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FLORA OF PERU 



BY 

J. FRANCIS MACBRIDE 

ASSOCIATE CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 




THt UBK/\BV Of THE 

JUL 25 1941 

UNIVERSITY OFJLLINOIS 



S NATURAL 
HISTORY 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IV, NUMBER 1 

JUNE 30, 1941 

PUBLICATION 496 



FLORA OF PERU 



BY 

J. FRANCIS MACBRIDE 

ASSOCIATE CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 



THE UBKARY OF THE 

JUL251941 

UNIVERSITY OF ILLINOIS 




BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART IV, NUMBER 1 

JUNE 30, 1941 

PUBLICATION 496 



PRINTED IN THE UNITED STATES OF AMERICA 

BY FIELD MUSEUM PRESS 

With WPA Assistance 



O u 



FB 



3 
V 

FLORA OF PERU 



J. FRANCIS MACBRIDE 



ELATINACEAE. Elatine or Waterwort Family 
Reference: Niedenzu in Pflanzenfam. ed. 2. 21: 270-276. 1925. 
Small herbs, at least the Peruvian, with opposite leaves, thin 
stipules and minute or inconspicuous axillary actinomorphic hermaph- 
rodite 2-5(-6)-merous flowers. Sepals free or united toward the 
base. Petals and stamens on the receptacle, the inner stamens some- 
times aborted. Styles distinct, the stigmas capitate. Fruit capsular 
with axillary placentae. Seeds longitudinally and transversely 
striate. E. nivalis Speg., Argentinian, has no stipules, according to 
the author. 

ELATINE L. 

Glabrous herbs of shallow pools or wet places, the 2-3-merous 
flowers usually solitary. Sepals united toward the base (at least 
the Peruvian species), membranous as the capsules. The similar 
genus Bergia L. characterized by firm sepals and capsules, the former 
often cusped by the sharp midrib, may be found as an introduction 
since B. verticillata Willd. of Egypt and India has apparently been 
collected on the southern coast of Ecuador. It is a rather coarse 
herb with verticillate pentamerous flowers. 

Elatine peruviana Baehni & Macbr. Candollea 8: 21. 1940. 

Glabra, humifusa; caulibus repentibus et adscendentibus ad 
nodos radicantibus; foliis late ovatis vel obovatis, acutis, in petiolum 
late attenuatis, 2 mm. longis, vix 1.5 mm. latis vel inferioribus fere 
sessilibus et suborbiculatis, 2 mm. latis, integris; floribus solitariis; 
pedunculis 3-4 mm. longis; sepalis 3, ovato-acutis, integerrimis 0.5 
mm. longis; petalis late ovalis minutissime ciliolatis 1.5-2 mm. longis; 
staminibus 3, filamentis e basi valde dilatatis; capsulis depresso- 
globosis; seminibus oblongis leviter curvatis 0.75 mm. longis, longi- 
tudinaliter 6-7-lineatis, transversim lineis elevatis multo-clathrata. 
E. peruviana seems to be nearest E. Lindbergii Rohrb. in Mart. 
Fl. Bras. 14, pt. 2: 321. pi. 72. 1872, but the sepals are entire and 
the leaves not cordate. 

Since writing the above, Fassett, Rhodora 41: 367-376. 1939, 
has presented a revision of the North American species and shown 



4 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

that the fundamental character is found in the seeds. The seeds 
of E. peruviana are about 732 n long, 300 n thick, with about 15 
pits in each row. The type formed mats in a sunny mossy bog, 
the flowers faintly green-tinted, the anthers black. 

Huanuco: Mito, 3,000 meters, 1544, type, Field Museum. 

Elatine triandra Schkuhr, Bot. Handb. 1: 345. pi. 109b. 1791. 

Diminutive herb similar vegetatively to E. peruviana but the 
2-3-celled capsules sessile; leaves linear to spatulate, often emarginate 
at tip; cf. Fassett, I.e. 369. The Peruvian form is probably variety 
andina Fassett, I.e. 374, the seeds 460-680 /* long, 160-280 M thick, 
with 8-10 rows of 12-19 pits each. Type from Sorata, Bolivia. 

Puno: Sachapata, Lechler 2687 (probably, from range but material 
seen meager). Bolivia; Chile. 

FRANKENIACEAE. Frankenia Family 

Reference: Niedenzu in Pflanzenfam. ed. 2. 21: 276-281. 1925. 

More or less suffrutescent herbs, or shrubby, with crowded 
often opposite leaves that, revolute, sometimes appear ericoid. 
Flowers small, solitary or cymose, hermaphrodite, the 4-6 connate 
sepals enclosing the valvular capsule, the same number of imbricate 
petals clawed and with (often) a scale-like appendage within. 
Stamens usually 6, hypogynous. Ovary superior, 1-celled with 2-4 
parietal placentae, the ovules many. 

FRANKENIA L. 

Stems round. Foliage glands rather superficial. Flowers cymose- 
paniculate. Possibly occurring is Anthobryum Phil., a densely matted 
ligneous "cushion" plant with quadrate stems, deeply sunk glands 
and solitary terminal flowers; two species approach southern Peru, 
A. aretioides Phil, of northern Chile, its stamens 5 and A. triandrum 
(Re~my) Surgis, Rev. Ge"n. Bot. 34: 455. 1922 by inference and ex 
Ndz. I.e. 281 (the combination overlooked by bibliographers), its 
stamens 3, Bolivian. 

Frankenia chilensis Presl in Roem. & Schult. Syst. 7:1618. 1830. 
F. campestris Schauer, Nov. Act. Acad. Leop. -Carol. 19: Suppl. 1: 
480. 1843. F. Nicoletiana Phil. var. aspera (Phil.) Reiche, Anal. 
Univ. Chile 90: 922. 1895. F. peruviana Schellenb. Bot. Jahrb. 50: 
Beibl. Ill: 10. 1913?; cf. note below. 

Shrubby, the nodose stems and thick revolute leaves beneath 
ashy-puberulent (Peruvian form); leaves ovate, blunt, roundish or 



FLORA OF PERU 5 

minutely cordate at base but generally so revolute that the form is 
concealed, characteristically about 5 mm. long, 2 mm. wide; sepals 
narrowly lanceolate, acute, to 4 mm. long or longer; style more or 
less trifid at apex. The Peruvian plant seems to be the pubescent 
form, var. aspera (Phil.) Johnst. Contrib. Gray Herb. 85: 77. 1929, 
whose interpretation of the species as very variable I follow; however, 
cf. Ndz. key, I.e. 281, in which, without placing F. peruviana he 
maintains F. aspera Phil, with style lobes 1-1.5 mm. long and 
F. campestris Schauer and F. chilensis Presl with style lobes shorter 
than 1 mm. The latter he separates on a minute difference in size 
of calyx but especially in the absence of a ligule in the flower of 
F. campestris. This is a point best worked out in the field. F. 
peruviana was described with calyx to 8.5 mm. long, the petal claw 
appendaged. Flowers white according to Weberbauer, pink or 
pinkish fide Johnston, who describes the plants as forming low 
shrubby growths on dry plains near the sea or on adjacent hillsides, 
this observation corresponding to Weberbauer, 148. F.M. Negs. 
35062; 35063. 

Arequipa: Islay, D'Orbigny. Mejia (Guniher &Buchtien 167, det. 
Bruns as F. Nicoletiana Phil. var. aspera (Phil.) Reiche). Mollendo, 
300 meters, Weberbauer 386 (type, F. peruviana}. 

FLACOURTIACEAE. Flacourtia Family 

Reference: Gilg in Pflanzenfam. ed. 2. 21: 377-457. 1925. 

Trees or shrubs with alternate stipulate leaves, the stipules often 
promptly caducous, the leaves often pellucid-dotted. Flowers dioe- 
cious or hermaphrodite, borne in axillary clusters, sometimes in pani- 
cles or otherwise disposed, the flower-parts 4-several, spirally or 
serially arranged. Stamens 8 to many, a disk usually more or less 
developed. Ovary generally superior, 1-celled or 2-several-celled by 
complete fusion of the 2-8 parietal placentae. Styles free or connate, 
sometimes simple. Fruit frequently capsular, sometimes more or 
less berry-like. Seeds often arillate, not infrequently pubescent. En- 
dosperm if present usually starchy or oily. 

Besides the above reference I acknowledge my indebtedness to 
the helpful work of Sleumer as published recently, particularly in 
Notizbl. Bot. Gart. Berlin as cited on occasion below. 

Baehni in Candollea 5: 405-412. 1934 was constrained to suggest 
placing Mollia, Nettoa and Trichospermum in this relationship, 
genera generally included in the Tiliaceae. The chief character on 
which he based his conclusion was drawn from the ovary, this being 



6 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pluricellular in Tiliaceae and 1-celled in Bixa, Mollia, Nettoa and 
certain species of Trichospermum. Since it seems to us that most 
of the morphological characters oppose such a classification, the 
family Tiliaceae may after all be a homogeneous ensemble; however, 
to conserve the genera named in this family it is necessary to modify 
its character to read ovary 1-several-celled, or even partially divided, 
as Burret does. 

The family (Tiliaceae) is variable then in this character just as 
is the Flacourtiaceae, so nearly related to it through the genera Bixa 
and Cochlospermum. But now one cannot more speak of the char- 
acter of the placentation as a general character of the Malvales. As 
soon as one diminishes the importance of the established separation 
between Parietales and Centrospermae one perceives these interest- 
ing analogies. Thus the situation of genera such as Mollia and 
Goethalsia (see Baehni, Candollea 6: 44-45. 1935 and Record, Trop. 
Woods 42: 21. 1935) in this arrangement becomes clearer. The 
single technical character that forced some botanists to place these 
genera in the Parietales having lost its importance, one can, because of 
general characters undeniably important, leave them in the Tiliaceae, 
reinterpreted. Baehni's research was thought-provoking, showing, 
once again, among other things, that taxonomy can only approx- 
imately valuate the nuances in plant relationships; that, above all, 
for taxonomy to remain practical it must perforce remain often 
"unsatisfactory," so to speak, either failing in one direction or in 
another; the situation is usually complicated, too, by the personal 
viewpoint of the author. Considering the disposition of Mollia in 
this light I am leaving it in Tiliaceae. 

Key (based on Gilg) 

Sepals and petals dissimilar or spiraled and similar or the petals 
more numerous than sepals, not appendaged. 

Styles 1 ; fruit wingless. 

Anthers obovoid; albumen starchy 2. Bixa. 

Anthers oblong or linear; albumen starchy or oily. 

Anthers laterally dehiscent; leaves simple 4. Lindackeria. 

Anthers opening by pores; leaves palmately lobed. 

3. Cochlospermum. 

Styles 3-7 5. Mayna. 

Sepals and petals similar, about the same number or the petals 
lacking. 



FLORA OF PERU 7 

Corona present; petals lacking; flowers yellow, in elongate ter- 
minal racemes; pubescence branched 6. Abatia. 

Corona lacking, sometimes also the petals. 
Petals present. 
Scandent shrub, each inflorescence with 1 sterile hooked 

peduncle 1. Ancistrothyrsus. 

Shrubs, but lacking modified peduncles. 

Stamens indefinite in number, not in bundles. 
Pubescence simple. 

Anthers long-linear; flowers large in spike-like racemes. 

7. Neosprucea. 
Anthers small, roundish; flowers panicled or not in 

spike-like racemes. 
Ovary at anthesis 3-5-celled ; flowers in short racemes ; 

stipules large 8. Prockia. 

Ovary 1-2-celled or falsely several celled by the 
intrusion of the placentae; stipules small. 

9. Banara. 

Pubescence stellate 10. Pineda. 

Stamens in bundles opposite the petals or solitary before 

each petal 11. Homalium. 

Petals none. 

Leaves not lucid-punctate; stamens hypogynous. 

Style extremely short, if obvious 12. Xylosma. 

Style elongate, divided 13. Ryania. 

Leaves usually not opaque, obviously lucid-punctate (usually) ; 

stamens more or less clearly perigynous. 
Sepals in bud connected, finally 2-5 separating . 14. Lunania. 
Sepals at least at tips free. 

Stamens same number as sepals 15. Tetrathylacium. 

Stamens more numerous than the sepals. 

Stamens 6-12 alternating with staminodial appendages. 

16. Casearia. 

Stamens usually 10 or more and without appendages. 

17. Laetia. 

Artificial key 

Scandent shrub, each inflorescence with 1 sterile hooked peduncle. 

1. Ancistrothyrsus. 



8 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Shrubs, but the peduncles if present never modified. 
Inflorescence notably branched, often paniculate. 

Inflorescence candelabriform 15. Tetrathylacium. 

Inflorescence not so formed. 

Flowers small, the parts similar 9. Banara. 

Flowers medium to large, the sepals and petals dissimilar. 

Anthers obovoid; flowers medium in size 2. Bixa. 

Anthers narrow; flowers showy 3. Cochlospermum. 

Inflorescence little if at all branched or in any case very narrow 

(the large-flowered Cochlospermum might be sought here). 
Flowers racemose or cymose, the inflorescence sometimes short, 

or the flowers solitary. 
Racemes notably elongating, the flowers numerous. 

Leaves opposite; racemes terminal, simple 6. Abatia. 

Leaves alternate; racemes often axillary, often with 1-2 

branches; leaves 3-nerved 14. Lunania. 

Racemes or cymes rarely 1 dm. long, the flowers few or rela- 
tively few, or panicled, clustered or solitary, but then 
5 mm. wide or wider. 

Leaves markedly 3-nerved from base (3-5-nerved). 
Anthers subglobose. 

Flowers in terminal panicles or axillary fascicles. 

9. Banara. 
Flowers in racemes or only 1 or 2 8. Prockia. 

Anthers linear; flowers large, in pseudospikes. 

7. Neosprucea. 

Leaves not distinctly 3-nerved (cf . sometimes Banara with 

branched inflorescence). 

Racemes often 1 dm. long; flowers medium size, normally 
10 or more; leaves glabrous 11. Homalium. 

Racemes or inflorescence rarely 8 cm. long, the flowers 
1-8; leaves usually puberulent. 

Leaves ashy pubescent with branched hairs. 

10. Pineda. 

Leaves not so pubescent but trichomes stellate in 
Ryania. 

Petals and sepals 3, persisting; stipules large. 

8. Prockia. 



FLORA OF PERU 9 

Petals none or at least twice as many as the sepals; 

stipules small or caducous. 
Petals none; sepals 4-5. 
Anthers linear; flowers few, rather large. 

13. Ryania. 
Anther ovoid or short-oblong, tiny; flowers 

small 17. Laetia. 

Petals 4-12; sepals 2-3. 
Style 1; fruit warty-echinate; leaves ample. 

4. Lindackeria. 
Styles 3-7; fruit prickly, soft appendaged or 

smooth 5. Mayna. 

Flowers in sessile or shortly pedicelled clusters, small or less 

than 5 mm. wide (rarely corymbose, Laetia). 
Leaves not lucid-punctate; no appendages between stamens, 

these usually indefinite 12. Xylosma. 

Leaves usually lucid-punctate. 
Flowers with staminodia; younger parts often pubescent, 

the leaves usually drying greenish 13. Casearia. 

Flowers without staminodia; Peruvian sp. mostly glabrous, 
the leaves net-veined, heavy, drying reddish-brown. 

