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Full text of "Flora of Peru"

y OF 

ILLINOIS LIBRARY 

AT URBANA-CHAMPAIGN 

BIOLOGY 

06 1996 



BOTANICAL SERIES 



FIELD MUSEUM OF NATURAL HISTORY 

FOUNDED BY MARSHALL FIELD, 1893 

VOLUME XIII 



FLORA OF PERU 

PART VI 



BY 

J. FRANCIS MACBRIDE 

ASSISTANT CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 



B. E. DAHLGREN 

CURATOR, DEPARTMENT OF BOTANY 
EDITOR 



PUBLICATION 364 



S NATURAL 
HISTORY 




CHICAGO, U.S.A 

SEPTEMBER 18, 1W 



TV,p nerson charging this material is re- 
Ip h ons P iblefor its rlt/rn to the Hbragrfrom 
which it was withdrawn on or before the 
Latest Date stamped below. 



the University. 



L161 0-1096 



BOTANICAL SERIES 



FIELD MUSEUM OF NATURAL HISTORY 

FOUNDED BY MARSHALL FIELD, 1893 

VOLUME XIII 



FLORA OF PERU 

PART VI THE uBRARY OF THE 

OCTS-1936 

UNIVERSITY OF ILLINOIS 
BY 

J. FRANCIS MACBRIDE 

ASSISTANT CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 



B. E. DAHLGREN 

CURATOR, DEPARTMENT OF BOTANY 
EDITOR 



PUBLICATION 364 




CHICAGO, U.S.A. 
SEPTEMBER 18, 1936 



PRINTKD IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 



FLORA OF PERU 



J. FRANCIS MACBRIDE 



RUBIACEAE. Coffee Family 
By Paul C. Standley 

Trees, shrubs, or herbs with stipules; leaves opposite or verticillate, 
entire or rarely pinnatifid; flowers small or large and showy, perfect 
or rarely unisexual, usually regular; calyx tube adnate to the ovary 
(hypanthium), the limb of the calyx usually dentate or lobate, one 
or more of the lobes occasionally enlarged and foliaceous; corolla 
gamopetalous, variable in form; stamens as many as the corolla 
lobes, inserted commonly in the tube or throat of the corolla, the 
anthers 2-celled; ovary 1-10-celled, inferior; fruit 1-10-celled, usually 
baccate, drupaceous, or capsular; seeds 1 to many, small or large, 
often winged. 

The family is represented generously in most tropical regions, 
and it is one of the largest of the Peruvian flora. Most of the Peru- 
vian plants of the family are confined to the eastern slopes of the 
Andes. Those occurring in the temperate regions belong chiefly to 
the genera Galium, Relbunium, and Arcytophyllum. 

Key to the Tribes 

Ovary with 3 or more ovules in each cell. 
Fruit dry. 

Flowers in very dense, globose heads. Plants armed with 
hooked spines, more or less scandent, woody . . I. Naudeeae. 
Flowers not in dense globose heads. 

Seeds winged, vertically imbricate II. Cinchoneae. 

Seeds not winged or, if winged, horizontal. 

Corolla lobes imbricate or contorted, never valvate. 

IV. Rondeletieae. 
Corolla lobes valvate. 

Seeds horizontal; stipules entire or bifid; trees or large 
shrubs III. Condamineae. 

Seeds peltately attached; stipules often fimbriate; plants 
commonly herbs or low shrubs V. Hedyotideae. 

3 



4 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Fruit fleshy. 

Corolla lobes valvate in bud VI. Mussaendeae. 

Corolla lobes imbricate or contorted in bud. 
Seeds many, minute, pitted or rarely tuberculate. 

VII. Hamelieae. 
Seeds usually few and large, the testa smooth or fibrous. 

VIII. Gardenieae. 
Ovules 1 or 2 in each cell of the ovary. 

Ovules 2 in each cell of the ovary. Fruit drupaceous, 5-7-celled. 

IX. Retiniphylleae. 
Ovules 1 in each cell. 
Ovules pendulous. 
Stamens inserted in the throat of the corolla tube. 

X. Guettardeae. 
Stamens inserted at the base of the corolla tube. 

XL Chiococceae. 
Ovules erect or ascending. 

Corolla lobes contorted XII. Ixoreae. 

Corolla lobes valvate. 
Ovules affixed to the base of the ovary. 
Ovary 1-celled, or 2-celled but with a very thin septum. 

XIII. Coussareae. 

Ovary with two or more cells, the septum thick. 
Flowers commonly dioecious; stamens usually inserted 
at the base of the corolla; plants chiefly herbaceous 

and prostrate XVI. Anthospermeae. 

Flowers perfect; stamens inserted usually in the throat 
of the corolla; plants usually trees or shrubs, 
sometimes scandent. 
Fruit drupaceous; plants not scandent. 

XIV. ' Psychotrieae. 
Fruit dry, dehiscent; plants scandent. 

XV. Paederieae. 

Ovules affixed to the septum. Stipules fimbriate or leaf-like; 
plants herbaceous or chiefly so. 

Stipules fimbriate, not leaf -like XVII. Spermacoceae. 

Stipules resembling the leaves and forming whorls with them. 

XVIII. Galieae, 



FLORA OF PERU 5 

I. NAUCLEEAE 
A single genus in Peru 1. Uncaria. 

II. CINCHONEAE 
Corolla lobes valvate. 

Placenta pendulous from the apex of the cell; flowers in elongate 

spikes, the corolla small 2. Alseis. 

Placenta ascending or erect, or adnate to the middle of the septum; 

flowers not spicate. 
Placenta ascending or erect from the base of the septum ; plants 

scandent, chiefly or wholly herbaceous 3. Manettia. 

Placenta adnate to the middle of the septum; plants not 

scandent. 
Leaves varnished, with a metallic appearance. 

4. Stilpnophyllum. 
Leaves not varnished, not with metallic appearance. 

Flowers solitary. A low shrub; corolla with a very long 
and narrow tube 5. Lecanosperma. 

Flowers numerous, in several- or many-flowered inflores- 
cences; trees or large shrubs. 

Capsule splitting from below upward ... 6. Cinchona. 
Capsule splitting from above downward. 

Corolla lobes bifid ; fruit dehiscing by 4 spirally coiled 
valves 7. Joosia. 

Corolla lobes entire; fruit not dehiscent by 4 spirally 
coiled valves. 

Capsule loculicidal 8. Macrocnemum. 

Capsule septicidal. 

Valves of the capsule bifid 9. Remijia. 

Valves of the capsule entire. 

Capsules small, barely 1 cm. long; flowers 

densely clustered 10. Pimentelia. 

Capsules large, commonly more than 2 cm. 
long, often much larger; flowers rather 
loosely clustered and usually pediceled. 

11. Laderibergia. 



6 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Corolla lobes imbricate or contorted. 

Calyx lobes unequal, one of them expanded into a large red limb. 

12. Capirona. 
Calyx lobes equal or nearly so, none of them expanded into a 

colored limb. 

Corolla lobes contorted, or one external in Calycophyllum. 
Calyx caducous; seeds not winged, bearing a tuft of hairs at 

one end 13. Hillia. 

Calyx persistent; seeds winged. 

Plants epiphytic; corolla tube 6-9 cm. long; leaves fleshy. 

14. Cosmibu&na. 
Plants terrestrial trees or shrubs; corolla smaller; leaves 

not fleshy. 
Corolla lobed to the middle or more deeply. 

15. Calycophyllum. 

Corolla with short lobes much shorter than the tube. 
Stamens equal, the anthers included .... 16. Loretoa. 
Stamens unequal, the anthers exserted. 

17. Ferdinandusa. 
Corolla lobes imbricate. 

Corolla regular, with a slender tube 18. Exostema. 

Corolla somewhat zygomorphic, with broad tube. 

19. Coutarea. 

III. CONDAMINEAE 

Calyx lobes unequal, one of them in some of the flowers expanded 

into a large red limb 20. Pogonopus. 

Calyx lobes equal or nearly so, none of them expanded and colored. 

Calyx deciduous; corolla large 21. Condaminea. 

Calyx persistent; flowers small 22, Chimarrhis. 

IV. RONDELETIEAE 

Corolla lobes imbricate. 

Calyx lobes unequal, one of them expanded into a large red limb. 

23. Warscewiczia. 

Calyx lobes equal or nearly so, none of them expanded into a 

colored limb. 
Leaves densely white-tomentose beneath 24. Rondeletia. 



FLORA OF PERU 7 

Leaves not tomentose. 

Flowers in terminal trichotomous cymes. Leaves subsessile; 

corolla 12-15 mm. long 25. Dolichodelphys. 

Flowers in usually large and many-flowered panicles. 

Seeds large; capsule loculicidal 26. Sickingia. 

Seeds small; capsule septicidal 27. Bathysa. 

Corolla lobes contorted. 

Plants low herbs 28. Sipanea. 

Plants shrubs or trees. 
Flowers large, the green corolla almost 5 cm. long. Plants 

glabrate 29. Macbrideina. 

Flowers small, the colored or white corolla less than 2 cm. long. 
Stipules persistent, exuding resin; leaves glabrate. 

30. Elaeagia. 

Stipules caducous, not resinous; leaves densely hairy. 

31. Phitopis. 
V. HEDYOTIDEAE 

Seeds angulate; plants annual 32. Oldenlandia. 

Seeds plano-convex; plants chiefly perennial and often suffrutescent. 

33. Arcytophyllum. 
VI. MUSSAENDEAE 
Inflorescence terminal. 
Flowers arranged in involucrate heads; plants epiphytic. 

34. Schradera. 
Flowers paniculate; plants not epiphytic. 

Panicles spike-like; flowers small 35. Gonzalagunia. 

Panicles thyrsiform; flowers large 36. Isertia. 

Inflorescence axillary. 

Plants creeping herbs. Flowers usually in dense heads. 

37. Coccocypselum. 

Plants shrubs or trees, or sometimes herbaceous and scandent. 
Leaves with many close striolae in the meshes of the veins. 

Inflorescences 1-3-flowered 38. Hippotis. 

Inflorescences few-many-flowered. 

Leaves small and thin; plants much branched. 

39. Sommera. 



8 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves very large, coriaceous; plants chiefly simple or 
nearly so 40. Pentagonia. 

Leaves without striolae in the meshes of the veins. Plants 
scandent 41. Sabicea. 

VII. HAMELIEAE 

Ovary 2-celled 42. Hoffmannia. 

Ovary 4-5-celled. 

Corolla lobes contorted in bud 43. Bertiera. 

Corolla lobes imbricate in bud. 

Corolla tubular 44. Hamelia. 

Corolla short-funnelform 45. Bothriospora. 

VIII. GARDENIEAE 

Corolla somewhat irregular, the buds curved 46. Posoqueria. 

Corolla regular, the buds not curved. 
Flowers perfect. 

Inflorescences terminal or terminal and axillary. 

Flowers in few- or many-flowered cymes 47. Tocoyena. 

Flowers mostly solitary or fasciculate. 

Corolla tube equaling or scarcely exceeding the limb, 
usually shorter 48. Sphinctanthus. 

Corolla tube usually much longer than the limb . 49. Randia. 
Inflorescences lateral. 

Corolla tube villous in both throat and base; stigma tapering; 
testa of the seeds fibrous; trees 50. Genipa. 

Corolla tube villous in throat or base but not in both; stigma 
not tapering; shrubs or small trees. 

Ovary 2-celled; native species .49. Randia. 

Ovary 1-celled; cultivated species 51. Gardenia. 

Flowers dioecious. 

Stipules forming a conic cap, connate into a sheath, deciduous 
above a circular slit, leaving a persistent basal caruncle. 

Staminate flowers in cymes, the pistillate usually solitary, 
rarely 2-3 in a head 52. Duroia. 

Staminate and pistillate flowers both in cymes . 53. Amaioua. 



FLORA OF PERU 9 

Stipules free or united only near the base, not forming a cap 
that is dehiscent by a transverse slit. 

Staminate flowers lateral; plants usually armed with spines. 

49. Randia. 

Staminate flowers terminal ; plants unarmed . . 54. Alibertia. 

IX. RETINIPHYLLEAE 
A single genus in Peru 55. Retiniphyllum. 

X. GUETTARDEAE 
Corolla lobes imbricate in bud. 

Fruit separating into 2 narrow dry cocci 56. Machaonia. 

Fruit drupaceous, indehiscent 57. Guettarda. 

Corolla lobes valvate. 

Tube of the corolla short and thick 58. Malanea. 

Tube of the corolla slender and elongate. 

Corolla lobes corniculate outside at the apex 59. Chomelia. 

Corolla lobes not corniculate 60. Anisomeris. 

XL CHIOCOCCEAE 
A single genus in Peru 61. Chiococca. 

XII. IXOREAE 

Calyx calyculate at the base; cultivated plants 62. Coffea. 

Calyx naked at the base; native plants 63. Ixora. 

XIII. COUSSAREAE 

Seeds vertical; ovules connate, borne on a common basal column. 

64. Coussarea. 

Seeds horizontal; ovules separate in a 1-celled ovary, collateral, 
basilar 65. Faramea. 

XIV. PSYCHOTRIEAE 

Ovary partly or almost wholly superior 66. Pagamea. 

Ovary inferior. 

Flowers in elongate, interrupted spikes, crowded in dense groups 
at the nodes of the spikes, each group surrounded by a short 
involucre of united bracts 67. Stachyococcus. 



10 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers not in interrupted spikes. 

Inflorescence an involucrate head (heads often compound in 

Cephaelis). 
Plants creeping herbs; seeds flat on the ventral side. 

68. Geophila. 
Plants usually large shrubs; seeds deeply furrowed on the 

ventral side 69. Cephaelis. 

Inflorescence not involucrate, or very rarely so. 
Fruiting carpels much compressed laterally, the fruit didy- 
mous; plants low, herbaceous or suffrutescent. 

70. Declieuxia. 
Fruiting carpels not compressed laterally; plants mostly trees 

or shrubs. 
Seeds with inrolled ventral surface; stipules pectinately 

lobed or with setiform appendages 71. Rudgea. 

Seeds not with inrolled ventral surface; stipules mostly 

entire or bilobate. 
Corolla tube straight, not gibbous at the base. 

72. Psychotria. 

Corolla tube elongate, more or less curved, gibbous at 
the base. Branches of the inflorescence usually 
red or yellow 73. Palicourea. 

XV. PAEDERIEAE 
A single genus in Peru 74. Paederia. 

XVI. ANTHOSPERMEAE 

Fruit leathery, pyriform, deeply costate, splitting into 2 cocci. 

75. Corynula. 
Fruit a succulent, red, berry-like drupe 76. Gomozia. 

XVII. SPERMACOCEAE 

Flowers in panicled umbels; plants scandent 77. Emmeorrhiza. 

Flowers not umbellate; plants usually not scandent. 
Fruit not separating into cocci, the whole fruit circumscissile. 

78. Mitracarpus. 
Fruit separating into cocci. 
Cocci indehiscent. 

Cocci 3-4 . . . . 79. Richardia. 



FLORA OF PERU 11 

Cocci 2 80. Diodda. 

Cocci, at least one of them, dehiscent. 
Cocci opening only at the base. Flowers axillary. 

81. Hemidiodia. 
Cocci opening at the apex. 
Cells of the fruit unlike, one opening, the other remaining 

closed 82. Spermacoce. 

Cells of the fruit alike, both opening 83. Borreria. 

XVIII. GALIEAE 

Calyx lobes developed, lanceolate 84. Sherardia. 

Calyx lobes obsolete. 

Flowers surrounded by a calyx-like involucre 85. Relbunium. 

Flowers not involucrate 86. Galium. 

1. UNCARIA Schreb. 

Scandent or trailing shrubs, armed with stout recurved spines; 
stipules interpetiolar, entire or bifid; leaves opposite, petiolate; 
flowers sessile or pedicellate, in solitary or racemose, pedunculate, 
spherical heads, 5-parted; calyx tubular or funnelform, dentate; 
corolla funnelform, the throat glabrous, the short lobes valvate in 
bud; stamens inserted in the corolla throat, exserted; fruit 2-celled, 
dry, fusiform, septicidally bi valvate; seeds few or numerous, winged 
and bicaudate. 

Flowers pedicellate; leaves glabrous U. guianensis. 

Flowers sessile; leaves tomentulose beneath or strigose on the veins. 

U. tomentosa. 

Uncaria guianensis (Aubl.) Gmel. Syst. 370. 1796. Ourouparia 
guianensis Aubl. PI. Guian. 177. pi. 168. 1775. 

A large shrub with scandent or recurved branches; leaves short- 
petiolate, the blades oblong to broadly elliptic, 6-12 cm. long, 
obtusely short-acuminate, truncate to obtuse at the base; flower 
heads 2.5 cm. in diameter; calyx 4-5 mm. long, turbinate, tomen- 
tulose; corolla white, 11 mm. long, densely pilose, the lobes obtuse; 
fruit 3.5-4 cm. long, glabrate. "Unganangi." 

Ayacucho: Choimacota Valley, 800 meters, Weberbauer 7537. 
Cuzco: Cosnipata, 700 meters, Weberbauer 6953. Loreto: Yuri- 
maguas, in forest, Williams 4170, 7844. La Campuya, W. Fox 94. 
Rio Itaya, Williams 251. Puerto Arturo, in pasture, Williams 5193. 



12 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Tarapoto, Williams 6525. Bolivia to Brazil and the 
Guianas. 

Uncaria tomentosa (Willd.) DC. Prodr. 4: 349. 1830. Nauclea 
aculeata HBK. Nov. Gen. & Sp. 3: 382. 1819, non Willd. A 7 , tomen- 
tosa Willd. ex R. & S. Syst. Veg. 5: 221. 1819. Ourouparia tomentosa 
Schum. in Mart. Fl. Bras. 6, pt. 6: 132. 1889. 

