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Flora of Panama. Part VII. Fascicle I Source: Annals of the Missouri Botanical Garden, Vol. 45, No. 1 (Feb.

, 1958), pp. 1-91 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2394626 Accessed: 06/08/2010 17:48
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FLORA

OF
Part VII.

PANAMA
Fascicle 1P

PASSIFLORACEAE
1. PASSIFLORA L.
PASSIFLORA L.

Sp. P1.955. 1753; Killip,in FieldMus. Publ.Bot. 19:1-613. 1938.

[Tourn.ex Rupp.] Adans.Fam. 2:408. 1763. Granadilla 97. 1787. Cieca Medik.Malvenfam. [Tourn.] Medik.loc. cit. 1787. Murucuja Gen.398. 1789. Juss. Tacsonia Neck. Elem.2:235. 1790. Erndelia 6:396. 1805. Paris Mus. inAnn. Juss. Distephana Psilanthus Juss. loc.cit. 1805.

Mag. 267. 1818,nonTorr.& Gray. Monthly Raf. in Amer. Baldwinia in Ann.Gen. Sci. Phys.2:138. 1819. Bory, Astephananthes Bory, loc. cit. 1819. Monactineirma Bory, loc. cit. 139. 1819. Anthactinia 5. 1822. DC. in Mem.Soc. Phys.Geneve1:43 Polyanthea 78. 1824. Disemma Labill.Sert.Austro-caled. 3:335. 1828. Salisb.ex DC. Prodr. Distephia

132. 1828. Consp. Rchb. Astrophea


Raf. loc. cit. 1836. Macrophora

Raf. Fl. Tellur.4:103. 1836. Blephistelma

Raf. loc. cit. 1836. Meioperis Raf. loc. cit. 1836. Tripsilina Raf. loc. cit. 1836. Xerogona Peremis Raf. loc. cit. 104. 1836. Raf. loc. cit. 1836. Pericodia Raf. loc. cit. 1836. Synactila Raf. loc. cit. 1836. Odostelma Steud.Nom. 1:521. 1840. Distephania 2:149. 1846. Monogr. Roem.Synops. Dysosmia Pentaria Roem.loc. cit. 131,187. 1846. Decaloba Roem.loc. cit. 131, 152. 1846. 236. 1847. Lem.Fl. desSerres Distemma

28:438. 1856. inLinnaea Karst. Poggendorffia Rathea Karst.Fl. Colomb.1:77. 1859.

Rech.Fl. Amer.Centr.18. 1863. Oerst. Ceratosepalum

erectherbs, rarely by tendrils, or woodylianas,usuallyclimbing Herbaceous simpleor to peltate, basifixed petiolate, or smalltrees. Leaves alternate, shrubs, with large irregular entireto deeplylobed,occasionally veryrarelycompound, withfew to several areas (ocellate) upon theblade; petiolefrequently glandular axillaryor solitary, simple, glands. Tendrilsvolubile, moreor less conspicuous
*

The NationalScienceFoundation. Assisted by a grantfrom (1)

IssuedMarch 20, 1958.

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produced fromthe inflorescence. Inflorescence axillary, usually 1-flowered, less frequently cymoseand few-flowered, the pedunclejointedand thereproducing usually3 inconspicuous and setaceousto large and petaloidinvolucrate bracts. Flowershermaphrodite, actinomorphic, frequently large and showy,perigynous. Hypanthium shallow to relatively deep, thesepals5, valvate, usually witha corniculate process toward the tip in the large-flowered species;petals5, rarely absent, inserted at the marginof the hypanthium, greento highlycolored;coronaof numerous petaloid filaments in 1 to several centripetally decreasing inserted series on themargin of thehypanthium at the base of the corolla. Pistiland stamens borneupon a moreor less elongatecommonandrogynophore; stamens 5, borne immediately beneaththe pistil,the anthers 2-celled,versatile; pistil 3- or very rarely 4-carpellate, theovary1-loculate with3 or veryrarely 4 parietal placentas bearing numerous ovules,the 3-4 stigmas usuallyquite sessile, rather elongate, radial. Fruitan inflated berry, frequently large,globose to fusiform, containing numerous small hard lenticular seedsimmersed in abundantmucilaginous pulp derived from thetesta. the Passifloraceae includea few smallgeneraof the Old Worldas Although well,thelargegenusPassiflora is confined to theAmericas in its indigenous distribution, although numerous species withshowy flowers and ediblefruits have been introduced into the tropics of the EasternHemisphere and are frequently cultivatedin greenhouses in the colderlatitudes. In America nativespecies are found from thesouth-central UnitedStatesto Argentina. Theirextremely complicated and frequently flowers largeand brightly colored have attracted severalbotanists to the specialstudyof the Passifloras, the most recent of whomis E. P. Killip. In his studyof Passiflora, Killip recognizes 353 whicharedistributed species amongst no lessthan22 subgenera, chiefly basedupon ratherintricate differences in flower structure. Twenty-six specieshave been in Panamato whichat leasta few additions discovered will be made undoubtedly in the future. The present accounthas been drawnverylargely fromthat of Killip'smonumental work,although the keysare original and the species concept has beenchanged in a fewminor instances.The authors of thisFloraare grateful to Mr. Killipforthegenerous cooperation whichhe has giventhem in thenaming of their study material overmanyyears. Early Spanishmissionaries to the New World saw in the radiantflowers of a symbol of theCrucifixion, and from thisinspiration theSpanish Passiflora popular namepasionaria and theEnglish"Passionflower" have had their origin. Less reverent Spanish namesof frequent application to various species are caizoncillo and ala de murcielago. The large pulpy berries the of variousspecies, particularly P. edulisand P. quadrangularis frequently cultivated areknownas granadillas and areeatenrawor moreoftencookedwithmilkto makea kindof dessert.
a. Flowerstypicallyrathersmall, mostlyless than 5 cm. in diameter, at the nodes (usuallysolitary in P. punctata); usuallypairedor several toward the tip; bracts small and setose,very sepals not corniculate 2 whenpresent. inconspicuous; petiolar glands usually

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1958] FLORA OF PANAMA

(Passifloraceae)

b. Flowers 2 upon a common elongate peduncle, the associated tendril terminal; plants glabrous and very glaucous; leaves small, suborbicular,entire or nearly so; petiole with 2 foveate glands at the juncture with the blade.1. P. TRYPHIOSTEMMATOIDES .-.-... bb. Flowers 1 or several upon paired (occasionally solitary) peduncles, the associatedtendrilbasal. c. Petioleseglandular. d. Flowers few or several upon each peduncle; plants densely pubescent. -2. P. SEXFLORA dd. Flowers solitary upon each peduncle; plants glabrous or very inconspicuously pubescent. e. Leaves basifixed;stipulesminuteand setose. f. Peduncles jointed at about the middle or somewhat below; flowers 3. P. BIFLORA . usually paired at the nodes. ff. Peduncles jointed far above the middle, almost directly beneaththe flower. g. Leaves about twice as long as broad or nearly so, very occasionbroadly 3-lobed with the central lobe stronger, ally nearlyentire; flowers . 4. P. LANCEARIA usually paired at the nodes gg. Leaves about as broad as long; flowersusually paired at the nodes. h. Leaves with 2 stronglyascending lateral lobes, rarely with a weak centrallobe; petals absent ........................... 5. P. APETALA hh. Leaves about equally 3-lobed; petals present. -.... 6. P. PANAMENSIS ggg. Leaves much broader than long, with 2 nearly transverse lateral lobes; flowers ............... 7. P. PUNCTATA usually solitaryat the nodes ee. Leaves eccentricallypeltate, entire or if 3-lobed the central lobe far stronger; stipules large and foliaceous, amplexicaul, ................................................ 8. P. HAHNIT ciliate-serrate,deciduous cc. Petiolesglandular. d. Leaves eccentrically peltate, deeply 3-lobed, the lateral lobes and as strongas the central to much stronger; nearly transverse P. CORIACEA -------------------------------------------9. stipules inconspicuousand subsetaceous. dd. Leaves basifixed. e. Flowers typically 2 or more upon a short common peduncle; leaves broadly 3-lobed, the lobes rounded, occasionally mucronulate,the central far stronger;petiolar glands inconspic10. P. HOLOSERICEA ...............1... ............ ........... uous, broadlypoculiform ee. Flowers solitary upon each peduncle; leaf -lobDes acute to elevated. acuminate; petiolarglands conspicuously f. Leaves entire or subentirewith the lateral lobes reduced to inconspicuousangles; petiolar glands very large and auricular, bornetowardthe base of the petiole .................................. 11. P. AURICULATA ff. Leaves definitely 3-lobed; petiolar glands rather inconspicuous, borne about midwayor higherupon the petiole. g. Petiolar glands borne about midway upon the petiole; gg. Petiolar glands borne almost directly beneath the leaf ......................... 13. P. blade; petals absent aa. Flowers typicallyratherlarge,mostlymore than 5 cm. in diameter, solitary at the nodes (except occasionallyin P. adenopoda); sepals usually conspicuouslycorniculatetoward the tip (except in P. pulchella and P. mnembranacea); bracts conspicuousand involucrate,foliaceousor petalaceous; petiolarglands usually more than 2 when present. b. Involucral bracts entire,serrulate, or simplylacerate. c. Sepals not corniculate;petiolesnot glandular. d. Leaves cuneately2-lobed, occasionallywith a small centrallobe; stipules inconspicuousand subsetaceous; involucral bracts obovate, obtuseor acute at the base, 1.0-1.5 cm. long; flowers about 4-6 cm. in diameter .. .................................................... 14. P. dd. Leaves suborbicular,indistinctly3-lobed; stipules conspicuously foliaceous; involucral bracts suborbicular,deeply cordate, about 3-5 cm. long; flowersabout 7-8 cm. in diameter ......................... 15. P. cc. Sepals corniculate; petiolesglandular.
petals present, linear......................... 12. P. PEDICULATA
SUBEROSA

PULCHELLA

MEMBRANACEA

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d. Leaves palmately 3- to 7-parted to below the middle, the central lobe somewhatcontracted toward the base. e. Involucral bracts much shorter than the sepals, free and never envelopingthe flowerbud, serrateor lacerate. f. Leaves 5- to 7-lobed (very rarely 3-lobed) petiolar glands very large and prominentlystipitate, borne almost directly beneath the leaf blade; involucral bracts lacerate; flowers 16. P. ADENOPODA ....................................................... purple and white ff. Leaves 3-lobed; petiolar glands inconspicuous and sessile, borne toward the base of the petiole; involucral bracts glandular-serrulate; flowersscarlet and yellow....................... 17. P. VITIFOLIA ee. Involucral bracts almost as long as the sepals, united toward the base and completely enveloping the flower bud, entire; flowersviolet and white....................................................... 18. P. WILLIAMSII dd. Leaves entire, or palmately 3-lobed above or about the middle and with the central lobe not constrictedtoward the base. e. Leaves 3-lobed. f. Plants more or less pilose; leaves lobed above the middle, the central lobe much stronger; petioles with 2 to several pairs of glands; involucral bracts rather narrowly lanceolate, foli19. P. MENISPERMIFOLIA aceous ....................................................... ff. Plants glabrous; leaves lobed to about the middle, the 3 lobes subequal; petioles with 1 pair of glands; involucral bracts ....................................... 20. P. SUBPELTATA broadly ovate, somewhat petalaceous ee. Leaves entire,deeply cordate in some species. f. Stipules very conspicuously foliaceous. g. Stipules equilateral, deciduous; involucral bracts about as long as the sepals. h. Stems not winged; leaves deeply cordate; petioles with 21. P. LIGULARIS filiformglands. ....................................................... hh. Stems prominentlywinged; leaves rounded at the base . 22. P. QUADRANGULARIS to obscurely cordate; petioles with ovoid glands .. gg. Stipules strongly inequilateral,persistent;involucral bracts about half as long as the sepals ............................................. 23 P. OERSTEDII ff. Stipules inconspicuous and subsetaceous or barely foliaceous. g. Leaves obtuse or very slightly cordate at the base; involucral bracts free. h. Leaves entire; petioles glandular at the middle or ....................... 24. P. AMBIGUA below; flowersmaroon, purple and white hh. Leaves serrateor serrulate; petioles glandular above the ..................................... 25. P. NITIDA middle; flowerswhite and purple gg. Leaves deeply cordate at the base; petioles glandular directly beneath the leaf blade; involucral bracts united at the base; flowerspurple and white ................................... 26. P. SEEMANNII bb. Involucral bracts complexly 2- or 3-pinnatifid,each filiformdivision gland-tipped; plants hirsutulose; petioles with scattered gland-tipped ............... 27 P. FOETIDA hairs; flowersgreenishwhite more or less tinged with purple

Other species, at present knownfromCosta Rica and Colombia,are to be in Panamaas well. expected
1. PASSIFLORA TRYPHOSTEMMATOIDES

Harms, in Engl. Bot. Jahrb.18: Beibl. 46:6. 1894; Killip,in FieldMus. Publ. Bot. 19:81. 1938.

Passiflora gracilimmaKillip, in Journ.Wash. Acad. Sci. 14:112. fig. 2b. 1924.

Plantsslender, climbing, glabrous and veryglaucous throughout. Leaveslongpetiolate, simple, entire, suborbicular, about1.5-4 cm. longand 1.5-3.0 cm. broad, darker very pale beneath, above;petioles veryslender, about 1-2 cm. long,bearing 2 rather inconspicuous foveate glandsat thejuncture withtheleafblade; stipules setose,veryinconspicuous.Inflorescence of 2 greenish whiteflowers upon an (4)

1 9 58 ]

FLORA

OF PANAMA

(Passifloraceae)

elongate common peduncle, the associated tendrilterminal; peduncle 0.5-4.5 cm. long; pedicels 1.5-4.0 cm. long, jointed somewhat above the middle, the bracts minutelysetose,deciduous. Flowers 1.5-4.0 cm. in diameter;sepals oblong, about 0.7-1.5 cm. long, not corniculate; petals narrowlyoblong, 0.5-1.0 cm. long; filamentsof the corona in a singleseries, filiform, 3-5 mm. long. Berrysubgloboseto ovoid, 3-6 cm. long, greenish yellow. Panama and Colombia, in wet forests of middleelevations.
uncertain,Hart I04.
BOCAS DEL TORO:

Fish Creek Hills, Von Wedel 242I,

38o.

CHIRIQUi:

exact locality

There seems to be little room for doubt that P. tryphostemmatoides and P. gracilimma are conspecific (cf. Killip, in Field Mus. Publ. Bot. 19:82. 1838), since they are not closely related to other species, their charactersoverlap or at least intergrade, and both have been reportedfrom closely adjacent stationsboth in Panama and Colombia. No intermediatestations between Bocas del Toro (probably Hart's "Chiriqui") and the Atrato valley in Colombia have been discovered as yet, but they are to be expected since the Caribbean coast of Panama still is known so poorly.
2.
PASSIFLORA SEXFLORA Juss. in Ann. Mus. Hist. Nat. 6:110. 1805; Killip, in Field Mus. Publ. Bot. 19:129. 1938.

pA.37. fig. 2.

Macf. Fl. Jam. 2:149. 1850. Passifloratriflora Passiflora miraflorensis Killip, in Journ.Wash. Acad. Sci. 14:109. 1924. isotrilobaCuf. Archivio Bot. 9:196. 1933. Passiflora

Lem.Fl. desSerres Passiflora floribunda 4:335b. 1848.

Decaloba sexflora(Juss.) M. Roem. loc. cit. 164. 1846.

Cieca pannosa(J. E. Sm.) M. Roem.Fam.Nat. Syn.2:148. 1846.

Passiflora pannosa J.E. Sm. in Rees, Cycl. 20: Passiflora no. 28. 1819. Passifloracapsularisvar. geminiflora DC. Prodr. 3:325. 1828 (as geminifolia). Meioperispannosa (J. E. Sm.) Raf. Fl. Tellur. 4:103. 1838.

in wet forests, 1300-2000


BOCAS CHIRIQUI: DEL TORO:

Plants ratherslender,climbing,denselypubescent throughout. Leaves petiolate, simple, basifixed,subtrapezoid,the top saliently and acutely 2-lobed, occasionallywith a smaller central lobe, the base rounded or slightlycordate, 3-9 cm. long, 4-10 cm. broad; petioles 1-3 cm. long, eglandular; stipules subulate, about 5 mm. long. Inflorescencebearing 2-10 greenish-white, purple-tingedflowers upon a common peduncle 2-3 cm. long, the associated tendrilbasal; pedicels 1.01.5 cm. long, subtendedby an inconspicuoussubulate bract. Flowers 2-3 cm. in diameter; sepals oblong-lanceolate,8-15 mm. long, not corniculate; petals linear, about 10 mm. long; filaments of the corona in 2 series,the outer about as long as the petals. Fruit subglobose,5-10 mm. in diameter,greenishyellow. SouthernFlorida; Mexico to northernColombia; Greater Antilles. Clearings
m. in Panama.

Bajo Mono, Boquete District, Davidson 49I; vicinity of Bajo Mona and Quebrada Chiquero, Woodson & Schery 5i9; valley of the upper Rio Chiriqui Viejo, vicinity of MonteLirio, Seibert aboveEl Boquete, I80; Rio Ladrillo, Pittier 3286.

Robalo Trail, northern slopes of Cerro Horqueta,Allen 4928.

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

tVot. 4S

Fig. 1.

Passiflora sexflora

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t958]

FLORA OF PANAMA 3.
PASSIFLORA BIFLORA

(Passifloraceae)

Lam. Encycl. 3:36. 1789; Killip, in Field Mus. Publ. Bot.

19:185. 1938. Passiflora lunataJ.E. Sm. Icon. P1. Rar. 11: pI. I. 1790, non Juss.nec Vell. necque Poepp. & Endl. Passiflora glabrata HBK. Nov. Gen.& Sp. 2:135. 1817. Cieca glabrata(HBK.) M. Roem.Fam.Nat. Syn.2:143. 1846. Decalobabiflora (Lam.) M. Roem.loc. cit. 2:161. 1846. var. major Decalobabiflora M. Roem.loc. cit. 1846. var. mexicana Decalobabiflora M. Roem.loc. cit. 1846. lunatavar. costata Passiflora Mast. in Mart. Fl. Bras. 131:552. 1872. Passiflora spathulata Mast.loc. cit. 1872. brighami S. Wats. in Proc.Amer. Acad. 21:473. 1887. Passiflora Mast. in Bot. Gaz. 16:7. 1891. transversa Passiflora Plants ratherslender,climbing,essentially glabrous throughout, the stems conspicuouslystriate. Leaves rathershortlypetiolate,simple, basifixed,subtrapezoid, the top salientlyand obtusely 2-lobed, occasionally with a smaller central lobe or mucro, the base obtuse to broadly rounded, 1-10 cm. long, 5-10 cm. broad, conspicuously glandular-ocellate; petioles 0.5-3.0 cm. long eglandular; stipules minutely setose. Inflorescences bearing solitarygreenishwhite or yellow flowers upon paired peduncles, the associated tendrilbasal; peduncles about 1.0-1.5 cm. long, jointed at about the middle or somewhatbelow; bracts very inconspicuous, subsetaceous. Flowers 3-4 cm. in diameter; sepals ovate-lanceolate,about 1 cm. long, not corniculate; petals oblong-lanceolate,slightly shorter than the sepals; filamentsof the corona in 2 series,the outer about as long as the petals. Fruit subglobose,1-2 cm. in diameter,greenishyellow. Mexico to Colombia and Venezuela; Bahamas; fromnear sea level to about 1,500 and common in Panama. m. Widely distributed
BOCAS DEL TORO: Shepherd Island,Von Wedel2694, 2737; WaterValley,Von Wedel Point,Von Wedel 2645; vicinity of Chiriqui Lagoon, Von Wedel384, 598, I697; Snapper CANAL ZONE: Allen & Seibert I218, I305. AnconHill, Woodson, 7I6; Chagres, Fendler I2I; Ancon, Pittier 2578, Standley I2I07; Culebra, Pittier 209I, Standley 25982; Empire Standley to Mandinga, Piper5520; Fort Sherman, 3I040, 3II84, StevensI046; Frijoles, Piper5806; Barro Colorado Island, Bailey & Bailey 3II, Wetmore & Abbe I40, Woodworth& Vestal307. CHIRIQUI: El Boquete,Killip 3608; David, Killip 3640. COCLE: Williams226. COLON: Porto Bello, Pittier 246I; Catival,Standley Penonome, 30349. PANAMA: PanamaVieja to Bella Vista,Allen 826; Las Sabanas, Standley 25835; Taboga

Island, Standley 27844, Pittier 3607; San Jose Island, Erlanson I83, 2I9, Johnston809, II89, 549; Tumba Muerto Road near Panama', Standley 29802; Trapiche Island, Miller Miller 20II. I893; Taboguilla Island,

Popular names of this weedy climber are guate-guate, camacarlata, and calzoncillo.

4.

PASSIFLORA LANCEARIA Mast. in Journ. Bot. 23:114. Mus. Publ. Bot. 19:160. 1938.

1885; Killip, in Field

talamancensis Passiflora Killip,in Journ. Wash.Acad. Sci. 12:260. 1922. Rather massive climbers,essentiallyglabrous throughout. Leaves ratherlongpetiolate, simple, basifixed,oblong-ellipticand entire to the tip or very broadly

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3-lobed withthe central lobe stronger thanthe lateral, obtuseto rounded at the rather base,5-12 cm. long,2-9 cm. broad, membranaceous to subcoriaceous, conspicuously glandular-ocellate; petioles 1-4 cm. long,eglandular; stipules subsetaceous, inconspicuous. Inflorescences bearing solitarygreenish-white, faintly the associated flowers tendril purple-tinged upon usuallypairedpeduncles, basal; farabovethemiddle almost peduncles about2-5 cm.long,jointed beneath directly the flower; bractsinconspicuously subsetaceous.Flowers3-4 cm. in diameter; sepalsovate-lanceolate, 1.0-1.5 cm. long,not corniculate, glabrous to moreor less as longas thesepals;filaments puberulent within; petalsabouthalfto one third of thecorona in 2 series, theouteraboutas longas thepetals. Fruitsubglobose, 2-3 cm. in diameter. from CostaRica and Panama, nearsea levelto about 1,700m. altitude.
BOCAS DEL TORO: Old BankIsland, Von Wedel2I20. District, Davidson 336.
CHIRIQUf:

Bajo Chorro, Boquete

theaffinities Thisis a representative of which of a very of species complex group HBK. are farfrom to P. mollis clear. Perhaps our plantsshould be referred
5.
PASSIFLORA APETALA

Killip, in Journ. Wash. Acad. Sci. 12:255.

1922.

Plants rather slender, climbing, glabrous throughout. Leaves rather longpetiolate,simple,basifixed, broadlycuneate-subtrapezoid, with 2 strongly ascending laterallobes,rarelywith a weak centrallobe, broadlyobtuse or roundedat the base, 3-10 cm. long and broad, inconspicuouslyglandular-ocellate; petioles 1-5 cm. long, eglandular; stipulesminute and subsetaceous. Inflorescences bearing solitary yellowish green flowersupon paired peduncles, the associated tendril basal; peduncles about 1.5-4.0 cm. long, jointed above the middle; bracts inconspicuousand subsetaceous. Flowers about 2 cm. in diameter;sepals narrowlyoblong-lanceolate, about half as long as the sepals or less. Fruit globose,about 1 cm. in diameter, dark purple. Costa Rica and Panama, in clearingsof forestsat 1,000-2,000 m.

about1 cm.long,notcorniculate; petalsabsent; filaments of thecorona in 1 series,

CmpRIQUI:trail fromBambitoto CerroPunta, Allen 3i6; Chiquero, Davidson5II; Bajo Mona,mouth of QuebradaChiquero, Allen d Seibert Woodson, ioi6; Bajo Mona and QuebradaChiquero, Woodson d Schery of "New Switzerland", central 583; vicinity valley of Rio ChiriquiViejo,Allen I346; valleyof upperRio ChiriquiViejo,vicinity of Monte Lirio, Seibert i6o, G. White 25; AltoLino,Bro.Maurice 835. 6.
PASSIFLORA PANAMENSIS

Killip, in Journ.Wash. Acad. Sci. 12:259. 1922.

Plants rather slender, climbing, glabrous throughout. Leaves long-petiolate, simple,basifixed, broadly 3-lobed, the lobes ascending,acute, and essentially equal, broadly rounded at the base, 5-12 cm. long and broad, conspicuouslyglandularocellate; petioles 1.5-3.0 cm. long, eglandular; stipules inconspicuous and subsetaceous. Inflorescences bearing solitary greenish lavender-tingedflowers upon paired peduncles, the associated tendril basal; peduncles 2-4 cm. long, jointed almostimmediately beneaththe flower;bracts minute,subsetaceous. Flowers about

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19bJb8

FLORA

OF PANAMA

(Passifloraceae)

3 cm. in diameter;sepals oblong-lanceolate,1.0-1.5 cm. long; petals oblanceolate, of the corona in 2 series,the outer about about half as long as the sepals; filaments 2 cm. in diameter,purple. about Fruit globose, as long as the petals. in lowland forests. Colombia, adjacent and Panama Eastern
DARIEN:

5556. Pittier River, alongSambui


PUNCTATA

7.

PASSIFLORA

19:183. 1938.

L. Sp. P1. 957. 1753; Killip, in Field Mus. Publ. Bot.

HBK. Nov. Gen.& Sp. 2:136. 1817. misera Passiflora in Ann.Gen.Sci. Phys.Brux.2:149. pI. 24. 1819. Bory, maximiliana Passiflora Ker,Bot.Reg. 7: pI. 597. 1821,nonL. vespertilio Passiflora Link& Otto,Icon.P1. Select.13: pI. 5. 1828. discolor Passiflora Hook. Bot.Misc.3:325. 1833. retusa Passiflora Decalobapunctata(L.) M. Roem.Fam. Nat. Syn.2:156. 1846. (Link& Otto) M. Roem.loc. cit. 140. 1846. Cieca discolor (HBK.) M. Roem.loc. cit. 1846. Cieca misera Mast.in Mart.Fl. Bras.131:556,593. 1872. microcarpa Passiflora WashAcad. Sci. 14:110. 1924. in Journ. Killip, laticaulis Passiflora in Ann.N. Y. Bot.Gard.7:309. 1927,in part. Rusby, translinearis Passiflora Annex.5: Bot. 1:71. pI. 63. Hoehne,in Linh. Telegr.Matto Grosso, longilobis Passiflora
1910.

Slenderclimbers,glabrousor essentiallyso. Leaves petiolate,simple,basifixed, lateral lobes and usually a slender much broaderthan long, with 2 nearlytransverse cm. long, 3-10 cm. broad, 0.5-3.0 lobe, central central mucro or rarelya small cm. long, eglandular; stipulesminute 1-6 petioles conspicuouslyglandular-ocellate; white, a singlegreenish and bearing solitary usually and subsetaceous. Inflorescence cm. 3-10 tendril basal; peduncle associated the flower, occasionallypurple-tinged and bracts inconspicuous the flower; beneath immediately long, jointed almost cm. 1-2 sepals ovate-lanceolate, in cm. diameter; 2-4 about setaceous. Flowers of the corona in 2 series, long; petals narrowlyoblong, 8-12 mm. long; filaments cm. in diameter, 1.0-1.5 Fruit globose, the petals. equaling the outer almost purple. Panama to Argentina,at low elevationsin thicketsand forestclearings. F Vestal457, 7I7, Bailey& Bailey BarroColoradoIsland,Woodworth CANAL ZONE: 574, HunnewellI9029; MonteLirio, & Abbe 26, Bangham 456, Wetmore 258, Kenoyer 3998. Pittier Maxon684I; between Frijoles and Monte Lirio,KillipI2I87; Trinidad River, 687. Stevens Mindi,Cowell I69; Rio Mindi,SuttonHayes 596; Baillemona, PANAMA: It is questionablewhetherP. misera can be maintainedas a speciesdistinctfrom P. punctata since their general aspect, variational tendency,and geographicaldistributionhave so very much in common. They were kept separate by Killip, are linear,broadly however,who was of the opinion that the innercorona filaments or capillary,rarely dilated at the apex and often lobulate in P. miseraand filiform minutelycapitellatein P. punctata.
8. PASSIFLORA HAHNII

(Fourn.) Mast. in Mart. Fl. Bras. 131:569. 1872.

HahniiFourn.in Rev. Hort.41: pI. 430. 1869. Disemma S. Wats.in Proc.Amer.Acad. 22:473. 1887. guatemalensis Passiflora (9)

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ANNALS OF THE MISSOURI BOTANICAL GARDEN

[Vol.

45

Ratherslender climbers, glabrous throughout.Leaves petiolate, simple, very eccentrically peltate, broadly ovate,essentially entire or if 3-lobed thecentral lobe farstronger and the lateralrelatively inconspicuous, the base truncate or broadly rounded, 5-8 cm. longand broad;petioles 2-3 cm. long,eglandular; stipules large and foliaceous, inequilaterally reniform, amplexicaul, ciliate-serrate, about 1 cm. long and 2 cm. broad,deciduous. Inflorescences bearing solitary yellowish-white flowers upon pairedor solitary the associated peduncles, tendril basal; peduncles 1.5-2.0 cm. long,jointedalmostimmediately beneath the flower; bractsminute andsubulate. Flowers about4-6 cm.in diameter; sepals oblong, 2-3 cm. long,not corniculate; petals similar to thesepals;filaments of thecorona in 2 series, theouter about 1.5 cm. long. Fruitglobose, 3-4 cm. in diameter, deeppurple. Southern at low to middle Mexicoto Colombia, elevations.
COCLE:

mountains beyond La Pintada, 400-600 m.,Hunterd Allen629.


Juss. in Ann. Mus. Hist. Nat. 6:109.

9.

Killip, in FieldMus. Publ.Bot. 19:83. 1938.

PASSIFLORACORIACEA

39, fig.2. 1805; pA.

difformis HBK. Nov. Gen.& Sp. 2:136. 1817. Passiflora J.E. Sm. in Rees,Cycl. 26: Passiflora no. 20. 1819. Passiflora clypeata Monactineirma coriacea(Juss.)Bory, in Ann.Sci. Gen. Phys.Brux.2:138. 1819. Cieca difformis (HBK.) M. Roem.Fam.Nat. Syn.2:140. 1846. Cieca coriacea(Juss.)M. Roem.loc. cit. 148. 1846. sexocellata Schlecht. in Linnaea27:521. 1854. Passiflora Passiflora obtusifolia Sesse& Moc. pl. Nov. Hisp. 156. 1887.

