5 teeth about as broad as long; corolla of 5 Uncar-oblong or linear- spatulate petals, these about 2 mm. long, glabrous; stamens 10, free, 3-4 mm. long, glabrous, the anthers ovate, somewhat less than 1 mm. long, pubescent or glabrous; ovary inequilateral, glabrous, short-stipitate, the style stout, the stigma obliquely truncate. Legume broadly linear, about 12 cm. long and almost 3 cm, wide, flat and thin, glabrous. Panama, CANAL zone: Barro Colorado Island, Bailey & Bailey 281; Miratlores, P, White IJj (Gray Herb., type); G. Wbiie 165 (Gray Herb., cotype). darien: RIo Chico, Allen 4642, 50SS. It is not clear just how closely this species is related to other South American species such as L. canescem Benth. and L. frichodcs (Jacq.) Bcnth. or to a num- ber of similar or little-known northern Central American and Mexican species described by Britton and Rose and others, many of them doubtfully tenable. It has been impossible, however, to find anything matching the cited specimens in the larger herbaria of the United States. The Darien specimens are somewhat dif- ferent from the Canal Zone specimens, but it is not deemed wise to consider them yet another species. The species is particularly distinguished by the inflorescence, which forms a thrice-branched terminal panicle of heads. No other species of the genus possesses this very branched Inflorescence, and in some cases the pedunculate heads are merely axillary. Nor Is the dense but fine pubescence of the lower leaf surface In L. multicapitula similar to that of any other species of which I have been able to examine specimens. Most species have glabrous leaves or a looser, longer pubescence. 12. PROSOPIS L. Prvosopis L. Mantissa, 10. 1767. Mitostax Raf. Sylva Tell. 120. 1838, fide Dalla Torre &: Harms. Neltiima Raf. loc. cit. 119. 1838. Pleuronienes Raf. loc. cit. 144. 183 8, fide Dalla Torre & Harms. Spirolohiiim Orblgny, Voy. Amcr. Mcrid. 7, p. 1, Sort. Patagon., ^ /J. 1839, fide Dalla Torre & Harms. Algarohia Benth. Pi. Hartweg. 13. 1839. Sfrnmbocarpa Engelm. & Gray, In Bost. Jour. Nat. Hist. 5:243. 1845. Soprolm Britt. & Rose, In N. Am. Fl. 23:182. 1928. (290) 19 50] FLORA OF PANAMA (Lcguminosae) 305 Trees or shrubs, armed or rarely unarmed. Leaves bipinnate, the pinnae 1 to 4 several pairs, opposite, the leaflets few or many pairs per pinna; petiole prominent, usually glandular; rachis present or obsolete; leaflets generally small, glabrous or less frequently pubescent; stipules small, caducous, modified as spines, or in some species apparently lacking. Inflorescence of axillary, pedunculate heads, spikes, or even racemes. Flowers small, 5-parted, sessile or substipitate, yellowish or whitish; calyx synscpalous, usually shallowly and distantly dentate; petals free or connate near the middle; glabrous or pilose within; stamens normally 10, free, exsertcd; anthers com.paratlvely large, bearing terminally a small or moderate, often caducous gland; ovary pubescent or glabrous, the style stout, the stigma small. Legume very variable, linear to spiral, flat or turgid, indehisccnt, with a pulpy mcsocarp, the endocarp septate or not septate. Chiefly New World from United States to Patagonia; a few species are Asiatic or African. A polymorphic genus, very variable and difficult to delimit precisely, containing many intergrading and difficultly distinguishable species. The genus exhibits a wide range of legume forms, which ordinarily in the Mimosoideae would be oc- casion for generic segregation. Strangely there has been a minimum of such segre- gation by responsible taxonomists, and some of the segregates (as SfrombocarpUy here included among the synonyms) appear to merit generic consideration. Within Prosop/s in the broad sense are found characteristics prominent among a number of different genera: hence it Is not surprising to find species now accepted as Prosopis having been described or included in Mnnosa, Acacia^ DesrnanthnSy etc. Neither the limits of the genus ncr of most of the species have been accurately defined. Bentham (Trans. Linn. Soc. 30:376-3 88) listed 16 species in 4 sections. Burkart (Darwiniana 4:57—128) recognizes 33 species and an additional 2 sections. Britton and Rose (N. Am. Fl. 23:184-187) Kst 14 species (In segregate genus Ncltuma) as occurring north of South America, but It Is generally conceded that few of these are tenable. Recent opinion indicates that but a single species occurs in Panama and all Central America. 1. Prosopis juLirxoRA (Sw,) DC. Prodr. 2:447. 1825. Mimosa juliflora Sw. Prodr. Yeg. Tnd. Occ. 85. 1788. Mimosa piliflora Sw. Fl. Ind. Occid. 2:986. 1800, fide Burkart. Acacia juliflora Willd. Sp. PL 4:1076. 1806. Ncltuma juliflora Raf. Sylva Tell. 119. 1838. Algarobia juliflora Bonth. ex Heynhold. Nom. Bot. Hort. 2:18. 1840, fide Britt. & Rose. Ncltuma Bakcri Britt. & Rose, in N. Am. Fl. 23:185. 1928. Bentham lists 29 additional synonyms for this species (Trans. Linn. Soc. 30:377. 1875), including (under a diversity of the generic names above given) the following specific epithets: affhiis, bractcolata, cumana^ cumanensis, dipt era, domingucnsis^ dulcis, ficxuosa, fruficosa, furcata, glafuUilosaj horrida, incrmis, laevigata, odorata, pallida, salinarum, siliquastrum. Small shrub to moderate tree, the branchlets terete, glabrous, flexuous, ob- scurely lenticellate, usually armed at the "nodes" with prominent paired spines up (291) 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. Z7 to a few cm. long. Leaves arising from very condensed short-shoots, moderate or moderately large, bipinnate, the pinnae mostly 1-2 pairs, opposite, the leaflets few to many pairs per pinna; petiole prominent, most frequently 2-5 cm. long, terete but flattened above, normally bearing a small, sessile, subconic gland apically above; rachis shorter than the petiole or obsolete, glabrous, apiculate terminally from be- low; pinnae 3-10 cm. long, the rachis glandular at the distal 1 to few rachial nodes; leaflets linear to oblong, 5-20 mm. long and 1-5 mm. wide, usually rounded or obtuse apically and basally, scarcely inequilateral, dull, glabrous or subglabrous, the venation remote and moderately prominent below; stipules linear, subsetiform. Inflorescence a slender, elongate, briefly pedunculate, axillary spike inserted on the short-shoots; pedimcles mostly 1-2 cm. long, often sparsely pubescent and minute- ly bracteatc above the middle; spike commonly about 6 cm. long, rather densely flowered, the floral bracts minute. Flowers sessile, whitish; calyx cupular, about 1 mm. long, glabrous or pubescent, the 5 teeth narrow and remote; petals 5, sub- linear, 2-3 mm. long, free (at least in age), pilose within especially distally; stamens normally 10, up to 6 mm. long, free, glabrous; anthers ovate, less than 1 mm. long, gland-tipped (gland often caducous); ovary pubescent; style stout. xceeding the stnmens. Legume linear or lincar-subtorulose, not uncommonly 20 or more cm. long and 8 mm. wide or wider, flattened or turgid, pulpy, indchiscent. In a broad sense United States to Argentina; in narrow sense West Indies, Venezuela, and Colombia. PANAMA: Bella Vista, Sfaudley 2JJQI; Maxotj ^ Yalcufinc 6g$8; Panama Vicja, Allcti S24. darijIn: La Palma, Pit tier 6616. This species is said to be the only one of the genus in Central America. Specific delimitation is exceedingly uncertain, and this difficulty is reflected in the unusually large number of synon)'ms which have accumulated. Certainly tremendous vari- ability in vegetative and legume characters is readily discernible in an)' folder of ''Prosopis jitl/flora^' in any herbarium collection. Bentham could find no defining characters witliin the complex, and hence regarded the species very broadly. More recently Burkart (Darwiniana 4:105. 1940) has concluded that delimitation in the past has been excessively broad, and regards the species in a much stricter sense. However, Standley and Steyermark, with wide experience in the flora of the Amer- icas, question (Fieldiana: Bot. 24:87. 1946) whether P. jnli flora can even be sep- arated from the older P. cLnleus'is (Molina) Stuntz of southern South America. Certainly herbarium sheets labeled P. chilcusis seem, at least upon superficial ex- amination, to fall within the limits of variability exhibited by sheets labeled P. jidiflora. The species, whatever may eventually prove to be its correct name and specific boundaries, Is a locally important plant of xerophytic and subxerophytic regions. In Central America it seems confined to thickets on the Pacific slopes where a marked dry season is usually to be had. It there may reach proportions of a small tree. In its broad sense It is better known, however, as the mcscpiifc of northern Mexico and the southwestern United States, where It is usually of smaller propor- (292) 1950] FLORA OF PANAMA (Lcguminosac) 307 ticns. A great many common names arc had for the species, among which arc found cashaiv and algarroba. The stem and roots provide wood for fuel, or oc- casionally for fence-posts, cross-ties, etc. The wood is reported to finish well, and to be resistant to decay. The pods are pulpy, and the pulp after drying and grind- ing forms an edible meal said to be consumed by Mexican Indians, The pods are excellent for feeding stock. In addition, the plant reportedly provides excellent honey-flowers, a tannin from the bark, and a gum from the trunk. 13. STRYPHNODENDRON Mart. The genus StryUmodendron Mart. (Flora 20:BeibL 2:117. IS 37) is known both from Costa Rica and Colombia, and well may occur in Panam.a although no record from there has as yet become known. The genus consists of unarmed trees, with large, bipinnate leaves and mostly alternate leaflets. The inflorescence is of axillary spikes, the flowers having 10 stamens and gland-tipped anthers. It is similar to Prosopis in having an indehiscent legume. 14. PIPTADENIA Benth. piPTADENiA Bcnth. in Hook. Jour. Eot. 4:3 34. 1842. Stachychrysnm Boj. Hort. Maurit. 114. 1837, fide Ind. Kew. Gohlmauia Rose ex Michell, in Mem. Soc. Bat. Geneve 34:274. 1903, fide Britt. & Rose. Niopa Britt. & Rose, in Addisonia 12:37. 1927. Piiyrocarpa Britt. & Rose, in N. Am. Fi. 23:190. 1928, Shrubs or trees, erect or sometimes decumbent, the branchlcts pubescent or glabrous, aculeate or unarmed. Leaves usually large, bipinnate, the pinnae usually several, multifoliolate, both the pinnae and leaflets usually opposite; petiole nor- mally bearing one or more sessile glands; rachis glandular or eglandular at the pinnular nodes; leaflets almost invariably small; stipules usually slender, caducous. Inflorescence of solitary or few-fasciculate, pedunculate heads or spikes, sometimes appearing paniculate by insertion at distal non-foliate nodes; floral bracts small or minute. Flowers 5-parted, sessile or subsessilc; calyx more or less campanulate, the teeth commonly smal!, glabrous or pubescent; corolla of 5 free (or often con- nate to middle), narrow petals well exceeding the calyx, valvate; stamens nor- mally 10, free except often inserted on a short tubular base or androgynophore, exserted; anthers in all except one or two species (not Panamanian) bearing an apical gland, this usually caducous in age; ovary glabrous or pubescent, often stipitate. Legume broadly linear, flat, not curved, stipitate or sessile, naembra- naceous or coriaceous, 2-valvate, the valve margins straight or constricted between the seeds; seeds compressed, exalbuminous. New World tropics and subtropics; Asia and Africa. A moderate-sized genus, abundant In middle and northern South America. It is not particularly "natural" or clear-cut, and except for the confusion that would result there might be justification in recognition of certain segregates (viz. Niopa). roo-^ ) \ ^^ [Vol. 37 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN ( ^ Certain species of Piptaderjia, at least superficially, mucli resemble otKer genera such as LcHcacri'ty Acacia, and Mimosa. It thus frequently becomes difficult to identify accurately poor or incomplete specimens. The bark of some species is said to be quite astringent, and the seeds are reported highly narcotic. A single species has so far been reported from Panama. 1. PiPTADENiA FLAVA (Sprcng.) Benth. in Trans. Linn. Soc. 30:37L 1875. Acacia flara Spreng. ex DC. Prodr. 2:469. 182 5. PiptaJcrjia siiavcolcus Griseb. Fl. Brit. W. Ind, 710. 1864; nom. nudum, not Miq. PipfaJcfjia Icptocarpa Rose, in Contr. U. S. Nat. Herb. 1:325. 1895. Mimosa biiceragenia B. L. Robins., in Proc. Am. Acad. 43:23. 1907, fide Britt. & Rose. Erect to procumbent small tree or shrub, the branchlcts glabrous or sparsely pubcrulcnt, inconspicuously lenticellatc, often longitudinally striate or ridged, visually sparsely armed with rather small, scattered, recurved thorns. Leaves large, bipinnate, the pinnae 6—11 pairs, more or less opposite, the leaflets many (frequent- ly 25 or more) pairs per pinna; petiole 1—4 cm. long, subtercte, sulcate above, glabrous or pubescent, bearing a shallow, oblong, cupulate gland usually near the middle; rachis up to 25 cm. long, usually pubcrulcnt above, eglandular except sometimes bearing a cupulate gland at insertion of terminal pair of pinnae; pin- nular rachis up to 8 cm. long, usually bearing a small gland at insertions of distal 1-2 pairs of leaflets, pubescent, subalate; leaflets linear, 4-8 mm. long and 1-2 mm. wide, broadly acute apically, asymmetrically rounded or truncate basally, glabrous, the costa excentric baseward, the secondary venation remote, ascending; stipules llnear-setiform, as much as 6 mm. long. Inflorescence of elongate, pedunc- ulate, axillary spikes, these solitary or few-fasciculate, near the tips of twigs arising from nodes not yet foliate and hence appearing somewhat paniculate; peduncles usually less than 1 cm. long, subtomentose; spike commonly about 6—7 cm. long, the axis pubescent, the flowers moderately dense, the floral bracts minute. Flowers sessile, reported yellow; calyx cupulate, about 1 mm, long, 5-lobcd, glabrous; corolla of 5, free, nearly linear petals about 3 mm. long, glabrous; stamens nor- mally 10, free, inserted atop a short tube or androgynophorc, 5-6 mm. long; anthers broadly oblong, almost 1 mm. long, bearing an apical gland, glabrous; ovary glabrous, conspicuously stipitate, apparently abortive in some flowers; stigma narrow. Legume broadly linear, 5—9 cm. long and about 15 mm. wide, thin, glabrous, glandular when young, the surface irregular because of maturing seeds "but not constricted between the seeds. Mexico to northern and western South America, cocLife: Aguadulce, Pit tier 48go, Vegetatively (or even in fruit) P. flava resembles species of Leucaena and Acacia. The elongate spike, however, tends to distinguish it from similar species of these other genera in Panama. P. viridiflora and/or P. communis may with comprehensive study prove to be synonyms of this species. (294) 1950] FLORA OF PANAMA (LegMmhiosae) 309 15. ENTADA Adans. Entada Adans. Fam. Pi. 2:318. 1763, nom. conserv. r Gigaloblinn P. Br. Nat. Hist. Jamaica, 362. 1756. PerimaV.^i, Sylva Tellur. 118. 1838. Stnpsilobus Raf. loc. cit. 117. 183 8. AilefwpoJia Presl, Epim. Bot. 206. 1850, fide Hook. Jour. Bot. 4:286. 1852. Pusactha [L. 1747] Ktze. Rev. Gen. PI. 1:204. 1891. Entadopsis Britt., in N. Am. Fl. 23:191. 1928. PscuJocritada Britt. & Rose, loc. cit. 1928. Woody vines or climbing shrubs or trees, usually very large, unarmed (in Panama) or aculeate, sometimes cirrhiferous. Leaves large, bipinnate, tbe pinnae few pairs (in Panama) or less frequently several to many, opposite, the leaflets few to many pairs; petiole and usually the rachls eglandular, the rachis sometimes terminating in a bifurcate tendril; leaflets large or small; stipules small. Inflores- cence of usually elongate spikes, these soUtary and borne from foliate branches or clustered terminally in a large raceme; floral bracts minute. Flowers small, usually briefly stipltate, 5-partcd, valvate, variously colored, often with a disagreeable odor; calyx small, synsepalous, broadly and shallowly cupulate; corolla of 5 free or slightly coherent petals; stamens 10, free, briefly exserted; anthers tipped with a (caducous) gland; ovary several- to many-ovulate; style slender; stigma terminal, truncate. Legume often very large (Panamanian species), flat, straight or curved, the valves in time breaking Into segments, the margins continuous and persistent. At least one species pantropical; others chiefly African and tropical American. A comparatively small genus of less than 20 species, many of which are dis- similar to the others and a cause for past generic segregation. It seems scarcely advisable, however, to accept division of Entada into several smaller genera based on these differing species, for such a procedure, if followed generally here and in the MiMOsoiDEAE as a whole, could continue ad infinitum until practically no "genus" could be readily located. The Panamanian species of Entada suggest, at least superficially, giant liana-type Mimosas, but can be readily distinguished by the long spikes of flowers which are not found in any of the Panamanian species of Mimosa, The leaves are sometimes reported sensitive, folding following dis- turbance, such as Is the case with a number of Mimosa species. Two species of Entada are known to occur in Panama, both distinctive vines of lowland areas. a. Inflorescence a supra-axillary spike; leaves often cirrhiferous; legume very large, the seed 3—5 cm. in diameter 1. E. gigas aa. Inflorescence a terminal raceme of spikes; leaves seldom with tendrils; legume smaller, the seeds only about 1 cm. long 2. E. polystachya 1. Entada gigas (L.) Fawc. & Rendle, Fl. Jamaica 4:124. 1920. Mimosa gigas L. PL Jam. 22. 1759. Entada gigalobium DC. Prodr. 2:424, 1825. Large vine of lowland areas, often climbing to the tops of forest trees, tKe branchlets essentially glabrous and inconspicuously lenticellate. Leaves very large, bipinnate, commonly cirrhiferous, the pinnae usually 2 pairs, opposite, the leaflets (295) [Vol. Z7 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3—6 pairs per pinna; petiole usually 2-4 cm. long, eglandular, frequently puberu- lent, callous basally; rachis similar to the petiole, up to 20 cm. long, eglandular, commonly terminating in a stout, branched tendril, especially on new growth; pinnae up to 15 cm. long, the axis similar to the rachis; leaflets asymmetrically oblong, the terminal ones as much as 8 cm, long and 3 cm. wide, the basal ones as small as L5 cm. wide and 1 cm, broad, usually blunt or even cmarginate apically, inequilaterally obtuse basally, glabrous except on the costa, somewhat glossy above, dull below, the venation conspicuous, the secondary lateral veins essentially parallel; stipules linear, about 4 mm. long. Inflorescence an elongate spike, supra-axillary (usually inserted 5-10 mm. above the axil), the peduncular portion only 1-5 cm. long, the floriferous portion 7-25 cm. long, the floriferous bractlets squamiform. Flowx^rs cream, odorous (often described as unpleasantly so, or with a "sickly odor"), short-stipitate; calyx shallowly cupulate, about 1 mm. deep, rather Ir- regularly 5-lobed, glabrous; corolla of 5 narrowly elliptic petals about 3 mm. l^'^S^ glabrous, free or somewhat connate basally; stamens 10, 6-7 mm. long, the anthers usually bearing a (caducous) gland; ovary glabrous. Legume very large, as much as 2 m. long (commonly several dm. long) and about 1 dm. broad, more or less oblong, curved, flat, transversely jointed but the margins continuous, glabrous, the segments 1-seeded; seeds orbicular, 3-5 cm. in diameter, flattened, blackish, glossy. Central America, northern South America, and West Indies; west Africa. BOCAs DEL TORO: Bocas del Toro, Read s, n. canal zone: Barro Colorado Island, Stafullcy 31264; Stevens 416, Panama: San Jose Island, Erlanson Ij, 82; Jolmdon 6oo^ A tendrilcd vine, very remarkable for the enormous size of the legume, prob- ably not equalled by that of any other species of the family in this hemisphere. It is the so-called "sea-bean," the shiny black seeds of which are carried long distances by ocean currents. The species is a lowland plant, frequently found near the sea- shore, and its wide distribution can perhaps be explained by discharge of the seeds into the ocean. Dr. I. M. Johnston (mss.), who studied "sea-beans" intensively during the war years, shows that on the basis of the fruit £. gigas Is quite distinct from the Old World £. pbascohides {^Mimosa Entada L. Sp. Pi. 518. 1753, fide Merrill; Lens pbaseolohles L. In Stickman, Herb. Amboin. 18. 1754; Mimosa scan d cm L. Sp. Pi. ed. 2, 1501. 1763; Adenanthera scandcjis (L.) Forster, Fl. Ins. Aust. Prodr. 33. 1786; Acacia scandens (L.) Willd. Sp. PL 4:1057. 1806; Ejitada Pursaefba, E, momntachyay and £. Adcfianthera DC. Prodr. 2:424. 1825, fide Bcnth.; E. Khccdii and E. Parr ana Spreng. Syst. 2:3 25. 1825; Strepsilobns scandots (L.) Raf. Sylva Tell. 118. 1838; Entada scanJcns (L.) Benth. in Hook. Jour. Bot. 4:332, 1841; E. Gandn Hoffm. ex Walp. Rep. 1:858, 1842, fide Benth.; £. Rump/m Scheff. In Natuurk. Tijdschr. Ned. Ind. 32:90. 1872, fide Benth.; Gigalobiiim scandens (L.) Hitchcock, in Rept. Mo. Bot. Card. 4:82. 1893), a name heretofore applied to a group of similar species and including the American £. gigas. (296) 1950] FLORA OF PANAMA (LegMmlnosae) 311 Fig. 103. EfitaJa polystachya Entada polystachya (L.) DC. Mem. Legum. 434. 1825. Mimosa polystochia L. Sp. Pi. 520. 1753. Mimosa bipinnafa Aubl. PI. Guian. 2:946. 1775. Mimosa cauJafa Vahl, Eclog. 3:3 5. 1807, fide Benth. Mimosa chiliantha G, F. W. Mey. Prim. Fl. Esseq. 163. 1818, fide Benth. Adcnanthera BouplafuUafia Kunth. in HBK. Nov. Gen. & Sp. 6:311. 1824. Acacia caudata DC. Prodr. 2:456. 1825. Entada Plumieri Spreng. Syst. 4": 164, 1827. Entadopsis polystachya Britt. in N. Am. Fl. 23:191. 1928. Large, woody liana up to 15 or more m, tall, the branchlets glabrous, longi- tudinally striate, the old stems rather shaggy-barked. Leaves large, bipinnate, the pinnae commonly about 4 pair, opposite, the leaflets mostly 5—8 pairs per pinna; (297) [Vol. 37 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN petiole mostly 4-8 cm. long, eglandular, glabrous, flattened above; rachis similar to the petiole, eglandular; pinnae usually 6-15 cm. long, the axis often puberulent above; leaflets oblong, 15-40 mm. long and 5-18 mm. wide, usually rounded aplcally and obliquely so basally, glabrous or sometimes sparingly pubescent below or along the margins, glossy above, the venation prominently reticulate; stipules linear, small, caducous. Inflorescence a large, terminal raceme (often 2 dm. long) of slender spikes, the latter 6-7 cm. long, inserted singly or in pairs; floral bract- lets not apparent; axis usually puberulent. Flowers small, very briefly stipitate, reported reddish, pinkish, whitish or greenish, with a disagreeable odor; calyx shallowly cupulate, scarcely 1 mm. deep, briefly dentate, glabrous; corolla of 5, free, narrowly oblong, glabrous petals about 2 mm. long; stamens 10, somewhat exceeding the petals, glabrous, the anthers (at least in youth) gland-tipped; ovary glabrous. Legume large, oblong, commonly 25-30 cm. long and 6 or 7 cm. wide, curved, flat, thin, glabrous, lustrous, the valves in time breaking into segments, the margins persistent; seeds comparatively small, oblong, no more than 1 cm. long. Mexico to middle South America; West Indies. BOCAs DEL TORO: Chiriqui Lagoon, H. von Wedel 1062, ^^^4, ^495, ^743- canal zone: Frijoles, Bro. llcriherto II J; Miraflores Lake, P. White 2^6, hfrrera: Peso, Allen 804. Panama: R. La Macstra, Allen J4; Juan Diaz, Sfandley 30^18. The species is an interesting tall vine of the lowland areas, with a large, term- inal, disagreeably odoriferous inflorescence and large, persistent legumes. Macerated root or stem reportedly produces a soap-like lather with water, suitable for washing. 