17. Laetia. 

1. ANCISTROTHYRSUS Harms 

Liana with alternate leaves and axillary often elongate peduncles 
trifid at apex, the middle branch modified into a sterile hooked hold- 
fast. Flower pedicel ed, whitish. Receptacle obsolete. Sepals and 
petals 4, similar. Disk or corona extrastamineal, tubular, laciniate 
at margin. Stamens 8, lightly connate, the anthers small, narrowly 
oblong, versatile. Ovary shortly stipitate, 1-celled, with 4 placentae, 
the ovules 2-3 in each partition. Styles 4 with thick spreading 
stigmas. Member of the Paropsieae according to the author, which 
group connects the family with the Passifloraceae. It is interesting 
to note the narrow 1-celled ovary. 

Ancistro thyrsus Tessmannii Harms, Notizbl. Bot. Gart. 
Berlin 11: 147. 1931. 

Younger parts puberulent or lightly pubescent becoming glabrate 
or glabrous; petioles about 1 cm. long; leaves rather elliptic-obovate 
or oblong, entire, mostly 10-15 cm. long, 5-8 cm. broad; peduncles 
4-8 cm. long, the sterile one with hook 2.5-3 cm. long; fertile 



10 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

peduncles bracteolate, the puberulent pedicels articulate, 1.5-2.5 
cm. long; petals acuminate, 3-3.5 cm. long; corona pubescent, 8 mm. 
high; greenish filaments glabrous, with the dark orange anthers 
about 14 mm. long; young fruits densely hirsute, subglobose. A 
beautiful plant, the showy white flowers campanulate (but divided 
to base). According to the discoverer the liana climbs to 3 dm., 
the stem sometimes thicker than 5 cm. Illustrated, I.e. opposite 
page 149. 

Loreto: In flood-free wood, mouth of the Santiago, Tessmann 
4127; 4479. Mishuyacu near Iquitos, Klug 650. 

2. BIXAL. 

Reference: Pilger in Pflanzenfam. ed. 2. 21: 313-315. 1925; 
Ducke, Arquivos Serv. Florestal 1: 36-37. 1939. 

Tree-like shrubs or large trees with colored sap, broad, entire, 
alternate leaves and terminal panicles of rather large roseate or 
white flowers. Stipules promptly caducous. Sepals and petals 5, 
imbricate. Flowers bisexual. Stamens many, free or nearly so, the 
horseshoe-shaped anthers opening by apical slits. Ovary 1-celled 
with two parietal placentae. Fruit smooth to echinate, 2-valved, 
the obovoid seeds with an outer colored covering or pulp that 
dissolves in water. 

The common and variable species B. Orellana L. is well known, 
at least in cultivation, in tropical and subtropical regions for the 
colored pulp that surrounds the seeds and which furnishes a color 
for foods such as rice, soups, and beverages, in which connection it 
is interesting to note that it has been found to be a source of vitamins 
D and A. It is known in commerce as annatto, which finds industrial 
application as a color for butter and butter substitutes. It was 
formerly employed as a dye for silk and wool. Among various 
Indian tribes it is used as body paint, and in civilized communities 
as lipstick. 

The method of separation of the pulp from the seeds is by im- 
mersion in hot or cold water; after agitation the pulp settles and the 
water is gently poured off; the sediment, dried, is the product. This 
at one time was believed to have excellent diuretic properties; the 
leaves and the roots according to Pilger have been used as a digestant. 

Pods ovoid-conical, often pointed, longer than wide, densely echinate 
or smooth B. Orellana. 

Pods spherical or reniform or wider than long, pubescent and 
echinate or tuberculate. 



FLORA OF PERU 11 

Pods papillose or shortly and sparsely spinulose tuberculate, 

compressed B. platycarpa, B. arborea. 

Pods more or less densely echinate; little compressed, seeds red- 
brown. 

Pods eglandular, the spines fine, numerous or few . . B. Urucurana. 

Pods glandular and more or less echinate with fragile but 

coarse spines B. excelsa. 

Bixa excelsa Gleason & Krukoff, Phytologia 1: 107. 1934. 

A tree with the spherical reniform pods of B. Urucurana, to 4.5 
cm. wide, 3 cm. long, but reddish-brown glandular- tomentose and 
coarsely echinate at least below, the fragile but thick-based spines 
breaking off in age; seeds strongly flattened, the raphe costate. 
Attains, according to Ducke, 10-30 meters in clay terrain above 
inundation. Has been distributed from region of Tapajos, Brazil, 
as B. arborea and B. Orellana. 

Rio Acre: Near Rio Blanco (Ducke). Brazil. "Urucurana da 
matta." 

Bixa Orellana L. Sp. PI. 512. 1753. B. odorata R. & P. ex G. 
Don, Gen. Syst. 1: 295. 1831. 

Shrub-like or low spreading tree with broadly ovate, acuminate 
leaves truncate or more or less cordate at base, usually somewhat 
pitted beneath; inflorescence scurfy pubescent; pods characteristi- 
cally ovoid-pointed, longer than broad, densely long-echinate; seeds 
red (yellow), papillose, the endocarp detaching. The forma leiocarpa 
(Kuntze) Macbr., comb. nov. (Orellana americana var. leiocarpa 
Kuntze, Rev. Gen. 1: 45. 1891), has the pods completely smooth. 
Small tree commonly cultivated in the montana, but according to 
Ducke not persisting in Amazonia when an area is retaken by forest. 
According to Weberbauer, 252, found in Peru between 1,800-2,200 
meters in bush- wood; according to Ruiz and Pavon, cultivated in 
the whole of Peru but apparently not found wild. A common escape 
in departments of Huanuco and Cuzco below 1,000 meters (Stork, 
Horton & Vargas). Illustrated, Mart. Fl. Bras. 13, pt. 1. pi. 87. 

Lima: Callao, Gaudichaud 125. Huanuco: Region Monzon, 
Weberbauer, 286. Cuchero, Ruiz & Pavon. Junin: Huacapistana, 
Weberbauer 2334; 252. La Merced, 5275; 5274; 5342 (f. leiocarpa). 
San Martin: Tarapoto, Williams 5549; 7182; 7694 (f. leiocarpa). 
Vchiza, Ruiz & Pavon. Loreto: Pebas, Williams 1583. Iquitos, 
Tessmann 5069; Mexia 6435. Balsapuerto, Klug 3040. Near 
Iquitos, Williams 1462. Caballo-Cocha, Williams 2193. Cuzco: 



12 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Prov. Convention, Santa Ana, Calca, Valle de Lares, Paucartambo 
(fide Herrera). Valle del Urubamba, 2,200 meters, Herrera 3217. 
Prov. Convention, in abandoned cultivations, Stork, Horton & 
Vargas 10475. "Achiote," "achote," "sacha achiote," "achihuiti," 
"huantura," "urcu," "achite amarillo," "achiote Colorado," 
"shambu," "shambu shambu," "shambu huayo," "shambu quiro." 
Warm regions. 

Bixa platycarpa R. & P. ex G. Don, Gen. Syst. 1: 294. 1831. 

Similar to B. Orellana and B. Urucurana with which it has been 
united but its pods are strongly flattened and merely roughened 
with scattered spinulose-tuberculate scabrosities; seeds papillose. 
Attains, according to Ducke, 30 meters and grows in clay terrain 
that is above inundation. Another tree apparently similar and 
similar in habitat has been described as distinct :B. arborea Huber, 
Bol. Mus. Goeldi 6: 87. 1910, with regular reniform strongly com- 
pressed pods merely muricate in age, the seed blue with a narrow 
red zone, papillose only on upper end (Pilger). The Tessmann and 
Williams collections match the type perfectly. The natives use the 
seeds to dye various things and to color food as they do also with 
B. Orellana (Ruiz & Pavon). F.M. Neg. 13629. 

Huanuco: Pozuzo, Ruiz & Pavon, type. Loreto: Yurimaguas, 
Williams 1+615. Rio Yanayacu, Pampa del Sacramento (Huber 
1552}. Mouth of Rio Napo (Ducke distr. 21, 279). Yarina Cocha, 
Tessmann 5434 (det. Pilger). Brazil. "Shambu-huayo," "max- 
pachin," "achote de monte." 

Bixa Urucurana Willd. Enum. Hort. Berol. 565. 1809. B. 
Orellana L. var. Urucurana (Willd.) Kuntze ex Pilger, I.e. 315. 
B. sphaerocarpa Triana, Bull. Soc. Bot. Fr. 5: 369. 1858. 

Like B. Orellana but pods spherical or flattened-spherical and 
smaller, densely to sparsely spiny, the seeds red-brown. According 
to Ducke, habitant of marl terrain that may be flooded. Ducke, I.e. 
37, apparently with reason, insists that this is a "good species"; 
Pilger, Notizbl. Bot. Gart. Berlin 11: 145. 1931, thought there were 
two species but apparently changed his mind when preparing the 
account for Pflanzenfamilien. Tree, 8 meters, native and planted 
for pulp surrounding the seeds, serving as paint for Indians and 
for coloring foods (Mexia). All the following were distributed as 
B. Orellana. 

Junin: Wooded valley, La Merced, Killip & Smith 23518. 
Loreto : Rancho Indiana, Mexia 6435. Caballo-Cocha, Williams 2082. 



FLORA OF PERU 13 

Middle Ucayali, Tessmann 5472. Leticia, Williams 3063. Rio 
Acre: Krukoff 5394- "Urucu-rana," "achiote bianco." Brazil; 
Venezuela. 

3. COCHLOSPERMUM Kunth 
Amoreuxia Moc. & SessS ex DC. Prodr. 2: 638. 1825. 
Reference: Pilger in Pflanzenfam. ed. 2. 21: 316-320. 1925. 

Shrubs or trees with palmately lobed or divided leaves and few, 
loosely borne large yellow flowers. Stamens sometimes shorter on 
one side or rarely some shorter on both. Anthers narrow, opening 
at top or at top and bottom by pores or chinks. Style slender. 
Ovary 1-celled medially but with 3-5 lateral placentae or 3-celled. 
Seeds sometimes long-hairy. Maintained by Engler (cf. Pilger, 
I.e. 316) as a family distinct from Bixaceae by the narrow anthers 
and by the oily endosperm. The name has been written Cochlio- 
spermum and Cochleospermum; it was conserved against Maximil- 
leanea Mart. & Schrank to which nameKuntze transferred the species. 

Leaves palmately 5-7-foliolate; anthers 2-pored. 
Stamens on one side of flower shorter. 

Sepals tomentose both sides C. Williamsii. 

Sepals glabrescent C. potentilloides. 

Stamens all the same length C. orinocense. 

Leaves palmately parted; anthers 1-pored C. vitifolium. 

Cochlospermum orinocense (HBK.) Steud. Nomencl. ed. 2, 
pt. 1: 393. 1840. Bombax orinocense HBK. Nov. Gen. & Sp. 5: 301. 

1822. 

A large tree with scarred branchlets, 7-5 digitate suborbicular 
leaves and terminal panicles of showy flowers; leaflets lanceolate, 
attenuate both ends, even petioled at base where somewhat unequal, 
entire, membranous, sparsely pubescent only on the nerves beneath; 
petioles elongate; calyx olive-green, puberulent-tomentulose, the 
broad sepals about 8 mm. long; petals broadly obovate, deeply 
emarginate or even bilobed, 4-4.5 cm. long; anthers about 3 mm. 
long, opening near tip by 2 short slits or pores; fruit not surely 
known. At least according to Pilger's interpretation (probably cor- 
rect) the following material is referable here; surely it is not C. 
Parkeri Planchon in Hook. Lond. Journ. Bot. 6: 310. 1847 under 
which name Klug 3120 was distributed; that British Guiana spe- 
cies has extremely obtuse leaflets and secund flowers. 



14 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Tarapoto, Williams 6664. Loreto: Yurimaguas, 
Poeppig 2473 (det. Szyszys.); Tessmann 3768 and Killip & Smith 
28176 (det. Pilger). Mission de Sarayacu, Castelnau. Between 
Balsapuerto and Moyobamba on Rio Cachiyacu, Klug 3120. Upper 
Itaya, Williams 3485. Lower Huallaga, Williams 3991. Near 
Iquitos, Klug 1468. Venezuela. "Huiiia caspi," "quillo-sisa," 
"huimba." 

Cochlospermum potentilloides (Pilger), comb. nov. Amo- 
reuxia potentilloides Pilger, Notizbl. Bot. Gart. Berlin 13: 255. 1936. 

Stems slender with a few short branchlets, 1.5 dm. long, shortly 
papillose toward the tips; leaves few (petioles 6-8 cm. long) orbicu- 
late-reniform, 3-3.5 cm. long, deeply 7-parted, the middle divisions 
2-2.5 cm. long, all obovate-cuneate, rounded at tip, serrate-dentate 
only above; stipules subulate; flowers few, borne laxly; sepals gla- 
brescent, the younger sparsely pubescent, narrowly ovate, 15 mm. 
long; petals to 3.5 cm. long; longer stamens to 15 mm. long, the 2- 
pored anthers 3.5-4 mm. long. Perhaps related to A. colombiana 
Sprague, Kew Bull. 104. 1922, with pubescent calyx, the seeds with 
a wide shallow sinus, the arrilode short-pubescent. A. unipora v. 
Tiegh. Journ. Bot. 14: 48. 1900, Bolivian and to be expected, has 
3-lobed leaves and 1-pored anthers; interesting to note C. tetrasporum 
Hall, f., also Bolivian, with 4-pored anthers, 2 pores at each end. 

Amazonas: Jacuanga and Bagua (Raimondi 631, type). 

Cochlospermum vitifolium (Willd.) Spreng. Syst. 2: 596. 1825. 
Bombax vitifolium Willd. Enum. Hort. Berol. 720. 1809. 

Smooth shrub or tree with long-petioled deeply 5-lobed leaves 
that are cordate at base, their lobes repandly serrulate or serrate, 
oblong-lanceolate, acuminate, nearly glabrous; flowers to 1 dm. 
broad; sepals puberulent both sides, about 15 mm. long; fruit 7-8 
cm. long, softly puberulent the many reniform seeds covered with 
a cottony down. Commonly planted in the West Indies; flowers 
when only 1 meter high (Standley). A common tree of the xerophytic 
forest of Cerro Viento about 35 miles east of Talara (Haught). 

Piura: Negritos, Haught 203. North to Mexico. 

Cochlospermum Williamsii Macbr. Candollea 5: 388. 1934. 

Tree; petioles 1.5 cm. long and longer, glabrous and striate-angled ; 
leaflets 6, entire, strongly unequal, the smaller about 6 cm. long, 4 cm. 
wide, the larger to 20 cm. long and nearly half as broad, all elliptic, 
cuneate to base, shortly acuminate, chartaceous-membranaceous, 



FLORA OF PERU 15 

glabrous and slightly lustrous above, opaque and shortly, sparsely 
pilose on nerves beneath, the 16 lateral nerves rather marked there; 
panicles densely reddish-tomentose with widely divaricate few- 
flowered branchlets; flowers 5 cm. long, expanded 7-8 cm.; pedicels 
1 cm. long, fruiting nearly 2.5 cm. long; sepals brown-tomentose 
both sides, elliptic, the outer two a little shorter, about 17 mm. 
long; petals obovate, deeply bilobed, 5 cm. long, 3.5 cm. wide; 
stamens unequal, those on one side about 2 cm. long those on the 
other scarcely half as long, the anthers 3 mm. long, more or less 
curved and apically 2-pored; style 12 mm. long, villous at base; 
capsule young, truncate-depressed, deeply silky brown-tomentose. 
Possibly it is a Bombax or of that affinity. However I allied it to 
C. Wentii Pulle of Surinam, a species with the sepals glabrous within. 
Described by the collector as a magnificent tree. 