Rather extensive climbers inconspicuously puberulent to glabrate. Leavespetiolate,simple, veryeccentrically peltate,deeply3-lobed,the laterallobes rather abruptly acuminate and nearly transverse, as strong as the central lobe to much stronger, 2 ocellate 3-7 cm. long,6-25 cm. broad;petioles 2-4 cm. long,bearing glands nearthejuncture withtheleafblade;stipules narrowly linear, about0.5 cm. 2- to several-flowered with slightly long. Inflorescence folia terminal raceme, aceous,frequently 3-lobedbractssubtended by pairedocellateglands; pedicels about5 mm.long. Flowers or yellowish 2.5-3.5 cm. in diameter, greenish white; 1.0-1.5 cm. long,not corniculate; sepalsoblong-lanceolate, petalsnone; filaments theoutermost 1-2 cm. in of thecoronain 2 series, 7-8 mm.long. Fruitglobose, diameter, deepblue. Mexicoto British Guiana and Bolivia,fromnearsea level to about 2,000 m.
CANAL ZONE: Miraflores CocoliIsland, Lake,P. White86, G. White133; Las Cascadas, to Mandinga, Stawdley Standley 29594; Darien Station, 3I617; Empire Piper5479; Barro & Vestal 5oi. El Valle de Anton, ColoradoIsland,Shattuck57, Woodworth COCLE: AllenI667, 4473. PANAMA: Alhajuela, Pittier 3456.

In somepartsof Central America thisspecies is knownas inurcielagoand ala de murcielago withreference to thepeculiar shapeof theleaves.

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1958]

FLORA

OF PANAMA

(Passifloraceae)

11

10.

PASSIFLORA HOLOSERICEA

19:99. 1938.

L. Sp. P1. 958. 1753; Killip, in Field Mus. Publ. Bot.

Decalobaholosericea (L.) M. Roem.Fam. Nat. Syn.2:164. 1846. Rather coarse climbers, densely puberulent throughout. Leaves petiolate, simple,basifixed, broadly 3-lobed, the lobes rounded and occasionallymucronulate, the central lobe far stronger,the base rounded to somewhat cordate, 5-10 cm. long, 4-8 cm. broad; petioles 1-3 cm. long, the glands ratherinconspicuousand broadly poculiform; stipules inconspicuous and filiform. Inflorescences usually paired,corymbiform, typicallybearing 2 or more greenishwhite or yellow flowers upon a veryshortcommonpeduncle,the associatedtendrilbasal; commonpeduncles 2-4 mm. long; pedicels about 1 cm. long; bracts inconspicuous and filiform. Flowers about 2-4 cm. in diameter,sepals oblong-elliptic,about 1.5 cm. long; petals broadly elliptic, about 1 cm. long; filaments of the corona in 2 series,the outer nearlyas long as the petals, flushedwith yellow and purple. Fruit globose, about 1.5 cm. in diameter. SouthernMexico to Colombia and Venezuela; Cuba. Lowland thicketsto about 700 m. 8i6. CHIRIQUI: vicinity of Puerto & Schery Armuelles, Woodson
11. PASSIFLORA AURICULATA HBK. Nov. Gen. & Sp. 2:131. 1817; Killip, in Field Mus. Publ. Bot. 19:122. 1938.

Passiflora appendiculata G. F. W. Mey.Prim.Fl. Esseq.223. 1818. Desv. in Ham. Prodr. 48. 1825. Passiflora cyathophora Passiflora Rohrii DC. Prodr.3:326. 1828. cinerea Poepp.& Endl.Nov. Gen.& Sp. 2:57. pI. I77. 1838. Passiflora Cieca auriculata (HBK.) M. Roem.Fam.Nat. Syn.2:143. 1846. Cieca appendiculata (G. F. W. Mey.) M. Roem.loc. cit. 145. 1846. Cieca cinerea(Poepp.& Endl.) M. Roem.loc. cit. 148. 1846. DecalobaRohrii(DC.) M. Roem.loc. cit. 156. 1846. Decalobacyathophora (Desv.) M. Roem.loc. cit. 157. 1846. Kegeliana Garcke, in Linnaea22:60. 1849. Passiflora torta Mast.in Mart.Fl. Bras.131:548. 1872. Passiflora cayaponioides in Bull. N. Y. Bot. Gard.8:107. 1912. Rusby, Passiflora cryptopetala Hoehne,in Comm. Linh. Telegr.Matto GrossoAnn. Bot. 5:76. Passiflora
pI. 112. 1915.

Rather coarse climbers, finely puberulent when young, becoming glabrate. Leaves petiolate,simple,basifixed, broadlyovate to ovate-lanceolate,entireor subentirewith the lateral lobes reduced to inconspicuousangles, 4-15 cm. long, 2-10 cm. broad, conspicuouslyocellate; petioles 1-3 cm. long, bearing 2 very large and auriculate glands near the base; stipules setaceous, up to about 1 cm. long. Inin pairs,bearinga singlegreenish florescences yellow,purple-tinged flower;peduncle 1 about cm. long, jointed above the middle; bracts inconspicuous and setose. Flowersabout 2 cm. in diameter, greenish yellowor white and purple-tinged toward the base; sepals narrowlyoblong-lanceolate,about 1 cm. long, not corniculate; petals linear,about half as long as the sepals; filamentsof the corona in 2 series, the outer about as long as the sepals. Fruit globose, 1.0-1.5 cm. in diameter, greenishyellow.

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BOTANICAL

GARDEN

Nicaragua to Bolivia and Brazil, fromabout sea level to 1,200 m.


TORO: vicinity of ChiriquiLagoon,Von Wedel I202; WaterValley,Von WedelI703. CANAL ZONE: Chagres, Fendler Sutton I22; Gatun, Hayes463; Agua Clara, Ft. Sherman, Standley 30949; Agua Clara Reservoir, Stevens 595; BarroColoradoIsland, Standley 41026, Zetek4364. DARIEN: Cana, Williams 964. BOCAS DEL

12.

PASSIFLORA

PEDICULATA

Mast. in Bot. Gaz. 23: 247.

1897; Killip, in Field

Mus. Publ. Bot. 19:113. 1938. Rather coarse climbers, glabrousthroughout. Leaves petiolate,simple,basifixed, broadly 3-lobed, the central lobe stronger, more or less deeply cordate at the base, 4-9 cm. long, 5-10 cm. broad, inconspicuously ocellate; petioles 1.5-5.0 cm. long, rather inconspicuouslyglandular about midway or above; stipules inconspicuous and subsetose. Inflorescences solitaryor paired, bearingsolitarygreenishyellow or white,purple tingedflowers;pedunclesabout 2-4 cm. long, jointed about midway; bracts inconspicuousand subsetaceous. Flowers about 3 cm. in diameter; sepals oblong-lanceolate, about 1.5 cm. long, not corniculate; petals linear,about 1 cm. long; filaments of the corona in 1 series,somewhatshorterthan the petals. Fruit not seen. Costa Rica and westernPanama, near sea level to about 1,500 m.
BOCAS DEL TORO:

WaterValley, Von Wedel854. L. Sp. P1. 958. 1753; Killip, in Field Mus. Publ. Bot.

13.

PASSIFLORA

SUBEROSA

19:88. 1938.

pallidaL. loc. cit. 1753. Passiflora hirsuta L. loc. cit.958. 1753. Passiflora minima L. loc. cit.959. 1753. Passiflora Passiflora nigra Jacq.Obs. 2:27. pI. 46, fig.3. 1767. glabraMill.Gard.Dict. ed. 8. Passiflora no. 4. 1768. Passiflora peltata Cav. Diss. 10:447. pI. 274. 1780. Passiflora Cieca viridis Medic.Malvenfam. 97. 1787. Cieca nigra(Jacq.) Medic.loc. cit. 1787. angustifolia Sw. Prodr. 97. 1788. Passiflora Sw. loc. cit. 1788 Passiflora parviflora heterophylla Dryand.in Ait. Hort.Kew. 3:309. 1789,nonLam. Passiflora Passiflora Lam. Encycl.3:38. 1789. hederaefolia longifolia Lam. loc. cit.40. 1789. Passiflora hederacea Cav. Diss. 10:448. 1790. Passiflora Granadilla suberosa (L.) Gaertn. Fruct.& Sem.2:480. 1791. Cieca heterophylla Method. (Dryand.) Moench, Suppl.101. 1802. Ciecasuberosa (L.) Moench, loc. cit. 102. 1802. Cieca minima (L.) Moench, loc. cit. 1802. litoralis HBK. Nov. Gen.& Sp. 2:138. 1817. Passiflora peltata(Cav.) Raf. Amer. Baldwinia Monthly Mag. 267. 1817. Monactineirma in Ann.G6n.Sci. Phys.Brux.2:138. 1819. angustifolia (Sw.) Bory, minima Mionactineirma loc. cit. 1819. (L.) Bory, Monactineirma suberosa loc. cit. 1819. (L.) Bory, peltata(Cav.) Bory, Monactineirma loc. cit. 1819. Monactineirma hederacea loc. cit. 1819. (Cav.) Bory, Vell.Fl. Flum.9: pI. 83. 1827. oliviformis Passiflora

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1958]

FLORA

OF PANAMA

(Passifloraceae)

13

globosa Vell. loc. cit. pl. 85. 1827. Passiflora Kohautiana Presl,Fl. Bemerk. 72. 1836. Passiflora villosa Macfad. Fl. Jam. 2:151. 1837, non Vell. Passiflora Meioperispeltata (Cav.) Raf. Fl. Tellur. 4:103. 1838. Meioperissuberosa (L). Raf. loc. cit. 1838. Meioperisminima (L.) Raf. loc. cit. 183 8. Meioperispallida (L.) Raf. loc. cit. 1838. Meioperisangustifolia(Sw.) Raf. loc. cit. 1838. Meioperishederacea (Cav.) Raf. loc. cit. 1838. PassifloraWarei Nutt. in Silliman's Journ.ex Torr. & Gray, Fl. N. Amer. 1 :539. 1838. Passifloralimbata Tenore, Ind. Sem. Hort. Neapol. 12: 1839. PassifloraflexuosaGardn. in Lond. Journ.Bot. 1:174. 1842. Fisch. Ind. Sem. Hort. Bot. Petropol. 9:82, ex Walp. Rep. 2:934. Passiflorapseudo-suberosa 1843. Cieca peltata (Cav.) M. Roem. Fam. Nat. Syn. 2:141. 1846. Cieca hederacea (Cav.) M. Roem. loc. cit. 1846. Cieca pallida (L.) M. Roem. loc. cit. 1846. Cieca angustifolia(Sw.) M. Roem. loc. cit. 143. 1846. Cieca olivaeforinis(Mill.) M. Roem. loc. cit. 144. 1846. Cieca globosa (Vell.) M. Roem. loc. cit. 1846. Cieca litoralis (HBK.) M. Roem. loc. cit. 145. 1846. Cieca pseudo-suberosa(Fisch.) M. Roem. loc. cit. 146. 1846. CieccaWarei (Nutt.) M. Roem. loc. cit. 1846. Cieca minima (L.) M. Roem. loc. cit. 147. 1846. Cieca flexuosa (Gardn.) M. Roem. loc. cit. 148. 1846. Cieca limbata (Tenore) M. Roem. loc. cit. 1846. PassifloralinearilobaHook. f. in Trans. Linn. Soc. 20:222. 1851. Hook. f. loc. cit. 1851. Passifloratridactylites puberula Hook. f. loc. cit. 223. 1851. Passiflora Passiflorasuberosavar. divaricata Griseb. in Bonplandia 6:7. 1858. suberosavar. minima (L.) Mast. in Trans. Linn. Soc. 27:630. 1871. Passiflora suberosavar. hirsuta (L.) Mast. loc. cit. 1871. Passiflora suberosavar. angustifolia(Sw.) Mast. loc. cit. 1871. Passiflora suberosavar. pallida (L.) Mast. loc. cit. 1871. Passiflora suberosavar. hederacea (Cav.) Mast. loc. cit. 1871. Passiflora suberosavar. lineariloba (Hook. f.) Mast. in Mart. Fl. Bras. 131:579. 1872. Passiflora suberosavar. longiloba Tr. & Planch. in Ann. Sci. Nat. V. Bot. 17:157. 1873. Passiflora suberosavar. longipesS. Wats. in Proc. Amer. Acad. 25:149. 1890. Passiflora Passifloracalliaquatica Krause, in Bot. Centralbl. Beih. 322:340. 1914.

Slenderclimbers, glabrous to denselypuberulent,the base of the stemsmore or extremely variable,usually definitely less corky. Leaves petiolate,simple,basifixed, 3-lobed,the centrallobe longer,rarelyalmostentire3-12 cm. long, 2-9 cm. broad, not ocellate or very inconspicuouslyso; petioles 1-2 cm. long, inconspicuously glandularnear the juncture with the leaf blade; stipulesinconspicuousand linear. flower;peduncles usually paired,each bearinga single small, greenish Inflorescences filiform,1-2 cm. long, jointed almost directly beneath the flower; bracts very minute. Flowers about 1-3 cm. in diameter; sepals lanceolate, 0.5-1.5 cm. long, of the corona in 2 series,the outer about not corniculate; petals absent; filaments half as long as the sepals. Fruit globose,deep purple, 0.5-1.5 cm. in diameter. PeninsularFlorida and southernTexas; Mexico to Brazil and Argentina; Bahamas; Antilles; introducedin the Old World tropics. Lowland thickets. MeWoncillo; huevo de gallo.

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14
30733.
PANAMA:
SAN

[Vol. 45 ANNALS OF THE MISSOURI BOTANICAL GARDEN

BLAS:

25357, Killip I2039; Bella Vista, Standley 4402. Puerto Obaldia,Pittier

Nuevo San Francisco, Standley

to the ubiquity and variabilityof P. The preposterous bibliographytestifies collected in Panama. suberosa. It is odd that it has been so infrequently
14.
PASSIFLORA PULCHELLA

HBK. Nov. Gen. & Sp. 2:134. 1817.

no. 13. 1768. bicornis Mill. Gard. Dict. ed. 8. Passiflora Passiflora rotundifolia Jacq.Obs. 2:26. pI. 46, fig.I. 1767,nonL. Passiflora DC. Prodr. 3:326. 1828. Passiflora rotundifolia var. Jacquinii DecalobaJacquini (DC.) M. Roem. Fam. Nat. Syn. 2:156. 1846. divaricata Griseb. in Bonplandia 6:7. 1858. Passiflora Mast.in Engl.Bot. Jahrb. 8:220. 1887. var. bifidata pulchella Passiflora Rather stout climbers, glabrous throughout. Leaves rather long-petiolate, simple,basifixed, broadly and cuneately 2-lobed, occasionally with a small central lobe, 2-6 cm. long, 3-9 cm. broad, rather inconspicuouslyglandular-ocellate; petioles1-3 cm. long, not glandular; stipuleslinear,5-10 mm. long. Inflorescences leaves,bearing solitary or rarelypairedin the axils of the greatlyreduceduppermost the associatedtendrilsbasal; peduncles solitaryinvolucrateblue or lavenderflowers, involucrate, 5-8 cm. long, jointed directlybeneaththe flower;bracts conspicuously broadlyobovate, cuneate at the base, entire,1.0-1.5 cm. long, pale greenoccasionally tinged with purple. Flowers about 4-6 cm. in diameter;sepals oblong, about 2 cm. long, not corniculate; petals ovate-lanceolate,about 1.5 cm. long; filaments of the corona in severalseries,the outer about as long as the petals. Fruit globose, about 1 cm. in diameter. SouthernMexico to Colombia and Venezuela; cultivated in Hawaii. Thickets near sea level.
PANAMA: Bella Allen & SeibertI42I. CANAL ZONE: Fort Kobe Road, Woodson, Standley 2I8; Las Sabanas, 4078I. Vista, KillipI2028, Bro.Heriberto alThis is one of the showiestPanamanian passionflowers because the flowers, raceme-likeclusters at the though not large, are borne in the very characteristic tips of the stems.

15. PASSIFLORA MEMBRANACEA Benth. P1. Hartw. 83. 1841; Killip, in Field Mus. Publ. Bot. 19:234. 1938. Cieca membranacea (Benth.) M. Roem.Fam. Nat. Syn.2:140. 1841. Rather stout climbers,glabrous throughout. Leaves rather shortlypetiolate, 3-lobed,the tip and lateral simple,barelypeltate,suborbicularand very indistinctly lobes mucronulate, 5-10 cm. long and about as broad; petioles 2-3 cm. long, and strongly eglandular; stipulesconspicuouslyfoliaceous,deeply cordate-reniform solitary at the nodes, bearing a amplexicaul, 1.5-3.0 cm. broad. Inflorescences white or cream flower;peduncles 10-15 single very strikingly involucrategreenish cm. long, jointed almostimmediately beneaththe flower;bracts very broadlyovate or suborbicular, very deeply cordate,4-5 cm. long and broad, pale green strongly suffused with rose. Flowers about 7-8 cm. in diameter; sepals oblong-lanceolate,

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1958]

FLORA

OF PANAMA

(Passifloraceae)

15

notcorniculate; petals oblong-elliptic, about as long as the sepals; about 4 cm.long, oblongtheouter about1 cm. long. Fruits in 2 series, filaments of thecorona
at thebase,4-6 cm. long and 2-3 cm. thick, yellowish ovoid,tapered green.
SouthernMexico to Panama, in highland forests. Muletoto summit, CHIRIQUf: Potrero Volcin de Chiriqui,Woodsond Schery389; Viejo,Allen1394. "New Switzerland", central valleyof Rio Chiriqui and attractivespeciesof the genus by virtue This is one of the most distinctive of the elongate peduncles and large petalaceous bracts which almost equal the flowers. 16.
PASSIFLORA ADENOPODA

DC. Prodr. 3:330. 1828; Killip, in Field Mus. Publ.

Bot. 19:222. 1938. Schlecht. & Cham.in Linnaea5:89. 1830. acerifolia Passiflora (Schlecht. & Cham.) M. Roem.Fam. Nat. Syn.2:151. 1846. Dysosmia acerifolia micranthumOerst. Amer.Centr.pI. 17. 1863. Ceratosepalurn Mast.in Trans.Linn.Soc. 27:630. 1871. ceratosepala Passiflora aspera Sesse& Moc. Fl. Mex. 227. 1887. Passiflora basiminutelyhispidulousto glabrate. Leaves long-petiolate, Coarse climbers, acuminate, fixed,deeply (3-) 5- to 7-lobed, the central and upper lobes stronger, deeply cordate, 7-15 cm. long, 8-17 cm. broad, membranaceous,hispidulous to stipitateglands glabrate; petioles 3-8 cm. long, bearing 2 very large prominently near the juncture with the leaf blade; stipulesbroadlyovate, amplexicaul,denticupaired, late, 1.0-1.5 cm. long and broad, foliaceous, persistent. Inflorescences bearing a single large purple and white flower; peduncle 3-4 cm. long, jointed somewhatbelow the middle and therebearing 3 lacerate subfoliaceousbracts about 3-4 cm. 1 cm. long. Flowers about 4-7 cm. in diameter;sepals oblong-lanceolate, 1.0-1.5 narrowly lanceolate, at the tip; petals long, prominentlycorniculate or the petals cm. long; filaments of the corona in a single series,about as long as densely in greenish yellow, cm. diameter, slightlylonger. Fruit globose, 2.0-2.5 puberulent. SouthernMexico to Venezuela and Peru at middle to higherelevations.
CHiIRIQUI:

El Boquete, Seemann I626.

The Seeman collection was found by Killip at both Kew and the British Museum; it was not examined for the presentaccount. 17.
PASSIFLORA VITIFOLIA HBK. Nov. Gen. & Sp. 2:138. Mus. Publ. Bot. 19:319. 1938.

1817, Killip, in Field

no. 45. 1819. J.E. Sm.in Rees,Cycl. 26: Passiflora sanguinea Passiflora R. & P. ex DC. Prodr.3:329. 1828. Pu-nicea Passiflora Tacsonia (J. E. Sm.) DC. loc. cit. 334. 182 8. sanguinea (J. E. Sm.) Raf. Fl. Tellur.4:103. 1838. Macrophora sanguinea 6:7. 1858. in Bonplandia serrulata var.pubescens Griseb. Passiflora Karst.in Linnaea30:163. 1859. serviteizsis Passiflora Lem.Ill. Hort. 14: pI. 519. 1867. Tacsonia Buchanani throughout. more or less denselyferruginous-pubescent Rather coarse climbers,

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[Vol. 45

16

ANNALS

OF THE

MISSOURI

BOTANICAL

GARDEN

deeply3-lobed,the serrulate, ratherinconspicuously basifixed, Leaves petiolate, 7-15 cm. long,8-17 cm. broad; cordate, moreor lessdeeply central lobestronger, the rather inconspicuous 2 to several glandstoward 2-6 cm. long,bearing petioles a single solitary, bearing 3-5 mm.long. Inflorescences subsetaceous, base; stipules 4-8 cm. long,jointedalmostdiflower; peduncle yellowand scarlet veryshowy petaloid, or somewhat and foliaceous bractsconspicuous theflower; rectly beneath
to nearlyentire,1.5-2.5 cm. long. Flowers 10-15 cm. lanceolate,glandular-serrate in diameter; sepals about 6-10 cm. long, corniculate; petals narrowlylanceolate, of the corona in 3 series,the outer 1.5-2.0 nearlyas long as the sepals; filaments yellow,denselypuberulent. cm. long. Fruits broadlyovoid, 3-6 cm. long, greenish

Nicaragua to Venezuela and Peru; Cuba and Jamaica; in lowland forests. Known as guate-guate and Pasionariain Panama. ColoradoIsland,Wetmore CANAL ZONE: Las CrucesTrail,Hunter& Allen456; Barro Fendler Abbe 76; between Summit and Gamboa, Greenman &f Greenman 5220; Chagres, & Schery 87. mountains beyondLa COCLE: ii8. CHIRIQUI: San Bartolom , Woodson Allen i66o. COLON: Camp Pifia, Allen 558; El Valle de Anton, Hunter Pintada, & Allen II50; Chepigana, Terry & Terry I374; Rio Sabana, 343I. DARIEN: Marraganti, NWilliams Maxon& Harvey6733, 6678, JuanDiaz region, LeopoldIII I26. PANAMA: Tapia River, TecumenRiver,Killip 33I3; OrangeRiver,Killip 3335; Pacora, 28IOI; 6683, Standley Bro.Paul 292. The dazzling flowersof P. vitifolia are frequent and widespreadin lowland thicketsand open foreststhroughoutthe Republic.
d

18.

Killip, in Journ.Wash. Acad. Sci. 12:262. PASSIFLORA WILLIAMSII Field Mus. Publ. Bot. 19:355. 1938.

1922; in

Coarse climbers,softly and densely puberulentthroughout. Leaves petiolate, deeply 3-lobed to somewhatbelow the middle, the central lobe stronger basifixed, and somewhat contracted toward the base, inconspicuouslyserrulate,8-15 cm. long, 9-16 cm. broad; petioles4-6 cm. long, with 2-3 ocellate glands toward the base; stipules filiform,inconspicuous. Inflorescencessolitary, bearing a single handsome violet flower; peduncle 1.0-1.5 cm. long, jointed directly beneath the flower;bracts involucrate,united toward the base and completelyenvelopingthe flowerbeforeanthesis,2.5-3.5 cm. long, denselypuberulent,pale green. Flowers about 6-7 cm. in diameter; sepals oblong, 3-4 cm. long; petals oblong-spatulate, of the corona in several series,the outermostless than 2.0-2.5 cm. long; filaments half as long as the petals. Fruits unknown. Known only fromPanama, at middle and low elevations.
4352.
CANAL ZONE: COCLE:

Paul 308.

and Mandinga, Colorado Island,Zetekl 548I; Barro Piper between Empire 585. PANAMA: Las Sabanas, Bro. Bismarck, above Penonome, Williams

This species bears 4 ratherconspicuousocellate glands upon the leaf blade, on each lobe toward the base of the two sinuses; the centrallobe thus with two glands, the lateral with one apiece.

(I 6)

1958]

FLORA 19.

OF PANAMA

(Passifloraceae)

17

PASSIFLORA MENISPERMIFOLIA HBK. Nov. Gen. & Sp. 2:137. 1817; Killip, in Field Mus. Publ. Bot. 19:457. 1938.

Passiflora villosaDombey,ex Triana& Planch.in Ann. Sci. Nat. ser. 5. 17:154. 1873, in synon. Rather coarse climbers,more or less densely pilose to glabrate throughout. Leaves petiolate, basifixed,broadly 3-lobed to somewhat above the middle, the central lobe much stronger,usually inconspicuouslyserrulatetoward the cordate base, 7-15 cm. long, 5-13 cm. broad; petioles2-5 cm. long, with 2 to severalpairs of slender stipitate glands; stipules conspicuously foliaceous, ovate-subreniform, deeply amplexicaul, 1.5-3.5 cm. long, glandular-denticulateto subentire. Inflorescences solitary,bearing a single white and purple flower; peduncle 4-6 cm. long, jointed directlybelow the flower;bracts rathernarrowlylanceolate, 1-2 cm. long, foliaceous,with ratherfew glandulardenticulationsto entire. Flowers about 6 cm. in diameter;sepals narrowlyoblong, 2-3 cm. long, corniculate;petals linearoblong, about equaling the sepals; filamentsof the corona in several series, the outermostabout half as long as the petals. Fruits broadly ovoid, 4-6 cm. long. Nicaragua to Peru and northernBrazil, from near sea level to about 1500 m. vicinity of ChiriquiLagoon,Von Wedel434, ii95, i406. CANAL MaddenDam, Allen 2007; Gatuncillo, Piper5623; BarroColoradoIsland,'Woodworth & Vestal 637. CHIRIQUI: Boquete,Terry I28I. VERAGUAS: Sona', 'Woodson,
BOCAS DEL TORO:
ZONE:

Seibert & Allen 5I5.

20.

PASSIFLORA SUBPELTATA Ortega, Nov. Rar. P1. Hort. Matrit. 6:78. Killip, in Field Mus. Publ. Bot. 19:436. 1938.

1789;

alba Link& Otto,Icon. PI. Rar. 65. p1.33. 1828. Passiflora Passiflora adenophylla Mast. in Mart. Fl. Bras. 131:568. 1872. Passiflora atomaria Planch.ex Mast.loc. cit. 570. 1872. Rather slender climbers,glabrous throughout. Leaves shortlypetiolate, basifixed,verybroadly3-lobed to about the middle,the 3 lobes subequal, rounded,with 2-3 pairs of inconspicuousserrulateglands in the sinuses,rounded to subcordateat the base, 4-9 cm. long, 5-12 cm. broad; petioles2-6 cm. long, with 1 pair of inconspicuous glands at about the middle; stipules conspicuouslyfoliaceous,broadly ovate, 1-4 cm. long and about half as broad. Inflorescences solitary,bearing a single purple and white flower; peduncle 2-5 cm. long, jointed directlybeneath the flower; bracts conspicuouslyinvolucrate, broadly ovate, about 1.0-1.5 cm. long and broad, abruptly acuminate, pale green somewhat tinged with rose. Flowers about 5-7 cm. in diameter; sepals oblong, about 3 cm. long, very conspicuouslycorniculate; petals narrowlylanceolate, 2.0-2.5 cm. long; filamentsof the corona in several series,the outermostabout equaling the petals. Fruits subglobose, 3-4 cm. in diameter. Mexico to Venezuela and Colombia, near sea level to about 2,800 m., in thicketsand open forest.
CANAL ZONE: Miraflores, White& White45; Red Tank,Maxon6574. Diaz, Standley 3205I; Taboga Island, Standley 2797I.

PANAMA:

Juan

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[Vol. 45

18

ANNALS

OF THE

MISSOURI

BOTANICAL

GARDEN

21.

PASSIFLORA LIGULARIS Juss. in Ann. Mus. Hist. Nat. 6:113. PA.40. Killip, in Field Mus. Publ. Bot. 19:344. 1938.

1805;

DC. Prodr. 3:328. 1828. serratistipula Passiflora

basifixed, Rather stout climbers,glabrous throughout. Leaves long-petiolate, entire,broadly ovate, deeply cordate, abruptly subcaudate-acuminate,8-17 cm. long, 6-15 cm. broad; petioles5-15 cm. long, with 3-5 pairs of elongatefiliform solitary,bearing a glands; stipulesfoliaceous,ovate, 1-3 cm. long. Inflorescences flower;peduncles 2-5 cm. single showy pale green,purple- and lavender-tinged long, jointed directlybeneaththe flower;bracts involucrateand foliaceous,ovate, 2-4 cm. long. Flowers about 7-9 cm. in diameter;sepals oblong, 3-5 cm. long; of the corona in several series, petals narrowlylanceolate,3-4 cm. long; filaments the outermostabout as long as the petals. Fruits broadly ovoid, 6-8 cm. long, purplishyellow. SouthernMexico to Venezuela and Bolivia, in highland forestsbetween 1,000 and 3,000 m.
CHIRIQUI:

vicinityof Finca Lerida, Woodson & Schery238.

22. PASSIFLORA QUADRANGULARIS L. Syst. Nat. 1248. 1759; Killip, in Field Mus. Publ. Bot. 19:335. 1938.
Granadilla quadrangularis(L.) Medic. Malvenfam. 97. 1787. tetragonaM. Roem. Fam. Nat. Syn. 2:165. 1846. Passiflora macrocepsMast. in Gard. Chron. 1869:1012. 1869. Passiflora

winged stems,glabrousthroughout. Leaves Coarse climberswith prominently entire,broadlyovate, abruptlyand shortlyacuminate,rounded petiolate,basifixed, at the base to broadlycordate, 10-20 cm. long, 8-15 cm. wide; petioles2-5 cm. long, with about 3 pairsof ovoid glands; stipulesfoliaceous,ovate, acuminate,2-4 cm. long. Inflorescences solitary,bearing a single greenishwhite flowerflushed with rose or purple; peduncles 1.5-3.0 cm. long, jointed directly beneath the so, flower;bracts delicatelyfoliaceous,broadlyovate, cordate,entireor essentially 2-4 cm. long and about as broad. Flowers about 8-12 cm. in diameter; sepals broadly oblong, conspicuouslycorniculate,4-5 cm. long; petals ovate-lanceolate, about equaling the petals. Fruits of the corona in 5 series, 3-5 cm. long; filaments with rose. broadlyoblongoid,up to 3 dm. long, pale yellow somewhatflushed Widely cultivated and escaping throughouttropical America; of uncertain
origin.
CANAL ZONE:

Chagres,Fendler 1Ig.