4 I 16. PENTACLETHRA Benth. Pentaclethra Benth. in Hook. Jour. Bot. 2:127. 1840. Trees, the branchlets coarse, the leaves few-ranked. Leaves very large, bipin- nate, the pinnae many pairs, opposite, the leaflets many pairs per pinna; petiole and rachis eglandular; leaflets small, numerous, inequilateral or falcate; stipules small. Inflorescence of stout, elongate spikes, apparently ebracteate. Flowers moderately small, 5-parted, rather coriaceous; calyx cupulate, the rounded lobes somewhat imbricate in bud; corolla of 5 petals connate below the middle, valvatc; stamens 5, cxsertcd, united below (with stnminodia) into a very short tube; staminodia 5 (in Panama), 10 or 15, usually about 3 times as long as the stamens; anthers rela- tively large, gland-tipped; ovary subsessile, the stigma expanded, terminal. Legume large, flat, thick, woody, broadest above the middle, elastically 2-valvate, longi- tudinally striate. Tropical America; Africa. World It is allied to Park/a (South American and Old Wo subcapitate inflorescence and 10 fertile stamens. (298) 19.5 0] FLORA OF PANAMA (Leguminosac) 313 Fig. 1 04. Fentaclcthra macroloba 1. Pentaclethra macroloba (Willd,) Ktze. Rev, Gen. PL 1:201, 1891. Acacia macroloba Willd. Sp. PL 4:1060. 1806. Mimosa macroloba Poir. In Lam. EncycL Meth. SuppL 1:66. 1810. Acacia aspidioiJcs G. F. W. Mcy. Prim. FL Esseq. 165. 1818, fide Benth. Pentacletlyra filamentosa Bcnth. in Hook. Jour. Bot. 2:127. 1840. Pentaclethra brcvipila Benth. loc. cit. 12 8. 1840, fide Benth. Caillea macrostachya Steud. in Flora 26:759, 1843, Entada Wrbcana PresI, Epim. Boc. 206. 18 50, ex char, fide Benth. Unarmed tree, the branchlcts stout, longitudinally striate, puberulent when young, callous in age. Leaves very large, bipinnate, the pinnae many (usually 15—20) pairs, opposite, the leaflets many pairs per pinna; petiole 1—5 cm. long, eglandular, terete except shallov/ly sulcate above, callous basally; rachis as much as 3 dm. long, pubescent above, eglandular; pinnae 2-10 cm. long, eglandular; (299) [Vol. 37. 1950] 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN leaflets small, linc\u--subfalcate, mostly 6-9 mm. long and 1-2 mm. wide, sharply acutc-acuminate apically, obliquely rounded or truncate and more or less auricu- latc on the proximal side basally, essentially glabrous, darker and lustrous above, the venation fairly prominent below; stipules small, linear. Inflorescence of few or several terminal and subterminal, elongate spikes; peduncular portion of spike 1-5 cm. long, pubescent; florifcrous portion usually 15 or more cm. long at maturity, pubescent, ebracteatc. Flowers dense, sessile, 5-parted, white; calyx cupulate, 1-2 mm. long, the broad, rounded lobes shorter than the tube and some- what imbricate in bud; petals linear-elliptic, about 5 mm. long, usually coherent for about the lower third or half of their length, glabrous, valvate; stamens 5, 6-7 mm. long, alternating with 5 thin, elongate staminodia about 2 cm. long, stamens and staminodia briefly united at the base; anthers about 1 mm. long, bearing a prominent apical gland or appendage; ovary subsessile, pubescent; style slightly exceeding the stamens; stigma expanded. Legume llnear-subspatulate, 2-3 dm, long and up to 4 cm. wide, flattened but thick, ligneous, narrowed baseward, lustrous, glabrous, longitudinally striate, 2-valvate, the valves coiling backwards after dehiscence. Southern Central America, West Indies, north-central South America. BOCAS DFi. TORO: Changulnola Valley, Cooper ^ Slater 126; Guabito, Skutcb IT. dari6n: El Real, Pit tier 6jOO; Pinogana, Allen pjj. A species quite different from other Mimosoideae in Panama, apparently con- fined to the lowlands. The large spikes of white flowers and the unusual legum.e make P. m-acroloba quite striking. (Lcgumtnosac to he cou eluded w Part V, Fasc, j) (300) F THE U -fc ¥' N Dire€i<^ Gechig£ T. Mooue t.v VOK SCHEENX Fatliologist Jhsse M. Grebnmabt, Curator Emttnt^^B -at the Herbarium Car-rgll W, Dcdg-Es Edgar x^NBEaso^'^ Geneticist Robert E, Wr-"isoN, Jr. Curator of th^ Hey1:aritms Paleobota. i-*-A -.- w Gekaiu> Uulio GUSTAY A, I Research Horticultusist RoLiA %L Tryok, Assistant Cnt^t^r of the Herbanam Georgs B- Vak Schaaoc, Hoaorarr Curattrr of G«tH5Ci JuriAH A. SXEYEaMAlUt, __ HcnonuT Itea«a5rli Asawjckte Nell C^Hornb% Librarian snd Editor of Fofelicstiona Busiiaess OAHD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN Presidemi Dahibl IL L* Rat Cartes DU0I.ET J0HK S. LeHMANH A. 'L»GM" Joseph M. Daest, » riXIAM g|. :#r> ^ ^ % WitttAM i'rf ^ .^, |.^«i»' WfWW* Ml lOMit- ? ^ - r -. ^ - L . - y^ V, Volume XXX V 11 Number 3 nals of the X950 ce m& can bpecies of Tritixnfett& I^.. Carfa «^ « '^ rO. It: -V. * *. ^ ■»t «■ « 337-40 ■-I Tl ^ > i:a.ti0iis * *^ dK ^V^" f f " L ^ mm QF TSXS^TM .MIS r J. ^^a^r^sd. wm^- m^&ss^nfy^^ ms^w^ m --iis«- •'-. iC^- "^ _ ^ - . \. /\nnci CI the J issoun Jotanical Garden A Qi^arterly Journal containIi:g Sci, :ti£c Ccntrib\itic--s from ^ ! ^_o, J^JBsscmri Botanical Garden and gton UmTemty Garden rleary in altiu^tion with Informatmn ^v^ ^.-;.-t^* Scnool ot Bct.;r»v of Mis:.:: ::ri Botanica. Tfe.e An-NALS " OF THB dtmiig' tW calendar -r-esr: Z TOil Ma" Set^-tembec, and pTCa^ four "t"" ones Four Salm:.dptioB ^.SlO, t^=^ ^mgie JSt*5A? ^r'i-Cll per vomir.e ^S.WBEi^' a-re listed in ti:^ Agricaim:rai ^Indes, jyuHish^d by tfae H* W- "Wllsoa T f ^ l^ . -1^ ^^i^^^^t:"; ■ . -. -t - . ^^ ,' C- L" ■ - " JM' ' ■_^. : ^■". _.>. Annals of the Missouri Botanical Garden Vol. 37 SEPTEMBER, 1950 No. 3 THE AMERICAN SPECIES OF TRIUMFETTA L.^ KO KO LAY = Introduction The genus Trhnnfetta, one of the largest of the Tillaceae, is pantropic in its distribution; the total number of valid species in the genus may approximate 150 or more. Species are especially abundant in Latin America, but extend from tropical Florida through the West Indies, Mexico, and Central America, southward to Argentina. The western-hemisphere species of the genus have received no special attention from systematists. K. Schumann's^ treatment of the South American species, though still a standard reference, is however pathetically out of date. Standley's'* classification of the Mexican species Is generally more acceptable, but an array of new species has been established since his publication. About 150 species have been named for this hemisphere, and it was thought desirable to monograph the genus for the New World in order to establish the proper status of the species. It is realized that the present splitting of the genus with regard to its occur- rence in the eastern and v/estern hemispheres, though convenient, is definitely not natural. At least three species which are represented in Latin America are also known to occur In the eastern hemisphere. There is a possibihty that other species may have similar distribution. Since this study was undertaken with a view to clarify the species of the western hemisphere, the total range and the remaining species of the genus have of necessity been Ignored. ^An investigation carried out In tlic graduate laboratory of the Henry Shaw School of Botany of Washington University and submitted as a thesis in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Washington University, -Lecturer in Botany, University of Rangoon, Burma. Formerly Burma State Scholar in the Henry Shaw School of Botany of Washington University. ^K. Schumann, in Mart. Fl. Bras. 12''*:131. 1886. ^Standley, in Contr. U. S. Nat. Herb. 233:741. 1923. Issued September 30, 1950, (315) [Vol. 37 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN A large quantity of specimens from nearly all the major herbaria of Europe and of both North and South America has been available for the study. About fifty plants of three species were also grown in a greenhouse, but the living plants were not amenable to such conditions and though they grew steadily none flowered. Hence this study is based entirely on the standard herbarium methods. In tryhig to understand the morphology and to clarify the basic species con- cept in the genus, I realized, as with some previous systematists/'''* ^' that the char- acters derived from the fruits are easily ascertainable and highly constant for most species. After a primary classification of all the fruiting specimens, an attempt was made to correlate them with the flowering ones. This is rendered imperative, as it is but rarely that one finds both the fruiting and flowering materials on the same specimen. It soon became apparent that certain species are exceedingly distinct in both fruit and flower, while others show considerable variation in the floral characteristics. The species that show the greatest diversity in floral char- acters are also found to possess extremely polymorphic leaves. To this category belongs the semitriloba complex, wherein the greatest number of species have been named. Mass collections, kindly placed at my disposal by Dr. Edgar Anderson, were studied for two species: T. semitriloba and T. CaldcrouL Though no conclusive proof was arrived at, the study gave suflFicient indication that introgression be- tween the species is rampant. The species T, semitriloba tends to approach T. Lapljuld, on the one hand, in its characteristic, more or less pandurate leaves and very condensed nodose cymes by the foreshortening of the peduncles and pedicels; and, on the other, T. bogofcnsis by its larger flowers, more showy petals, more densely pubescent fruits, and also by the presence of short simple hairs on the leaves. The variation in the types of inflorescences, and especially the abortion of spines on the surfaces of the fruits (with their presence only on the lateral mar- gins) , tend to merge T. Calderoni with the genus Heliocarpus. Were it not for the presence of the urceolus, certain specimens of T. Calderoni would definitely break the generic separation between Heliocarpiis and Triumfctta. Despite the great quantity of material available for study, at least a third of the species are very imperfectly known; six species being represented only by the types. The genus further presents a problem in distribution, a great many species being strict endemics while a few are pantropic weeds. A number of species of widespread distribution, as T. semitriloba, T, bogoteiishy T. Bartramia and others, apparently have an infinite capacity for variation in minor characters of leaf size, shape, and indument. The extremes often seem quite dis- tinct in themselves but in a study of large numbers of collections they appear to lose their seeming significance. Wherever such intergradations have been found, I have reduced a number of previously described species to synonymy. At the same time I have described a number of novelties which in my opinion are new to "Masters in Oliver, Fl. Trop. Africa 1:254. 1868. ^Spraguc Si. Hutchinson, in Jour. Linn. Soc. Bot. 39:231. 1909. 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 317 science. With greater abundance of material in the future and intensive study of the genus further reduction may be in order. History The genus Trinmfetta was proposed by Plumier"^ in 1703, in honor of John Baptiste Triumfetti of Bolognc, doctor in medicine, lecturer in botany, and direc- tor of the botanical garden at Rome. Linneaus^ in 1737 adopted the genus and described both calyx and corolla, which were depicted in Plumier's rough illustra- tion. In the same year Linneaus^ gave a detailed description under the same generic name of a plant growing in Clifford's garden, received from either Jamaica or Bermuda, and described it as apetalous. In 1742^*^ he altered the description of the genus and stated that it lacked the calyx (corolla). In the first edition of 'Species Plantarum'/^ the description In 'Hortius CUffortianus' was cited for T. LappuIiJj the only species named under the genus. Linneaus had already estabhshcd the genus Bartraviia^- which, according to Fawcett and Rendlc, was based on a plant In Hermann's Herbarium (now in Herb. Mus. Brit.) with both calyx and corolla. The species Bartrainia indica was pub- lished In 'Species Plantarum' on an earlier page (p. 3 89) than Trinmfetta Lappula, As synonyms under B. ifulica Linneaus cited, among others, one of Plukcnet's"^^ drawings which is rather aptly done and quite diagnostic. Six years later, Lin- neaus^^ included Bartramia in Trhnnfetta and altered the trivial by naming the species T. Bartramia, De Candolle^'"\ being unable to determine the identity of T. Bartramia on the ground that the synonyms cited include many diverse species, adopted Jacquin's name T. rhomboidea}^. Later botanists followed De Candolle till recently when Fawcett and Rcndle^'^ established that T. Bartramia and T. rhomboidea are con- specific. It is, however, to be noted that the earliest trivial is iiidica, but as Lamarck^^ subsequently described T. indica, the identity of which is doubtful, it is desirable to adopt the Linnean name T. Bartramia, Gaertner^**^ maintained the two genera as distinct and further distinguished them by the presence of indivisible fruit with 1 -seeded loculi in Trinmfetta, and divisible fruit and 2-seeded loculi in Bartramia. The fruit of T. Bartramia is not really dehiscent, but it could be made to separate into component valves by prolonged boiling. "^Nov. Pi. Amer. Gen. 40, t. 8, 1703. **Gen. PI. ed. 1. 344. 1737. "Hort. Cliff. 210. 1737. ^^Gcn. PL cd. 2. 243. 1742. "Sp. PI. ed. 1. 444. 1753. ^-Fl. Zeylanica, 77. 174S. ^•"^ Almagest. Bot. 206, /. 41, f. 5. 1691. ^■^Syst. PI. ed. 10. 1044. 1759. ^^Prod. 1:508. 1824. =^^Enum. Pi. Carib. 22. 1760. ^^Jour. Bot. 59:224. 1921. ^^Encyc. Meth. 3:420. 1789. ^^Fruct. & Sem. 2:137. 1791. [Vol. 37 "^ 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN De Candollc"^ finally united the two genera and established two sections Bartramia and Lappula based on Gaertner's diagnosis, where Lappula is sep- arated from Bartramia by the lack of petals and by its indivisible fruits. End- licher^i adopted the sections of Dc Candolle but defined them entirely on the fruit characters- Baillon"^ substituted the name Eutriumfetta for Lappula and still distinguished it from Bartramia by the indchiscent fruit. K. Schumann^^ was dissatisfied with the classification based entirely on the characters of the fruit. To him the fruits (and also the leaves) were extremely polymorphic and did not offer any constant characters. He redefined the sections, basing his classification entirely on floral characters: Eutriumfetta being apetalous and without a glandular gonophore, and Bartramia with manifest petals and glandular gonophorc. Sprague and Hutchinson^* disagreed with K. Schumann's classification* as it Lappiila. Tl ruits certain characters derived from the indument of the sepals. According to them, the genus is divided into the following four sections: 1, Lepidocalyx: Calyx scaly without; stamens numerous (25-60); ovarj^ 10- oody 1 -seeded, the spines fusiform. 2. Propa: Calyx not scaly; stamens numerous (25-40); ovary 6- to 10-loculate, each with 1 ovule; fruit spheroid, woody, 6- to 10-loculate, each 1 -seeded, the spines constricted at the base and the tip. 3. Lasiothrix: Calyx not scaly; stamens usually numerous (20-50), rarely few; fruit spheroid, not woody, very light, indehiscent, often 1 -seeded, the spines weak, with plumose tip. 4. Lappula: Calyx not scaly; stamens numerous or few; fruit spheroid or ovoid, dehiscent or indehiscent, many- or 1 -seeded, the spines rigid, the tips rarely P 1 umose. Since the above classification, there has been no appreciable study of the genus as a whole. Hochrcutiner's^^ work Is more In the nature of reorganization of the tribe Grewleae than of the genus itself. Geography Trlnmfetta is pantropic in Its distribution. There are 43 species in the western hemisphere, about 50 in Africa and 10 In Australia. The number of valid species in Asia Is not known, but may approximate 40 or more. T. Bartramsa and T. LaplmJa are equally at home in both the eastern and western hemispheres, T. sonitriloba Is abundantly represented in the western hemisphere and to a lesser 20Loc. cit. 506. 1824. 2iGcn. PL 1008. 1840. 22Hist. Pl. 4:195. 1873. ^^Marc. Fl. Bras. 123:131. 1886. 2'' Jour. Linn. Sac. Bot. 39:233. 1909. 2^"^ Ann. Conscrv. Jard. Bot. Geneve 18:92. 1914. 1950] KO KO LAY AMERICAN TRIUMFETTA 319 34, T. SEMITRILOBA 30. T. BOGOTENSIS «6. T. UPPOU 27. T. GftAMDIFLORA i9. T. ABUTILOIOES 39. T. BARTRAMIA 6. T. fiOUMANt 26. T. DISCOLOR Fig. 1, Geographical distribution of species of Triumfctta, extent in Africa and eastern and soutKeastern Asia. In the western Iiemisphcre the species extend from tropical Florida to the West Indies, Mexico, and Central America southward to northern Argentina, Species are especially abundant in Mexico and Central America, There are six species with very extended ranges found in both North and South Americas. The three very closely related species, T. bogotemis, T. semi- triloba, and T. Lappnla, have very similar ranges, each extending from Mexico and the West Indies through Central America southward to northwestern South America, finally to central and eastern South America. T. graiidiflora, one of the more primitive species, has similar north-south distribution, but is rather sparsely represented throughout Its range. T. Bariramia, whose affinities with the western- hemisphere species Is rather doubtful, has an altogether distinct distribution. It 320 [Vol. 37 MISSOURI 41. T. PAMICULAU 42. T. GALEOTTIANA 32. T. ACRACWTHA ) 4. T. CUCULLATA 5. T. CORIACEA 17. T. PALNERI 25. T. BARBOSA 26. T. COlUMt^ARlS 8. T. UINTONII 10. T. FERRUSIMEA 14. T. ATTErSUATA 15. T. (KXIAE 16. T. BREVIPES Fig. 2. Gcograpliical distribution of species of Triumfvtia. is known in the West Indies nnd eastern South America, and except for one locality In British Honduras and in Florida, where introduced; It is not represented on continental North America. T. abutiloides approximates the distribution of its closely allied species T. bogotctnh, except that it Is not known in continental North America. There are nine species known only from South America. T. althacoldes, which frequents lowland sandy areas, is austral In its distribution, and is found exten- sively in northern South America wherever suitable habitat is available. T, obsaira, which is the counterpart of T. acracantha from Mexico, is confined to south- eastern Brazil. T. Sampaioi, a species closely related to T. Barfram'ia, is known only In the lowlands of northeastern Brazil. The three closely related species, T. hn\;/coma, T. mulf'docularts, and T. wollissima, are found exclusively in south- eastern Brazil, Peru, and Colombia respectively. T. candata, T. senrata, and T. pcrsimilh occupy narrow ranges in Colombia. J 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 321 ©1. T. STZLLATA •2. T, POLTAKDRA 3. T. SPECIOSA ►3. T. CALDERONI 11. T. ARBORESCENS 33. T. ALTHAEOIOES 21, T. MULTILOCUUSIS • 7. T. CAUDATA •37. T. SERICATA • 23. T. L0N6IC0HA *31. T. OBSCURA 22. T. MOLLISSIMA 40. T. SAMPAIOB Fig. 3. Geographical distribution of species of Triumfctta, The majority of the species are found in Mexico, and among them are a few that extend into Central America. The two more primitive species, T. polyainfra and T. spcciosa, though found sparsely in any particular locality, extend southward into Central America. Most of the species are known only from central and western Mexico. T. discolor, T. Goldmani, T, acracantbay T. pantciihta, and T. Galeottianu are the most prominent species of western Mexico, while T. hrcripcs^ T. VaJiucri^ and T, columuarls are found rather abundantly in central and south- western Mexico. The rest of the species are endemics, being known from very restricted areas. The series Geniculatae is best represented in Mexico and Central America. Of its fifteen species only two (T. candata and T. persim/Iis) are from Colombia. Because of the presence of the majority of species in central Mexico and northern Central America, including most of the primitive ones and also many very re- stricted endemics, it is assumed that the primary centre of dispersal of the species is probably from that area. [Vol. 37 322 ANNALS OF THE MISSOURI Morphology ■fetta trees or slirubs. The series Geniculatae constitutes species wherein the tree habit is most predominant, although no tree in the genus is known to exceed 5 m. The stems are terete in all the species except T. paniciilata where they are ridged or irregularly angled, the ridging occurring as a result of dccurrent petiole bases. Certain plants, especially those in the seint branch at almost every node of the main axes, while others (as represented by T. Caldcronl) are sparsely, if ever, branched. The bark is prominently lenticellate, with cither small white or large brown lenticels. The leaves are petiolate. The length of the petiole is usually correlated with the shape of the lamina, the broadly ovate leaves being long-petiolate- while the large elliptic ones are shortly so. T. obscura is an exception as it has elliptical leaves which are long-petiolate. The lobing of the leaf generally is a good primary character in certain species, but in others, such as T. scmitriloba, T. bogotcmis, and T. Bartramia, the general leaf-shape Is so variable that it is relatively useless as a definitive character. In general, weedy species of the genus have variously indented or lobed leaves and show much greater variation than the non-weedy ones. The margin is serrated, the serrations usually being very Irregular. Most species have small glands on the basal serrations from which the mucilaginous sap exudes. In three species, T. althacohlesy T. cncullata, and T, corjacca, these glands are found on the petioles at the basal sinus. Because of their extreme constancy they offer an easy means of identifying these species. The term "bract" has been used in this paper to mean the terminal leaves of a branch which subtend the cymes. The bracts arc usually very prominent in plants with axillary inflorescences. They are invariably smaller and more shortly petioled than the vegetative leaves and are generally elliptical in outline. No measurements of the bracts are given, as it is difficult to determine from dried material whether they have attained their maximum growth or not. AH species have more or less coarsely pubescent leaves, except T, acracantba and r. obscura which have glabrescent ones. The leaves usually possess three tiers of induments, two of which consist of stellate hairs, one closely appressed to the surface and the other usually much larger and spreading irregularly, the third tier of simple hairs which are usually much longer and generally more prominent. The nature of the Indument offers good primary characters in certain species. How- ever, the use of this character requires great judgment as the pubescence changes considerably during the course of development. Young leaves may be covered rather densely with both stellate and simple hairs, while the mature ones on the same plant may have hairs of one kind only. The leaves are stipulate. The stipules are usually large and linear and tend to dry from the tip to the base. They are rarely caducous; in most species they 1950J KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 323 r persist even after the shedding of the leaves. No diagnostic importance has been attached