Loreto: Pebas on the Amazon, Williams 1964, type; also 1778. 
Caballo-Cocha, Williams 2090. 

4. LINDACKERIA Presl 

Unarmed shrubs or trees with several- to many-flowered axillary 
racemes of medium-sized polygamous flowers. Sepals 3, the petals 
6-12. Stamens sometimes in a column, the anthers narrow. Ovary 
1-celled, warty. Could well be treated as a section of the genus 
Mayna, the chief difference being in the way in which the flowers 
are borne. But Mayna in turn is essentially Oncoba Forsk. except 
that the latter is often thorny and the petioles little if at all thick- 
ened below the leaf blade as in this genus and in Mayna; the matter 
deserves study. 

Lindackeria maynensis Poepp. & Endl. Nov. Gen. & Sp. 3: 
63. pi. 270. 1845. Oncoba maynensis (Poepp. & Endl.) Eichler in 
Mart. Fl. Bras. 13, pt. 1: 441. 1871. 

Smooth tree or shrub with ovate- or nearly oblong-elliptic more 
or less caudately acuminate leaves and medium-sized white or yel- 
lowish polygamo-monoecious flowers borne in short axillary racemes 
on the loosely spreading branchlets; younger nodes often glutinous; 
leaves usually a dm. or two long and less than half as wide, entire, 
opaque; racemes several- to 25-flowered, the lower male flowers short- 
the upper female long-pedicellate; flowers to 12 mm. wide; sepals 
oblong; filaments free, equaled by the narrow anthers; fruit globose, 
thickly warty tuberculate-echinate, brownish-yellow; seeds 2, rarely 
3, lustrous. L. latifolia Bth. andL. pauciflora Bth., both Amazonian, 
have respectively lanceolate and suborbicular sepals, the filaments 



16 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

of the former united; the latter moreover has 9 petals and the racemes 
are only 2-4-flowered. Rarely to 12 meters high. F.M. Neg. 13617. 

Loreto: Iquitos, Tessmann 5385; Williams 8005; 3753; Killip 
& Smith 27316; 27083. Mishuyacu, King 765; 977; 804; 437. 
Yarina Cocha, Tessmann 5414; 5396; 5422. Lower Rio Nanay, 
Williams 425. Balsapuerto, Klug 3037 (det. Standl.). Yurimaguas, 
King 2816; Poeppig; Williams 4681; 4705. San Martin: Pongo de 
Cainarachi, Klug 2661 (det. Standl.). Brazil. "Caracana," "hua- 
capu," "huicho caspi," "lluicho caspi," "quinillia Colorado." 

5. MAYNAAublet 

Shrubs or small trees with entire or toothed leaves, deciduous 
stipules and fragrant flowers, the male in short axillary fascicles, 
the female solitary or few. Sepals 3, the longer petals 6-9. Stamens 
free, pubescent, the anthers linear. Ovary 1-celled, stiffly hairy, the 
styles 3. Fruit berrylike, dry, prickly. The petioles are thickened 
below the leaf blade. At least for the purposes of this work, here 
may be included Carpotroche Endl. which is in general similar but 
sepals 2-3, petals 4-12, styles 4-8 and fruit a large capsule or rather 
small, smooth or with 8-16 wing-like appendages borne from the base 
to the persistent short styles or the fruit merely lacerate-appendaged. 
In some species the flowers are polygamo-dioecious. One Brazilian 
species has smooth fruits. The appendages of the fruit if present are 
various; cf. illustrations, An. da Reun. S.-Amer. Bot. 3: 93-96. 1938. 

Group is of special interest because at least one species, notably 
C. brasiliensis Endl., has been found to be a good source of Chaul- 
moogra oil. 

Leaves glabrous or merely puberulent beneath. 
Leaves ample, cuneate-obovate. 

Petioles 1-2.5 cm. long; fruit aculeate, 15 mm. thick. 

M. echinata. 

Petioles 4-6 cm. long; fruits large the wing-like appendages 
interspersed with lacerate ones M. longifolia. 

Leaves medium size; fruits to 2.5 cm. thick, only with soft tri- 
angular rarely cleft appendages M. parvifolia. 

Leaves densely soft-pilose beneath M. amazonica. 

Mayna amazonica (Mart.) Macbr., comb. nov. Carpotroche 
amazonica Mart, ex Eichler in Mart. Fl. Bras. 13, pt. 1: 437. 1871. 
C. mollis Macbr. Candollea 5: 390. 1934 (?). 



FLORA OF PERU 17 

Small tree more or less densely and softly pilose even to the 
pedicels except the leaves above, these glabrous and somewhat 
lustrous between the nerves that are rather prominent beneath; 
petioles to about 2 cm. long; leaves chartaceous, remotely and 
minutely apiculate-dentate, elliptic to obovate, the base obtuse, the 
apex broadly acuminate, to 3 dm. long, 12 cm. wide; pedicels about 
7 mm. long; male flowers few in short axillary racemes, 1-2 
cm. broad, the white segments silky pilose without, 6 mm. long, or 
typically longer, the petals and sepals subequal. The female and 
polygamous flowers are solitary or few in the axils. Fruits smooth. 

Loreto: Mishuyacu near Iquitos, Klug 1120 (type, C. moms'). 
Brazil; Colombia. 

Mayna echinata Spruce, in herb. Dendrostylis echinata (Spruce) 
Benth. Journ. Linn. Soc. Bot. 5: App. 2: 82. 1861. 

Shrub with lightly hirsutulous branchlets (toward the tip) 
petioles and pedicels, also somewhat puberulent; leaves oblong- or 
obscurely obovate-elliptic, 1-2 or even 3 dm. long, 4-9 cm. wide, 
cuneate at base, more or less abruptly caudate at tip, typically well- 
undulate, membranous or in age chartaceous, the veins becoming 
rather prominent beneath but fine; petioles 10-25 mm. long, slender; 
stipules subulate, 4-5 mm. long; petals about 7 mm. long (female 
flowers), the male smaller; fruit pubescent, about 15 mm. thick, 
sparsely to densely aculeate, the triangular-based aculei about 3 
mm. long. As remarked by Eichler, I.e. 444 and by Gilg, I.e. 404, 
this is possibly better treated as a variety of M. odorata Aubl., 
which, however, appears, at least typically, to have much thicker 
leaves with coarse venation, a denser almost tomentulose puberu- 
lence on the branchlets, and stiffer narrower prickles on the glabrate 
fruits; there may of course be intermediate forms. F.M. Neg. 24100. 

San Martin: Tarapoto, Spruce 4499, type; Williams 6552; 6557; 
6531; 5822. Junin: Rio Perene", Killip & Smith 25105. Puerto 
Bermudez, Killip & Smith 6677. Loreto: Balsapuerto, Killip & 
Smith 28590. Mishuyacu near Iquitos, Klug 866; Mexia 6398 (det. 
A. C. Sm.). Puerto Arturo, Williams 5021; 5206; 5328. Rio Itaya, 
Williams 235. Pongo de Manseriche, Tessmann 4583. Pongo de 
Cainarachi, Klug 2758 (det. Standl.). Rio Acre: Krukoff 5693. 
Amazonian region. "Shamshu huayo," "congo caspi," "sapote 
yacu." 

Mayna longifolia Poepp. & Endl. Nov. Gen. & Sp. 3: 64. pi. 

271. 1845. 



18 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Carpotroche longifolia (Poepp. & Endl.) Benth. Journ. Linn. Soc. 
Bot. 5: Suppl. 2: 82. 1861. 

Shrub or tree with shortly rusty- tomentulose branchlets; petioles 
4-6 cm. long, markedly articulated below the leaf -blade, this 3-4 or 
even 6 dm. long, 1-2 dm. wide, obovate-oblong, long-attenuate at 
base, shortly caudate acuminate, remotely sinuate, somewhat 
pubescent on both sides but soon glabrate above, chartaceous; stipules 
6-10 mm. long; flowers dioecious, the male 10-12, the female 15-16 
mm. wide (Eichler), both sorts fascicled on the trunks; sepals 2; 
petals 6-7 (9-10); capsules woody, to 6 cm. long, 4 cm. thick, the 
soft wings with many soft lacerated crests between them; seeds 
irregularly shaped, obtusely angled, about 6 cm. thick, 10 mm. long. 
The flowers, according to Poeppig, are odorless. The leaves 
simulate those of Clavija sp. The white fruit, turning green, is 
borne on stubs from the main trunk (Mexia). The "pulp" of the 
seeds is edible, the name "Huira guayo" meaning "lard fruit" 
(Klug). C. amazonica Mart, and C. grandiflora Spruce have the 
male flowers in short racemes; the petals and sepals of the former 
are subequal, the flowers less than 2.5 cm. broad; in the latter the 
petals are longer and the flowers attain a diameter of 4 cm. The 
fruit of the former is a dehiscent gray-tomentose capsule to 6.5 cm. 
long, 5.8 cm. wide, with 5 stout and 5 very weak protuberances; 
that of the latter is a capsule that opens apically star-like and that 
was described by Eichler as being the fruit of the former (Ducke). 
There are also C. integrifolia Kuhlm. the leaves really entire, the 
flowers small, and C. crispidentata Ducke, Archiv. Inst. Biol. Veg. 
Rio Jan. 4: 55. 1938, with the fruit of M. longifolia but green, long- 
peduncled and densely armed with small soft crisped appendages; 
the male form of this simulates C. brasiliensis. Another Amazonian 
species that may occur is C. apterocarpa Kuhlm. Archiv. Inst. Biol. 
Veg. Rio Jan. 2: 87. pi. 5. 1935, the fruits wingless; east to Rio 
Tapajoz on non-inundated lands (Ducke). 

San Martin: Tarapoto, Spruce 4255. Junin: 400 meters, Puerto 
Jessup, Killip & Smith 26256; and Puerto Bermudez, 26491; 26675 
Loreto: Mouth of Rio Santiago, Mexia 6131 (det. A.C.Sm.). Caballo- 
Cocha, Williams 2131. La Victoria, Williams 2711. Soledad, Tess- 
mann. Pebas, Wittiams 1801. Lower Rio Nanay, Williams 625. 
Balsapuerto, Killip & Smith 28551. Iquitos, Williams 8017. Mishu- 
yacu, Klug 1124; 259; 12; 97; 1121; 4; Killip & Smith 29864. 
Cerro de Conchohuayu (Huber 1379). Brazil. "Huira huara," 
"huira guayo," "zapoto del mono." 



FLORA OF PERU 19 

Mayna parvifolia (Macbr.) Sleumer, Repert. Sp. Nov. 45: 
12. 1938. Carpotroche parvifolia Macbr. Candollea 5: 390. 1934. 

Completely glabrous shrub or perhaps tree; branchlets slender, 
pale, flexuous; petioles 5-10 mm. long, grooved above; leaves elliptic 
or oblong-elliptic, shortly acute at base, abruptly acuminate, 10-12 
cm. long, 4-4.5 cm. wide, often smaller, membranous and rather 
densely lineate-punctate-lucid, pale green and slightly lustrous both 
sides; lateral nerves 4-5, moderately prominent with the reticulate 
veins beneath; fruiting pedicels thick, nearly 1 cm. long; capsule 
globose to 2.5 cm. thick, densely covered with triangular-subulate 
puberulent soft crests 5 mm. long. Otherwise unknown. 

Loreto: Puerto Arturo, Yurimaguas, Williams 5260. "Casha 
huayo." 

6. ABATIA Ruiz & Pavon 

Myriotriche Turcz. Bull. Soc. Nat. Mosc. 36: 554. 1863. 
Slender shrubs with opposite estipular leaves and long racemes 
of medium-sized yellow flowers. Petals none, the short-tubular 
calyx with 4-5 teeth, the base persisting about the globose dehiscing 
capsule. Stamens many, the slender filaments with filiform append- 
ages, the anthers short. Ovary 1-celled, or partly 2-celled, with 1 
slender style. Seeds minute, dorsally winged or keeled. Named 
for Don Pedro Abat, professor of botany at Seville. 

Calyx subsessile, even in fruit, 1-2.5 mm. long A. boliviana. 

Calyx well-pedicellate, somewhat to much longer. 
Pedicels (in flower) and sepals 2-5 (7) mm. long with gray-fulvous 
trichomes. 

Pubescence stellate- tomentulose A. parviflora. 

Pubescence pilose A. canescens. 

Pedicels and sepals soon 7-10 mm. long or longer, the pubescence 

rather brown. 

Pedicels geniculate well below the middle, to 1 cm. long in fruit. 

A. rugosa. 

Pedicels geniculate slightly below the middle, 2 cm. long in 
fruit A. spicata. 

Abatia boliviana (Mandon & Wedd.) Britton, Bull. Torrey 
Club 17: 214. 1890. Graniera boliviana Mandon & Wedd. ex Benth. 
& Hook. f. Gen. PI. 1: 799. 1867; name. 

Straggling or loosely slender-branched shrub; petioles 2-5 mm. 
long; leaves ovate-lanceolate, minutely cordate at base, acuminate, 



20 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

finely denticulate, ashy scurfy-stellate beneath but soon glabrate, 
especially above; racemes with peduncle only about 7 cm. long, often 
shorter, densely flowered at least above, spike-like; rachis densely 
stellate- tomentose; calyx lobes puberulent without, triangular-acute; 
fertile stamens few; capsule globose, stellate puberulent, the seeds 
puncticulate. F.M. Neg. 24096. 

Puno: 2,700 meters, Cuyocuyo, Sandia, Weberbauer 876; 237. 
Bolivia. 

Abatia canescens Sleumer, Notizbl. Bot. Gart. Berlin 11: 951. 
1934. 

About 2 meters high, lightly ashy pilose except the style and the 
leaves above where scarcely so, beneath more densely, especially on 
the nerves; leaves opposite, oblong-obovate or oblong, narrowed to 
the 1 cm. long petiole, often minutely acuminate, denticulate, mem- 
branous, 7-9 cm. long, 3-4 cm. wide; racemes terminal, 16-18 cm. 
long with often some lateral ones from the axils of the younger 
leaves; bracts acuminate, 3-5 mm. long, about as long as the pedicels; 
sepals 4, ovate-lanceolate, 3-4 mm. long; stamens about 35; ovary 
densely pilose, attenuate to the glabrous style. 

Lambayeque: 1,500 meters, Olmos (Weberbauer 7099, type). 

Abatia parviflora R. & P. Syst. 1: 136. 1798. A. verbascifolia 
HBK. Nov. Gen. & Sp. 5: 358. pi. 486. 1823? 