A large, more or less scandent shrub, armed with stout spines; 
leaves short-petiolate, the blades thin, oval to ovate or broadly 
oblong, 10-15 cm. long, acute or short-acuminate, usually pale 
beneath and minutely tomentulose, often glabrate and strigose on 
the veins; flowers whitish, fragrant, the dense heads 2 cm. in diam- 
eter; corolla 8-10 mm. long, densely tomentulose; capsule 6-8 mm. 
long. 

Loreto: Mouth of Rio Santiago, Tessmann ^663. Also in Colom- 
bia, the Guianas, Trinidad, and Central America. 

Of the collection cited I have seen only a fragment. Evidently 
it represents a species distinct from U. guianensis, but I do not feel 
certain that it is U. tomentosa, which is not known to occur between 
northern Colombia and Peru. 

2. ALSEIS Schott 

Trees or shrubs with opposite leaves; stipules short or elongate; 
flowers small, white or yellow, spicate, the spikes simple or branched, 
axillary and terminal; hypanthium obconic; calyx 5-lobate, the lobes 
broad or narrow, deciduous; corolla cylindric, short, pilose within, 
the limb 5-lobate, the lobes valvate; stamens inserted at the base 
of the corolla tube, the filaments villous; capsule oblong- turbinate, 
2-celled, septicidally bivalvate from the apex; seeds linear-fusiform, 
the testa produced at each end. 

Alseis peruviana Standl. Field Mus. Bot. 11: 179. 1936. 

A tree, the branchlets fulvous-hispidulous; stipules deciduous, 
ovate, acuminate, 1-1.5 cm. long; leaves short-petiolate, membrana- 
ceous, the blades rhombic-oblanceolate or cuneate-obovate, 21-37 
cm. long, 7-18 cm. wide, acute or acuminate, long-attenuate to the 
base, short-pilosulous above, densely hispidulous beneath; spikes 
6-17 cm. long; calyx 1.2 mm. long, glabrate, the lobes unequal, lance- 
oblong, acute; corolla 2.5-3 mm. long, minutely fulvous-hispidulous. 

Loreto: Puerto Arturo, Yurimaguas, 200 meters, in forest, 
Williams 5030, type. San Martin: Juan Guerra near Tarapoto, 
Williams 6916. "Mishu-quiro," "palo bianco." 



FLORA OF PERU 13 

3. MANETTIA L. 

Plants scandent, herbaceous or suffrutescent; stipules inter- 
petiolar, sometimes dentate or fimbriate; leaves opposite, herbaceous 
or coriaceous; flowers 4-parted, solitary and axillary or disposed in 
cymes or panicles, often brightly colored ; calyx with 4 or 8 segments, 
these equal or unequal, persistent; corolla tubular or funnelform, 
the lobes valvate in bud, the tube usually pilose within above the 
base, the throat glabrous or villous; stamens included or exserted, 
the anthers linear; capsule 2-celled, septicidally dehiscent; seeds 
small, discoid, winged. 

Calyx lobes 8, usually broad and foliaceous, united at the base into 
a short tube. 

Leaves glabrous or practically so. 

Corolla glabrous M. glandulosa. 

Corolla sparsely or densely pubescent outside. 

Capsule pyriform, acute at the base; leaf blades rounded at 
the base M. tarapotensis. 

Capsule globose, rounded at the base; leaf blades acute at 
the base M. coccinea. 

Leaves copiously pubescent beneath, at least when young. 

Plants villous-pilose throughout with long, weak, shaggy hairs. 

Corolla tube dilated above; calyx lobes broad; capsule 
globose M. hispida. 

Plants short-pilose. 

Corolla tube dilated above; leaves soon glabrate. 

M. vacillans. 

Corolla tube cylindric; leaves permanently pubescent. 

Calyx lobes narrowly lanceolate, long-attenuate; capsule 
obovoid, acute at the base; flowers racemose. 

M. racemosa. 

Calyx lobes lanceolate or ovate; capsule globose, rounded 
at the base; flowers subumbellate M. Schunkei. 

Calyx lobes 4. 
Leaves tomentose beneath with matted hairs. 

Calyx lobes minute, scarcely 1 mm. long M. tomentulosa. 

Calyx lobes 1.5-4 mm. long. 
Corolla glabrous outside M. poliodes. 



14 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Corolla villous or tomentose. 

Calyx lobes triangular-ovate, 1.5-2 mm. long, in fruit as 
much as 4 mm.; tomentum of the lower leaf surface 
yellowish M. peruviana. 

Calyx lobes lanceolate, 2-4 mm. long, in fruit as much as 

1 cm. ; tomentum whitish M. Weberbaueri. 

Leaves not tomentose. 

Corolla large, 3-5 cm. long M. cordifolia. 

Corolla usually less than 1.5 cm. long. 

Calyx lobes minute, ovate to lanceolate, erect or suberect, less 

than 2 mm. long, inconspicuous, never much thickened. 

Young branches glabrous. Corolla tube 3 mm. long; 

flowers laxly paniculate M. paniculata. 

Young branches pilose or villous. 

Calyx lobes broadly ovate, obtuse M. thysanophora. 

Calyx lobes triangular or lanceolate, acute or acuminate. 
Capsule obtuse at the base; flowers racemose-paniculate. 

M. veronicoides. 
Capsule acute at the base; flowers subumbellate. 

M. modica. 

Calyx lobes large or elongate, usually much more than 2 mm. 
long, sometimes recurved, often conspicuous and leafy, 
frequently much thickened. 
Corolla villous or pilose outside. 

Tube of the corolla about 6 mm. long; leaves oblong- 
lanceolate M. acutifolia. 

Tube of the corolla 12 mm. long; leaves elliptic. 

M. asclepiadacea. 
Corolla glabrous outside. 

Calyx lobes united at the base into a short tube, recurved. 

M. divaricata. 

Calyx lobes free or nearly so, not recurved. 
Calyx lobes ovate or orbicular, obtuse or rounded at 
the apex. 

Corolla tube 12-14 mm. long M. dubia. 

Corolla tube 5-6 mm. long M. umbellata. 

Calyx lobes lanceolate or linear-lanceolate, acute or 
acuminate. 



FLORA OF PERU 15 

Leaves large, about 6 cm. wide M . glandulosa. 

Leaves smaller, 3 cm. wide or less. 

Calyx lobes 6-10 mm. long; leaves coriaceous. 

M. Albert-Smithii. 
Calyx lobes 3-3.5 mm. long; leaves thin. 

M. leucantha. 

Manettia acutifolia R. & P. Fl. 1: 58. pi. 89, f. b. 1798. M. 
acutiflora Pers. Syn. PI. 1: 134. 1805. Lygistum acutifolium Kuntze, 
Rev. Gen. 1:287.1891. 

A large, slender vine; stipule lobes short, acute; leaves short- 
petiolate, thin, the blades 1.5-3 cm. wide, long-acuminate, rounded 
or obtuse at the base, glabrous or nearly so; flowers congested in the 
leaf axils; hypanthium pilose, the calyx lobes linear-lanceolate or 
linear-oblong, long-attenuate, erect; corolla purple or white, densely 
pilose outside, the tube 6 mm. long. Neg. 218. 

Cuzco(?): Type collected between Acomayo and Pati, Ruiz & 
Pavdn (photo, and fragm. of authentic material seen, ex hb. Berol.). 
Huanuco: Rio Huallaga Canyon below Rio Santo Domingo, climbing 
over shrubs, 1,200 meters, Macbride 4255. 

Manettia Albert-Smithii Standl. Field Mus. Bot. 8: 151. 1930. 

A scandent herb, the stems glabrous; stipules acute, sometimes 
fimbriate, 2-2.5 mm. long; leaves short-petiolate, coriaceous, the 
blades oblong-lanceolate to ovate, 1.5-3 cm. wide, acuminate, obtuse 
at the base, glabrous; inflorescence racemiform, the flowers short- 
pedicellate; hypanthium glabrous; calyx lobes erect, green, coriaceous, 
lanceolate, acute or acuminate; corolla white, glabrous outside, 
the tube 6-7 mm. long, the lobes 6-7 mm. long, long-villous within; 
capsule subglobose, 6-7 mm. long, obtuse or abruptly contracted at 
the base. 

Ayacucho: Carrapa, 1,200 meters, wooded hillside, Killip & 
Smith 22418, type. Aina, 750-1,000 meters, in thickets, Killip & 
Smith 23104. 

Manettia asclepiadacea Standl. Field Mus. Bot. 8: 332. 1931. 

Stems retrorse-pilosulous; stipules setulose-pectinate; leaves 
petiolate, the blades membranaceous, elliptic, 4.5-5.5 cm. wide, 
long-acuminate, acute at the base, glabrous, the veins obscure, paler 
beneath ; flowers umbellate in the leaf axils, the pedicels 5-9 mm. long; 
hypanthium densely pilosulous; calyx lobes lanceolate or narrowly 
ovate, 8-10 mm. long, green; corolla white-pilose outside, the white 



16 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

tube 12 mm. long, the lobes yellow with purple tips, 4.5 mm. long; 
capsule obovoid, acute or attenuate at the base. 

Loreto: San Antonio, Rio Itaya, 110 meters, dense forest, Killip 
& Smith 29476, type. Also in Ecuador. 

Manettia coccinea (Aubl.) Willd. Sp. PL 1: 624. 1797. Nacibea 
coccinea Aubl. PL Guian. 96. pi. 37, f. 1. 1775. 

Stems usually glabrous; stipules truncate, denticulate; leaves 
herbaceous, petiolate, the blades lanceolate to ovate, 3-10 cm. long, 
acute or acuminate, acute to obtuse at the base; flowers chiefly 
axillary; calyx lobes 8, linear or oblanceolate, 4-14 mm. long, foli- 
aceous, ciliate, recurved; corolla red, 17-25 mm. long, more or less 
pilose; capsule subglobose, 6-12 mm. long, rounded at the base. 

Huanuco: Posuso, 600 meters, over sunny shrubs, 4592. Loreto: 
Between Rio Nanay and Rio Napo, in forest, Williams 686. Widely 
distributed in the lowlands of tropical America, ranging northward 
to Mexico. 

Manettia cordifolia Mart. Denkschr. Acad. Muench. 9: 95. 
pi. 7. 1824. Guagnebina ignita Veil. Fl. Flum. 45. pi. 115. 1825. 
M. ignita Schum. in Mart. Fl. Bras. 6, pt. 6: 170. 1889. 

A large, herbaceous vine; leaves short-petiolate, membranaceous, 
ovate to lance-oblong, long-acuminate, rounded or obtuse at the 
base, finely pubescent beneath; flowers chiefly axillary and solitary, 
long-pedicellate; calyx lobes 4, oblong or lanceolate, usually unequal, 
large and foliaceous; corolla bright red, glabrous outside, 3-5 cm. 
long, the tube dilated above, the lobes very short. Neg. 193. 

Huanuco: Posuso, Lopez to Cueva Blanca, Pearce 215 (teste 
Wernham). San Martin: San Roque, abandoned land, Williams 
7773, 7479. Department unknown: Without locality, Weberbauer 
6560; Pav6n; Mathews 1343. Andes at 1,500-2,400 meters, Pearce 
590 (teste Wernham). Bolivia to Brazil and Argentina. 

Manettia cordifolia var. attenuata (Nees & Mart.) Wernham, 
Gen. Manettia 20. 1919. M. attenuata Nees & Mart. Act. Soc. Nat. 
Cur. 12: 14. 1825. 

Similar to the species, but the upper leaves, at least, narrowly 
lanceolate or linear-lanceolate, acute at the base, finely pubescent 
beneath. 

San Martin: Lamas, Williams 6406. Near Tarapoto, Spruce 
3929. Also in Brazil. 



FLORA OF PERU 17 

Manettia cordifolia var. glabra (C. & S.) Standl. Field Mus. 
Bot. 7: 263. 1931. M. glabra C. & S. Linnaea 4: 159. 1829. M. 
micans P. & E. Nov. Gen. & Sp. 3: 24. 1845. 

Like the species, but the leaves glabrous. Neg. 192. 

Huanuco: Posuso, rocky wooded slopes, 600 meters, 4722. 
Puerto Victoria, 300 meters, wooded river banks, Killip & Smith 
26823. Junin: La Merced, 600 meters, river cliffs, 5585. Vitoc, 
Ruiz. Chanchamayo Valley, 1,800 meters, Schunke 480, 1519, 
1430. San Ramon, 900-1,300 meters, edge of woods, Killip & Smith 
24748. Huacapistana, Weberbauer 1967. La Merced, Killip & 
Smith 28519. Rio Peren4, 600 meters, in forest, Killip & Smith 
25213. Loreto: Type of M. micans from Yurimaguas, Poeppig. 
Yurimaguas, 135 meters, dense forest, Killip & Smith 27990. Bal- 
sapuerto, 220 meters, King 3080. San Lorenzo, Killip & Smith 
29200. Madre de Dios(?): Seringal San Francisco, Rio Acre, 
Ule 9864. San Martin: Between Moyobamba and Tarapoto, 
Raimondi 490. Tarapoto, Mathews 1343. Rio Huallaga, Spruce 
4592. Zepelacio, 1,200-1,600 meters, King 3541- Pongo de Cai- 
narachi, 230 meters, Klug 2606. Ranging to Brazil and Argentina. 

The variety is much more frequent in most parts of the range 
than the typical form of the species. It must be a handsome and 
showy plant, with its large and brilliantly colored flowers. 

Manettia divaricata Wernham, Gen. Manettia 41. 1919. 

A large, herbaceous vine, glabrous throughout or nearly so; 
stipules small and inconspicuous, deciduous; leaves slender-petiolate, 
blackish when dried, papyraceous, the blades elliptic to ovate- 
oblong, acutely acuminate, acute at the base; flowers few, axillary 
and solitary or in 3-flowered cymes; calyx lobes fleshy, ovate or 
lanceolate, acuminate, 3-5 mm. long; corolla lilac-rose or with 
purplish green tube and dull pink lobes, the stout tube 15 mm. long, 
the rather short lobes recurved; capsule large, obovoid, acutish at the 
base or obtuse. 

Junin: Puerto Bermudez, 375 meters, dense forest, Killip & 
Smith 26615. Loreto: Lower Rio Nanay, Williams 564- Masisea, 
275 meters, open woods, Killip & Smith 26848. Mishuyacu, 100 
meters, in clearing, Klug 849. Iquitos, in forest, Killip & Smith 
29858. Also in Bolivia. 

Manettia dubia Wernham, Gen. Manettia 31. 1919. 
Stipules deltoid, acuminate; leaf blades elliptic, 6.5-8.5 cm. long, 
obtuse or acuminate, rounded at the base, glabrous; calyx lobes 



18 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ovate-orbicular, mucronate-acute, 6 mm. long; corolla blue, glabrous 
outside, the tube 12-13 mm. long, the lobes 4-5 mm. long. 

Ayacucho: Type from Puitac, at 3,000-3,300 meters, Pearce. 
Known to the present writer only from description. 

Manettia glandulosa P. & E. Nov. Gen. & Sp. 3: 24. 1845. 
Lygistum glandulosum Kuntze, Rev. Gen. 1: 287. 1891. 

Stipules deltoid, glandular-ciliate; leaves glabrous, subcoriaceous, 
the blades as much as 12.5 cm. long, acuminate, shining, especially 
beneath; peduncles axillary, 1-several-flowered; calyx lobes 4 or 
8, lanceolate, half as long as the corolla; corolla small, glabrous, pink; 
capsule oval, didymous, glabrous. 

Loreto: Type from forests of Yurimaguas, Poeppig. Reported by 
Wernham (Gen. Manettia 38. 1919) from Rio Huallaga, Yurimaguas, 
Spruce 3874- Known to the present writer only from description. 

Manettia hispida P. & E. Nov. Gen. & Sp. 3: 24. pi. 228. 1845. 
Lygistum hispidum Kuntze, Rev. Gen. 1: 287. 1891. 

A large, suffrutescent vine, villous-pilose throughout with long, 
spreading, whitish hairs, the stems straw-colored; stipules oblong, 
acute; leaves on short, stout petioles, the blades broadly elliptic to 
oblong, acuminate, acute at the base; flowers few and axillary, long- 
pedicellate, often umbellate; calyx lobes 7, oval-spatulate; corolla 
red or pinkish red, copiously long-pilose, the tube 15 mm. long; 
capsule ovoid or globose, villous. Neg. 216. 

Huanuco: Cerro de San Cristobal, near Cochero, Poeppig. 
Loreto: Yurimaguas, Poeppig 2321 (photo, and fragm. ex hb. Berol.). 
Fortaleza, Yurimaguas, edge of forest, Williams 4299. Puerto Arturo, 
in forest, Williams 5315. Balsapuerto, 220 meters, Klug 3024- 
Santa Rosa, 135 meters, dense forest, Killip & Smith 28906. Yuri- 
maguas, 135 meters, in forest, Killip & Smith 29115. San Martin: 
Lamas, in forest, Williams 6483. Department unknown: Without 
locality, Poeppig 235; Pavdn (labeled "Besleria sp. nova")- 

Manettia leucantha Krause, Repert. Sp. Nov. 19: 308. 1924. 

A slender, herbaceous vine, the stems glabrous or nearly so; 
stipules subulate-aristate; leaves short-petiolate, the blades lanceo- 
late or ovate-lanceolate, 5-8 cm. long, long-acuminate, acute or obtuse 
at the base, glabrous; calyx lobes linear-lanceolate, acute, 3-3.5 mm. 
long; corolla white, the tube 4-6 mm. long, the lobes acute, slightly 
shorter than the tube; capsule subglobose, glabrate. Neg. 210. 



FLORA OF PERU 19 

Huancavelica: Rio Mantaro, below Huancamayo, in forest, 1,500 
meters, Weberbauer 6573, type collection. 

Manettia modica Standl. Field Mus. Bot. 4: 275. 1929. 