to the stipules. The inflorescence consists of compound aggregate dichasia. In the more primitive species, the cymes, each consisting of 1—3 cymules, are borne in the axils of the bracts. In the more advanced species the cymes are terminal and consist of a shortened sympodial branch system, each bearing 4-6 cymules. There is usually more than one cyme per bract, the cymes being clustered rather densely at the nodes, and the whole inflorescence is usually large and spreading. In both types of inflorescences the cymule is either 2- or 3 -flowered. Each cymule is sub- tended by a pair of fugacious bractcoles. The number of flowers per cymule is not constant even on the same inflorescence, probably due to the abortion of one of the lateral flowers in early ontogeny. The inflorescences are hermaphrodite or, in a few species, gynodioecious. The species of the genus have considerable range in the size of flowers. Slightly before anthesis the floral buds of T. polyandra and T. speclosa are as large as 3-4 cm. in length, while those of T. Lappida and T*. hcliocarpoides rarely, if ever, exceed 6 mm. The size of the flower is generally a good constant character for most species. However, in T. bogotensis the length varies from 7 to 13 mm, and Is not correlated with the variations in either the fruit or vegetative characters. The floral buds in the majority of the species are oblongoid, but in the Galcottiana and Caldcroni complexes they are obovoid in the hermaphrodite flowers. The sepals afford good specific characters. All plants of the genus bear a sub- apical appendage on the sepals, the lengths of which are generally constant for a given species. T. Vurpiisii and T. falcifera possess the longest appendages, usually about 5 to 7 mm. long, and T, Caldcroni the shortest, being nearly obsolete. Unless otherwise mentioned, the lengths of the sepals given in this paper include the apical appendages. The indument of the sepals may also be used as a primary character in certain species. However, as with the pubescence of the leaves, its use in the identification of a species needs caution. There is a tendency in the genus towards a completely apctalous condition. The more primitive species, such as T. polyandra, T. widfilocnlaris, T. hrevipes and others, have large showy petals, while the more advanced species (T, Lappula^ T. Galeottiaua, and T. hcliocarpoides) are either apetalous or possess very small ones. The petals have a short ciliate claw, which is hairy on the outside and some- times has a transverse band of stellate hairs on the inner surface at the base of the blade. The androgynophore or gonophore, with its five spherical or elliptical glands, contributes some important diagnostic characters. The shape of the glands, their size, and whether they are contiguous or otherwise, are usually constant for a species. Because of their extreme smallness, these characters, though mentioned in the specific descriptions, have not been used in the key. The gonophore is slightly accrescent in fruit. T. Lappiila is the only species known which does not possess a glandular gonophore. [Vol. 37 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN The urceolus Is a small, ratlicr membranaceous ring with a cihate margin, which is borne on the gynophore and completely surrounds the base of the stamens. It is usually 5-lobed, sometimes 10-lobed, and very rarely unlobed. T. Lappula and T. Sanipaioi do not possess an urceolus. The stamens are borne cyclically on the gonophore, and the number is of great importance in specific criteria. Certain species have as high as 60 (T. attcfiuatUy T. arhorcscais) y others have about 40 (T, polyaiulray T. P^irpusiiy T. panicidata) ^ while the majority of the species possess between 15 and 25. T. Sampaio} is the only species in the western hemisphere v/hich has only 5 stamens. The smaller numbers are rather rare in Triunifetta. At least three species are known to be gynodioecious. In these the pistillate flower either has staminodcs or lacks them completely. No staminate flower is known for the whole genus. The filaments have retrorse serrations in both the scmifriloha and Bartramia complexes. Other species have smooth filaments. In T. cbihuahucnsis the bases of the filaments are lightly stellate-pubescent. The anthers do not yield any character of diagnostic value. They are always dorsi-fixcd, consisting of two thccac, each with two introrse, longitudinally dehiscing cells. The pollen grains are ellipsoidal, and about 50 /i long and 3 5 /x wide. The surface is minutely rugulose to very nearly smooth. The number of cells of the ovary is a good specific character. In the scries Geniculatae, the ovary is invariably 2-mcrous, and in most species of Unctnatae It is 3-, and rarely 2-locuIate. Where the ovary is too small for convenient dis- section, the number of locules may be ascertained by dissecting a very small floral bud, and counting the number of stigma lobes. In more mature buds the stigmas, in most species, fuse and do not separate easily. In certain species, however, the stigmas remain separate throughout. The descriptions of the stigma under each species In this paper is that of the later stage. The style Is usually much longer than the stamens, and the stigma Is generally exscrted above the anthers. However' in T. dioica and the species of the Galcottiaua complex the style is shorter than the stamens and the stigma is included by the anthers. It may be of interest to note that these species are the only ones known in the genus which exhibit gynodloecism. The spinules arc small transparent epidermal outgrowths of the ovaries. They arc the first to difi^ercntiate on the ovaries and retain their shape and size on the mature fruits. They are probably secretory in function, as there are large quanti- ties of mucilage In the early stages of ovary differentiation. In the series Stellatae and Geniculatae these spinules arc always straight-pointed, while in Unctnatae they are arcuately recurved. After fertilization, the spines develop froni the pericarp, each at the base of a splnule. Soinctimes, as In T. candata and especially in T. Hiutoniiy spines fail to develop from every spinulc. The spinules arc thus carried at the tips of the spines, and usually harden slightly after anthesis. What- ever their degree of rigidity, the spinules are always transparent and hyaline. The nature of the spinules on young fruits, slightly after fertilization, is the best 195 0] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 325 criterion for determining to which series the plant belongs. The fruit is the best single character for determining the species. The shape is generally constant for a species. T. caudata^ T. Hintonii^ and T, socorrensis possess oblongoid fruits, while the rest of the species have cither spheroid or sub- orbicular ones. It may be noted that these three species possess much fewer spines than the others. It is often difficult to speak with certainty regarding the dehiscence of the fruits. In at least three species (T. polyandray T. grandifloray and T. discolor) the fruits are easily dehiscent. In certain others, such as T. seviitriJoba^ T, miilfi- locularisy and T. altbaeoideSj they are quite indehiscent. In a few the fruit can be made to separate Into valves by prolonged boiling, but there is no evidence that it actually dehisces in nature. The number of cells is constant for a given species. Each cell is potentially 2 -seeded, but there Is a general tendency in most species to the formation of false scptae and production of double the number of cells in mature fruit, each with one seed. The reverse is true In three species, T. HintonU^ T. Bartramla, and T. Lappiila^ where one or more cells abort early in the process of fruit development. The size of the mature fruit body shows little variation. However, care is needed in the use of this character, as it often is difficult to know from dried material whether a fruit Is mature. In general, species with large showy flowers tend to have relatively large fruits. p exceptions as they possess rather large fruits but have very small flowers. The nature of the pubescence on the fruit is less variable than that of the leaves and stems. Certain species, such as T. Mex/ac and those in the GaJeottiana complex, possess glabrous fruits. Sometimes two rather closely allied species may be essentially different only In the pubescence of the fi'uit body (T, Purpiisii and T, falc'ifera). In T*. Lappula a few fruits have been noticed to have densely pubescent bodies and long plumose hairs on the spines when either diseased or attacked by insects. This may be interpreted as traumatic reversion to an ancestral character. While the general nature of the splnules affords characters for series, those of the spines themselves are characteristic of individual species. The spines may be glabrous or pubescent. When pubescent, they may bear many long hairs (plumose) or short rather irregularly arranged hairs (irregularly pubescent) or short hairs which are all retrorsely pointed (retrorsely pilosulose). Further, the length of the spines and the number per fruit being reasonably constant for a given species, they have been found satisfactory in primary separation of the species. Section Lappula is characterized by rigid spines; the only known American species with more or less flexible spines is T, lougicoma. The seeds are cither ovoid or lenticular, and have a prominent funicular groove. There Is a considerable variation In the size of the seeds among the species. Measure- ments of the mature seeds are given under every specific description. 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN [Vol. 37 27 Generic and Subgeneric Relationships Trhimfctta Is commonly referred-^ to tlie tribe Grcwieae of the family Tiliaceae and usually is recognized by its characteristic bur-like fruits. The genus Is dis- tinct from the other genera of Grewieae except HeUocarpus. Some of the more important distinguishing characters between the two genera have been enum- erated in my revision of the genus Heliocarp74s. In their treatment of the African species of Trhimfetta Sprague and Hutchin- son"^ divide the genus Into four sections based on the texture of the fruits, the nature of the spines and the inflorescences, and some characters derived from the indument of the sepals. According to their classification all the American species belong to section Lappula, under which they tentatively proposed the three series Stellatae, Geniculatae and Uncinatae. Before considering the scries in detail it is desirable to summarize here the char- acters of the genus which have been considered primitive: 1. Arborescent habit as opposed to shrubs or annuals. 2. Dense tomcntum of both stellate and simple hairs on leaves and branches, usually associated with plumose hairs on spines versus glabrcscent condition of leaves and branches together with glabrate or retrorsely pilusolosc spines. 3. Simple inflorescences consisting of 1 or 2 cymulcs in the axils of the bracts as compared with compound aggregate dichasia where the cymes are borne in dense nodose clusters of 12-20 flowers each. 4. Large showy flowers with many stamens, and multiloculate large fruits versus small flowers having few stamens with a tendency towards dioccism; also marked tendency towards reduction of floral parts such as gonophore, gland, urceolus, and petals, and the general progress towards 1-loculate, small, glabrous fruits. The series Stellatae Is known to me from only one specimen, and I am in no position to comment on its phylogcny or relative antiquity, except to say that it IS very closely related to Geniculatae. The splnules on the ovary are straight and undivided and resemble those In Geniculatae; but in fruit they consist of 4 to 5 per spine and are stellately arranged. The series Geniculatae is a relatively small group consisting of 14 species, most of which are rather poorly represented in the herbarium. The species of this scries generally possess a tree habit, 2-merous ovary, usually numerous stamens, and ovaries bearing spinules which are either straight or slightly dcflcxed in fruit, but never recurved. Each spine is terminated by one spinule only. These char- acters, together with the obvious consideration that a hooked spinule of Uncinatae is definitely more efficient in the dispersal of fruits than the straight one of Geniculatae, lead one to the conclusion that Geniculatae is more primitive. ^*'K. Schumann in Engl. & Prantl, Nat. Pflanzenfam. 3'^:28. 1895. -"Ann. Mo, Bot. Card. 36:508. 1949. 28 Jour. Linn. Soc. But. 39:233. 1909. 1950] KO KO LAY AMERICAN TRIUMFETTA 327 UNCINATAE T. Lappula « Bartramia complex" tc SEMITRILOBA complex" f V Galeottiana complex" T. falcijcra T. Purpusii T. grandiflora T. discolor A T. Pa!?neri T. socorrensis] coll nun arts barbosa chibnahiicnsis Pn T, brevipcs longicoma fnollissima T, wtdtilocularis GENICULATAE STELLATAE r. dioica t T, Mcxiae T, Caldcrom 7*. Hhifoui T. Cauda fa T. Goldnianl T, coriacca T. cucullata T, speciosa T, polyandra r, stcllata > Triumfetta polyandra is apparently the most primitive species of the series since it possesses very large showy flowers (the largest known for the genus), numerous stamens, and also very large fruits with very densely plumose spines. The character of the spines In turn relates it to Lasiothrix, a section considered to be more primitive than Lappula by Sprague and Hutchinson, [Vol. 37 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN The derivation of the species of Geniculatae from T. polyamlra is appar- ently quite simple, T. spcciosa^ with its very large flowers and fruits and its long plun^ose spines, could be derived from T. polyafidray from wliich it differs in having fewer stamens and much smaller petals. By the reduction in the sizes of flowers and fruits and also in the number of spines, T. caudata could be conveniently arrived at from T, spcciosa, T. caudata possesses very plumose spines and densely tomcntose fruits, leaves, and branches. T. Hiiitonii culminates this line of reduc- tion, as it has glabrescent leaves and branches, much smaller flowers, very few short spines, and a 1-loculate ovary due to the abortion of one loculc. In T. caudata^ and especially in T. Hintouiiy the number of spines per fruit is usually much less than the number of spinules on the ovary. Another line of evolution which stems from T, spcciosa leads to the Calderoni complex, consisting of seven species. The more primitive species of this complex are characterized by the possession of plumose spines and small but densely tomcntose fruit bodies. They all have very small flowers usually borne on com- pound aggregate dichasia, in which each cyme is composed of 12—20 flowers. The derivation of the flowers of this complex from T. spcciosa is not too clean The three species, T. Calderoni, T. fcrrugifica, and T, arhoresccnSy are considered more primitive than the rest of the species in the complex, since they possess numerous stamens, plumose spines, and densely tomcntose fruits. T. Mexiac is the only species of the genus which has a small gynophore borne upon the gonophore. This character, in conjunction with the glabrate fruits and spines and fewer stamens as compared with the three species mentioned above, makes T, Mexiac appear definitely more advanced. T. attcnuata is very closely related to T. Mexiac. Dioecism Is attained in T. dioica, a species ^hich shows close affinity with T. Caldc A second line of evolution from T. polyavdra leads to T, coriacca and T. cnciillata. It is unfortunate that no fruiting specimen of T. citciiUata is available. However, in its vegetative and floral characters it is strikingly similar to T. coriacca^ a species which has numerous stamens and large flowers with showy petals. It diff^ers from T. polyandra in its smaller fruits and fewer and less pubescent spines. Except that T. Goldmafii has fewer stamens, it has much the same characters as T. coriacca. The scries Uncinatae is rather well represented in the herbarium and consists of 27 species in the western hemisphere and about 25 in the eastern. Unlike Geniculatae, this series is not a compact group, and is apparently polyphylctic in origin, as some of the species with uncinate spinules could be derived more easily from species in Geniculatae than from those in Uncinatae. This scries is characterized by the members having spinules which are continuously recurved both on the ovary and the spine, by their shrubby habit, by the predominance of 3-merous ovaries, and, except in the more primitive species, by the presence of only 5-25 stamens. roni. 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 329 The most obvious direct line of evolution is from T. breiipes wKich, though advanced in many respects, could be easily referred to T, polyandra on such primitive characters as the size of the flowers, the presence of numerous stamens, and the large, more or less tomcntose fruits with numerous though short spines. This species apparently gives rise to three Unes of evolution, of which two end in strict endemism and the third attains a high degree of success and complexity. In possessing relatively large, showy flowers, and large, more or less tomentose fruits and spines, the endemic species T, socorrensh is obviously related to T. brei'ipes; however, it is more advanced than the latter as the number of spines and stamens is considerably less. T. Piirpusn^ another endemic species, has the same type of fruits as T. brcvipcs, though they are slightly smaller and armed with fewer spines. Its flowers still possess numerous stamens, but are rather oddly specialized in having excessively long apical appendages of the sepals. The rela- tionship between T. Purpusii and T. falcifera is so close that it is sometimes slightly difficult to distinguish the two species when in flower. The fruits of T. falcifera and T, Purpii-sii are alike In general shape and size but those of the former arc nearly glabrous. The more successful line of evolution from T. breiipes leads to T, Pahneri, An Interesting specialization appears here for the first time. The spines arc clothed lightly with short hairs which are all retrorsely pointed. This feature, together with hooked spinulcs, is of definite advantage in the dispersal of the fruits by animals. Though T. Pahncri has retrorsely pllosulose spines, it still possesses large flowers and fruits. The highly successful and widely distributed semitriloba complex may be derived from this type by the reduction in the sizes of flowers and fruits, and also by the inflorescences becoming slightly more compound. Further reductions in the floral parts leads to T, Lappithj a very widely distributed species of the genus. There Is a considerable confusion with regard to the delimitation of T. semi- triloba. Some authors-'*' ^^ treat it in a very inclusive sense, to include all plants which have retrorse hairs on their spines but are not T. Lapfnila. This inclusive interpretation Is not satisfactory, because some of the segregate species are distinct even to a casual observer. It is more usual to split the complex into many species, chiefly distinguishing them on the size of the flowers, shape and size of the leaves, and on pubescence. This treatment does not solve the problem either, as most of the so-called species appear to be connected by numerous intermediate forms which defy classification In the herbarium. The difficulty is further enhanced by the Incomplete nature of the specimens, classification being frequently uncertain in the absence of mature fruit. The variation in the size and shape of the leaves is of such great magnitude that it Is usually Impossible to match two specimens with any degree of certainty by ^^K. Schumann in Mart. Fl. Bras. 12^:135. 1886. ^•^Uittien in Pullc, FL Surinam 3:56. 1932. [Vol. 37 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN tlic use of leaf characters only. Other characters being very similar, the leaves usually may vary from undivided to deeply and profoundly 3-lobedj the lobing being either above or below the middle of the lamina. The size of the flowers is relatively less variable, but there are instances, especially in T. bogotcnshy where it does not correlate with the other trends. The nature of the pubescence is slightly more constant, though there is a very considerable variation in the degree. Apparently the kind of hairs, whether stellate, simple, or both, is generally constant for a species, as it tends to vary with the general shape and size of the petals and fruits together with the number of locules and the nature of the spines. On some of these more constant characters, I have recognized six species in the scmitriloba complex, with the hope that it would be more satisfactory from both the practical and taxonomic standpoints. Some of the more important characters which help in the identification of the species are discussed briefly under each, besides those mentioned in the key. The three South American species, T. vinltilocularis, T. molliss/may and T. longicomay are related in but a superficial way to T. brcvipcs. The very definitely 2-merous ovaries, the general tendency towards a tree habit, and the large fruits with very plumose spines relate them more closely with T. polyandra. Among themselves, the relationships to one another are extremely strong. T. miilti- loculariSf though a very distinct species, has hitherto been mistaken for T. F njoIJissima in the herbarium. T. longicoma is the most advanced member of the th ree. Triumfcfta chihuahnensisy T. barhosa, and T. colnmitarh are three distinct but very closely related species. Although they have a few characters In common with T. Talnieriy they do not exhibit any close affinity to that species and are much more closely related to T. coriacea and T. GoldmanL Without the uncinate spinule, one would certainly experience difficulty in identifying T. coluwnaris and T. GoldmanL It is most probable that these three species of Uncinatae are de- rived directly from the three geniculate species, T. ciindlata, T. coriacea, and T. Goldman}, T. grand/flora and T, discolor are definitely related in having dehiscent fruits and rather large showy flowers with about 20 stamens. They are in turn related to T. mollissima. The Galeotfiana complex (consisting of the three species, T, panicalafa, T, Galeottiana^ and T. heliocarpoides) is of great interest in that though the three are definitely uncinate, they do not show any close affinity with any other species of the series. However, there is a very remarkable similarity between this com- plex and the Calderoni complex in Geniculatae. The two complexes have the same type of inflorescence, flowers of the same shape and structure, and fruits whose superficial resemblance is rather striking despite the fact that all species of the Galeottiana complex have essentially glabrous fruits. All species in the Calderoni complex have 2-merous ovaries, and T. panicnlata bridges this difference in possessing a 4-loculatc fruit. The affinity is further strengthened by the 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 331 tendency of T. pauiciilata to a tree habit, Its numerous stamens, and its relatively large fruit. T. Galcottiana and T. heViocarpoidcs are not only closely related species but arc also very closely allied with T. panicnlata. Though the three are distinct, sometimes the delimitation of the species becomes very difficult especially when immature fruiting specimens are being considered. Both T. Galcottiana and T, beliocarpoicles attain gynodioecism. The Bartramia complex, consisting of four species, T. sericafa, T. brachista- cantha, T. Bartranua, and T, Sampaioiy is really problematic in its relationships. In the possession of cymes in dense nodose clusters the species resemble those of the Galcottiana complex, but, on the other hand, the oblongoid floral buds and pubescent fruits make them resemble the scmitriloba complex. The picture is further complicated by the fact that T. Bartramia is a pantroplc weed which is equally at home in both the eastern and western hemispheres. Though the com- plex does not attain gynodioecism, there is considerable reduction in the floral parts and in the size of the fruits, which are the smallest known for the American species. The fact that T. Sampaioi lacks an urceolus and has only five stam.ens places it In approximately the same degree of advancement and specialization as T. Lappula, T. scvicafa resembles T. hogotensis in having the same general type of flowers and indument, but has much smaller fruits with glabrous spines. T. hrachistacautha has characters which arc about intermediate between those of T, sericata and T. Bartramia. The relationships of the species of the Bartramia complex with those of the western hemisphere arc not too apparent. Its affinity may be with either the scmitriloba or Galcottiana complexes, or possibly with both. From the above discussion of some of the salient characters and relationships of the species, it is quite obvious that no definition of the series could be given which would suit all the species. The characters are superposed in such a manner that hardly any two are correlated. I am unable to give any precise definition of the series as there are nearly as many exceptional forms as there are typical ones. Further, an attempt at classifying the genus based on characters derived from the size of the flowers, types of inflorescences or the nature of the stigmas (as sug- gested by Hochreutiner^^) leads one into similar dilemmas. Because of these apparent reticulate combinations of the characters, I am led to the conclusion with the original authors that the ''series, though very useful for the purposes of a clavis, do not coincide with natural groups, but appear to represent three of the phylogenetic stages through which many species of section Lappula have passed, the stellate arrangement being the most primitive and the uncinate the most highly modified. "^^ The three series arc here maintained not only for the sake of convenience, but because they arc the nearest approach to the real relationship of the species. 