Gray stellate-tomentulose, especially the young striate-angled 
branchlets and the younger leaves beneath these more or less densely 
soft-pilose above with 2-3-parted trichomes, oblong-elliptic, acute 
at base, very shortly acuminate, the 10-12 lateral nerves with the 
veins reticulate and prominent only beneath, in age sparsely stellate- 
pubescent beneath and subcoriaceous, to about 10 cm. long, 5 cm. 
wide; petioles to 1.5 cm. long; racemes to 2 dm. long; pedicels 3 mm. 
long or the lower even 6-9 mm. long, these remote; sepals glabrous 
within, ovate, acute, 2-5 mm. long; stamens about 20; ovary globose, 
the style glabrous. The Colombian plant of HBK. may be distinct 
from the Peruvian, at least varietally, which here is described from 
the typical form from Muna. F.M. Neg. 34907. 

Huanuco: Muna, Ruiz & Pawn, type; 4300; 2075. Above Mito, 
flowers bright or lemon yellow, 1594 (det. Sleumer). Pampayacu, 
Kanehira 74 (det. Sleumer). Yanano, 4926. Ecuador. "Taucca- 
taucca." 



FLORA OF PERU 21 

Abatia rugosa R. & P. Syst. 1: 136. 1798. 

Branches gray-brown, nearly smooth; branchlets compressed at 
the nodes, compactly stellate- tomentose; petioles 1-1.5 cm. long; 
leaves densely floccose stellate-tomentose beneath, sparsely so or 
soon glabrate above, thick-chartaceous, closely denticulate, oblong- 
elliptic, unequally and slightly cordate, minutely acuminate, the 
older 12 cm. long, 4 cm. wide or larger, the upper smaller; nerves 
and veins conspicuously impressed-reticulate above, beneath con- 
cealed by the pubescence; racemes floriferous nearly to base, about 
2 dm. long; pedicels and sepals yellow stellate-pubescent, the former 
8-10 mm. long, geniculate 2-3 mm. above the base, the latter 
narrowly ovate, acuminate, about 6 mm. long; glabrous within; 
stamens about 25 or 30; capsule included in the calyx, globose with 
acute tip, the seeds wing-keeled. F.M. Neg. 24097. 

Huanuco: Rondos, Pillao and Nauyen, Ruiz & Pavon, type. 
Above Huanuco (near Pillao), 3,000 meters, 2075. Chinchapalca, 
1594. Ghaglla, Weberbauer 6694- Below Cerro de Pasco, Sawada 
P92. "Yoriturpi," "galgaretama," "tauhac tauhac," "retamo 
cimmarona," "taucca-taucca." 

Abatia spicata (Turcz.) Sleumer, Repert. Sp. Nov. 45: 13. 1938. 
Abatia macrostachya Killip, Journ. Wash. Acad. Sci. 24: 50. 1934. 
Myriotriche spicata Turcz. Bull. Soc. Nat. Mosc. 36: 555. 1863. 

Densely stellate-tomentose throughout except the sepals within 
and the style, the ovate, auriculate-based leaves sparsely so above; 
racemes lax, the slender pedicels divaricate, 1.5-2 cm. long, genicu- 
late near the middle; sepals narrowly lanceolate, 7-8 mm. long, 
attenuate to the subulate tip; fertile stamens about 35. The racemes 
may attain 2.5 dm. but it is not clear to me that the plant is distinct 
from A. rugosa because there appear to be intermediate collections; 
for example, Weberbauer 4986, also from Cuzco region, has pedicels 
even of old lower flowers barely 10 mm. long yet the geniculation is 
about at the middle. On the other hand it is true that the pedicels 
of typical A. rugosa from central Peru are geniculate well below the 
middle. Under these circumstances more material will be necessary 
to decide the matter. The type is by Mathews from Andinamarca 
(not seen). 

Cuzco: Yanamanchi to Amaibamba (CooA; & Gilbert 1136, type, 
A. macrostachya}; Valle de San Miguel, Herrera 2015 (det. Sleumer). 
Urubamba Valley, Herrera 1568 (det. Killip). Convention, Weber- 
bauer 4986 (det. Sleumer). Apurimac: 2,900 meters, Weberbauer 
5838. Huancavelica: Forest edge, 2,900 meters, Ampurco, Stork & 



22 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Horton 104.19. Without locality, misit Mathews, Gaudichaud 1204. 
"Escobilla." 

7. NEOSPRUCEA Sleumer 

Spruceanthus Sleumer, Notizbl. Bot. Gart. Berlin 13: 362. 1936, 
not Verdoorn, 1934. 

Shrub or small tree with elliptic-oblong 3-nerved leaves and 
spike-like racemes of rather long flowers. Leaves 3-nerved from the 
biglandular base. Stipules minute, caducous. Sepals and petals 4, 
persisting at base of the dry fruit. Anthers long-linear, the short 
filaments filiform. Ovary 1-celled or semi-5-6-celled by the prominent 
placentae. Genus name a highly just compliment to the great 
English botanist Spruce. 

Neosprucea grandiflora (Spruce) Sleumer, I.e. 14: 47. 1938. 
Banara grandiflora Spruce ex Benth. Journ. Linn. Soc. Bot. 5: 
Suppl. 2: 93. 1861. Hasseltia grandiflora (Spruce) Sleumer, I.e. 11: 
960. 1933. Spruceanthus grandiflorus Sleumer, I.e. 13: 363. 1936. 

Petioles about 1 cm. long; leaves broadly cuneate or subrounded 
at base, acuminate, chartaceous, 13-25 cm. long, 5-6 (-9) cm. wide, 
remotely and obtusely toothed or subentire; flowers white; rachis, 
pedicels (2 mm. long), sepals and petals tomentose, the latter 8-10 
mm. long, 3.5 mm. wide; sepals ovate, 4-6 mm. wide, 6-7 mm. 
long; anthers 8-10 mm. long, the filaments 2 mm. long. F.M. 
Negs. 24085; 34873. 

San Martin: In rocks along streams, Tarapoto, Spruce 4897, 
type. Loreto: Pampas de Ponasa, 1,200 meters, Vie 6712. Colom- 
bia; Brazil. 

8. PROCKIA P. Br. 

Shrubs or small trees with pinnately nerved toothed leaves, 
mostly persistent conspicuous stipules and few-flowered usually 
terminal racemes of rather large blossoms. Sepals and petals 
similar, 3, rarely 4, persisting. Stamens many, free, the anthers 
small. Ovary completely 3-5-celled, the style simple. Capsules not 
dehiscing, the small seeds angled and in a white pulp. The leaf- 
teeth are gland-tipped as are the leaves at the base. This was 
included at one time as Hasseltia in the Tiliaceae because of the 
multicellular ovary; cf. remarks above at beginning of family. 

Prockia Crucis L. Sp. PI. ed. 2. 745. 1762. 
Grayish-villous, especially the young leaves and branchlets; 
leaves rather unequally cordate-ovate, characteristically thin but 



FLORA OF PERU 23 

varying considerably in size and coarseness of serration, 3-5 (more 
or less distinctly) -plinerved, acuminate; corymbs few (-12, Eichler) 
-flowered; pedicels about 1 cm. long, usually ebracteolate; sepals 
ovate-cordate, about 6 mm. long; capsule somewhat depressed- 
globose, hirsutulous. The upper stipules, arcuate-lanceolate, den- 
tate, are a prominent feature of this shrub or small tree. Our 
common form is the var. septemnervia (Spreng.) Sleumer, Notizbl. 
Bot. Gart. Berlin 14: 47. 1938, the leaves rather obviously 7-nerved, 
the pedicels sometimes bracteolate. Illustrated, Moric. PI. Nouv. 
Amer. pi. 97; Hook. Icon. pi. 94. 

San Martin: Tarapoto, Williams 5818. Juanjui, Klug 3829; 
3865. Amazonas: Chachapoyas, Mathews. Huanuco: Huallaga, 
1,600 meters, Weberbauer 6817; Mexia 8314 (det. Standl.). Pozuzo, 
Ruiz & Pavon (det. Sleumer as the sp.). Junin: La Merced, 1,000 
meters, Weberbauer 1940; Isern 2123; Schunke 336. Loreto: Yuri- 
maguas, Williams 4727; 5048; 5220; 4285; Killip & Smith 28070. 
Santa Rosa, Killip & Smith 28806. Rio Acre: Ule 9364 (the sp.). 
Brazil to Argentina. "Charapilla." 

9. BANARAAublet 

Hasseltia HBK. Nov. Gen. & Sp. 7: 231. pi. 651. 1825. Hassel- 
tiopsis Sleumer, Notizbl. Bot. Gart. Berlin 14: 46, 49. 1938. 

Trees or tree-like shrubs frequently with gland-toothed pinnately 
nerved leaves, or these more or less distinctly 3-5-nerved, minute 
or small caducous stipules and usually many rather small flowers 
borne in terminal or axillary cymes, panicles or compound umbels. 
Sepals and petals similar, 3, or rather less frequently 4-6, in the 
former case always persisting, in the latter in some species falling 
from the base of the young fruit. Stamens many, free, the anthers 
minute. Ovary 1-celled at anthesis, the placentae more or less 
intruded or rarely 2-celled by their complete union. Capsule inde- 
hiscent or tardily and irregularly dehiscent, often berry-like, some- 
times hard and apiculate by the persistent style, the seeds 1-several. 

The genus Hasseltiopsis recently has been proposed by Sleumer 
to care for certain species with the ovary-placentae (3-5) little 
intruded, the flower parts falling from the young fruits, the leaves 
3-nerved from base. Unfortunately he has selected as type of his 
group Banara dioica Benth. Journ. Linn. Soc. 5: Suppl. 2: 94. 1861, 
which, as is evident from examination of cotype material of the two 
collections cited by Bentham, is a Banara; at least its sepals are 
persistent about the mature capsules. Therefore it seems that a 



24 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

new name must be given to include the species actually conforming 
to Sleumer's diagnosis. As I had long questioned the distinctness 
of Hasseltia HBK. which Sleumer's exacting research proves con- 
clusively is only a Banara with ovary with 2 placentae that ulti- 
mately are connate (but I would record that I had retained the 
genus even as Hasseltiopsis at first, to avoid transferring names) I 
now prefer to follow my belief, and the more simple course, letting 
Banara include these units rather than give a name to a segregate 
which seems evidently to be artificial. Equivalent differences are 
to be found within generic lines in many families, as in Lobeliaceae 
and in Carica, to mention only two. And in the related Melasto- 
maceae the number of ovary cells may vary even in the same species. 
Certainly the characters defining the groups may be said to be vari- 
able or to represent merely degrees in a common development. How- 
ever, compare Sleumer, I.e. 46, for a careful and illuminating analysis 
of them upon which I have drawn. I follow his most useful work in 
leaving Prockia and Pineda conveniently marked entities and Neo- 
sprucea Sleumer, at least as far as this group is concerned, as 
outstanding. 

Leaves 3-nerved from base. 

Sepals lanceolate, promptly reflexing and with petals persisting; 
leaves usually subentire, acute at base B. laxiflora. 

Sepals ovate, spreading or tardily reflexing; leaves more or less 
dentate and rounded at base. 

Sepals 2-2.5 mm. long; lower panicle branches to 2 cm. long. 

B. glabrata. 

Sepals 3-4 mm. long; lower panicle branches elongate. 

B. leucothyrsa. 

Leaves pinnately nerved even from the base (Peruvian species). 
Leaves subsessile, typically soft-pilose beneath even at maturity, 

closely serrate B. mollis. 

Leaves markedly petioled at least if pubescent, usually glabrous 
or glabrate beneath, subentire or somewhat toothed. 

Petioles rarely 5 mm. long, mostly shorter B. guianensis. 

Petioles (6) 8-12 mm. long. 
Leaves ovate-elliptic-oblong, fully twice as long as broad. 

Leaves opaque, regularly toothed, or subentire. 

B. amazonica. 

Leaves lustrous, the teeth unequally remote B. nitida. 



FLORA OF PERU 25 

Leaves ovate-oval, scarcely twice longer than broad. 

B. Tessmannii. 

Banara amazonica Sleumer, Notizbl. Bot. Gart. Berlin 12: 
50. 1934. 

Bark yellowish-brown the lenticels small; petioles minutely 
pubescent or glabrate, 10-12 mm. long, the binate glands at the 
base of the leaf, this subrotund or narrowed at the base, broadly 
acuminate, chartaceous, glabrous, opaque both sides, 8.5-11 cm. 
long, 3.5-5 cm. wide, the regularly placed teeth obtuse, scarcely 0.5 
mm. high, nearly glabrous or the prominent midnerve obscurely 
pilose beneath; panicles axillary and terminal, many-flowered, 8-13 
cm. long, the lower branches to 8 cm. long; pedicels glabrous, the 
bractlets puberulent; sepals ovate, acute, puberulent without, 4-4.5 
mm. long, subequaling the yellow rotund-ovate petals; ovary gla- 
brous, the fruit lustrous. Tree 40 meters high or higher (Mexia); 
medium-sized tree with trunk 25 cm. in diameter (Tessmann). F.M. 
Neg. 28938. 

Loreto: Yarina Cocho, 155 meters, Tessmann 3219, type. Rancho 
Indiana, left bank Maranon, Mexia 6404 (distr. as B. laxiflora). 
Bolivia. "Teareo." 

Banara glabrata Sleumer, Notizbl. Bot. Gart. Berlin 11: 
952. 1934. 

Branchlets glabrous, the lenticels round; petioles pubescent, 8-10 
mm. long; leaves rounded or narrowed at base, acuminate, subcoria- 
ceous, subentire or most minutely and remotely denticulate, 10-12 
cm. long, 4.5-6 cm. wide; prominent 3 (-5) nerves beneath a little 
pubescent toward base of leaf, the binate glands obvious; panicles 
axillary and terminal, many-flowered, yellowish-tomentulose, 4-7 
cm. long, the lower branchlet 1.5-2 cm. long, the pedicels about 1 
mm. long; sepals ovate, acuminate, 2 mm. long; petals green, rounded, 
tomentulose, slightly longer; ovary glabrous. Tree in flood-free 
wood. F.M. Neg. 28941. 

Loreto: Puerto Melendez below the Pongo de Manseriche (Tess- 
mann 3916, type). Rio Acre: Krukoff 5770. 

Banara guianensis Aublet, PI. Guian. 1: 548. pi 217. 1775. 

Low shrub or tree with oblong, regularly glandular-serrate leaves 
and gray-puberulent panicles of small yellow flowers; petioles short 
with 1 or 2 cupulate glands at the tip or at base of the leaf, this 
rounded or subcordate, the tip shortly acuminate; leaves somewhat 
pilose on the nerves beneath, chartaceous, usually about 8-12 cm. 



26 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, 4-6 cm. wide; pedicels 3-5 mm. long; ovary glabrous, the 5-8 
placentae lamelliform; fruit to 8 mm. in diameter, pulpy within, 
the black seeds sulcate-ribbed. Sometimes 15-30 meters (Tess- 
mann); a 7 meter tree or taller used for firewood and posts (Mexia). 

There are often one or two glands placed irregularly near the 
junction of the petiole and leaf -blade; sometimes the glands are 
regularly opposite each other at the tip of the petiole and this vari- 
ation seems to be associated with somewhat smaller leaves so that 
designation seems to be called for: var. isadena Standley, var. 
nov. in herb, foliis plerumque oblongo-ellipticis, 6-8 cm. longis, circa 
3 cm. latis, petiolo apice regulariter biglanduloso. Florida, Klug 
2246, type; 2358. 