Suffrutescent, with stramineous branches, the young branchlets 
villosulous; stipules truncate, bearing a few subulate setae; leaves 
short-petiolate, the blades lanceolate or ovate-lanceolate, 2-3.5 
cm. long, long-acuminate, obtuse at the base, sparsely villous, pale 
beneath, the margins revolute; flowers umbellate-paniculate, pedi- 
cellate; calyx lobes broadly ovate, acute; capsule obovoid, 5 mm. long, 
acute and long-attenuate at the base. 

Huanuco: Vilcabamba, Rio Chinchao, 1,800 meters, climbing 
over shrubs, Macbride 5154, type. 

Manettia paniculata P. & E. Nov. Gen. & Sp. 3: 24. 1845. 
Lygistum paniculatum Kuntze, Rev. Gen. 1: 288. 1891. 

A slender, herbaceous vine, glabrous; stipules broad, short, 
acutish; leaves on short, slender petioles, the blades lance-oblong, 
about 7.5 cm. long and 2.5 cm. wide, acuminate, acute to rounded at 
the base, glabrous, paler beneath; flowers arranged in lax, open 
panicles, the flowers long-pedicellate; calyx lobes oblong, obtuse, 
thick, half as long as the corolla, somewhat spreading; corolla tube 
3 mm. long. Negs. 25688, 202. 

Loreto: Forests of Yurimaguas, Mainas, Poeppig 2462 (photo, 
and fragm. of type material, ex hb. Berol.). Wernham (Gen. Manet- 
tia 30. 1919) reports also a specimen collected by Poeppig at Casapi 
(Huanuco). 

Manettia peruviana Standl. Field Mus. Bot. 4: 274. 1929. 

A scandent shrub, the branchlets densely villous; stipules tri- 
angular, acute, 2.5 mm. long; leaves almost sessile, the blades thick, 
oblong-ovate, 2.5-4.5 cm. long, acute or short-acuminate, rounded 
at the base, scabrous and villosulous-puberulent above, densely 
tomentose beneath; flowers umbellate, long-pedicellate; calyx lobes 
triangular-ovate, 1.5-2 mm. long; corolla dark red, short- villous, the 
tube 8 mm. long, the obtuse lobes 2 mm. long; capsule didymous- 
globose, 5-6 mm. wide. "Pushpu-hoirer." 

Huanuco: Mito, 2,700 meters, twining on shrubs of sunny slopes, 
Macbride & Featherstune 1395, type. Tambillo, 2,400 meters, 3571 . 
Junin: Mountains west of Huacapistana, 2,400 meters, Weberbauer 
2287. 



20 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Manettia poliodes Standl. Field Mus. Bot. 8: 151. 1930. 

A large vine, the young branchlets densely villosulous-tomentose; 
stipules 2 mm. long, 3-denticulate; leaves short-petiolate, the blades 
membranaceous, ovate-elliptic or oblong-ovate, 4-8 cm. long, 
acuminate, rounded or obtuse at the base, green above, sparsely 
villosulous, densely tomentose beneath; flowers umbellate-cymose, 
slender-pedicellate; calyx lobes ovate, acute or acuminate; corolla 
violaceous, the tube 7-10 mm. long, the triangular-ovate lobes 
2-2.5 mm. long; capsule subglobose, 5 mm. long, obtuse at the base. 

Cuzco: Marcapata Valley, near Chilechile, 2,500 meters, in 
thicket, Weberbauer 7863, type. Below Lares, 3,100 meters, in 
thicket, Weberbauer 7890. Valle del Apurimac, Mollepata, 2,750 
meters, Herrera 1230. 

Manettia racemosa R. & P. Fl. 1: 58. pi. 89, f. a. 1798. M. 
mutabilis Pers. Syn. PI. 1: 134. 1805. Nacibea mutabilis Poir. in 
Lam. Encycl. Suppl. 4: 55. 1816. Lygistum racemosum Kuntze, Rev. 
Gen. 1: 288. 1891. 

A large, herbaceous vine; stipules acuminate, ciliate; leaves 
slender-petiolate, the blades membranaceous, ovate to elliptic- 
oblong, more or less villous or pilosulous, at least beneath along 
the nerves, acute or acuminate, acute to obtuse at the base; flowers 
few, cymose or axillary and short-racemose; calyx lobes usually 
8, linear or lanceolate, large and elongate, herbaceous, often re- 
curved; corolla purplish or red-violet, villous; capsule ellipsoid, 
sparsely villous or pilosulous, 12 mm. long, acute at the base. 
Neg. 221. 

Huanuco: Described from Cochero, Chinchao, Cassape, Macora, 
Marimarchahua, and Hualqui, Ruiz & Pavdn (photo, and fragm. 
seen, ex hb. Berol.). Loreto: Mishuyacu, 100 meters, in marsh, 
Klug 116 If. Locality unknown: Pawn; Dombey 553. San Martin: 
Zepelacio, 1,200-1,600 meters, mountain forest, Klug 3609. 

Manettia Schunkei Standl. Field Mus. Bot. 4: 273. 1929. 

Plants scandent, herbaceous or suffrutescent, the slender stems 
at first reflexed-hirtellous; stipules triangular, acute; leaves short- 
petiolate, firm-membranaceous, the blades lance-ovate or ovate- 
elliptic, 2.5-5.5 cm. long, acuminate, rounded or obtuse at the base, 
sparsely scaberulous above, densely scaberulous beneath; cymes 
axillary, umbelliform, the flowers slender-pedicellate; calyx lobes 8, 
foliaceous, lanceolate or ovate, 3-4 mm. long; corolla red, sparsely 



FLORA OF PERU 21 

puberulent, the tube 13 mm. long, the ovate lobes 6 mm. long; 
capsule small, didymous-globose, obtuse or rounded at the base. 

Junin: Chanchamayo Valley, 1,200 meters, Schunke 362, type; 
at 1,200-1,500 meters, Schunke 1428, 1461, 1444- Colonia Perene", 
600 meters, in forest, Killip & Smith 25202. 

Manettia tarapotensis Wernham, Gen. Manettia 38. 1919. 

A large, herbaceous vine, the stems puberulent; stipules truncate, 
glandular-denticulate; leaves slender-petiolate, the blades sub- 
coriaceous, ovate or ovate-elliptic, acuminate, rounded at the base 
and abruptly contracted, glabrous, 6-9 cm. long; flowers in rather 
dense and many-flowered umbels, long-pedicellate; calyx lobes 8, 
linear, 5-6 mm. long or more; corolla puberulent, the tube 12-13 
mm. long; capsule pyriform, 1 cm. long or larger, thinly tomentulose. 

San Martin: Type from Tarapoto, Spruce 4076, 4426. Rumizapa, 
Williams 6764. Tarapoto, in forest, Williams 6105, 6528. 

Manettia thysanophora Wernham, Gen. Manettia 28. 1919. 

A rather coarse vine; stipules united, reflexed, fimbriate; leaves 
short-petiolate, membranaceous, the blades elliptic or broadly ovate, 
3.5-5 cm. long, acuminate, acute at the base, glabrate and green 
above, pale beneath and when young sparsely hirtous; flowers 
numerous, subumbellate in the axils, slender-pedicellate; calyx lobes 
4, broadly ovate, usually obtuse, glabrous, 1 mm. long; corolla 
glabrous outside, the tube 5 mm. long, the lobes 2 mm. long. 

Peru, without locality, Mathews 1501, type collection. 

Manettia tomentulosa Standl. Field Mus. Bot. 4: 273. 1929. 

Young branchlets densely puberulent or short- villous; stipules 
subtruncate, laciniate-dentate; leaves petiolate, thick-membrana- 
ceous, the blades oblong-ovate to lance-oblong, 3-7 cm. long, acumi- 
nate, rounded or obtuse at the base, sparsely appressed-pilose above 
or glabrate, minutely and densely tomentulose beneath; flowers in 
axillary umbels, slender-pedicellate; capsule subglobose, 3.5 mm. long, 
minutely puberulent, rounded at the base; calyx lobes oblong or 
ovate, obtuse. 

Junin: La Merced, 1,200 meters, climbing over bushes, Mac- 
bride 5772, type. Huacapistana, 1,800-2,400 meters, in thickets, 
Killip & Smith 24141. 

Manettia umbellata R. & P. Fl. 1: 58. pi. 90, f. a. 1798. Lygis- 
tum umbellatum Kuntze, Rev. Gen. 1: 288. 1891. 



22 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A large, glabrous vine, herbaceous or suffrutescent; stipules short, 
triangular-apiculate; leaves short-petiolate, subcoriaceous, the blades 
ovate to oblong or elliptic, acuminate, obtuse or rounded at the 
base; flowers umbellate or short-racemose in the leaf axils, slender- 
pedicellate; calyx lobes 4, ovate, obtuse or acute, thick; corolla blue, 
glabrous, the stout tube 6-7 mm. long, the lobes short, triangular- 
ovate; capsule globose-ovoid, 7 mm. long. Neg. 199. 

Ayacucho: Choimacota Valley, 2,800 meters, Weberbauer 7572. 
Huanuco: Described from Muna, Ruiz & Pavdn (photo, and fragm. 
of authentic material seen, ex hb. Berol.; labeled as from Parrasa- 
huas). Vilcabamba, 1,800 meters, 5153. Panao, 2,700 meters, a 
vine over shrubs, 3614- Pampayacu, Kanehira 104. Madre de 
Dios: Seringal San Francisco, Rio Acre, Vie 9863. 

Manettia vacillans Standl. Field Mus. Bot. 4: 272. 1929. 

Scandent, suffrutescent, the young branchlets densely puber- 
ulent; stipules acute, 2 mm. long; leaves short-petiolate, membra- 
naceous, the blades elliptic or broadly ovate, 2.5-3.5 cm. long, 
short-acuminate, rounded or obtuse at the base, sparsely appressed- 
pilose above at first, beneath sparsely tomentulose on the veins 
or almost glabrous; inflorescences subumbellate, axillary, few- 
flowered; hypanthium tomentulose; calyx lobes unequal, lanceolate 
or ovate, 2.5-5 mm. long, acuminate, spreading or recurved; corolla 
reddish white, turning rose, the tube 11-12 mm. long, the ovate, 
obtuse lobes 3-5 mm. long. 

Ayacucho: Mountains northeast of Huanta, 3,100 meters, moist 
ravine, Weberbauer 7509, type. 

Manettia veronicoides Standl. Field Mus. Bot. 4: 272. 1929. 

A slender vine, the young stems sparsely puberulent; stipules 
broadly triangular, acuminate; leaves almost sessile, the blades 
ovate or ovate-lanceolate, 2.5-4.5 cm. long, long-acuminate, rounded 
or obtuse at the base, glabrous above, sometimes tomentulose 
beneath along the nerves; flowers umbellate-paniculate, short- 
pedicellate; calyx lobes oblong- triangular, acute, 1-1.5 mm. long; 
corolla glabrous, the tube 2.5 mm. long, the lobes of the same length; 
capsule obovoid-globose, 3-4 mm. long, obtuse at the base. 

Cuzco: Between the tambos Tres Cruces and Tambomayo, 2,400 
meters, Weberbauer 6971, type. 

Manettia Weberbaueri Krause, Repert. Sp. Nov. 19: 309. 1924. 
Stipules acuminate; leaves short-petiolate, the blades lance- 
oblong to ovate or elliptic, 2-4 cm. long, short-acuminate, acute or 



FLORA OF PERU 23 

acutish at the base, sparsely appressed-pilose or glabrate above, 
pale-tomentose beneath; flowers loosely cymose; calyx lobes lanceo- 
late or ovate-lanceolate, acute, pilose, at first 2-4 mm. long, later 
accrescent; corolla pink, the tube 6-9 mm. long, appressed-pilose, 
the lobes 2-2.5 mm. long; capsule subglobose, 5 mm. long, broadly 
obtuse at the base, sparsely pilose or almost glabrous. Neg. 222. 

Cajamarca: San Pablo, 2,400-2,700 meters, Weberbauer 3808, 
type collection (photo, and fragm. seen, ex hb. Berol.). 

4. STILPNOPHYLLUM Hook. f. 

Glabrous shrubs; stipules caducous; leaves opposite, petiolate, 
coriaceous; flowers small, in small, axillary panicles, 5-parted; 
calyx cupular, 5-dentate, eglandular, persistent; corolla campanu- 
late-funnelform, coriaceous, the tube villous within at the base, the 
lobes half as long as the tube, valvate in bud ; stamens inserted in the 
corolla throat, the anthers linear; ovary 2-celled, with numerous 
ovules in each cell. The genus consists of a single species. 

Stilpnophyllum lineatum Hook. f. Icon. PI. pi. 1147. 1873; 
Schum. in Mart. Fl. Bras. 6, pt. 6: 158. pi. Ill, f. 2. 1889. Elaeagia 
lineata Spruce ex Schum. loc. cit. 

A shrub 3 meters high; leaf blades oblong-lanceolate orobovate- 
oblong, 5-9 cm. long, short-acuminate, acute at the base, lustrous; 
panicles 2 cm. long, much shorter than the leaves; hypanthium 
turbinate; calyx 1 mm. long, the teeth ovate-triangular, minutely 
pilosulous; corolla 5-6 mm. long, minutely puberulent, the lobes 
oblong, acute, recurved. 

San Martin : Summit of Cerro Pelado near Tarapoto, Spruce 4568, 
type collection. 

5. LECANOSPERMA Rusby 

Stiff, much branched shrubs; stipules short, persistent; leaves 
opposite, small, short-petiolate ; flowers terminal, solitary; calyx 
5-parted, the lobes narrow, foliaceous; corolla salverform, with a 
slender, elongate tube and broad, spreading lobes, the tube villous 
within at the insertion of the stamens; stamens inserted in the throat 
of the corolla, the anthers sessile; capsule subglobose, loculicidally 
bivalvate; seeds rather few, compressed, imbricate, narrowly winged. 
The genus consists of a single species. 

Lecanosperma lycioides Rusby, Bull. Torrey Club 20: 431. 
pi. 168. 1893. 



24 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A dense shrub 2 meters high or less, the branchlets often sub- 
spinose; leaf blades elliptic-oblong, 3-12 mm. long, glabrous or nearly 
so; corolla white, the tube nearly 2 cm. long; capsule 6 mm. long. 

Apurimac: Between Amorayana and Sanaica, 2,700 meters, 
Weberbauer 7174- Also in Bolivia. 

6. CINCHONA L. 

Trees or shrubs, glabrous or variously pubescent; stipules large, 
distinct, caducous; leaves opposite, petiolate; inflorescence terminal, 
paniculate, with opposite branches, usually many-flowered; flowers 
5-parted; calyx campanulate, dentate or lobate; corolla salverform, 
pubescent outside, the spreading lobes valvate in bud, more or less 
villous within along the margins; stamens semi-exserted or included, 
with short or elongate filaments, the anthers linear; capsule ovoid 
or oblong to subcylindric, bisulcate, dehiscent from the base to apex, 
many-seeded; seeds broadly winged. 

The genus contains some of the most important drug plants of 
the world, the source of quinine, the universal remedy for malaria. 
In former years large amounts of cinchona bark were exported from 
Peru as well as from other regions of the Andes, particularly Bolivia 
and Ecuador. The gathering of the bark involved the destruction 
of the trees, and the supply of wild trees was finally almost exter- 
minated. The greater part of the quinine now used in medicine is 
obtained from trees cultivated in the East Indies. 

There is a vast literature relating to the taxonomy and economic 
applications of the genus. Those interested particularly in medicinal 
properties of the plants made intensive studies of them, describing 
a great number of forms as species. Many forms of the trees were 
distinguished by the men who collected the bark, and various races 
that differ but slightly in characters of foliage and flowers seem to 
vary greatly as to their quinine content. Because of the large number 
of names concerned, the synonymy of the genus is exceedingly 
involved. The differences between the supposed species are poorly 
marked and imperfectly understood, or perhaps rather confused by 
the excess of attention paid to the forms. It is, therefore, dis- 
couraging to attempt to present a reasonable account of the species. 

The following enumeration of the Cinchona species is far from 
satisfactory, but it will perhaps permit a sufficiently fine segregation 
of the species. The greater number of the names in the genus relate 
to forms of C. officinalis and C. pubescens. The species vary greatly, 



FLORA OF PERU 25 

but even after the examination of a large amount of authentic 
material of their segregates, it does not seem possible to recognize 
more than a few true species in the genus. The writer believes that 
the greater part of the names published under the genus pertain to 
forms of scarcely or not at all more significance botanically than 
horticultural varieties of common garden vegetables. 

Because of the involved synonymy, it is by no means certain that 
all the names listed below are correctly placed. The most of them, 
however, are of little importance, and it matters little what disposal 
is made of them. 
Leaves hirsute, hispid, or hispidulous beneath, at least on the costa 

and nerves, the hairs stiff and spreading. 
Leaves firm-membranaceous or subcoriaceous, usually dull, hirsute 

or hispidulous beneath over the whole surface. 
Leaves glabrate on the upper surface, long-hirsute beneath. 

C. Humboldtiana. 
Leaves appressed-pilose or glabrate on the upper surface, densely 

short-pilose beneath C. Delessertiana. 

Leaves mostly thick-coriaceous, lustrous, hirsute or hispid chiefly 

on the costa and nerves. 
Calyx 1.5-2 mm. long, densely appressed-pilose. 

C. glandulifera. 

Calyx 4-5 mm. long, glabrous or glabrate C, hirsuta. 

Leaves glabrous beneath, or pilosulous or villosulous with weak or 

short, often appressed hairs, never hirsute or hispid. 
Corolla 14-17 mm. long; leaves mostly firm-coriaceous, relatively 

small C. officinalis. 

Corolla 6-12 mm. long; leaves comparatively thin and large. 
Corolla 10-12 mm. long; leaves usually copiously pubescent 
beneath C. pubescens. 

Corolla 6-8 mm. long; leaves glabrous or glabrate beneath except 
along the costa. 

Calyx lobes linear-subulate C. amazonica. 

Calyx lobes deltoid C. micrantha. 