31 Ann. Conscrv. Jard. Bot. Geneve 18:92. 1914. ^-Spraguc £c Hutch. Jour. Linn. Soc. Bot. 39:240. 19C9. [Vol, 37 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chromosome Counts Chromosome counts were made for the following three species: Triumfetfa Bartramia 2n 32 Triumfetta Calderoni 2n 32 Triumfetta semi triloba 2n 32 The counts were made from root-tips of plants raised in a greenhouse at Mis- souri Botanical Garden. The seeds for T, Calderoni were collected by Dr. Edgar Anderson from Honduras, and those for T. Bartramia and T. semiiriloba were sent to me by Dr. Brandao-Joly from Sao Paulo, Brazil. As none of the plants flowered mciotic counts could not be made. Acknowledgments I have been able to examine specimens, including many valuable types, from several major herbaria in Europe and North America and South America. The herbaria where specimens have been obtained for study, together with the symbols^^ employed in their citations are as follows: BR Jardin Botaniquc de I'Etat Bruxcllcs. C Universltctcs Botaniske Museum, Copenhagen. FM Chicago Natural History Museum, Chicago. G Institut dc Eotanique Systcmatique de rUnlversitc, Geneve. GH Gray Herbarium of Harvard University, Cambridge, Mass. K Herbarium Royal Botanical Gardens, Kcw. MICH University of Michigan Herbarium, Ann Arbor. Mo Herbarium Missouri Botanical Garden, St. Louis. NY Herbarium New York Botanical Garden, New York. P Museum National d*Histoire NaturcHc, Paris. R Museu Nacional, Rio de Janeiro. S Naturhistoriska Riksmusect, Stockholm. TRIN Herbarium Department of Agriculture, Port of Spain, Trinidad. U Botanisch Museum en Herbarium, Utrecht. UC University of California Herbarium, Berkeley. US U. S. National Herbarium of Smithsonian Institution, Washington, D. C. Dot maps have been prepared from most of the specimens from which the exact locality could be determined. In the Index to Exsiccatae all the numbered and most of the unnumbered collections have been cited. I wish to acknowledge my indebtedness to the curators of the institutions mentioned above, and to the Director of Missouri Botanical Garden, where this study was made. For illustrations and aid in chromosome counts I am indebted to Marilyn Amy Gage. Particular thanks are due to Dr. R. E. Woodson, Jr., for his advice, guidance, and constructive criticism. Taxonomy Triumfetta L. Sp. Pi. ed. 1. 444, 1753; DC. Prod. 1:506. 1824; K. Schum. In Mart. Fl. Bras. 12^:131, 1886, in Engl. & PrantI, Nat. Pflanzenfam. 3^:28. 1895; Sprague & Hutch, in Jour. Linn. Soc. Bot. 39:231. 1909; Hochr. in Ann. Conserv. Jard. Bot. Geneve 18:92, 1914. Bartramia L. Sp. Pi, ed. 1. 389. 1753. ^^The symbols used are as suggested by Lanjouw in Cliron. Bot. 5:143. 1939. 19 5 0] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 333 Z m ipr^ A^CLjm 8 Fig. 4. Illustrations of taxonomic criteria for Series Stellatae and Geniculatae 1. Small orbicular fruit of T. CalJeront with densely plumose spines. 2. Suborbicular fruit of T. Mexiae borne on a short gynophore above the gonophore. 3. Suborbicular to oblongoid fruit of T, caudata with about 25 plumose spines. 4. Oblongoid fruit of T. Hintonii with few short spines, 5. Fruit of T. stellata with 3—5 spinules per spine. 6. Orbicular fruit of T. Goldmani with falcate spines. 7. Large fruit of T. speciosa with densely plumose spines. 8. Large, easily dehiscent fruit of T, polyandra with large, very densely plumose spines. [Vol. 37 334 ANNALS OF THE MISSOURI BOTANICAL GARDEN Shrubs or trees. Leaves alternate, petiolate, stipulate, the stipules usually large, persistent, rarely fugacious, the blade 3- to 5-lobed or undivided, the venation palmate, 5- to 7-costate at the base. Inflorescences hermaphrodite or gynodioecious, usually axillary (terminal in few species), consisting of numerous aggregate dichasla. Flowers either hermaphrodite or pistillate, staminatc ones not known, hypogynous, 5-mcrous; sepals 5, valvatc, free, with apical appendages; petals 5, sometimes absent, free, imbricate, 3- to 5-nervcd, shorter than the sepals, usually cihatc at the base (claw) ; gonophore bearing 5 spherical or elongate glands mani- • fest, absent in some species, usually crowned by a ciUate, rather membranaceous ring (urceolus) surrounding the stamens; stamens 5-60 in hermaphrodite flowers, staminodial or absent in the pistillate, borne cyclically on the enlarged gonophore, with 2 thecous, 4-celled, introrse, longitudinally dehiscent anthers, the filaments free, usually glabrous, sometimes pubescent at the base, often with retrorse serra- tions; ovary wholly superior, sessile on the gonophore (except in T. Mcxiae, with very short gynophore on the gonophore), ellipsoid to orbicular, covered with many short, cither erect or recurved, hyaline spinules, 2- or 3-loculate, each with 2 anatropous collateral pendulous ovules, the style filiform, the stigma briefly 2- or 3 -parted, sometimes capitate. Fruits dry, indchiscent or loculicidally dehiscent, 2- or 3-loculatc, each cell 2-seeded, or falsely 6- to 9-loculate, or each cell 1-seedcd, or 1-loculatc with 1 seed due to abortion, usually sessile on the slightly accrescent gonophore, lappaceous, the spines elongating from the pericarp at the bases of the spinules; seeds compressed-ovoid or pyriform, with a more or less prominent funic- ular groove, the cotyledons folded, the endosperm oily- Standard species: Triumfctta Lappula L. KEY TO THE SERIES A. Hyaline spinules of ovary straight, In fruit cither straight or slightly deflcxcJ, or consisting of 3-5 stellatcly arranged; ovaries 2-Tncrous; stamens 20-60, most of the species with 40 or more; usually small trees 3-5 m. talL B. Each spine with 3-5 stellately arranged spinules; mature floral buds panduratc Stfllataf (p. 334). BB. Each spine with 1 straight or slightly dcflexed spinule; mature floral buds oblongold or obovoid Geniculatae (p. 3 36) AA. Hyaline spinules of ovaries and fruits arcuatcly recurved; ovaries 3- or rarely 2-merous; stamens 5-40, most of the species with less than 30; usually shrubs 1-3 m. high Uncinatae (p. 349) Series Stellatae Sprague 8: Hutcli. in Jour. Linn. Soc. Bot. 39:240. 1909. 1. Triumfetta stellata Ko Ko Lay, sp. nov. Arbores parvac 3 m. altae. Ramuli graciles dense stellato-pubescentcs. Folia anguste ovata interdum obscure 3-lobata 5-6 cm. longa 2-3 cm. lata ad basim subtruncata vel subcordata longe acmuinata subdupliciter serrata utrinquc pilis sparsls stcllatis; pctiolis 1-3 cm. longis pilis fulvis stellatis pubescentibus. In- florcscentiae axillares, cymis cymulis 4-5, pedunculis plerumquc 4-6 mm. longis, pedicellis 2-3 mm. longis. Flores hcrmapKroditi, alabastro maturo pandurato 8-9 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 335 / Fig. 5. Triumfctfa stcllata mm. longo; sepalis ellipticis circiter 10 mm. longis extra minute stcllato-puberu- Icntls, appendiculis circiter 0.25 mm. longis; petalis obovatis brevissimis 2-3 mm. longis, unguilo 1.0-1.5 mm. longo extra breviter villoso; gonoplioro circiter 1.5 mm. longo, glandulis ellipticis circiter 1.0 mm. longis 0.5 mm. latis; urceolo 0.3 mm. alto 5-lobato breviter ciliato; staminibus 25-30, filamentis usque ad 5-7 mm. longis glabris; ovario breviter oblongo-ellipsoideo circiter 1 mm. longo, spinulis 3 5-40 erectis, stylo glabro 2-3 mm. longo, stigmate breviter 2-fido. Fructus globosi non debiscentes, corpore ipso 3 mm. diametro 2-loculari, loculis 1-spermis cano-tomentosis, aculcis 2-3 mm, longis inaequilater breviter pubescentibus, spinulis 3-5 per aculco; seminibus maturis baud visis. Mexico: Mexico: Tcmascaltepec, Las Mesas, alt. 2000 m., Oct. 25, 1933, Hinton 4460 (HOLOTYPE in Herb. Kcw.). [Vol. 37 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN The aflfinities of this very distinct species are doubtful. It has a superficial resemblance to T. Galcottiana, but can be recognized easily in fruit by the 3 to 5 stcllatcly arranged spinules on the spines, and in flower by its panduratc floral buds with very small obovate petals, which also have small foveae at about the tip of the claw. J 1909. KEY TO THE SPECIES A. Sepals at anthesls 2-4 cm. long; mature fruit body 4-10 mm. in diameter. B. Sepals at anthesis 3.5-4.0 cm. long, the apical appendages 1-3 mm. long; spines 4 mm. long or more, densely plumose. C. Leaves broadly elliptical to obovate, the petioles about 1 cm. long or less; petals broadly obovate, subcqualling the sepals; fruit easily dehiscent, 1.2-3,0 cm. wide including the spines, the spines 0.6-1.0 cm. long; southern Mexico and Honduras 2. T. polyaudra CC. Leaves ovate to 3-lobed, the petioles 3-5 cm. long; petals ellip- tical, less than half the length of the sepals; fruit indehiscenc, 1.0-1.5 cm. wide including the spines, the spines 3-5 mm, long; southern Mexico to Panama 3. T. spccjosa BB. Sepals at anthesis 2.0-2.5 cm. long, the apical appendages about 1 mm. long; spines 2-3 mm. long, lightly pubescent to glabrescent (not known in T. cucullata) , C. Leaves 11-15 cm. long, the petioles with 2-4 prominent spherical glands at the basal sinus; petals broadly obovate, about 22 mm. long. D. Leaves elliptical, acute, the petioles about 1.5 cm. long; stamens 20; coastal west-central Mexico 4. T. cucuUafa DD. Leaves broadly ovate to 3-lobcd, acuminate, the petioles 4—6 cm. long; stamens 40; southwestern Mexico 5. T. corliicca CC. Leaves 5-7 cm, long, the petioles without glands at the basal sinus; petals elliptical to narrowly obovate, 1.2—1.6 cm, long; western Mexico ^. T, GolJmaiii AA. Sepals at anthesis 0.4-1.6 cm. long; mature fruit body 2-3.5 mm. in diameter. B. Sepals at anthesis 1.0-1.6 cm. long; fruit body oblongoid, the spines 2 5 or fewer. C. Stem and petioles densely ferruginous-tomcntose; leaves pubescent on both surfaces, densely so beneath; sepals densely tomentose without; mature fruit tomentose, 2-loculatc, the spines 3-5 mm. long; Colombia 7. T. canJata CC Stem and petioles glabrescent; leaves essentially glabrous on both surfaces; sepals nearly glabra te without; mature fruit glabrate, I-loculate, the spines 1-2 mm. long; central Mexico.... 8. T. Hinfonii BB. Sepals at anthesis 0.4-1.0 cm. long; fruit body globose or sub- globose, the spines 50 or more. C. Petals 1-2 mm. lung; leaves densely tomentose; spines plumose, the fruit densely tomentose (not known in T. fcrruginca) . D. Apical appendages of the sepals nearly obsolete; leaves cuncate to rounded at the base, rarely subcordate; Honduras and El Salvador ...- 9. T. Cahlcroni DD. Apical appendages of the sepals manifest; leaves distinctly cordate. E. Inflorescences hermaphrodite; sepals 7—10 mm. long; style 4—6 mm. long. F. Fluwerin;; peduncles 2-4 mm, long; floral buds obovold; stamens 40-60. G. Apical appendages of the sepals 2-3 mm. long, plumose; stamens about 60; stigma undivided; southern Mexico.. 10. T. fcrrn^iuca 19 50] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 337 GG. Apical appendages of the sepals about 1 mm. long, lightly pubescent ; stamens 40-5 ; stigma 2-lobcd ; Costa Rica and Panama 11. T. arboresccns FF. Flowering peduncles about 1 mm. long; floral buds ob- longoid; stamens about 20; Colombia 12. T. persimills EE. Inflorescences gynodioecious; pistillate sepals 4-5 mm. long; style about 2 mm. long; southern Mexico 13. T. dioica CC. Petals 3—7 mm. long; leaves glabrescent; spines and fruits glabratc (not known in T. attcnuuta), D. Leaves iong-attcnuate, the veins on lower surfaces glabrate; sepals essentially glabrous without, the apical appendages about 2 mm. long; stamens 60; style about 6 mm. long; southern Mexico 14. T. aitenuata DD. Leaves shortly acute to acuminate, the veins on the lower surfaces pubescent; sepals lightly stellate-pubescent without, the apical appendages less than 1 mm. long to nearly obsolete; stamens 40; style about 2 mm. long; southwestern Mexico.... 15. T. ls\cx'iac m 2. Triuaifetta polyandra DC. Prod. 1:508. 1824. (T.: based on Sessc & Mocino's tab, II J in Herb. Geneve). 34 Triumfctta obovata Schlecht. et Cham, in Linnaca 5:288. 1830. (T.: Schlcde Gf Deppc Trmmfetta lnsig7ih S. Wats. In Proc. Am. Acad. 22:400. 1887. (T.: £. Calmer 1^2). Small trees or large sbrubs about 3-4 m. high; branches and inflorescence axes very densely tomcntose with long ferruginous hairs. Leaves broadly elliptical to obovate, 13—15 cm. long and 10—12 cm. wide, the base rounded or subcordatc, the tip abruptly acuminate, doubly and regularly serrate, both surfaces scabrous, rather densely tomentose with both simple and stellate hairs; petioles short, stout, about 1 cm. long, covered rather densely with long hairs. Inflorescences on axillary and terminal branches, the cymes of 2—3 cymules, the flowering peduncles 5—9 mni. long, the pedicels 3-5 mm. long. Flowers hermaphrodite, the buds oblongoid, 30—3 5 mm. long slightly before anthesis, the apical appendages 1—3 mm. long; sepals narrowly obovate, about 3 6 mm. long, covered rather densely with many long coarse hairs without; petals broadly obovate, showy, 2.7-3.2 cm. long and 5—6 mm. wide, the claw 5—6 mm. long, densely plumose ; gonophorc short and stout, about 0.75 mm. long and 2.0 mm. thick, the glands oblong, subcqualling the gonophore; urceolus about 0.75 mm. high, many-lobed, lightly cihate; stamens 40—50, the filaments glabrous; ovary spheroid, 3 mm. long, the style 25-28 mm. long, the stigma acute. Fruit spherical, easily dehiscent, the body 6-10 mm. In diameter, densely plumose, 4-Ioculate, each 2-seeded, the spines numerous, 6—10 mm. long, densely plumose with long hyaline or ferruginous hairs, the spinules erect; seeds ovoid, about 2 mm, long and wide. Rather rare plants which grow in open forests, usually upon rocky mountain slopes at altitudes of 800-1500 m. in southern Mexico and northern Honduras; flowering from August to September, the fruit maturing from late October and persisting on the plant until early December. h 34 Hochr. in Ann. Conserv. Jard. Bot. Geneve 18:110. 1914. [Vol. 37 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mexico: chiapas: Chacomusclo, alt. 800 m., MatuJa 44j6{}Ao). jalisco: Rio Blanco, E. Pahncr Ij2 (Mo, US), 2j2 (US); slopes of barranca near Guadalajara, Pringle 446Q (BR, Mo); bluffs near Guadalajara, alt. 5000 ft., Prifigle g6()4 (Mo, US); on tKe road between Colotlan and Bolanos, Kose 2822 (NY). Mexico: Tcmascaltepcc, Luvianos, alt. 1300 m., Hhiton 5goo (US), nayarit: Tepic, between Pedro Paula and Acuponcta, Kose ^92J (NY), vera cruz: Mirador and vicinity, Lkhmann ^T2, J2I (US), alt. 3500 pod., Lifhlcn Suppl 26 (BR), 47 (BR, G) ; Cordillera, Zacuapan, alt. 2000-3000 ped., Galeofti 7062 (BR). Honduras: yoro: Aguan River valley, vicinity of Coyoles, alt. 1000 ft., Yuuckcr, Koepper & Wagnrr S167 (Mo). This species is very distinct and can be easily recognized by its large showy flowers, the largest known for the genus, and also by the extremely characteristic large, easily dehiscent fruits with many long very plumose spines. De Candollc based his description on the sketch of Scsse and Mocino. I have seen a photograph of the original sketch and have no difficulty in recognizing that T. insignh of S. Watson IS conspcciflc with it. I have not seen the type of T. olwvafii, but my interpretation of that species is based on the original description, which was later amplified by Turczanlnow on Linden's Supplementary Collection No. 47, which I h ave seen. 3. Triumfetta speciosa Seem. Bot. Voy. Herald, 86. 18 53. (T.: Sccmann 1240). Tr'iumfcifa ynacrocalyx Turcz. in Bull. Soc. Nat. Moscou 31^:230, 1858. (T.: Linden SuppJ,2j).^ Triumfetta mlcropetula Hochr. in Ann. Conserv. Jard. Bot. Geneve 18:98. 1914. (T.: Linden Sup pi . 25). Small trees or large shrubs 3-4 m. high; branches and inflorescence axes scurfy, densely covered with ferruginous tomentum. Leaves broadly ovate, generally 3- lobcd, 6-9 cm. long and 5—7 cm. wide, the base rounded or subcordate, the middle lobe long-attenuate, the laterals short and blunt, the serrations very irregular, usually blunt and often glandular, the upper surface dark green, coarse with many short stellate hairs, the lower pallid, densely tomentose; petioles 3-5 cm. long, lightly pubescent. Inflorescences axillary, the cymes of 1—2 cymules, usually opposite the bracts, the flowering peduncles 4-5 mm. long, the pedicels 6-8 mm. long. Flowers hermaphrodite, large, the buds oblongoid, 30-3 5 mm. long slightly before anthesis, the apical appendages about 2 mm. long; sepals narrowly elliptical, about 3 5 mm. long with many separate tufts of long stellate hairs without; petals ovate, short, about 6 mm. long, the claw about 3-4 mm. long, vcry^ densely plumose; gonophore about 1 mm. long and 2 mm. wide, the glands spherical, sub- equalling the gonophore; urceolus short, about 0.5 mm. high, many-lobed, lightly ciliate; stamens 20, the filaments glabrous; ovary ovoid, about 1.5 mm. long, the style 3 0—34 mm. long, with many short separate tufts of stellate hairs, the stigma acute. Fruit orbicular, the body 6-8 mm. in diameter, rather densely pubescent, 4-loculate, each cell 2-seeded, on maturity with 8 distinct 1 -seeded locules, the spines numerous, 100 or more, each 3-4 mm. long, densely plumose with short hairs, the spinules erect; seeds ovoid, about 2.0 mm. long and wide. 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 339 A species of relatively scattered distribution, which extends from southern Mexico to Panama; usually growing in open areas on slopes of mountains, or along the rocky banks of streams at altitudes of 13 00-1700 m.; flowering from late July to September, the fruit persisting on the plants until late February or March. MExrco: chiapas: Siltepec, Matuda 1 581 (Mo). Jalisco: San Sebastian, trail to El Ranchito, alt. 1500 m., Uexia 1444 (Mo, US), vera cruz: Mirador, Linden Siippl 2j (BR, FM photo); Zacuapdn and vicinity, Purpus IQlS (Mo, NY, US), 3749 (Mo, UC); Cordillera, locality not mentioned, Galeotti ip^J (BR), 4162 (BR). British Honduras: Vaca Falls District, Record s, n, (US). Guatemala: alta verapaz: Patal, alt. 1600 m., Titerckbcim II 1546 (Mo); San Cristobal, Tejada 265 (US). Honduras: morazan: La Montanita, alt. 1400 m., Williams ^ Molina R. 10636 (Mo); Mt. Uyuca, alt. 1600 m., Williams ^ Molina R. 10252 (Mo). Panama: chiriqui: Boquetc, Seernann 1240 (K) ; Volcan de Chiriqui, Boquete Distr., alt. 7000 ft., Davidson 515, 8g5 (Mo); Finca Lerida to Boquete, alt. 1300-1700 m., Woodson, Allen ^ Scibert II 1 5 (Mo). This species is characterized by its very large flowers and disproportionally short petals, and also by its large fruits with briefly plumose spines. It is rather interesting to note that both the species of Turczaninow and Hochreutiner are primarily based on Linden SnppL 25, a sheet which was apparently examined also by Seernann, as he cites a Linden collection from Vera Cruz, Mexico. 4. Triumfetta cucullata Fernald, in Bot. Gaz. 20:532. 1895. (T.: Lamb 579)^ Shrubs about 2-3 m. high; lower branches sparingly pubescent, the upper and the inflorescence axes densely cinereous-tomentose. Leaves elliptical, about 13-15 cm. long and 4-5 cm. wide, the base obtuse, with 4 spherical glands at the basal sinus, the serrations generally glandular, the upper surface Hghtly stellate-pubescent, the lower more densely so; petioles 1.0-1.5 cm. long, densely tomentose. In- florescences axillary, the cymes of 3 cymules, generally condensed, the flowering peduncles about 3 mm. long, the pedicels 3-5 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, 22-24 mm. long slightly before anthesis, constricted sHghtly above the base, the apical appendages slender, 0.5-1.0 mm. long, very lightly pubescent; sepals oblong, 22-25 mm. long including the apical appendages, steUate-tomeatose without, glabrcscent within; petals broadly obovate, about 23 mm. long nnd 6-7 m.m, wide, the claw about 4-5 mm. long, densely plumose; gonophore short and stout, about L5 mm. long and 2-3 mm, wide, the glands quadrangular, broad, about 1.0 mm. long and 1,5 mm. wide; urceolus short, about 0,5 mm. high, distinctly many-lobed, briefly cihate; stamens 20, the filaments smooth, densely pubescent at the base; ovary orbicular, about 1.5 mm. long, the spinulcs about 50, erect, the style about 22 mm. long, the stigma acute. Fruit not sQcn. Known only from the type. Flowers in February. Mexico: nayarit: Tepic, Zopelote, alt. 2000-3 000 ft., Lamb 57g (Mo, US). Though very imperfectly known, this species may be easily identified by its short-petiolate elliptic leaves, by its large showy flowers, and also by its possession of only 20 stamens with the filaments densely pubescent at the base. [Vol. 37 340 ANNALS OF THE MISSOURI 5. Triumfetta coriacea Hochr. in Ann. Conserv. Jard. Bot. Geneve 18:108. 