Huanuco: Region Churubamba, 1,100 meters, Mexia 8159 (det. 
Standl.). Junin: Puerto Yessup, Killip & Smith 26303; 26436 (det. 
Standl.). San Martin: Juanjui, 400 meters, Klug 3908. Loreto: 
Lower Rio Huallaga, Williams 4978. Mouth of Santiago, Tessmann 
4508; 3976. Fundo Indiana, Mexia 6387 (toward the var.). Yuri- 
maguas, Williams 5321; 5365; 4014- Costa Rica to Brazil and 
Guiana. "Raya caspi," "machu-mangua," "linque," "oco cireyda," 
' 'machinmangua. ' ' 

Banara laxiflora Benth. Journ. Linn. Soc. Bot. 5: Suppl. 2: 
91. 1861. Hasseltia laxiflora (Benth.) Eichler in Mart. Fl. Bras. 13, 
pt. 1: 502. 1871. H. peruviana Pilger, Verh. Bot. Ver. Brandenb. 
47: 160. 1905. 

A tree or shrub with slender branchlets, typically glabrous but 
usually the younger parts and the leaves, especially on the nerves 
beneath, somewhat pubescent; petioles 7-10 mm. long; leaves ellip- 
tic-oblong, lightly sinuate-dentate or subentire, acute at base, 
acuminate, usually 1-2 dm. long, 4-7 cm. wide, with 2 glands at the 
base, lustrous, chartaceous-coriaceous; panicle lax, the branches 1-4 
times divaricately divided, often about 1 dm. long and even as wide, 
sometimes much smaller; sepals barely 2 mm. long, lanceolate, 
slightly tomentulose, exceeded by the stamens, these about 30, 
equaled by the petals; ovary glabrous, slightly attenuate at base; 
placentae 2-3, completely connate below the middle (Benth.). H. 
peruviana seems to be the typical form. H. floribunda HBK., found 
as near as Ecuador, has coarsely dentate leaves, the inflorescence 
densely flowered. Illustrated, Mart. Fl. Bras. 12, pt. 3. pi. 34. 
F.M. Negs. 13649; 13650. 

San Martin: Pongo de Cainarachi, Ule 6416, type H. peruviana. 
Frequent on the banks of the Maranon and Pastasa, Spruce 4964, 



FLORA OF PERU 27 

type. Junin: Vitoc, Ruiz & Pavon (det. SI. & Mansf.). Loreto: 
Puerto Arturo, Killip & Smith 27894. Soledad, Tessmann 5226. 
Mouth of Santiago, Tessmann 4306; 3420. Yurimaguas, Williams 
4976; 4556; 4611; 5196; 5028; 4292; 4293. Mishuyacu, King 1462. 
La Victoria, Williams 2945; 2937; 2779. Lower Rio Nanay, Williams 
479; 588; 632. Rio Monzon, Williams 8176. Lower Rio Huallaga, 
Killip & Smith 29264? (10 meter tree, leaves 3.5 cm. wide). Balsa- 
puerto, King 2992; 3234 (det. Standl.). Rio Acre: Krukoff 5770; 
5395. "Tamamaru," "tama-nara," "okuchi-uasi," "okuchi huasi," 
"ratan-caspi." 

Banara leucothyrsa (Sleumer), comb. nov. Hasseltiopsis leu- 
cothyrsa Sleumer, Notizbl. Bot. Gart. Berlin 14: 51. 1938. 

Tree, glabrous except for the petioles, gray-tomentulose sepals, 
petals and ovary; petioles 1-2.5 cm. long; leaves broadly oblong- 
elliptic to elliptic, more or less rounded at base, where biglandular, 
and caudate at tip, 7-14 cm. long, 4-7 cm. wide, subcoriaceous, lus- 
trous both sides but paler beneath, remotely crenately glandular-ser- 
rate; basal nerves 3; panicles 9-12 cm. long, the lower branches to 
8.5 cm. long, the rachis and pedicels rather stout, rigid; sepals and 
petals 4-6, 3.5-4 mm. long, ovate-oblong, subacuminate, falling 
from the fruit, slightly longer than the stamens; ovules many; seeds 
unknown. The author apparently rightly has distinguished this tree 
from B. dioica Benth. (Hasseltia dioica Sleumer, I.e. 12: 55. 1934); it 
is rather similar to B. mexicana Gray, with narrower leaves cuneate 
at base and longer stamens. He further separates a Colombian form 
(Hasseltiopsis albomicans Sleumer, I.e. 51) with nearly twice as large 
panicles. 

Loreto : In flood-free wood below Puerto Melendez, Tessmann 4802, 
type. Pongo de Manseriche, Tessmann 4835. Mouth of Santiago, 
Tessmann 3976; 4472. 

Banara mollis (Poepp. & Endl.) Tul. Ann. Sci. Nat. ser. 3. 7: 
290. 1847. Kuhlia mollis Poepp. & Endl. Nov. Gen. & Sp. 3: 74. pi. 
285. 1845. B. Tulasnei Macbr. Candollea 5: 389. 1934, probably. B. 
guianensis Aublet, var. mollis (Poepp. & Endl.) Eichler in Mart. 
Fl. Bras. 13, pt. 1: 501. 1871. 

Shrub or small tree with many spreading branches, the lax rusty- 
pilose branchlets terminated by often long and narrow panicles of 
small flowers; leaves subsessile, usually oblong, with a rounded base, 
this bicupulate-glandular, and a short acumination, regularly and 
prominently glandular-serrulate, to 2 dm. long or longer and 5 cm. 



28 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or so wide, or the upper smaller, glabrate above but usually at least 
on the pinnate nerves softly sericeous-pilose beneath; sepals ovate, 
acutish, subglabrous within, 2 mm. long, the rounded petals a little 
shorter; ovary glabrous; fruit ellipsoid, apiculate, pulpy within. 
Perhaps better treated as a variety of B. guianensis, following Eichler. 
Eichler designated Spruce 4894 as a small-flowered form but this 
number as seen by me is only in bud. F.M. Negs. 18229; 24088; 
24089. 

San Martin: San Roque, Williams 7031. Tarapoto, Spruce 4894; 
Vie 6589 (det. Pilger). Pongo de Cainarache, King 2680 (det. 
Standl.). Huanuco: Cochero, Poeppig, type; Dombey; Ruiz & 
Pavdn. Huallaga, Weberbauer 6818. Chinchao, Ruiz & Pavdn. 
Junin: La Merced, 526 1; 5717. Chanchamayo, Weberbauer 1832. 
Pichis Trail, Kittip & Smith 25718 (var.? petioles longer). Near 
Moyobamba, Weberbauer 4518; Klug 3301 (det. Standl.). Cerro de 
Canchahuayo (Huber 1467). Cuzco: Gay. Brazil; Colombia? 
"Borracho sisa," "galgaretama." 

Banara nitida Spruce ex Benth. Journ. Linn. Soc. Bot. 5: 
Suppl. 2: 93. 1861. 

Glabrate or in age glabrous; petioles 6-10 mm. long, minutely 
strigillose, grooved above, often with one cupulate gland on the 
junction to the leaf-blade, this ovate-oblong, inequilateral at the 
narrowed base, well-acuminated, coarsely but remotely crenate, 
chartaceous-coriaceous, lustrous above, to 2 dm. long, 7 cm. wide, 
mostly smaller; sepals ovate, acute, about 3 mm. long, the broader 
petals a little longer; panicle puberulent in age, about 12 cm. long, 
the lowest branches 6 cm. long, widely divaricate, the fruiting pedi- 
cels to 7 mm. long; capsules ellipsoid, indehiscent, apiculate, the few 
seeds faintly lineate. B. regia Sandw. Kew Bull. 154. 1930 (B. 
magnifolia Sleumer, Notizbl. Bot. Gart. Berlin 11: 952. 1934), from 
Ecuador, has leaves 3-5 dm. long, the teeth even 2.5 mm. high. 
F.M. Neg. 24090. 

San Martin: Tarapoto, Spruce 4512, type; Williams 6530; 6986. 
Loreto : Pro on the Amazon, Williams 1 965a. 

Banara Tessmannii Sleumer, Notizbl. Bot. Gart. Berlin 14: 
48. 1938. 

Tree-like shrub with smooth sparsely lenticellate branches; 
petioles 10-12 mm. long, ashy-pilose; leaves ovate or rarely oblong- 
ovate, rounded at base, sometimes more or less inequilaterally, 
broadly attenuate at tip but obtuse, chartaceous, glabrous except the 



FLORA OF PERU 29 

puberulent pilose nerves, regularly sinuate, glandular-denticulate, 
9-12 cm. long, 5-7 cm. wide; panicles terminal, lax, about 15- 
flowered, and 6 cm. long, gray-pubescent; pedicels slender, about 
1 cm. long; sepals broadly deltoid, obtuse, 3.5 mm. long, the pale 
green petals slightly smaller. In flooded area. 

Loreto : Upper Ucayali, edge of Lake Mosote, Tessmann 3359, type. 

10. PINEDA R. & P. 

Christannia Presl, Rel. Haenk. 2: 91. pi 67. 1835. 

Shrub with alternate entire blunt oblongish leaves, these ashy- 
green by reason of the felty covering of branched trichomes. Flowers 
rather large, long-pediceled, the persisting sepals and petals usually 
5. Stamens many, the anthers small. Ovary 1-celled, the placentae 
thick. Style short, thick, crowning the dehiscing 2-7-seeded capsule. 

Pineda incana R. & P. Prodr. 76. pi. 14. 1794. Christannia 
salicifolia Presl, Rel. Haenk. 2: 91. pi. 67. 1835. Banara incana 
(R. & P.) Benth. Journ. Linn. Soc. Bot. 5: Suppl. 2: 94. 1861, and 
var. Jamesonii Benth. I.e. P. Lehmannii Hieron. Bot. Jahrb. 20: 
Beibl. 49: 58. 1895, fide Sleumer. 

Low shrub often many-stemmed, with short petioled oblongish 
subentire to distinctly toothed more or less white stellate-pubescent 
leaves or these beneath in age only arenose between the veins, 
usually about 2.5 (-3.5) cm. long; flowers few in short corymbiform 
racemes, the pedicels bracteolate, 6-12 mm. long, yellow fading 
reddish; sepals oblong-lanceolate, the similar petals somewhat longer, 
the stamens included; ovary glabrous. The leaves supply a black 
dye (Herrera). The exceptionally strong wood is used for walking 
sticks, stools, and baskets and in construction of hanging bridges 
(R. & P.). 

Lima: San Rafael, 2282; Sawada P110; Mathews891. San Mar- 
cos, prov. Huari, Raimondi. Huanuco: (Haenke). Monzon, 2,700 
meters, Weberbauer 3730. Huanuco, Ruiz & Pavdn, type. Ambo, 
3147. Junin: Huacapistana (Weberbauer, 247). Cuzco: Gay. 
Paucartambo and Quispicanchi ( Herrera}. Urubamba (Weberbauer, 
175); Soukup. Ecuador. "Lloque," "llogui," "lloqque." 

11. HOMALIUM Jacq. 

Reference: Blake, Contr. U. S. Nat. Herb. 20: 221-235. 1919. 

Trees or shrubs with pinnately nerved usually toothed but 

eglandular leaves and mostly axillary racemes, rarely panicles, of 



30 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

short-pedicelled hermaphrodite flowers. Calyx tube turbinate, the 
usually 5-7 parts persisting in fruit as also the similar petals. Sta- 
mens opposite the petals, solitary or fascicled and bearing cushion- 
like processes at base, just between the petals. Anthers globose. 
Ovary more or less united to calyx, 1-celled, the styles 2-6. Capsule 
somewhat leathery, the seeds without aril. 

Homalium pedicellatum Spruce ex Benth. Journ. Linn. Soc. 
Bot. 4: 36. 1860; 231. 

Branches rather rough, the young branchlets, petioles and leaf- 
nerves beneath sparsely pilose or glabrate; petioles 3-8 mm. long; 
leaves oblong-ovate or obovate-elliptic, mostly rounded at the 
acutish base, shortly and bluntly attenuate at apex, coarsely crenate- 
dentate, fugaciously barbellate in the nerve-axils beneath, charta- 
ceous, lustrous above, reticulate-veiny on both sides, mostly 8 cm. 
long, 4-5 cm. wide; racemes terminal on axillary branchlets, ashy- 
pubescent, lax, simple or apparently sometimes few-branched; lower 
pedicels 5-7 mm. long; flowers 5-6-merous, more or less ashy- 
puberulent, the narrowly lanceolate sepals to 3.5 mm. long, the 
elliptic- or oblong-ovate petals 3.5-4.5 mm. long, nearly equaled 
by the glabrous stamens, these mostly in threes; appendages puberu- 
lent; ovary loosely or sparsely villous; styles typically united below 
or the ovary tip extended as the style-column. Resembles closely H. 
guianense (Aubl.) Warb. with petals 2.5-3.4 mm. long and is very simi- 
lar to H. eleutherostylum Blake, I.e. 232, the leaves more pubescent, 
the styles distinct to the base; it, in turn, is near H. racemosa Jacq., 
with glabrous branchlets. Unfortunately Tessmann 5203 seems 
intermediate in characters to these forms or is itself another species; 
it is nearly glabrous, has branching racemes, styles not united. As 
there is other material difficult to place, it may be that the species 
have been drawn on too fine lines. An Amazonian species better 
marked is H. densiflorum Spruce, the flowers subsessile, 2-2.4 mm. 
long, crowded in panicles with suberect branches. Illustrated, 
Mart. Fl. Bras. 13, pt. 1. pi. 101. 

Loreto : 17 meters high, Soledad, Tessmann 5208. Flowers yellow, 
Iquitos, Klug 1297. Brazil; Venezuela. 

12. XYLOSMA G. Forst. f. 

Myroxylon J. & G. Forst. Char. Gen. 125. 1776. Hisingera 
Hellen. in Vet. Akad. Handl. Stockh. 32. pi. 2. 1792. Hiesingera Endl. 
Gen. Suppl. 5: 47. 1850. 



FLORA OF PERU 31 

Frequently thorny and more or less pubescent shrubs or small 
trees with shortly petioled usually toothed estipulate leaves and 
small flowers borne in bracteolate axillary racemes, often reduced 
to appear as axillary clusters or glomerules. Flowers usually dioe- 
cious, the sepals 4-5 (6-7), somewhat united at base, the petals none. 
Stamens many from a ring or glandular disk, the long filaments 
free. Ovary lacking in male flowers, free, 1-celled. Style nearly 
obsolete to well-developed, the stigma sometimes lunate and more 
or less lobed. Fruit a berry with few seeds. 

The following key is not satisfactory and I doubt the validity of 
some species including those proposed by me; X. minutiflorum 
Macbr., Candollea 5: 392. 1934, omitted, is evidently, from fruiting 
material of the same collection in Herb. Delessert, Geneva, a species 
of Phyllanthus, probably P. guianensis Aublet or affine. The type 
specimen is only in flower, but of course the resemblance even so is 
only superficial. 

Leaves pilose beneath, at least on the nerves. 

Petioles 3-5 mm. long; leaves to about 8 cm. long X. pilosum. 

Petioles about 10 mm. long; leaves 12-18 cm. long. . X. Ruizianum. 
Leaves glabrous or essentially so. 

Leaves cordate-ovate X. cordatum. 

Leaves distinctly acute at base. 
Leaves ample; male disk-lobes many; style lunately several- 

lobed X. Tessmannii. 