Cinchona amazonica Standl. Field Mus. Bot. 8: 334. 1931. 

A medium-sized tree; leaves petiolate, the blades thick-mem - 
branaceous, obovate-elliptic, 15-25 cm. long, 8-12 cm. wide, obtuse 
or acutish and abruptly short-acuminate, narrowed to the base, 



26 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glabrous above, beneath sparsely and minutely sericeous or almost 
glabrous; panicle large and broad, the branches densely puberulent 
or sericeous, the flowers mostly sessile; hypanthium densely seri- 
ceous; calyx 5-parted, 1.5-2 mm. long, minutely sericeous, the lobes 
linear-attenuate; corolla 3.5-4 cm. long, densely tomentulose, the 
lobes equaling the tube. 

Loreto: Pebas, on the Amazon, Williams 1747, type. Also in 
adjacent Brazil. 

Cinchona carabayensis Wedd. Ann. Sci. Nat. III. 10: 9. 1848. 
C. Pahudiana Howard, Nueva Quinol. Pav. pi. 21. 1862. C. cara- 
bayensis var. lanceolata Miq. Ann. Mus. Bot. Lugd. Bat. 4: 270. 
1868-69. (?) C. Hasskarliana Miq. op. cit. 4: 266. 1868-69. 

A shrub 1-3 meters high, the trunk 3-4 cm. thick; stipules obo- 
vate-oblong, much longer than the petioles; leaf blades ovate-elliptic 
or lanceolate, 8-12 cm. long, 4-7 cm. wide, acute at each end or 
rarely obtuse, coriaceous, glabrate above, pubescent-tomentose 
beneath; calyx teeth triangular-lanceolate, acute; capsule oblong- 
lanceolate or ovate-lanceolate, pubescent. 

Puno : Described from thickets on the summits of the mountains 
between the valleys of the Province of Carabaya, especially near San 
Juan del Oro. Formerly, at least, planted in Java as a source of 
quinine. 

Weddell did not describe the flowers of this species, and its 
position is consequently uncertain. It may be only a form of 
C. officinalis, although in that species the leaves ordinarily are nearly 
glabrous. Weddell states that the plant is called "cascarilla de las 
lomas," and that it probably withstands a greater degree of cold 
than any other member of the genus. 

Schumann in the Flora Brasiliensis (6, pt. 6: 141. pi. 97} describes 
and illustrates C. carabayensis, but since the writer has seen no 
material of the species, even with his account of it there is difficulty 
in placing the plant. If his description of the flowers is correct, the 
species is close to C. pubescens, differing principally in its smaller 
leaves. 

Cinchona coronulata Miq. Journ. Bot. Neerl. 1: 140. 1861. 
Described from Peru, but of uncertain status. 

Cinchona Delessertiana, Standl., sp. nov. 
Ramuli crassi obtuse tetragoni densissime brunneo-pilosi, inter- 
nodiis elongatis; stipulae late ovali-ovatae 12 mm. longae deciduae 



FLORA OF PERU 27 

obtusae extus densiuscule hispidulae intus glabrae, erectae; folia 
mediocria petiolata subcoriacea, petiolo crasso 1-2 cm. longo densis- 
sime pilis brunnescentibus hispidulo-tomentoso; lamina ovalis ad 
rotundato-elliptica 6.5-17 cm. longa 5-8.5 cm. lata apice rotundata 
vel obtusa et subapiculata, basi truncata vel late rotundata, supra 
primo sparse adpresso-pilosa cito glabrata nervis nervulisque pro- 
funde impressis, subtus undique pilis brevibus patentibus brun- 
nescentibus vel flavidis dense hispidula, costa crassa elevata, nervis 
lateralibus utroque latere circa 14 prominentibus fere rectis, infimis 
angulo recto superioribus angulo paullo angustiore divergentibus, 
venulis paucis elevatis laxe reticulatis; inflorescentiae terminales et 
ex axillis superioribus nascentes densissime multiflorae longe pedun- 
culatae basi foliaceo-bracteatae, floribus dense aggregatis sessilibus, 
bracteis ultimis linearibus vel lanceolatis 5-8 mm. longis; hypanthium 
2.5 mm. longum dense fulvo-tomentosum; calyx 3-3.5 mm. longus 
dense adpresso-hispidulus, ad medium dentatus, dentibus triangular- 
ibus erectis acutis; corolla aperta non visa, in alabastro fere 1 cm. 
longa dense fulvo-strigosa; capsula immatura anguste lanceolato- 
oblonga 1.5 cm. longa sparse fulvo-hispidula. 

Amazonas: Chachapoyas, Mathews (Herb. Delessert, type). 

In a genus in which very numerous species have been described, 
based upon differences of slight importance, a group to which much 
attention has been given because of its economic importance, it is 
surprising to discover a form so conspicuously distinct from other 
species of the genus as this. It is hard to understand why this 
Mathews collection, made long ago, has not been named, but I can 
find no mention of it in literature. The species is a well-marked one, 
characterized by the very dense pubescence of short, spreading hairs 
that covers all parts of the plant. 

Cinchona glandulifera R. & P. Fl. 3: 1. pi. 224. 1802. C. 
undulata Pa von ex Howard, 111. Nueva Quinol. Pav. 2. pi. 6. 1862. 

A shrub 2-4 meters high with 2-4 trunks, these as much as 7 cm. 
thick; bark ashy gray, the branchlets pilose or hirsute ; stipules oblong, 
obtuse or acutish, villous; leaves short-petiolate, the blades ovate- 
lanceolate to lanceolate or oblong-elliptic, 8-12 cm. long, acute at 
each end, glabrous and shining above, hirsute beneath, especially 
on the nerves, the hairs somewhat glandular at the base; panicles 
leafy, terminal and axillary, many-flowered, the flowers short- 
pedicellate; calyx lobes subulate, short, both calyx and hypanthium 
densely pilosulous; corolla pinkish white, 6-7 mm. long, tomentose 
outside; capsule oblong, 1.5 cm. long. Neg. 102. 



28 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Described from the mountains of Chicoplaya, Ruiz & 
Pavdn; photo, and fragm. of authentic material seen ex hb. Berol., 
also material in hb. Deless. Reported also from Monzon, Panata- 
huas, and Cochero. "Cascarilla negrilla." 

Cinchona govana Miq. Journ. Bot. Neerl. 1: 140. 1861. The 
status of this species, described from the Department of Puno, is 
uncertain. 

Cinchona hirsuta R. & P. Fl. 2: 51. pi 192. 1799. C. pubescens 
var. hirsuta DC. Prodr. 4: 353. 1830. C. pelalba Pa von ex DC. Bibl. 
Univ. 41: 152. 1829. C. pubescens var. heterophylla DC. Prodr. 4: 
118. 1830. C. heterophylla Pavon ex t)C. loc. cit. in syn. 

A slender tree 4-5 meters high, the trunks usually several, com- 
monly 15 cm. or less in diameter; leaves on stout petioles, the blades 
ovate or ovate-elliptic, 6-10 cm. long, obtuse, cuneate to obtuse or 
rounded at the base, shining above, more or less hirsute beneath; 
panicles corymbiform, usually rather few-flowered, but sometimes 
many-flowered, the branches pubescent; calyx deeply lobate, the 
lobes linear-lanceolate, glabrate; corolla purplish, as much as 2 cm. 
long, tomentulose; capsules about 3 cm. long (described by Weddell 
as 10-14 cm., probably in error), ovoid-oblong, Negs. 91, 94, 95, 98. 

Huanuco: Described from the mountains near Pillao and Aco- 
mayo, Ruiz & Pavon; authentic material seen in hb. Deless., also 
photo, and fragm. ex hb. Berol. Authentic material of C. pelalba 
and C. heterophylla represented in hb. Field Mus. by photos, and 
fragm. ex hb. Berol. "Cascarilla delgada," "cascarilla delgadilla." 

Cinchona Humboldtiana Lamb. 111. Cinch. 7. 1821. C. villosa 
Pavon ex Lamb. loc. cit. in syn. C. conglomerata Pavon ex Howard, 
111. Nueva Quinol. Pav. pi. 15. 1862. C. Humboldtiana var. con- 
glomerata Wedd. Ann. Sci. Nat. V. 12: 45. 1869. 

Branchlets pilose or hirsute; leaves short-petiolate, the blades 
rather thin, elliptic to lance-oblong, 8-13 cm. long or larger, acute or 
acutish at each end, glabrate above, long-hirsute beneath; panicles 
mostly small and rather dense, pilose; calyx small, shallowly lobate, 
the lobes broadly triangular, acute, the hypanthium densely tomen- 
tose; corolla 1 cm. long; capsule ovate or oblong, 1.5-2.5 cm. long, 
tomentose-pilose or glabrate. Negs. 101, 105. 

Cajamarca: Type from Jae*n, Pavdn; photo, of authentic specimen 
seen, ex hb. Berol. Also in Bolivia, and perhaps in Ecuador. "Cas- 
carilla colorada." 



FLORA OF PERU 29 

In the herbarium of Field Museum there is a photograph of a 
specimen in hb. Berol, " Kunth 23," from Jauja, Junin, labeled 
Cinchona purpurascens Wedd., that may be referable to this species. 

Cinchona legitima Ruiz ex Lamb. Bull, de Pharm. 293. 1810. 
Described from Peru; a species of uncertain position. 

Cinchona micrantha R. & P. Fl. 2: 52. pi. 194. 1799. C. 
affinis Wedd. Ann. Sci. Nat. III. 10: 8. 1848. C. micrantha var. 
rotundifolia Wedd. Ann. Sci. Nat. III. 11: 270. 1849. C. micrantha 
var. oblongifolia Wedd. loc. cit. C. Reicheliana Howard, Nueva 
Quinol. Pav. pi. 5. 1862. C. micrantha var. huanucensis, C. mi- 
crantha var. affinis, C. micrantha var. Reicheliana, C. micrantha var. 
calisayoides Howard, op. cit. 1862. C. Pavoniana Kuntze, Monogr. 
Cinch. 29. 1878. 

A medium-sized tree, the branchlets sericeous or glabrate; 
stipules ovate, obtuse, glabrous; leaves petiolate, large, oval to 
obovate, rounded, or elliptic, obtuse to rounded at the apex, acute 
to rounded at the base, nearly glabrous but usually pilose beneath 
along the costa; panicles large and many-flowered, rather open, 
puberulent or tomentulose, often leafy; hypanthium minutely 
tomentulose; calyx minute, the teeth acute, purplish; corolla pink, 
about 6 mm. long, densely tomentulose, the lobes much shorter than 
the tube; capsule oblong, acute, about 1.5 cm. long. Negs. 108, 109. 

Huanuco: Type from San Antonio de Playa Grande, Tafalla; 
photo, and fragm. of authentic material seen, ex hb. Berol. Type of 
C. Reicheliana from the region of Huanuco. Cochero, Poeppig 
1339, 1737, 1708. Cajamarca: Valley of Rio Tabaconas, 1,200 
meters, Weberbauer 6271. Junin: La Merced, 1,000 meters, Weber- 
bauer 1843. San Ramon, 900-1,300 meters, edge of woods, Killip 
& Smith 24767. San Martin: Zepelacio, 1,100 meters, King 3650. 
-Without locality: Mathews 1953; Poeppig 3047, 1707, 2834. 
Reported by Weddell from Carabaya, Department of Puno. The 
type of C. affinis is from Peru. Also in Bolivia and Ecuador. 
"Cascarilla provinciana," "cascarilla provinciana blanquilla," "cas- 
carilla provinciana negrilla," "motosolo." "Quepo cascarilla," 
"cascarilla verde" (Bolivia). 

Weddell states that immediately after the bark is removed from 
the tree it assumes a blood-red color, and that it is exceedingly 
bitter. This species was formerly a more or less important source 
of the drug cinchona. 



30 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cinchona officinalis L. Sp. PI. 172. 1753. C. land/olio, Mutis, 
"Periodico de Santa Fe," 465. 1793. C. nitida R. & P. Fl. 2: 50. 
pi. 191. 1799 (type from Pampamarca, Chacahuassi, Casape, Casa- 
pillo, Cayumba, Sapan, Cochero, Ruiz & Pavdn; authentic material 
seen ex hb. Berol. and hb. Deless.). C. lanceolata R. & P. Fl. 2: 51. 
1799 (described from Mima, Panao, Pillao, Cochero, Ruiz & Pavdn; 
authentic material seen, ex hb. Berol. and hb. Deless.); 3: 1. pi. 
223. 1802. C. angustifolia Ruiz, Quinol. Suppl. 14. 1801. C. Con- 
daminea H. & B. PI. Aequin. 1: 33. pi. 10. 1808. C. colorata Lamb. 
Bull, de Pharm. 294. 1810. C. Condaminea var. lanceolata Lamb. 
111. Cinch. 2. 1821. C. stupea Pavon ex Lamb. loc. cit. in syn. 1821; 
Lindl. Fl. Med. 416. 1838. C. lancifolia var. nitida R. & S. Syst. 
Veg. 5: 9. 1825. C. lancifolia var. lanceolata R. & S. loc. cit. C. 
macrocalyx DC. Bibl. Univ. 41: 150. 1829 (authentic material seen, 
ex hb. Deless.). C. coccinea Pavon ex DC. Prodr. 4: 353. 1830, in 
syn. (authentic material seen, ex hb. Deless.). C. macrocalyx var. 
obtusifolia DC. loc. cit. C. obtusifolia Pavon ex DC. loc. cit. in syn. 
(authentic material seen, ex hb. Deless.). C. macrocalyx var. lucumi- 
folia DC. loc. cit. C. lucumifolia Pavon ex DC. loc. cit. in syn. 
(authentic material seen, ex hb. Deless.). C. macrocalyx var. Uritu- 
singa DC. loc. cit. C. Uritusinga Pavon ex DC. loc. cit. in syn. 
C. Bonplandiana Klotzsch in Hayne, Arzneigew. 7: pi. 37. 1846 
(photo, of authentic material seen, ex hb. Berol.). C. Calisaya Wedd. 
Ann. Sci. Nat. III. 10: 6. 1848. C. amygdalifolia Wedd. loc. cit. 
(described from Bolivia and Peru). C. boliviana Wedd. op. cit. 
7. 1848. C. Calisaya var. Josephiana Wedd. Ann. Sci. Nat. III. 11: 
269. 1849. C. Condaminea var. lanceolata Wedd. loc. cit. C. crispa 
Tafalla ex Howard, Nueva Quinol. Pav. pi. 2. 1862. C. Palton 
Pavon ex Howard, op. cit. pi. 13. 1862. C. parabolica Pavon ex 
Howard, op. cit. pi. 16. 1862. C. violacea, C. suberosa Pavon ex 
Howard, loc. cit. 1862. C. officinalis var. Uritusinga Howard, 
Rept. Internat. Bot. Congr. 201. 1866. (?)C. euneura Miq. Ann. 
Mus. Bot. Lugd. Bat. 4: 265. 1868-69 (type from Rio Grande, 
Carabaya, Department of Puno, Hasskarl). C. lucumifolia var. 
stupea Wedd. Ann. Sci. Nat. V. 11: 358. 1869. C. Calisaya var. 
boliviana Wedd. Ann. Sci. Nat. V. 12: 55. 1869. C. elliptica Wedd. 
op. cit. 60. 1869 (type from Province of Carabaya, Department of 
Puno). C. Weddelliana Kuntze, Monogr. Cinch. 29. 1878. Hindsia 
subandina Krause, Bot. Jahrb. 40: 431. 1908, nomen. 

A large or medium-sized tree or sometimes only a shrub, the 
trunk rarely as much as 1.5 meters in diameter, the bark rugose, 



FLORA OF PERU 31 

fuscous, the branchlets strigillose-pilosulous; stipules lanceolate or 
oblong, acute or obtuse, glabrous; leaves petiolate, lanceolate to 
elliptic or ovate, small, acute, acuminate, or obtuse, at the base 
rounded to attenuate, coriaceous, glabrous above and often lustrous, 
glabrous beneath or puberulent or short-pilose, especially on the 
veins, usually about 10 cm. long and 3.5-4 cm. wide, often scrobicu- 
late beneath; panicles terminal, leafy, rather small and dense, 
many-flowered; hypanthium strigose; calyx glabrous or nearly so, 
reddish, the teeth triangular, acute; corolla pink or red, sericeous, 
the lobes ovate, acute, the tube about 1 cm. long; capsule oblong, 
commonly 1.5-2 cm. long, glabrate. Negs. 96, 110, 112, 114-121, 572. 

Amazonas: Reported from Chachapoyas. Huanuco: Pam- 
payacu, Kanehira 352. Pan de Azucar, Sawada 69, 72. Junin: 
Jae"n, Rivera 588. Loreto: Canela Ucsha, 1,000 meters, Ule 6767 
(type of Hindsia subandina). Puno: Reported from the Sandia 
Valley. San Martin: Monte Campana near Tarapoto, Spruce 
4832 (det. doubtful). Reported to grow at 1,600-2,700 meters. 
Ranging from Bolivia to Colombia. "Cascarilla verde," "cascarilla 
verde morada," "cascarilla crespilla," "cascarilla calisaya," "ichu 
cascarilla." 

This species was formerly an important source of cinchona bark 
in the Andean region, and it was also the one first used in medicine. 
It was first made known from the region of Loja, in southern Ecua- 
dor. The species as treated here is a variable one, at least as regards 
leaf form, but no clear lines can be drawn to separate the forms into 
groups. It may well be, however, that with ample material available 
for study, if it ever can be brought together, there may be found 
some means of distinguishing several species in the material now 
referred to C. officinalis. 