1914, (T.: Lariglasse 815). Small trees or large shrubs 3-4 m. Klgh; branches and inflorescence axes gla- bresccnt, brownish. Leaves broadly ovate to 3-lobed, 11-14 cm. long and 8-12 cm. wide, the base cuneatc to rounded, the tip long-acuminatc; the serrations regular and usually double, the upper surface glabrate, the lower very lightly pubescent to glabrescent; petioles stout, 4-6 cm. long, with 2 or 4 prominent glands at the basal sinus. Inflorescences axillary, rarely terminal, the cymes of 2-3 cymulcs, borne very loosely In nodose clusters, the flowering peduncles 6-8 mm. long, the pedicels 2-4 mm. long. Flowers hermaphrodite, the buds narrowly oblongoid, about 22 mm. long sUghtly before anthesis, constricted in the middle, the apical appendages very short, about 0.5 mm. long; sepals elliptical, 22-24 mm. long, rather lightly stellate-pubescent without; petals broadly obovate, 20-22 mm. long and 6-8 mm. wide, the claw about 3 mm. long, densely plumose; gonophore slender, 1-2 mm. long, the glands elliptical, subequaUing the gonophore; urceolus about 0.5 mm. high, many-lobed, lightly dilate; stamens about 40, the filaments glabrous; ovary ovoid, about 1.5 mm. long, the splnules numerous, erect, the style about 22 mm. long, the stigma acute. Immature fruit orbicular, the body essen- tially glabrous, 2-IocuIate, each cell 1 -seeded, the spines numerous, rather short; mature fruit not seen. Southwestern Mexico, usually growing on hillsides in pine and oak forests at altitudes of 800-1000 m.; flowering from middle of December to late January. Mexico: guerrero: San Vincente de San Juan, alt. 920 m., Hinton IIIQJ (GH) ; Plan de Carrizo, Galcr.na, alt. 850 m., Hinton TIO56 (GH) ; Sierra Madre, alt. 800 m., LangJassc 815 (G, GH, US). This species can be recognized by its glabrescent, broadly ovate to 3-lobcd leaves v/Ith 2-4 spherical glands on the petioles at the basal sinus; and also by Its large, showy flowers with deep yellow broadly obovate petals. It is unfortunate that mature fruiting specimens are unavailable. 6. Triumfetta Goldmani Rose, in Contr. U. S. Nat. Herb. 12:285, 1909. (T.: Goldman 264). Triumfetta sanctae-Iuciae Sprague, in Kew Bull. 114. 1923. (T.: Sccmann 2141). Triumfetta qnercetonim Bullock, loc. cit. 295. 1937. (T.: Hinton 6746), Small trees or large shrubs 3.0-3.5 m. high; branches and inflorescence axes with many short stellate hairs, later becoming glabrate. Leaves elliptical to nar- rowly ovate, 5-7 cm. long and 2-3 cm. wide, the base cuncate or rounded, the tip narrowly and gradually long-attenuate, the serrations rather regular, both surfaces scabrous with many short stellate and simple hairs; petioles slender, 1.0-1.5 cm. long, rather lightly but coarsely pubescent; bracts of the same general shape as the vegetative leaves. Inflorescences axillary, rarely terminal, the cymes of 1-2 cymulcs, always in the axils of the bracts, the flowering peduncles 4-6 mm. long, the pedicels about 4 mm. long. Flowers hermaphrodite, the buds narrowly ob- longoid, 18-20 mm. long and 2-3 mm. wide sHghtly before anthesis, the apical 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 341 appendages slender, 1-2 mm. long; sepals narrowly ovate, about 20 mm. long, with few separate tufts of short stellate hairs without; petals elliptical to narrowly obovate, 12-16 mm. long and about 3 mm. wide, the claw 4-5 mm. long, densely plumose; gonophore large, about 2.5 mm. long and 1.5 mm. wide, the glands elliptical, subequalling the gonophore; urceolus short, about 0.2 5 mm. high, many- lobed, lightly ciliatc; stamens 20, the filaments glabrous, ovary orbicular, about 1.5 mm. long, the spinules about 75, erect, the style about 16 mm. long, the stigma either acute or briefly bifid. Fruit spheroid, the body 4-5 mm. in diameter at maturity, Hghtly pubescent, 4-loculate5 each cell 1 -seeded, the spines falcate, about 75, 2-3 mm. long, lightly pubescent with few short hairs; seeds pyriform, about 1.5 mm. long and wide. Rather common plant of western and central Mexico, usually growing in oak and pine forests in shady slopes under trees in moist places or on edges of meadows along streams at altitudes of 600 to 2000 m.; flowering in September and October, the fruit persisting until late January. Mexico: chihuahua: Arroyo Hondo, Sierra Charuco, Gentry 1^68 (Mo, US); San Jose dc Pinal, Rio Mayo, Gentry 2841 (Mo); Guicorichi, Rio Mayo, alt. 5000 ft.. Gentry 1967 (Mo); El Carrizo, alt. 800 m., Ortega 6011 (US); Balboa, Ortega 504] (US). DURANGO: Sierra Trcs Picos, alt. 3500 ft., Gentry S2q6 (Mo); La Bajada, Tamazula, alt. 300 m., Ortega 4252 (US). Guerrero: Mina, Zihuaglo-Ojo dc Agua, alt. 1640 m., Hinton gyio (US); MIna, Chlrlagua, alt. 1600 m., Hhrton 9847 (GH) , 10663 (GH, Mo, US); Pllas FIIo Mayor, alt. 1800 m., Hinton 10750 (GH, US); Chilacayote-Espadinas, alt. 15 00 m., Hinton 14927 (GH, US). Mexico: Temascaltepec: RIncon del Carmen, alt. 1340 m., Hinton 1953 (Mo); Nanchititla, Hinton 3082 (GH, K), 4962 (K) ; Punga- rancho, alt. 950 m., Hinton 5253 (Mo, US), 6746 (GH, K, Mo, US); Yperlcones, Hinton 6996 (Mo, US), mtchoacan: Coalcoman: Salltrc-mcsa, alt. 1780 m., Hinton 12492 (GH, US); Pto. Zarzamora, alt. 1660 m., Hinton 12262 (GH, US), sinaloa: Santa Lucia, alt. 1200 m., Seemann 2147 (GH, K); Sierra dc Choix, 50 miles n. e. of Choix, Goldman 264 (US); Cerro de la Sandia, northeast of Panuco, alt. 1800-2000 m., Pennell 20I2I (US); Qucbrada de Mansana, Sierra Suratato, alt. 4000-4500 ft.. Gentry 6382 (Mo); Sierra Madrc Mts., "El Bate!'*, alt. 4000 ft., Mexia 431 (Mo); Quebrada Chica, San Ignacio, alt. 13 80 m., Narvaez d Salazar 100 (US), sonora; Arroyo del Agua Blanco, Dist. Alamos, Gentry §o6M (US). The fruits of this species are very characteristic, being spheroid with about 75 falcate, lightly pubescent spines. Though the spines arc slightly recurved the spinvilcs are erect. The small elliptical to narrowly ovate leaves and the relatively large flowers with yellow, narrowly obovate petals make this species convenient to identify. The leaves of the type of T*. qjiercetornm are very slightly more pubes- cent than those of T. Goldmani, but in both they are nearly glabrate and the variation is only slight. 7. Tkiumfetta caudata Tr. & Planch, in Ann. Sci. Nat. Bot. IV, 17:353. 1862. (T.: "Rio Hacha", PnrJie s. n. in Herb. Kew.), Small trees or shrubs 2-3 m. high; branches and inflorescence axes covered very densely with rich orange-brown simple and stellate hairs. Leaves broadly ovate, 11-15 cm. long and 8-10 cm. wide, the base rounded to subcordate, the r tip gradually and narrowly long-attenuate, the serrations blunt, small, usually double and regular, the upper surface scabrous, lightly pubescent, the veins more [Vol. 37 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN densely so, the lower rather densely tomentosc with both stellate and simple hairs; petioles 3-5 cm. long, densely covered with orange-brown hairs. Inflorescences axillary, the cymes of 2-3 cymules, usually in the axils of the bracts, the flowering peduncles 4-5 mm. long, the pedicels 3-4 mm. long. Flowers hermaphrodite, the buds narrowly oblongoid, 10-15 mm. long slightly before anthesis, the apical appendages 1-2 mm. long, densely pubescent; sepals oblong, 12-16 mm. long with dense orange-brown pubescence without; petals narrowly obovate, subcqualling the sepals, the claw 2-3 mm. long, rather densely plumose; gonophorc short, about 0.5 mm. long, the glands small, spherical; urceolus about 0.5 mm. high, many- lobed, densely ciliate; stamens 20-2 5, the filaments glabrous; ovary ovoid, about 0,75 mm, high, the spines 40-50, erect, the style 7-8 mm. long, the stigma acute. Fruits oblongoid, the body 2-3 mm. in diameter and 3-5 mm. long at maturity, densely pubescent with many short stellate and simple hairs, 2-loculate, each cell 1 -seeded, the spines 20-25, each 3-5 mm, long, densely plumose with short grayish hairs; seeds ovoid, about 1.5 mm, long and wide. Apparently rare plants known only from Colombia, where they grow on the borders of forests and open grasslands at altitudes of about 750 m.; flowering from October to December, and the fruits maturing in January and February. Colombia: magdalena: pres de Molina, Rio Hacha, Vurdic s.n. (K); Minca, Apolifiar 62J (US), santa marta: Onaca, alt. 2500 ft., H. H. ^m'ltb l688 (G, Mo, NY, UC, US), 1907 (Mo, NY). This is an extremely distinct species, and is characterized by its branches and petioles being densely covered with lustrous ferruginous tomcntum, by the leaves which are profoundly cordate with long attenuate tips, by the large and spreading Inflorescences, and also by its small ellipsoid fruits with about 25, rather densely tomentose spines. 8. Triumfetta Hintonh Sprague, in Kew Bull. 294. 1937. (T.: Hinton 701S). Small trees 2.5-6.0 m. high; branches and inflorescence :ixcs very lightly pubescent, glabrcsccnt. Leaves broadly ovate to elliptical, 8-11 cm. long and 3-5 cm. wide, the base cuneate to rounded on younger ones, subcordate on more mature ones, the tip gradually and narrowly acuminate, the serrations small, blunt, usually double and regular, the upper surface dark green, very lightly pubescent to nearly glabrnte, the lower pallid, slightly more pubescent than the upper; petioles 2-3 cm. long, lightly pubescent. Inflorescences axillary, the cymes of 2 cymules, usually opposite the bracts, the flowering peduncles 3-5 mm. long, the pedicels 2—3 mm. long. Flowers hermaphrodite, the buds oblongoid, 9-11 mm. long at anthesis, the apical appendages very small, nearly obsolete; sepals oblong, 10-12 mm. long, lightly stellate-pubescent without; petals narrowly obovate, 6-7 mm, long, the claw very distinct with a constriction at the tip, about 3 mm. long, densely plumose; gonophore slender, about 2 mm. long, the glands oblong, about half the length of the gonophore; urceolus short, about 0.5 mm. high, 5-lobed, lightly ciliate; stamens 15—20, the filaments smooth, glabrous; ovary ellipsoid, about 1.5 mm. long, the spinules 40-50, erect, the style about 6 mm. long, the 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 343 stigma acute, sometimes briefly 2-lobed. Fru ed but only one maturing, tKe other aborting completely, the spines slightly falcate, short, 1-2 mm. long, only 10-16 remaining at maturity; seeds pyriform, about 2.5 mm. long and 1.5 mm. wide. Known only from central Mexico, growing in pine and oak forests, along streams at altitudes of 2000-2400 m.; flowering from June to September, the fruit maturing from October to November, Mexico: Mexico: Temascaltepcc: La Labor, alt. 2000 m., Hhifon 38^^ (K, US); Cajones, alt. 2480 m., Hinton 4224 (K) ; Nanchititia, Hinton 5324 (K, US); La Labor, Hinton 6594 (GH, Mo, US); Cumbrc dc Tejupllco, Hinton 7018 (K, Mo, US); Pantoja, Hlnfon J02J (Mo^ US), michoacan: Coalcoman, Sierra Torricillas, alt. 2140 m., Hinton J 2 343 (GH). This species has very characteristic fruits. The flower is quite normal for the genus, bearing a 2-loculate ovary with 40-50 straight-pointed spinules. After fertilization, most of the spines fail to develop at the bases of the spinules; thus the mature fruit body becomes rugose In places where the spines have not elongated sufficiently; furthermore, one of the locules aborts completely, and only one 1- seeded locule matures. In their oblongoid fruits and also in their marked tendency towards having a smaller number of spines on the fruit than of spinules on the ovaries, T. Hintonii and T. caiidafa are closely related. 9. Triumfetta Calderoni Standi, in Jour. Wash. Acad. 14:98. 1924. (T.: Calderon yS) . Small trees or large shrubs 6-8 m. high; branches and inflorescence axes covered densely with both fine stellate and long spreading coarse stiff hairs. Leaves broadly ovate to obscurely 3-lobed, 10-15 cm. long and 7-11 cm. wide, the base rounded or subcordatc, the tip gradually and narrowly long-acuminate, the serrations coarse and irregular, blunt and usually glandular, the upper surface dark green, lightly but coarsely pubescent with many short suppressed stellate hairs, the lower pallid, densely pubescent with long spreading stellate and simple hairs; petioles 5—7 cm. long, densely pubescent; bracts elliptical. Inflorescences terminal, large and spreading, with many small, caducous bracts, the cymes of 4—6 cymules. rather lax in flower though generally crowded in fruit, the flowering peduncles about 3—4 mm. long, the pedicels 3—5 mm. long. Flowers hermaphrodite, the buds broadly obovoid, constricted towards the base, about 5 mm. long at anthesis, the apical appendages very small to nearly obsolete; sepals elliptical, 5-7 mm. long, whitish and densely tomentose without, yellow-brown and glabrescent within; petals obovate, small, about 2 mm. long, the claw very short, lightly plumose; gonophore short and stout, about 0,5 mm. long and 1.5 mm. wide, the glands very small, spherical; urceolus about 0.25 mm. high, many-Iobcd, lightly ciliate; stamens 40, the filaments glabrous; ovary ovoid, about 1 mm. long, the spinules about 50, erect, the style 2-3 mm. long, the stigma acute. Fruit spheroid, the body 2-3 mm. in diameter at maturity, rather densely tomentose, 2-locuIate, each cell 1- [Vol. 37 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN seeded, the spines about 50, 4-5 mm. long, densely plumose with relatively long stiff white hairs; seeds pyrlform, about 1.0 mm. long and wide. Common weedy plant of Honduras and El Salvador, chiefly found in dry rocky thickets at altitudes of 450-1000 m.; flowering in October, the fruit maturing in late December to January. Honduras: choluteca: San Marcos, alt. 1000 m., pine forest area, WilUams ^ Molina R. I0Q2J (Mo), el pARAfso: 8 kms. west of Ojo de Agua, Williams ^ Molina R, Io6/6 (Mo), morazan: San Antonio, road near Zamorano, Williams ^ Molina R. IT160 (Mo). El Salvador: san Jose: Caldcron ig2g (GH, US), san Salvador: vie, Tonaca- tepeque, SfanJley 19475 (GH, US); vicinity of San Salvador, alt. 650-850 m., StanJlcy 22/82 (GH, US) ; San Salvador, Caldcron 78 (GH, US), J5^ (US), I2S7 (GH, Mo, US). DATA incomplete: Cboiissy 55 (US). This species can be recognized in fruit by its long slender, rather densely plumose spines, and in flower by the absence of apical appendages on the sepals, and also by the short petals. 10. Triumfetta ferruginea Ko Ko Lay, sp. nov. Ramuli graciles dense pubescentes pilis ferrugineis et stellatis et simpilicibus. Folia late ovata interdum 3-lobata 6-7 cm. longa 5-7 cm, lata base profunde cordata apice longe acuminata supra pilis sparsis stellatis, infra dense canescentia vel ferrugineo-tomentosa; petlolis 3-4 cm. longis dense ferrugineo-tomentosis. In- florescentiae terminales, cymis circiter 6 cymulis in axillis bractearum dense con- fertis, bracteis ellipticis vel lanccolatis, pedunculis plerumque 3 mm. longis, pedicellis 1-2 mm. longis. Florcs hermaphroditi, alabastro maturo obovoideo 6-7 mm. longo; sepalis ellipticis 7-3 mm. longis extra brcviter denseque stellato- pubcscentibus, appcndiculis circiter 2 mm. longis dense plumosis; petalis obovatis brevissimis 1.5-2.0 mm. longis, unguilo circiter 0.5 mm. longo extra dense plumosa; gonophoro circiter 0,5 mm. longo 1.5 mm. lato, glandulis quadrangular- ibus circiter 1.0 mm. longis; urceolo 0.5 mm. alto multilobato breviter ciliato; staminibus 60, filamentis usque 3-4 mm. longis glabris; ovario ovoideo circiter 1 mm. longo, splnulis 30-40 erectis, stylo glabro circiter 4 mm, longo, stigmatc acuto. Fructus maturi desunt. Mexico: chiapas: Las Pilas, June 2, 1904, Goldman I050 (holotype in U. S. Nat. Herb.). Related to T. arborescens from which it may be easily distinguished by its more numerous stamens and much longer apical appendages of the sepals. I have not seen any fruiting specimen of this species. 11. Triumfetta arborescens (Seem.) Sprague, in Kew Bull. 3 51. 1923. Hcliocarpiis arborescens Seem. Bot. Voy. Herald, 86. 1853. (T.: "Panama," Sccmann 5. «. In Herb. Kew.). Small trees 5-6 m. high; branches and inflorescence axes scurfy, densely tomcntose with short ferruginous stellate hairs. Leaves broadly ovate, usually 3- lobed, 10-13 cm. long and 7-10 cm. wide, the base cordate, the middle lobe long- 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 345 L•^' Fig. 6. Triumfctta fcrruglnca attenuate, tlic laterals acute, the serrations very irregular, the basal ones glandular, the upper surface scabrous, covered lightly with many short stellate hairs, the lower densely tomentose with both simple and stellate hairs, becoming less so at maturity; petioles slender, 6-9 cm. long, densely ferruginous-tomentose. Inflores- cences terminal, the cymes of 3—5 cymules, condensed in nodose clusters, the flowering peduncles 2-4 mm. long, the pedicels about 2 mm. long. Flowers hermaphrodite, the buds obovoid, 6—7 mm. long slightly before anthesis, con- stricted towards the base, the apical appendages about 1 mm. long; sepals broadly [Vol. 37 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN elliptical, 7-10 mm. long, pallid and densely tomentose without, ycUow-brown and glabrescent within; petals obovate, short, about 2-3 mm. long, the claw very short, briefly plumose; gonophore slender, about 0.75 mm. long, the glands ellip- tical, subequalling the gonophore; urceolus about 0.5 mm. high, indistinctly lobcd, briefly ciliate; stamens about 50, the filaments smooth, glabrous; ovary ovoid, about 0.75 mm. long, the spinules about 75, erect, the style about 5 mm. long, the stigma very briefly 2-fid. Fruit suborbicular, the body 2-3 mm. in diameter and 3-4 mm. long at maturity, cinereous-pubescent, 2-loculate, each cell 1 -seeded, the spines about 75, 3-4 mm. long, reddish-brown, shortly plumose; seeds pyriform, about 0.75 mm. long and wide. Costa Rica: guanacaste: vicinity of Tilaran, alt, 500-650 m., StamUcy ^ Valeria 45709 (US). Panama: cocle: between Paso del Arado and Ola, In savannahs and thickets, alt. 20-280 m., Pittier jOI/ (US), data incomplete: Scemann g6 (GH), s. w. (K); Cawell 239 (NY). Rather rare small trees of Costa Rica and Panama; flowering in December and the fruit maturing in January and February. 12. Triumfetta persimilis Ko Ko Lay, sp. nov. Frutcx circiter 1 m. altus. Ramuli graciles dense pubescentes, pilis ferrugincis et stcllatis et simplicibus. Folia late ovata vel 3-lobata 10-14 cm. longa 8-10 cm. lata base profunde cordata apice longe acuminata margine irregulariter serrata intcrdum glandulosa supra pilis sparse stellatis scabratis, infra dense canescenti- tomentosa; petiolis 3-4 cm. longis dense tomcntosis. Inflorescentiae termlnalcs, cymis 3-4 cymulis, bracteis ovatis vel obscure 3-lobatis, pedunculis circiter 1 mm. longis, pcdicelHs 6-8 mm. longis. Flores hermaphroditi, alabastro maturo ob- longoideo 6-8 mm. longo; sepalis cllipticis 7-9 mm. longis extra longe ct dense stellato-tomentosis, appendiculls circiter 2 mm. longis dense plumosis; petalis ellipticis, 6-7 mm. longis, unguilo 0.5-1.0 mm. longo extra breviter plumoso; gonophore circiter 0.25 mm. longo, glandulis quadrangularibus; urceolo 0.2 5 mm. alto, 5-lobato breviter ciliato; staminibus 20, filamentis usque ad 5-6 mm. longis glabris; ovario ovoidco circiter 1.5 mm. longo, spinulis 70—80 erectis, stylo glabro 3-4 mm. longo, stigmate breviter 2-fido. Fructus globosus non dehiscens, corporc ipso 2-3 mm. diametro, dense canescenti-tomentosus 2-locularis, loculis 2-spermis, aculeis circiter 75, 3—4 mm. longis dense plumosis; scminibus ovoideo-acutis, circiter 1.5 mm. longis et latis. Coeombia: santander: en terrcnos secos y arenosos a orillas del Rio Chicamocha, en pcscadero, alt. 500 m., Dec. 194S, Araque ^ Barklcy 18S.286 (holotype In Herb. Mo. Bot. Gard., isotype in U. S. Nat. Herb.). In flower this species has a superficial resemblance to T". candafa, but its fruits are much more strikingly similar to those of T. CalderonL It may be easily rec- ognized by its densely tomentose leaves, small oblongoid floral-bud with only 20 stamens, and by its spheroid fruit with many densely plum^ose spines. 19 50] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 347 13. Triumfetta dioica T, S. Brandg. in Univ. Calif, Publ, Bot. 10:412. 1924. (T.: PiirpMS p2j6). Small trees about 3 m. high; branches and inflorescence axes covered densely with many long grayish-brown hairs. Leaves broadly ovate to obscurely 3-lobed, 8-11 cm. long and 5—7 cm. wide, the base deeply cordate, the tip gradually and narrowly long-acuminate, the serrations very irregular, usually blunt, the basal ones glandular, the upper surface dark green, scabrous with many long appresscd simple and stellate hairs; petioles 4-6 cm. long, slender, densely clothed with long spreading hairs. Inflorescences gynodioecious, the pistillate terminal, the cymes of 4-6 cymulcs in dense nodose clusters, the flowering peduncles and pedicels about 2 mm. long. Hermaphrodite flower not seen, the pistillate small, the floral bud 3-4 mm. long slightly before anthesis, the apical appendages short, 1 mm. long or less; sepals narrowly elliptical, 4-5 mm. long, paUid and densely tomcntose without, yellow-brown and glabrescent within; petals elliptical to narrowly obo- vate, small, about 1.5 mm. long, the claw nearly obsolete; gonophore short, about 0.5 mm. long, the glands oblong, subcqualling the gonophore; urceolus about 0.5 mm. high, many-lobed, briefly ciliate; ovary ovoid, about 1.5 mm. long, the splnules about 50, erect, the style about 2 mm. long, the stigma briefly 2-fid. Fruit spheroid, covered with about 50 slender, densely plumose spines; mature fruit not seen. Known only from Chiapas, Mexico. Flowers in September. Mexico: chiapas: Hacienda Monserrate, Purpus Q2$6 (Mo, UC, US); Buena Vista, Purpus 928s (UC). It is unfortunate that this species is so imperfectly known. Due to the lack of mature fruits and hermaphrodite flowers its proper position is rather obscure. It is apparently related to T. Calderoni in the possession of small flowers and spheroid fruits with numerous plumose spines. 14» Triuaifetta attenuata Ko Ko Lay, sp. nov. RamuH glabri irregulariter punctati lenticellis multis parvis tecti. FoUa late ovata 10-14 cm. longa 6-9 cm. lata base subtruncata vel rotundata apice longe attenuata subdupliciter scrrata utrlnque glabrata; petiolis graciUbus 6-9 cm. longis glabris. Inflorescentiae terminales, cymis 4-5 cymulis, peduncuUs 5-7 mm. longis, pedicellis 4-5 mm. longis. Flores hermaphroditi, alabastro maturo obovato 6-7 mm. longo; sepalis ellipticis vel anguste obovatis 7-9 mm. longis extra glabrescenti- bus, appendiculis circiter 2 mm, longis glabris; petalis obovatis 6-7 mm. longis, unguilo circiter 1.5 mm. longo dense plumoso; gonophoro circiter 0.75 mm. longo 1.5 mm. lato, glandulis ellipticis 0.5-0.75 mm. longis; urceolo 0.5 mm, alto 5- lobato breviter ciliato; staminibus 60, filamentis usque ad 5-7 mm. longis glabris, spinulis circiter 75 erectis, stylo glabro circiter 6 mm. longo, stlgmate acuto. Fructus maturi desunt. Mexico: chiapas: Escuintla, Dec. 1936, Matnda 0488 (holotype in Herb. Mo. Bot. Card.; isotypes in Herb. Univ. Michigan, and U. S. Nat. Herb.). [Vol. 37 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species IS most closely related to T. Mexiae, from which it may be dis- tinguished easily by its glabrate leaves and floral buds, the much longer apical appendages of the sepals, and its style which is about 6 mm. long with the stigma cxscrted above the stamens. It is unfortunate that no fruiting material of this species is available. 