Leaves small to medium, rarely 10 cm. long or longer. 

Leaves mostly 3.5-6 cm. wide, rather coarsely toothed or 

crenate-dentate; petioles 2-5 mm. long. 
Male disk 6-10 gland-lobed; style shortly bifid, the 

stigma lunate X. Salzmannii. 

Male disk crenate; stigmas 2, subulate X. digynum. 

Leaves 2-3 (-4) cm. wide, minutely crenate-dentate; petioles 
1-2 (3) mm. long X. Benthami. 

Xylosma Benthami (Tul.) Eichler in Mart. Fl. Bras. 13, pt. 1: 
447. 1871. Flacourtia Benthami Tul. in Ann. Sci. Nat. ser. 3. 7: 291. 
1847. F. nitida Benth. in Hook. Journ. Bot. 3: 119. 1851? Xylosma 
armatum Macbr. Candollea 5: 391. 1934? 

Branches short, glabrous, sometimes smooth, sometimes armed 
with spines 1-3 cm. long; petioles scarcely 1 mm. long; leaves 
crowded, elliptic, shortly cuneate at base, obtuse or acutish, mostly 



32 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

about 4.5 (-7) cm. long, 2 (-3) cm. wide, coriaceous, not at all lucid- 
punctate, minutely denticulate, slightly lustrous above, faintly 
reticulate-veined; fascicles many, the pulverulent pedicels 2-2.5 
mm. long; sepals 4-7, broadly ovate, 1 mm. long; stamens strongly 
unequal, to 3 mm. long, usually only about 12, apparently sometimes 
as many as 30, glabrous; male disk 8-10 crenate-lobed ; style 
short, bifid at apex. The type of X. armatum is male; the 
leaves vary in degree of glandular-serration. But it seems to me 
doubtful if the Peruvian shrub is distinct. 

San Martin: San Roque, Williams 7237 (type, X. armatum). 
Loreto: Fortaleza near Yurimaguas, King 2807 (det. Standl.). Juan- 
jui, Klug 3835 (det. Standl. as X. armatum). 

Xylosma cordatum (HBK.) Gilg, Pflanzenfam. ed. 2. 21: 434. 
1925. Flacourtia cordata HBK. Nov. Gen. & Sp. 7: 241. 1825. 
Berberis rotunda Macbride, Candollea 6: 3. 1934. 

Branchlets unarmed or apparently sometimes armed, puberulent 
as the short (1-2 mm,) petioles; leaves cordate-ovate, acute, the 
larger 4 cm. long, coriaceous, epunctate, lustrous above, reticulate- 
veined, especially beneath, with minute incurving callus-teeth, the 
callus thus lying on the leaf -edge; bracts ciliate, equaling the pedi- 
cels, these 3-4 mm. long; male flowers 4-6 in the upper axils, 
the female solitary in the lower; former glabrous, the 5 sepals 
ovate-orbicular, equaled by the 35 glabrous stamens, the anthers 
oblong, the filaments free but accompanied by 25 (30) free hypog- 
ynous glabrous glands; fruit globose, red, to 6 mm. thick. Clos, 
Ann. Sci. Nat. se>. 4.8: 254. 1857, omits this shrub from the genus and 
probably from the family but Gilg by inference transfers it and 
associates it with X. digynum. My synonym probably belongs 
here; cf. note in this work under Berberis. Dr. Weberbauer has 
kindly supplied the data for the collection, data which never lacks 
and is in commendable completeness for all his collections. My 
phrase "Without locality" refers in this work only to the particular 
specimen seen. 

Piura: Ayavaca, Bonpland, type. Cajamarca: Above Namas, 
3,200 meters, Weberbauer 7218, male; 7220, female (type, B. 
rotunda). 

Xylosma digynum [Benth.] Eichler in Mart. Fl. Bras. 13, pt. 
1: 447. 1871. Flacourtia digyna Benth. in herb. 

Often a medium-sized tree and typically with abundant stout 
spines, these even to several cm. long; petioles 3-5 mm. long, canicu- 



FLORA OF PERU 33 

late above; leaves elliptic or elliptic-oblong, shortly attenuate 
at base, acuminate, 6-12 cm. long, 3-5 cm. wide, rarely somewhat 
larger, rather densely serrate-dentate, firm-membranous, usually 
lustrous above but opaque beneath ; fascicles congested at the nodes, 
few-flowered, sessile or subsessile, the peduncle thick; pedicels about 
3 mm. long (that of the female flowers shorter and thicker), articulate 
a little above the base, obsoletely puberulent as the ciliolate ovate 
1-1.5 mm. long bractlets; male flowers 3 mm. long, pubescent as 
the bractlets, the 4 broad sepals 1 mm. long; disk crenate, glabrous; 
stamens 15-20, twice as long as the calyx; female flowers 1.5 mm. 
long; stigmas 2, subulate, spreading-erect; capsule 5-6 mm. thick, 
with 3-4 trigonous lustrous seeds about 4 mm. long. According to 
Martius a common name, "auui-uva," means lignum acus; fide 
Mexia, "christo-casha," thorn of Christ. F.M. Negs. 6510; 13655. 
Loreto: Yurimaguas, Poeppig 2480; Williams 4752. Lower Rio 
Huallaga, Killip & Smith 29010. Florida, King 2276 (distr. as 
Casearia sp.). San Martin: Zepelacio, Klug 3285 (det. Standl.). 
Tarapoto, Ule 6757 (distr. as X. Salzmannii). Chazuta, Klug 4067 
(det. Standl.). Brazil. "Christo-casha," "umuruico" (Huitoto). 

Xylosma pilosum Macbr. Candollea 5: 391. 1934. 

Shrub with slender branchlets that are densely soft-pubescent 
with short trichomes; petioles 3-5 mm. long, shortly and densely 
pilose as the leaf-nerves beneath; leaves ovate-elliptic, broadly 
cuneate at base, subabruptly acuminate, to 8 cm. long, 3.5 cm. wide, 
chartaceous-membranous, epunctate, glabrous and somewhat lus- 
trous above, only the midnerve slightly pilose, deeply and obtusely 
serrate; pedicels lightly pilose, 1.5 mm. long; sepals 4, acute; stamens 
about 10, nearly 3 mm. long, glabrous, the disk rather prominent. 

Loreto: Puerto Arturo, Yurimaguas, Williams 4977, type. 
"Diablo-casha," "supay caspi." 

Xylosma Ruizianutn Sleumer, Notizbl. Bot. Gart. Berlin 12: 
476. 1935. 

Young branchlets densely tomentose, with yellowish hairs 0.5 
mm. long, as also the 1 cm. long petioles; leaves ovate-oblong, sub- 
rotund at base, gradually or shortly acuminate, the acumen 1-2 cm. 
long, obtuse, 12-18 cm. long, 6-10 cm. wide, rigid chartaceous, 
shortly pubescent above on the nerves, yellowish velutinous-tomen- 
tose beneath especially on the 7-8 lateral nerves, the veins laxly 
reticulate; margin coarsely subcrenate-dentate, the obtuse teeth 
1.5-2 mm. high, and about 5 mm. remote; axillary spines short, 6 



34 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

mm. long; fascicles dense, axillary; pedicels 1-2 mm. long; sepals 
4-5, ovate, obtuse, tomentose, about 2 mm. long, the indefinite 
glabrous stamens as long. Allied by the author to X. rubicundum 
(Karsten) Gilg with still larger leaves. 

Peru (?) : Without locality, Ruiz (Herb. Dahlem) ; Ruiz & Pavon 
(Herb. Madrid). 

Xylosma Salzmanni (Clos) Eichler, I.e. 448. Hisingera Salz- 
manni Clos, Ann. Sci. Nat. ser. 4: 8: 224. 1857. 

Nearly glabrous, the trunk and branches stoutly armed, the 
simple spines often lacking on the young branchlets; leaves mostly 
ovate-oblong and ovate, obtuse or obtusely produced, shortly atten- 
uate at base, more or less crenate-dentate, membranous, somewhat 
lustrous on both sides or opaque beneath, 4-10 cm. long, 2-5 cm. 
wide, often with 2-4 glands at the base of the blade; petioles 5-10 
mm. long; flowers umbellate-fasciculate, the 4-5 sepals ciliate, about 
1 mm. long, acute, the pedicels about 6 mm. long or much shorter; 
disk of male flowers 6-10 gland-lobed, the stamens 14-25, much longer 
than calyx; disk of female flowers reduced to a ring; style shortly 
bifid, the stigma semilunate. Stigma entire or more or less lobed 
and leaves variable according to Eichler, from whom this description, 
as others, is compiled. F.M. Negs. 13658; 24080. 

Huanuco: Pozuzo, 4551. San Martin: San Roque, Williams 
7976. Tarapoto, Ule 6757. Juanjui, King 4165 (det. Standl.). 
Loreto: Yurimaguas, Williams 4487? Mouth of Santiago, Tessmann 
3574? Rio Mazan, Schunke 174; 83. Brazil to Uruguay. "Costa- 
dosache," "cunchucra," "cunshi-cashan." 

Xylosma Tessmannii Sleumer, Notizbl. Bot. Gart. Berlin 12: 
477. 1935. 

Tree with numerous spines, these even to 10 cm. long; branchlets 
terete, somewhat pubescent; petioles about 1 cm. long; leaves ellip- 
tic-oblong, broadly cuneate at base, attenuate to the apex, this itself 
abruptly attenuate-acuminate and 1.5-2.5 cm. long, papery, gla- 
brous or nearly, 14-22 cm. long, 6.5-10 cm. wide; margin coarsely 
and regularly subcrenate-dentate, the teeth obtuse, mostly 5 mm. 
remote and 2 (-4) mm. high; lateral nerves 6-7, with the reticulate 
veins little prominent; fascicle many-flowered, the slender puber- 
ulent pedicels 5-6 mm. long; sepals ovate-acuminate, 4-5, puberulent 
and ciliate, yellowish-green, scarcely 1.5 mm. long; disk lobes many, 
scalelike, glabrous; stamens indefinite, 3 mm. long; style bifid, the 



FLORA OF PERU 35 

stigmas flabellately lobed. Known to attain 12 meters, branching 
at 4 meters. 

San Martin: Pongo de Cainarachi, Klug 2632. San Roque, 
Williams 7463. Loreto: Lower Rio Nanay, Williams 476. Flood- 
free wood, Iquitos, Tessmann 5153, type. Balsapuerto, Klug 3026 
(distr. as Casearia macrophylla). Brazil. "Supai kasha," "umuriuco." 

13. RYANIA Vahl 

Patrisia L. C. Rich. Act. Soc. Hist. Nat. Paris 1: 110. 1792. 

More or less stellately pubescent shrubs or trees with entire 
epunctate leaves, short petioles, caducous stipules and rather large 
hermaphrodite flowers borne solitary or few in the leaf-axils on 
geniculate pedicels that are scale-bracteate at base. Calyx 5-parted 
nearly to base, the 2 inner of the lanceolate divisions smaller. Petals 
none. Stamens many in 1-2 rows, the anthers linear. Ovary 1- 
celled with 2-6 placentae and the style as many-parted at tip. 
Fruit a berry-like capsule, often spongy-appendaged. There is a 
cupulate ring or disk between the ovary and the stamens. The 
name Ryania has been conserved; nevertheless that of Richard has 
been employed in Pflanzenfamilien, ed. 2. 

Ryania pyrifera (L. C. Rich.) Uittien & Sleumer in Pulle, Fl. 
Surinam 3: 286. 1935. Patrisia pyrifera L. C. Rich. Act. Soc. Hist. 
Nat. Paris 1: 111. 1792. R. speciosa Vahl, Eclog. Am. 1: 51. 1796. 

Slender branched tree, the younger branchlets, petioles, pedicels 
and sepals rusty- tomentose; stipules narrow, 4-8 mm. long, equaling 
the petioles; leaves nearly oblong or rarely a little obovate, obtuse 
and rounded or narrowed at the slightly unequal base, more or less 
abruptly acuminate, firm, rusty-tomentose at least on the mid-rib 
above, mostly 10-15 cm. long, 4-6 cm. wide; flowers 1 or 2 in the 
upper leaf-axils, subsessile, the lanceolate, acuminate, spreading 
sepals 2.5-3 cm. long but later erect and enclosing the large, densely 
corky-rugose, stellate capsule; stamens about as long as the sepals, 
the linear anthers 4-6 mm. long, the filaments filiform. The variety 
tomentosa (Miq.) Sleumer has the leaves densely rusty-tomentose 
beneath. Leaves rather variable in size and shape. Often only 
4-7 meters high. F.M. Negs. 34867; 34868 (var.). 

San Martin: Pongo de Cainarachi, Klug 2635. Loreto: Pampa 
del Sacramento, Castelnau. Rio Itaya, Williams 3443. Iquitos and 
Mishuyacu, Klug 7; 162; 795; 953; Tessmann 5361; Williams 8229; 
Killip & Smith 29900; 27113. Mouth of Santiago, Tessmann 4096. 
To Trinidad and the Guianas. 



36 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

14. LUNANIA Hook. 

A tree with entire lucid-punctate 3-nerved estipulate leaves 
and almost minute flowers borne on greatly elongate usually axillary 
racemes that are sometimes sparsely branched. Calyx short-tubular 
with 2-5 lobes. Petals none. Stamens 6-12, frequently with as 
many scales around a cupulate disk, the filaments mostly very short. 
Ovary free, 1-celled, pointed, with 3 broad placentae and 3 short 
styles or these nearly obsolete. Capsule coriaceous, globose, dehis- 
cent, the seeds small, punctate. 

Lunania parviflora Spruce ex Benth. Journ. Linn. Soc. Bot. 
5: Suppl. 2: 90. 1861. L. cuspidata Warb. in Pflanzenfam. 3, 6a: 
47. 1893. 

Slender, glabrous or nearly glabrous tree; petioles 1-1.5 cm. long; 
leaves chartaceous, lustrous above, finely net-veined, 3 (-5) -nerved 
from the rounded base, rather abruptly caudate-acuminate, gen- 
erally 12-15 cm. long, 5-6.6 cm. wide; racemes often 2 dm. long, 
the puberulent rachis notably slender, the small flowers shortly 
pedicellate or subsessile; sepals reflexing; stamens nearly always 10, 
subsessile, the short cupulate disk minutely and obtusely ciliolate- 
lobed; style short, conical, 3-parted on the young capsule. Warburg 
distinguished his species, based on Poeppig 2163, by its 8 stamens, 
disk lobes lacking, pubescent branchlets and 3-nerved leaves; I have 
not been able to see these differences, or they are not concomitant. 
Both the Poeppig and Spruce collections, however, seem to be smaller- 
flowered than most of the other material collected since and certainly 
this, although mostly determined as L. cuspidata, has a typically 
developed disk. I do not know the Poeppig locality. F.M. Negs. 
21317; 13639. 