PANAMA:

JuanDiaz, Standley32057.

and is popularly known This is one of the most popular edible passionfruits, in Brazil. It as granadilla its range in Spanish America and maracuja throughout quality, and although capable of has a ratherinsipid fruityflavor of indefinite being eaten out of hand it usually is made into a conserveor a thin pudding after boiling in milk.

(18)

FLORA

OF PANAMA

(Passifloraceae)

19

23.

Mast. in Mart. Fl. Bras. 131:562. 1872; Killip, in Field Mus. Publ. Bot. 19:418. 1938.
PASSIFLORA QERSTEDIT

populifoliaTriana& Planch.in Ann.Sci. Nat. ser.5. 17:150. 1873. Passiflora Wash.Acad. Sci. 12:261. 1922. in Journ. Purpusii Killip, Passiflora Passifloradispar Killip, loc. cit. 330. 1922.

Rather slenderclimbers,glabrous throughout. Leaves rathershortlypetiolate, entire,ovate or oblong-ovate,acuminate, rounded to deeply cordate at basifixed, the base, 6-13 cm. long, 3-9 cm. broad; petioles 1-3 cm. long, bearing several pairs of small stipitateglands; stipules conspicuouslyfoliaceous,stronglyinequilateral, narrowlyovate, caudate-cuspidate,entire, 1-4 cm. long. Inflorescences solitary, bearing a single showy purple or lavender-roseflower; peduncles 3-6 cm. long, jointed directlybeneath the flower; bracts delicately foliaceous,ovate, roundedor cordate at the base, 1.0-1.5 cm. long. Flowers about 6-8 cm. in diameter; sepals ovate-lanceolate, 3-4 cm. long, conspicuously corniculate; petals of the corona in many series,the about 1.5 cm. long; filaments linear-oblanceolate,

4~~~~~

oerstedui Fig. 2. Passiflora

outermost equaling the petals or somewhat longer. Fruits broadly ovoid, 4-6 cm. long. Southern Mexico to Colombia and Venezuela, from near sea level to about 1500 m., in open forest. of El Valle WaterValley, Von WedelI756. COCLE: La Mesa,north Woodson d Schery Bajo Mona and QuebradaChiquero, CHIRIQUI: deChiriqui, Terry I303. 59I; Volcain
Allen 2399. de Anton,
BOCAS DEL TORO:

24.

PASSIFLORA AMBIGUA Hemsl. in Curt. Bot. Mag. p1. 7822. Field Mus. Publ. Bot. 19:3 63. 1938.

1902; Killip, in

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[Vol. 45

ANNALS

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BOTANICAL

GARDEN

so. Leavespetiolate, glabrous throughout or essentially Rather coarse climbers, or rounded at thebase, acuminate, obtuse basifixed, entire, ovateto oblong-elliptic, petioles 2-3 cm. long, thickly coriaceous; 10-20 cm. long,5-9 cm. broad,rather 2 thickocellate filiform, relatively inconthebase; stipules bearing glandstoward rose,or purple solitary, bearing a singleshowymaroon, spicuous. Inflorescences the flower; bractsfree, beneath flower; peduncles 5-7 cm. long,jointeddirectly or somewhat petalaceous. Flowers subfoliaceous ovate, 3-6 cm. long, delicately inconspicuously 8-12 cm.in diameter; narrowly oblong, 4-6 cm.long,rather sepals of the coronain lanceolate, 3-4 cm. long; filaments corniculate; petalsnarrowly broadly ovoid, theoutermost abouthalfas longas thepetals. Fruits several series, 10-12 cm. long,greenish yellow. nearsea levelto about 1000 m.,in thickets Southern Mexicoto Panama,from and openforest.
COCLE:

8o. Wedel 25.

BOCAS

Von Wedel2324; ColumbusIsland,Von Fish Creek Mountains, Sutton 466. Gatuin, Hayes 430; BarroColoradoIsland,Bangharn Allen i686. north rimof El Valle de Anton,
DEL TORO: CANAL ZONE:

Publ.Bot. 19:374. 1938.

PASSIFLORA NITIDA HBK.

Nov. Gen.& Sp. 2:130. 1817; Killip,in FieldMus.

nymphaeoides Karst.in Linnaea30:165. 1859. Passiflora

Rathercoarseclimbers, Leavesrather shortly petiolate, glabrous throughout. rather serrate or serrulate, at least towardthe base, broadly basifixed, distantly rounded at thebase,9-18 cm. long,6abruptly acuminate, elliptic to lanceolate, with2 largeovoid glands cm. 10 cm. broad, petioles 1.0-1.5 long, subcoriaceous; and filiform.Inflorescences somewhat above the middle;stipules inconspicuous whiteand purple flower; peduncles 3-6 cm. long, showy solitary, bearing a single theflower; 3-4 cm. long,folibeneath bractsfree, ovate,entire, jointeddirectly sepalsoblongaceous or somewhat petalaceous. Flowers8-9 cm. in diameter; petalsaboutas long as the 4.0-4.5 cm. long,inconspicuously corniculate; elliptic, theoutermost abouthalfas longas in several of thecorona series, sepals;filaments thepetals. Fruits obovoid, 4-6 cm. long. and open Brazil and adjacentPeru,in lowlandthickets Panamato northern forests.
CANAL ZONE:

Lindsay 230. Island, Shattuck 675; Summit, Barro Colorado Griseb.in Bonplandia 6:7. 1858; Killip, in Field Mus.

26.

PASSIFLORA SEEMANNII

Publ. Bot. 19:347. 1938.

incanaSeemann,ex Mast. in Journ.Linn. Soc. 20:40. 1883, non Kerr, in synon. Passiflora in Ann. Sci. Nat. ser.5. 17:150. 1873. Planch.& Linden, Passiflora orbifolia

Leavespetiolate, basifixed, entire, stoutclimbers, glabrous throughout. Rather and shortly 5-10 cm. long, abruptly apiculate-acuminate, broadly cordate-ovate, sessile 3-7 cm. long,with2 rather inconspicuous glands 5-7 cm. broad;petioles below the leaf blade; stipuleslinear,about 1 cm. long. Inflorescences directly

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1958]

FLORA

OF PANAMA

(Passifloraceae)

21

solitary, bearing a singlehandsome whiteor creamflower strongly suffused with purple orviolet;peduncles 6-10 cm. long,jointed directly belowtheflower; bracts moreor lesspetalaceous, unitedat the base and completely enveloping the flower before anthesis, 3-5 cm.long. Flowers about8-10 cm.in diameter; sepals narrowly
ovate, 4-5 cm. long, ratherinconspicuouslycorniculate; petals oblong-lanceolate, 3-4 cm. long; filamentsof the corona in 2 series,the outer about 1 cm. long. Fruits ovoid, 3-5 cm. long and broad. SouthernMexico to Colombia, at near sea level to about 1,500 m.

BOCAS DEL TORO: LincolnCreek,Rio Changuinola, Dunlap 422; vicinity of Chiriqui Lagoon,Von WedelI260, I366; WaterValley,Von Wedel i679. CANAL ZONE: Bohio, Maxon4765; Gamboa, Pittier 25i5, Allen3930; between Summit and Gamboa, Greenman d Greenman 522I; Gatuin, Bro. Heriberto 53; betweenMt. Hope and Sta. Rita Trail, Cowell 59; Las Cascadas,near Summit, Standley 29566; East Paraiso,Standley29838; QuebradaCulebra, Dodge d AllenI7048; Las CrucesTrail,Hunters. n.; Chagres,.Fendler I20; Miraflores, Whited White42. COLON: between FranceField and Catival,Standley 30286. DARIEN: Tucute,Chepigana District, Terryd TerryI393. PANAMA: hillsabove Campana, Alleni694; Tapia River, Maxond Harvey6730, Standley 28I79, Killip3297; Juan Diaz, Standley 30634.

This handsomespecieshas been introducedinto cultivationin Mexico, Nicaragua, and Hawaii, according to Killip. According to Killip also, the leaves occasionallyare lobed, but those of all the specimenscited above are entire.
27.
PASSIFLORA FOETIDA

19:481. 1938.

L. Sp. P1. 959. 1753; Killip, in Field Mus. Publ. Bot.

vesicaria L. Amoen. Passiflora Acad. 5:3 82. 1760. hibiscifolia Lam.Encycl.3:39. 1789. Passiflora ciliataDryand.in Ait. Hort.Kew. 3:310. 1789. Passiflora Granadilla foetida(L.) Gaertn. f. Fruct.& Sem.1:289. 1790. Passiflora gossypifolia Desv. in Hamilt.Prodr. Fl. W. I. 48. 1825. Tripsilina foetida(L.) Raf. Fl. Tellur.4:103. 1838 (as fetida) . nigelliflora Passiflora Hook. Bot. Mag. pA. 3635. 1839. hastata Passiflora Bertol. Fl. Guat.427. 1840. Dysosmia foetida(L.) M. Roem.Fam.Nat. Syn.2:149. 1846. Dysosmia gossypiifolia (Desv.) M. Roem.loc. cit. 1846. hastata(Bertol.)M. Roem.loc. cit. 1846. Dysosmia Dysosmia hibiscifolia (Lam.) M. Roem.loc. cit. 1846. Dysosmia ciliata(Dryand.) M. Roem.loc. cit. 1846. M. Roem.loc. cit. 150. 1846. Dysosmia fluminensis Dysosmia nigelliflora (Hook.) M. Roem.loc. cit. 151. 1846. Lem.in Illustr. Hort. 8: pA. Baraquiniana 276. 1861. Passiflora Liebmanni Mast.in Mart.Fl. Bras.131:547. 1872. Passiflora DC. ex Triana& Planch.in Ann.Sci. Nat. ser.5. 17:172. 1873. hispida Passiflora Moritziana Planch.loc. cit. 175. 1873. Passiflora muralis Barb.Rodr.in Contr. Bot. Rio Jan.1:29. 1891. Passiflora Jard. Balansae Chod.in Bull.Herb.Boiss.ser.2. 2:744. 1902. Passiflora in Bull.Torrey Bot. Club 44:19. 1917. pseudociliata Passiflora Britton, varietal and formal (Also verynumerous designations). Rather slenderclimbers,commonlymore or less yellow-pubescent throughout, less frequently glabrateor glabrous. Leaves petiolate,basifixed, extremely variable, in Panama chiefly broadlyand acutely 3-lobed to about the middle,the centrallobe

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ANNALS

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BOTANICAL

GARDEN

usually about 4-12 cm. long and wide; petiolesusually about 2-6 cm. far stronger, long, usually beset with mixed glandular and eglandular hairs; stipules subsetasolitary,bearing a single greenishwhite or ceous, inconspicuous. Inflorescences flower;peduncles 3-7 cm. long, jointed directly yellow,more or less purple-tinged each filiform beneath the flower;bracts foliaceous,complexly 2- or 3-pinnatifid, divisiongland-tipped,usually 2-4 cm. long. Flowers about 4-6 cm. in diameter; sepals oblong-lanceolate,about 2-3 cm. long, shortlycorniculate; petals oblongoblanceolate, slightlyshorterthan the sepals; filamentsof the corona in several series,the outermostabout 1 cm. long. Fruits subglobose,yellow or orange,about 2-3 cm. in diameter. Common throughouttropical America in lowland thickets and open forest, becoming a weed. Von Wedel2I42; WaterValley,Von WedelI7i8, BOCAS DEL TORO: Old Bank Island, of ChiriquiLagoon,Von Wedel 1562. CANAL ZONE: Las CrucesTrail, 183I; vicinity Fendler 117; Ft. San Allen 899; Chagres, of Rio Chagres, Hunter & Allen 724; nearmouth Sutton Pittier 6954; GatunStation, 7OI3; MatiasHernandez, Maxond Valentine Lorenzo, 2678; Rio Pittier Cowell393; valleyof Masambi, Hayes597; AhorcaLagartoto Culebra, Grande,near Culebra,Pittier2089. CHIRIQUI: Ilanos west of Gualaca, Allen 5048. and Yavisa,Allen292. Pinogana COLON: Porto DARIEN: trailbetween 2474. Bello,Pittier 3569; Tapia River,Maxon6719. 2042, Pittier Miller PANAMA: Taboga Island,
Panama. Passiflora foetida must be frequent at low elevations in every province of The species, although unmistakable, is extremely variable in small details,

and Killip recognized no less than 42 named varieties. All of the specimens to var. isthmiaKillip. above he referred enumerated

CARICACEAE
shrubsand small trees,usually dioeciousor monoecious Soft-woodedlaticiferous branchingtrunksornaand with disproportionately thick, simple or infrequently mentedwith large leaf scars. Leaves spiral,usually quite large and long-petiolate, exstipulate,the blade palmately compound or simple and variously lobed, rarely axillary at the uppermostnodes, the staminatean extensive entire. Inflorescence the pistillatemuch less compoundedand with fewer and highlycompound thyrse, and larger flowers,the fruits developed at the base of the young growth and appearing cauliflorous. Flowers dichlamydeous,5-merous, regular, hypogynous, so in the pistillate;calyx lobes 5, imbricateor but usually imperfectly sympetalous or infundibuliusually inconspicuous;corolla salverform nearlyopen in aestivation, form and with a well-developedtube in the staminate,more or less campanulate and ratherimperfectly sympetalousin the pistillate,the limb 5-lobed with contortedaestivation; stamens 10 in 2 whorls insertednear the orificeof the corolla tube, the antepetalous usually with somewhat shorterfilamentsthan the antesepalous, the filamentsoccasionally more or less petaloid or with petaloid appendages,reduced to staminodiaor absent in pistillateflowers;pistil superior,3to 5-carpellate, usually 1-celled and with parietalplacentation,rarely3- to 5-celled

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1958 1

FLORA

OF PANAMA

(Caricaceae)

23

the stigmas3-5, the ovulesnumerous, anatropous, and with axile placentation, reducedto an rarelysubcapitate, so, usuallydichotomous, sessileor essentially small flowers.Fruita very largeto rather in staminate or absent pistillode acicular and funicles. seedswithpulpytestas thenumerous berry, 1- to 5-celled succulent
The Caricaceae are a small familyof four genera,all tropical American except the bitypic Cylicomorpha of Africa. The plants are of strikinglyantediluvian aspect with theirstout,usually simple trunksheavilyornamentedwith large spiral leaf scars and their terminalcrowns of fern-likeleaves. The latex tubes, which traverseall parts of the plant, are associated with various proteolyticenzymes, particularlypapain.
.-------------1. JACARATIA ovary5-celled a. Leavespalmately petalsantesepalous; compound; --2. CARICA ovaryi-celled. lobed; petalsalternisepalous; aa. Lcavespalmately

1. JACARATIA Marcgr. ex Endl.


JACARATIA

Marcgr. ex Endl. Gen. 933. 1839 (as Jaracatia).

Pileus Ramirez, in An. Inst. Med. Nac. Mex. 5:29. 1901.

Dioecious shrubs and trees, branching rather frequentlyfor the family, the trunk becoming very thick. Leaves palmately compound, with 3-12 petiolulate, in the axils of the uppermostleaves, entire or undulate leaflets. Inflorescences the pistillatereduced and flowered, many and extensive the staminate thyrsiflorous, Corolla salverformor inconspicuous. 5-lobed, Calyx with few larger flowers. Stamens 10, the filaantesepalous. 5-lobed, limb the narrowlyinfundibuliform, Ovary 5-celled, con1petaloid. or less conspicuously more ments and connectives rather elongate,5-celled. usually Berry placentas. axile on ovules tainingnumerous Paraguay. to Mexico from extending dozen species a Probablyabout
1. J. DOLICHAULA leaflets 3-5, greenbeneath ................................ a. Stems unarmed; aa. Stems armed with conspicuousspines; lcafiets5-), very pale and 2. J. COSTARICENSIS glaucous beneath
1. JACARATIA DOLICHAULA

(Donn. Sm.) Woodson, in Ann. Missouri Bot. Gard.

- 37:404. 1950. in Bot. Gaz. 23:247. 1897. Donn.-Sm. Caricadolichaula Trees up to 12 m. tall, the trunk usually thickened toward the base, the branchesglabrousand unarmed. Leaves palmatelycompound with 3-5 petiolulate broadly elliptic or obovate, caudate-acuminate leaflets6-15 cm. long, 2-6 cm. broad, greenbeneath,the petiolulesabout 1 cm. long; petiolesslender,up to about much shorterthan the subtendingpetioles,the sta15 cm. long. Inflorescences the pistillateusually 1- to 3-flowered. Staminate minate up to about 30-flowered, flowers: calyx lobes broadly ovate, about 1 mm. long; corolla slenderand salverform,the tube 3-5 cm. long, about 3 mm. in diameter,scarcely dilating toward obtuse or acute, white,the lobes oblong-spatulate, pale greenor greenish the orifice, with pink toward tinged white occasionally 1.0-1.25 cm. long, slightlyspreading, antepetalous the exserted, tube, the corolla of the orifice at the tip; stamensinserted

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24

ANNALS

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BOTANICAL

UVol. 45

GARDEN

Fig. 3. facarai4 dolichaula

with very shortfilaments, the alternating whorl sessile,the anthers narrowly about 5 mm.long,theconnective oblong, and somewhat thanthe petaloid longer anther;pistillode narrowly acicular,about 1 cm. long. Pistillate flowers:calyx lobes broadly ovate,about 1-2 mm. long; corollashortly the tube salverform, about 1.5 cm. long,about 5 mm.in diameter at thebase,contracting towardthe

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FLORA OF PANAMA

(Caricaceae)

25

middle, then dilated to about7 mm.at theorifice, white, thelobesnarrowly oblong, about 1.5 cm. long,slightly spreading, white;staminodia absent or inserted at the orifice of the corollatube,sessile, like the stamens essentially of the staminate flower but the anthers somewhat smaller and the petaloidconnectives somewhat larger;ovaryoblong-ovoid, about 5 mm.long,5-celled, the stigma with 5 ovate essentially entirelobes. Berries ovoid-fusiform, about 10 cm. long and 5 cm. broad, persistently 5-celled, numerous containing ovoidseeds7-8 mm. compressed long; fruiting peduncle about 3.5 cm. long. CostaRica and Panama, in forests, 200-1000 m. elev. Popular names recorded in Costa Rica are papaya, by Standley palo de barril, papayade monte, papayillo, who explains thelast as follows:"The palo de barril, whichI believe referable to thisspecies, is common in Guanacaste, where it is a largetreewithbroadcrown and a verythicktrunk. This is so softor spongy that witha few slashes of a it is possible machete to fella largetree. If a largetrunk is thuscut, and leftfor someweeksto dry,it is possible to separate a cylinder of bark. Sections of this are usedin place of barrels, forstoring maizeand otherarticles."(Fl. Costa Rica
2:724.
VERAGUAS:

1937).
DEL TORO: Fish Creek,vicinity of ChiriquiLagoon,Von Wedel I084, forested of CerroTute,vicinity of SantaFe, Allen4392. slopes

BOCAS

2207.

2.

JACARATIA COSTARICENSIS

I. M. Johnston, in Contr. Gray Herb. 70:79.

1924.

Trees up to 20 m. or more, the branches armed with numerousshort stout spines, glabrous. Leaves palmately compound with 5-9 very shortlypetiolulate oblanceolate shortlyacuminate entire leaflets7-12 cm. long and 2-3 cm. broad, dark greenabove, very heavily glaucous beneath,the petiolulesabout 1.0-1.5 cm. long; petiolesslender,10-20 cm. long. Inflorescences shorterthan the subtending petioles, the staminate racemose-paniculate,many-flowered, the pistillate erect, long-pedunculate, 1-flowered. Staminateflowers about 17 mm. long, very glaucous without,narrowlyclavate in bud, the lobes about 7 mm. long, linear,obtuse, the tube narrowly cylindric,about 1.2 mm. broad, sparselyvillous within; stamens strigose, stronglyunequal, the filaments conspicuouslyconnate,the antherssimilar, without a ligulate connective,with a small erect spiniformprocess. Berry pyriform, up to about 7 cm. long and 3 cm. broad, orange, the fruitingpeduncle pendulous,up to about 10 cm. long. Costa Rica and Panama, in forests near sea level to about 700 m. elev. Tonduz reportsthe vernacularname in Costa Rica as papayillo de venado. vicinity of CampPifia, alt. 25 m.,Allen3679. The description of the flowers is taken fromJohnston;our specimenis fruiting. Jacaratia costaricensisis closely related to J. spinosa (Aubl.) DC., as noted by Johnston. Both species are very glaucous, particularlythe lower surface of the leaves, and the chief distinguishing characterobvious from our meagre herbarium representation appears in the length of the fruitingpeduncles,which are only up to about 2 cm. and suberectin J. spinosa.
COLON:

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[Vol. 45

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2. CARICA L.
CARICA

L. Spec. PI. 1036. 1753.

Papaya[Tourn.]Adans.Fam. P1.2:357. 1763.

Vasconcellea St. Hil. 2e. Mem. Resed. 13. 1837.

Dioecious or monoecious trees and shrubs, the thick succulent trunk usually unbranched. Leaves simple and palmatelylobed in the Panamanian species,longthe in the axils of the uppermostleaves, thyrsiflorous, petiolate. Inflorescences the pistillate reduced and with fewer staminate extensive and many-flowered, in the staminate largerflowers. Calyx 5-lobed,inconspicuous. Corolla salverform campanulate in the pistillate,the limb 5-lobed, alternisepalous. Stamens flowers, 10, the connectiveoccasionallyproducedat the tip. Ovary usually 1-celled,usually septate at the base, containing numerous ovules on parietal placentas. Berry i-celled. broadlyovoid-pyriform, Although the papaya is one of the most common and favorite fruits of the tropics,appearingto the outlanderlike some celestialmelon, the genus is in a state of botanical confusion with at least 50 poorly defined "species" ranging from Mexico to Argentinaand in the Antilles. It has been introducedinto the Eastern escapes. Hemispheresince long ago and frequently
a. Leaveswith7 palmate primary veins,usuallywith7 deeplyand irreguwith 5 dichotomous flowers pistilof pistillate segments; larlypinnatifid flowers with broadlydeltoid or trichotomous stigmalobes; staminate 3-4 cm. long. 1. C. calyxlobesabout 1 mm.longand yellowor whitecorollas with5 shallowsegments each with aa. Leaveswith5 palmate primary veins, with 5 linearunflowers 1-2 broad laterallobes; pistil of pistillate divided lobes (presumptive forC. chiriquensis). stigma so, the middlesegb. Leaves and inflorescences glabrousor essentially the4 lateralsegments unlobed;(staminate mentof theleaves3-lobed, with whitecorollasabout 4 cm. long and ovate calyx lobes flowers . . ..... 2. C. about 1 mm. long?).............. ................ rather bb. Leavesand inflorescences villous,the middlesegment sparsely with a singlebroadly of the leaves 3-lobed,the 4 lateralsegments corollas1.5-2.5 cm. long flowers withpale green acutelobe; staminate calyx lobesabout 3 mm.long....................................... 3. C. and lanceolate

PAPAYA

CUCURBITIFOLIA

CHIRIQUENSIS

1. CARICA PAPAYA L. Sp. P1. ed. 1. 1466. 1753.


1824. Papaya sativa Tussac, FI. Ant. 3:45. p1. I-II. Carica Mamaja Vell.Fl. Flum.10: p1. I3I. 1827. Papaya vulgarisA. DC. in DC. Prodr. 151:414. 1864. Blanco, Fl. Filip. 3:212. 1879. Carica hermaphrodite

Dioecious or occasionally polygamous or monoecious trees 2-8 m. tall with stout succulent and unbranchedtrunks,usually glabrousin all parts characteristic or essentiallyso. Leaves in a dense terminalcrown, long-petiolate;blade simple and broadlypinnatifid but palmatelydivided into usually 7 more or less irregularly up to 4-5 dm. in diameter;petiole usually variable in size but frequently segments, compound,many-flowered, repeatedly up to 5-7 dm. long. Staminateinflorescences than the subtendingpetioles,spreadingor pendulous,the pistillate somewhatshorter far shorterthan the subtendingpetioles,usually only 1- to 3-flowered. Staminate

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Fig. 4. Carica Papaya

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flowers: calyxlobesvery broadly deltoid, obtuse, about1 mm.long;corolla salverform whiteor yellow, thetubeabout1.5-2.0 cm. long,about2 mm.in diameter, the lobes oblong,1.0-1.5 cm. long, slightly spreading; stamens inserted at the orifice of the corollatube,exserted, theantepetalous anthers about 2.5 mm.long, nearly sessile and descending basally intothecorollatube,theantesepalous anthers erect, wholly exserted, about2 mm.long,withapically 2-lobed filaments of nearly equal length;pistillode acicular,about 1 cm. long. Pistillate flowers far larger than the staminate, irregularly campanulate, whiteor pale cream: calyx lobes broadly deltoid, about 1 mm.long; corollaessentially polypetalous or veryweakly thesegments united, about5-7 cm. long,1.5-2.0 cm.broad, broadly acute,irregularlyreflexed toward thetips; staminodia lacking, rarely minute and hypogynous; ovary ovoid-ellipsoid, usually about3 cm. long (including theindefinitely attenuate style) and 2 cm. broad,the stigmaessentially sessile, of 5 thicksegments about 1 cm. long fimbriately di- or trichotomous towardthe tip. Berryextremely variable, up to 3 dm. longand 2 dm. thick, green or yellow whenripe. Papayosare grownthroughout the tropics of the world,chiefly at low elevations,and tend to becomeescapeseverywhere. In Panama theyare one of the commonest weedtrees, and the qualityof their fruits variesfromthe sublime to the ridiculous. Someof the mostinferior producefruits scarcely largerthan a hen'segg. The vast man-made distribution of Carica Papayais quiteat variance withtheveryrestricted natural ranges of theother published species of thegenus, and eventheapproximate siteof theprogenitors of thecultivated papayais now to ascertain:it may well be the eastern impossible slopesof the Peruvian Andes where mostof thewild species withsimilar leafforms occur today. Undersuch the citationof herbarium circumstances is apt to becomesomewhat specimens but is appended absurd, to thisaccountforgood measure. Associated withthe latexof all partsof the plantare various enproteolytic zymes, particularly papain, whichareisolated and usedforvarious pharmaceutical and gastronomic purposes.In Panamaleavesof papayosometimes are cookedwith toughmeat,or theraw meatsoftened by wrapping in green leavesand leftovernight. HernandoCortesis said to have observed thisuse of papayoduringhis and fromthis sourcehas conquestof Mexico in the earlyeighteenth century, in whichpapainis of meat tenderizing, the recently developed booming process a prominent agent. is dioecious. In somedistricts Carica papayanormally of Panamatheinhabithatbarren(i.e. staminate) treesof papayocan tantsadhere to a "superstition" in the bark the Sign of the Crosswitha be made fruitful by deeply inscribing which do indeed machete. I have been shownexamples of such incisedtrees, also has it thatsuch a treedoes appearto become polygamous.Local experience fruitful but mustbe wounded The response notbecome permanently periodically. foundation reversion. wouldappearto have scientific upon thebasisof traumatic
BOCAS DEL TORO: Careening Cay, Von Wedel 573. CANAL ZONE: Barro Colorado & Greenman of Ancon, Greenman Island,Shattucki65; low ground,outskirts 5073; CocoliIsland, of Miraflores vicinity Lake,P. White 290.

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Fig. 5. Carica cucurbitifolia

2. CARICA cucurbitifolia Woodson, spec. nov. Arboresca. 3-6 m. altae. Folia longe petiolata glabra lamina simplicipalmatim 5-divisa basi 5-venosa segmentocentralimaximo profundelateque 3-partitousque 15 cm. longo et lato vel ultra (speciminisnostrijuvenalis) segmentis2 mediis ca. 12 cm. longis 6 cm. latis atque 2 basalibus 6 cm. longis 3 cm. latis omnino integris.

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Inflorescentia mascula jam ignota feminea valde contracta usque ca. 7-flora pedunculo ca. 2 cm. longo. Florum femineorum calycis laciniae minute trigonales ca. 2 mm. longae. Corolla ejusdem generistubulosa ca. 2.5 cm. longa tubo ca. 3 mm. diam. basi videturlacteo lobis anguste oblongis ca. aequilongiserectis apicem versus roseo-tinctis. Staminodia nulla. Ovarium in floreovoideum ca. 6 mm. longum glabrumin styloangusto ca. 3 mm. longo abruptecontractumstigmatibus 5 linearibusintegris10-12 mm. longis. Bacca ignota. ridgeknownas the COCLE: in heavyforest belowgrass, region north of El Valle,grass Bot. Gard., Loma del Tigre,alt. 2800 ft.,April8, 1947,P. H. Allen4478 (Herb. Missouri
HOLoTYPE).

The ovary of C. cucurbitifolia, with its slenderstyle,recalls that of C. stylosa Heilborn of Peru, the leaf segmentationof which, however, correspondsmore closely with that of C. papaya. It is unfortunatethat staminateflowersare not known. In the herbarium of the MissouriBotanical Garden is a staminatespecimen of papayo collected by Skutch (no. 4797) in Costa Rica which appears to be related to C. cucurbitifolia, although with a differently divided leaf more similarin outline to that of the succeeding species. The Skutch plant has white corollas is not about 4 cm. long and ovate calyx lobes about 1 mm. long. It definitely C. papaya as presently determined. 3. CARICA chiriquensis Woodson, spec. nov.

Arboresusque 7 m. altae vel ultra. Folia longe petiolata lamina simplici palmatim 5-divisabasi 5-venosa segmentocentralimaximo profundelateque 3-partito usque 35 cm. longo nisi ultra segmentismediis ca. 25 cm. longis 12 cm. latis margineinferiore late auriculato segmentis2 basalibus ca. 20 cm. longis 10 cm. sparsiuscule latis margineinferiore acute auriculato membranaceapagina inferiore villosula. Inflorescentia femineajam ignota mascula effusethyrsiflora pedunculis gracilibus dense villosulis. Florum masculorum calycis laciniae lanceolatae apice acuminatae laxe reflexae ca. 3 mm. longae margineminute ciliato. Corolla florum ejusdem generispallide viridismagnitudinevalde variabilistubo anguste cylindrico extus sparsepilosulo ca. 1.5 mm. diam. tum 5 mm. tum 15 mm. longo lobis linearibus ca. 1 mm. latis extus apice versusdensiuspilosulis tum 7 mm. tum 12 mm. longislaxe patulis. Stamina 10 ad ostium corollaetubi posita exsertaantepetalorum antheriscum appendice apicali ca. 1.5 mm. longis filamento ligulariintegroca. 0.5 similised 2 mm. longo. Pistilmm. longo alternorum vix 1 mm. longis filamento ca. 8 mm. longum. Bacca ut diciturca. 15 cm. longa lodium angusteaciculiforme 13 cm. lata 5-angulata.
CHIRIQUI: QuebradaVelo, vic. Finca Lerida,alt. 5000 ft.,July24, 1947,P. H. Allen Bot.Gard.,HOLOTYPE). 4675 (Herb. Missouri

Perhaps it is temptingfate to describe two new dioecious species, apparently areas as Chiriqui and Cocle when only the closely related,fromsuch neighboring as indicated in the key sexual formsare known. The formsdiffer complimentary is glabrouswhile C. chiriquensis and the descriptions, however,and C. cucurbitifolia (30)

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is villous. The leafform of chiriquensis is quitesimilar to thatof theSkutch plant fromCosta Rica which was discussed under cucurbitifolia, but the staminate flowers arestrikingly dissimilar.