Cinchona pubescens Vahl, Skrivt. Naturh. Selsk. 1: 19. 1790. 
C. Morado Ruiz, Quinol. 67. 1792. C. purpurea R. & P. Fl. 2: 52. 
pi. 193. 1799 (type from Pati, Ruiz & Pavdn; authentic material 
seen, ex hb. Berol. and hb. Deless., from Chinchao, Pillao, etc.). 
C. ovata R. & P. op. cit. 52. pi. 195. 1799 (described from Posuso 
and Panao; authentic material seen, ex hb. Berol.). C. grandifolia 
Mutis ex Humb. Ges. Naturf. Freund. Berlin Mag. 1: 117. 1807. 
C. scrobiculata H. & B. PI. Aequin. 1: 165. pi. 47. 1808 (type from 
Jae"n, Department of Junin, Humboldt &Bonpland; type material seen 
in hb. Deless.). C. rotundifolia Pavon ex Lamb. 111. Cinch. 5. 1821. 
C. pubescens var. ovata DC. Prodr. 4: 353. 1830. C. pallescens Ruiz 
ex DC. loc. cit. in syn. C. tennis Ruiz ex DC. loc. cit. in syn. C. dis- 



32 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

color Hayne, Arzneigew. 14: sub pi. 14- 1846. C. Delondriana Wedd. 
Ann. Sci. Nat. III. 10: 7. 1848 (type from Peru). C. rufinervis Wedd. 
op. cit. 8. 1848 (type from Peru). C. Pelletieriana Wedd. loc. cit. 
1848 (described from Bolivia and Peru). (?)C. purpurascens Wedd. 
loc. cit. 1848 (described from Bolivia and Peru). C. scrobiculata 
var. genuina Wedd. Ann. Sci. Nat. III. 11: 270. 1849. C. scrobicu- 
lata var. Delondriana Wedd. loc. cit. 1849. C. pubescens var. pur- 
purea Wedd. loc. cit. 1849. C. ovata var. vulgaris Wedd. loc. cit. 
1849. C. ovata var. erythroderma Wedd. loc. cit. 1849, nomen nudum. 
C. Lechleriana Schlecht. Linnaea 26: 728. 1853-55 (type from San- 
gaban, Department of Puno, Lechler 2347; type material seen in lib. 
Deless.). C. succirubra Pavon ex Klotzsch, Abh. Akad. Berl. 1857: 60. 
1858 (type material (?) seen in hb. Berol., from Tucora in the hills of 
San Antonio). C. lutea Pavon ex Howard, Nueva Quinol. Pav. pi. 14 
1862. C. obovata Pavon ex Howard, op. cit. pi. 18. 1862 (type from 
region of Huanuco, Pavdri). C. decurrentifolia Pavon ex Howard, 
op. cit. pi. 23. 1862. C. peruviana Howard, op. cit. pi. 27. 1862 
(type from Cochero). C. subcordata Pavon ex Howard, loc. cit. 
1862. C. viridiflora Pavon ex Howard, loc. cit. 1862. C. subsessilis 
Miq. Ann. Mus. Bot. Lugd. Bat. 4: 272. 1868-69 (type from Peru, 
Hasskarl). C. caloptera Miq. op. cit. 273. 1868-69 (grown in Java 
from Peruvian seed). C. erythroderma Wedd. Ann. Sci. Nat. V. 11: 
362. 1869 (type from Rio Vilcomayo north of Cuzco). C. platy- 
phylla Wedd. loc. cit. 1869. C. elliptica Wedd. Ann. Sci. Nat. V. 
12: 60. 1869. C. Howardiana Kuntze, Monogr. Cinch. 30. 1878. 

A medium-sized tree, the trunk about 30 cm. in diameter, the 
branchlets pubescent; stipules large, ovate, obtuse or acute, sericeous 
or almost glabrous; leaves slender-petiolate, the blades usually 
large, commonly broadly ovate to orbicular, rounded to acute at 
the apex, cordate to acute at the base and often decurrent, usually 
glabrate above, beneath densely short-pilose or tomentose to glabrate, 
often scrobiculate; panicles usually large, often leafy, many-flowered, 
open, the flowers subsessile; hypanthium densely sericeous; calyx 
appressed-pilosulous, the teeth short and broad, acute; corolla red 
or pink, sericeous, 10-12 mm. long, the lobes half as long as the tube; 
capsule lanceolate or oblong, glabrate, commonly 1.5-2.5 cm. long. 
Negs. 93, 103, 106, 107, 571, 22802. 

Cajamarca: Huambos, 2,400 meters, Weberbauer 4202, 4195. 
Cuzco: San Miguel, Urubamba Valley, 1,800 meters, Cook & Gilbert 
1173,1781. Torontoy, 2,400 meters, Cook & Gilbert 1 099. Huanuco : 
Pan de Azucar, Sawada 75. Rio Posuso, 2,300 meters, Weberbauer 



FLORA OF PERU 33 

6782. Casapi, Mathews 1952. Junin: Jae"n, Bonpland. Chan- 
chamayo Valley, 1,600 meters, Schunke 1490, 546. Puno: Mountains 
near Sangabdn, Lechler 2347. Province of Carabaya, Weddell 4353. 
Bolivia to Colombia. "Cascarilla," "cascarilla boba," "cascarilla 
delgada," "cascarilla pata de gallinazo," "cascarilla fina," "cascarilla 
morada," "cascarilla pata de gallareta," "palo bianco," "quina de 
Chito y Ynta," "quina blanca," "cascarilla colorada," "cascarilla 
de Santa Ana," "quina amarilla," "carua-carua," "cargua-cargua," 
"cascarilla mula," "cascarilla zamba morada," "cascarilla de 
Carabaya." 

This species is almost as variable as C. officinalis. It is and has 
been one of the most important sources of the cinchona bark 
employed in medicine, if not the principal one. 

Cinchona umbellulifera Pavon ex Howard, 111. Nueva Quinol. 
Pav. pi. 22. 1862. Described from Peru, the type collected by 
Pavon. I have seen no material of the species, whose status is 
uncertain. 

7. JOOSIA Karst. 

Trees with bitter bark; stipules caducous; leaves opposite, 
petiolate, herbaceous; inflorescence cymose or verticillate-umbellate, 
the flowers white, 5-parted; calyx large, 5-dentate, in bud open or 
subvalvate; corolla salverform, deeply lobed, each lobe provided 
with 2 petaloid, crispate appendages, the tube glabrous within; 
stamens inserted in the tube, the filaments short, the anthers linear; 
capsule linear, bisulcate, 2-celled ; seeds numerous, imbricate, winged. 
The genus consists only of the three species listed here. 

Capsule valves 1.5-2.5 cm. long, not spirally twisted after dehiscence. 

J. Dielsiana. 

Capsule valves 3.5-6.5 cm. long, spirally twisted after dehiscence. 
Leaves glabrous on the upper surface; inflorescence cymose- 

umbellate; capsule appressed -pilose J. umbellifera. 

Leaves appressed-pilose on the upper surface; inflorescence strictly 
dichotomous; capsule glabrous J. dichotoma. 

Joosia dichotoma (R. & P.) Karst. Fl. Columb. 1: 9. 1858. 
Cinchona dichotoma R. & P. Fl. 2: 53. pi. 197. 1799. Ladenbergia 
dichotoma Klotzsch in Hayne, Arzneigew. 14: sub pi. 15. 1846. 
Cosmibuena dichotoma G. Don, Hist. Dichl. PI. 3: 479. 1834. 

A nearly glabrous shrub or tree 2-8 meters high; stipules oblong- 
obovate, obtuse, longer than the petioles, caducous; leaves short- 



34 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

petiolate, oblong-lanceolate to elliptic, acute, at the base acute or 
obtuse, sericeous-pilose beneath; inflorescence rather few-flowered, 
the flowers secund, short-pedicellate; corolla about 17 mm. long, 
with a slender tube. Neg. 6641. 

Huanuco: Type from Chicoplaya, Tafalla in 1797. Loreto: 
Pampas de Ponasa, 1,200 meters, Ule 6795. 

Joosia Dielsiana Standl. Field Mus. Bot. 11: 219. 1936. 

A tree 6 meters high, the trunk 12 cm. in diameter; stipules 
narrowly oblong, 1 cm. long, obtuse, glabrate, deciduous; leaves 
9-18 cm. long, petiolate, elliptic-oblong or narrowly elliptic, acutely 
acuminate, acute at the base, glabrous above, at first laxly appressed- 
pilose beneath, soon glabrate; inflorescence long-pedunculate, rather 
laxly few-flowered; corolla densely appressed-pilose, the slender 
tube greenish yellow, 11-12 mm. long, the lobes of equal length; 
capsule oblong, subterete, glabrate. 

Loreto: Upper Maranon, mouth of Rio Santiago, in upland forest, 
160 meters, Tessmann 4478, type. 

The fruit is so unlike that of the other species that the tree might 
be considered worthy of generic segregation, but the curious corolla 
is very similar to that of the other species, and quite unlike the corolla 
of any other genus of the family. 

Joosia umbellifera Karst. Fl. Columb. 1: 9. pi. 5. 1858; Schum. 
in Mart. Fl. Bras. 6, pt. 6: 159. pi. 100. 1889. 

A medium-sized tree; stipules linear-oblong, acute or subobtuse, 
1.5 cm. long; leaves short-petiolate, oblong to obovate-oblong or 
oblong-lanceolate, 10-22 cm. long, long-acuminate, acute to rounded 
at the base, appressed-pilose beneath on the veins; inflorescence 
15 cm. long; calyx lobes triangular-ovate, acute; corolla subsericeous, 
12-14 mm. long, the lobes ovate, each bearing at the apex 2 large, 
oval appendages; capsule 3.5-6.5 cm. long, 3 mm. wide. 

San Martin: Reported by Schumann from Tarapoto, Spruce 
3944. Loreto: Balsapuerto, 220 meters, King 2943. Also in 
Colombia. 

The plant is described as a shrub of 2-3 meters or as a tree of 
7-11 meters. The .flowers are pure white and sweet-scented. 

8. MACROCNEMUM L. 

Shrubs or trees; stipules usually caducous; leaves opposite, 
petiolate; inflorescences axillary, often leafy, paniculate, the flowers 



FLORA OF PERU 35 

usually showy, 5-parted; calyx persistent, short or elongate, 5-den- 
tate or lobate; corolla salverform or funnelform, the lobes valvate 
in bud, the tube pilose at the insertion of the stamens; stamens 
unequal, attached to the middle of the tube, exserted; capsule 2- 
celled, oblong or cylindric, bisulcate, loculicidal; seeds numerous, 
minute, compressed, narrowly winged. 

Leaves glabrous beneath but barbellate in the axils of the nerves, 
sometimes minutely appressed-pilose when young, but soon 
glabrate M . roseum. 

Leaves more or less densely pilose beneath with short, spreading 
hairs. 

Calyx lobes obtuse M. cinchonoides. 

Calyx lobes acute M. pilosinervium. 

Macrocnemum cinchonoides Wedd. Ann. Sci. Nat. IV. 1: 
76. 1854. Lasionema cinchonoides Wedd. Hist. Quinquin. 99. 1849. 

Leaves short-petiolate, the blades obovate or oblanceolate- 
oblong, acute, narrowed to the base, glabrous above, short-pilose 
beneath, the veins glabrate; calyx lobes short, obtuse; branches of 
the panicle densely rusty-pilose; capsule linear-clavate, arcuate, 
1.5-2 cm. long. "Cascarilla bruta." Neg. 183. 

Puno: Humid forests, Tambopata, Province of Carabaya, 
Weddell 4349, type material. 

Probably both this and M. pilosinervium are only variants of 
M. roseum, and not worthy of specific rank. 

Macrocnemum pilosinervium Standl. Field Mus. Bot. 4: 
326. 1929. 

Branchlets densely appressed-pilose; leaves short-petiolate, the 
blades obovate or oblong-obovate, 15-22 cm. long, acute, acute at 
the base or attenuate, glabrous above, densely spreading-pilose 
beneath and with dense, subappressed hairs along the veins; inflo- 
rescences long-pedunculate, densely many-flowered, the branches 
appressed-pilosulous; corolla glabrous, the tube 1 cm. long, the 
rounded lobes 2-2.5 mm. long; immature capsule 8 mm. long. 

Type from Peru, without locality, Mathews in 1862. 

Macrocnemum roseum (R. & P.) Wedd. Ann. Sci. Nat. IV. 
1: 76. 1854. Cinchona rosea R. & P. Fl. 2: 54. pi. 199. 1799. C. 
fusca Ruiz ex Vitm. Summa Suppl. 1 : 262. 1802. Lasionema roseum 
Don, Edinb. Phil. Mag. 2: 377. 1833. C. Tarantaron Pav. ex Wedd. 



36 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Hist. Quinquin. 64. 1840, in syn. M. Sprucei Rusby, Mem. Torrey 
Club 6: 44. 1896. 

A shrub or tree, as much as 15 meters high and probably larger, 
nearly glabrous; leaves petiolate, the blades obovate to oblong, 
obtuse, cuneately narrowed to the base, firm, mostly 6-15 cm. long 
and 3-7 cm. wide; inflorescence rusty-pilosulous or sometimes 
glabrate; corolla pink, glabrous outside, about 12 mm. long; capsule 
1.5-2 cm. long. Negs. 185, 564, 6643. 

Amazonas: Mouth of Rio Santiago, Tessmann 3509. Maranon 
Valley, Tessmann 3835. Huanuco! Described from Posuso and San 
Antonio de Playa (type material seen). Posuso, 600 meters, Mac- 
bride 4588. Pampayacu, Sawada 15. Rio Posuso, Weberbauer 
6751. Loreto: Lower Rio Huallaga, 200 meters, Williams 1+957. 
Cahipuerto, 250 meters, Klug 3123. Puerto Arturo, 135 meters, 
dense forest, Killip & Smith 27898. Pumayacu, 600-1,200 meters, 
Klug 3202. Madre de Dios(?): Seringal San Francisco, Rio Acre, 
Ule 9862. San Martin: Tarapoto, Williams 5468; Spruce 3972. 
Campana, near Tarapoto, Spruce 4367. Pongo de Cainarachi, 230 
meters, Klug 2631. Juan Jui, Alto Rio Huallaga, 400 meters, 
Klug 3788. Without locality, Pavon. Also in Bolivia and Brazil. 

"Palo de San Juan," "asmonich," "cascarillo pardo." Ruiz and 
Pavon state that the flowers were used for decoration in the 
churches. 

9. REMIJIA DC. 

Shrubs or trees; stipules binate or quaternate; leaves opposite 
or verticillate, petiolate, usually coriaceous; inflorescence axillary, 
paniculate, with opposite branches, 5-parted; calyx dentate or 
lobate; corolla salverform, the tube glabrous within, the lobes mostly 
coriaceous, glabrous on the margins, valvate in bud ; stamens inserted 
on the tube, included or subexserted, the anthers linear; capsule 
2-celled, loculicidal, the valves woody, bifid; seeds peltately attached, 
winged. 
Capsules 8-10 mm. long; leaf blades attenuate to the narrow base; 

hypanthium glabrous or nearly so R. megistocaula. 

Capsules 12-15 mm. long; leaf blades merely acute at the base or 

often obtuse; hypanthium densely appressed-pilose. 

R. peruviana. 

Remijia megistocaula Krause, Bot. Jahrb. 40: 319. 1908. 

A tree up to 30 meters high, the branchlets puberulent; stipules 

3-3.5 cm. long, ovate, acuminate, glabrate, soon deciduous; leaves 



FLORA OF PERU 37 

short-petiolate or almost sessile, the blades coriaceous, lanceolate 
to obovate-lanceolate, 25^10 cm. long, 12-16 cm. wide, or sometimes 
smaller, glabrate, short-acuminate; inflorescences large and many- 
flowered, ferruginous-pilose; capsule narrowly pyriform, glabrous. 
Negs. 161, 162. 

Huanuco: Between Monzon and Rio Huallaga, 700 meters, 
Weberbauer 3687, type collection. 

Remijia peruviana Standl. Field Mus. Bot. 8: 156. 1930. 

A shrub or tree as much as 10 meters high; stipules deciduous, 
oval or oblong, obtuse or rounded at the apex; leaves opposite, 
short-petiolate, the blades coriaceous, elliptic or broadly ovate, 
8-17 cm. long, 4-9 cm. wide, acute, beneath minutely appressed- 
pilosulous, especially on the veins, or glabrate; panicles long-peduncu- 
late, many-flowered, about equaling the leaves, densely fulvous- 
pilosulous; calyx lobes ovate or lanceolate, acute; corolla white, 
densely fulvous-sericeous, the tube 8-11 mm. long, the attenuate 
lobes 5-6 mm. long; capsule narrowly oblong, sparsely appressed- 
pilosulous. "Cascarilla," "asar lisa," "collar lisa." "Quina" 
(Colombia). 

Loreto: Balsapuerto, 220 meters, King 3072. Near Marana, 
vicinity of Iquitos, Williams 1512 (type), 1511. Forest between 
Nanay and Napo rivers, Williams 675. Punchana, Williams 1333, 
8022. San Juan, in forest, Williams 3738. Morona-cocha, 115 
meters, Mexia 651 2a. Palta-cocha, Alto Nanay, Williams 3190. 
Nauta, Rio Ucayali, Spruce 3857. Mishuyacu, 100 meters, Klug 
1340, 1203. Iquitos, Tessmann 3661, 5095; Killip & Smith 26973, 
27296. Rio Nanay, Williams 340. San Martin: Tarapoto, Spruce 
4581, 4263; Williams 6107, 6670, 6322. Also in Colombia. 

10. PIMENTELIA Wedd. 

Trees; stipules free, covered at the base with resin; leaves oppo- 
site, short-petiolate, subcoriaceous; inflorescence axillary, paniculate, 
small, the flowers sessile; hypanthium turbinate, puberulent; calyx 
short, persistent, the teeth triangular; capsule linear-oblong, short, 
2-celled, septicidally dehiscent from apex to base, the valves entire; 
seeds numerous, minute, linear-lanceolate, winged. 

The genus was named for Don Pablo Pimentel, once governor of 
the Province of Carabaya. It consists of a single species. 