Huanuco: Pozuzo, Weberbauer 6752. San Martin: Tarapoto, 
Spruce 3909, type. Pongo de Cainarachi, Klug 2640; 2687 (det. as 
L. cuspidata). Junin: Pichis Trail, Killip & Smith 26217. Loreto: 
Yurimaguas, Ule 6295; 6294; Williams 5059 (pedicels long). Pebas, 
Williams 1895. Balsapuerto, Klug 2901 (det. L. cuspidata). Cacho- 
puerto, Klug 3135 (det. L. cuspidata). Mishuyacu near Iquitos, 
Klug 158; 622 (abnormal or var.?). Without locality, Poeppig 
2163 (type, L. cuspidata). Rio Acre: Ule 9630; Krukoff 5227; 5275. 
Brazil. "Pina-quiro," "charapa-huatana." 

15. TETRATHYLACIUM Poepp. & Endl. 
Edmonstonia Seem. Bot. Voy. Herald 98. pi. 18. 1853. 



FLORA OF PERU 37 

A shrub-tree with ample, opaque, remotely serrate or subentire 
leaves and small, connately bracteolate flowers borne in crowded 
spikes panicled from the trunks and branches. Sepals none or minute. 
Petals 4, connate into a globose tube. Stamens 4, the anthers cordate. 
Ovary with 4 placentae, the style obsolete. Capsule coriaceous, 
tardily dehiscent. 

Tetrathylacium macrophyllum Poepp. & Endl. Nov. Gen. 
& Sp. 3: 34. pi. 240. 1845. Edmonstonia pacifica Seem. I.e. 

Small tree; petioles robust, 5-12 mm. long; leaves oblong, deeply 
cordate to truncate at base, shortly acuminate, opaque, subcoria- 
ceous, sometimes slightly pubescent beneath, even to 3.5 dm. long, 
16 cm. wide, many smaller; inflorescence broom-like, the rachis 
flexuose, sometimes 14 cm. long, 10 cm. across at top; flowers sessile, 
crowded, yellowish-tomentulose within as the style, the cordate 
anthers conspicuously exserted; bractlets persisting. The only 
species and unmistakable. 

San Martin: Pongo de Cainarachi, King 2702 (det. Standl.). 
Caballo-Cocha, Williams 2130 (det. Harms). Loreto: Yurimaguas, 
Ule 6293; Poeppig, type; Spruce 3881; Williams 4010; 3922. Pongo 
de Manseriche, Tessmann 3892. "Llaja." 

16. CASEARIA Jacq. 

Reference: Eichler in Mart. Fl. Bras. 13, pt. 1: 457-488. 1871. 

Tree or tree-like shrubs with entire or toothed usually lucid- 
punctate and lucid-lineate leaves, small deciduous stipules and small 
hermaphrodite flowers fascicled or capitate in the leaf -axils. Branch- 
lets rarely thorny. Pedicels very short with scaly bracts, sometimes 
two united bracteoles. Calyx tube short or long with 4-6 teeth, 
persistent. Petals none. Stamens 6-12-25 in a single row with as 
many staminodia, alternating free or attached to each other or with 
the filaments sometimes forming a stamineal corona. Filaments 
slender, anthers small. Ovary free, 1-celled with 2-3 placentae. 
Style simple or trifid, often short. Capsule dehiscent with 1-several 
often hairy seeds. I acknowledge with appreciation my indebtedness 
after Eichler to Sleumer's research, notably in Notizbl. Bot. Gart. 
Berlin 11: 953-960. 1934 and 12: 52-54. 1934. Besides the following 
there are two collections in herbaria. Williams 2738 from La 
Victoria once given an herbarium name by me is obscure; Klug 3343 
was distributed as C. sylvestris but the precocious flowers are without 
disk processes and have only 4 stamens; it suggests Celastraceae. 
C. Mathewsii Turcz. Bull. Soc. Nat. Mosc. 36, pt. 1: 608. 1863 based 



38 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

on Mathews 1649 without locality from Peru has not been identified 
from the description, which gives no essential character. Williams 
records for an unknown species the native name "quillo bordon." 

Branchlets exuding resin at the leaf -nodes; lateral nerves distinctly 

perpendicular-ascending near the margins C. resinfera. 

Branchlets not resiniferous; lateral nerves horizontal-ascending. 
Stamens 8, more or less connate with the disk lobes; stigma 
capitate; flowers cylindrical; stipules caducous. . .C. aculeata. 
Stamens 10-25, the disk lobes free or rarely a little connate with 
the filaments, or the former united into an intrastamineal 
crown. 

Leaves nearly always lucid-punctate or (and) lineolate at least 
obscurely against strong light; disk lobes, if connate, not 
into an intrastamineal crown; stamens 10-15. 
Calyx lobes reflexing from the base at anthesis; anthers 

glabrous. 
Leaves glabrous or glabrate. 

Filaments glabrous; pedicels elongate C. Pavoniana. 

Filaments villous below; pedicels to 10 mm. long. 

C. decandra. 

Leaves rusty- tomentose beneath C. Zahlbruckneri. 

Calyx lobes erect or erect-spreading at anthesis, sometimes 
widely spreading as in C. macrophylla but this with 
hirsute anthers. 

Style simple, the stigma capitate. 
Anthers glabrous and without a dorsal gland. 
Leaves softly pubescent beneath. 

Leaves drying light green or brown; ovary glabrous 

or strigose above C. obovalis. 

Leaves drying blackish; ovary pilose. 

Pedicels 6-7 mm. long; stamens subequal. 

C. nigricolor. 
Pedicels to 2 mm. long; stamens unequal. 

C. nigricans. 
Leaves glabrous or glabrate (or early pubescent, 

C. arguta). 
Leaves entire or essentially. 

Leaves to 1 dm. long, 4 cm. wide, often smaller; 
pedicels longer than flowers. 



FLORA OF PERU 39 

Stipules subulate, caducous C. petiolaris. 

Stipules lanceolate, subpersistent.C. prunifolia. 
Leaves usually 1.5 dm. long and one- third to one- 
half as wide; pedicels shorter than or about 
as long as flowers. 

Stipules conspicuous, subpersistent; stamens un- 
equal C. combaymensis. 

Stipules apparently small, caducous; stamens 

equal C. fasciculata. 

Leaves serrulate, sometimes minutely. 

Serrulations of leaves minute; calyx about 4 mm. 
long . . .C. tarapotina. 

Serrulations of leaves conspicuous; calyx 5-6 mm. 

long C. arguta. 

Anthers usually barbate, sometimes minutely, or at 
least with a dorsal gland. 

Dorsal gland glabrous (typically); leaves oblong- 
elliptic, lineate-pellucid C. Cambessedesii. 

Dorsal gland barbate, sometimes minutely. 
Flower-fascicles sessile; leaves thin, subentire; sepals 
(3) 4.5-6 mm. long C. macrophylla. 

Flower-fascicles often peduncled; leaves firm, more 

or less serrulate; sepals 2-4.5 mm. long. 
Buds rather oblong-ovoid; sepals oblong, about 
4 mm. long C. arborea. 

Buds globose or subglobose; sepals suborbicular, 
about 2 mm. long C. Blanchetiana. 

Styles trifid (minutely), the flowers almost tiny. 

Filaments somewhat pilose; anthers glandular. 

C. sylvestris. 

Filaments glabrous; anthers eglandular(?) . . . .C. ovoidea. 

Leaves not pellucid-punctate; disk lobes united into a crown; 
stamens 10-25; style trifid. 

Leaves glabrous; sepals reflexed C. javitensis. 

Leaves softly pilose beneath; sepals erect C. iquitosensis. 

Casearia aculeata Jacq. Enum. PL Carib. 21. 1760; 463. 
C. spinosa (L.) Willd. Sp. PI. 2: 626. 1799. Samyda spinosa L. Sp. 



40 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

PI. ed. 2. 557. 1762. C. berberoidea Rusby, Mem. N. Y. Bot. Gard. 
7: 307. 1927, fide Sleumer. 

Branchlets spinescent, usually glabrous or less frequently (in 
Peru), as the younger leaves beneath, rusty-pubescent; petiole 
sulcate above, 3-5 mm. long, the stipules minute and promptly 
caducous; leaves ovate or obovate-elliptic and oblong, obtuse or 
obtusely acuminate, acute at base, subentire or remotely crenate- 
dentate, firm or membranous, sometimes subcoriaceous, opaque 
both sides, generally pellucid-punctate and lineolate, 4-10 cm. long, 
2-5 cm. wide; lateral nerves 4-5, sometimes tardily glabrate beneath; 
fascicles sessile, 5-10 (-30) flowered; pedicels 3-6, rarely 8-9 mm. 
long, the rusty-pubescent or glabrate, whitish calyx 4 mm. long, 
nearly cylindrical, the 5 oblong lobes erect; stamens 8, glabrous, 
connate below with the tomentose oblong disk lobes; ovary villous, 
about equaled by the glabrous style, the stigma capitate; capsule 
baccate, about 1 cm. thick, the 3-10 seeds in a pulp. The var. 
Tafallana (Eichler) Macbr., comb, nov., the type as C. spinosa var. 
Tafallana Eichler in Mart. Fl. Bras. 13, pt. 1: 464. 1871, from Guaya- 
quil, has subcoriaceous leaves, obscurely observable pellucidity, 
nerves prominent beneath and fascicles 10-30-flowered. Twigs 
become thorns (Mexia). 

As Eichler remarked, the species is highly variable in pubescence 
and in form, dentation and size of leaves; however Sleumer has 
helpfully called attention to the fragility of the long slender pedicels, 
readily breaking in herbaria, and to the cylindrical (somewhat 
enlarged at base) yellowish flowers, "characters as yet unknown to 
me in any other species." But similar technically and probably to 
be found within Peru is C. guyanensis (Aublet) Urban, espinose, 
the leaves glabrate, papery, crenate, nerves 6-8, ovary sparsely 
pubescent. F.M. Negs. 13711 (var.); 13690. 

San Martin: Tarapoto, Spruce 4271; Ule 6649 (det. Pilger). 
Juanjui, Klug 3903; 3858. Loreto: near Iquitos, Mexia 6413 (det. 
Standl.); Tessmann 4962. Pongo de Manseriche, Mexia 6260 (det. 
Standl.). Florida, Klug 2326 (det. Standl.). Lower Rio Huallaga, 
Williams 5199. Tarapoto, Williams 5523; 5497; 5774; 5574; 6744; 
6895; 6158. Iquitos, Williams 8039. Yurimaguas, Williams 5199. 
Widely distributed in South America to Central America and the 
West Indies. "Espino del demonio," "espina cacha," "supiecacha" 
(Huitoto name), "naranjilla," "espuela casha," "supai cashi." 

Casearia arborea (L. C. Rich.) Urban, Symb. Ant. 4: 421. 1910; 
478. Samyda arborea L. C. Rich. Act. Soc. Hist. Nat. Paris 1: 109. 



FLORA OF PERU 41 

1792. C. stipularis Vent. Choix PI. 46. 1803. Chaetocrater capitatum 
R. & P. Syst. 1: 108. 1798. C. capitata (R. & P.) Spreng. Syst. 
Veg. 2: 355. 1825. C. Poeppigii Eichler in Mart. Fl. Bras. 13, pt. 1: 
475. 1871. 

Branches, at least when young, white-lenticellate; young branch- 
lets and leaves beneath minutely but densely ashy- or rusty-tomen- 
tulose, tardily glabrate or the leaves glabrate; petioles subterete, 
2-4 mm. long, the sometimes persisting stipules about twice as long; 
leaves oblong, acute at base, attenuate to tip, densely serrate-dentate, 
4-12 cm. long, 1-3.5 (-4) cm. wide, firm-membranous or subcoria- 
ceous, resinous or pellucid-punctate, glabrous and lustrous above; 
lateral nerves (4) 6-10; axillary umbels sessile or shortly peduncled, 
20-30-flowered, the pedicels 2-4 mm. long, equaling the typically 
ashy-tomentose calyx (in Peruvian plant glabrate), its lobes oblong; 
stamens 10, free, the anthers glandular-barbellate, the spathulate- 
oblong disk lobes half as long as the glabrous filaments; ovary 
strigose or glabrous, the style villous at base, the stigma ovoid- 
capitate; capsule angled with 3-4 foveolate seeds. Killip & Smith 
26977 with herb, name is a form, ovary glabrous, flower-cluster 
sessile. The type of C. Poeppigii is very young. F.M. Neg. 13687. 

Huanuco: Cuchero, Chinchao and Pozuzo, Ruiz & Pavon, type, 
C. capitata. San Martin: Tarapoto: Williams 6078; 6612; 6116. 
Loreto: Yurimaguas, Poeppig 3100, type; Williams 4741; 4579; 
4529. Near Iquitos, King 365; Killip & Smith 26977. Amazonian 
Brazil to Central America and the West Indies. "Llajas." 

Casearia arguta HBK. Nov. Gen. & Sp. 5: 364. 1823; 470. 
C. Fockeana Miq. Ann. Sci. Nat. se>. 3. 1: 39. 1844, fide Sleumer. 

Young branchlets and leaves, the latter, especially beneath on 
the nerves, gray-puberulent, glabrate in age; petioles 2-5 mm. long, 
the stipules scarcely longer, caducous; leaves elliptic- or lanceolate- 
oblong, little if at all unequal at the shortly acute base, acuminate, 
8-15 (-18) cm. long, 3-5.5 cm. wide, serrate-dentate, the mem- 
branous pellucid dots minute if obvious, the lateral nerves mostly 
6-9; umbels sessile, 20-30-flowered, the pedicels 3-6 (-8) mm. long, 
ashy-tomentose as the calyx, this 5-6 mm. long, the narrow sepals 
not reflexed; stamens 10, free, the filaments glabrous or villous 
below, the disk lobes hirsute, the anthers eglandular, glabrous, 
elliptic; style glabrous, globose, velvety- tomentose, or glabrate; 
seeds 6 mm. long, the arils fleshy. F.M. Neg. 13659. 

Rio Acre: Tree, 10 meters, Krukoff 5623; Ule 9624; 9625. 



42 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Casearia Blanchetiana Miq. Linnaea 22: 801. 1849; 475. 
C. celtidifolia Eichler in Mart. Fl. Bras. 13, pt. 1: 477. 1871. 

Similar to C. arborea but glabrous, the leaves thin, the buds 
ovoid-subglobose, the fascicles and flowers subsessile and only 
6-15-flowered, the sepals suborbicular. Some of the stipules may 
persist. The few hairs on the small anthers are often visible only 
under the binocular. The calyx is papery, drying yellowish or pale 
brown. F. M. Negs. 13661; 13665. 

Loreto: Florida, Rio Putumayo, Klug 2294 (det. Standl.). Rio 
Itaya, Williams 3393. Iquitos region, Tessmann 3910. Yurimaguas, 
Ule 6756; Poeppig 2263, type, C. celtidifolia. La Victoria, Williams 
2758. Balsapuerto, Klug 2982. Brazil. "Ullu-mullaca," "uchu 
mullaca." 

Casearia Cambessedesii Eichler, I.e. 475. C. Bangii Rusby, 
Mem. Torrey Club 3, no. 3: 34. 1893, fide Sleumer. 

Allied to C. arborea but the branchlets and leaves soon glabrate, 
the leaves abundantly punctate and, or only, resinous-lineate with 
7-12 lateral nerves, the buds ovoid, the calyx lobes broadly oblong- 
ovate, 4 mm. long, the disk lobes subequaling the barbate-based or 
glabrate filaments; anthers typically glabrous, sometimes sparsely 
hirsute as ovary or this with style strigose. The leaves and calyx 
are much firmer than in C. Blanchetiana. F.M. Neg. 13662. 