Fig. 6. Carica chiriquensis

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LOASACEAE
Mostly perennialor annual herbs,sometimes quite massive,erect or clambering, infrequently twining or climbing, characteristicallystrongly hispid and with stinging hairs and glochids. Leaves opposite or alternate,variously incised and occasionally palmatifidor pinnatifid; stipules lacking or vestigal. Inflorescence terminalor extra-axillary, basically cymose but frequentlymonopodial and occasionally reduced to a solitary flower. Flowers dichlamydeous,hermaphrodite, epigynous,5- or rarely4-, 6-, or 7-merous; sepals foliaceous,imbricateor open in aestivation; petals free or united at the base, imbricated or contorted; stamens rarely5-10 or fewer,usually very numerous,the filaments more or less united to the base of the corolla, all fertileor partly staminodialand more or less petaloid; ovary inferior, usually 3- to 5-carpellate and 1-celled with numerous ovules on parietal placentas, rarely 1-carpellatewith a single pendulous ovule (Gronovia), the style simple with a capitate or obscurely divided stigma. Fruit a variously dehiscentcapsule. Except for the peculiarmonotypicgenusKisseniaof Africa and southern Arabia, the Loasaceae are an American familymost frequentin arid or semi-aridregions with greatestdiversification in Mexico and the southwestern United States and in Chile. Those who have encounteredthem in the fieldremember them henceforth chiefly because of theirpainfullystinginghairs which seem to have been specially contrivedto protect the elaborateand frequently very showy flowers.
stamens 10 to many, frequently in part; erector sprawling staminoidial herbs;leavesvariously incised or lobed,sometimes pinnatifid or pinnate. b. Stamens all fertile or theouterbecoming centrifugally petaloid. 1. MENTZELIA bb. Fertile stamens antesepalous, staminodia antesepalous. c. Flowers 4-merous; staminodia freeor unitedonlyat thebase - . 2. KLAPROTHIA cc. Flowers 5-(rarely6- to 7-)merous;staminodia unitedinto strongly a moreor less saccatepetaloid scale. -.-----------------------------.-.-....3. LOASA aa. Ovary 1-carpellate, witha singleapical pendulous ovule; stamens 5, all slender leavesverydeeplyand palmately fertile; twiners; 4. GRONOVIA divided -----------a. Ovary 3- to 5-carpellate, with numerous ovules or parietal placentas;

The genus Sclerothrixextends from southernMexico to Bolivia and is to be expected in Panama. It resemblesKlaprothia strongly,but may be recognized readilyby its spirallycontortedfruits. 1. MENTZELIA Deutsch. Akad. Naturf. 76:22. 1900. L.

MENTZELIA L. Syst. ed. 10. 1076. 1759; Urb. in Nova Acta Abh. K. Leop.-Carol.

Bartonia in Sims, Pursh, Bot.Mag. sub t. I487. 1812, nonMuhl. Acrolasia Presl, Rel. Haenk.2:39. 183 5. Torreya Eaton,Man.Bot. ed. 7. 560. 1836, nec Arm. nec alior. Creolobus Sver.OdR.Vext. 67. 1839. Lilj. Fl. Oefver Trachyphytum Nutt. ex Torr. & Gray, Fl. N. Am. 1:533. 1840. Chrysostoma Lilj. in Linnaea 15:263. 1841. Hesperaster Cockerell, in Torreya 1:142. 1901.

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Fig. 7. Mentzeliaaspera

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Herbaceous annualsor perennials, rarely smallshrubs or trees, the stemsand foliagescabrous and hispid, but without stinging hairs. Leaves alternate or opposite,variously incisedor lobed. Inflorescence terminal, cymose, usuallyfewflowered. Flowers largeto mediocre; sepals foliaceous, 5; petals5-10, free orbarely united at thebase; stamens 10 to verynumerous, free or united at thebaseof the filaments, whollyfertile or the outermost becoming petaloidand sterile;ovary inferior, 1-celled, withfewto numerous ovulesborne upon 3-7 parietal placentas, thestyle elongate, filiform, thestigma scarcely dilated, capitate or obscurely divided. Fruitan apically dehiscent 3- to 7-valvedcapsule. A genusof nearly100 species, principally in the western UnitedStatesand but extending Mexico, to Patagonia. A alongthePacificcoastof SouthAmerica single species is knownfrom Panama.
1. MENTZELIA
ASPERA L.

Spec. P1. ed. 1. 516. 1753.

Mentzelia stipitata Presl, Rel. Haenk. 2:40. 183 1. Mentzelia pedicellata Presl, Epim. Bot. 246. 1849. Acrolasia squalida Hook. f. in Trans. Linn. Soc. 20:222. 1851.

Slender erector sprawling annuals. Stemsslender, dichotomous, up to about 6 dm.long,rather thinly covered withshort weakglochidiate hairs. Leavesalternate, rathersparse,shortly but evidently petiolate, the blade ratherhastately 3-lobed and irregularly serrate, acuminate, broadly obtuseat thebase,about8 cm. longand 4 cm. broador somewhat larger toward thebaseof thestem, diminishing gradually towardthe tip, lax and thinly membranaceous, weaklyand variously strigillose withsimple hairs aboveandbelow, thepetiole about5 mm.long. Flowers in few-flowered leafy-bracted cymesat the tips of the branches, mediocre, pale or salmon, orange sessile or essentially so; hypanthium narrowly cylindrical, about 1 cm. long and 1 mm.broadat anthesis, densely covered withslender glochidiate hairs,the calyx lobes broad trigonalat the base, very narrowly subcaudateacuminate, nearlyas long as the hypanthium, sparsely strigillose but without glochidiate hairs;petals5, obovate, up to 1 cm. long and 5 mm.broad, spreading, or salmon, pale orange united at thebase; stamens slightly about20-30, all fertile, but theoutermost somewhat and withenlarged longer somewhat petaloid filaments, somewhat shorter thanthe petals. Capsulenarrowly cylindrical but slightly attenuate towardthe base,up to about 2.5 cm. long and 5 mm. broad,3-valved, covered withthepersistent glochidiate hairs. One of the mostwidespread of the genus, fromArizonato species occurring in drythickets Argentina.Ratherfrequent uponthePacific slopeof Panama.
L CANAL ZONE: Balboa Heights, Greenman & Greenman 5055; AnconHill, Greenman Greenman Standley26360; between Corozal and Ancon, Pittier2209; Gamboa, 5II4, Pittier 4793; Sosa Hill, Balboa, Standley 25256, 32I50; Darien station,Standley 3I52I.

2. KLAPROTHIA HBK.
KLAPROTHIA

HBK. Nov. Gen. & Sp. 6:96.

1823.

Herbaceous annuals, the stemsand foliageratherinconspicuously hispidulousto

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essentiallyglabrous,without stinginghairs. Leaves opposite, distinctlypetiolate, serrate. Inflorescence terminal,diffusely cymose, bearing rather numerous small 4-merous flowers. Hypanthium subglobular,the calyx lobes 4, broadly trigonal; petals 4, free; stamens in antepetalous groups of 4-5, with slender minutely papillate filaments;staminodiain antesepalousgroups of 4-5, with slenderpilosulose filamentsbarely united at the base and incised flabelliform sterile anthers; ovary 4-carpellary,1-celled,with 1-2 ovules upon the 4 parietal placentas; style the stigma very obscurely divided. Fruit a small septicidally4-valved filiform, capsule. A monotypicgenus of relativelyhigh elevationsextendingfromCosta Rica to Venezuela and Bolivia.

nzentzelioides Fig. 8. Klaprothia

1.

KLAPROTHIA

MENTZELIOIDES

HBK. Nov. Gen. & Sp. 6:96.

1823.

Weak erect or sprawlingherbaceousannuals. Stems slender,branchinginfrequently,up to about 6 dm. long, usually less, inconspicuouslypilosulose to essentially glabrous. Leaves opposite, rather sparse, lax and long-petiolate,the blade elliptic to ovate or lanceolate,acuminate,obtuse or somewhatattenuateat the base, ratherevenly serrate,up to about 10 cm. long and 5 cm. broad, delicatelymembranaceous,minutelystrigillose upon both surfaces,the petioleslender,up to about 3 cm. long, commonlyhorizontallyspreadingor somewhatreflexed. Inflorescence terminal,laxly cymose, bearing rather numerous small white shortlypedicellate flowers. Hypanthium subglobose, about 2 mm. broad, minutely and densely the calyx lobes broadlytrigonal,about 1 mm. long; petals broadlyoval hirsutulose, or obovate, somewhat cucullate, 6-8 mm. long. Stamens and staminodia of the

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genus, about as long as the petals. Capsules oblongoid, about 4-6 mm. long and 2-4 mm. broad, persistently hirsutulose. Rather frequentin the moist highland forestsat elevations of 1500-2000 m.
BOCAS DEL

CHIRIQUI:

TORO: Robalo Trail,northern slopes of CerroHorqueta,Allen 49I7, 5002. Bajo Chorro, 65o, Davidson82. Woodson & Schery

3. LOASA Adans.
LOASA

Adans. Fam. 2:501. 1763.

Ortiga Neck. Elemn. 2:400. 1790. H-uidobriaC. Gay, Fl. Chil. 2:43 8. 1846.

Annual or perennialherbs or subshrubs,erect or sprawling, rarely twining, besetin nearlyall parts with scabrous,stinginghairs. Leaves alternateor opposite, variouslyincised and lobed. Inflorescence cymose,terminalor lateral,occasionally reduced to a solitaryflower. Flowers mediocre to large, usually 5-merous,rarely 6- to 7-merous; sepals foliaceous,open in aestivation; petals valvate or imbricate, somewhatclawed; stamensnumerous,in antepetalousclusters,the filaments elongate, united at the base; staminodiaunited into conspicuous more or less saccate antesepalouspetaloid scales; ovary inferior, i-celled, with usually numerousovules borne upon 3-5 parietal placentas, the style usually shortlycolumnar, the stigma obscurelydivided. Fruit a 3- to 5-valved capsule. 75-100 species,extendingfromMexico to Patagonia, but particularly numerous in Peru and Chile. The Loasas sting probablymore painfullythan any nettle,and the quality sometimes is not completelylost fromherbariumspecimens.
a. Leavespinnate or verydeeply pinnatifid above; flowers mediocre, about 2.5 cm. in diameter, thepetalswhite........................................................... 1. L. aa. Leavesonlymoderately lobedand incised. b. Leaves somewhatlonger than broad, somewhatpinnatelylobed;
flowersabout 5 cm. in diameter,the petals pale green ........................... 2. L. bb. Leaves about as long as broad, somewhat palmately lobcd; flowers
RUDIS

GRANDIS SPECIOSA

about 15 cm. in diameter, the petalsbright ............................... 3. L. orange

1. LOASA RUDIS Benth. P1. Hartw. 75. 1839. Loasa rhoeadifoliaSchlecht. in Linnaea 14:382. 1840. Loasa bipinnataDonn. Sm. in Bot. Gaz. 23:7. 1897. Loasa triphyllaJuss.var. 0 rudis (Benth.) Urb. & Gilg, in Nova Acta, Abh. K. Leop.-Carol. Deutsch Akad. Naturfor. 76:239. 1900.

Weak annual herbs up to about 1 m. tall. Leaves alternate,rather sparse, pinnateor very deeplypinnatifid above, the uppermostgreatlyreduced and simple, the lower shortlypetiolate,up to about 15 cm. long and 9 cm. broad, with 3-5 pairs of oblong-ellipticcoarselyserrateleafletsor lobes. Flowers several in laxly monochasial,leafy-bracted terminalcymes; pedicels 5-10 mm. long; calyx lobes 5, broadlyovate-trigonal, obtuse, 6 mm. long and broad; petals 5, broadly spatulate, 1 cm. long, widely spreading, white; staminodiaprofoundlysaccate, about 5 mm. long, stripedwith red or pink. Capsules erect upon the accrescentpedicels,rather narrowly campanuate, up to 2 cm. long and 1 cm. broad, densely hispid, the persistent calyx lobes accrescentand about half as long as the capsule.

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Fig. 9. Loasa rudis

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Southern Mexicoto Panama,wherethe plantis knownlocallyas pringamoza or pringamosca. A frequent herbin pastures and forest clearings at elevations of
1500-2000 m.
CHIRIQUi: upperRio ChiriquiViejo valley,near Cerro Punta, G. White39; Bajo Chorro, BoqueteDistrict, Davidson277, Woodsond Schery 645; CasitaAlta, Volcainde Chiriqui, Woodson, Allend Seibert 936.

This plantprobably is not conspecific with trueL. triphylla of Ecuadorand Peru which,as the name suggests, has palmately dividedleaves. However,our plantmay eventually proveto be conspecific with L. papaverifolia HBK. which extends from Colombia and Venezuela to Chile; in thiscase,L. rudiswill have to be superceded by theearlier name.
2. LOASA GRANDISStandl.in Journ. Wash.Acad. Sci. 17:12. 1927.

to 4 dm. long and 3 dm. broad, graduallyreduced above, the petiole 3-8 cm. long. Flowers several or ratherfew in leafy-bracted, lax terminalcymes; pedicels about 2 cm. long; calyx lobes 5, broadlyovate, about 1 cm. long, sparsely serrate-dentate; petals 5, pale green,cucullate, broadlyobovate, about 2.5 cm. long, 1.5 cm. broad; scales broadly oval-cucullate, 5-6 mm. long, entireor broadly 3-lobed at the tip (exappendiculate?) 3-nerved, the opposite staminodia narrowly acicular, longacuminate, minutely puberulent immediatelyabove the slightly enlarged base, about 1 cm. long; stamensin groups of about 12 opposite the petals, the filaments about 1.5 cm. long, the anthersabout 2 mm. long; roof of the ovary about 1 cm. in diameter,denselyarachnoid-villous;style about 5 mm. long. Capsules broadly turbinate, about 2 cm. long, dehiscingwith 5 broad valves alternatewith the calyx lobes; seeds innumerable, about 1 mm. long or somewhatless. Panama and Costa Rica (?), in clearingsof moist forest,70-1000 m. elev.
CANAL ZONE: forest alongbanksof QuebradaLa Palma and Canion of Rio Chagres, Dodged AllenI7336. COCLi: north Alleni658. rimof El Valle de Anton,

Verylargecoarse herbs up to about4 m. tall. Leavesalternate, rather sparse, long-petiolate, broadly oval and divided into 6-8 broadcoarsely serrate lobes,up

The type of L. grandis comes from Guanacaste, Costa Rica.

I have not been

able to detect the three small conic appendages below the apex of the scales describedby Standley for the Costa Rica plants. 3. LOASA SPECIOSA Donn. Sm. in Bot. Gaz. 23:8. 1897. Large coarse weak herbsup to 2 m. tall, abundantlycovered with long brown stinginghairs. Leaves opposite or subopposite,long-petiolate,the blade about as long as broad, somewhatpalmatelylobed and serratein the mannerof a maple leaf, up to 15 cm. long and broad or somewhatlargerbelow, the petiole up to 12 cm. terminalcymes,the pedicelsup to 6 cm. long. Flowerssolitaryor in few-flowered long; calyx lobes 5, ovate-lanceolate, acuminate,about 3 cm. long; petals 5, bright orange,broadlyoval, 5-6 cm. long, flat or essentiallyso, slightlyspreading;scales the tip unequally 4-lobed, the about 1.8 cm. long, broadly truncate-triangular,

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Fig. 10. Gronaviascandens

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lateral lobessomewhat longer, with2 broadly oval sacs at thebase,thestaminodia 2, about2.5 cm. long,narrowly acicular but greatly bulbose at thebase,minutely and closely aculeolate; stamens in groups of about 14 opposite thepetals,thefilaments glabrous, about2 cm. long,theanthers about2 mm.long; style slender, 1.5 cm. long,minutely hispidulose toward the base,thestigma subcapitate.Capsules narrowly campanulate, about 2 cm. long and 1 cm. broad,bearing innumerable reticulate seedsabout 1 mm.broaduponusually3 parietal placentas. HighlandCosta Rica and western at about 2000Panama,in forest clearings
2500 m. elev.
CHIRIQUI:

3486.

Bajo Chorro, BoqueteDistrict, Davidson445; vicinity of CerroPunta,Allen

With its brightorange flowers15 cm. or more in diameter,this is one of the most showy herbs of the Chiriqui mountains,but is also one of the most painful ortigas.

4. GRONOVIA L.
GRONOVIA

L. Sp. P1. ed. 1. 202. 1753.

Slenderannual twinerscovered with weak uncinate hairs on all parts. Leaves alternate,long-petiolate, the blade very deeply and palmatelydivided and coarsely dentate, broadly ovate-reniform,deeply cordate. Inflorescenceextra-axillary, several times dichasial with the ultimate branchesmonochasial,bearing numerous rathersmall sessileflowers. Calyx lobes 5, subfoliaceous,valvate; petals 5, free, open in aestivation;stamens 5, antesepalous,all fertile;ovary 1-celled, containing a solitarypendulousovule, the styleratherslender,the stigma subcapitate. Fruit a small 5-winged cypsela crowned by the persistent calyx. Two species,extendingfromMexico to Venezuela and Peru.
1. GRONOVIA SCANDENS L. Sp. P1. ed. 1. 202. 1753.

Slenderannual twiners, in all parts. Leaves rathersparse, weaklyuncinate-pilose long-petiolate, the blade broadlyovate-reniform, deeply cordate,up to 8 cm. long and 9 cm. broad, deeply and palmatelydivided into usually five unequal, narrowly acuminate lobes, the middle somewhat longest, entire, the lateral decreasingly smaller and with 1 to several broad dentate incisions,the petiole up to 7-8 cm. long. Inflorescence about equaling the leaves, compound,bearingnumerousrather small sessileflowers. Flowers greenishyellow; sepals oblong-lanceolate,acute, 5-6 mm. long; petals rathernarrowlyspatulate,acuminate,about 3 mm. long; stamens about 4 mm. long, the filaments scarcelylonger than the anthers. Fruits pentagonal-obovoid,about 3-4 mm. long, decurrently 5-wingedfromthe tip, crownedby the persistent, somewhataccrescentsepals. SouthernMexico to Venezuela and Ecuador, in lowland thickets.
PANAMA:

O Schery of Old Panami,Woodson vicinity 948.

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19581

FLORA OF PANAMA

(Begoniaceae)

41

BEGONIACEAE
BY LYMAN B. SMITH
AND

BERNICE G. SCHUBERT L.

1. BEGONIA
BEGONIA

L. Sp. P1. 1056. 1753; A. DC. in DC. Prod. 151:278. 1864.

Lindl. Nat. Syst. ed. 2, 440. 1836. Eupetalum Raf. Fl. Tellur. 2:91. 1836. Trilomisa Gaud. Bot. Voy. Bonite, pl. 32. 1839. Mezierea FalkeaKoenig ex Steud. Nomencl. Bot. ed. 2, 1:194. 1840. Lindl. Veg. Kingd. 319. 1847. Diploclinium T. Moore in Moore & Ayers,Gard. Mag. Bot. 2:153. 1850. Nom. prov. Platyclinum Hassk. in Versl. Acad. Wetensch. Amsterdam4:139. 1855. Sphendanthera K1. in Ind. Sem. Hort. Berol. App. 1. 1855. Nephromischus Huszia K1. in Monatsber.Akad. Berlin 121. 1854. K1. in Monatsber.Akad. Berlin 122. 1854. Saueria BaryaK1. in Monatsber.Akad. Berlin 122. 1854. K1. in Monatsber.Akad. Berlin 122. 1854. Knesebeckia K1. in Monatsber.Akad. Berlin 123. 1854. Gaerdtia K1. in Monatsber.Akad. Berlin 123. 1854. Trendelenburgia EwaldiaK1. in Monatsber.Akad. Berlin 123. 1854. K1. in Monatsber.Akad. Berlin 123. 1854. Gurltia K1. in Monatsber.Akad. Berlin 123. 1854. Scheidweileria LepsiaK1. in Monatsber.Akad. Berlin 123. 1854. K1. in Monatsber.Akad. Berlin 123. 1854. Riessia K1. in Monatsber.Akad. Berlin 123. 1854, as "Mitcherlichia". Mitscherlichia RachiaK1. in Monatsber.Akad. Berlin 124. 1854. K1. in Monatsber.Akad. Berlin 124. 1854. Petermannia K1. in Monatsber.Akad. Berlin 124. 1854. Augustia K1. in Monatsber.Akad. Berlin 125. 1854. Gireoudia K1. in Monatsber.Akad. Berlin 125. 1854. Rossmannia K1. in Monatsber.Akad. Berlin 125. 1854. Magnusia Haagea K1. in Monatsber.Akad. Berlin 125. 1854. Kl. in Monatsber.Akad. Berlin 126. 1854, as "Titelbachia". Tittelbachia K1. in Monatsber.Akad. Berlin 126. 1854. Pritzelia K1. in Monatsber.Akad. Berlin 126. 1854. Wageneria K1. in Monatsber.Akad. Berlin 126. 1854. Doratometra Kl. in Monatsber.Akad. Berlin 127. 1854. Pilderia DonaldiaK1. in Monatsber.Akad. Berlin 127. 1854. K1. in Monatsber.Akad. Berlin 127. 1854. Moschkowitzia K1. in Monatsber.Akad. Berlin 127. 1854, as "Stiradotheca". Stibadotheca K1. in Monatsber.Akad. Berlin 127. 1854. Casparya K1. in Monatsber.Akad. Berlin 127. 1854, as "Isopteris". Isopteryx SasseaK1. in Monatsber.Akad. Berlin 128. 1854. K1. in Abh. Akad. Berlin 1854:54. 1855. Reichenheimia K1. in Abh. Akad. Berlin 1854:64. 1855. Steineria K1. in Abh. Akad. Berlin 1854:82. 1855. Trachelanthus K1. in Abh. Akad. Berlin 1854:100. 1855. Cyathocnemis Kl. & Oerst. ex Kl. in Abh. Akad. Berlin 1854:119. 1855. Weilbachia LaucheaK1. in Abh. Akad. Berlin 1854:121. 1855. K1. in Abh. Akad. Berlin 1854:123. 1855. Platycentrum K1. in Abh. Akad. Berlin 1854: 134. 1855. Putzeysia Muell. in Walp. Ann. Bot. 4:909. 1857. Trachelocarpus K1.ex A. DC. in DC. Prod. 151:278. 1864. Cladomischus Irma Bout. ex A. DC. in DC. Prod. 151:406. 1864.

(41)

I Vol. -15

42

ANNALS

OF THE

MISSOURI

BOTANICAL

GARDEN

Begoniella Oliver, in Trans. Linn. Soc. 28:513. 1873. SemibegoniellaC. DC. in Bull. Herb. Boiss. II. 8:327. 1908.

Annual or perennialherbs, shrubs or rarely small trees. Stern elongate and succulent or woody or a cylindricalrhizome or sometimesreduced to a tuberous rhizome, sometimesclimbing. Leaves usually alternate or rarely subverticillate, entire,serrate, lobed or digitatelycompound,usually asymmetric, sometimes peltate, stipules2, free,persistent or deciduous. Peduncles mostlyaxillary. Inflorescence usually cymose,sometimes 1-flowered or racemose. Cymes regularor strongly onesided, usually dichotomous,unisexual or bisexual. Flowers monoecious. Tepals freeor rarelyconnate, usually 4 staminatein two pairs and 5 pistillate. Stamens indefinite, insertedon the receptacle,filamentsfree or united. Styles usually 3, free or connate, usually bifid,ovary inferior,placentae axile, simple or divided. Fruit capsular,usually bearing 3 unequal wings.
a. Plantsacaulescent or subacaulescent with the leavesclustered near the apexof therhizome, or withslender creeping stolons if theinternodes are elongate. b. Leavespeltate. c. Cymesstrongly unilateral;capsule-wings slightly unequal; leaves 2-12.5 cm. long 1. B. CONCHIFOLIA ................ cc. Cymesregular; capsule-wings veryunequal;leaves17-40 cm. long. 2. B. NELUMBIIFOLIA bb. Leavesbasifixed. c. Cymes strongly unilateral. d. Stipules ecarinate,coarsely reticulate-nerved; staminateand as highas wide ........... 3. B. PLEBEJA pistillate tepals2; largest capsule-wing dd. Stipules alate-carinate withfineinconspicuous nerves;staminate widerthanhigh ............................... 4. B. GARAGARANA tepals4; largest capsule-wing cc. Cymesregular. of the cyme3 or 4 at each node; inflorescence d. Branches manyflowered . ........ ...........................,.,.,,., 5. B. QUATERNATrA of thecyme2 at each node. dd. Branches e. Plantsor at leastthepetioles densely vestite. f. Indument of linear of thepetioles consisting trichonmes. thestaminate g. Cymefew-flowered; tepals pubescent, 4, the withelongate stolons often internodes pistillate 3; slender 6. B. CARLETONII ........ present;capsule reflexed gg. Cyme many-flowered; tepals glabrous," cach; capsule 7. B. VILLIPETIOLA erect . ,. .. tepals2. g. Cymenot at all umbelliform; capsuleerect h. Largestcapsule-wing narrower thanhigh; leavesentire or slightly ciliate ................................... 8. B. serrate, coarsely hh. Largestcapsule-wing much wider than high; anthers 9. B. leavesciliate-denticulate..... than the filaments; shorter gg. Cyme umbelliformwith greatly reduced terminal branches; capsule decurved; antherslonger than the 10. B. .... filaments ee. Plantscompletely glabrous. f. Pistillatetepals 2 or 3; pistillatebracteolesapparently lacking. staminate g. Tepalsand upperbracts glandular; tepalsnearly 11. B. on a column;pistillate tepals2............ equal; stamens staminate gg. Tepals and bractseglandular; tepalsvery unequal; stamens free,antherslongerthan the filaments; 12. B. pistillate tepals3. ff. Pistillatetepals 4; pistillatebracteolesorbicular,entire; staminatetepals very unequal; anthersshorterthan the B. . .13. filaments internodes. stemand distinct aa. Plantswithan erector ascending
ff. Indument of thc petioles consisting of laccrate scales;

STRIGILLOSA STIGMOSA

VESTITA

BREVICYMA

MUCRONISTIPULA

DAVIDSONIAE

(42)

19581

FLORA

OF PANAMA

(Begoniaceae)
GLABRA

43

. 14. B. ovate b. Leaves nearlysymmetric, ..................................... asymmetric. bb. Leavesdistinctly c. Softlow annuals;tepalsnotover3.5 mm.long. d. Capsule-wingssubequal, lunate; stamens free; leaves soon

on a shortcolumn;leaves veryunequal; stamens dd. Capsule-wings 16. B. FILIPES above ....................................... appressed-pilose cc. Coarseherbsor shrubs;tepalslarger. oblique. or strongly transverse d. Leavespalminerved, tepals tepals5; staminate pistillate fimbriate; e. Bractspersistent, 17. B. FISCHERI 4................................................... tepals2 or 3. pistillate entire; ee. Bractsdeciduous, equal or short;capsule-wings peduncles few-flowered; f. Cymes tepals3. subequal;pistillate of the leaves; at least on the underside g. Plant pubescent B. OAXACANA ..18. if at all beaked capsulescarcely at the apex exceptfora ringof trichomes gg. Plant glabrous B. UDISILVESTRIS beaked ..19. of the petiole;capsuleslenderly peduncleselongate; capsule-wings ff. Cymes many-flowered; 2. tepals unequal;pistillate very branchesmuch reducedmakingthe infloresg. Secondary enclosedby younginflorescence cence appearumbellate, .20. B. INVOLUCRATA two largebracts;leavespubescent and distinctly normal;inflorescence branches gg. Secondary 21. B. MULTINERVIA leavesglabrous ......................................... cymose; regularly oblique. or slightly straight penninerved, dd. Leaves fromthe upperpart of e. Ovary and capsuleequally 3-horned few-flowered. theangles;cymes f. Leaves dimidiateat base; antherslinear; pistillatetepals 22. B. URTICAE .............. 5; stylesmultifid pistillate subglobose; ff. Leavesdeeplycordateat base; anthers 23. B. HEYDEI ... bifid tepals3; styles 3 wingswhichare attachedthe full lengthof ee. Ovary bearing unequal. distinctly and areusually theangles acusubequal; leaves falcate, oblanceolate, f. Capsule-wings 24. B. TONDUZIT .............................................. minate unequal. strongly ff. Capsule-wings cymesglobose,many-flowered; g. Leaves subulate-dentate; ............................................. 25. B. OPULIFLORA staminate tepals4, subequal teeth. acutenotsubulate gg. Leaveswithflat staminate tepals4; h. Pistillate tepals5; stylesmultifid; 26. B. GUADUENSIS ............................. very unequal. ........... tepals2 (or if 3; staminate tepals2 or rarely hh. Pistillate bifid). merely 3 or 4 thenthestyles i. Internodevery short; plants scandent; staminate 27. B. ESTRELLENSIS ...................................... .. 4 tepals elongate;plantsusuallyerect; staminate ii. Internodes 3 or 4). tepals2 (rarely of the wings; j. Capsule 10-12 mm. highexclusive 28. B. SEEMANNIANA ................................... floriferous cymesmoderately jj. Capsule 6-10 mm. high exclusiveof the wings; floriferous. densely cymes k. Leaves 5-7.5 (-11) cm. long, 2.5-3.7 (-4.7) .............................29. B. CARPINIFOLIA cm. wide, serrate-denticulate kk. Leaves8.5-15 cm. long,4-7.5 cm. wide,some30. B. CONVALLARIODORA ................................................... whatdenticulate 1. BEGONIA CONCHIFOLIA A. Dietr. in Allg. Gartenzeit. 19:258.

......... glabrous

15. B. SEMIOVATA

1851.