Pimentelia glomerata Wedd. Monogr. Cinchon. 94. pi. 27, 
B. 1849. 



38 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tree 5-6 meters high, the trunk 20-30 cm. in diameter, the 
branchlets glabrous, resinous; stipules ovate, obtuse, glabrous; leaf 
blades oblong-obovate, 20-25 cm. long, 10-13 cm. wide, rounded to 
short-acuminate, acute at the base, pilosulous beneath in the axils 
of the nerves, otherwise glabrous; panicles opposite, rounded, many- 
flowered, the short branches glabrate; capsule 1 cm. long, 3 mm. 
wide, glabrate. Neg. 160. 

Puno: Type collected in the mountains of Carabaya, growing 
with various species of Cinchona and Ladenbergia, Weddell; photo, 
seen of specimen in hb. Berol. "Ueno-ueno," "beno-beno." 

11. LADENBERGIA Klotzsch 

Shrubs or small trees, pubescent or almost glabrous; stipules 
free or united, caducous; leaves opposite, petiolate, small or large, 
often coriaceous; flowers large or medium-sized, disposed in terminal 
panicles, these leafless and with opposite branches; calyx commonly 
5-dentate or 5-lobate; corolla salverform, pilose, the lobes valvate, 
short-papillose on the margins or over the inner surface; stamens 
included, the anthers linear; capsule usually cylindric, dehiscent 
from the apex, woody or coriaceous; seeds broadly winged. 

The species of this genus, closely related to Cinchona, are almost 
as difficult of discrimination as in the latter. Since they are of little 
economic importance, less attention has been given to them by 
botanists and pharmacists, and their synonymy is, therefore, far less 
involved. 

Corolla 7-10 mm. long;' capsule 1-2 cm. long. Leaves acuminate, 
pilose beneath in the axils of the nerves, otherwise glabrous or 

nearly so L. pedunculata. 

Corolla 1.8-5.5 cm. long; capsules usually larger. 
Leaves conspicuously bullate, oblanceolate-oblong, short-hispid 
beneath on the veins. Capsules about 4.5 cm. long .L. bullata. 
Leaves not bullate, not short-hispid beneath, usually broadest at 

or below the middle. 

Calyx 7 mm. wide or more in anthesis; corolla tube very thick 
and broad ; stipules connate into a cap. Leaf blades acute 

at the base, densely pubescent beneath L. crassifolia. 

Calyx much narrower; corolla tube slender; stipules free or 

nearly so. 

Leaves densely pubescent or tomentose beneath, the pubes- 
cence persistent. 



FLORA OF PERU 39 

Leaf blades acute at the base, oblong or obovate-oblong. 

L. Riveroana. 
Leaf blades rounded or subcordate at the base, broadly 

oval to elliptic or rounded. 
Lobes of the corolla almost equaling the tube, the corolla 

less than 2 cm. long L. gavanensis. 

Lobes of the corolla much shorter than the tube, the 
corolla more than 3 cm. long. 

Corolla 3.5 cm. long L. malacophylla. 

Corolla 4.5-5 cm. long L. Carua. 

Leaves glabrous or glabrate beneath except sometimes on the 

veins, the pubescence, if any, chiefly deciduous. 
Calyx repand-dentate or shallowly lobate, the lobes shorter 

than the tube. 
Leaves glabrous, paler beneath, 2-4.5 cm. wide. 

L. discolor. 

Leaves pubescent beneath on the veins, broader. 

L. ferruginea. 

Calyx deeply lobate, usually to below the middle. 
Calyx lobes rounded-ovate, obtuse. . . .L. heterophylla. 
Calyx lobes ovate or lanceolate, acuminate to acutish. 
Hypanthium glabrous. Leaf blades 5-6.5 cm. long, 

rounded-obovate, glabrous L. coriacea. 

Hypanthium densely pilose or sericeous. 
Corolla 2 cm. long or less. 

Leaf blades oblong-lanceolate, acute at the base. 

L. stenocarpa. 
Leaf blades chiefly elliptic or ovate and obtuse to 

rounded at the base L. magnifolia. 

Corolla 3-3.5 cm. long. 

Leaf blades rounded at the base; pubescence of 

the hypanthium spreading. . . .L. acutifolia. 

Leaf blades acute or obtuse at the base; pubescence 

of the hypanthium appressed . .L. graciliflora. 

Ladenbergia acutifolia (R. & P.) Klotzsch in Hayne, Arznei- 
gew. 14: sub pi. 15. 1846. Cinchona acutifolia R. & P. Fl. 3: 1. 
pi. 225. 1802. Cascarilla acutifolia Wedd. Ann. Sci. Nat. III. 10: 
11. 1848. Buena acutifolia Wedd. Journ. Linn. Soc. 11: 186. 1869. 



40 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A shrub or tree 3-7 meters high, the branches glabrate; stipules 
ovate, acute; leaves petiolate, the blades ovate-lanceolate to lance- 
oblong, sometimes 20 cm. long, acute or acuminate, rounded at the 
base, coriaceous, glabrous and shining above, pilosulous beneath on 
the veins; inflorescence long-pedunculate, small, many-flowered, 
the branches tomentulose, the flowers subsessile; calyx lobes lan- 
ceolate; corolla white, 3 cm. long, the lobes shorter than the tube; 
capsule oblanceolate, about 3 cm. long, pubescent. Negs. 136, 6642. 

Huanuco: Type from Chicoplaya, Ruiz & Pavon; authentic 
specimen seen in hb. Deless., and photos, ex hb. Berol. "Cascarilla." 

Ladenbergia bullata (Wedd.) Standl. Trop. Woods 34: 41. 1933. 
Cascarilla bullata Wedd. Ann. Sci. Nat. III. 10: 11. 1848. Buena 
bullata Wedd. Journ. Linn. Soc. 11: 187. 1869. 

A shrub or tree 3-5 meters high, the branchlets glabrate; stipules 
oblong; leaves short-petiolate, the blades oblanceolate-oblong, 15-20 
cm. long, 4-8 cm. wide, acute or acutish at each end, conspicuously 
bullate, almost glabrous, coriaceous, shining above; panicle few- 
flowered, corymbose, the branches ferruginous-pilose; calyx lobes 
lanceolate; capsule oblong, 4-8 cm. long, 6-10 mm. wide, glabrate. 
Negs. 127, 25714. 

Puno: Described from the valleys of Tambopata and San Juan 
del Oro, at 2,000-2,500 meters, Weddell; authentic material seen in 
hb. Deless. Between Yuncacoya and Ramospata, Raimondi 10068. 
"Cargua-cargua," "cargua-cargua chica." 

Ladenbergia Garua (Wedd.) Standl. Field Mus. Bot. 7: 274. 
1931. Cascarilla Carua Wedd. Ann. Sci. Nat. III. 10: 12. 1848. 
Cinchona Carua Miq. Ann. Mus. Bot. Lugd. Bat. 4: 275. 1868-69. 
Buena Carua Wedd. Journ. Linn. Soc. 11: 187. 1869. 

A tree, the branchlets ferruginous- tomentose; stipules large, 
obtuse or rounded, tomentose; leaves short-petiolate, the blades 
oblong or oval, very large, 30-40 cm. long and more, obtuse or 
acute, cordate or rounded at the base, pubescent or glabrate above, 
ferruginous- tomentose beneath; panicles large, rather few-flowered, 
the white flowers very fragrant; calyx teeth short, triangular; 
corolla densely sericeous, 4.5-5 cm. long, the lobes more than half 
as long as the tube; capsule 3.5-5 cm. long, tomentose. 

Reported by Weddell from the departments of Cuzco and Puno 
(Province of Carabaya). Also in Bolivia. "Cargua-cargua," "car- 
gua-cargua grande." "Carua," "cascarilla de mula" (Bolivia). 



FLORA OF PERU 41 

Ladenbergia coriacea Krause, Bot. Jahrb. 40: 318. 1908. 

A shrub or small tree 4 meters high; stipules triangular, acute, 
4-6 mm. long; leaves short-petiolate, stiff-coriaceous, the blades 
rounded-obovate or broadly elliptic, 5-6 cm. long, 3-3.5 cm. wide, 
obtuse to rounded and apiculate at the apex, acutish to rounded at 
the base, glabrous; panicles many-flowered, rather dense; hypan- 
thium glabrous, the triangular calyx lobes acute; corolla puberulent, 
the tube 15-18 mm. long, the lobes oblong, 8-10 mm. long; capsule 
rather broadly oblong, 15-18 mm. long, glabrous. Neg. 128. 

Amazonas: Molinopampa east of Chachapoyas, 2,000-2,300 
meters, Weberbauer 4331, type; photo, and fragm. seen, ex hb. Berol. 
Chachapoyas, Mathews. 

Ladenbergia crassifolia (Pa von) Standl. Field Mus. Bot. 7: 
200. 1931. Cinchona crassifolia Pavon ex DC. Bibl. Univ. 41: 150. 
1829; Prodr. 4: 354. 1830. Cascarilla calyptrata Wedd. Ann. Sci. 
Nat. III. 10: 13. 1848. Buena crassifolia Wedd. Journ. Linn. Soc. 
11: 187. 1869. 

A tree, the branchlets densely ferruginous-tomentose; stipules 
connate into a cap that encloses the buds; leaves petiolate, the 
blades coriaceous, oblong to narrowly elliptic, mostly 15-20 cm. 
long, acutish, attenuate to the base, glabrous and lustrous above, 
densely puberulent beneath; panicles corymbiform, few-flowered, 
the branches ferruginous-tomentose; calyx short, the broadly tri- 
angular lobes acute or obtuse; corolla tube thick, densely sericeous, 
2.5 cm. long, the narrow lobes slightly shorter; capsules 4-7 cm. 
long, 1 cm. broad, becoming glabrate. Neg. 25716. 

Junin: Chanchamayo Valley, 1,500 meters, Schunke 297. 
Without locality, Pavdn. Also in southern Ecuador, where it was 
first collected by Ruiz and Pavon in the region of Loja. 

The Peruvian plant has been confused with the closely related 
L. macrocarpa (Vahl) Klotzsch, and the latter has been reported 
from Peru, apparently in error, the collections so reported being 
really from southern Ecuador. 

Ladenbergia discolor Schum. in Mart. Fl. Bras. 6, pt. 6: 
146. 1889. 

A shrub 1-4 meters high, the branchlets at first ferruginous- 
tomentulose, soon glabrate; leaves short-petiolate, coriaceous, the 
blades oblong or ovate-oblong, 7-10 cm. long, 2-4.5 cm. wide, acute 
or subobtuse at each end, lustrous, glabrous, paler beneath; inflores- 



42 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cence dense, 4-5 cm. long, the branches pilosulous, the flowers sub- 
sessile; calyx repand-dentate or undulate; corolla white, sericeous, 
18 mm. long, the lobes half as long as the tube, linear, acute. 
Neg. 140. 

Loreto: Canela Ucsha, 1,000 meters, Vie 6767. San Martin: 
Near Tarapoto, Spruce 4%47, type collection. Between Tarapoto 
and Chasuta, Raimondi 1195. 

Ladenbergia ferruginea Standl. Field Mus. Bot. 8: 336. 1931. 

A shrub 4 meters high, the branchlets densely ferruginous-his- 
pidulous; leaves slender-petiolate, subcoriaceous, the blades oblong- 
elliptic or lance-oblong, 14-25 cm. long, 5.5-9 cm. wide, acute or 
short-acuminate, narrowed to the acute base, ferruginous-pilose 
beneath on the veins; panicles dense and many-flowered, the branches 
ferruginous- tomentose, the flowers pedicellate; calyx 3.5 mm. long, 
sparsely pilose, the teeth triangular, acute or apiculate; corolla 
white, appressed-pilose, the tube 10-12 mm. long, the lobes 6-7 mm. 
long. 

Puno: Chunchusmayo, in forest near the river, 900 meters, Weber- 
bauer 1175, type. 

Ladenbergia gavanensis (Schlecht.) Standl., comb. nov. 
Cascarilla gavanensis Schlecht. Linnaea 26: 730. 1854. 

Leaves petiolate, subcoriaceous, the blades rounded-oval, 25 
cm. long or more, obtuse or rounded at each end or sometimes 
acutish, densely ferruginous-tomentose beneath; panicles rather open, 
many-flowered, the branches puberulent; hypanthium densely 
pubescent, the calyx teeth broadly triangular, acute; corolla densely 
tomentose, the tube 8 mm. long, the narrow lobes 7 mm. long. 
Neg. 126. 

Puno: Mountains near Sangaban, Lechler, type; photo, seen ex 
hb. Berol. 

The species was based upon scant material and its status is 
uncertain. 

Ladenbergia graciliflora Schum. in Mart. Fl. Bras. 6, pt. 6: 
145. 1889. 

A small tree, the branchlets rufous- tomentulose; stipules lance- 
oblong, more than 2 cm. long, attenuate; leaves slender-petiolate, 
subcoriaceous, the blades oblong or ovate, 11-13 cm. long, 5-6.5 
cm. wide, acute or acuminate, obtuse or rounded at the base, lustrous, 
at first puberulent but soon glabrate; panicles 6-8 cm. long, dense, 



FLORA OF PERU 43 

many-flowered, the branches puberulent, the flowers short-pedicel- 
late; calyx lobes triangular-lanceolate, glabrous or nearly so; corolla 
white, 3-3.5 cm. long, tomentulose, the narrow lobes less than half 
as long as the tube; capsule linear, glabrous, 4-5 cm. long. Negs. 
154, 141. 

San Martin: Near Tarapoto, Spruce 4581, type collection. Also 
in Goyaz, Brazil. 

Ladenbergia heterophylla (Wedd.) Standl. Field Mus. Bot. 
7: 18. 1930. Cascarilla heterophylla Wedd. Ann. Sci. Nat. III. 10: 
10. 1848. Buena heterophylla Wedd. Journ. Linn. Soc. 11: 186. 1869. 

Branchlets pubescent; stipules ovate or ovate-oblong, obtuse; 
leaves short-petiolate, submembranaceous, the blades rounded- 
ovate, 10-30 cm. long, obtuse or acutish, glabrate above, pilosulous 
beneath in the axils of the nerves; panicle corymbose, the branches 
tomentulose; calyx lobes rounded-ovate, obtuse; corolla twice as 
long as the calyx, tomentulose; capsule 10-15 cm. long, 12-15 mm. 
wide. 

Reported by Weddell from Peru, without indication of the local- 
ity; also in Colombia. 

I have seen no material representative of this species. 

Ladenbergia magnifolia (R. & P.) Klotzsch in Hayne, Arz- 
neigew. 14: sub. pi. 15. 1846. Cinchona magnifolia R. & P. Fl. 2: 
53. pi. 196. 1799. Cinchona grandifolia Poir. Encycl. 6: 38. 1804. 
Cinchona caduciflora H. & B. PI. Aequin. 168. 1813. Cascarilla 
magnifolia, C. caduciflora, C. rostrata Wedd. Ann. Sci. Nat. III. 10: 
10. 1848. Cascarilla magnifolia var. vulgaris, var. caduciflora, var. 
rostrata Wedd. Hist. Nat. Quinq. 79. 1849. Cinchona lutescens Ruiz 
ex Vitm. Summa, Suppl. 1: 262. 1802. 

A small or medium-sized tree with cinnamon-brown bark; 
stipules 2-3 cm. long, acute, sericeous; leaves long-petiolate, cori- 
aceous, the blades elliptic to oblong-ovate, 10-30 cm. long, 7-20 cm. 
wide, acute or obtuse, acute to rounded at the base, glabrous above 
and often shining, beneath pubescent or usually almost glabrate but 
barbellate in the axils of the nerves; panicles usually large and open, 
many-flowered; calyx lobes triangular-ovate, acute, tomentulose; 
corolla white or pink, sericeous, 1.5 cm. long, the lobes about equaling 
the tube; capsule linear-oblong or lanceolate, 2.5-7 cm. long, 7-10 
mm. broad, glabrate. Negs. 156-158, 575. 

Huanuco: Described from Chinchao, Cochero, and Chacahuasi, 
Ruiz & Pavdn; authentic material seen, ex hb. Berol. Posuso, 600 



44 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

meters, 4705. Cajamarca: Jae"n, Humboldt & Bonpland, type 
material of C. caduciflora; seen in hb. Deless. Junin: La Merced, 
1,200 meters, 5759, 5725. Chanchamayo Valley, 1,500 meters, 
Schunke 270. Loreto: Balsapuerto, 220 meters, Klug 3035. Puma- 
yacu, Klug 3142. Iquitos, in forest, Williams 3781. Mishuyacu, 
Klug 1345, 1387; Killip & Smith 29952. San Martin: Tarapoto, 
Mt. Campana, Spruce 484$- Bolivia to Colombia. "Cascarilla boba," 
"cascarilla amarilla," "cascarilla flor de azahar." 

The tree once was believed to furnish quinine, but it was found 
later that its bark possessed no value as a drug. The tree is said 
to be highly ornamental when covered with its handsome white 
flowers, which exhale an odor suggestive of orange or jasmine 
blossoms. 

Ladenbergia malacophylla Standl. Field Mus. Bot. 8: 157. 
1930. 

A tree 6-12 meters high, the branchlets densely tomentose; leaves 
petiolate, firm-membranaceous, the blades broadly oval or elliptic, 
22-26 cm. long, 14-18 cm. wide, obtuse or almost rounded at the 
apex, broadly rounded or shallowly cordate at the base, pilosulous 
above, velutinous-pilose beneath; panicles rather large and many- 
flowered, the branches fulvous-tomentose, the flowers sessile or 
short-pedicellate; hypanthium densely pilose, the calyx 2-2.5 mm. 
long, the lobes ovate or triangular, acute or subobtuse; corolla white, 
densely sericeous, the slender tube 22-24 mm. long, the lobes half 
as long; capsule oblong, 2.5-4.5 cm. long, sparsely puberulent or 
pilosulous. 