15. TiuuMFETTA Mcxiac C. V. Morton & Ko Ko Lay, sp. nov. Arborcs parvae 5-6 m. altae. Ramuli inferne glabrati apicem versus tomcn- tosi pilis multis brevibus stellatls. Folia late ovata 8-11 cm. longa 6-9 cm. lata base cuncata vcl rotundata apice longe acuminata subdupllcltcr serrata utrinquc pilis sparsis stcllatis mox glabrescentia, nervis subtus palHdiorlbus pllis brevibus stellatis; petiolis 3—5 cm. longis sparse stellato-pubesccntibus. Inflorcsccntlae terminales, cymls cymulis 4—5, pedunculis 2—3 mm. longis, pcdicellis 3—4 mm. longis. Flores hermaphrodlti, alabastro maturo obovoidco 5—6 mm. longo; scpalis angustc obovatis 5—8 mm. longis extra breviter stellato-pubcscentibus; pet alls ellipticis ve! anguste obovatis 4—6 mm. longis, unguilo clrciter 2 mm. longo dense plumoso; gonophoro circitcr 0.75 mm. longo 1.0 mm. lato, gland ulis ellipticis circiter 0.75 mm. longis; urceolo 0.25 mm, alto multilobato breviter ciliato; staminibus 40, filamcntis usque ad 3—4 mm. longis glabris; ovario ovoideo circitcr 1.5 mm. longo, stlgmate breviter 2-fido. Fructus globosi non dehiscentes, gyno- phoro brcvi supra gonophorum, corporc ipso 3-4 mm. diametro glabro 2-loculari, loculls 2-spcrmis, aculeis 75-100 graciUbus glabris; seminibus ovoideo-acutis circitcr 2 mm. longis et latis, A rather common species of southwestern Mexico, growing at altitudes of 1000 to 1200 m.; flowering in September and early October, the fruits maturing from late October to November. Mflxico: GUERRERO: Sicrra Madre del Sur, north of Rio Balsas, Distr. Adama, Tcmisco, barranca below Ore Mill, alt. 1265 m., Mcxia SSj4 (holotype in U. S. Nat. Herb.; iso- types In Gray Herb., Herb. Mo. Bot. Gard., and Herb. Univ. Calif.); mountains near Iguab, alt. 2500 ft., Pringlc 9228 (GH, MICH, US); Iguala Canon, ?riugJc JOodgYz (GH, MICH, US); between Sochi and Tlalkinsala, alt. 3600-4800 ft.. Nelson 2036 (NY, US) ; Coyuca-Chacamerito, Hinton 6621 (US), michoacan: Zitacuaro-La Mora, alt. 1325 m., Hintoti 13230 (GH), There is a great superficial resemblance In the aspects of the flowering specimens of this species to those of Heliocarpus occidcntalis Rose. It is, however, quite distinct from other species of Trhunfetta and can easily be recognized. Series Uncinatae Sprague & Hutch, in Jour. Linn. Soc. Bot. 39:240. 1909 KEY TO THE SPECIES A. Flowers hermaphrodite, the buds oblongold, the style at least 3 times the length of the ovary; fruits distinctly pubescent. B. Petioles usually less than 1.5 cm. long, the leaves elliptical, at least twice as long as broad; southwestern Aiexico. C. Fruit with more than 100 short, straight, glabrous spines, each about 1 mm. long; sepals at anthesis 13—16 mm. long; stamens 25-30 1(5. r. brnipcs 19 50] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 349 J/ ^/^, y. '4^ V T; )^^^. >C, M-^4¥ - .^ r-^ j\ yr-.-'^K ' V. ' y ^' J ^ i>V. ^ V Ki ■:;v * * L» * 1 3 4 2 4^ # -^ ;( ^■/ -.y h ■ .J , I ':* ^ 6 7 8 9 10 f^a^ &^ Fig. 7. Illustrations of taxonomic criteria for Series Uncinatae: 1. Narrowly oblongoid floral bud of T. Palmcrl with relatively short apical appendages 2. Narrowly oblongoid floral bud of T, Purpiisii with very long apical appendages. 3. Obovoid floral bud of T, Galrottiana. 4. Younger floral bud of T. Bartramia with very deeply cucullate sepals. 5. Broadly obovate petal of T. bogotends, 6. Elliptical to narrowly obovate petal of T. obscura, 7. Retrorsely pilosulose spine of T. semifriloba, 8. Densely plumose spine of T, socorrens'n, 9. Irregularly pubescent spine of T. discolor with few short hairs. 10. Glabrate spine of T. panicnlata. [Vol. 37 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN CC. Fruit with less than 75 falcate, pubescent spines, each about 2—3 mm. long; sepals at anthesis 15—20 mm. long; stamens 20.... 17. T. PaJmcri BB. Petioles generally much longer than 2.0 cm., the leaves ovate (ex- cept in r. obscura)^ usually much less than twice as long as broad. C. Stamens 20-40 (except T. Lappula with 5—15); mature fruit (including spines) 7—15 mm. thick. D. Spines plumose or irregularly pubescent; sepals at anthesis 1.2—3.0 cm. long, E. Fruit body oblongoid with about 20 spines, each ^-10 mm, long; sepals at anthesis 12—14 mm. long; Revilla Gigedo Islands, Mexico 18. T. socorrcnsh HE. Fruit body globose with 40 or more spines, each usually 2—5 mm. long; sepals at anthesis 1.5-3.0 cm, long. F. Fruit with 40-50 spines; apical appendages of the sepals 5—10 mm. long; stamens about 40. G. Spines densely plumose; apical appendages of the sepals 7-10 mm. long; southeastern Mexico 19. T. Purpusii GG. Spines slightly pubescent to nearly glabrate; apical appendages of the sepals 5—7 mm. long; southwestern Mexico 20. r. falcifcra FF. Fruit with 75-200 spines; apical appendages of the sepals shorter than 3 mm.; stamens 15—30. G. Leaves densely tomentose on both surfaces; mature fruit 6- to 10-loculate, each cell 1 -seeded. H. Mature fruit 8- or 10-loculatc; sepals at anthesis 1.8-2.7 cm. long, the apical appendages 2—3 mm, long. I. Mature fruit 5- to 10-locuIate, the spines and body very shortly pubescent; sepals broadly obovate, the stamens 25-30; Peru 21. T. viulfilocularh IT. Mature fruit 8-loculate, the spines and body with lon.^ plumose hairs; sepals elliptical to narrowly obovate; stamens 20; Colombia 22. T. inoUhnma HH. Mature fruit 6-loculate; sepals at anthesis about \.7 cm. long, the apical appendages about 1 nim. long; northeastern Brazil 23. T. lorigicama GG. Leaves densely tomentose on lower surfaces only or both surfaces lightly pubescent to glabresccnt; fruit with 3, 1- to 2-seedcd locules. H. Sepals at anthesis 22-3 mm. long; fruit indehisccnt insofar as known. I. Leaves 11-16 cm. long and 6-9 cm. wide, or slightly larger; fruit stellate-pubescent, spines brief- ly but distinctly pubescent. J. Leaves 3-lobcd, without tufts of hairs at axils of the veins on the lower surface; filaments lightly stellate-pubescent at the base; spines very lightly pubescent with short hairs; northwestern Mexico 24. T. chihnahucnsh JJ. Leaves not lobed, with many long tufts of hairs at the axils of the veins on the lower surface; filaments not pubescent; spines rather densely pubescent; west-central Mexico 25. T. harhosa IL Leaves 6-8 cm. long and 3—4 cm. wide; fruit tomentose, the spines glabrescent; central and southwestern Mexico 26. T. columnarh HH. Sepals at anthesis 15-20 mm. long; fruit easily dehiscent. I, Leaves distinctly longer than broad, gradually and narrowly acuminate, both surfaces lightly pubescent tc glabrescent; fruit with 100 or more slender spines, each about 4 mm. long; southern Mexico and West Indies to northern South America 27. T. granJijlora 1950] KO KO LAY — AMERICAN SPECIES OF TRIUMFETTA 351 II. Leaves nearly as broad as long, shortly acute to acuminate, tomentose beneath; fruit with about 75 relatively thick spines, each about 2-3 mm. long; northwestern Mexico 28. T. discolor DD. Spines retrorsely pilosulosc; sepals at anthcsis 3-9 mm, long (except T. bogoicnsis and T. abutiJoldes sometimes up to 13 mm. long), E. Petals present; fruit and flower with a 5-glandular gono- phore, the urceolus manifest. F, Leaves with both stellate and simple hairs, the latter prominent on both surfaces; sepals at anthesis 8-13 mm. long, the. petals broadly obovate, G. Leaves densely tomentose on both surfaces; apical ap- pendages of the sepals nearly obsolete or less than 1 mm, long; fruit with 6, 1 -seeded locules at maturity; Haiti and northern South America 29. T. abutiloidcs GG. Leaves lightly pubescent on both surfaces; apical ap- pendages of the sepals 2-3 mm. long; fruit with 3, 2-seeded locules at maturity; pantropical 30. 7*. bogofcjish FF. Leaves with stellate pubescence only; sepals at anthesis 5—9 mm. long, the petals linear to narrowly obovate. G. Leaves and bracts narrowly elliptical, at least twice as long as broad; northeastern South America 31. T.obscura GG. Leaves ovate, generally 3-lobed, less than twice as long as broad, the bracts elliptical. H. Leaves slightly pubescent, glabrcscent; petals less than half the length of the sepals; mature fruit body about 3 mm, thick; western Mexico 32. T*. acracantha HH. Leaves coarsely stellate-pubescent ; petals subcqual- ling the sepals; mature fruit body usually 4 mm. thick or more. I, Petioles with 4 prominent glands on the basal sinus; mature fruit 8- to 10-loculate, the spines numerous, about 200; Trinidad and northern South America.. 3 3, T*. althacoides II. Petioles without glands at the basal sinus; fruit 3-loculate, the spines 75 or fewer. J. Apical appendages of the sepals 2 mm. long or less; fruit body lightly to densely pubescent, spines about 75; pantropical 34. T . semitriloba JJ. Apical appendages of the sepals 3 mm. long or more; fruit body tomentose, the spines 2 5—40; Ecuador and Peru 3 5. T, oligacantha EE. Petals absent; gonophore and urceolus obsolete in fruit and flower, the glands absent; pantropical 36. T, Lappula Stamens 5—15; mature fruit (including spines) 4-6 mm. thick. D. Sepals very slightly cucullate in younger floral buds; fruit very lightly pubescent. E. Leaves densely tomentose on both surfaces, 9-12 cm. long and 7—10 cm. wide; fruit with few, 50 or less, spines, each about 2 mm. long; Colombia and Ecuador 37. T". scricafa EE. Leaves lightly pubescent on both surfaces, glabrcscent, about 2 cm, long and 1 cm. wide; fruit with numerous, 75—100 short spines, each about 1 mm, long or less; south- western Mexico 3 8. T. brachhfacantha DD. Sepals cucullate about half their length In young floral buds; fruit densely pubescent. E. Floral buds 5—8 mm. long slightly before anthesis, the urceolus manifest, the stamens 1 0—1 5 ; fruit body densely cinereous-tomentosc, the spines glabrcscent; West Indies and northern South America 39. T. Barframla EE. Floral buds 3—4 mm. long slightly before anthesis, the urceolus absent, the stamens 5; fruit body and spines densely plumose; northeastern Brazil 40. T. Sampaioi [Vol, ^7 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN AA, Flowers either hermaphrodite or pistillate, the hermaphroditic buds obovoid, the style about twice the length of the ovary; fruits es- sentially glabrous. B. Leaves 15-18 cm. long and 16-20 cm. wide, 3- to 5-Iobed; stamens 30-40; mature fruit body 5-6 mm. thick, the spines 3-5 mm. long; western Mexico 41. T. i)aiiicnldta BB. Leaves usually 7-15 cm. long and 5-12 cm. wide, not lobed to s!Ightly 3-lobed; stamens 20-30 in hermaphroditic flowers, stam- inodial or absent in pistillate; mature fruit body 2-4 mm. thick. C. Pistillate flower apetalous; (hermaphroditic floral bud 5-7 mm. long slightly before anthesis, the petals about 3-4 mm. long); fruit body 3-4 mm. thick, the spines 30-40; southern Mexico... 42. T. Galcotfiana CC. Pistillate flower with petals, each about 1 mm. long; (hermaphro- ditic flower not seen); mature fruit body about 2 mm. thick, the spines about 20; central Mexico 43. T, hciiocarpoiJcs 16. Triumfetta brevipes S. Wats, in Proc. Am. Acad. 22:400. 1S87. (T.: E. Pahfier 323) , Spreading busliy sKrubs 1-2 m. high; branches and inflorescence axes rough, scurfy, fcrruginous-tomcntose. Leaves broadly elhptical, about 11-13 cm. long and 3-4 cm. wide, the base obtuse, the tip gradually acuminate, the serrations unequal and blunt, usually glandular, the upper surface lightly but coarsely stellate-pubescent, the lower densely stellate-tomcntose; petioles 1.0-1.5 cm. long, densely tomentose. Inflorescences axillary, the cymes of 2, rarely 3, cymules, gen- erally condensed, the flowering peduncles 3-4 mm. long, the pedicels short, about 2 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, 1.0-1.4 cm. long at anthcsis, sHghtly constricted in the middle, the apical appendages slender, about 2-3 mm. long, slightly pubescent; sepals oblong, about 13-16 mm. long including the apical appendages, densely tomentose without, glabresccnt within; petals narrowly obovate, about 10 mm. long, the claw about 4 mm. long, shortly plumose; gonophorc about 1.5 mm. long and nearly as wide, the glands broadly oblong, 1.0-1.5 mm. long and 0.75-1.0 mm. wide; urceolus large, about 1 mm. high or more, indistinctly lobed, slightly ciliatc; stamens 2 5-3 0, the filaments smooth, without serrations; ovary orbicular, about 1.5 mm. long, the splnules numerous, recurved, the style about 8-9 mm. long, the stigma acute. Fruit or- bicular, the body about 6-7 mm. in diameter at maturity, matted cinereous- tomentose, 3-loculate, each cell 1-seeded, the spines numerous, 100 or more, straight, glabrous, each about 1 mm. long; seeds pyriform, 3.0-3.5 mm. long and 2.5-3.0 mm. wide. A common ground cover in pine and oak forests or rocky hillsides at altitudes of 1000 m. or more in southwestern Mexico; flowering in August and September, the fruit maturing from late September to mid-October, Mfxico: GUERRi kg: Tierras Blancas, MIna, alt. 1400 m., Hinfon gg22 (GH, US). JALISCO: Rio Blanco, £. Palmer 323 (GH, US); rocky hills under oaks near Guadalajara, VrUigle 4302 (BR, GH, Mo, US); barranca of Rio Blanco, near Guadalajara, alt. 4500 ft., Vr'niglc I13S8 (GH, US); near Chapala, Rose ^ Pafvfcr T6g3 (GH, US), michoacan: road from Tancitaro to Apatzingan, alt. 3400 ft., Leavenworth I'^JI (Mo); vicinity of Morclia, Loma Sta. Maria, alt. 1950 m., Arshie 2316 (K, Mo, P, US); vicinity of Morelia, vers la Huerta, alt. 1950 m., Arseue 5131 (GH, Mo, US); vicinity of Morclia, alt. 1900 m., Aru'uc 334^ (GH, Mo, US). 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 353 Tlie shortly petiolate elliptic leaves, the orbicular fruit with numerous very short glabrous spines, and the oblongoid floral bud 10-14 mm, long at about anthesis, are characters which distinguish this species. 17. Triumfetta Palmeri S. Wats, in Proc. Am. Acad. 22:400. 1887. (T.: £. Palmer 330). Low shrubs about 1 m. high; branches and inflorescence axes scurfy, ferru- ginous-tomentose. Leaves narrowly elliptic, about 6-8 cm. long and 2—3 cm. wide, the base obtuse, the tip gradually acuminate, the serrations rather regular and double, the upper surface lightly stellatc-pubcscentj the lower more densely so; petioles very short, not longer than 1 cm., densely tomentose, nearly obsolete in the bracts. Inflorescences axillary, the cymes usually of 3 cymulcs, rarely 4, each cymule 2-, rarely 3 -flowered, the flowering peduncles variable in length, usually 4-6 mm. long, the pedicels about 2-3 mm, long. Flowers hermaphrodite, the buds narrowly oblongoid, 14-18 mm. long slightly before anthesis, usually falcate, the apical appendages short, slender, 1—2 mm. long, slightly pubescent; sepals narrowly ovate, about 15-20 mm. long including the apical appendages, densely tomentose without, glabrcscent within; petals about 8—9 mm. long, narrowly obovatc, with a claw about 2 mm. long, very densely plumose; gonophore slender, 1.0—1.5 mm. long, the glands oblong, about 1 mm. long and 0.5 mm. wide; urceolus very short, about 0.25 mm. high, rather obscurely 5-lobcd, briefly ciliate; stamens 20, the filaments without serrations, slightly pubescent at the base; ovary orbicular, about 1 mm. long, the spinules numerous, recurved, the style 10—12 mm. long, the stigma acute. Fruit orbicular, the body about 4-5 mm. in diameter at maturity, with many short, stellate hairs, 3-loculate, each cell 1-seeded, the spines falcate, about 75, lightly pubescent to rctrorscly pilosulose, 2-3 mm. long; seeds pyriform, about 3.0 mm. long and 2.5 mm. wide. Southwestern Mexico, usually at altitudes of 1000-2000 m.; flowering in mid- August, and the fruit maturing from mid-September to early October. Mexico: durango: Sierra Madrc, Rose 35^^ (GH). jalisco: Rio Blanco, £. 'Palmer SJO (Mo, US) ; on the road between Colotlan and Bolanos, Rose 2838 (US) ; Sierra Madre, west of Bolanos, Rose 3701 (US) ; near Guadalajara, Rose ^ Painter 7447 (US) ; barrancas near Guadalajara, Pringle 4493 (BR, Mo, US); barranca near Rio Blanco near Guadala- jara, alt. 4500 ft., Pringle Il38g (US). This species is closely related to T. hreiipes, but may be distinguished from the latter by its floral buds, which are 3-4 mm. longer at about anthesis, and also by Its falcate, relatively longer, pubescent spines. 18. Triumfetta socorrensis T. S. Brandg. in Erythea 7:1, 1899. (T.: Anthony 37 S) . Handsome shrubs about 1.5-2.0 m. high; branches and inflorescence axes rough, scurfy, ferruginous-tomcntose. Leaves broadly ovate to rhombic-ovate, about 5-6 cm. long and 4-5 cm, wide, the base rounded or cordate, the tip abruptly acumi- nate, the serrations very irregular, unequal and blunt, the upper surface coarsely stellate-tomcntose, the lower more densely so; petioles about 2.0 cm. long, stout, [Vol. 37 354 ANNALS OF THE MISSOURI BOTANICAL GARDEN densely tomcntose. Inflorescences axillary, the cymes of 3-4, very rarely 2, cymules, generally condensed, the flowering peduncles very short, 1—2 mm. long, the pedicels 2—3 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, about 9-11 mm. long slightly before anthesis, very slightly constricted near the base, the apical appendages short, about 1 mm. long; sepals broadly elliptical, about 12-13 mm. long at anthesis including the apical appendages, densely tomen- tose without, glabrescent and greenish within; petals broadly obovate, about 8-9 mm. long, the claw about 2-3 mm. long, densely plumose; gonophore short and stout, about 0.75 mm. long and 1.5 mm. thick, the glands small, sph.erical; urceolus short, about 0.5 mm. high, distinctly many-lobed, slightly ciliate; stamens about 20, tlie filanients glabrous; ovary orbicular, about 1.5 mm. long, the splnulcs about 3 0, recurved, the style about 3-4 times the length of the ovary, the stigma very briefly 2-fid. Fruit oblongoid, the body about 3-4 mm. in diameter and 5-6 mm. long at maturity, densely stellate-tomentose, 3-loculate, each cell 1-seeded, the spines about 15-20, 6-10 mm. long, densely pubescent v/ith many short spreading hairs; seeds pyriform, 2.0-3.0 mm, long and 1.5-2.0 mm. wide. Known only from Socorro Island adjacent to the mainland off the coast of Baja California, Mexico; flowering and fruiting from March to June. Mexico: revilla gigedo islands: I. Socorro: Anthony 3j8 (Mo, NY, UC, US); Barkclcw IQ7 (Mo, UC, US); Grayson's Cove, Mason l6j2 (Mo, US); dry slopes, north anchorage, Howell 8444 i^US). A very distinct species and an endemic, it is conveniently recognizable by its densely tomentose, rhombic-ovate leaves with very short petioles, and also by its very characteristic oblongoid fruits with few densely plumose, long spines. In its vegetative and floral characters, it is closely related to T. bveilpes, a common species of southwestern Mexico, and in its fruit characters to T. Pnrpnsii, an endemic from Vera Cruz, Mexico. 19. Triumfetta pL^RPUsn Standi, in Field Mus. Publ. Bot. 11:167. 1936. (T.: Pur pits goog). Tall sparingly branched shrubs about 1.5 m. high; branches and inflorescence axes rough, scurfy, ferruginous-tomentose. Leaves broadly ovate, about 8-9 cm. long and 4-5 cm. wide, the base subcordate, the tip gradually and narrowly acuminate, the serrations unequal and blunt, generally glandular, the upper surface lightly stellate-pubescent, the lower densely tomentose; petioles 3-5 cm. long, densely stellate-pubescent. Inflorescences axillary, the cymes of 2, rarely 3, cymules, generally condensed, the flowering peduncles about 2-3 mm, long, the pedicels of about the same length. Flowers hermaphrodite, the buds broadly oblongoid, 8—12 mm. long slightly before anthesis, the apical appendages very slender, about 6—10 mm. long, usually reflexcd vipon the bud, densely plumose; sepals oblong, about 16-20 mm. long including the apical appendages, densely long-tomentose without, glabrescent within; petals obovate to narrowly so, about 6—8 mm. long, the claw about 4 mm. long, densely plumose; gonophore about 1.0 mm. long and 1.5-2.0 mm. wide, the glands large, oblong, 1.0—1.5 mm. long and 0.75—1.0 mm. wide; 195 0] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 355 urceolus small, about 0.5 mm. high, very deeply many-lobed, lightly ciliate; stamens about 40, the filaments smooth; ovary orbicular, about 1 mm. long, the spinules about 50, recurved, the style about 8-9 mm. long, the stigma acute. Fruit suborbicular, the body about 5 mm. wide, very densely tomentose, 3-loculate, each cell I -seeded, the spines straight, about 40, each about 2 mm. long, with many long plumose hairs; seeds pyriform, 2.0-2.5 mm. long and 1.5-2.0 mm. wide. Known only from Vera Cruz, Mexico. Flowers and fruits from November to March, Mexico: vera cruz: Mata 2ar2a, Purpus QOOQ (FM, GH, Mo, NY, UC, US); barranca de Panoaya, Pjirpus 86og (GH, Mo, NY, UC, US) ; Bancs de Azufre, Purpns 8274 (GH, Mo, NY, UC, US); Remudatero, Vtirpm Iiogg (Mo, US). The presence of very long appendages of the sepals makes T. Purptisii and T. falcifera very distinct from the rest of the species. This species can be distinguished from the latter, in flower by its calycine apical appendages, which are not only longer by about 2 mm. or more but are also more densely plumose; and in fruit, by the densely plumose body and the spines. 20. Trtumfetta falcifera Rose, in Contr. U. S. Nat. Herb. 12:28 5. 1909. (T.: E. Painter dj). Bushy shrubs about 2 m. high; branches and inflorescence axes smooth with many long, spreading, stellate hairs. Leaves narrowly to broadly ovate, 12-15 cm. long and 6—9 cm. wide, the base obtuse to subcordate, the tip narrowly and gradually acuminate, the serrations rather regular, generally glandular, the upper surface lightly stellate-pubescent, the lower more densely and softly so; petioles 5-7 cm. long with long, spreading, stellate tomentum. Inflorescences generally axillary, sometimes terminal, the cymes of 1 cymule, the flowering peduncles about 8 mm. long and the pedicels about 6 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, 6-10 mm. long at anthesis, the apical appendages very slender, 5-7 mm. long, usually spreading, rather densely plumose; sepals oblong, about 12-16 mm. long including the apical appendages, lightly long-plumose without, brownish and glabrescent within; petals obovate, about 3-4 mm. long, the claw about 1 mm. long, lightly plumose; gonophore about 1 mm. long, nearly 2 mm. wide, the glands narrowly elliptical; urceolus rather short, 0.25-0.5 mm. high, distinctly many-lobed, lightly ciliate; stamens about 40, the filaments glabrous; ovary orbicular, about 1.5 mm. long, the spinules about 40, recurved, the style 6-7 mm. long, the stigma acute, sometimes very briefly 2-fid. Fruit orbicular, about 4 mm. in diameter at maturity, very lightly pubescent, usually 3-locuIate, sometimes 4- to 6-celIed by false-septation, each cell 1 -seeded, the spines about 40, straight, essentially glabrous, each about 2 mm. long; seeds pyriform, 2.0-2.5 mm. long and 1.5-2.0 mm. wide. Southern Mexico; flowering and fruiting in September and October. Mexico: guerrero: In Acapulco and vicinity, £. Palmer 6j (US), 266 (Mo, UC, US); Baqucta, alt. 200 m., Langlassc §02 (P, US); Acapulco, presqu'lles Griffon, Le Jolts s.n. (G). tabasco: Jalapa, Tequisatlan, Lkbmann j02 (US), data incomplete: Liebrnann 534A (US). I [Vol. Z7 356 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2L Triumfetta multilocularis Hochr. in Ann. Conserv. Jard. Bot. Geneve 18:104. 