Meddell.1852:9. 1853. Liebm.in Kjoeb.Vidensk. scutellata Begonia Akad.Berlin125. 1854. KI. in Monatsber. conchifolia Gireoudia 8. 1864, nomen in synon. Hort.ex A. DC. in DC. Prod.151:33 warscewicziana Giroudia 1929. Bot. Gard.27:322. 1940,nonIrmscher, in Ann.Missouri Standl. pumilio Begonia Herbaceous. Rhizome oblique, 5 mm. thick, internodesvery short. Leaves

(43)

[Vol. 45

44

ANNALS

OF THE

MISSOURI

BOTANICAL

GARDEN

ia Econchaefol
111 W

fnelunbiifokia
[\ _

andB. nelumbiifolia Pig.11. Begonia conchifolia


peltate,broadlyovate, acuminate,7-8-nerved, entireor dentate at the ends of the nerves, 2-12. 5 cm. long, nearly or quite glabrous above, rufous-pilosebeneath especiallyon the nerves,fleshy, petioles 3-10 cm. long, rufous-pilose, stipulespersistent, ovate, setose-acuminate,brown., glabrous or sparsely pilose. Peduncles much exceedingthe leaves,to 15 cm. long, rufous-pilose. Cymes bisexual,strongly unilateral and often much longer than wide, few- to many-flowered. Bracts deciduous, small, lanceolate to obovate, entire,setose-ciliate. Pedicels 4-12 mm. 4-5 mm. long, entire,pink. Stamens few, long. Staminatetepals 2, suborbicular, free. Pistillate bracteoles elliptic, nearly equaling the ovary. Pistillate tepals 2, like the staminate. Ovary 3-celled, placentae bilamellate,ovuliferous-throughout, styles connate at base, stigmas lunate-capitate. Capsule erect, subglobose,wings slightlyunequal, sublunate,angled near apex. Honduras, Costa Rica, Panama.
COCLE: of La Mesa,hillsnorth of El Vtalle de Anton,alt. 1000 m., epiphytic, vicinity Allen 2297; vicinity of El Valle de Anto6n, Allen 292.5. PANAMA': on boulders, Cerro Allen2089. Campana,

2. BEGONIA

NELUMBII1FOLIA

Schlecht. & Cham.in Linnaea5:604. 1830. (44)

1958]

FLORA 5: misc. 355. 1844.

OF PANAMA

(Begoniaceae)

45

Begoniaderycxiana Lem. in Hortic. Univ. Ki. in Monatsber. Gireoudianelumbiifolia Begonia caudilimbaC. DC. in Smithson.
Misc. Coll. 69: no. 12:9. 1919. Akad. Berlin 125. 1854.

B.plebeja

Herbaceous. Rhizome repent,short, 15 mm. thick, setose, internodes extremelyshort. Leaves peltate,obliquely very broadly ovate or subelliptic,7-9nerved, evenly rounded except for the ! proabruptlyacuminateapex or slightly duced at the ends of the nerves, 17-40 cm. long, 10-28 cm. wide, remotely denticulate, soon glabrous, the margin ciliate and sometimes purple, thin, petioles 15-45 cm. long, to 8 mm. thick, becoming glasparsely rufous-hirsute, lanceolate,over brous,stipulespersistent, 2 cm. long, entire, rather firm,pilose. Peduncle usuallyexceedingthe leaves, to 66 cm. long, 6 mm. thick,soon glabrous. Cyme regular, much branched, diffuse, 2-5 dm. broad. Bracts deciduous,ovate, obtuse. Pedicelsslender,9-22 mm. long. Staminate tepals 2, suborbicular, 6-8 mm. long, white. Stamensfew, anthers narrowly obovate, equaling or longer than the filaments. Pistillatebracteoles lacking. Pistillatetepals 2, like the staminate. Ovary 3-celled,placentae bifid, ovuliferous stylesconnateat throughout, base, distinctly divided, the stigmatic surface linear,spiral,continuous. Capsule erect or somewhat nutant, broadly ovoid, 6-10 mm. long, wings very unequal, the largest ovate, obtuse, to 15 mm. wide, seeds ellipsoid,blunt. SouthernMexico to Colombia.
CANAL ZONE: on rocky bank, forest nearsea level, alongtheRio Indiode Gatuin, Maxon4866. DARIEN: Chepigana District,

Cana-Cuasi Trail (Camp 1), alt. 240 m., Terry I623.

Fig. 12.

Begoniaplebeja

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46
3.
BEGONIA

[Vol. 45 ANNALS
PLEBEJA

OF THE

MISSOURI

BOTANICAL

GARDEN

Liebm. in Kjoeb. Vidensk. Meddel. 1852:8. 1853.

Begonia glandulosa sensu J. D. Smith,Enum. PI. Guat. etc. 4:182. 1895; 5:34. 1899. Begonia uvana C. DC. in Smithson.Misc. Coll. 69: no. 12:4. 1919.

Herbaceous, fuscous-villouswhen young. Rhizome usually repent, to 8 cm. long at least, 8-10 mm. thick,internodes veryshort. Leaves oblique, broadlyovate or elliptic, acute, palmately 7-8-nerved, denticulate, angulate-dentateor almost evenly rounded,cordate, 7-20 cm. long, thin, soon glabrous,petioles erect, 6-28 cm. long, usuallysoon glabrous,stipulespersistent, imbricate, lanceolate,acuminatesetiferous, 1-2 cm. long, entire,heavily and closely reticulate-nerved. Peduncles exceeding the leaves, up to 22 cm. long. Cymes bisexual, stronglyunilateraland usuallymuch longerthan wide, few- to many-flowered.Bracts deciduous,obovate, ciliate. Staminate tepals 2, broadly ovate, 5-9 mm. long, white or pale pink. Stamens about 15, filamentsshort,anthersoblong. Pistillate flowersbracteolate. Pistillatetepals 2, like the staminate. Ovary 3-celled, placentae bifid,ovuliferous on all sides,stylesshort-connate, stigmaslunate-capitate. Capsule suberect,oblongellipsoid,12 mm. long, wings unequal, the largestsubtriangular and as high as wide. SouthernMexico and Central America.
between Madden Dam and Saddle 11 near Alajuela, alt. 90-100 m., Dodge, Steyermark& Allen I6503; WesterlyArm of Quebrada Salamanca, alt. 70 m., Dodge,Steyermark & Allen I7008; forest along banks of Quebrada Fea, Q. Pura and Canion of Rio Chagres, alt. 70-100 m., Dodge & Allen I7408; forestalong telephone trail between the Rio Indio Hydrographic Station and the Natural Bridge of Rio Puente, Dodge & Allen I7482. COCLE: vicinity of El Valle, alt. 800-1000 m., Allen 94. PANAMA: Old Panama, Rose I8508; Taboga Island,Killip 3I67; Standley27966, Allen I264; Rio Tapia, Standley 26172, 28288; Juan Diaz, Standley 30583; along road between Panama and Chepo, Dodge, Hunter,Steyermarl & Allen i6683; Rio La Maestra, alt. 0-25 m., Allen 58; San Jose Island, Johnston 83, 207, 322, 386, 436. VERAGUAS: Isla de Uva, Contrerasgroup, Pittier
295I3; 5IO9.
CANAL ZONE:

Las CascadasPlantation, near Summit, Standley 25683, 25759, 2578i,

4.

BEGONIA

GARAGARANA

C. DC. in Smithson.Misc. Coll. 69: no. 12:2. 1919.

Herbaceous, completelyglabrous. Rhizome repent,only a small terminalsection known, internodesvery short. Leaves oblique or transverse,palminerved, strongly asymmetric,broadly ovate or elliptic, abruptly acuminate, shallowly cordate at base, to 24 cm. long, entire,petioles erect, 25 cm. long, stipulesdecidwith fine membranaceous uous, lanceolate,acute, 2 cm. long, entire,alate-carinate, inconspicuous nerves. Peduncle much exceeding the leaves, to 75 cm. long. Cymes dichotomous, strongly asymmetric,lax, many-flowered,15 cm. long. Bracts deciduous, unknown. Pedicels 24 mm. long. Staminate tepals 4, entire, glandular-punctate,the outer broadly obovate, 7-9 mm. long, the inner very narrow. Stamens few on a low torus, filamentsvery short, anthers narrowly obovoid, 2.5 mm. long, connective scarcely if at all produced. Pistillate flowers unknown. Capsule ovoid or ellipsoid, 8-10 mm. long, wings very unequal, the seeds largest26 mm. wide, stylesdeciduous,placentae bifid,ovuliferous throughout, stalked,ellipsoid. Endemic.

(46)

1958]

FLORA

OF PANAMA

(Begoniaceae)

47

B. aragarana

B.quaterna-t

Fig. 13. Begoniagaragarana and B. quaternata


DARIEN: Cerrode Garagarai, Sambu' Basin,southern Darien, alt. 500-974 m., Pittier VERAGUAS: forested slopesof CerroTute, vicinity of SantaFe, alt. 750 m., Allen

5672.
4375.

5.

BEGONIA

QUATERNATA

figI,,P-s.

Smith & Schubert,in Journ.Wash. Acad. Sci. 40:244.

1950.

Stemlesswith the leaves clusterednear the apex of the erect densely rooting rhizome,internodesvery short; leaves transverse, very broadly elliptic, obliquely acuminate and also cuspidate opposite the principal nerves, deeply and narrowly cordate at base, 17 cm. long, 14 cm. wide, nearly glabrous above, below densely piloseon the nervesand sparsely elsewhere, petiolesslender,15-18 cm. long, densely vestite with reflexedlacerate fuscous scales, stipules early deciduous, unknown. Peduncles 4 dm. long, much exceeding the leaves, slender,glabrous. Inflorescence regularly cymose, 3-4-radiate, many-flowered, 13 cm. broad, glabrous. Bracts quickly deciduous,small,elliptic,entire. Flowerswhite (! Allen) but yellowwhen dry. Staminate tepals 2, suborbicular,5 mm. long, entire. Stamens insertedon a short column, anthers ellipsoid, slightly shorterthan the filaments. Pistillate bracteolesapparentlylacking. Pistillate tepals 2, to 2.5 mm. long. Styles bifid, ovary 3-winged,wings equal, broadlyovate.
Endemic. 1800-2000 m., January6-14, 1939, Allen I336.
CHIRIQUI:

of "New Switzerland", centralvalleyof Rio ChiriquiViejo, alt. vicinity

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[Vol. 45

48

ANNALS

OF

THE

MISSOURI

BOTANICAL

GARDEN

and B. villipetiola Fig. 14. Begoniacarletonii

of that of B. stigmosa. The generalhabit of Begonia quaternata is reminiscent However, the flowersof B. quaternata are much smaller,and its cyme has four rays,a characterthat is veryrareif not unique in North AmericanBegonia. primary 6.
BEGONIA CARLETONII

Standl. in Journ.Wash. Acad. Sci. 17:314. 1927.

Herbaceous. Rhizome slenderwith short internodesbut also producing very slenderstolons, with internodes5-8 cm. long, setose-pilose. Leaves asymmetric, oblique, broadlyovate, abruptlyacuminate,cordate at straightor oftenerstrongly on ciliate, sparselysetose-pilose base, 6-16 cm. long, thin, finelysinuate-dentate, stipules both sides, paler beneath, petioles slender,5-16 cm. long, reflexed-setose, narrowlylanceolate, 9 mm. long, entire,thin, pilose. Peduncles shorteror longer pilose. Bracts subpersistent, than the leaves, slender,pilose. Cymes few-flowered, ovate or elliptic, obtuse, 3-5 mm. long, entire. Pedicels to 18 mm. long. Staminate tepals 4, entire,white, the outer broadly ovate, obtuse, 6-12 mm. long, sparselypilose, the inner narrowlyoblanceolate, shorter,glabrous. Stamens on a conlow torus,numerous,anthersobovate, somewhat shorterthan the filaments, wanting. Pistillatetepals 3, nective not produced. Pistillatebracteolesapparently like the staminate. Ovary 3-celled,pilose,stylesslightlyconnate at base, otherwise ellipsoid,7 mm. long, wings the stigmaticsurfacelunate. Capsule sharplyreflexed, fromthe lowerhalf of the capsule,oblong,obtuse, veryunequal, the largestreflexed 5 mm. wide, the othersnarrowlylunate. Endemic.

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1958]

FLORA

OF PANAMA

(Begoniaceae)

49

BOCAS DEL TORO: Changuinola Valley,Dunlap 206; localityuncertain, Carleton 206. vicinity of Chiriqui Lagoon, von Wedel 227I; Fish CHIRIQUl: Fish Creek Mountains,

Creek, von Wedel 2222. 7.

BEGONIA VILLIPETIOLA C. DC. in Smithson.Misc. Coll. 69: no. 12:5.

1919.

Herbaceous. Rhizome shortwith extremely short internodes, rooting densely. Leaves strongly asymmetric,transverselyovate, abruptly acuminate and often with a second cusp or lobe more nearlyopposite the petiole,cordate at base, 15-21 cm. long, ciliate-denticulate, thin, sparselypilose above, more densely so beneath especiallyon the nerves,petiolesslender,15-25 cm. long, denselyvillous with linear trichomes, stipulesdeciduous,triangular-ovate, acuminate, 15-20 mm. long, pilose. Peduncle 18-37 cm. long, exceeding the leaves when in fruit,becoming glabrous. Cymes bisexual, regularlydichotomous,many-flowered, subdense,glabrous except the bracts. for Bracts quickly deciduous,the lowest large, broadlyelliptic,entire,
pilose. entire. Flowers pink or white. Staminate tepals 2, suborbicular, 10 mm. long, Stamens free, numerous, anthers obovate, shorter than the filaments, con-

nective not produced. Bracteoles lacking. Pistillate tepals 2, like the staminate but smaller. Ovary 3-celled, placentae bilamellate,ovuliferousthroughout,styles connate at base, stigmatic tissue linear, spiral, continuous. Capsule erect, subglobose, 4-5 mm. long, wings distinctlyunequal, broadly ovate, the largest to 11 mm. wide. Endemic.
CANAL ZONE:
COCLE:

Allen2I3, 238, II84, I664, 2906, 42i6.

Mojinga Swamp near mouthof Rio Chagres,alt. 1 m., Allen 864. Bismark abovePenonome, Williams 309; vicinity of El Valle, alt. 600-1000 m.,
1851.

8. BEGONIA STRIGILLOSAA. Dietr. in Allg. Gartenzeit. 19:330.

Gireoudia strigillosa Kl. in Monatsber.Akad. Berlin 125. 18 54. Begonia daedalia Lem. in Ill. Hortic. 7: misc. 54. 1860. 8: pI. 269. 1861. Begonia barbana C. DC. in Bull. Soc. Bot. Belg. 35: pt. 1:261. 1896.

Herbaceous. Rhizome varyingfromshortand compact with very shortinternodes and imbricate stipules to slender and branching with distinct internodes. Leaves oblique, broadlyovate, acuminate,7-17 cm. long, entireor slightlyserrate, rounded to angulate-dentate,coarsely ciliate, glabrous above, sparselylong-pilose below especiallyon the nerves,often spottedor mottled,petioles to 18 cm. long, often spotted,bearingnumerousreflexed fimbriate scales, stipuleslanceolate with a setiferous apex and oftenfimbriate keel, persistent. Peduncle usually exceedingthe leaves, sparselypilose,often purple-spotted. Cyme 2-4-branched,subsymmetrical, mostlyglabrous,diffuse.Bracts deciduous,elliptic,entire,mucronulate. Staminate tepals 2, suborbicularto obovate, 6-9 mm. long. Stamensfew, the anthersoblong, obtuse. Pistillatebracteoleslacking. Pistillatetepals 2, like the staminate. Ovary connate at base, 3-celled,placentae bifid,ovuliferousthroughout,stylespersistent, locules to somewhatbifid,stigmaslinear. Capsule suberect,ovoid or elliptic-ovoid, or subobtuse to acuminate. 10 mm. long, the two largerwings subtriangular Guatemala, Costa Rica.

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B. .

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[Vol. 45 MISSOURI BOTANICAL GARDEN

HX q :P42Bstimo S
7,~~~~~~~~~~~~~.
C

Fig. 15. Begonia strigillosa and B. stigmosa

Indefinite. Seed no. 7 fromMr. Clayton Kelly of Long Beach, California. Mr. Rudolf Ziesenhennewho sent us the specimensays that it is called water begonia for it grows only near streamsin Panama. He considersit a distinct species but so far we are unable to separateit satisfactorily from B. strigillosa. B. strigillosa has been collected frequentlyin Costa Rica so that its occurrence in Panama is to be expected. 9. BEGONIA STIGMOSALindl. in Bot. Reg. 31: misc. 32. 1845. sensu nonLiebm. 1853. Begonia squarrosa Seemann, Bot.Herald 128. 1853, nomen, Gireoudia Ki. in Monatsber. Akad.Berlin125. 1854. stigmosa Herbaceous. Rhizome repent, to 12 cm. long, 1 cm. in diameter,coarsely lepidote, internodesvery short and covered by the stipules. Leaves oblique and

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19581

FLORA

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51

stronglyasymmetric, very broadly ovate or suborbicular,abruptly acuminate and usually with a second cusp more nearlyoppositethe petiole,cordate at base, 15-30 cm. long, scarcely if at all lobed, ciliate-denticulate, thin, glabrous above or with a few small white trichomes, the nervesbeneathbearingtrichomes like thoseon the petiole but smaller,petioleserect, to 38 cm. long, 4-10 mm. in diameter,covered with spreadingto reflexed narrowlytriangularlacerate pink scales 3-5 mm. long, stipules tardily deciduous, lanceolate, pilose, membranaceous,15-20 mm. long. Peduncle 2-5 dm. long, 7 mmn. in diameter, soon glabrous. Cymes bisexual,nearly regular, few- to many-flowered, 7-22 cm. long, lax, glabrous. Bracts quickly deciduous, oblong, acute, fimbriate. Pedicels slender,7-23 mm. long. Flowers white or pink. Staminatetepals 2, suborbicular, cordate at base, 10-15 mm. long. Stamens free, very numerous, filamentsmuch longer than the broadly oblong anthers,connectiveapiculate-produced. Bracteoleslacking. Pistillatetepals 2 like the staminateor rarelya smallerone inside the others. Ovary 3-celled, placentae bifid,ovuliferousthroughout,styles bifid,the stigmatic tissue linear, spiral, continuous. Capsule erect, broadly ovoid, 6-8 mm. long, wings very unequal, the largest oblong or subdolabriform, 10-15 mm. wide. the othersnarrowlymarginiform. SouthernMexico, Central America and Colombia. aroundEl Boquete,alt. 1000-1300 Dodge d Allen 17368. CHIRIQUI: forests Hato del Joboand CerroVaca, eastern m., Pittier 2896; Maxon4947; between Chiriqui, alt. 700-1000 m., Pittier 54II.
70-80 m.,
CANAL ZONE:

forest along banks of Quebrada La Palma and Caflon of Rio Chagres,

10. BEGONIA VESTITA C. DC. in Bull. Herb. Boiss. II. 8:315.

1908.

Herbaceous. Rhizome repent,short,internodes very short. Leaves transverse, ellipticor ovate, abruptlyacuminate,cordate at base, 7-18 cm. long, palminerved, entire,glabrous above, brown-pilosebeneath especiallyon the nerves,petioles 1525 cm. long, subdenselypilose with fimbriatescarious scales, stipules deciduous, 10-15 mm. long. Peduncle not much exceedingthe leaves ovate, membranaceous, until in fruit,to 35 cm. long, bearingscales like those of the petioles. Cymes bisexual, few-flowered, subumbellate with the ultimate branches greatly reduced. Bracts deciduous. Pedicels 5-15 mm. long. Flowers white or pink, sometimes red-punctate. Staminate tepals 2, suborbicular, 5-7 mm. long, rather fleshy, sparsely pilose. Stamens free, few, anthers oblong, longer than the filaments. Pistillatebracteoleslacking. Pistillatetepals 2, like the staminate. Ovary 3-celled, placentae bilamellate,ovuliferousthroughout,styles connate at base, bifid, stigmatic tissue spiral. Capsule sharply deflexed,ellipsoid, 10-12 mm. long, pilose, wings veryunequal, the largestsubreflexed, broadlyovate, obtuse,to 10 mm. wide, the othersnarrowlylunate or marginiform. Costa Rica, Panama.
north of El Valle de Ant6n, alt. 1000 m., Allen 2906.

alt. 1800 m., Davidson7I; epiCHIRIQUI: rain forest, Bajo Chorro, BoqueteDistrict, of Bajo Chorro,alt. 1900 m., Woodson d Schery62I. COCLEf: region phyte, vicinity

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[Vol. 45

52

ANNALS

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B-vestit?

5.brevicyma

B.davidsoniae

Fig. 16. Begonia

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19 58 ]

FLORA

OF PANAMA

(Begoniaceae)

53

11.

BEGONIA

BREVICYMA

C. DC. in Smithson.Misc. Coll. 69: no. 12:3. 1919.

Herbaceous, glabrous. Rhizome 6-7 mm. in diameter,internodesvery short. Leaves oblique, palminerved,strongly asymmetric,broadly subelliptic, abruptly acuminate with smaller cusps at the end of the principal nerves,cordate at base, broadly ovate, acu10-14 cm. long, petioles 16-24 cm. long, stipules persistent, minate,laceratenear apex, 15 mm. long. Peduncles to 26 cm. long. Cyme regular, few-flowered. Bracts tardily deciduous, the lower ample and covering the the upper much shorterthan the pedicels, glandular. Pedicels to young flowers, 25 mm. long. Staminatetepals 4, subequal, obovate, 17 mm. long, entire,pinkish white, glandular,the outer somewhatbroader. Stamens on a column, numerous, connectivenot produced. Bractethan the filaments, anthers obovate,much shorter oles wanting. Pistillate tepals 2, broadly ovate, glandular. Wings of the ovary very unequal. Endemic. vicinity of Bajo Chorro,alt. 1900 m., Woodsond Schery620; epiphyte, CHIRIQUI: slopeof Cerrode la Horqueta,Chiriqui, humidforest aroundLos SiguasCamp,southern alt. 1700 m., Maxon54I7. 12.
BEGONIA MUCRONISTIPULA

C. DC. in Smithson. Misc. Coll. 69: no. 12:3.

1919. internodes very Herbaceous, completelyglabrous. Rhizome 6 mm. in diameter, asymmetric, broadlyelliptic,apiculate strongly short. Leaves oblique, palminerved, entire,cordateat base, 10 cm. long, petioles oppositethe principalnerves,otherwise erect, 16 cm. long, stipules persistent,broadly ovate, mucronate, nearly 2 cm. long (! C. DC.), entire,thin. Peduncle much exceeding the leaves, 37 cm. long. Cyme regular, few-flowered. Bracts deciduous, unknown. Pedicels to 15 mm. long. Staminatetepals 4, entire,the outer broadlyobovate, 20 mm. long, one red and one white, the inner subspatulate,15 mm. long, white. Stamens free,rather numerous,anthersoblong-obovate,slightly longer than the filaments. Pistillate bracteoleslacking. Pistillatetepals 3, the outer suborbicular,8 mm. long (possibly immature), the inner one broadlyovate, much smaller. Ovary 3-celled,placentae styles3, connate at base, bifid,stigmatictissue bilamellate,ovuliferousthroughout, spiral. Capsule-wingsvery unequal, the largest narrowlytriangular,much wider than high, the othersnarrowlylunate. Endemic. slopeof Cerrode the Rio Ladrilloand Los SiguasCamp,southern CHIRIQUI: between la Horqueta, alt. 1200-1700m.,Pittier 3I72. 13.
BEGONIA DAVIDSONIAE Standl. ex Smith & Schubert,in Journ. Wash. Acad. Sci. 40:242. fig.I, I-o. 1950.

Stemlesswith the leaves clusterednear the apex of the short prostratedensely rooting rhizome, glabrous, internodesvery short; leaves oblique, broadly ovate, abruptlyacuminate and also cuspidate at the ends of the principal nerves,deeply and narrowlycordate at base, 10 cm. long, 7 cm. wide, petioles slender, 17 cm.

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EVol. 45

long, stipulesdeciduous,triangular-ovate, acuminate, 12 mm. long, lacerate toward apex, thin. Peduncles slender, to 26 cm. long, exceedingthe leaves. Cymes regular, few-flowered, dense. Bracts quickly deciduous. Flowers white or pink (! Davidson). Staminate tepals 4, entire,the outer suborbicular,12 mm. long, the inner elliptic, 10 mm. long. Stamensnumerous,anthersobovate, much shorterthan the filaments, connectivenot produced. Pistillate bracteolesorbicular,entire. Pistillate tepals 4, like the staminatebut smaller. Stigmatic tissue spiral,wings of the ovary very unequal, the largest oblong, obtuse, much wider than high, the others narrowlymarginiform.
Endemic.
CHIRIQUI:

Davidson 38.

rain forest,Bajo Chorro, Boquete District, alt. 1800 m., January 5, 1938,

This species appears to be related to Begonia brevicymaand B. inucronistipula as indicatedin the foregoing key, however,it has not been possibleto ascertainthe detailedstructureof the stylesand ovary without dangerof seriously damaging the type,so theremay be greaterdifferences than appear.
14.
BEGONIA GLABRA

Aubl. PI. Guian. 2:916. PI. 349. 1775.

Begonia lucida Otto & Dietr. in Allg. Gartenz. 16:162. 1848. Begonia moritzianaKunth & Bouche in Ind. Sem. Hort. Berol. 16. 1848. Begonia physalifolia Liebm. in Kjoeb. Vidensk. Meddell. 1852:19. 1853. Wageneria lucida Kl. in Monatsber.Akad. Berlin 126. 1854.

Begonia HBK. Nov. Gen.& Sp. 7:180, pl. 641. 1825. elliptica

Begonia scandensSw. Prodr. 86. 1788.

moritziana Ki. in Monatsber. Akad.Berlin126. 1854,nomen. Wageneria

Begonia locellata A. DC. in Ann. Sci. Nat. IV. 11:137. 1859.

Wageneria glabraKl. in Monatsber. Akad.Berlin126. 1854. Ki. in Abh. Akad.Berlin1854:113. 1855. Wageneria deflexa Wageneria Montana Ki. in Abh. Akad.Berlin 1854:115. 1855.

Succulent herb to 9 mn. high. Stem scandent, rooting at the nodes, glabrous. Leaves nearlysymmetrical, broadlyovate, 4-15 cm. long, short-acuminate, rounded or barely cordate at base, sparsely serrate and ciliate to entire, often undulate, glabrous,petioles1-8 cm. long, stipulespersistent, ovate-oblong,mucronate, entire, 10-24 mm. long, membranaceous, red-brown. Peduncles axillary,6-20 cm. long. diffuse. Bracts persistent, Cymes many-flowered, minute. Pedicels slender,6-16 mm. long. Staminate tepals 4, white, the outer broadly obovate, 3-8 mm. long, the innernarrowlyelliptic. Stamens free,few, anthersoblong. Pistillatetepals 5, 4-6 mm. long. Styles 3, 2-parted,completelycovered by stigmaticpapillae, placentae simple. Capsule 6-9 mm. long, largest wing oblong to triangular,10-14 mm. wide, the other two marginiform, very narrow. SouthernMexico and the West Indies to Guiana, Bolivia and Peru.
BOCAS DEL TORO: FishCreek, of Chiriqui von Wedel2204; FishCreek vicinity Lagoon, von Wedel2278, 2345; FarmSix,Changuinola Mountains, Valley,Dunlap 500. CHIRIQUI:

Ilanos on slopes of Volcain de Chiriqui Viejo and along Rio Chiriqui Viejo, alt. 1200 m., Allen 982. COCLE: vicinity of El Valle de Anton, alt. 600-1000 m., Allen 221, II94, PANAMA: forestsalong Rio Boquer6n above Peluca Hydrographic Station, alt. 90 i655. m., Hunter d Allen 655. VERAGUAS: forestedslopes of Cerro Tut6, vicinity of Santa F6, alt. 900 m., Allen 4357.

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1 958:1

FLORA

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(Begoniaceae)

55

B. cIbra

Q B5e miovatat

B. fizzes

er~?1

f iipe
Fig. 17. Begonia (Lower right: B. fischeri)

i
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15.

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[Vol. 415

BEGONIA SEMIOVATA Liebm. in Kjoeb. Vidensk. Meddel. 1852:22.

1853.

Begonia spruceana A. DC. in Ann. Sci. Nat. IV. 1:142. 1859. Begonia flexuosaA. DC. in Ann. Sci. Nat. IV. 11:142. 1859. Begonia guyanensisA. DC. in Ann. Sci. Nat. IV. 11:142. 1859. Hoff-mannella rosea Ki. in A. DC. in DC. Prod. 151:299. 1864. Nomen. Begonia rosea (KI.) A. DC. in DC. Prod. 151:299. 1864. Begonia guyanensisvar. glaberrimaC. DC. in Bot. Gaz. 20:540. 1895.

Herbaceous, slender,2-6 dm. high. Stem simple or slightlybranched,essentially glabrous or glabrescent,prostrate to ascending, sometimesrooting at the ovate) acuminate, asymetric, straight(or rarelytransversely nodes. Leaves strongly unequally cuneate to cordate at base, subpinnatelynerved, occasionally shallowly margins, 4.5-7 cm. long, 1.5-2 cm. wide, essentially lobed with ciliate-serrate glabrous on both surfaces,petioles 0.5-2 cm. long, glabrous, stipules persistent, lanceolate,up to 2 cm. long, entire,glabrous,acuminate. Peduncles axillary,2-3 lanceolate to cm. long, glabrous. Cymes laxly few-flowered. Bracts persistent, 1-2 mm. long. Staminatepedicelsveryslender, to ciliate-serrulate, ovate, subentire to 6 mm. long. Staminatetepals 2, ovate to suborbicular,2 mm. long and broad. Stamens free, few, antherselliptic or ovate, the connective produced. Pistillate to almost bracteoles persistent,obovate to orbicular, entire to ciliolate-serrulate lacerate,about 3 mm. long. Pistillatepedicels to 5 mm. long. Pistillate tepals 5, elliptic to ovate, 2-3 mm. long. Styles 3, 2-partedwith the stigmatictissueforming a continuouslinear spiral band, ovary 3-celled, placentae variable even in the same ovary. Capsule subelliptic to orbicular, glabrous, 6-14 mm. long, with subequal roundedwings, seeds stalked,oblong and obtuse, truncateat the base. Mexico and Guiana to Peru. alt. 0-50 m., Carleton 54; von Wedel 370; vicinity of Nievecita, BOCAS DEL TORO: of ChiriquiLagoon,von Wedel I33I; ChanguiWoodson, Allen & Seibert I8I6; vicinity nola Valley, Dunlap I67. CANAL ZONE: Chagres, Fendler 297.
16. BEGONIA FILIPES Benth. Bot. Voy. Sulph. 101. 1845.