Junin: Porvenir, Pichis Trail, in dense forest, 1,500-1,900 meters, 
Killip & Smith 25914, type; 25434- Ecuador. 

Ladenbergia pedunculata (Karst.) Schum. in Mart. Fl. Bras. 
6, pt. 6: 146. 1889. Cinchona pedunculata Karst. in Koch & Fint. 
Wochenschr. 2: 30. 1859; Karst. Fl. Columb. 1: 53. pi. 36. 1859. 
Remijia pedunculata Flueck. Chinarinde 17. pi. 6. 1883. 

A small tree 3-4.5 meters high, the young branches sericeous; 
stipules obovate, sericeous-pilose; leaves petiolate, coriaceous, 
the blades lanceolate to elliptic or ovate, 6-17 cm. long, 3-8 cm. wide, 
acuminate, at the base attenuate to subcordate, when young minutely 
appressed-pilose, pilose beneath in the axils of the nerves; inflores- 
cences terminal and axillary, long-pedunculate, the branches pilo- 
sulous, the flowers subsessile; calyx lobes ovate or subulate, glabrous, 
sometimes ciliate; corolla white, 7-10 mm. long, appressed-pilose, 



FLORA OF PERU 45 

the lobes equaling the tube; capsules 1-2 cm. long, 6 mm. broad, 
glabrous. 

San Martin: Tarapoto, Spruce 4263. Also in Colombia. 

Ladenbergia Riveroana (Wedd.) Standl. Field Mus. Bot. 7: 201. 
1931. Cinchona oblongifolia Lamb. 111. Cinch. 12. 1821, non Mutis, 
1793. Cascarilla Riveroana Wedd. Ann. Sci. Nat. III. 10: 11. 1848. 

A tree, the branchlets ferruginous-tomentose or finally glabrate; 
stipules ovate-oblong, tomentose; leaves thin-coriaceous, short- 
petiolate, the blades oblong-lanceolate or ovate-oblong, 15-25 cm. 
long, 9-15 cm. wide, acute at each end or more rarely obtuse, puberu- 
lent or glabrate above, tomentose or puberulent beneath, with rusty 
pubescence; panicles rather large and many-flowered, ferruginous- 
tomentose; hypanthium tomentose, the calyx teeth ovate, acutish, 
tomentulose; corolla pilose, the tube 7-15 mm. long, the lobes 6-8 
mm. long; capsule linear-lanceolate, 4-6 cm. long, ferruginous- 
tomentose. Neg. 113. 

Junin: Huacapistana, 1,800-2,400 meters, Killip & Smith 24214, 
a shrub or tree 3-3.5 meters high. San Martin: Zepelacio, 1,200- 
1,600 meters, Klug 3515. Bolivia to Ecuador. "Azahar macho," 
"azahar hembra." 

Klug describes the species as a tree of 5 meters with white flowers. 

Ladenbergia stenocarpa (Lamb.) Klotzsch in Hayne, Arz- 
neigew. 14: sub pi. 15. 1846. Cinchona stenocarpa Lamb. 111. Cinch. 
13. 1821. Cascarilla stenocarpa Wedd. Hist. Nat. Quinq. 81. 1849. 
Buena stenocarpa Wedd. Journ. Linn. Soc. 11: 186. 1869. 

Branchlets glabrate; stipules ovate; leaves short-petiolate, sub- 
membranaceous, the blades oblong-lanceolate, acute at each end, 
12-15 cm. long, glabrous or pilose beneath along the costa; panicle 
open, many-flowered, the branches puberulent; hypanthium tomentu- 
lose, the calyx teeth ovate, acutish; corolla puberulent, the tube 13- 
15 mm. long, the lobes 9-10 mm. long; capsule linear, 3.5 cm. long. 
Neg. 135. 

Cajamarca: Type from Jae"n, Pavdn; photo, of authentic material 
seen, ex hb. Berol. "Cascarilla azaharito." 

12. C API RON A Spruce 

Trees with large stipules; leaves large, opposite, petiolate; 
inflorescence terminal, paniculate, the flowers showy, 5-parted, 
zygomorphic; calyx cupular, 5-dentate, one of the teeth often 



46 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

expanded into a large, foliaceous, colored limb; corolla somewhat 
gibbous from a tubular base, the limb campanulate and subbilabiate, 
the short lobes contorted in bud ; stamens inserted above the base of 
the tube, the filaments villous at the base, the anthers linear; capsule 
clavate or obovate, 2-celled, septicidally bivalvate; seeds numerous, 
winged. 

Capirona decorticans Spruce, Journ. Linn. Soc. 3: 200. 1859; 
Schum. in Mart. Fl. Bras. 6, pt. 6: 195. pi. 107. 1888. 

A tree 13-16 meters high with reddish brown bark; stipules more 
or less foliaceous, acute or acuminate, 3-6 cm. long; leaves almost 
sessile, oblong to oval, 22-45 cm. long, obtuse or short-acuminate, 
very obtuse to subcordate at the base, almost glabrous; panicles 
15-20 cm. long or more; calyx 3-4 mm. long, the expanded lobe obo- 
vate-spatulate, 5-7 cm. long, red, attenuate at the base; corolla 
2.5-3 cm. long, the lobes ovate, obtuse; capsule 2-3 cm. long, woody. 
"Capirona negra." 

San Martin: Near Tarapoto, Spruce 4202, type collection. 
Tarapoto, in forest, Williams 6070. Also in Colombia. 

13. HILLIA Jacq. 

Shrubs, usually epiphytic, glabrous, with thick branches; stipules 
membranaceous, caducous; leaves opposite, fleshy, drying thick and 
hard; flowers large and showy, solitary or clustered, surrounded by 
an involucre of leaves or bracts, 4-6-parted ; calyx none or composed 
of 2-6 free or short-connate segments, persistent or caducous; corolla 
salverform or funnelform-tubular, the lobes contorted, the throat 
glabrous; stamens inserted below the throat of the corolla, included, 
the anthers oblong; fruit a cylindric capsule, somewhat attenuate 
at each end, septicidally bivalvate; seeds fusiform, bearing a tuft of 
hairs at one end. 

Leaves small, mostly 7-8 mm. wide -. H. Macbridei. 

Leaves comparatively large, commonly 2-8 cm. wide. 

Corolla tube conspicuously dilated in the throat and 1.5-2.5 cm. 

wide ; calyx lobes persistent. 
Leaf blades rounded or very obtuse at the base, mostly 2-3.5 cm. 

wide; corolla about 4 cm. long H. Ulei. 

Leaf blades acute at the base, 4-7 cm. wide; corolla about 5.5 

cm. long H. illustris. 

Corolla tube slender, scarcely dilated in the throat and less than 
1 cm. wide; calyx lobes deciduous. 



FLORA OF PERU 47 

Veins of the leaves conspicuous; lobes of the corolla about half 
as long as the tube H. Killipii. 

Veins of the leaves obscure or obsolete; lobes of the corolla less 
than one-third as long as the tube. 

Leaf blades elliptic or oblong-elliptic, rounded to acute at the 
base H. parasitica. 

Leaf blades narrowly oblanceolate-oblong, attenuate to the 
base H. Weberbaueri. 

Hillia illustris (Veil.) Schum. in Mart. Fl. Bras. 6, pt. 6: 202. 
1889. Saldanha illustris Veil. Fl. Flum. 3: pi. 157. 1827; text, ed. 
Netto 134. 

Leaves short-petiolate, thick and hard when dried, the blades 
elliptic or broadly elliptic-ovate, 10-16 cm. long, abruptly acuminate; 
flowers solitary, subtended by 2 large stipules, these 1.5-2 cm. long, 
obtuse; calyx lobes 6, linear-subulate, herbaceous, 1.5-2 cm. long; 
corolla white, the short lobes ovate-rounded, recurved; capsule 10 
cm. long and 1 cm. thick. Neg. 234. 

Loreto: Rio Itaya, in forest, Williams 99. Extending to Brazil 
and the Guianas. 

Hillia Killipii Standl. Field Mus. Bot. 8: 159. 1930. 

Described as a tree 3-4.5 meters high; stipules narrowly oblong, 
rounded at the apex, 2.5 cm. long; petioles slender, 1-1.5 cm. long; 
leaf blades comparatively thin, elliptic-oblong, 7.5-9.5 cm. long, 
3-3.5 cm. wide, abruptly long-acuminate, acute at the base; flowers 
terminal, solitary, sessile, subtended by 2 stipules; calyx obsolete; 
corolla white, the slender tube 8 cm. long, the 6 lobes lance-linear, 
4-4.5 cm. long. 

Junin: Between San Nicolas and Azupizu, in dense forest, 650- 
900 meters, Killip & Smith 26085, type. Loreto: Pumayacu, 
100-600 meters, a liana on a tree, Klug 3145. 

Hillia Macbridei Standl. Field Mus. Bot. 4: 277. 1929. 

A small epiphytic shrub; stipules lanceolate or linear-lanceolate, 
caducous, 1.5-2 cm. long; petioles 3-5 mm. long; leaf blades lanceo- 
late, 2-3 cm. long, narrowed to the subobtuse apex, acute at base, 
the veins obsolete; flowers terminal, solitary, sessile; calyx persistent, 
the 5 lobes oblong-ovate, obtuse, 5 mm. long; capsule narrowly 
clavate, 5.5-6.5 cm. long, 1 cm. wide. 



48 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Junin: La Merced, 1,200 meters, Macbride 5760, type. Chan- 
chamayo Valley, 1,500 meters, Schunke 436. Above San Ramon, 
1,300-1,700 meters, Schunke AlOO. 

Hillia parasitica Jacq. Enum. PI. Carib. 18. 1760. Cosmibuena 
acuminata R. & P. Fl. 3: 4. pi. 226. 1802. Buena acuminata DC. 
Prodr. 4: 356. 1830. H. odorata Krause, Bot. Jahrb. 40: 321. 1908. 

A glabrous, epiphytic shrub, sometimes as much as 2 meters high, 
with stout branches; stipules 8-12 mm. long or more, obtuse; pet- 
ioles thick, 2-7 mm. long; leaf blades mostly 5-11 cm. long, short- 
acuminate; flowers solitary, sessile; calyx lobes linear-lanceolate, 
obtuse, 4 mm. long; corolla white, the tube 6.5-12 cm. long, the 6 
lobes linear-lanceolate to oblong; capsule cylindric, 7-9 cm. long. 
Negs. 235, 236 (H. odorata}. 

Amazonas: East of Chachapoyas, Weberbauer 4364- Huanuco: 
Type of Cosmibuena acuminata from Chicoplaya, Ruiz & Pavon. 
Junin: Chanchamayo Valley, Schunke 290. Huacapistana, 1,800 
meters, Weberbauer 2152, type of H. odorata (photo, and fragm. seen, 
ex hb. Berol.). Ranging to Brazil, Mexico, and the West Indies. 

Hillia Ulei Krause, Verh. Bot. Ver. Brandenb. 50: 97. 1908. 

A glabrous epiphyte with stout branches; leaves short-petiolate, 
the blades broadly ovate or elliptic, 4-6.5 cm. long, obtusely short- 
acuminate or acute, rather conspicuously palmate-nerved, very 
thick; flowers terminal, solitary, pedicellate; calyx lobes 6, linear, 
10-12 mm. long; corolla grass-green, the tube gradually narrowed 
from throat to base, the lobes short and broad ; capsule 8-10 cm. long. 
Neg. 237. 

Loreto: Near Yurimaguas, Ule 6303, type (photo, and fragm. 
seen ex hb. Berol.). Mishuyacu, near Iquitos, 100 meters, Klug 
1182. 

Hillia Weberbaueri Standl. Field Mus. Bot. 11: 214. 1936. 

A glabrous epiphyte with stout branches; stipules oblong, 
obtuse, 18 mm. long; leaves short-petiolate, the blades narrowly 
oblanceolate-oblong, 8.5-13.5 cm. long, 2.5-3.5 cm. wide, abruptly 
rather long-acuminate, long-attenuate to the base, with about 8 
pairs of lateral nerves; flowers terminal, solitary; calyx none or 
caducous; corolla 6-parted, the very slender tube 7 cm. long, the 
lobes oblong, obtuse, 1.5 cm. long. 

Without locality: Weberbauer 6955, type. 



FLORA OF PERU 49 

14. COSMIBUENA R. & P. 

Glabrous shrubs, usually epiphytic, often scandent; stipules 
caducous; leaves opposite, thick and fleshy; flowers large and 
showy, pedicellate, solitary or clustered, 5-6-parted; calyx 5-6- 
lobate, circumscissile at the base; corolla salverform, the lobes 
contorted, fleshy, the tube rather slender and much elongate; stamens 
inserted below the throat of the corolla, the anthers linear; capsule 
oblong or cylindric, 2-celled; seeds numerous, oblong or lanceolate, 
winged, the wings fimbriate or erose at the apex. 

Cosmibuena grandiflora (R. & P.) Rusby, Bull. N. Y. Bot. 
Card. 4: 368. 1907. Cinchona grandiflora R. & P. Fl. 2: 54. pi. 
198. 1799. C. obtusifolia R. & P. Fl. 3: 3. pi. 198. 1802. Buena 
obtusifolia DC. Prodr. 4: 356. 1830. 

A small or large, epiphytic shrub; leaves on stout petioles, the 
blades elliptic to obovate, 8-13 cm. long or larger, rounded at the 
apex, acute at the base, very thick, the nerves inconspicuous; calyx 
10-12 mm. long, tubular below, soon deciduous; disk large and 
conspicuous; corolla white, the tube 6-9 cm. long, the lobes oblong, 
obtuse, about 2.5 cm. long; capsule 4-5 cm. long. Neg. 240. 

Huanuco: Type from Posuso, Ruiz & Pavon. Monzon, 900 
meters, Weberbauer 3502. Junin: Reported by Ruiz and Pavon 
from Pueblo Nuevo de San Antonio de Chicoplaya. Loreto: 
Moyobamba, 800 meters, Weberbauer JtflS. San Martin: Near 
Tarapoto, Spruce 4179. Department uncertain: Without locality, 
Pavdn; Maclean. Ranging from Colombia to Bolivia. 

15. CALYCOPHYLLUM DC. 

Trees with opposite leaves; stipules caducous; flowers small, 
4-8-parted, arranged in terminal cymes, the bracts and bractlets 
large and membranaceous and enclosing the buds; calyx almost 
obsolete or of short segments, one of the segments in some of the 
flowers often expanded into a large, colored limb; corolla short- 
funnelform, lobed to the middle or more deeply, the segments 
imbricate in bud, the throat villous; stamens inserted in the throat, 
long-exserted ; capsule costate, elongate, 2-celled, septicidally bival- 
vate; seeds small, winged. 

Calycophyllum Spruceanum (Benth.) Hook. f. ex Schum. in 
Mart. Fl. Bras. 6, pt. 6: 191. pi. 106. 1889. Eukylista Spruceana 
Benth. Kew Card. Misc. 5: 230. 1853. 



50 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tree 15-27 meters high, with brown bark; leaves petiolate, 
the blades oblong to oblong-ovate, acute or obtuse, acute to obtuse 
at the base, 9-17 cm. long, minutely puberulent beneath at first but 
soon glabrate, barbate in the axils of the nerves; cymes dense and 
many-flowered, the inflorescences at first wholly enclosed by the 
thin bracts; calyx 6-9-dentate, the lobes all minute; corolla white, 
6-7 mm. long, the lobes spreading; hypanthium densely white- 
pilosulous; capsule oblong, 8-11 mm. long, densely appressed- 
pilose. Neg. 8601. 

Loreto: La Victoria on the Amazon, in forest, Williams 3010, 
2855. Lower Rio Nanay, Williams 455. Iquitos, Williams 8033. 
Yurimaguas, Williams 4486. Rio Huallaga, Spruce. Rio Ucayali, 
Tessmann 3378. Balsapuerto, 220 meters, Klug 3005. Rancho 
Indiana, 110 meters, Mexia 6458. Florida, Rio Putumayo, riverside 
forest, 180 meters, Klug 2190. Fortaleza, 140 meters, Klug 2814. 
Also in eastern Bolivia and Amazonian Brazil. 

"Capirona." Huber reports the tree as common along the 
lower Ucayali, where it forms forests known as capironales. The 
Brazilian name is "pao mulato." The wood is said to be strong, 
yellowish blackish, and useful for making various utensils and for 
construction purposes. It also is used extensively as firewood. Mrs. 
Mexia states that the bark is glossy and dark red, and that the 
flowers are slightly fragrant. 

16. LORETOA Standl. 

Large, glabrous trees; stipules large, persistent; leaves opposite, 
petiolate, coriaceous; flowers large, 5-parted, sessile or pedicellate, 
arranged in a large, terminal, sessile panicle composed of numerous 
open cymes; calyx cupular, shallowly 5-6-dentate, densely sericeous 
within; corolla clavate-funnelform, the short lobes contorted; anthers 
basiflxed, on slender filaments inserted above the base of the tube; 
ovary 2-celled, the ovules peltately imbricate, winged. The genus 
consists of a single known species. 

Loretoa peruviana Standl. Field Mus. Bot. 11: 222. 1936. 

A tree 20 meters high; stipules 3.5 cm. long, very obtuse or 
acutish; leaf blades broadly elliptic or suborbicular, 15-18 cm. 
long, 12 cm. wide, rounded at the apex, very obtuse or rounded at 
the base; panicle 25 cm. long, with numerous large, spreading bracts, 
the pedicels bracteolate; calyx 3 mm. high and 7-10 mm. wide; 
corolla rose-garnet, almost 4 cm. long, the tube 12 mm. wide near the 
middle, the rounded lobes 8 mm. long. 



FLORA OF PERU . 51 

Loreto: Florida on the Rio Putumayo, 200 meters, in forest, 
King 2022, type. 

Called "meta guais." The fruit of this tree is not known, and 
there is consequently some doubt regarding the proper position of the 
genus. 