1914. (T.: Pavon s.n. in Herb. Geneve). Small trees or shrubs 2-4 m. high; branches and inflorescence axes rough, rather densely tomentose, with relatively long, ferruginous, stellate hairs. Leaves broadly ovate, usually 3-lobed, 8-10 cm. long and 6-7 cm. wide, cordate, narrowly and gradually acuminate, the serrations very irregular and rather large, not gland- ular, the upper surface coarsely stellate-pubescent, the lower more softly and densely so; petioles 4-6 cm. long, rather stout, with many spreading, long, stellate hairs. Inflorescences axillary, the cymes usually of 1, rarely 2, cymulcs, loose and spreading, the bracts usually broadly ovate, not lobed, the flowering peduncles 8-12 mm. long, the pedicels about 6 mm. long. Flowers hermaphrodite, the buds broadly oblongold, 22-25 mm. long and 3-4 mm. wide slightly before anthcsis, the apical appendages rather stout, 2-3 mm. long, lightly pubescent; sepals broadly obovate, about 27 mm. long including the apical appendages, stellate-pubescent without, glabrescent within; petals broadly obovate, showy, about 24 mm. long and 4-6 mm. wide, the claw about 4 mm. long, densely plumose; gonophorc large, about 1.5-2.0 mm. long and wide, the glands quadrangular, large, about 1.5 mm. long and 1.0 mm. wide; urceolus 10-lobcd, about 0.75 mm. high, distinctly ciliate; stamens 2 5—30, the filaments retrorsely 2-serrate, slightly pubescent at the base; ovary ovoid, 1.5—2.0 mm. long, the spines numerous, recurved, the style about 22 mm. long, pubescent at the base, the stigma acute. Fruit orbicular, the body about 6-8 mm. in diameter at maturity, lightly and briefly stellate-pubescent, 3- to 5-loculate, each cell 2-seeded, later falsely 8- to 10-loculate, each cell with 1 seed, the spines numerous, short, 1-2 mm. long, very shortly pubescent; seeds ovoid, about 2 mm. long and broad. Peru; growing at altitudes of 2000-3 000 m.; flowering and fruiting from June to December. Peru: ayacucho: between Huanata and Rio Apurimac, alt. 2200 m., Killip ^ Sniilh 223OP (US), cuzco: San Miguel, Urubamba Valley, alt. 1800 m., Cook & Gilbert 1126, Il6j (US); Prov. Quispicanchis, Dist. Marcapata, Vargas 9685 (Mo); below Machu Plcchu, lower valley slopes, alt. 2100 m.. West 64/3 (Mo), huanuco: "Fl. dc Huaya- quil", Pavon s. ft. (G); *'Herbier dc Ventcnat", 5. «. (G, FM photo); Acomayo, upon banks of Rio Acomayo, alt. 2100 m., Woytkowski 34013 (Mo), junin: Prov. Tarma, Carpapata, alt. 2400 m., Ochoa 2Sj (US); Carpapata, above Huacapistana, alt. 2400 m., K////> ^ Smith 24361 (US); Huacapistana, Rose d Rose iSjjO (NY, US); Prov. Tarma, between Palca and Carpapata, alt. 2900 m., S/o?7^ lOp/'O (Mo), lima: Escucla Nac'l. Agric, from Prov. Sandia &: Huacapistana and Tarma, Mexia 4^^^5 (Mo), data incom- plete: Amazonian slope of the Andes, alt. 10,000-11,000 ft., B. Ehle s. n. (NY). Ornamental shrub or small tree with long wlilp-like brandies. Though suf- ficiently distinct from T. vwUissima^ it is usually misidentificd with it. The much larger flowers with broadly obovate, rather showy petals, and the pubescent to nearly glabrescent fruits, with numerous very short spines, clearly distinguish this species from the former. 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 357 22. Triumfetta mollissima HBK. Nov. Gen. & Sp. 5:345. 1821. (T.: "S. Fc dc Bogota/* Bonpland s.n. in Herb. Paris.). Tr'ni7)ifetta aciunhiata HBK. loc. cit. 344, 1821. (T.: Bonpland s.n, in Herb. Paris,). Shrubs about 1-2 m. high; branches and inflorescence axes rough, very densely long ferruginous-tomentose. Leaves broadly ovate to 3-lobed, 12—15 cm. long and 6-8 cm. wide, rounded or subcordate at the base, gradually and narrowly acuminate, the serrations very irregular, blunt and generally glandular, the upper surface scabrous, very pubescent, the lower smooth, densely tomentose; petioles 3—5 cm. long, rather stout, with long spreading ferruginous tomentum. Inflores- cences axillary, the cymes of 2-3, rarely 1, cymules, rather lax, the bracts broadly lanceolate to elliptical, long-acuminate, the flowering peduncles about 8 mm. long, the pedicels about 6 mm. long. Flowers hermaphrodite, the buds narrowly ob- longoid, 15-20 mm. long and about 2 mm. wide slightly before anthesis, the apical appendages slender, 1-2 mm. long, very lightly pubescent; sepals narrowly ellip- tical, 18-22 mm. long at anthesis, with many long spreading stellate hairs without, glabrescent within; petals linear, 16-18 mm. long, the claw 3-4 mm. long, densely plumose; gonophore rather large, about 1 mm. long and 1-2 mm. wide, the glands quadrangular, large, subcqualling the gonophore; urceolus about 1 mm. high, dis- tinctly 5-lobed, densely ciliate; stamens 20, the filaments slightly pubescent at the base, smooth; ovary orbicular, about 2 mm. long, the spinules numerous, recurved, the style 16-17 mm. long, the stigma acute. Fruit orbicular, the body 6-8 mm. in diameter at maturity, densely tomentose with long spreading hairs, 4-loculatc, each cell 2-seedcd, later falsely 8-Ioculate with 1 seed to a cell, the spines numerous, 100 or more, each 2-3 mm. long, long-plumose at the base; seeds ovoid, about 2 mm. long and broad. Known only from Colombia; growing in dry open areas, at altitudes of 1200— 2000 m.; flowering and fruiting from May to November. Colombia: antioquia: vie. Medcllin, Toro IT/', 50T, yoi (NY); Dabclba, Johnson & Barkley i8c 6/6 (US); Bello, alt. 1500-1800 m., Archer 2l8 (US), caldas: Supb, Cordillera Occidcntnl, ah. 1200-1500 m., Venncll lO/lo (NY), cauca: forests in high- lands of Popayan, alt. 1700-2000 m., Lehinann 5521 (NY, US); Tolima, Lchmann 2ig6 (G); Rio Sucio to iRio Piedras, west of Popaydn, alt. 1500-1700 m., Venncll ^ Killip 822J (NY); La Manuelita, near Palmira, eastern side of Cauca Valley, alt. 1100-1300 m., Vitticr 818 (US); Municipio Popaydn, alt. 1800 m., Agrcdo 2j2 (US); El Tambo, Verticnte Oriental, alt. 1700 m., Sncrdern 4743, 5559, 56JI (US); Popaydn, matorraies en Rio Blanco, alt. 1800 m., Arbelaez & Ciiatrecasas 5761 (US); El Tambo, Garcia B. 4418 (US), cundinamarca: Estacion Santana, arriba do Sasaima, alt. 1600-1700 m., Dugand e> Jaramillo 3845 (US); cntre Pandi c Icononzo, alt. 980-1340 ni., Garcia B. 11995 (US); Villete, Arhte-Joscph s,n., 5373 (US), distrito federal; S. Fc dc Bogota, Bonpland s. n. (P, Mo photo, NY); exact locality lacking, Bonpland s. n, (P, Mo photo), valle: Palmira, Cordillera Central, alt. 1200-1800 m., Pcnncll ^ Killip 6101 (NY); Pichindo, bank of Rio Cali, alt. 1600 m., Duqne 1474 (US); La Cumbre, Cordillera Occidental, alt. 1500-1700 m., Venncll 5214 (US); Rio Bolo, alt. 1000 m., Dryandcr 253, 254 (US), data incomplete: Triana 32^0 (US); MufiK 2j88, 417 1 (US); Picdra dc Moler, Holton 77S (NY); Linden s, n. (NY). [Vol. 37 358 ANNALS OF THE MISSOURI BOTANICAL GARDEN The reason for the discrepancy in the description of the leaf -shape of T. actnn'niata is very obviously due to the fact that the authors mistook the bracts for vegetative leaves. I have seen the types of both T. mollhsima and T. acuminata and am convinced that they are conspeciiic. The type of T. mollissima has both the vegetative leaves and the bracts, while that of T. acuminata has only the bracts. 23. TRiUMrETTA LONGicoMA St. Hil. Fl. Bras. Mcrid. 1:221. 1825. (T.: St. Hilatre B. IQ) . Older branches smooth, glabrescent, the younger ones and inflorescence axes longitudinally ridged, clothed with long stellate hairs. Leaves broadly ovate, about 8 cm. long and 4-6 cm. wide, cordate, narrowly and gradually long-acuminate, both surfaces with scattered tufts of long stellate hairs, especially on the veins; petioles 2-3 cm. long, slender, with few long stellate hairs. Inflorescences axillary, the cymes of 2-3 cymules, the flowering peduncles 3-4 mm. long, the pedicels 2-3 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, about 15 mm. long and 2-3 mm. wide slightly before anthesis, the apical appendages short, about 1 mm. long, glabrescent; sepals oblong, about 17 mm. long, densely pubescent without with long stellate hairs, glabrescent within; petals obovatc, about 14 mm. long and 3 mm. wide, the claw about 2-3 mm, long, briefly ciliate; gonophore stout, about 1.0 mm. long and 1.5 mm. wide, the glands narrowly elliptical; urceolus deeply 5-lobcd, about 0.5 mm. high, densely ciliate; stamens about 20, the filaments smooth, glabrous; ovary ovoid, about 1 mm. long, the spinulcs numerous, recurved, the style about 10 mm. long, the stigma acute. Fruit lightly stellate-pubescent, the body about 5 mm. in diameter at maturity, 3-loculate, each cell 2-seeded, later 6-celled each with 1 seed by false septation, the spines num- erous, slender, each about 4 mm. long, lightly pubescent; seeds ovoid, about 2 mm. long and broad. Apparently a rare species of northeastern Brazil, usually growing along stream- banks in wooded areas at altitudes of about 1000 m.; flowering and fruiting from May to July. Brazil: minas gerais: Cobras, S/. Ililaire B. ig (P, FM photo, US); Distr. Diaman- tina, Chnstais, near Corriga Duas Pontes, alt. 1160 m., Mexia 5821 (NY, US), data incomplete: Allcmao & Sysnciros 1491 (R). The characters that help in the identification of this species are the long golden- brown silky stellate tomentum, which clothes the younger branches, the Inflores- cence axes, the outside of the sepals, the leaves and the petioles, the relatively large flowers, and the comparatively long but weak spines. It is quite likely that T. lougicoma is conspecific with the Old World T. velntina Vahl. I have not seen the type of the latter, but have seen a specimen from India in Hooker's Herbarium at Kew, labelled T. vchttina, which matches well T. longicoma, 24. Triumfetta chthuahuensts Standi, in Contr. U. S. Nat. Herb. 23:744. 1923, (T.: M. E, Jones 5, n. In U. S. Nat. Herb.). Slender spreading shrubs 1-2 m. high; older branches Hghtly pubescent to 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 359 '■\ Fig. 8. Triumfctta chihuahucftsss * I [Vol. 37 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN nearly glabratc, punctate with prominent white lenticcls, the younger stems and inflorescence axes densely pubescent with short stellate and simple hairs. Leaves broadly ovate, generally 3-lobed, 11—14 cm. long and 6-9 cm. wide, distinctly cordate, the middle lobes narrowly and gradually acuminate, the laterals acute to shortly acuminate, both surfaces nearly glabrous, only the primary veins lightly pubescent; petioles slender, 6-7 cm, long, briefly stellate-pubescent. Inflorescences axillary, the cymes of 2-3 cymules, usually opposite the bracts, the flowering peduncles about 5 mm. long, the pedicels 3-5 mm. long. Flowers hermaphrodite, showy, the buds broadly oblongoid, 24-26 mm. long and 3-4 mm. wide slightly before anthesis, the apical appendages about 1 mm. long, glabrcscent; sepals oblong, 2 5—3 mm. long including the apical appendages, scattered stellate-pubescent without, glabrescent within; petals broadly obovate, 20-22 mm. long and 4-5 mm. wide, the claw about 3—5 mm. long, densely but very briefly pubescent; gonophore large, about 2.5 mm. long and 1.5 mm. wide, the glands oblong, about 2 mm. long and 1 mm. wide; urceolus short, distinctly many-lobed, briefly ciliatc; stamens 20, the filaments smooth, lightly stellate-pubescent at the base; ovary orbicular, about 2 mm. long, the spinules numerous, recurved, the style about 22 mm. long, the stigma very briefly 3 -parted. Fruit orbicular, the body about 6 mm. in diameter at maturity, very lightly stellate-pubescent, 3-loculate, each cell 1 -seeded, the gonophore and glands prominent, the spines about 75—100, lightly pubescent with few scattered hairs; seeds ovoid, about 3.5 mm. long and 3.0 mm. wide. Known only from southwestern Mexico, where it grows along steep, moist, shady canyons in pine forests at altitudes of 1000-1500 m.; flowers and fruits in September and October. Mexico: chihuahua: Guayanopa Canyon, Sierra Madre Mts., alt. 5000 ft., M. £. Jofjcs s. w. (Mo, US); near Batopilas, alt. 5500-6500 ft., Goldman 211 (GH, US); Arroyo Hondo, Sierra Charuco, Gentry lySo (Mo, US), sinaloa: Ocurahui, Sierra Surotato, alt. 6000-7000 ft., Gentry 6240 (Mo), sonora: Sanguarlbo, Rio Mayo, alt, 5500 ft., Gentry 212 J (Mo, US). This species is rather closely related to T. columnar i$, but may be distinguished from it by the larger, usually 3-lobed leaves and also by the pubescent spines. 25. Triumfetta barbosa Ko Ko Lay, sp. nov. Frutex circiter 2—4 m. altus. Ramuli graciles longitudinaliter sulcati 4-angulati glabrescentes. Folia late ovata non-lobata 14—16 cm. longa 7—10 cm. lata base cuneata vel rotundata apice longe attenuata subduplicitcr serrata pilis sparse stellatis mox glabresccntia infra breviter stellato-pubescentia, in axillis nervorum dense longi-pubescentia. Inflorcscentiae axillares, cymis cymulis 2, pedunculis circiter 4 mm. longis, pedicellis 8—10 mm. longis. Flores hcrmaphroditi, alabastro maturo oblongoideo 1 8—20 mm. longo; sepalis ellipticis 20—22 mm. longis extra breviter pubesccntibus, appendiculis circiter 2 mm. longis glabris; petalis ellipticis vel obovatis 14-16 mm. longis, unguilo 2—3 mm. longo dense plumoso; gonophoro circiter 1.5 mm. longo, glandulis ellipticis 0.75—1.0 mm. longis; urccolo circiter 0.75 mm, alto, 5-lobato breviter ciliato; staminibus 20, filamcntis usque ad 12-14 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 361 Fig. 9, Triumfetta harhosa mm. longis glabris; ovario globoso circlter 1.5 mm. longo, splnulis 75-100 rc- curvatis, stylo glabro 15—16 mm. longo, stigmate acuto. Fructus globosus non dehiscens, corporc ipso 4-6 mm. diametro, breviter stellato-pubescens 3-locularis, loculis 1-spermis, aculeis circitcr 75, 3—4 mm. longis breviter pubescentibus; seminibus ovoideo-acutis circiter 2 mm, longis et latis. [Vol. 37 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN West-central Mexico, Mexico: jalisco: Sierra Madre Occidental, Real Alto, trail to Arroyo del Jaguay, alt. 2500 m., Mcxia 1753 (holotype in Herb. Mo. Bot. Gard.; isotypes in Herb. Univ. Mich, and U. S. Nat, Herb.). A species rather closely allied to T. cbihuaJjjiensis^ but may be distinguished by its undivided long-attenuate leaves with dense tufts of hairs In the axils of the veins on the lower surface, and also by its more densely pubescent spines and glabrous filaments. 26. Triumfetta columnaris Hochr. in Ann. Conscrv. Jard. Bot. Geneve 18:111. 1914. (T.: Hartweg448). Triumfetta pscudocolumnaris Hochr. loc. cit. 103. 1914, (T.: Galcofti 41 JT), Triumfetta coUimnarioides Bullock, in Kew Bull. 292. 1937, (T.: Wnton 4917). Shrubs about 2 m, high; older branches glabrescent, longitudinally ridged, the younger ones and inflorescence axes scurfy, with long spreading stellate and simple hairs. Leaves broadly ovate to obscurely 3-lobed, usually small, 6-8 cm. long and 3-4 cm. wide, rounded or subcordate at the base, gradually and narrowly long- acuminate, both surfaces lightly stellate-pubescent, slightly more so on the veins; petioles relatively stout, about 2—3 mm. long, coarsely long stellate-pubescent. Inflorescences axillary, the cymes of 2 cymulcs, usually opposite the bracts, the flowering peduncles 4—6 mm. long, the pedicels 3-5 mm. long. Flowers hermaph- rodite, the buds broadly oblongoid, 22-25 mm. long and about 3 mm. wide slightly before anthesis, the apical appendages slender, 1—2 mm. long, glabrescent; sepals oblong, about 25 mm. long including the apical appendages, glabrescent and brownish without, light yellow within; petals narrowly obovate, about 18 mm, long and 3—4 mm. wide, the claw 4-5 mm. wide, densely but briefly ciliate; gono- phore large, about 2 mm. long and 1 mm. wide, the glands oblong; urceolus about 0,5 mm. high, 5-lobed, briefly ciliate; stamens 20, the filaments smooth, long stellate-pubescent at the base; ovary orbicular, about 2 mm, long, the spines num- erous, recurved, the stigma very briefly 3 -parted. Fruit orbicular, the body about 6 mm. in diameter at maturity, lightly stellate-pubescent, 3-loculate, each 1- seeded, the spines about 75—100, each 1-2 mm. long, lightly pubescent to nearly glabratc, the gonophore and glands prominent; seeds ovoid, about 2.5 mm. long and 1.5—2.0 mm. wide. Central and southwestern Mexico, usually growing in oak forests, at altitudes of 1500—2500 m.; flowering and fruiting from October to December. Mexico: guerrero: Mina, Fresnos, alt. 2160 m., Hififon 9749 (GH, US); Rio Frio Diamentes, Mina, Hinton 107J4 (GH, Mo, US). Mexico: Comunidad, Temascaltcpec, alt. 2350 m., Hinton 4S91 (GH, K, Mo, US), alt. 2300 m., 4917 (Mo, US), michoacan: Sierra TcrricHIia, Coalcoman, alt. 2420 m., Hhtton 12^49, ^5^44, ^5^70 (GH). morelos: Cuernavaca, Holway 532 (NY), oaxaca: Cordillera, alt. 5000-6000 ped., Galeofti 4131 (BR, FM photo); Tokontcpcque (Tehuantcpec?) , Harturg 448 (NY, FM photo); "les montagnes ombragier en tcrre tempiric,** Ghiesbreght 17S (F)* The small leaves and large flowers tend to confuse this species with T. Gold- mani, when examined superficially. 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 363 27, Triumfetta grandiflora Vahl, Eclog. Amer. 2:34. 1796. (T.: Ryan s.n. in Herb. Copenhagen). Trhunfctta ncmoralis St. Hll. Fl. Bras. Merld. 1:221. 1825. (T.: S/. Hilairc B. /. loyo in Herb, Paris.). Triumfetta longhcta DC. Pi. Rar. Jard. Geneve, 64. 1829. (T.: Burat s.n. "Havanne" in Herb. Deless.). Triumfetta longicuspis Turcz. in Bull. Soc. Nat. Moscou 31^:229. 1858. (T.: Jorgensen 882). Triumfetta cymosa Tr. & Planch, in Ann. Sci. Nat. Bot. IV, 17:3 53. 1862. (T.: "Quindio," Triana s,n. in Herb. Paris.). Triumfetta cymosa var. glabrescef7s Tr. & Planch. loc. cit. 3 54. 1862. Triumfetta cymosa var. hirsuta Tr. & Planch, loc. clt. 3 54. 1862. (T.: 'Tampalona," Triana s. w. in Herb. Paris.). Triumfetta grandiflora var. brasiliensis Hachr. in Ann. Conserv. Jard. Bot. Geneve 18:100. 1914. (T.: Glazioup3s6). Triumfetta boyacana Moldenke, in Phytologia 1:10. 193 3. (T. : Lawrance /). Triumfetta Schunkei Macbr. in Candollea 5:3 81. 1934. (T.: Schunke /jp/). Shrubs 1—2 m. high; older branches glabrous, smooth, punctate v/ith small white lenticelsj the younger stems and inflorescence axes rough, generally clothed with many long stellate and simple hairs. Leaves broadly ovate, usually about 14 cm. long and 9 cm. wide, sometimes slightly larger, rounded or cuneatc at the base, narrowly and gradually long-acuminate, both surfaces glabrescent or lightly scabrous with short stellate hairs, especially on the nerves; petioles slender, about 6-10 cm. long, usually lightly stellate-pubescent. Inflorescences terminal, the cymes of 1—2 cymules, usually opposite the bracts, the flowering peduncles about 6 mm. long, the pedicels 3—5 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, 14-18 mm. long and 3-4 mm. wide slightly before anthesis, the apical appendages slender, 1-2 mm. long, glabrescent; sepals linear-ovate, 15-20 mm. long including the apical appendages, glabrescent or lightly ferruginous-pubescent without; petals lanceolate to narrowly obovate, subcqualling the sepals, the claw 3-5 mm. long, densely plumose; gonophore about 1.5 mm. long and 2.0 mm. wide, the glands spherical; urceolus about 0.5 mm. high, many-lobed, briefly ciliate; stamens 20, the filaments glabrous, smooth; ovary orbicular, about 2 mm. long, the spinules numerous, recurved, the style 14-16 mm. long, the stigma very briefly 3 -parted. Fruit lightly pubescent, the body 5-6 mm. in diameter at maturity, rather easily loculicidally dehiscent, 3-loculate, each cell 2-scedcd, the spines num- erous, about 200, slender, 4-5 mm. long, lightly pubescent at the base, sometimes with few plumose hairs; seeds pyriform, glabrous, about 2-3 mm. long and 1.5-2.5 mm. wi ide. Rather widely distributed throughout the American tropics, though not abundant at any single locality, usually growing at altitudes of about 1000 m,; flowering and fruiting In its northern range from September to December, and in its southern range from March to June. Mexico: oaxaca: Sierra Snn Pedro Nolasco, Talea, Jorgenseyt 882, g6o (G) ; Chinantia, Galeotti 4162 A (BR, NY, US), vera cruz: Sierra Madre Misantia, Vtirpus 600Q (UC). Guatemala: alta verapaz: Coban, alt. 13 50 m., Tuerckbeim II 861 (BR, GH, Mo), alt. 43 00 pp., Tuerckbeim 342 GH). [Vol. 37 364 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panama: chiriqui: vicinity of El Valle de Anton, Allc?? 20^2 (Mo), Cuba: Havana: pros de la Havana, M. Sa^itos-Burat s. n, (P, Mo photo). Martinique: Sicber /p (Mo); Duss 2jlS (Mo, NY). Colombia: boyaca: region of Mt. Chapon, northwest of Bogota, Laurance J (NY, US) . santander: Eastern Cordillera, between El Roble and Tona, Ki\h\> ^ Sniifb 19399 (US). Brazil: minas gerais: Ponte dos Paulistas, Comarca do Cerro do Frio, S/. Ilihurc B. L lOJO (P) ; Bclo Horizontc, Barrcto 7923 (R). rig de Janeiro: Scrra de Estrclla pres de Botequin, Glaziou 9356 (BR, G); environs de Rio de Janeiro, Glaziou 174/0 (G). DATA incomplete: Ryan s.n, (C, FM photo). Ecuador: pichincha: Pucnte Gloria dc Maria, west of Quito, alt. 1200 m., AspIufiJ 7250 (US). Peru: junin: Chanchamayo Valley, alt. 1500 m., Schnnkc 1597 (FM). puno: Sandia, alt. 2100-2300 m., Wcbcrbaucr 507, 508 (G). data incomplete: ?()cp[ng 2053 (FM photo). Bolivia: cochabamba: Incachaca-S, Antonio, alt. 1500 m., WcrJermann 2127 (Mo). LA PAZ: Prov. South Yungas, Bnssin Rio Bopi, San Bartolome, near CaHsaya, alt. 750-900 m., Krukoff 10246 (Mo); Mapiri, Baf7g 1499 (Mo, US); Achiquin, between Mapiri and Apolo, air. 750 m., Cardenas IIIO (NY); San Carlos, Mapiri, Buchficn 1823 (US). TiPUANi: Hacienda Casana, Buchticn 73^^ (US). The best single character for the identification of this species is its fruits, which are usually rather easily dehiscent at maturity and have numerous, very long slender spines. In flower, it may be recognized by its broadly oblongoid floral buds, which are about 14-18 mm. long and 3 mm, wide at about anthcsis. The leaves are broadly ovate and the base is never cordate. There is considerable vari- ation in the pubescence of the leaves; in general, though not always, plants from South America tend to be more pubescent than those from North America. 28. Triumfetta discolor Rose, in Contr. U. S. Nat. Herb. 12:285. 1909. (T.: Rose 1979). Triumfetta dehhceus Rose, loc. cit. 285. 1909. (T.: Rose 1698). Low, spreading, bushy shrubs about 8 dm. high; branches and inflorescence axes rough, scurfy, with many long, spreading, stellate hairs. Leaves broadly ovate to rhombic-ovate, about 7—9 cm. long and 6—8 cm. wide, the base rounded, rarely subcordate, the tip abruptly acute, the serrations unequal, blunt and irreg- ular, the upper surface greenish, with few scattered, short, coarse, stellate hairs, the lower pallid, smooth, with dense stellate tomcntum; petioles 3—5 cm. long, clothed with long, spreading, stellate hairs. Inflorescences axillary, rarely terminal, the cymes of 2 cymules, rather lax, the flowering peduncles 3—4 mm. long, the pedicels about 2 mm. long, the subtending bracteoles prominent. Flowers hermaph- rodite, the buds oblongoid, about 15—20 mm. long slightly before anthesis, the apical appendages short, abovit 1 mm. long, lightly pubescent; sepals oblong, about 18 mm. long, brownish and lightly pubescent without, greenish and glabrcsccnt within; petals broadly obovate, about 15 mm. long and 3—4 mm. wide, deep yellow, the claw about 2-3 mm. long, slightly plumose; gonophore broad, about 1 mm. long and 2-3 mm. wide, the glands quadrangular, subcqualling the gono- 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 365 pliore; urceolus prominent, about 0.75 mm, high, many-lobcd, densely ciliate; m Stamens usually 20, rarely 25, the filaments glabrous; ovary orbicular, about 2 mm. long, the spinulcs numerous, recurved, the style 15-17 mm. long, the stigma acute. Fruit orbicular, the body about 6-7 mm. in diameter at maturity, slightly stellate- pubescent, loculicidally dehiscent, 3-loculate, each cell 2-seeded; spines about 75, about 2 m.m, long, lightly pubescent; seeds ovoid, about 1.5-2,0 mm. long and 1.0—1.5 mm. wide, Western Mexico, usually growing on rocky shaded slopes under oaks or in oak-savannahs at altitudes of 1000-1700 m.; flowering and fruiting from July to September. Mexico: durango: Huasemote, Rose & Hough 35^3 (GH, US); along trail from Pueblo Nuevo to Cucva, Sierra Madrc Occidental, alt. 1500-1700 m,, Harvey 18486 (US); exact locality lacking, Rose 22 jj (US), jalisco: in the foothills of Sierra Madre, near Colomos, Rose l6g8 (GH, US), nayarit: in the foothills of Sierra Madre, Terr, de Tepic, between Pedro Paulo and San Blascito, Rose IQJQ (GH). sinalqa: Cerro Colo- rado, alt. 2500 ft., Gentry 512J (GH, Mo); Las Mesas, Sierra Surotato, alt. 3000 ft., Gentry 6ljO (GH, Mo), 61 jl (Mo, US), Sljla (Mo); Qucbrado de Mansana, Sierra Surotato, alt. 4000-4500 ft., Gentry 6526a (GH, Mo); Las Mesas, Sierra Surotato, alt. 3000 ft.. Gentry 66§2 (GH, Mo), sonora: Sierra de Alamos, vicinity of Alamos, Rosey Stamiky & Russell 12881 (NY, US). The ligbtly stellate, easily dehiscent fruits, the large showy flowers, and the rhombic-ovate leaves with dense tomentum on lower surfaces, characterize this species. 