Begonia hygrophilaC. DC. ex Th. Dur. & Pittierin Bull. Soc. Bot. BeIg. 351:265. 1896. Begonia hygrophilavar. puberula C. DC. ex Th. Dur. & Pittier in Bull. Soc. Bot. BeIg. 351: 266. 1896. Begonia mameiana C. DC. in Smithson.Misc. Coll. 69: no. 12:4. 1919. Begonia leptopodaC. DC. in Smithson.Misc. Coll. 69: no. 12:6. 1919. C. DC. in Smithson.Misc. Coll. 69: no. 12:8. 1919. Begonia chepoensis

heterodonta Rusby, Descr.New Sp. S. Am. PI. 66. 1926. Begonia Herbaceous, 2-6 dm. high. Stem simple to slightlybranched,glabrous,finely semiovateto obliquely or transversely ovate, ridged. Leaves stronglyasymmetric, acuminate,mostlytruncateat base, palmatelynerved with the chief veins branchand ciliate, 4-10 cm. ing divaricatelyabove and with the marginscrenate-dentate above with long tapering long, 2-4 cm. wide, ratherabundantly appressed-pilose hairs,essentially glabrouson lower surface,petiole 0.5-4 cm. long, glabrous,stipules oblong-ovate,acute to acuminate to the setulose tip, 6-12 mm. mostlypersistent, long, 2-4 mm. wide, very thin, entire. Peduncles axillary, up to 5 cm. long, ovate-acute with setulose tip, glabrous. Cymes few-flowered. Bracts persistent,

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19581

FLORA

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57

up to 7 mm. long and 2 mm. wide. Pedicels glabrous,4-10 mm. long. Staminate tepals 2, ovate, to 3.5 mm. long and 2 mm. wide. Stamens few, attached to a to short column, connectiveslightlyproduced. Bracteolesobovate, serrate-ciliate, ovate or 4 5, Pistillate occasionally tepals 3.5 mm. long, more or less persistent. to orbicular,ca. 2 mm. long, slightlymore than 1 mm. wide. Styles 3, 2-parted, with the papillate stigmatic surface formingspiral bands. Ovary 3-celled, placentae variable. Capsule glabrous,4-6 mm. long, the largestwing mostlyobtuse, to 1 cm. wide, the other two subequal, narrow, seeds stalked, oblong, obtuse, truncate at base. Costa Rica and Panama to Colombia. between forest Frijolesand Monte 225I; Mamei,alt. 10-30 m., Pittier ZONE: CANAL 84; Las Gatu'n,Ostenfeld along smallrivuletin forest, Lirio,alt. 30 m., Killip I2I43; Cerro Gordo, near Standley25706, 25723, 295I2; near Summit, Cascadas Plantation, 27222; hillsnorth Standley 26003; hillswestof the canal,nearGatu'n, Standley Culebra, 28406; Rio Paraiso,above East Paraiso, 27568; Gamboa,Standley Standley of Frijoles, 3I743; Barro 30952; Obispo,Standley Standley of FortSherman, 29882; vicinity Standley & Vestal 39I; drowned 40909, 41020, Woodworth Standley Lake, Gatu'n Island, Colorado MaddenDam and alt. 70-75 m.,Dodge d Allens. n.; between of QuebradaCulebra, forest & Allen i6.504; on tree, Saddle 11, near Alahuela,alt. 90-100 m., Dodge, Steyermark withRio Pequeni and Rio Indio,alt. junction between alongRio Chagres forest drowned of San Felix,alt. 0-120 in., Pittier & Alleni6795. CHIRIQUI: vicinity 66 m.,Steyermark wooded swamp betweenFrance Field and Catival, Standley30I94. COLON: 52I5. along ChavareRiver, of Pinogana,alt. 20 m., Allen 937. PANAMA: vicinity DARIEN: 47I3; rockybanksof Pacara River,KilliP3I56; wet nearChepo,alt. 50-200 m., Pittier 27888; moistforest, 26I55, 28234; Taboga Island,Standley Rio Tapia, Standley forest, io8. Johnston Island, 294II; San Jose Standley Rio Tecumen, 17. BEGONIA FISCHERI Schrank var. TOVARENSIS (Kl.) Irmscher,Bot. Jahrb.76: 23. 1953. KI. in Abh. Akad.Berlin1854:31. 1855. tovarensis Begonia sensuLiebm.in Kjoeb. Vidensk.Meddel. 1852:16. 1853, non HBK. Begoniapopulifolia 1825. K1. in Abh. Akad. Berlin1854:31. 1855,nonKunth& Bouche,1848. moritziana Begonia 03. 1864. A. DC. in DC. Prod. 151:3 , ocanensis tovarensis Begonia Misc.Coll. 69: no. 12:5. 1919. C. DC. in Smithson. ciliibracteola Begonia to glabrous. Stem erect,red, Herbaceous, 2-12 dm. high,sparselybrown-pilose broadly ovate or suborbicular, branches short. Leaves more or less asymmetric, cilicrenate-serrate, acute or rounded,cordate at base, 2-9 cm. long, palminerved, ate, glabrous above, more or less brown-pilosebeneath especially on the nerves, petioles 5-45 mm. long, stipulesdeciduous,ovate-oblong,to 10 mm. long, ciliateserrulate. Peduncles axillary, 15-55 mm. long. Cymes bisexual, few-flowered. ovate, 2-4 mm. long, fimbriate. Pedicels 5-20 mm. long. StaBracts persistent, minate tepals 4, the outer orbicular, 8 mm. long, the inner smaller, narrowly short,anthersoblong, the connective obovate. Stamens free,numerous,filaments produced,obtuse. Pistillatebracteoleslike the bracts. Pistillate tepals 5, obovate, 3-6 mm. long. Styles 3, bifid,the stigmatic tissue linear, spiral, continuous,placentae bilamellate,ovuliferousthroughout. Capsule 10-15 mm. long, its wings often hooked, veryunequal, the largesttypicallyascendingand tapering, decurrent, to 23 mm. wide, seeds fusiform.

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ANNALS OF THE MISSOURI BOTANICAL GARDEN Cuba, Mexico, Central America, Colombia, Venezuela, Peru, Bolivia.

L Vul. 45

or floating island, ZONE: AhorcaLagartoto Culebra,Cowell 388; on marshy d Vestal696. COCLE: shaded 460, Bailey644, Woodworth Island, Kenoyer Barro Colorado of El Valle de Ant6n,alt. 600 m., Allen 2777. near Rio Anton,floor margins of marsh alt. 26-50 m., lowerRio Trinidad, of Arenoso, PANAMA: on grassyjams in lake,vicinity Seibert 617.
CANAL

18. BEGONIA OAXACANAA. DC. in DC. Prod. 151:3 12. 1864. A. DC. in DC. Prod.151:312. 1864. Begonia oaxacana3 pilosula Begonia luxiiC. DC. in Bot.Gaz. 20:541. 1895. serrulatoala C. DC. in Bull.Herb.Boiss.II. 8:321. 1908. Begonia Begonia pubipedicella C. DC. in Smithson. Misc.Coll. 69: no. 12:7. 1919. terrestrial or epiphytic,suberect or climbing,very Herbaceous to suffruticose, variable in all its parts, 3-15 dm. high. Stems 2-12 mm. thick,glabrousto pilose, or oblique or rarelysome almost straight,broadly greenor red. Leaves transverse or narrowlyovate, acuminate, palmately 5-7-nerved, cordate at base, slightlyor sparselyhirsute usually not at all lobed, 6-18 cm. long, 3-11 cm. wide, serrulate, 4-15 cm. long, sparsely on both sidesor nearlyglabrousabove, thin,petiolesslender, hirsute to glabrous, stipules deciduous, oblong, 12 mm. long, setose-mucronate, bisexual. Bracts entire. Peduncles axillary,2-5 cm. long. Cymes few-flowered, tardily deciduous, broadly ovate or suborbicular,obtuse, 6-12 mm. long, entire. Pedicels 10-15 mm. long. Staminate tepals 4, the outer suborbicularor broadly ovate, 10-15 mm. long, pink, glabrous or at times somewhatpubescent,the inner longerthan obovate, white. Stamensfree,numerous,the filaments slightlyshorter, the obovoid anthers. Pistillatebracteoleswanting or quickly deciduous. Pistillate tepals 3, the outer two suborbicular,8-14 mm. long, the innerone much smaller, narrowlyobovate or oblong. Ovary 2-3-celled, placentae bifid,ovuliferouson all bilobed, the stigmatic surface linear, spiral, continsides, styles 3, short-connate, uous. Capsule erect, broadlyelliptic, 10-15 mm. long, hirsuteto glabrous,wings 3 or sometimesup to 6, subequal, narrow, angled at middle or somewhat above seeds giving the fruit a rhombic or obovate outline, entire to fimbriate-serrulate, ellipsoid,broadlyobtuse, short-stalked. SouthernMexico, Guatemala, Panama. slopeof Cerrode la Horhumidforest of Cuestade Las Palmas,southern cHIRIQuS: alt. 1800 Boquete District, Bajo Chorro, 3248; rainforest, queta,alt. 1700-2100m., Pittier alt. 1500-2000m., Woodson, of CasitaAlta,Volcain de Chiriqui, 55; vicinity m., Davidson of CerroHorqueta, alt. 1500Allen& Seibert slopes 959; Bajo Mono,RobaloTrail,western 2100m.,Allen4809. 19. BEGONIA UDISILVESTRISC. DC. in Smithson.Misc. Coll. 69: no. 12:9. 1919. Herbaceous, 1 m. high, branching,completelyglabrous except for a ring of or oblique, ovate, cordate trichomesat the apex of the petiole. Leaves transverse to 11.5 cm. long, serrulate,petiolesslender, at base, long-acuminate, palminerved, regular, to 11 cm. long, verruculose,stipules deciduous. Cymes few-flowered, about equaling the leaves or shorter. Bracts deciduous,entire. Staminate tepals 4, entire,the outer suborbicular,5 mm. long, the inner narrowlyobovate, shorter.

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1958]

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59

B.oaxacana

B.udisiIvestris

B.involucrata

(/,

B.muV Iinerva

i B.tonduzi
B.heydeil

Fig. 18. Begonia

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BOTANICAL

[Vol. 45 GARDEN

Stamens free,anthers oblong,shorter thanthe filaments.Pistillate tepals3, the outertwo suborbicular, 5 mm. long,the innerone muchsmaller and narrower. Ovary 3-celled,placentaebilamellate, ovuliferous throughout, stylesdeciduous, connate at base,bifid. Capsuleerect, ovoid,attenuate into a slender beak,wings equal,verynarrow. Costa Rica and Panama.
BOCAS DEL TORO: Robalo Trail, northern slopesof Cerrode la Horqueta,alt. 18002100 m., Allen 4938. CHIRIQUI: humid forestof Cuesta de Las Palmas, southernslope of Cerro de la Horqueta, alt. 1700-2100 m., Pittier 3249; Volcain de Chiriqui, Boquete District, alt. 2700 m., Davidson 1025; trail from Cerro Punta to headwatersof Rio Caldera, alt. 2250-2500 m., Allen I440; Casita Alta to Cerro Copete, alt. 2300-3300 in., Woodson & Schery 349; PortreroMuleto to summit,Volca'n de Chiriqui, alt. 3500-4000 m., Woodson & Schery 444.

20.

BEGONIA

INVOLUCRATA

Liebm. in Kjoeb. Vidensk. Meddell. 1852:15. 1853.

Begonia broussonetiifolia A. DC. in Ann. Sci. Nat. IV. 11:133. 1859. Begonia laciniosa A. DC. in DC. Prod. 151:340. 1864.

Fruticose,1-2 m. high. Stem fulvous-tomentose or becomingglabrous. Leaves oblique, cordateat base, 2-5-lobed at apex with acuminatepoints, 15-28 cm. wide, to 20 cm. high, denticulate,evenly pilosulous above, more denselyon the nerves beneath,petioles 12-18 cm. long, fulvous-tomentose or becomingglabrous,stipules deciduous,ovate, setose-apiculate, 14-25 mm. long, entire,brown,membranaceous, glabrous or sparsely pilose. Peduncles axillary, exceeding the leaves, fulvoustomentose becomingglabrous. Cymes many-flowered, dense,theirprimary branches short and theirultimate branchesso nearly aborted that the inflorescence appears almost umbellate. Bracts deciduous,entire,the outer ovate, ample and completely enclosing the young inflorescence. Pedicels 5-25 mm. long. Staminate tepals 2, suborbicular,6-11 mm. long, entire,sparselypilose to glabrous,white. Stamens numerous, anthers oblong, about equaling the filaments,connective produced, broadlyobtuse. Pistillatebracteoleslacking. Pistillatetepals 2, like the staminate. Ovary 3-celled, placentae bilamellate, styles connate at base, bifid. Capsule ellipsoid, very unequally 3-winged, the largest wing elliptic or falcate, obtuse, 8-10 mm. wide. Guatemala, Nicaragua, Costa Rica, Panama.
moist shaded places, valley of the upper Rio Chiriqui Viejo, alt. 1300-1900 CHIRIQUI: m., White 34; rain forest,Bajo Chorro, Boquete District, alt. 1800 mn., Davidson I08; vicinity of "New Switzerland", central valley of Rio Chiriqui Viejo, alt 1800-2000 m., Allen I349. 21. BEGONIA MULTINERVIA Liebm. in Kjoeb. Vidensk. Meddell. 1852:18. 1853.

in Bull. Soc. Bot.Belg. 351:260. 1896. C. DC. ex Th. Dur. & Pittier Begonia cuspidata 2-3 m. high. Stem flexuouswith elongate internodes,brownSuffruticose, pilose. Leaves transversely elliptic, abruptly acute, cordate at base, palminerved, 12-22 cm. long, entire,glabrous,petioles divergent,6-16 cm. long, brown-pilose, 2-3 cm. long, stipulesdeciduous,lance-oblongor narrowlytriangular, setose-acute,

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61

sulcate, soon glabrous. Cymes bisexual, regularlydichotomous,12-25 cm. broad, glabrous. Bracts deciduous,suborbicular, entire,the lowest 17 mm. long. Pedicels 7-15 mm. long. Flowerswhite. Staminatetepals 2, broadlyovate or suborbicular, 8 mm. long, entire. Stamens numerous,anthers obovoid, shorterthan the filaments,connectivescarcelyif at all produced. Pistillatebracteoleslacking. Pistillate tepals 2, like the staminate. Ovary 3-celled,placentae bilamellate,ovuliferous throughout, stylesmuch connate at base, bifid,the stigmatic tissue linear, spiral, continuous. Capsule subglobose,7 mm. long, wings veryunequal, the largestovate or elliptic,to 14 mm. wide, the othersnarrowlylunate, seeds stalked,ellipsoid. Costa Rica, Panama. Pittier 5420.
22.
BEGONIA CHIRIQUI:

entire, membranaceous. Peduncles axillary, 15-38 cm. long, exceeding the leaves,

between Hato del Joboand CerroVaca, eastern Chiriqui, alt. 700-1000 m.,
URTICAE

L. f. Suppl. 420. 1781.

Begonia urticifolia J.E. Smith, P1.Icon. Ined. 2: P1.45. 1790,proerrore. Begonia columnaris Benth. P1.Hartweg.131. 1844. Begonia trachyptera Benth. P1.Hartweg.184. 1845. Stiradotheca trachyptera KI. in Monatsber. Akad.Berlin127. 1854., Begonia coccinea Ruiz ex KI. in Monatsber. Akad. Berlin127. 1854,e. p., nomen in synon. Sasseaurticae KI. in Monatsber. Akad.Berlin128. 1854. Sasseacolumnaris KI. in Monatsber. Akad.Berlin128. 1854. Begonia glabra Ruiz ex KI. in Monatsber. Akad.Berlin128. 1854,nomen in synon. KI. in Abh.Akad. Berlin1854:128. 1855. coccinea Casparya Stibadotheca trachyptera KI. in Abh.Akad.Berlin1854:131. 1855. SasseaglabraKI. in Abh.Akad.Berlin1854:134. 1855. Begonia cucullata Ruiz ex KI. in Abh.Akad.Berlin1854:134. 1855,nomen in synon. Casparya trachyptera A. DC. in DC. Prod.151:274. 1864. columnaris Casparya A. DC. in DC. Prod. 151:274. 1864,nonKI. 1854. I glabra columnaris Casparya A. DC. in DC. Prod. 151:274. 1864. urticae A. DC. in DC. Prod.151:274. 1864. Casparya urticae A. DC. in DC. Prod.151:274. 1864. Casparya ,/hispida Sasseahoffinanniana KI. ex A. DC. in DC. Prod.151:275. 1864,nomen in synon. Begonia C. DC. in Bull. Herb.Boiss.II. 8:325. 1908. monticola Begonia torresii Standl.in Journ. Wash.Acad Sci. 17:313. 1927. Begonia chiriquensis Standl. in Woodson & Schery in Ann.Mo. Bot.Gard.27:321. 1940. Begonia columnaris var. glabraSmith & Schubert in Macbride, Fl. Peru,in FieldMus. Pub. Bot. 134:1 87. 1941. Herbaceous to suffruticose.Stem branching,decumbent,rootingat the nodes, 15-40 cm. high -orrarelyto 2 m., the youngerpartsusually ferruginous-puberulent. Leaves straightor nearly so, stronglyasymmetric, ovate or elliptic, acute or acuminate,base acute on the adaxial side, obtuse and decurrenton the other, 3-8 cm. long, penninerved,doubly serrate,ciliate, sparsely hirtellous to glabrous above, denselypubescenton the nervesto wholly glabrousbeneath,petiole2-15 mm. long, stipulesdeciduous,ovate, obtuse, setose,2-6 (rarely to 12) mm. long. Peduncles axillary,erect, 12-45 mm. long, 1- to few-flowered. Bracts deciduous, elliptic, setaceous-dentate at apex. Pedicels 6-18 mm. long. Staminatetepals 4, subequal, 3-8 mm. long, the outer elliptic,entire,red, often pilose, the innerobovate, white. Stamenson a slendercolumn 2 mm. high, filaments short,antherslinear,the con-

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5, subequal, elliptic, 3-5 mm. long, entire, red or white, the outer often pilose. Styles 3 with many shortbrancheswholly covered by stigmaticpapillae, placentae bilamellate, ovuliferous on all sides. Capsule very variable, broadly turbinate, obtuse at base, with 3 ascendinghorns on the angles, the tips of the horns often deciduous, the apical column well developed,angled or terete,slender,conical or cylindricalor subclavate, from longer than the horns to rarelyshorter. Costa Rica to Peru.
BOCAS DEL TORO: alongstreams, RobaloTrail,northern slopes of Cerrode la Horqueta, alt. 1800-2100m., Allen4980. CHIRIQUI: rainforest, BoqueteDistrict, alt. Bajo Chorro, 1800 m., Davidson107; epiphytic, trailfrom CerroPunta to headwaters of Rio Caldera, alt. 2250-2500m.,Allen1435; vicinity of Cerro Punta,alt. 2000 m.,Allen3484.

nective slightly produced. Pistillate bracteoles deciduous, elliptic. Pistillate tepals

23.

BEGONIA

HEYDEI

C. DC. in Bot. Gaz. 20:540. 1895.

Begonia Pittieri C. DC. in Bull.Herb.Boiss.II. 8:316. 1908. Begonia triloba C. DC. in Bull.Herb.Boiss.II. 8:322. 1908. Erect, to 6 dm. high, branching with elongate internodes,glabrous. Leaves oblique, cordate at base, narrowlyovate, acuminate, penninerved, 6-10 cm. long, 2-3.5 cm. wide, remotelydenticulate,petioles 10-34 mm. long, stipulesdeciduous, lance-ovate, subulate, 5 mm. long. Peduncles axillary,15-28 mm. long. Cymes few-flowered, exceeded by the leaves. Bracts deciduous. Pedicels 8-10 mm. long. Staminate tepals 4, entire,obtuse, the outer ovate or elliptic, 7 mm. long, setose toward base, the innerslightlyshorter. Stamens many on a shortcolumn, anthers obovate, nearly equaling the filaments. Pistillate bracteoleselliptic, entire,about equaling the ovary. Pistillate tepals 3, elliptic or obovate, entire,setose toward base. Ovary 3-celled, placentae simple, styles deciduous, connate at base, bifid. Capsule rhomboidal, attenuateat base, bearingthreeequal slenderacuminate horns 9-11 mm. long. Guatemala, Costa Rica, Panama.
COCLE: of El Valle de Anton,alt. 600 m., Allen 2078; epiphytic, epiphytic, vicinity La Mesa,Allen2722.

24. BEGONIA TONDUZII C. DC. in Bull. Soc. Bot. Belg. 351:264. 1896. allenii Standl.in Woodson & Seibert Begonia in Ann.Mo. Bot.Gard.25:83 0. 1938. Caulescent,28-48 cm. high. Stem erect,simpleor rarelyfew-branched, woody at least toward base, rufous-pilose becoming glabrous with age, internodes mostly short but distinct. Leaves nearly straight, strongly asymmetric,oblanceolate, penninerved, acuminate, cuneate, falcate, 6-12 cm. long, 2-5 cm. wide, coarsely sinuate-dentate near apex, glabrousor pilose on the midnervebeneath,petioles5-9 mm. long, pilose to glabrous,stipulesmore or less persistent, broadlyovate, setoseacuminate, 10-25 mm. long, entire,thin. Peduncles few, axillary,3-7 cm. long. Cymes few-flowered, regular,ratherdense,pilose at first. Bracts persistent, small, suborbicular,serrate. Pedicels 5-16 mm. long. Flowers pink or red. Staminate tepals 2, broadly ovate, 5-8 mm. long, more or less pubescent. Stamens free,

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63

connective produced, obtuse. Pistilanthers oblong, aboutas longas thefilaments, placentae tepals5, equal. Ovary3-celled, latebracteoles likethebracts. Pistillate tissuelinear, throughout, styles connate at base,stigmatic bilamellate, ovuliferous wings subequal,veryvariablein spiral,continuous. Capsule erect,subglobose, width. CostaRica, Panama.
COCLE: El Valle de Anton, along Rio Indio trail,alt. 500-700 m., Hunter 3' Allen 327; northrim, vicinity of El Valle, Allen234, i663; banksalongtrailto La Mesa,hills of El north of El Valle de Anton,alt. 800-1000 m., Allen 2286; CerroPajita,hillsnorth alt. 1000-1200m.,Allen4I74. Valle de Anton, 25.
BEGONIA OPULIFLORA

Putz. in Fl. des SerresI. 10: P1.995. 1854-55.

strongly over 1 m. high,glabrous. Leaves straight, Suffruticose, branching, asymmetric, ovate or elliptic,acute, 8-9 cm. long, roundedat base, pinnate, oblong-cordate, much shorter than the blades,stipules subulate-dentate, petioles

ora B.opulifr

Fig. 19. Begoniaopuliflora

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long-aristate, membranaceous. Peduncles axillary, about 15 cm. long. Infloresglobose, 50-60-flowered. Pedicels elongate. Flowers white. cences umbelliform, Staminate tepals 4, subequal, broadly elliptic, obtuse, entire. Stamens about 30, slightlyconnate at base, anthersoblong. Pistillatetepals 5, oblong or lance-oblong, subacuminate. Styles3, connate,stigmatictissuelunate. Fruit 3-winged,the wings very unequal, the largest much wider than high, twice the width of the others. Endemic Province of Soto whichcentered fromcultivation fromthe former COCLE: Described on Penonome. 26.
BEGONIA GUADUENSIS

HBK. Nov. Gen. & Sp. 7:178.

1825.

Repert. Bot. Syst.2:212. 1843. Begonia ottonis Walpers, Begonia walpersii Heynh.Nom. 2:63. 1846. Akad.Berlin127. 1854,nomen. Donaldiaottonis Ki. in Monatsber. in synon. laurina Begonia Hort.ex A. DC. in DC. Prod.151:292. 1864,nomen C. DC. in Smithson. Begonia serratifolia Misc.Coll. 69: no. 12:7. 1919. scandent to 2 m. high, glabrous. Stem erect, branched. Leaves Suffruticose, straight, asymmetric,pinnate-nerved,lanceolate to elliptic-lanceolate,acute or acuminate,unequal at base with the adaxial side cuneate and the otherroundedand ciliate, memdecurrent,5-9 cm. long, 16-40 mm. wide, doubly crenate-serrate, oblong-ovate, branaceous,petiole2-8 mm. long, stipulesdeciduousor subpersistent, 8-10 mm. long, scarious. Peduncles terminaland axillary, acuminate-setiferous, 6-15 cm. in diameter. few- to many-flowered, 4-5 cm. long. Cymes dichotomous, 4-7 mm. long, ovate, obtuse,acute or mucronate, Bracts deciduousor subpersistent, scarious. Pedicels 4-19 mm. long. Flowers pinkish white. Staminate tepals 4, the outer elliptic-ovate,entire or dentate at apex, 8-15 mm. long, the inner obovate, distinctlyshorter. Stamens free,very numerous,anthersoblong, mostly shorterthan the filaments,connective produced, apiculate or obtuse. Pistillate bracteoles deciduous, ovate to narrowlyobovate, 6 mm. long, entire. Pistillate tepals 5, subequal with the two outer slightlysmaller,elliptic, acute, 6-10 mm. long. Styles 3, bifid nearly to base, each main branch with 3-4 short spiral branches, placentae bifid, ovuliferous throughout. Capsule ellipsoid, the largest wing triangular,horizontal or slightly ascending, 15 mm. wide, the other two marginiform. Panama, Colombia, Venezuela. between Gorgonas and Gatun,alt. 10-50 m., CANAL ZONE: sidecuts,railroad relocation alt. 70-75 m.,Dodge & Allen; westerly Pittier forest of QuebradaCulebra, 2258; drowned armof QuebradaSalamanca, alt. 70 m.,Dodge,Steyermark & Allen; nearMaddenDam at trailbe& Allen I6822; forest along telephone Alahuela,alt. 70 m., Dodge, Steyermark tweenthe Rio Indio Hydrographic Stationand the Natural Bridgeof the Rio Puente, of QuebradaAncha,alt. 70 m., Dodge & Steyerforest Dodge & Allen I748I; drowned mark. CHIRIQUI: of San Felix, eastern Chiriqui,alt. 0-120 m., PittierSI26. vicinity VERAGUAS: hillswestof Sona,alt. 500 m., AllenI034.
27.
BEGONIA ESTRELLENSIS

C. DC. in Bot. Gaz. 20:540.

1895.

Woody vine. Stem elongate,glabrous. Leaves pinnatelyveined, ovate-elliptic,

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B.guaduensis
tS /1a

B.seemaniana

B.carpinifolia

lariodora B. conval

Fig. 20. Begonia

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ratherabruptly narrowed to a slender acumination (to 1 cm. long), acute to cuneateat base withthe broader half produced 1 mm.beyond the narrower, remotely denticulate, glabrous, foveolate, 5-6.5 (-9.5) cm. long,1.7-2.3 (-3.3) cm. wide,petioles to 12 mm.long,glabrous, stipules earlydeciduous, onlyscarsseen. Inflorescences pedunculate and cymose, terminal and axillary. Staminate pedicels to 18 mm.long. Tepals regularly 4, theouterovate,obtuse, to 6 mm.long and 5.5 mm.broad, theinner elliptic, to 5 mm.longand 2 mm.wide. Stamens numertheconnective ous,free, produced. Pistillate pedicels slender, glabrous, to 20 mm. long. Tepals 2, ovate,broadly acute,to 10 (-16) mm. long and 8 mm. wide. Ovary glabrous, puncticulate.Styles3, bifid, broadly expanded above the base. Capsuleunequally tri-alate, the largest wing to 10 mm. wide,ascending, acute. Costa Rica, Panama.
COCLE:

vicinity ofEl Valle, north rim, alt.800-1000 m.,Allen 220. DC. in Ann. Sci. Nat. IV. 11:133. 1859.

28.

BEGONIA SEEMANNIANA A.

Begonia chiriquina C. DC. in Smithson. Misc.Coll. 69: no. 12:8. 1919.

Shrubto 3.5 m. high. Stemslender, glabrous, finely ribbed. Leaves asymmetric, elliptic-obovate, penninerved, serrulate-denticulate with setiferous teeth, acuminate, unequally obtuseat basewiththebroader sideextended on the petiole thenarrower, beyond glabrous to minutely puberulent throughout, foveolate, 9.514 cm. long,4.5-6 cm. wide,petioles glabrous, rugose, 5-15 mm. long,stipules glabrous, ovateto elliptic, obtuseto acuminate and mucronate, truncate to nearly at base,15-25 mm.long,4-6 mm.wide. Peduncles clasping to 7 cm. long. Inflorescences the staminate cymose, terminal and axillary. Bractsinconspicuous, thin, elliptic, ca. 1 mm.long,thepistillate veryearly deciduous.Pedicels glabrous, to 2 cm. long. Staminate to semi-orbicular, slender, tepals2, ovate-obtuse usually cordate at base,5-6 mm.long,3-8 mm.wide. Stamens withconnumerous, free, nectives produced and exceeding the shorter outerfilaments.Pistillate tepals2 (rarely a smaller third onepresent), obovate andobtuse to elliptic, 5-8.5 mm.long, withreddish sometimes dotson surface. Styles 3-5.5 mm.wide. Ovaryglabrous, free. Capsule tri-alate, the largestwing with 3, bifidor irregular, essentially the thirdonly truncate obtuseapex, to 15 mm. wide, one wing marginiform, lessthan0.5 mm.long. or ellipsoid, broader. Seedsobovoid slightly Costa Rica, Panama.
queta, alt. 1700-2100 m., Pittier; vicinity of El Boquete, alt. 1000-1300 m., Maxon 4948;
CHIRIQUI:

humidforest of Cuestade Las Palmas,southern slopeof Cerrode la Hor-

Pittier 3152.

29.