17. FERDINANDUSA Pohl 

Trees or shrubs, glabrous or sparsely pubescent; stipules caducous; 
leaves opposite, often coriaceous, short-petiolate; flowers 4-5- 
parted, cymose-paniculate; calyx cupular, shortly dentate; corolla 
funnelform, the lobes sometimes recurved, bifid at the apex, con- 
torted in bud, glabrous; anthers sessile, exserted or included; ovary 
turbinate or oblong, 2-celled ; capsule globose to cylindric, bisulcate, 
bivalvate from the apex; seeds winged. 

Corolla tube 3-4.5 cm. long; leaves with short scattered hairs on the 

lower surface F. loretensis. 

Corolla tube less than 1.5 cm. long; leaves glabrous. . .F. chlorantha. 

Ferdinandusa chlorantha (Wedd.) Standl. Trop. Woods 
34: 41. 1933. Gomphosia chlorantha Wedd. Ann. Sci. Nat. III. 10: 
14. 1848. G. laxiflora Benth. in Hook. Kew Journ. 5: 232. 1853. F. 
andina Wedd. Ann. Sci. Nat. IV. 1: 78. 1854. 

A shrub or tree 3-20 meters high or more, glabrous throughout; 
stipules lanceolate; leaves very shortly petiolate, the blades coria- 
ceous, ovate to oblong, 7-12 cm. long, acuminate, rounded to acutish 
at the base; panicles small and dense, leafy at the base; calyx lobes 
very short, acute or acuminate; corolla glabrous, white or greenish, 
the lobes short, rounded; capsule oblong, 1.5-2.5 cm. long, 5-7 mm. 
wide. Neg. 249. 

Loreto: Forest between Rio Nanay and Rio Napo, Williams 
697. Mishuyacu, near Iquitos, 100 meters, in forest, Killip & Smith 
29902; King 294, 139.Puno: Province of Carabaya, Weddell (photo, 
of type seen, ex hb. Berol.). Perhaps also in Bolivia. 

"Louro-micuna," "guacamayo." It is somewhat uncertain 
whether the tree of Loreto (which seems to be F. Paxii Winkl.) is 
really identical with F. chlorantha, but a reading of Weddell's descrip- 
tions and examination of a photograph of the type reveal no impor- 
tant differences. Weddell states that the tree. is a typical one of the 
regions in which it grows, its clean, silvery trunk rising well above 
the surrounding trees. 



52 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ferdinandusa loretensis Standl. Field Mus. Bot. 8: 337. 1931. 

A tree 6 meters high, the branchlets sparsely puberulent or 
glabrate; leaves coriaceous, short-petiolate, the blades oblong- 
elliptic, 11-15 cm. long, abruptly short-acuminate, rounded at the 
base or abruptly contracted; inflorescence small and few-flowered, 
short-pedunculate, the flowers pedicellate; calyx teeth triangular, 
acute; corolla white, glabrous, the 4 lobes 8-10 mm. long; capsule 
oblanceolate-oblong, 3.8 cm. long, 1.4 cm. wide. 

Loreto: Mishuyacu, near Iquitos, 100 meters, in forest, Klug 
1348, type. 

18. EXOSTEMA Rich. 

Shrubs or trees; stipules distinct or short-connate, persistent or 
deciduous; leaves opposite, petiolate; flowers small or large, axillary 
and solitary or arranged in panicles, 5-6-parted ; calyx campanulate, 
lobate; corolla salverform or funnelform, the tube long or much 
elongate, glabrous or pilose in the throat, the lobes linear, imbricate 
in bud; stamens exserted, the anthers linear; capsule 2-celled, cori- 
aceous or woody, cylindric to obovate, septicidally bivalvate; seeds 
imbricate, winged. 

Corolla 10-12 cm. long E. maynense. 

Corolla 1.5-2.5 cm. long. 

Leaf blades truncate or subcordate at the base . . . . E. peruvianum. 

Leaf blades mostly acute or acutish at the base . .E. corymbosum. 

Exostema bicolor P. & E. Nov. Gen. & Sp. 3: 32. 1845. 

A small tree; leaf blades very broadly ovate or elliptic, acuminate, 
about 10 cm. long and 7.5 cm. wide, glabrous; flowers pubescent, in 
terminal, trichotomous corymbs; calyx lobes linear; corolla purplish 
green, pubescent, about 2.5 cm. long, the lobes oblong, equaling the 
tube; capsule rounded, compressed. Neg. 8603. 

Huanuco: Type from Cerro San Cristobal, near Cochero, 
Poeppig. Without locality: Poeppig 1352 (probably the original 
collection). 

The species is not included in the key because I am unable to 
separate it, by description, from E. peruvianum and E. corymbosum. 
Probably it is not specifically distinct from E. peruvianum. 

Exostema corymbosum (R. & P.) Spreng. Syst. Veg. 1: 706. 
1825. Portlandia corymbosa R. & P. Fl. 2: 49. pi. 190, j. a. 1799. 



FLORA OF PERU 53 

A tree 4-6 meters high ; leaves short-petiolate, subcoriaceous, the 
blades ovate to oblong, mostly 7-10 cm. long, acute or acuminate, 
glabrous or sparsely pubescent beneath; flowers in terminal, leafy, 
many-flowered corymbs; calyx lobes elongate, linear; corolla white or 
purplish white, pubescent outside, about 2 cm. long, the lobes linear, 
equaling the tube; capsule ferruginous, 8 mm. long. Neg. 8602. 

Cajamarca: Province of Hualgayoc, Weberbauer 1+166. Huan- 
cavelica: Tayacaja, Weberbauer 6482. Huanuco: Described from 
Chaclla and Muna, Ruiz & Pavdn. Mufia, 2,100 meters, dry steep 
slope, 3972. 

Exostema maynense P. & E. Nov. Gen. & Sp. 3: 31. 1845. 

A tree, sometimes 20 meters high, with red wood ; leaves petiolate, 
rather thin, oblong to broadly ovate, acuminate, glabrous, 7-15 
cm. long; flowers cymose-corymbose, terminal; calyx 5-6-dentate, 
the teeth broad, short-acuminate; corolla 10-12 cm. long, the lobes 
linear, one-third as long as the tube; capsule obovoid, acute at the 
base, 3 cm. long. Neg. 251. 

Loreto: Forests of Mainas near Yurimaguas, Poeppig (photo, of 
type seen, ex hb. Berol.). Mainas, Poeppig D2392. Middle Mara- 
fion, Tessmann 4888. Balsapuerto, 220 meters, Klug 3064; a tree of 
15 meters with white flowers. Without locality: Poeppig 3076, 
probably the type collection. 

"Puca yanta." Poeppig reports that the very bitter bark was 
used in domestic medicine. The flowers are said to have the odor 
of orange blossoms. 

Exostema peruvianum Humb. & Bonpl. PI. Aequin. 1: 133. pi. 
38. 1808. 

A shrub 3-3.5 meters high, the trunk 10 cm. in diameter; leaves 
ovate or broadly oblong, coriaceous, acute, 5-7.5 cm. long, glabrous, 
the upper ones sessile and cordate at the base; flowers fragrant, 
in dense, leafy, terminal corymbs; calyx lobes lanceolate, acute; 
corolla pink, pubescent, 2 cm. long, the lobes linear, spreading, equal- 
ing the tube. Neg. 563. 

Huanuco: Casapi, Mathews 1450. Type from the Andes of Peru 
(photo, seen, ex hb. Berol.). Department uncertain: Callcate, 
Jelski 366. 

Probably this is not distinct from E. corymbosum. 



54 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

19. COUTAREA Aubl. 

Shrubs or trees, the branches with conspicuous elevated lenticels; 
stipules persistent; leaves opposite, short-petiolate, rather thin; 
inflorescence terminal or axillary, the flowers mostly solitary or 
ternate, usually large and showy, asymmetric, 5-8-parted; calyx 
cupular, lobate; corolla tubular-campanulate, often gibbous and 
somewhat bilabiate, the lobes contorted or imbricate, the throat 
glabrous; stamens inserted at the base of the tube, included or 
exserted, the anthers linear; capsule compressed contrary to the 
partition, obovate, woody or coriaceous; seeds broadly winged. 

Coutarea hexandra (Jacq.) Schum. in Mart. Fl. Bras. 6, pt. 6: 
196. 1889. Portlandia hexandra Jacq. Sel. Stirp. 63. pi. 182, f. 20. 
1763. C. speciosa Aubl. PI. Guian. 1: 314. pi. 122. 1775. Bignonia 
triflora Pav. ex DC. Prodr. 9: 148. 1845, in syn. 

A shrub or small tree; leaves lanceolate to ovate or elliptic, 
mostly 7-13 cm. long, acute or acuminate, acute to rounded at the 
base, nearly glabrous; flowers chiefly in terminal clusters of 2-3; 
corolla whitish or greenish, 4-5 cm. long; capsules 2-3.5 cm. long, 
broadly rounded at the apex, marked with numerous pale lenticels. 

Loreto: Rio Masana, Williams 158. Without locality, Pavdn; 
type material of Bignonia triflora. Widely distributed in tropical 
America. 

The bark is bitter, and in some regions it has been employed in 
domestic medicine as a substitute for quinine. In Brazil the species 
is called "quina do Piauhy," "quina do Pernambuco," and "murta 
do mato." 

Coutarea hexandra var. tarapotensis Schum. (in Mart. Fl. Bras. 
6, pt. 6: 198. 1889) was based upon Spruce 4&43 from Tarapoto. It is 
merely a form with unusually large leaves. 

20. POGONOPUS Klotzsch 

Shrubs or trees; stipules almost free, deciduous; leaves opposite, 
herbaceous, petiolate; flowers showy, in opposite-branched panicles, 
5-parted ; calyx tubular, lobate, one of the lobes often expanded into 
a large, brightly colored limb; corolla tubular, the short lobes 
valvate in bud, the tube somewhat curved, pilose within above the 
base; stamens inserted in the upper part of the tube, unequal, the 
anthers sagittate; capsule subglobose, 2-celled, loculicidally bival- 
vate; seeds numerous, minute, horizontal, compressed and marginate. 



FLORA OF PERU 55 

Pogonopus tubulosus (DC.) Schum. in Mart. Fl. Bras. 6, 
pt. 6: 265. 1889. Calycophyllum tubulosum DC. Prodr. 4: 367. 1830. 

A shrub or tree 2-5 meters high or larger; leaves short-petiolate, 
the blades ovate, obovate, or elliptic, large, short-acuminate, densely 
pubescent or glabrate; stipules 3-4 mm. long; calyx 2 mm. long, the 
lobes subulate; corolla pale lilac, 3.5 cm. long, minutely puberulent or 
glabrate; enlarged sepal with a rounded, pink blade as much as 12.5 
cm. broad ; capsule woody, 7 mm. long. 

Cuzco: Chanchamayo, Lares Valley, 1,000 meters, Weberbauer 
7940. Junin: La Merced, 700 meters, in thickets, Killip & Smith 
23811. Madre de Dios: Seringal San Francisco, Rio Acre, Ule 
9848. Also in Brazil, Bolivia, and Argentina. "Quina" (Bolivia). 

The tree is an exceedingly showy one when in flower, because of 
the enlarged and brilliantly colored calyx lobes. 

21. CONDAMINEA DC. 

Shrubs or trees; stipules often bipartite; leaves opposite, often 
large and coriaceous; flowers mostly large, 5-parted, disposed in 
large, terminal panicles; calyx campanulate, truncate or dentate, 
deciduous; corolla salverform, pubescent in the throat, the lobes 
valvate; stamens inserted in the corolla throat, the filaments subulate, 
pilose at the base; capsule usually pyriform and truncate at the 
apex, bisulcate, loculicidally bivalvate, more or less woody; seeds 
horizontal, minute, angled, compressed. 

Calyx scarcely 1 mm. long. Leaves petiolate C. microcarpa. 

Calyx 5-7 mm. long. 

Leaves sessile or nearly so, acuminate C. corymbosa. 

Leaves long-petiolate, very obtuse C. macrophylla. 

Condaminea corymbosa (R. & P.) DC. Prodr. 4: 402. 1830; 
Schum. in Mart. Fl. Bras. 6, pt. 6: 257. pi. 124. 1889. Macrocnemum 
corymbosum R. & P. Fl. 2: 48. pi. 189. 1799. 

A shrub or small tree 1-6 meters high, glabrous or nearly so, with 
stout branches; leaves mostly sessile and clasping, oblong to obovate, 
20-60 cm. long or larger, cordate at the base, coriaceous; cymes 
corymb-like, large, long-stalked, many-flowered; corolla 2.5 cm. long, 
reddish outside, whitish or cream-colored within; capsules about 
1.7 cm. long, obtuse or acutish at the base. Neg. 567. 

Huanuco: Described from Chinchao, Acomayo, Pillao, and 
Muna, Ruiz & Pavdn. Yanano, 1,800 meters, open hillside, 3662. 



56 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pampayacu, Kanehira 107. Junin: Huacapistana, 2,000 meters, 
Killip & Smith 24130. Chanchamayo, Martinet 1412. La Merced, 
700 meters, wooded valley, Killip & Smith 23506. San Martin: 
San Roque, 1,400 meters, in forest, Williams 7794- Bolivia to 
Panama. 

"Sauco" (Williams) ; "ccaratu" (Ruiz & Pavon). Ruiz and Pavon 
state that the bark, although not very bitter, was used as an adulter- 
ant of cinchona bark. 

Condaminea corymbosa var. pubescens Spruce ex Schum. in 
Mart. Fl. Bras. 6, pt. 6: 258. 1889. C. angustifolia Rusby, Mem. 
Torrey Club 6: 45. 1896. 

Leaves densely and finely pubescent beneath, often short-petiolate; 
inflorescence finely and densely pubescent. 

Ayacucho: Carrapa, 1,000 meters, wooded hillside, Killip & 
Smith 22449. Cuzco: San Miguel, Urubamba Valley, 1,800 meters, 
Cook & Gilbert 913. San Martin: Near Tarapoto, Spruce 4579, type. 
Zepelacio, 1,100 meters, a tree of 5 meters, Klug 3672. Also in 
Bolivia. 

Condaminea glabrata DC. Prodr. 4: 402. 1830. Macrocnemum 
glabratum Bartl. ex DC. loc. cit. as syn. 

Leaves obovate, short-acuminate, cuneate-attenuate at the base, 
glabrous; panicle glabrous, the flowers glomerate. 

Huanuco: Type from the mountains of Huanuco, Haenke. 

The species is known to the writer only from the brief original 
description. 

Condaminea macrophylla P. & E. Nov. Gen. & Sp. 3: 30. 1845. 

A tree 10-13 meters high; leaves long-petiolate, the blades oval, 
30-40 cm. long, about 25 cm. wide, very obtuse, rounded or cordate 
at the base, glabrous; inflorescence 20-30 cm. long, the flowers 
pedicellate; calyx irregularly dentate, sericeous within. 

Loreto: Type from forests of Yurimaguas, Poeppig 2463. 

Schumann (in Mart. Fl. Bras.) states that this plant may be a 
Rustia rather than a Condaminea. 

Condaminea microcarpa (R. & P.) DC. Prodr. 4: 402. 1830. 
Macrocnemum microcarpum R. & P. Fl. 2: 49. pi. 188, f. a. 1799. 

A slender shrub or small tree; leaves petiolate, the blades elliptic 
or oblong-elliptic, obtusely acuminate, obtuse at the base, glabrous 
above, short-pilose beneath; flowers racemose-paniculate, sessile, 



FLORA OF PERU 57 

clustered; bracts ovate, obtuse, ciliate; corolla small, white; calyx 
cupular, 0.8 mm. long, shallowly dentate; capsule turbinate or oval, 
3.5-4 mm. long, the seeds minute. 

Huanuco: Described from Chinchao and Cochero, Ruiz & Pavdn. 

Although I have seen a specimen of this species, in the Delessert 
Herbarium, I am doubtful as to its proper generic position. It is 
scarcely a species of Condaminea. 

Condaminea venosa (R. & P.) DC. Prodr. 4: 402. 1830. 
Macrocnemum venosum R. & P. Fl. 2: 49. pi 190, f. b. 1799. 

A shrub 3.5 meters high; leaves petiolate, the blades oblong- 
elliptic, acutely acuminate, conspicuously nerved, puberulent on the 
nerves; flowers small, white, sessile, in terminal panicles. 

Cuzco: Acomayo and Pati, Pavdn. 

Scarcely a true Condaminea; perhaps a species of Chimarrhis. 

22. CHIMARRHIS Jacq. 

Trees; stipules interpetiolar, acuminate, caducous; leaves oppo- 
site, short-petiolate; flowers very small, in pedunculate, axillary, 
corymbiform panicles, 5-parted; calyx cupuliform, dentate or 
truncate; corolla short-funnelform, the lobes valvate, usually 
longer than the tube, villous within; stamens inserted in the corolla 
throat, the filaments elongate, villous below, the anthers dorsifixed ; 
capsule small, oblong or subglobose, 2-celled, septicidally dehiscent; 
seeds numerous, minute, horizontal, angulate, reticulate, margined. 

Calyx deeply dentate; leaves not barbate beneath in the axils of the 
nerves C. dioica. 

Calyx shallowly dentate; leaves barbate beneath in the axils of the 
nerves. 

Stipules glabrous; calyx and hypanthium minutely puberulent. 

C. Hookeri. 

Stipules densely sericeous outside; calyx and hypanthium glabrous. 

C. Williamsii. 
Chimarrhis dioica Schum. & Krause, Bot. Jahrb. 40: 312. 1908. 

A shrub 3 meters high, probably also a tree ; leaves short-petiolate, 
the blades oblong or oblong-elliptic, 8-20 cm. long, 4-12 cm. wide, 
acuminate, acute at the base, often bullate and with strongly im- 
pressed veins, pilose or subtomentose beneath on the veins; flowers 
"dioecious," in panicles 3-6 cm. long; calyx lobes ovate, acu