29. Trtumfetta abutiloides St. HiL Fl. Bras. Merid. 1:223. 1825. (T.: St. Hilaire s, n., ''Itajuru de San Miguel de Mato Dentro" in Herb, Paris.), Sparsely branched, more or less trailing shrubs 1—3 m. high; older branches terete, scurfy, rather densely pubescent with long stellate and simple, frequently gland-tipped hairs, the younger ones and inflorescence axes very densely tomentose with long, spreading, stellate and simple hairs. Leaves broadly ovate, markedly 3-lobate, the lobes usually above the middle of the lamina, 12—14 cm. long and 8-10 cm. wide, the base subcordate, the terminal lobe long-acuminate, the laterals usually blunt, very rarely short-acuminate, the serrations very irregular, the basal ones always glandular, both surfaces densely pubescent with both stellate and simple hairs, the veins densely tomentose, the lower ones very prominent; petioles 3—4 mm. long, stout, densely long-tomcntose, usually flattened and forming a callus at the sinus. Inflorescences on terminal branches, the cymes of 3 cymules, always in the axils of the bracts, rather lax, the bracts elliptical, the flowering peduncles about 3 mm, long, the pedicels 3-4 mm, long. Flowers hermaphrodite, the buds broadly oblongoid, about 8—11 mm. long slightly before anthesis, the apical appendages about 1 mm. long; sepals oblong, 8-11 mm. long, grayish and rather densely tomentose without, yellow-brown and glabrescent within; petals subequalling the sepals, broadly obovate, showy, 7—10 mm. long and about 3 mm. wide, the claw 2—3 mm. long, densely plumose; gonophore slender, about 0.5 mm. long, the glands spherical to oblong, subequalling the gonophore; urceolus short. [Vol. 37 366 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 0.5 mm. High, distinctly 5-lobcd, densely ciliate; stamens 20, the filaments retrorsely 4-serratc; ovary orbicular, about 1.5 mm. high, the spinules numerous, recurved, the style 5-6 mm. long, the stigma briefly 3 -parted. Fruit spheroid, the body rather densely pubescent, 3—5 mm. In diameter at maturity, 3-loculate, each cell with 2 seeds, on maturity with 6 definite 1-seeded locules, the spines 100 or more, each about 3 mm. long, retrorsely pilosulosc; seeds lenticular, about 2 mm. long and wide, the funicular groove prominent. A rather common widespread species of South America, with a few scattered representatives in the West Indies; commonly found in abandoned fields or as a roadside weed at altitudes from about 300 to 1500 m.; flowers and fruits from January to April, Haiti: Massif de la Sella, Port-au-Prince, alt. 300 m., Ekman f2l^ (S), 740J (S), Colombia: cauca: Las Suntas, alt. 300-500 m., Lebmann 2J4 (G). magdalena: Santa Marta, H. H. Smifb l68g (NY, US). Venezuela: distrito federal; in valley, on road from EI Vallc to Cua, VUiicr I1974 (G); between Cotiza and Los Venados, near Caracas, Allart 21 (US); Caracas and vicinity, Bailey Gf Bailey 2$y (US). Brazhl: minas gerais: Caldas, Regnell L 102 (S, US); Dist. Carangola, Fazenda da Gramma, about 6 km. north, Mexia 43^4 (Mo, US); Caldas, in campo, Lindberg 284^ (BR, S); Itajuru de San Miguel de Mato Dentro, S/. Hilaire s.n. (P); Passa Quatro, Sampaio 603J, 6118 (R); Domlngos Pisoni, Municlpio do Leopoldini, Barreto 7928 (R); Jard. Bot. dc Bclo Horizonte, Barreto 7Q22 (R); exact locality lacking, Clausscn 482 (BR). :iio DE JANEIRO: Nova Friburgo, alt. 900 m., Holivay ^ Holway J 459 (US). RIO GRANDE DO SUL: Tristcza, Porto Alegro, Malme 1494 (S) ; Canoas, Prov. Porto Alegro, Malme s.n, (S) ; Cascate, Prov. Gloria, Malme 1353 (S)- sao paulo: Carapicho, Moscn II 32 (S). SANTA catharina: Vie 81 J (US), data incomplete: Burcbell 2491 (P). Ecuador: guayas: Milagro, alt. 500 m., Hitchcock 19960^ 20240 (US), manabi: Portoviejo, Rose ^ Rose 2^409 (US). Peru: cuzco: Prov. Quispicanchis, Tio, Marcapata, alt, 2000 m., Vargas C. 3^79 (Mo); Torontoy, Urubamba Valley, alt. 2400 m,, Cook G? Gilbert 819 (US); Santa Ana, alt. 900 m., Cook ^ Gilbert 1437 (US), huanuco: Huanuco, Macbride 3521 (US); Macbride ^ Teathersto>7e 2055 (US), loreto: Mishuyacu, near Iquitos, alt. 1000 m., Killip 6 Smith 29894 (US), data incomplete: Grisar s. n, (P). Bolivia: la paz: Mapiri, alt. 2500 ft., Rtisby 1450 (US); Hacienda Simao sobrc el camino a Tipuani, alt. 1400 m., Buchtien 545^ (Mo, S, US), 5453 > 5454 (US) ; Caranaim, Tipuani, Buchtien 73^7 (S, US); Nord Yungas, Milluguaya, alt. 1300 m., Btichtien 7^^ (FM, Mo, S, US); Sirupaya bei Yanacachi, alt. 2100 m., Buchtien 509 (US), 510 (S); Villa Aspiazu, Holway & Holway 688 (US). Paraguay; \J*Ksur\c\on^ Bal ansa 2327 (P). Argentina: Formosa: Riocho dc Oro, Reales s. w. (Mo), misiones: Candclaria, Santa Ana, Schwindt 49 (Mo); Posadas, Ekman 199 (S, US), tucuman: Tafi, alt. 600 m.,Yentiiri 1767 (US). This species is very closely related to T. bogotensis^ with which it is often con- fused. It may, however, conveniently be distinguished from the latter by the very densely pubescent leaves, the short apical appendages of the sepals, and also by the presence of false septa in the fruit, which on maturity becomes completely 6- loculate, each with one seed. 30. Triumfetta bogotensis DC Prod. 1:506. 1824. (Based on T. pilosa HBK.). Triumfetta pilosa HBK, Nov. Gen. & Sp. 5:343. 1821, non Roth (T.: Bonpland s.n. in Herb. Paris.). 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 367 fat Triumfctta clnmeforum Schlecht. In Linnaea 11:377. 1837. (T.: Keerl s, n. in Herb. Brux.). Trhiwfctta bispida A. Rich, in Sagra, Hist. Nat. Cuba 10:204. 1839. (T.: de la Sagra s, n, in Herb. Paris.)- Triumfetta grossitlariae folia A. Rich. loc. cit. 205. 1839. (T.: de la Sagra s. n. in Herb. Paris.) . Triumfetta Lindemana Turcz. in Bull. Soc. Nat. Moscou 31^:229. 1858. (T.r Linden jg). Triumfctta Bottcriana Turcz. loc. cit. 32^:260. 18 59. (T.r Bottcri 773), Triumfetta Orizaba Turcz. loc. cit. 261. 1859. (T.: Botteri 77 1), Triumfetta Josefina Polnk. in Linnaea 41:552. 1877. (T.: Polakoivsky 5^J). Triumfetta vincentina Urb. Symb. Ant. 5:414. 1908. (T.: H. H. 6 G. W. Smith 1057). Triumfetta valenciensis R. Kunth, In Fedde's Rep. Sp. Nov. Bcih. 43:464. 1927. (T.: Pitticr8677). Triumfetta panamensis L M. Johnston, in Sargcntia 8:194. 1949. (Trjobnston 1 1 12), Shrubs about 1-2 m. high; branches and inflorescence axes with many long, spreading hairs. Leaves broadly ovate, usually 3-lobcd, 9-10 cm, long and 7-8 cm. v/ide, the base rounded to subcordate, the tip acuminate, the serrations very irregular, unequal and blunt, both surfaces lightly pubescent with both stellate and simple hairs, the latter prominent, long, either spreading or appressed; petioles about 4-6 cm, long, much shorter in the bracts, very densely tomcntose with long simple and stellate hairs. Inflorescences axillary, the cymes of 2 or 3 cymules, the bracts broadly elliptical, the subtending bracteoles prominent, the flowering peduncles 4-7 mm. long, the pedicels 3-4 mm. long. Flowers hermaphrodite, the buds oblongold, 7-11 mm. long slightly before anthcsis, constricted about the middle, the apical appendages slender, 2-3 mm. long, lightly pubescent; sepals oblong, 8-13 mm. long including the appendages, long-hirsute without, glabrescent within; petals broadly obovate, about 8-9 mm. long and 3-4 mm. wide, the claw about 2 mm. long, densely plumose; gonophore slender, about 1 mm. long and 0.5 mm, wide, the glands elliptical, subequalling the gonophore; urceolus short, 0,25-0.50 mm. high, not lobed, distinctly ciliate; stamens 25-30, the filaments glabrous, without serrations; ovary suborblcular, about 1.5 mm. long, the spinules numerous, recurved, the style 8-9 mm. long, the stigma acute. Fruit spheroid, the body about 3-4 mm. in diameter at maturity, 3-loculate, each cell 2-seeded, tbe spines slender, about 75, retrorsely pilosulose, about 3 mm, long; seeds lentic- ular, 2-3 mm. long and about 2 mm. wide. A widespread weedy species of tropical America, which grows in waste lands, cut-over forest areas, and on hill-slopes, up to an altitude of about 2000 m.; flower- J J range The appressed simple hairs on both surfaces of the leaves, the broadly obovate petals, and the densely pilosulose spines distinguish this species from T. semitriloba. Mexico: guerrf.ro: Galcana, Carrizo to Pie de la Cucsta, Hinton 14735 (MICH). JALISCO: Atoyac, Kerber 135 (G). Mexico: Temascaltepec, Bejucos, alt. 610 m,, Hinton 2021 (K). michoacan: Cordillera, alt. 3000 ped., Galeotti 4156 (BR, P) ; exact locality [Vol. Z7 368 ANNALS OF THE MISSOURI BOTANICAL GARDEN lacking, GaJcofti 40^3 (BR, P). oaxaca: near Etia, Vriuglc 5718 (MICH); exact locality lacking, alt. 1750 m., Conzatti 6 Gonzalez 10^4 (GH) ; Licbmann 515 (US), tamaul- IPAS: Rio de la Ternera, Viirpus 8^77 (NY, UC, US) ; Tampico, Bcrlaudier JI (G, FM photo), VERA CRUZ: Jnlapa, f. W. Johnson s. n, (NY); Jalapa, Linden s,n. (K), 59 (G, FM photo), 771 (G, K, P, FM photo. Mo), 772 (P), 773 (K, P, FM photo) ; Orizaba and vicinity, Bofferi Io8g (US), Bourgeau 2846 (BR, G, P), Uiiller 1600 (BR); Zacua- pan, Caleotfi 7040 (BR), yucatan: Izamal, Cctnmer QOI, lOOj (BR, Mo), 1440 (Mo); Tcmax, Gaumer 21 JO (Mo); Calotmul, Gauwer 21 JI (BR); Chichankanab, Gaumer 20^5 (Mo); Xnocac, Gaumer 2J46g (Mo), data incomplete: Linden 7g (BR), 211 (BR); Hahn 65, 66 (P) ; Keerl s.n. (P). British Honduras: exact locality lacking, Feck 240 (K). Guatemala: alta verapaz: Coban, alt. 13 50 m., Tiierckbeim II 1957 (BR, Mo), alt. 4300 pp., Tiierckbeim IJ5g (Mo); Samac to Chama, alt. 2500 ft., H. Johnson 781 (US). DATA incomplete: Savage $. n, (P). Nicaragua: secovia: environ de Granadc, alt. 40 m., Levy j8g (G, P). Costa Rica: alajuela: collins de San Pedro en San Ramon, alt. 1050-1075 m., Brenes 601 (FM). cartago: northwest slope of Cerro Carpintcra, above La Union, Tres Rios, alt. 1320-1500 m., Dodge 6 Thomas 4S28 (Mo), guanacaste: defrichcmcnts sur les bords du Tillri, Tonduz 7208 (BR, GH). heredia: Vera Blanca de Sarapiqui, between Poas and Barba volcanoes, a!t. 1710 m., Skiitch 3374 (Mo), san jose: vicinity of El General, alt. 975 m., Skuich 2gi4 (GH, Mo); heights of La Carpintera, vicinity of Tres Rios, Allen 524 (GH, Mo); Hacienda San Francisco de Guadeloupe, alt. 1150 m., Pitfier 8472 (BR, GH, Mo), 7160 (BR); San Jose, P/7//Vr 42 (BR); Piedra Blanca, Tonduz 1267 (BR), data incomplete: Tonduz iog25 (BR, GH, K, NY); Endress 33 (K). Panama: canal zone: vicinity of Miraflores, P. c^ G. White 52 (GH, Mo) ; Gamboa, Sfandley 28511 (GH). chiriqui: vicinity of Bajo Mona and Qucbrada Chiquero, alt. 1500 m., Woodson ^ Schery 5g2 (Mo); Boquctc, alt. 4000 ft., Dai'idson 861 (Mo). PANAMA: Agricultural Experimental Station at Matias Hernandez, Pit Her 6g25 (GH, Mo) ; along the Corozal road, near Panama, Standley 2678g (GH) ; San Jose Island, about 55 miles s. se. of Balboa, Johnston III2 (GH). data incomplete: S. Hayes 155 (GH), 325 (NY). Cuba: las villas: in Magdalena, Cayamas, Baker 24go (UC). oriente: vicinity of Baracoa, Pollard, Palmer & Palmer 32 (Mo); exact locality lacking, Ekman Si (S). pinar del rio: Sierra de Anafe, Wilson & Leon Il2g8 (Mo), isle of pines: near Nucva Gcrona, Curtiss 234 (Mo). Virgin Islands: saint croix: Bassin yard, Ricksecker 80 (Mo), 136 (Mo, NY). Saint Vincent: Chateau Bclair, Beard l3go (Mo); H. H. ^ G. W. Smith 1057 (GH), ig43 (GH). Tobago: Friendship, R. O. Williams IIO41 (TRIN). Trinidad: Lavcntillc Hills near Port-of-Spain, Broadway 7^^7 (G, Mo) ; Chancellor Road, St. Ann, Broadivay 5242 (Mo), R. O. Williams II726, Tig32 (TRIN); St. Ann, " Broadway 4415A, 44T5B (TRIN); Hort. Trin., Back Lookout, Broadway 36S1 (TRIN). Colombia: antioquia: Hatillo, Bro. Daniel ^72 (US) ; Tablaza, Caldas, Gartner N, 8 (US); vicinity of Medellin, Robledo & Yepes 24 (US); Medellln, alt. 1500 m., Archer 730 (US), atlantico: Barranquilla and vicinity, Bro, Elias 6gg, 1 353 (US); Piojo, Los Mamcyales, alt. 350-400 m., Dugand 6 Garcia B, 2408 (US); Los Pendales, camino a Santacruz, alt. 5-10 m., Dugand & Jaramillo 2832 (US). Bogota: prope S. Fe de Bogota, alt. 1360 hex., Bonpland s.n. (P, FM photo, Mo); Gachata, alt. 1600 m., Triana s.n. (P). bolivar: Carthagcna, Blllberg 4g6 (S). boyaca: region of Mt. Chapon, extreme western part of Dept. Boyaca, Lawrance 157 (US), caldas: Chinchina, alt. 13 50-1400 m., Cuatrccasas 2310O (FM). cauca: Popayan, alt. 1700-2000 m., Lehmann 5522 (US), alt. 1740 m., Lehmann 3553 (G, US); entre Popayan y Cajeti (Hacia Tanibo) , alt. 1700 m., Cuatrccasas I383g (FM). cundinamarca: La Vega, alt. 13 00 m., Arbclaez ^ Cuatrecasas 5344 (US) ; Sasaima, Bro. Jose 25E (US), magdalena: vicinity of Cristobal Colon, Broadway 2g4 (US), norte de santander: region del Sararc, Hoya del Rio Chitaga, sobrc La Cubaya, alt. 1600-1800 m., Cuatrecasas, Scbultes & Smith 12143 (US); Ocana, alt. 3500 pp., Scblim 210 (BR), santander: northern slope of Mesa de Los Santos, alt. 1000-1500 m.^ Killip 6 Smith 1538S (US), tolima: 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 369 La Virginia Libano, alt. 1200-1500 m., Poincll 3282 (US), valle: none de Alban, alt. 2100 m., Dugand & jaravi'iUo 3058 (US); La Cumbre, Cordillera Occidental, alt. 1500- 1700 m., Venncll j2IJ (US); Cordillera Occidental, Vertiente Occidental, alt. 1770-2260 m., Cuafrccasas 18544, 18623 (FM), 227^2 (FM); Alto del Dinde, entre Cartago y Alcala, alt. 1200-1600 m., Cuairecasas 22gig (FM). data incomplete: Cnatrecasas 4682, 7794 (US); Bro. Hcribcrfo 479 (US). Venezuela: aragua: Maracay, Cor;/r/;o (5^ (US),Vo^/5/ (S), carabobo: vicinity of Valencia, alt. 400-800 m., Vittier 8675, 8677 (US), distrito federal: Las Adjuntas, Allart 506 (G, NY, US); lower Catuche, wood above Caracas, Niftier 7535 (US); Caracas and vicinity, Bailey ^ Bailey 286 (US), merida: 3 miles west and 2 miles south of M6r!da, Keed 30S (US); Tabay, alt. 1800-1900 m., Gebriger 447 (G, US); prope coloniam Tovar, Tcndler 123 (Mo.) miranda: between Las Canales and El Encanto, Lasser 671 (US), zamora: La Union, Edo. Bolivar, L. Williams I1226 (US). Brazil: bahia: Carpicha, Blancbet 7gg (G) ; Jacobina, Blanchet 3749 (BR, G, Mo). ceara: exact locality lacking, St. Hilaire Al. Ijd (P) ; Alle^nao 5. «. (R) ; Kocha 6 (US). PARANA: vicinity of Parana, Diisen, 22 (S). pernambuco: Tapera, ?ickel 2727, 2727a (US). RIO de janiero: Serra ^o Itatlaya, Nopasto, Sajnpaio 4143 (R). Rio grange do sul: Porto Alcgre, Reiueck & Czermak 329 (P). sao paulo: vicinity of Sao Paulo, Puiggari 3328 (P), Vidal s. n, (R). data incomplete: Regnell I 102b (S); Nazarctb, Jobert II 17 (P). Ecuador: cuayas: vicinity of Guayaquil, Mille 8 (NY), 198 (US), Rose ^ Rose 22453 (NY, US) ; vicinity of La Chonta, Rose, Pacharw & Rose 23474 (NY, US) ; 8 km. north of Guayaquil, Beetle 8941 , 8988 (US) ; Milagro, alt. 50 m., Hitchcock 20220 (US) ; vicinity of Las Juntas, Rose, Vacbano ^ Rose 23302 (US); vicinity of Huigra, Rose 6 Rose 22572 (US). Peru: cajamarca: San Miguel, Urubamba Valley, alt. 1800 m., Cook c^ Gilbert 89I (US), cuzco: Prov. Urubamba, Machupiccbu, alt. 2100 m., Vargas 795 (Mo, US). Bolivia: tarija: Carapari, Fries 1349 (S). Paraguay: Villa Rica, dans les forcts, Balansa 2325 (G, P) ; Caaguazu, Balansa 2326 (BR, G, P); Centralis, near Lake Ypacari, Hassler II703 (Mo, US). Argentina: jujuy: exact locality lacking, Fries 52 (S). misiones: San Ignacio, Quiroga 7^74^7(^75 (S). 31. Triumfetta obscura St. Hil. Fl. Bras, Merid. 1:222. 1825. (T.: St. Hilaire C2S2 in Herb. Paris.). Slender shrubs 1-2 m, high; older branches terete, glabrous, the bark with many shallow, longitudinal ridges, minutely lenticellate, the younger stems and inflorescence axes scurfy, slender with few short, coarse, stellate hairs. Leaves elliptical, more than twice as long as broad, sometimes very obscurely 3-Iobed, 11-13 cm. long and 4-6 cm. wide, the base cuncate to rounded, never cordate, the tip gradually and narrowly long-acuminate, the serrations double and rather regu- lar, both surfaces very lightly pubescent with few short, stellate hairs, glabrescent; petioles 3—5 cm. long, slender, scurfy, with many short separate tufts of stellate hairs. Inflorescences axillary, rarely terminal, the cymes of 2-3 cymules, the sub- tending bracts of the same general shape as the vegetative leaves, the flov/ering peduncles about 3 mm. long, the pedicels 2-3 mm. long, the bracteolcs prominent. Flowers hermaphrodite, the buds narrowly oblongoid, 6-8 mm. long slightly before anthcsis, the apical appendages about 1 mm. long, lightly stellate-pubescent; sepals elliptical, 6-9 mm. long, greenish with many short separate tufts of stellate hairs without, brownish and glabrate within; petals linear to narrowly obovate, sub- equalUng the sepals, the claw 1-2 mm. long, lightly plumose; gonophore slender. [Vol. Z7 370 ANNALS OF THE MISSOURI BOTANICAL GARDEN about 0.5 mm. long, the glands spherical to slightly oblong, about half the length of the gonophorc; urceolus short, about 0.25 mm. high, distinctly 5-lobed, lightly ciHate; stamens 20, the filaments retrorscly 4- to 6-serrate; ovary ovoid, about 1 mm. long, the spinules numerous, recurved, the style 5-6 mm. long, the stigma acute. Fruit orbicular, the body 3-4 mm. in diameter at maturity, very lightly ■ pubescent, 3-loculate, each 2-seeded, the spines slender, 100 or more, about 3 mm. long, slightly rctrorsely pilosulose; seeds pyriform, small, about 1.5 mm. long and wide, the funicular groove prominent. Small trailing shrubs usually growing in forests at altitudes of 1000-1500 m., with scattered distribution; flowering and fruiting in its northern range from October to December and in its southern range from January to about the middle of April, Venezuela: miranda: San Antonla, near Las Mostazas, alt. 1200 m., Allart 266 (NY, US). MONAGAs: between La Sabana de Las Picdras and Ccrro Negro, northwest of Caripe, alt. 1200-1500 m., Stcyermark 6l822 (FM, Mo). Brazil: distrito federal: trail between Sylvcstrc and Paincira, L. B. Smith 22 jj (US). MiNAs gerais: Fazcnda da Cachoeira, Municipio de Tambos, Barreto 1620 (R). PARANA: Ponta Grossa, in sllva, Dushi 7S5^ (Mo, S). Rio de Janeiro: circa Sabastian- opol, S/. Uilaire C2. 82 (P) ; Sebastianopol, Martins 87 (BR), 88 (BR, Mo, NY, S) ; exact locality lacking, Luschnatb 5. w. (S) ; Vautljicr 4JT (G) ; WiJgren Q2 (S). Rio grande DO sul: Porto Alegre, Keineck 220 (S). sag paulo: opposite Santos, Litnlbcrg 6g2 (BR); Santos, Moscn 3264 (S); Santa Anna, Bracie SQOO (S). data incomplete: Ackcnnann s. ft. (BR); Bnrchell 2486 (BR); Fryer s. n. (S) ; RieJcl log (BR, S) ; Liischnath s.fi. (BR); Kaben 4^1 (BR). Argentina: misiones: Loreta, in silva, Ekman 200 (S), This species may be distinguished from T, semitriloba by its narrowly elliptic, nearly glabrate leaves and by the small apical appendages on the sepals. On a fuller knowledge, it is quite possible that T. obscura might be assigned a subordi- nate rank under T. seviitriloba. 32. Triumpetta acracantha Hochr. in Ann. Conserv, Jard. Bot. Geneve 18:111. 1914- {i:,: Langlasse 505). Small shrubs about 1.5 m. tall; branches and inflorescence axes smooth, brown- ish, glabresccnt. Leaves ovate-elliptic, slightly 3-Iobed, about 1 0-1 1 cm. long and 4-5 cm. wide, the base rounded or obtuse, the tip gradually and narrowly acuminate, rather equally and doubly serrate, both surfaces with very few small tufts of stellate hairs, nearly glabrate; petioles slender, about 4—6 cm, long, slightly pubescent. Inflorescences usually axillary, sometimes terminal, the cymes of 2 cymules rather lax, the bracts elliptical to narrowly ovate, the flowering peduncles 2—3 mm. long, the pedicels about 2 mm. long. Flowers hermaphrodite, the buds oblongoid, 7—8 mm. long slightly before anthesis, slightly constricted towards the base, the apical appendages slender, 2-3 mm. long, glabrescent ; sepals oblong, about 8—9 mm. long including the apical appendages, lightly stellate-pubescent without and glabrescent within; petals lanceolate, 3-5 mm. long, the claw about 2 mm. long, sparsely plumose; gonophorc stout, about 1 mm. long and 1.5 mm. thick, the glands spherical, about 0.75 mm. long; urceolus about 0.5 mm. high, 1950] KO KO LAY AMERICAN SPECIES OF TRIUMFETTA 371 indistinctly lobcd, lightly ciliatc; stamens 20, the filaments smooth, slightly pubescent towards the base; ovary ovoid, about 1 mm. long, the spinules numerous, recurved, the style slender, about 6 mm. long, the stigma briefly 3-Iobed. Fruit orbicular, the body about 3 mm. thick at maturity, lightly stellate-pubescent, 3- loculatc, each cell 2-seeded, the spines slender, about 75, retrorsely pilosulose. Small shrub of western Mexico growing at low elevations, usually between altitudes of 100 and 200 m.; flowering and fruiting from September to December. Mexico: colima: Manzanillo, E, Palmer go2 (NY), guerrero: Baqueta, alt. 200 m., Langlassc JOJ (G, GH, US); Atoyac, Galeana, alt. 160 m., Hintan logil (GH). sinaeoa: La Norla, foothills, ait. 800 ft., Ucxia JJJ (UC) ; Capule, alt. 200 m., Ortega 6083 (US); Ymala, vicinity of Culican, £. Palmer 1683 (US); vicinity of Culican, BranJegee s. n, (UC). The glabrcsccnt leaves and sepals, together with the regularity and the fineness of the serrations, clearly distinguish this species from T. scmitriloba. 33. Triumfetta althaeoides Lam. Encyc. Meth. 3:420. 1789. (T.: "Cayenne", Sfoupy s. n, in Herb. Paris.). Trmmfetta rnicrophylla Vahl, Eclog. Amer. 2:34. 1796. (T.: "Br. Guyana", von Rohr s. w. in Herb. Copenhagen). Trhimfctfa suriuamenfis Steud. Flora 26:755. 1843. (T.: Hostmann 6oj). Triumfetta dicbotoma Turcz. in Bull. Soc. Nat. Moscou 32^:261. 1859. (T.: Kegel Triumfetta semitriloba var. surinamcmh (Steud.) Hochr. in Ann. Conscrv. Jard, Bot. Geneve 18:95. 1914. Triumfetta semitriloba f. altbaeoides (Lam.) Uittien in Pulle, Fl. Surinam 3:56. 1932. Bushy, spreading shrubs about 2 m. high; older branches lightly ferruginous- pubescent, the younger ones and the inflorescence axes more densely so, slightly scurfy. Leaves broadly ovate, 12-14 cm. long and 11-13 cm. wide, generally not lobed, sometimes obscurely 3-lobcd, the base shallowly cordate, the tip gradually and narrowly acuminate, the serrations rather regular, generally glandular at the base, with 4 prominent glands on the petiole at the basal sinus, the upper surface lightly stellate-tomentose, the lower more densely and coarsely so; petioles 4-6 cm. long, stout, densely tomcntose. Inflorescences on terminal branches, the cymes of 3-4 cymulcs, usually in the axils of the bracts, condensed in dense nodose clusters, the flowering peduncles 3-5 mm. long, the pedicels short, usually 1-2 mm. long. Flowers hermaphrodite, the buds broadly oblongoid, about 6 mm. long at anthesis, the apical appendages about 1 mm, long, nearly obsolete; sepals broadly elliptical, 6-7 mm. long, densely ferruginous-tomentose without, glabrescent within; petals linear, 3-4 mm. long, the claw about 1-2 mm. long, hghtly plumose; gonophore stout, about 0.75 mm. long and 1.0-1.5 mm. wide, the glands spherical, small; urceolus short, less than 0.5 mm. high, 5-lobed, lightly ciHate; stamens 20, the filaments glabrous, retrorsely 6-scrrate; ovary orbicular, about 1.0 mm. long, the