BEGONIA CARPINIFOLIA

Liebm.in Kjoeb. Vidensk.Meddel.1852:20. 1853.

or subscandent.Stem glabrous, suffrutescent Herbaceous, greento reddish, to obliquelyelliptic,obliquely finely ridged. Leaves obliquelyovate-acuminate 1 mm.or more thenarrower, halfproduced at basewiththebroader obtuse beyond (66)

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67

with setiferousteeth, 5-7.5 (-11) cm. long, 2.5-3.7 (-4.7) cm. wide, petioles 3.5-10 mm. long, glabrous, stipules persistent,pinkish to greenishwhen fresh, linear to lanceolate or narrowlyovate with oblique base and acute apex, 7-13 mm. pedunculate, cymose,axillary and terminal. long, ca. 5 mm. wide. Inflorescences Bracts of the peduncles similar to the stipules, secondary bracts and bracteoles obovate to orbicular,obtuse to truncate. Pedicels glabrous,slender,to 1 cm. long in fruit. Staminate tepals 2, orbicular,cordate at base, 5-8 mm. long, 6-7 mm. connectiveproof the outer ones shorter, free,filaments wide. Stamensnumerous, orbicular,3-6 mm. long, 4.5-5 mm. wide. Styles duced. Pistillatetepals 2, chiefly 3, short-connate. Capsule ca. 6 mm. high, its largest wing ascending, acute to and subequal. Seeds minute, obtuse, ca. 5 mm. wide, the other two marginiform obovoid. slenderly Costa Rica, Panama. centralvalleyof Rio ChiriquiViejo, alt. of "New Switzerland", vicinity CHIRIQUI: 1800-2000m., AllenI408. 30.
BEGONIA
CONVALLARIODORA

serrate-denticulate prominently beneath, veined andprominently foveolate glabrous,

C. DC. in Bot. Gaz. 20:538.

1895.

Herbaceous, 1-15 m., scandent to erect. Stem slender,little branched,ribbed of the main axis 5.5-14 cm. long. Leaves pinnately and glabrous,the internodes somewhat denticulate,obliquely elliptic, abruptly acuminate, veined, asymmetric, the broaderhalf usually producedinto a conspicuouslobe at base, 8.5-15 cm. long, 4-7.5 cm. wide, petioles 12-40 mm. long, stipules membranaceous,obliquely elliptic, entire,acute and apiculate, truncate at base, 25 mm. long, 7 mm. wide. Peduncles 3-8 cm. long. Cymes regular,multi-flowered.Bracts early deciduous. Pedicels 6-16 mm. long. Tepals entire,white or pinkish,the staminate2 (rarely elliptic, 3 or 4), orbicular,cordate at base, 4-8 mm. long, the inner,when present, 1-3.5 mm. long. Stamens numerous,anthersoblong-elliptic,usually equal to or connective produced, obtuse. Pistillatebracteolessoon longer than the filaments, deciduous, entire, elliptic, obtuse. Pistillate tepals 2 (rarely 3), orbicular to rounded-ovate,6-9 mm. long, the third,when present,smaller. Styles 3, bifid. Placentae bilamellate, ovuliferous throughout. Capsule ellipsoid, 3-winged, the largestwing obtuse, 7-12 mm. broad. SouthernMexico to Panama. of MonteLirio,alt. 1300valleyof theupperRio ChiriquiViejo, vicinity CHIRIQUI: alongRio Caldera,alt. 207; Bajo Mona,mouthof QuebradaChiquero, 1900 m., Seibert of Bajo Mona and Quebrada Allen d SeibertIOI5; vicinity 1500-2000 m., Woodson, alt. 1900 m., of Bajo Chorro, alt. 1500 m., Woodson 533; vicinity s Schery Chiquero, , Schery of El Valle, alt. 600-1000 m., Allen vicinity Woodson 647. COCLE: epiphytic,
ii88, 1207.

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CACTACEAE
or epiphytic More or less massive and succulent, usually leafless terrestrial plants. Stemsusually (except in Pereskia) phylloidand jointed,the joints elongate the nodes with axillaryareas (areoles) tereteto angled or flattened, to suborbicular, usually armedwith spines,slenderbarbs (glochids) or hairsin variousnumbersand occasionally naked. Leaves alternateand laminate in Pereskia,but combinations, in the othergeneraminute and fugacious,or wholly lacking. Flowers pedunculate panicles in Pereskia,in the other genera solitaryand and in 1- to several-flowered sessile,very large to very small, frequentlynocturnal, epigynous (perigynousin to rotate, with or without a Pereskia); perianthsalverformor infundibuliform actinomorphicwhen erect and more or less conspicuous tube, monochlamydeous, zygomorphicby position when horizontal or pendulous, the segments petaloid, usually white to red or yellow, relativelyfew to exceedinglynumerous,the inner broaderand more petaloid than the outer and lower, the tube naked or with bracts inserted with elongatefilaments, or areolesor both; stamensusually verynumerous, within the perianthtube or upon the ovary in rotate flowers;ovary inferior(subin Pereskia), 1-celled,with few to severalparietalplacentasusually bearing inferior with an elaborate numerousovules, the style more or less elongate and filiform, radiate terminalstigma. Fruit a berryusually with numerousseeds immersedin succulent pulp. The Cactaceae are a typicallyWesternHemispherefamily,with only the genus Rhipsalis,the MistletoeCactus, doubtfullyindigenousto the Old World. Species of Opuntia, however,especially 0. ficus-indica,have been introduced since very early times and become naturalized in Africa and elsewhere. The genera vary in of individual authors,from only 20 (K. number according to the interpretation Schumann, in Engl. Nat. Pflanzenfam.III: 6. 1894) to over 100 (cf. Britton & Rose, in Carnegie Inst. Publ. No. 248. 1919-1923). Much criticismhas been made of the numeroussegregategenera proposed by Britton and Rose and much undoubtedly remains to be learned of their natural relationships. In Panama, however, their genera usually are quite readily distinguishable into apparently natural groups; consequently they are adopted here vs. splitters"in dealing of "tlumpers withoutmajor modification. The controversy with cacti dates at least as early as 1813, when Sims (in Curtis's Botanical Magainveighedagainst Haworth's generic segregatesand advozine 38: sub pl. I557) cated the returnof both Cereus and Opuntia to the Linnaean genus Cactus, now long abandoned. as a rule to those so familiar In humid Panama the cacti bear little resemblance in the landscapesof the arid regionsof the United States and Mexico. Here they are mostly clamberingsemilianeson rocks, or epiphytes,with only two or three to representthe Prickly species of Opuntia and Nopalea, possibly introductions, a cactus at all, with its shrubbyor tree-like habit, Pears. Pereskiascarcelyresembles teretestems,and conspicuous leaves. Cacti of the Rhipsalis and Cereus alliances

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often are encountered in Panama, but all to frequently in a sterilecondition. This, togetherwith the difficulty of preparingsuch succulent plants for the herbarium, accounts largely for the very few specimensfound in museums. Actually, the speciesare not at all as rare as the few citationswould indicate. The total number in Panama must be considerably greaterthan that of the presentaccount.
a. Stemsnot phylloid;leavesconspicuous and laminate, deciduous;flowers aa. Stemsphylloid;leaveslackingor inconspicuous and fugacious;flowers sessile, solitary, epigynous. b. Stemjointssuborbicular to broadly linguiform in our species, entire, the areolesamphigenous, with glochids;perianth rotateto campanulate. c. Petals erect, shorterthan the stamens; stamen filaments not erectat anthesis; sensitive, styleenlarged nearthe base into a disciform stylopodium. ,-,,..,..,,,,,, .. ,,,.. ,.. ,,..,.. ,,,,.2. cc. Petalsspreading, longer thanthestamens; stamen filaments sensitive, spreading at anthesis and inflexing upon stimulation; style only bb. Stem jointselongate, tereteto angled,or flattened and undulateor lobed, theareoles marginal upontheflowering without glochids. joints, c. Flowers verylarge,rarely mediocre, with a definite perianth tube; stamens adnate to theperianth tube. d. Flowering stemjointsribbedor angled, the areoles usuallymore or lessspiny and bristly. e. Stem joints usually 5-angled,with numerouslarge spines; plantsterrestrial, the stemserectat first, but finally deflexed and rooting at the tips; perianth tube much longerthan the the segments, adnate bractsinconspicuous and fugacious but with prominent spinyareoles.-,,,,,,,,,... 4. ee. Stemjointsusually3-angled, withinconspicuous and infrequent or unarmed; spines, plantsclambering uponrocksor epiphytic, rooting fromthe stemjoints; perianth laterally tube shorter thanthesegments. f. Flowersvery large; perianthtube with conspicuous and not bristly or hairy.-.,,,,. persistent bracts, ,,.,,,,.5. ff. Flowersmediocre;perianthtube with inconspicuous and fugaciousbracts subtending conspicuous persistent bristly or hairy areoles..,,-.,,..,,, ..,..,,..,,,,, ..,, ..,.....6. dd. Flowering stem joints flatand leaflike, lobed or undulate,the areoles nakedto minutely puberulent. e. Flowerslarge,with an elongateperianth tube, the segments stamens the filaments very numerous; very numerous, many or relatively mediocre ee. Flowers shortperismall,witha rather about 10-20; stamens anth tube, the segments about 20-30, the filaments not greatly thanthe anthers; lobes longer stigma
3-5 -..-.-.. times longer than the anthers; stigma lobes numerous-................ slightly thickened toward the base.,,.,,,,,,,,,,,,,,,,,,,,,,,,,,.,,.3.

pedunculate, often clustered, perigynous .

,,,.,,,,,,,,,,,,,,,,,,,,,-,,,,.,,,,,.1.

PERESKIA

NOPALEA

OPUNTIA

ACANTHOCEREUS

HYLOCEREUS

WEBEROCEREUS

7. EPIPHYLLUM

without a definite tubeor thesegments cc. Flowers verysmall, perianth adnate (in our species) to the marginof the quite free; stamens or essentially stemjointsterete flowering hypanthium; so, nakedor 8. withbristly . or hairyareoles .,,,,,,,

8.

WITTIA

RHIPSALIS

1. PERESKIA Mill.
PERESKIA Mill. Gard. Dict. ed. 4. 1754; Britton& Rose, in Carnegie Inst. Wash.

Publ. no. 248. 1:8. 1919.

Neck. Elem.Bot. 2:84. 1790,as Carpophillus. Carpophyllus Peireskia Mill. acc. to Steud.Nom. ed. 2. 2:282. 1841,var. typogr.

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ANNALS O1 THE

MISSOURI BOTANICAL GARDEN

tVol. 45

Leafytrees or shrubs, sometimes clambering; stems terete, branching, succulent above, woody below, armed withstoutspines in minutely pubescent axillary areoles. Leavesalternate, spiral, broadly laminate, deciduous. Flowers pedunculate, paniculate or solitary, both terminal and axillary;perianth rotate, with manyfreeor slightly unitedsegments, the outersomewhat shorter and less petaloidthan the inner;stamens numerous, inserted in a shallowtube adnateto the base of the perianth. Ovary subinferior. Fruita fleshy relatively few-seeded berry bearing themarcescent remains of theperianth. A genusof abouta dozenspecies, ranging from Mexicoand Floridato Argentina and in theAntilles. Verywidelycultivated in the tropics and subtropics of theworldandin greenhouses, and usedas an under stockof grafted Zygocacti and The plants Schlumbergeras. farmore resemble thenorthern OsageOrangein habit thana truecactus.
a. Clambering lianas; spinesshortand reflexed; berries globose, relatively aa. Shrubsor small trees; spineselongateand straight;berriesturbinate, ..2. relatively large 1. PERESKIA ACULEATA
small.............................................-... ..
1. P.
ACULEATA

P.

BLEO

Mill.Gard.Dict. ed. 8. 1768.

Cactus pereskiaL. Sp. P1. 469. 1753. Cactus lucidus Salisb. Prodr. 349. 1796. Pereskia longispinaHaw. Syn. P1. Succ. 178. 1812. Pereskiaaculeata P longispina (Haw.) DC. Prodr. 3:475. 1828. PereskiafragransLem. Hort. Univ. 2:40. 1841. Pereskiaundulata Lem. Illustr. Hort. 5: Misc. 11. 1858. Pereskia pereskia (L.) Karst. Deutsch. Fl. 888. 1882; Britton & Rose, in Carnegie Inst. Wash. Publ. No. 248. 1:10. 1919. PeireskiafoetensSpegg. in Weingart,Monatsschr.Kakteenk. 14:134. 1904. Pereskiagodsefliana Hort. Sand. in Gard. Chron. III. 43:257. 1908.

Clambering shrubs or lianas;branches about 3-10 m. terete, relatively slender, long,withpairsof stoutreflexed spines2-5 mm. long. Leavesshortly petiolate, ,thebladevariable, thebaseobtuse to obovate, obtuseto acuminate, usually elliptic to cuneate, 3-8 cm. long,1.5-4.0 cm. broad,thepetiole3-5 mm.long. Flowers in terminal or axillary usuallyfew or several about twiceas long as the panicles subtending about 0.5-1.5 cm. long,accrescent leaves;pedicels in fruit;perianth white, or pink,rotate, about 3-5 cm. in diameter, yellowish, the segments rather the outeroblong-linear; the innerobovate-oblong, numerous, stamens numerous, lessthanhalfas long as the perianth segments; stylerather stout,slightly longer than the stamens. Berryglobose,1.5-2.0 cm. in diameter, with inconspicuous whenripe. or orange thornless areoles, yellow from Panamaand Colombia;also reported southern Floridaand MexAntilles; cultivated. In the British as an escape. Frequently West Indiesthe ico, perhaps areeatenand madeintopreserves. knownas Barbados fruits, Gooseberries,
PANAMA:

Cerro Campana, alt. 800 m., Allen 3972.

(70)

19 58]

FLORA

OF PANAMA

(Gactaceac)

71

aculeata Fig. 21. Percskia

(71)

72
2.

ANNALS OF THE MISSOURI BOTANICAL GARDEN


PERESKIA BLEO

[Vol. 45

(HBK.) DC. Prodr. 3:475. 1828; Britton& Rose, in Carnegie Inst. Wash. Publ. No. 248. 1:17. 1919.

CactusbleoHBK. Nov. Gen.& Sp. 6:69. 1823. Pereskia panamensis Weber, in Bois, Dict. Hort. 739. 1898. Shrubs or small treesto 4-7 m. tall; branchesterete,ratherstout, armed with few to several slender straight spines usually about 1 cm. long. Leaves rather long-petiolate,the blade usually elliptic-obovate,acuminate, cuneate, 4-20 cm. long, 2-7 cm. broad, the petiole 0.5-3.5 cm. long. Flowers usually few or solitary in terminalor axillarypanicles shorterthan the subtendingleaves; pedicels about 5 mm. long; perianthpink to deep red, rotate, about 5-7 cm. in diameter,the segments rather numerous, the inner broadly obovate, the outer oblong-linear; stamensnumerous,about half as long as the perianthsegments;style ratherstout, slightlylonger than the stamens. Berryturbinate,5-6 cm. long and broad, with severalmore or less persistent bracts, yellow when ripe. South America. In Panama the plants are known Panama and northwestern as gajz' de culebra and iajz de espinas. Britton and Rose found that the plants to P. grandifoliaHaw., a widely cultivated as P. bleo actually should be referred native of Brazil. L34.
trail between Pinogano and Yavisa, Allen 28i; Marraganti, Williams DARIEN: 66i; PANAMA: Cana, Pittier I884. Chepo, Pittier 4704. SAN BLAS: Caledonia Harbor, Elmore

2. NOPALEA
NOPALEA

Salm-Dyck 1850; Britton & Rose, in

Salm-Dyck, Cact. Hort. Dyck. 1849:63. CarnegieInst. Wash. Publ. No. 248. 1:33. 1919.

Massive succulent shrubs; stems phylloid and jointed when young, the joints fleshy, compressed, oval to oblong-elliptic,bearing amphigenousand rather sparse areoles armed with hairs and glochids and usually 1 to few stout spines, stoutly cylindricwhen older. Leaves inconspicuous,acicular, fugacious. Flowers sessile, solitary,chiefly marginal upon the young stem joints; perianthshortlycampanulate, with a short and broad hypanthium,the segments rather few, the outer somewhatshorterand less petaloid than the inner,erect or very slightlyspreading; stamens very numerous,the filamentsmuch longer than the perianth,united in graded seriesat the base of the hypanthiumin a shallow, glandular-dentate cup; ovary broadly turbinate,deeply concave, more or less tuberculate,the areolesvery prominent;style longer than the stamens,enlarged shortlyabove the base into a conspicuousdisciformstylopodium. Fruit a fleshy berrywith numerousseeds. About a dozen speciesof the Antillesand Central America. The two speciesof Panama, if they are speciesindeed,both appear to have been introduced,although Brittonand Rose consideredN. dejecta as "perhaps native in Panama."
a. Areoleswithoutspinesor nearly so; ovary and berriesprominently tuberculate 1. N. COCHENILLIFERA ........................-------------------------------aa. Areoles with2-8 stoutspies; ovaryand berries ....... 2. N. DEJECTA tuberculate scarcely

(72)

19581

FLORA

OF PANAMA

(Cactaceae)

73

..**...~

~~~..

......

..

A\~~~~~~~~~~x

Fig. 22. Nopalea cochenillif era

(73)

74

ANNALS OF THE MISSOURI BOTANICAL GARDEN

LVA1.45

1. NOPALEA COCHENILLIFERA (L.) Salm Dyck, Cact. Hort. Dyck. 1849:64. 1850, as coccinellifera;Britton& Rose, in Carnegie Inst. Wash. Publ. No. 248. 1:34. 1919. Cactuscochenillifer L. Sp.P1.468. 1753. Opuntiacochinelifera (L.) Mill.Gard.Dict. ed. 8. 1768,sphalm. Stout shrubs to 4 m. tall, the older trunksbroadly cylindric,to 2 din. thick, the young stem joints elliptic-oblong,to 5 dm. long, green, the areoles without erect or nearly spinesor nearlyso. Flowerspink to deep red, the perianthsegments so, the outer 0.5-1.5 cm. long, deltoid, the inner broadlyoblong-obovate,1.5-2.5 cm. long, the hypanthiumnaked, broadly campanulate or urceolate, 1.0-1.5 cm. long, about 1 cm. broad at the orifice;staminalfilaments 2.5-3.5 cm. long, pink; ovary 2-3 cm. long, about 1.5 cm. broad, oblong-turbinate, truncate and deeply concave, tuberculate, the areoles unarmed. Berry oblongoid-turbinate, about 5 cm. long, deep red when ripe. Original habitat unknown,widely cultivated and escaped in tropical America. Occasionally encounteredin cultivation or thicketsin Panama, according to both Standley (Contr. U. S. Nat. Herb. 27:278. 1928) and Britton and Rose, but only one specimen of definitely Panamanian origin has been found during the preparationof the presentaccount. When the Spaniardsarrivedin Mexico in 1518 they found the natives using a brilliant purple, red, or orange dye, cochineal (Spanish, cochinilla), which they brushed from the surface of various cacti, particularlyNopalea cochenillifera. Their value quickly appreciated,the dye and the plants fromwhich it was gathered were distributed by the Spaniardsto otherparts of theirempireand to Spain itself as early as 1523. It was not until the year 1703 that the microscoperevealed the source of the was obtained by dye to be a small coccid scale infestingthe plants. The dye stuff brushingthe insectsfromthe parasitizedplant and roastingor boiling theirbodies, which later were dried and pulverized for the vat. Until the invention of the aniline dyes cochineal was of enormous importance as a dye stuff,more than 6,000,000 poundsvalued at about $4,000,000 beingproducedin 1868 by the Canary Islands alone. Little use of it is found at presentsince the dye is not permanent.
CANAL ZONE:

in I9I3. Bas Obispo, Verner

2.

NOPALEA

DEJECTA (Salm-Dyck) Salm-Dyck, Cact. Hort. Dyck. 1849:64. 1850; Britton& Rose, in Carnegie Inst. Wash. Publ. No. 248. 1:37. 1919.

OpuntiadejectaSalm-Dyck, Hort.Dyck. 361. 1834. Shrubs 1-2 m. tall, the old trunks stoutly cylindrical,very spiny, the young stem joints narrowlyelliptic-oblong, to 2 din. long, bright green,the areoleswith 2-8 stout spines. Flowers dark red to pink, the perianthsegmentserect or nearly so, the outer deltoid, 0.5-1.0 cm. long, the inner broadly oblong-obovate,1.5-2.0 cm. long, the hypanthiumnaked, broadly campanulate,about 1 cm. long and 0.7 cm. broad at the orifice;staminal filamentsdark red to pink, about 4 cm. long; (74)

1958]

FLORA

OF PANAMA

(Cactaceae)

75

concave,2.0-2.5 cm. long, 1.0-1.5 and deeply turbinate, truncate ovarybroadly or with 1-2 shortslender the areolesunarmed tuberculate, cm. broad,scarcely 3-4 cm. longand broad,deepredwhenripe. turbinate, spines. Berry as "perhaps suggested America, and escapedin tropical cultivated Commonly of during the preparation and Rose. Nevertheless, nativein Panama"by Britton Panamawas discovered. from specimen herbarium accountonlya single thepresent
CANAL ZONE:

between Empire and Culebra, Cowell 4i8a.

3. OPUNTIA [Tourn.] Mill.


OPUNTIA

[Tourn.]Mill. Gard.Dict. ed. 4. 1754.

Tephrocactus Lem. Cact. 88. 1868. Ficindica St. Lag. in Ann. Soc. Bot. Lyon 7:70. 1880.

Lem.Rev. Hort. 1862:174. 1862. Consolea

and jointedwhen phylloid stems and subshrubs; Succulent shrubs, smalltrees, in our species, linguiform thejointsfleshy, and oval to broadly compressed young, unveryrarely and spines, glochids, withhairs, armed areoles amphigenous bearing acicular and fugacious. Flowerssessile,chiefly armed. Leaves inconspicuous, witha shortand campanulate, broadly upon theyoungjoints;perianth marginal and less shorter theouterprogressively numerous, broadhypanthium, thesegments the filaments verynumerous, stamens widelyspreading; petaloidthanthe inner, glandular levelsintoa shallow at different united muchshorter thantheperianth, ovary uponstimulation; and inflexed sensitive deflexed at anthesis, cup, somewhat longerthan styleslightly prominent; concave,the areoles turbinate to cylindric, seeds. withnumerous berry to fusiform.Fruita fleshy terete thestamens, 0. but some,particularly to the Americas, all indigenous Verymanyspecies, of theEastand subtropics in thedrytropics and escaping introduced ficus-indica, pestin many a serious hasmadethem growth exuberent where their ernHemisphere species, is mostdiversified various and frequent, where thegenus areas. In Mexico, underthe nameof nopal 0. ficus-indica and 0. tuna,are cultivated particularly in tunasare marketed calledtunas, eatenas a delicacy. Although and thefruits, berries insipid rather the largeusually in thesouthwest, chiefly theUnitedStates, muchpopularity. havenot attained
1. OPUNTIA
ELATIOR

Inst.Wash.Publ.No. 248. 1:153. 1919.

Mill. Gard. Dict. ed. 8. 1768; Britton& Rose, in Carnegie

Cactus nigricansHaw. Misc. Nat. 187. 1803. Cactus elatior (Mill.) Willd. Enum. Hort. Berol. Suppl. 34. 1813. Cactus tuna I8 elatior (Mill.) Sims,loc. cit. sub pI. I557. 1813. Cactus tuna y nigricans (Haw.) Sims, in Curt. Bot. Mag. 38: pl. I557.

1813.

obovate to 5 m. tall,theyoungstemjointsnarrowly shrubs Denselybranched the areoles2-4 cm. apart,minutely to 4 dm. long,olive-green, to suborbicular, spines1-7 cm. brownblack-tipped hairyand with 1-8 unequal,terete-acicular deep yellowmoreor less tingedwith long. Flowersabout 5 cm. in diameter, about2 cm. long; staminal mucronate, theinner obovate, segments redor salmon, (75)

76

[Vol. 45

ANNALS

OF

THE

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BOTANICAL

GARDEN

filaments 1.5-2.0

cm. long, deeply tinged with red or salmon, yellow toward the

base; ovary obovoid-turbinate, about 2 cm. long and broad, the areolesunarmedor nearlyso. Berryobovoid, truncate,4-5 cm. long, deep red when ripe. Original habitat unknown; said to be frequentin Curagao, coastal Colombia, known stationsin Panama at present and Panama (Britton& Rose), but definitely are limitedto the islands of the Gulf of Panama. TaboguillaIsland,Howe 3792; San JoseIsland,on rocksof beach,ErlanPANAMA:
son I7I; locality indefinite, 207. Christopherson

4. ACANTHOCEREUS

(Engelm. ex A. Berger) Britton & Rose

ACANTHOCEREUS (Engelm. ex A. Berger) Britton & Rose, in Contr. U. S. Nat. Herb. 12:432. 1909; in CarnegieInst. Wash. Publ. No. 248. 2:121. 1920. in Ann. Rep. Missouri Bot. Gard. 16: Acanthocere-us Cereus subsect. Engelm. ex A. Berger,
75. 1905.

Succulent erect or clamberingshrubs; stems phylloid and jointed, the joints elongate, rooting adventitiouslyat the tips, 3- to 5-angled, the areoles marginal on the angles,shortly spiny. Leaves veryinconspicuous, pubescentand prominently acicular and fugacious or apparentlywholly lacking. Flowers nocturnal, sessile, with an elongate tube, borne singly at the areoles; perianthlarge, subsalverform, shorterand narrowerthan the segmentsvery numerous,the outer progressively the inner,widely spreading,the tube bearing several minute caducous bracts and prominentpubescent and spiny areoles; stamens very numerous, the filaments somewhat shorterthan the perianth,united at progressively deeper levels to the the areoles numerousand crowded,pubescent hypanthium;ovary cylindric-ovoid, and spiny; style filiform,terete, somewhat shorter than the outer and upper stamens. Fruit a fleshy berrywith numerousseeds. A genus of about a dozen species,most frequentin Central America. A single species is recordedfromPanama. 1. ACANTHOCEREUS PENTAGONUS (L.) Britton & Rose, in Contr. U. S. Nat. Herb. 12:432. 1909; in Carnegie Inst. Wash. Publ. No. 248. 2:123. 1920.
Cactus pentagonusL. Sp. P1. 467. 1753. Cactus pitajaya Jacq. Enum. P1. Carib. 23. 1761. Cereus pentagonus (L.) Haw. Syn. P1. Succ. 180. 1812. Cactus prismaticus Willd. Enum. P1. Suppl. 32. 1813. Cereus prismaticus(Willd.) Haw. Suppl. P1. Succ. 77. 1819. Cereus pitajaya (Jacq.) DC. Prodr. 3:466. 1828. Cereus acutangulus Otto, in Pfeif. Enum. Cact. 107. 1837. Cereus princepsPfeif. loc. cit. 1837. Cereus variabilisEngelm. in Boston Journ.Nat. Hist. 5:205. 1845, non Pfeif. Cereus nitidusSalm-Dyck, Cact. Hort. Dyck. 1849:211. 1850. Cereus vasmeriYoung, Fl. Tex. 276. 1873.

Cereus dussii Kakteen K. Sch. Gesamtb. 89. 1897. in Bull. Mus.Hist. Nat. Paris10:384. 1904. Cereus sirul Weberex Gosselin, Clambering terrestrial shrubs,erect when young, soon arching and rooting at the tips of the stems and formingratherextensive clones, the young stem joints (76)

1958]

FLORA

OF PANAMA

(Cactaceae)

77

Fig. 23. Acanthocereus Pentagonus

(77)

[Vol. 45

78

ANNALS

OF THE

MISSOURI

BOTANICAL

GARDEN

elongate,ratherslender,3- to 5-angled, the areolesmarginalon the angles, shortly pubescentand with few to several stout straightspines. Flowers very showy,the perianth tube 15-20 cm. long, about 5 mm. in diameter at the base, abruptly dilated near the orificeinto a shortobconic throatabout 4 cm. in diameter,armed with rather sparse spiny areoles, the lobes widely spreading,white or the outer segmentsmore or less suffused with greenishred, the inner oblanceolate,acute to acuminate, 4-7 cm. long; ovary cylindric-ovoid,truncate, about 2 cm. long, densely armed with spiny areoles. Berry oblong-ovoid, 4-5 cm. long, deep red when ripe,persistently spiny. Forming thicketsat the marginsof beaches from the keys of Florida and the coast of Texas southwardto Colombia and Venezuela; sporadicallyintroducedin the Antilles and perhapsindigenousin Guadeloupe.
Woodson, Allen & Seibert i490, Maxon d Valentine 6967; Bella Vista, Killip in I9ii; Gorgona Beach, G. WhiteI54. Panami, Hitchcock
CANAL ZONE:

vicinity of Miraflores Lake, G. White I45.

PANAMA:

Isla Taboga,
120I5;

Old

5. HYLOCEREUS
HYLOCEREUS

(A. Berger) Britton & Rose


1920.

(A. Berger) Britton & Rose, in Contr. U. S. Nat. Herb. 12:428.

1909; in Carnegie Inst. Wash. Publ. No. 248. 2:183.

Cereus subgen. Hylocereus A. Berger,in Ann. Rep. MissouriBot. Gard. 16:72. 1905.

Succulent shrubby root-climbers,frequently epiphytic; stems phylloid and jointed, rooting adventitiously at the nodes, the joints elongate, usually 3-angled, the areoles marginal on the angles, shortly pubescent and with infrequent and inconspicuous spines. Leaves inconspicuous and fugacious or wholly lacking. Flowers nocturnal,sessile,borne singly at the areoles; perianthlarge, broadly inthe segments the outer progressively fundibuliform, verynumerous, shorter and less petaloid than the inner,the tube ratherbroad and somewhatshorterthan the segments, bearing rather few conspicuous, persistentfoliaceous bracts but without well-defined areoles; stamensvery numerous,the filamentsshorterthan the perianth, united at progressively deeper levels to the hypanthium; ovary cylindricovoid, with few to numerouspersistentaccrescent foliaceous bracts but without well definedareoles; style filiform, somewhat longer than the stamens. Fruit a fleshy berrywith numerousseeds. About 20 speciesof Central America,northern South America,and the Antilles. Several speciesof Hylocereus are cultivated since they are amongstthe most handsome of the NightbloomingCereusesand some have escaped and naturalizedin the tropicsof the Old World.
a. Areoles unarmed or withfewand inconspicuous tubeand spines; perianth lobesdichotomous bracts;stigma ................... 1. H. ovarywithfewand distant tube and ovarywith aa. Areoleswith severalshortstout spines;perianth numerous imbricate lobesentire ..2. H. bracts;stigma
MONACANTHUS POLYRHIZUS

a sterilespecimen collectedby Cowell in Panama to H. triangularis but remarks (Brittonrefers "probablynot native there,however." The stem joints of Cowell's specimen, which we have are generally to thoseof H. polyrhizus but bear 8-10 spinesat the areolesrather similar examined, thanthe3-5 usualforthelatter species.)

(78)

1958]

FLORA

OF PANAMA

(Cactaceae)

79

11/

I'~~~I

Fig. 24. Hylocereusmnonacanthus

(79)

80 1.

[Vol. 45 ANNALS
HYLOCEREUS

OF THE

MISSOURI

BOTANICAL

GARDEN

MONACANTHUS

(Lem.) Britton& Rose, in Carnegie Inst. Wash.


1845.

Pubi. No. 248. 2:190.

1920.

Cereus monacanthusLem. Hort. Univ. 6:60.

Succulent shrubby root-climbers,usually epiphytic, the young stem joints elongate,ratherstout,usually sharply3-angled,the areolesmarginalon the slightly undulate angles, very minutelypuberulentand occasionallywith 1 to few inconspicuous spines. Flowers very showy, the perianth tube 12-15 cm. long, about 1 cm. in diameterat the base, abruptlydilating to a broadlyobconic throatabout foliaceous bracts 1-2 cm. 7 cm. long and broad, bearingfew to several persistent long, the segmentsgradually spreading,greenishwhite, the inner rather broadly 12-15 cm. long; ovary cylindric-ovoid,1.5oblanceolate,subcaudate-acuminate, 2.0 cm. long, persistentfoliaceous bracts 0.2-1.0 cm. long, accrescent; stigma about 8 cm. long, unarmedsave for the lobes dichotomous. Berrycylindric-ovoid, marcescentbases of the bracts. Epiphyticin thicketsand forestedriverbanks, Panama and Colombia.
CANAL ZONE: Gatu'n, Maxon 4689; vicinity of Summit, Bartlett & Lasser I7023; east of Gualaca, alt. 500 ft., Allen CHIRIQUf: Alhajuela, Chagres valley, Pittier in i9ii. Pinogana, Pittier 6653. PANAMA: Rio Tatare, 504I; Isla Parida, Maxon 4904. DARIEN: Woodson d Schery I02I; Bella Vista, Killip 12030; Tapia River, Juan Diaz region,Maxon & Harvey 6705; Urava Island, Howe 3I778; Taboga Island, Gaillard 2.

(Weber) Britton & Rose, in Carnegie Inst. Wash. 2. HYLOCEREUS POLYRHIZUS Publ. No. 248. 2:185. 1920.
Cereus polyrhizusWeber, in Schum. Gesamtb. Kakteen 151. 1897. Cereus trigonusvar. costaricensis Weber, in Bull. Mus. Hist. Nat. Paris 8 :457. 1902. Hylocereus costaricensis(Weber) Britton& Rose, loc. cit. 186. 1920. Cereus costaricensis(Weber) A. Berger,Kakteen 122. 1929.

Succulent shrubbyroot-climbers, frequentlyepiphytic,the young stem joints elongate, usually sharply 3-angled, the areoles marginal on the entire or slightly undulate angled, armed with several short stout spines. Flowers very showy, the perianthtube ratherstout, 1.0-1.5 dm. long, about 1 cm. in diameterat the base, gradually dilating to a broadly obconic throat about 6 cm. in diameter at the orifice,bearing numerouspersistentfoliaceous bracts particularlyimbricatingtoward the base, the segmentsgradually spreading,greenishwhite, the outermost tingedwith purple,the innerratherbroadlyoblanceolate,acuminate,about 1.5-2.0 foliaceousbracts dm. long; ovary cylindric-ovoid, about 4 cm. long, the persistent numerous and imbricate, 1-2 cm. long; stigma lobes entire. Berry oblongoid, about 10 cm. long. Colombia and Panama. "A common species on barren rocks, islets and cliffs especiallyon the along the coast [of San JoseIsland], and frequentas an epiphyte, branches of Bombax quinatum in the interiorof the island [I. M. Johnston,in Sargentia 8:217. 1949]."
PANAMA: San Jose Island, Perlas Archipelago, Johnston I126, Panama, Rose & Rose i8506.

Erlanson

I72;

Old

(80)

1958]

FLORA OF PANAMA

(Cactaceae)

81

andH. costaricensis andRosedistinguish H. polyrhizus BothBerger andBritton to Britton & Rose) of thestem joints, whether (according solely uponthethickness The short descripteonly 3 to 4 cm. thick"or "5 to 10 cm. thick,"respectively. almostsolelyin the similar, differing tionsof Britton and Rose are remarkably comments, partly Dr. Johnston's position of identical adjectives. In thisregard, "The plantvariesin its formand appearance quotedabove,are verysuggestive: it may be habitat. On flat exposedrock surfaces according to its particular stout,decumbent or of short, compactin growth, 1-3 dm. tall, and composed or fromthe fromhighbranches ascending stems3-6 cm. thick. Stemsdangling several meters or even topsof cliffs usually areonly1-3 cm. thickbutmaybecome as muchas 10 cm. on branches becomeflattened, morein length. Stemsrooted about the support. The planthas typically broad,and tendto wrap themselves it 4-angled onesmaybe found." Withsuchtestimony, 3-angled stems, but rarely are conspecific. thatH. polyrhizus and H. costaricensis appears entirely probable 6. WEBEROCEREUS Britton & Rose
WEBEROCEREUS Britton & Rose, in Contr. U. S. Nat. Herb. 12:431.

Carnegie Inst.Wash.Publ.No. 248. 2:214. 1920.

1909; in

stemsphylloidand Succulentshrubby root-climbers, frequently epiphytic; at thejoints, thejointselongate, rather indefinitely adventitiously jointed, rooting marginal on thewings, terete or flattened, the areoles winged, occasionally nearly with extremely few to essentially naked,seldom veryinconspicuously puberulent and inconspicuous lacking. Flowers nocturnal, wholly spines. Leavesapparently areoles, very bornesingly at theuppermost sessile, of mediocre size in our species, the tubeverybroadand somewhat shorter thanthesegbroadly infundibuliform, bractsand conspicuous numerous broadfugacious ments, bearing few to rather and hairs, the segments bristles areoles armed withnumerous slender subarachnoid and less petaloidthan the inner, the outerprogressively shorter verynumerous, inserted thantheperianth, thefilaments shorter stamens numerous, spreading; very at progressively ovary cylindric, bearing deeperlevelswithinthe hypanthium; armed withnumerous subarachnoid and areoles persistent crowded bracts fugacious thanthestamens.Fruita fleshy somewhat bristles and hairs;style filiform, longer seeds. withpersistent and numerous bristly areoles berry of Panamaand Costa Rica. Threeor fourquestionably distinct species
1. WEBEROCEREUS PANAMENSIS Britton & Rose, in Carnegie Inst. Wash. Publ.

No. 248. 2:215. 1920.

4-7 mm. the youngstemjointselongate, Succulent root-climbers, epiphytic flat, the areoles angled or winged and occasionally thick, ratherindefinitely to naked. puberulent indented, very inconspicuously marginal, inconspicuously at the theperianth tube2-3 cm. long,about4 mm.in diameter Flowers mediocre, at the to an obconicthroatabout 1 cm. in diameter dilating base,immediately deltoidbractsand areolesarmedwith slender orifice, bearingseveralfugacious

(81)

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pale suberect or slightly spreading, subarachnoid bristles and hairs, the segments occasionally yellowish orange tinged withpink,apparently white, oblong-elliptic, truncate, about 1 cm. long,bearing 1.5-3.0 cm. long; ovaryoblong-turbinate, broadly subarachnoid-bristly areoles.Berry minute fugacious bracts and persistent red when bright 2-3 cm. long,veryprominently tuberculate, obovoid-turbinate,
ripe.

Fig. 25. Weberocereus panamensis

Lowland streamside Panama. forests and thickets,


Allen 882; COLON: nearmouthof Rio Chagres, CALITY UNKNOWN: Hunter I7040, I704I. 3903. alongRio Fato,Pittier
LO-

4-angled of olderstemswhichare sharply They are quite sterile, however, consisting it is as a rule. Althoughwe believe that theyprobablyrepresent W. panamensis stem possibleof course that such actually may not be the case. The flowering which are beforeus are 2- to 3-angled. joints of W. panamensis 7. EPIPHYLLUM
EPIPHYLLUM

as Selenicereus inermis (Otto) B. & R. trees"have been determined previously

The Hunter numbers, with only the comment"In long strandsclimbingover

[Hermann] Haw.

[Hermann] Haw. Syn. PI. Succ. 197. 1812; Britton & Rose, in CarnegieInst. Wash. Publ. No. 248. 4:185. 1923.

PhyllocactusLink, Handb. Erkenn.Gewichse 2:10. 1831.

Nat. Bruxelles 112. 1839. Phyllocereus Miq. in Bull.Sci. Phys. stemsphylloidand Succulent shrubbyplants,usually epiphyticroot-climbers; usutypes: the primarytereteor subterete, jointed,usually of two intergrading and leaf-like,lobed or undulate, ally rooting,the secondary(flowering)flattened the areolesmarginal, naked or rarelyminutely puberulent. Leaves lacking. Flowers nocturnal or ephemeral,sessile,borne singly at the areoles; perianthusually the outerprogressively the segments shorter verynumerous, large,infundibuliform, the tube relativelyslender, or spreading, and less petaloid than the inner,reflexed bearing usually few and sparse,inconspicuous usually longer than the segments,

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the filaments very numerous, areoles;stamens bracts,usuallywithoutdefinite levelsto the deeper at progressively and united thantheperianth shorter somewhat naked or to few numerous with cylindric-ovoid, ovaryfrequently hypanthium; the than longer somewhat areoles; style filiform, puberulent rarelyminutely seeds. withnumerous berry stamens. Fruita fleshy in Cenconcentration withthegreatest America, of tropical About20 species cultivated Cereususually of Nightblooming tralAmerica. Theseare the species of most pattern growth as houseplantsin the UnitedStates. A characteristic rooting or less terete, more of the stemsinto is the differentiation Epiphyllums flowers the which joints upon secondary leaf-like jointsand flattened, primary secondary the upon regularity withgeometric appear areborne. Areoles eventually upon the primary.The two basic typesof joint may be but are erratic joints, vice versa,a prifroma primary, proceeding in anyway: a secondary combined a secondary. from or a secondary a primary, from mary
tube 6 or moretimes the perianth and slender, a. Flowersveryelongate narrowjointsrather stem (flowering) secondary thanthesegments; longer the areolesnaked in dessication, thickand callose-marginate ly serrate, ................-.....................-............-....-..-.-...... 1. E. or indefinitely papillate tube2-3 (or less) times theperianth broadand massive, usually aa. Flowers thanthesegments. longer about 8-15 cm. long,the smalland inconspicuous, relatively b. Flowers stem secondary tube 2-3 timeslongerthan the segments; perianth in dessicathickand callose-marginate serrate, narrowly jointsrather 2. E. ............. nakedor indefinitely papillate tion,the areoles tubelessthantwiceas long theperianth largeand handsome, bb. Flowers as thesegments. slender stemjointsrather about 15-25 cm. long: primary c. Flowers and terete. thinin dessication, serrate, narrowly stemjointsrather d. Secondary about 15-20 cm. long. 3. E. flowers the areoles puberulent; minutely dd. Secondarystem joints deeply crenate,the areoles minutely ..................................... 4. E. about 20-25 cm. long flowers puberulent; stem joints very stout and cc. Flowersabout 40 cm. long; primary 5. E. stemjointsunknown) ... or angled(secondary moreor lesswinged

PHYLLANTHUS

PITTIERI

LEPIDOCARPUM MACROPTERUM GIGAS

& 1. EPIPHYLLUM PHYLLANTHUS (L.) Haw. Syn.P1. Succ. 197. 1812; Britton 1923. 4:187. 248. No. Publ. Wash. Inst. in Carnegie Rose,
(L.) DC. Prodr. 3:469. 1828. Cereus phyllanthus Gewachse 2:11. 1831. (L.) Link,Handb.Erkenn. phyllanthus Phyllocactus (L.) K. Sch. in Mart.Fl. Bras.42:298. 1890,in part,as to basinym. phyllanthus Rhipsalis U. S. Nat. Herb. 16:240. 1913. Britton& Rose,in Contr. gaillardae Epiphyllum 23:87. 1913. Kakteenk. & Rose) Vaupel,in Monatsschr. (Britton gaillardae Phyllocactus Succulent epiphyticshrubs,the primarystem joints terete,elongate, about 5 mm. thick when young, probably thicker when old, the secondary stem joints usually obtuse to rounded at the tip, narrowlyoblong-ellipticto linear-elliptic, cuneately decurrentinto a terete subpetiolarbase, 3-5 dm. long, 2-8 cm. wide, in dessication,the areoles 3-5 rathernarrowlyserrate,thick and callose-marginate upon the upper papillate, borne superficially cm. distant, naked or indefinitely white, broad marginof the serrations. Flowersnocturnal,veryelongateand slender, (L.) Opuntia phyllanthus

L. Sp. PI. 469. 1753. Cactusphyllanthus

Mill. Gard. Chron. ed. 8. 1768.

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Fig. 26. Epiphyllum phyllanthus

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Porto COLON: Williams 423. northof El Valle, alt. 1000 m., Allen 2334; Penonome, headwaters of Rio Chico, alt. 500-750 ft., Allen 4609. DARIEN: Bello,Pittier in I9I2. of Pacora,alt. 40 m., Allen224I; JuanDiaz, Maxond Harvey6706. vicinity PANAMA: 2.
EPIPHYLLUM PITTIERI

about 3 mm. in ditubular, tube about 18-25 cm. long,narrowly the perianth shortconic throatabout dilatedinto an extremely at the base,abruptly ameter external inconspicuous and distant few very bearing 8 mm.broadat the orifice, spreading, acuminate, verynarrowly oblong-elliptic, segments theperianth bracts, tuberculate. inconspicuously the innerabout 3-4 cm. long,the ovaryoblongoid, 7-8 cm. about tuberculate-angulate, somewhat truncate, oblong-turbinate, Berry ripe. when red bright longand 2 cm. broad, Costa Rica to Brazil,Boliviaand Paraguay. MaddenLake,Dodge & Allen i7o56. COCLE: hills QuebradaCulebra, CANAL ZONE:

(Weber) Britton & Rose, in Contr. U. S. Nat. Herb.

16:258. 1913. in Bois,Dict. Hort. 2:957. 1898. Weber, pittieri Phyllocactus Succulent epiphyticshrubs,the primarystem joints tereteor weakly angulate, 5-10 mm. thick when young, probably thicker when old, the secondary stem acute to obtuse at the tip, decurrentto a short terete subjoints linear-elliptic, petiolar base, about 2-10 dm. long and 2-6 cm. broad, rathernarrowlyserrate, in dessication,the areoles 3-6 cm. distant, naked or thick and callose-marginate upon the upper broad marginof the serrapapillate,borne superficially indefinitely tions. Flowers nocturnal,relativelysmall and inconspicuous,white, the perianth tube about 6-10 cm. long, narrowlytubular,about 4 mm. in diameterat the base, short conic throat about 7 mm. broad at the abruptlydilated into an extremely bearingfew and distantinconspicuouspetaloidbracts,the perianthsegments orifice, linear-elliptic,acuminate, spreading, the inner about 4-5 cm. long, the ovary bracteate and tuberculate. Berryoblongoid, narrowlyoblongoid,inconspicuously about 2 cm. long, deep red when ripe. Costa Rica and Panama. WaterValley,Vol of ChiriquiLagoon,Von WedelI294; vicinity BOCAS DEL TORO: Bay,Von Wedel7; GarayCreek, WedelI528, I58o, I768A; QuebradaNigua, Almirante Von Wedel 2630. CANAL ZONE: DraytonTrail, BarroColoradoIsland,Shattuck603. DavidsonII89. Puerto Armuelles, forest, CHIRIQUI: 3. EPIPHYLLUM LEPIDOCARPUM (Weber) Britton & Rose, in Contr. U. S. Nat. Herb. 16:257. 1913. in Bull.Mus.Hist. Nat. Paris8:462. 1902. Weber, lepidocarpus Phyllocactus Succulent epiphyticshrubs,the primarystem joints weakly 3-angled, up to 1 cm. thick when young,becoming tereteand thickerwhen old, the secondarystem joints oblong- to narrowlyovate-elliptic,obtuse at the tip, decurrentto a very short weakly angled subpetiolarbase, about 2-7 dm. long and 3-7 cm. broad, the thin and more or less venose in dessication, remarkably rathernarrowlyserrate, upon the upper areoles 4-5 cm. distant,minutelypuperulent,borne superficially

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broadmargin of the serrations.Flowers nocturnal, rather broadand handsome, whitewiththeoutersegments flushed withrose, theperianth tubeabout7-8 cm. long,tubular, about4 mm.in diameter at thebase,rather gradually dilating to a narrowly conicalthroat about 1.5 cm. broadat theorifice, bearing a fewinconspicuous deciduous linear bracts subtending puberulent areoles toward thebase,the segments 7-8 cm. long, the outerlinear,flushed with rose,the innernarrowly obovate-elliptic, white,the ovarynarrowly turbinate, bearing crowded persistent ovatebractsand conspicuous puberulent areoles. Berry turbinate-fusiform, occasionally conspicuously beaked,6-9 cm. long, 2-4 cm. thick,coveredwith the accrescent bractsand puberulent hyaline areoles, purplish pinkwhenripe. HighlandCosta Rica and western Panama,epiphytic or upon rocks. 1900m.,Seibert 206; BajoChorro, Boquete District, alt.6000ft., Davidson 253. A species particularly noteworthy becauseof its peculiar scalyfruits. 4.
EPIPHYLLUM MACROPTERUM
CHIRIQUI:

valleyof the upperRio ChiriquiViejo,vicinity of MonteLirio,alt. 1300-

Publ.No. 248. 4:193. 1923.

(Lem.) Britton & Rose,in Carnegie Inst.Wash.

Phyllocactusmacropterus Lem. Illustr. Hort. 11: Misc. 73. 1864. Phyllocactus thomasianus K. Sch. in Monatsschr.Kakteenk. 5:6. 1895. Phyllocactus costaricensis Weber, in Bull. Mus. Hist. Nat. Paris 8:463. 1902. PhyllocactusmacrocarpusWeber, loc. cit. 464. 1902. Epiphyllum costaricense (Weber) Britton & Rose, in Contr. U. S. Nat. Herb. 16:256. 1913. Epiphyllumthomasianum(K. Sch.) Britton& Rose, loc. cit. 259. 1913.

the primary or weakly Massivesucculent stemjointsterete epiphytic shrubs, in age,the and 5-8 mm.thick whenyoung, and terete stouter 4-angled becoming secondary stemjointsnarrowly obovate-to oblong-elliptic, obtuseto rounded at thetip,broadly decurrent to a short weakly angled subpetiolar base,about6-8 dm. thinand rather and regularly long and 5-10 cm. broad,rather broadly crenate, in dessication, theareoles venose 2-5 cm. distant, conspicuously callose-marginate, thedeepindentations borne within of thecrenations. minutely puberulent, Flowers the perianth nocturnal, verylargeand handsome, tube 12-15 cm. long,tubular, to a conicalthroat about6 mm.broadat thebase,gradually about arcuate, dilating rather 3 cm. broadat theorifice, pale greenish rose, bearing conspicuous persistent the subfoliaceous about 8-12 cm. long,slightly bracts,the segments spreading, outerlinear-lanceolate, pale greenish rose,theinnerobovate-elliptic, white;ovary bracts. Berry about 3-4 cm.long,bearing oblongoid, manysubfoliaceous oblongfusiform, usuallybeaked,about 10-12 cm. long and 4-5 cm. thick,slightly red whenripe. tuberculate at theminutely puberulent areoles, and upon rocks,apparently Costa Rica and Panama,epiphytic usuallyat low altitudes.
of ChiriquiLagoon,Von Wedel1I64; Snapper BOCAS DEL TORO: vicinity Point,Von Wedel2639; WaterValley,Von Wedel736; Shepherd Island,Von Wedel 2722. CHIRIDavidson QuI: Puerto Armuelles, I190.

An extremely showy plant,but surpassed by theone whichfollows:

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5.

EPIPHYLLUM

gigas Woodson & Cutak, spec. nov.

Frutices succulenti scandentesradicantes,ramis ut dicitur ultra pedes centum in longitudinemmetientibus; articulis rami radicantibus validissimisancipitibus leviter undulatis nostris 2-3 cm. crassis; articulis florigeris infeliciterignotis. Flores maximi speciosissimi ut credituralbi noctiflori;perianthiitubo ca. 22-25 cm. longo basi ca. 7 mm. lato apicem versusgradatimdilatato ostio ca. 2.5-3.0 cm. linediam. extus bracteas linearispaucas gerente; perianthiisegmentis exterioribus aribus longe acuminatis 15 cm. longis vel ultra patulis, interioribusne latioribusin exemplonostrovix bene visis; ovario late cylindricoca. 1.5 cm. longo inconspicue bracteato areolas parvas paucas minute puberulentes gerente. Bacca ignota. Species inter species generisjam cognitas omnibus partibus longe maxima. 20, 1946,P. H. Allen3772 (Herb. Missouri Bot.Gard.,HOLOTYPE).
PANAMA:

Cerro Trinidad, alt. 800-1000 m., climbing rock faces of main peak, Oct.

This is a tremendous species,far surpassing in size of all parts any otherknown Epiphyllum; as such it is impossibleto ignore, even though one quite naturally would desirethe flowering stem joints and fruitsfor a completedescription. The ratherregularundulationsof the primary stem joints and the puberulentareolesof the ovary suggestan affinity with E. macropterum and E. oxypetalumft. Mr. Allen's statementthat the holotypespecimenwas a "giant liana, to several hundred feet in length" might seem totallyincrediblewere it not for the gigantic flowerwhich he preserved. 8. WITTIA K. Sch.
VITTIA K. Sch. in Monatsschr. Kakteenk. 13:117.

1903; Britton & Rose, in

Carnegie Inst. Wash. Publ. No. 248. 3:206. 1923.


Pseudorhipsalis Britton& Rose, loc. cit. 4:213. 1923.

Succulent shrubbyroot-climbers, usually epiphytic;stemsphylloidand jointed, usually of two intergrading typesas in Epiphyllum: the primarytereteor weakly winged, usually rooting, the secondary (flowering) flattenedand leaf-like,lobed or undulate, with marginal areoles covered with few to several minute scales and indefinitely papillate or naked. Leaves lacking. Flowers ephemeral (or nocturnal?), sessile,borne singly at the areoles; perianthmediocreor relativelysmall, tubular to tubular-salverform or campanulate,the tube about as long as the segments or shorter, bearing a few inconspicuousdeltoid bracts, the segmentsabout few to severaltimes 10-15, erect or spreading;stamensabout 30-45, the filaments insertedat progressively longer than the anthers, deeperlevels within the perianth tube, the outermost conspicuously shorter than the innermost; ovary ovoidfusiform, minutely tuberculate-bracteate. Fruit a fleshyberry with rather few seeds. It is extremely difficult to justifythe separationof Pseudorhipsalis from Wittia upon the basis of our scanty herbariummaterial,and surelyit is nearlyimpossible to do so even from the meagre descriptionsand rather irrelevantcomments of

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between the subthat there is an obvioustransition Britton and Rose. Granted as wellas thatof thepurpose of illustration tribes Epiphyllanae and Rhipsalidanae, recognized better by fewer gradient genera thanthose convenience wouldbe served by Britton and Rose. two species each for Wittiaand Pseudorhipsalis. Britton and Rose enumerate In addition in Panama,W. amazonica is knownfromextreme to the two species Jamaica. and Ps. alata from northeastern Peruand adjacentColombia,
with rose,the perianth a. Flowerstubular-salverform, whiteflushed segments stemjointsnarrowly ................. 1. W. repand widely spreading; flowering the perianth aa. Flowerstubular, segments deeppink tippedwithlavender,
erect; floweringstem joints narrowly serrate............................................. 2. W.
HIMANTOCLADA PANAMENSIS

1. WITTIA himantoclada (Roland-Gosselin) Woodson,comb.nov. himantoclada in Bull. Soc. Bot. France55:694. 1908. Rhipsalis Roland-Gosselin, & Rose,in Contr. Wittiacostaricensis Britton U. S. Nat. Herb. 16:261. 1913. Britton & Rose, in CarnegieInst. Wash. Pseudorhipsalis himantoclada (Roland-Gosselin) Publ.No. 248. 4:213. 1923.

the primary Succulentepiphytic stemjointstereteor weakly root-climbers, lanceolate or stemjointsnarrowly winged, up to about 1.5 m. long,thesecondary at thetip,cuneately obtuseto rounded oblanceolate, 2-5 dm. long,1-5cm. broad, in dessithickand coriaceous base,narrowly repand, decurrent intoa subpetiolar whiteflushed cation,the areoles1-2 cm. distant. Flowerstubular-salverform, theperianth tubeabout 5-6 mm.long,about 3 mm. withroseor orange without, in diameter at thebase,the perianth ellipticto linear, segments 10-15, narrowly stamens about 45, about 1 cm. long and up to 3 mm. broad,widelyspreading; exserted about7 mm.long, included; ovarybroadly cylindrical, slightly to barely unknown. deeprose. Berry Costa Rica and Panama.
COCLE:

north rimof El Valle,AllenI897.

2. WITTIA PANAMENSIS

Britton & Rose, in Contr.U. S. Nat. Herb. 16: 241. Inst.Wash.Publ.No. 248. 4:207. 1923. 1913; in Carnegie

or upon rocks,the primary Succulent scandent stemjoints shrubs, epiphytic oblongthe secondary stem joints very narrowly tereteor weaklyancipitous, decurrent into a veryshortsubpetiolar lanceolate, rounded at the tip, cuneately and closelycuneate-serrate, base, 3-5 dm. long, 3-5 cm. broad,verynarrowly in dessication, the areoles1.0-2.5 cm. distant, naked thickand callose-marginate or indefinitely scales,bornesuperficially papillatebeneaththe minuteimbricate Flowers rather of thenarrow serrations. small, ephemeral, upontheupper margin tube about 1.5 cm. long,about the perianth tubular, pinktippedwithlavender, 2 mm. in diameter the perianth dilatedtowardthe orifice, at the base,scarcely about 10, about 1 cm. long,erector essentially so; ovaryabout 3 mm. segments not seen. tuberculate-bracteate. Berry long,minutely and Venezuela. Eastern Panama;Colombia

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Fig. 27. Wittiapanarnensis

DARIEN: Marraganti, Williams698; PANAMA: Chepo, Pittier SAN BLAS: 457I.

trail betweenPinoganaand Yaviza, Allen 278. Caledonia, Elmore L4o.

Wittia panamensisis distinguishedfrom W. amazonica by Bntton and Rose solely upon the fruit,whether "smooth" or "roughened by small tubercles" respectively. I have seen no fruitof W. panamensisas yet, but the ovaries of the flowerscertainlyare minutelytuberculateand I rather suspect that the fruit of the two speciesmay have much in common. 9. RHIPSALIS Gaertn.
RHIPSALIs Gaertn. Fruct. Sem. 1:137.

1788; Britton & Rose, in Carnegie Inst. Wash. Publ. No. 248. 4:219. 1923, nom. conserv.

HariotaAdans.Fam. 2:243. 1763,nom.rejic. Rather slendersucculent root-climbers, usually epiphytic; stems phylloid and so in our species,the areolesspiral,naked to abundantly jointed,tereteor essentially hirsuteor bristly. Leaves lacking. Flowers verysmall, sessileor somewhatsunken within the stem joints, solitary;perianthrotate,the segmentsfreeor essentially so, relativelyfew, sometimes only 5, usually spreading;stamensabout twice as many as the perianth lobes, the filamentsinsertedon the margin of the hypanthium. Berrysmall, usually white,with ratherfew small seeds. A perplexinggenus of 50 or more species in tropical and subtropicalAmerica; also with severalspeciesdescribedfromthe Old World, at least some of which may be introducedfromAmerica. Known as MistletoeCactus in the United States. Besides the ubiquitous Rh. cassytha there are two very distinct species in Panama which are known only from sterile specimens. Both probably are undescribed,but of course cannot be provided with names until floweringplants are found.

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Fig. 28. Rhipsaliscassytha

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.------------------------ 3. RH. SP. aa. Sterile areoles with conspicuous tufts of spreading bristles

naked. puberulent to essentially minutely areoles a. Sterile naked ................... 1. RH. CASSYTHA b. Flowering to essentially areoles puberulent minutely floccose-hirsute bb. Flowering areoles ............................. 2. RH. SP. veryconspicuously 1788; Britton & Rose, in

1. RHIPSALIS CASSYTHA Gaertn. Fruct. Sem. 1:137.

Carnegie Inst. Wash. Publ. No. 248. 4:225. 1923, as cassutha.


77. 1788. Sw. Prodr. Cactuspendulus Haw. Syn.P1.Succ. 187. 1812. parasitica Rhipsalis (Gaertn.) Lem. Cact. Gen. Nov. Sp. 75. 1839. Hariotacassytha Pendulous, usually epiphyticsucculents up to 9 m. long; stem joints terete, or in false whorls,yellowishgreen,about 2-3 mm. thick when young, dichotomous the areolesvery inconspicuous,glabrous to minutelypuberulent. Flowers lateral, solitary,greenishwhite, the perianthsegmentsabout 2 mm. long. Berryglobose, white (rarelypink), about 5 mm. in diameter. Throughout tropical America, apparentlyalso in Ceylon and tropical Africa. Very frequentin Panama.

Von Wedel649, i670; LittleBocas,Von Wedel2529; BOCAS DEL TORO: WaterValley, of Von Wedel2584; Isla Col6n,Von Wedel2924. CANAL ZONE: vicinity River, Pumpkin MaddenDam and Alahuela,alt. 80-100 m., Hunterl! Allen 787; BarroColoradoIsland, Shattuck447; Bas Obispo, Verners. n.; Gatu'nLake, Rowlee d Rowlee 405; Balboa, 28546. Standley
2. RHIPSALIS SP.

This specieshas stem joints which are about half as thick as is usual for those of Rh. cassytha. The sterileareoles are glabrous or essentiallyso, but the young areolesare coveredwith exceedinglydense tuftsof yellow hairs 2-3 mm. flowering long. If this is a species of seriesFLOCCOSAE, as seems probable, it is the most of it. representative northern
BOCAS DEL TORO:

Von WedelI393. Lagoon, of Chiriqui vicinity

3. RHIPSALIS SP.

a Lycopodium,at firstglance. It is juvenile, apparThis little plant resembles ently,and the stems,which are somewhatthickerthan thoseof Rh. cassytha,have the sterileareolesbeset with conspicuoustufts of spreadingbristles1-2 mm. long. Its generalaspect agreesin general with the illustrationof Rh. aculeata given by Britton& Rose (in Carnegie Inst. Wash. Publ. No. 248. 4: p1.xxiv, fig. 8). On the other hand, a good many Rhipsalis seedlingsappear bristlyor spiny and later more mature growthbecomes almost glabrous.
BOCAS DEL TORO:

Van Wedel2572. River, Pumpkin

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