San Martin: Tarapoto, Williams 6116 (distr. as C. Poeppigii 
as also 6612, Alto Rio Huallaga). Loreto: Florida, Klug 2307. 
Balsapuerto, Klug 3112 (narrow leaves, some anthers sparsely 
hairy). Brazil to Colombia. 

Casearia combaymensis Tul. Ann. Sci. Nat. se>. 3. 7: 362. 
1847. Casearia singularis Eichler in Mart. Fl. Bras. 13, pt. 1: 473. 
1872, fide Sleumer. Casearia membranacea Britton, Bull. Torrey 
Club 17: 214. 1890, fide Sleumer. 

Branchlets and leaves glabrous; petiole 5-15 mm. long, stipules 
conspicuous, persisting on the branchlets toward their tips, green 
or pale, 5-8 mm. long; leaves obovate-oblong, nearly caudate- 
acuminate, attenuate at base, 15-25 cm. long, 5-10 cm. wide, often 
smaller, very thin to subcoriaceous, obscurely if at all resinous- 
pellucid, nearly opaque, entire, 6-8 lateral nerves prominent beneath ; 
fascicles axillary and lateral, about 10-flowered, the gray-puberulent 
pedicels (2-) 3-6 mm. long, and calyx subequal, the ovate-oblong 
suberect lobes of the calyx about twice as long as the subhemispheric 
tube; stamens 10, shortly connate with the disk lobes, the longer 



FLORA OF PERU 43 

filaments densely villous, their anthers glabrous, eglandulose, the 
alternate ones glabrous, their anthers larger; disk lobes linear- 
clavate, glabrous, about equaling the shorter stamens; ovary sub- 
globose, included in the calyx tube, glabrous as the very short 
style. C. singularis of the Guianas was described as having subulate 
caducous stipules and is probably distinct. F.M. Negs. 21322 
(singularis); 34893. 

Loreto: Ucayali region, Tessmann 3229. Junin: Puerto Ber- 
mudez, 375 meters, Killip & Smith 26448 (young). Loreto: Soledad, 
Killip & Smith 29578; 29781; 29692. San Antonio, Killip & Smith 
29358 (tree, 10 meters); 29517. Balsapuerto, Killip & Smith 28659. 
Rio Acre: Ule 9726. Brazil to Colombia and Guiana. 

Casearia decandra Jacq. Sel. Stirp. Am. Hist. 133. pi. 85. 1763; 
467. C. parvifolia Willd. Sp. PI. 2: 628. 1799. 

With the floral characters of C. arguta but the oblong-lanceolate 
calyx-lobes finally reflexing and the capsule always glabrate; otherr 
wise differs in the fugacious pubescence being rusty-tomentose, the 
ovate, rather abruptly acuminate leaves only 4-6 (-11) cm. long, 
1-3 (-4) finely serrate or subentire, the umbels sometimes fewer- 
flowered, the flowers 3-4 mm. long. With the distribution of 
C. arguta (Sleumer); pellucid dots and lines often many. Williams 
6219 not restudied. 

San Martin: Tarapoto, Williams 6219 (or aff.). Loreto: 
Pumayacu, 600-1,200 meters, King 3212 (det. A. C. Smith). Yarina 
Cocha, Tessmann 5415. Ucayali, Tessmann 3458. Florida, Klug 
2275; 2353. Mouth of Santiago, Mexia 6108. Warm Central and 
South America. "Fortuga caspi," "titibeguisi-ey" (Huitoto), 
"limon caspi." 

Casearia fasciculata (Ruiz & Pavon) Sleumer, Notizbl. Bot. 
Gart. Berlin 11: 955. 1934; as to name. Chaetocrater fasciculatum 
Ruiz & Pavon, Syst. 1: 107. 1798. Casearia maculata Pilger, Verh. 
Bot. Ver. Brandenb. 47: 161. 1905. 

Allied to C. combaymensis; petioles to 10 mm. long; leaves rather 
obovate-elliptic, acute at base, caudate-acuminate, to 2 dm. long, 
6 cm. wide, membranous, punctate-lineolate, pellucid, entire; 
fascicles dense, axillary, sessile; pedicels shorter than the ashy- 
puberulent flowers; calyx lobes rounded, about 2 mm. long; disk 
lobes narrow, somewhat villous, connate below with the slightly 
villous filaments; anthers broadly ovoid, glabrous; ovary glabrous, 
the stigma capitate. This follows Sleumer's interpretation of the 



44 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

species, accepting the type as the plant so labeled at Madrid; the 
specimen at Geneva, however, also with original label, is C. arborea. 
Of course the Madrid collection may best be taken as type. 

Huanuco: Chinchao about Hualqui, Ruiz & Pawn, type. Brazil. 

Casearia iquitosensis Macbr. Candollea 8: 22. 1940. 

Frutex vel arbuscula; ramulis foliisque subtus molliter et persis- 
tenter ferrugineo-pilosis; petiolis crassis 5-8 mm. longis; foliis 
chartaceo-coriaceis supra nervis impressis exceptis glabris, nitidis, 
utrinque prominenter laxeque reticulato-venosis, oblongo-ellipticis, 
basi rotundato-acutis, subabrupte breviterque acuminatis, plerumque 
2.5 dm. longis, 10 cm. latis plus minusve repando-crenato-denticu- 
latis; fasciculis florum plerumque axillaribus sessilibus; pedicellis 
ad 6 mm. longis adpresse pilosis; calycis tubo brevissimo, lobis 
erectis late oblongis, 5.5 mm. longis, fere 2.5 mm. latis, puberulis 
haud hirsutis; staminibus 10-15, glabris; disci lobis dense tomentosis 
filamento multo brevioribus in coronam intrastaminealem conni- 
ventibus; stylo inferne villoso, apice trifido; capsula trigono-ovidea 
10 mm. longa, extus dense ferrugineo-puberula, intus ferrugineo- 
floccosa; seminibus 1-3, subglobosis molliter tomentoso-pilosis. 
Nearly C. dentata (Aubl.) Eich. of Guiana with 6-angled glabrate cap- 
sules and C. lasiosperma Tr. & PL, Colombian, with leaf -nerves ele- 
vated above, hirsutulous calyces and glabrate capsules 15 mm. long. 

San Martin: Near Moyobamba, Klug 3560 (det. Standl.). 
Loreto: Iquitos, Williams 3708; 3737 (type fruit and flower); 7075; 
3749; 8015. Rio Acre: Killip & Smith 29836; Klug 545; 499; 3; 
98; Krukoff 5237. 

Casearia javitensis HBK. Nov. Gen. & Sp. 5: 366. pi 
479. 1823. 

Shrub or tree with widely spreading simple branches, the younger, 
as the leaves on the nerves beneath, puberulent but soon glabrate; 
petiole 3-10 mm. long; leaves usually oblong, obtuse at base, acumi- 
nate, subentire or more or less serrate-dentate, firm-membranous, 
not punctate, 6-30 cm. long, 2.5-10 cm. wide; lateral nerves 5-7; 
fascicles axillary, sessile, 20-60-flowered, the pedicels usually 4-7 mm. 
long, gray-puberulent as the calyx, this 3-4 (-5) mm. long, with very 
short tube, the oblong-obtuse lobes reflexing at an thesis; stamens 
10 or 15, glabrous, disk lobes linear-clavate, rusty-tomentose, 
connivent in a crown between the stamens; style tomentose, trifid, 
the stigmas capitate; capsules tomentose, the 1-2 globose seeds 
pubescent. F.M. Neg. 13675. 



FLORA OF PERU 45 

Loreto: Yurimaguas, Williams 3883; 4590 (pedicels longer). 
Caballo-Cocha, Williams 2317. Mishuyacu, King 227; 520. Iquitos, 
Mexia 6272 (det. A. C. Sm.); Kittip & Smith 27282. San Martin: 
Pongo de Cainarachi; Klug 2672 (det. Standl.). Rio Acre: Ule 
9626. North to Central America and the Guianas. "Capanca." 

Casearia macrophylla Vahl, Eclog. Am. 2: 32. 1798; 470. 

Glabrous or the growing parts sparsely puberulent; petioles 5-7 
mm. long; leaves elliptic-oblong or obovate-oblong, acute at base, 
acuminate, subentire, 1-2 dm. long, 4-10 cm. wide, usually minutely 
pellucid-puncticulate; umbels 10-20-flowered; anthers subrotund, 
glandular barbate-villous; ovary glabrous, style hirsute, the globose 
capsule tubercled but glabrous, the seeds smooth, irregularly 
flattened. Otherwise similar to C. arguta. 

A puzzling shrub, possibly distinct, may be named var. barbatula 
Macbr., var. nov., foliis subcoriaceis, floribus paucis, vix 3 mm. 
longis: Iquitos, Killip & Smith 26977 (type); and 27375. Some of 
the material cited here or under C. tarapotina that is in fruit may 
be misdetermined. F.M. Neg. 21323. 

San Martin: Tarapoto, Spruce 4887 (var.). Loreto: Mishuyacu 
near Iquitos, Klug 20. Yurimaguas, Williams 4456. Rio Nanay, 
Williams 735. Timbuchi, Williams 995 (with herb. name). Florida, 
Klug 2281; 2257; 2241 (flowers white). Balsapuerto and Pongo 
Cainarachi, flowers greenish-cream, Klug 2637; 2665; 3026 (all det. 
Standl.). Yurimaguas, Williams 4934; 4614; Killip & Smith 
29029; 28314 (this last tree, 8 meters). Brazil to Colombia and 
Guiana. "Usico-ey," "cuipe-ey," "sasishy-ey" (Huitoto), "achu- 
caspi," "uchu caspi," "oje de tucunare." 

Casearia nigricans Sleumer, Notizbl. Bot. Gart. Berlin 11: 
956. 1934. 

Rusty-pilose tomentose even to the calyx, except the leaves 
above; leaves lanceolate or oblanceolate, mostly broadest at the 
middle, strongly attenuate to the short petiole (3-4 mm. long), 
long-acuminate, minutely toothed, the teeth about 0.5 mm. high, 
3-4 mm. distant, subglabrous above; lateral nerves 10-12; fascicles 
axillary, 10-15-flowered, sessile; pedicels to about 1.5 mm. long; 
sepals 5, ovate-oblong, obtuse, blackish, about 4 mm. long; stamens 
10, the 5 longer 2.5 mm. long, the shorter alternating with the 
pilose disk lobes; ovary ovoid, densely pilose. Section Pitumba 
(Sleumer). Known to attain 5 meters. Leaves thin, drying black. 
Here perhaps belongs Tessmann 3569, region Maranon. 



46 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Mountain east of Moyobamba, 1,200 meters (Weber- 
bauer 4727, type). 

Casearia nigricolor Sleumer, Notizbl. Bot. Gart. Berlin 11: 

957. 1934. 

Shrub with sparsely pubescent branchlets and large, thin, sub- 
sessile leaves drying blackish, although rusty- tomentose beneath, 
subglabrous and lustrous above; leaves oblong-ovate or broadly 
oblong, rounded at base, shortly attenuate at tip, distinctly punctate 
against a strong light, subentire or the teeth minute, 8.5-16 cm. 
long, 4-7.5 cm. wide, the nerves little prominent even below; fascicles 
15-20-flowered, sessile; pedicels puberulent, 6-7 mm. long; sepals 5, 
ovate-oblong, obtuse, puberulent both sides, at anthesis spreading or 
reflexing, about 3 mm. long, stamens 10, all about as long as the 
sepals, the filaments glabrous, the anthers globose, the disk lobes 
villous; ovary ovoid, pilose. Section Pitumba (Sleumer). Type 

5 meters high. It has been distributed as a new species by me and 
under a valid name. The type sheet is at Field Museum, not 
"Dahlem." 

Libertad: Valley system of the Rio Mixiollo, 1,800 meters, 
Weberbauer 7051 (type, Field Museum). 

Casearia ovoidea Sleumer, Notizbl. Bot. Gart. Berlin 11: 

958. 1934. 

Small tree, glabrous except for a little pubescence on the ovoid 
ovary and the three-forked style; petioles about 7 mm. long; leaves 
ovate or ovate-oblong, broadest above the middle, equilateral, sub- 
obtuse at base, obtusely acuminate, membranous, punctate with 
lines and dots, somewhat lustrous above, minutely and obtusely 
dentate, the lateral nerves distinctly marked above; fascicles sessile, 
the pedicels 2 mm. long; sepals 5, oblong, obtuse, erect, 1.5 mm. 
long; stamens 10, glabrous, the filaments filiform, the disk lobes 
pubescent, short; style pubescent. Type 5 meters high. Section 
Crateria (Sleumer). This seems to be a species doubtfully distinct 
from C. sylvestris, or better treated as a variety. 

Junin: On open mountain above La Merced, 5260, type; Killip 

6 Smith 23672; 23809. 

Casearia obovalis Poeppig ex Griseb. Erlaut. Trop. Pflz. 27. 
1860; Eichler in Mart. Fl. Bras. 13, pt. 1: 472. 1871. C. obovata 
Poeppig ex Eichler, I.e., not Schlecht., 1839. C. commutata Briq. 
Ann. Conserv. Jard. Bot. Geneve 2: 65. 1898. 



FLORA OF PERU 47 

Leaves loosely grayish-brown pilose beneath, glabrate above 
unless somewhat puberulent on the nerves, oblong-elliptic or often 
obovate, usually acuminate or cuspidate, entire or subentire, opaque, 
green, membranous, 6-15 cm. long, 3-6 cm. wide, the lateral nerves 
about 5; petioles about 2 mm. long, sometimes longer, somewhat 
villous, as the young branchlets; fascicles axillary, sessile, several 
to many-flowered; pedicels 2-3 mm. long; calyx to 4 mm. long, the 
short tube turbinate, the longer lobes erect, oblong-lanceolate; 
stamens 10, glabrous, slightly unequal, the lower half connate with 
the apically barbate disk lobes; style becoming as long as the shorter 
stamens, glabrous, the stigma capitate; fruit orange-yellow, about 

2 cm. thick, the few seeds nearly 10 mm. long. Evidently Eichler 
inadvertently wrote the name "obovata," because he says "Poeppig 
mss." and Poeppig wrote, as quoted by Grisebach and also by 
Eichler in referring to the Grisebach reference, "obovalis," which 
was indeed Poeppig's choice of name. I therefore think that there 
is no need of a new cognomen if the original one is written correctly. 
In any case the identity of the species being definite, the Grisebach 
reference constitutes publication. F.M. Neg. 13668. 

San Martin: Tarapoto, Spruce 3894- Loreto: Yurimaguas, 
Poeppig 2438, type; Kittip & Smith 28015; 29097. Region Ucayali, 
Tessmann 3184. Rio Acre: Krukoff 5507. Brazil. 

Case-aria Pavoniana Sleumer, Notizbl. Bot. Gart. Berlin 11: 
958. 1934. 

Tree with brownish tomentulose or subglabrous branchlets and 
rather oblique-based regularly and acutely dentate leaves; petioles 
about 5 mm. long; leaves moderately acuminate, glabrous, some- 
what lustrous above, membranous and distinctly pellucid-punctate, 
the lateral nerves well-marked only beneath; fascicles 15-20-flowered, 
sessile; pedicels slender, tomentose, some scarcely 10 mm. long, or 
shorter than 1.5 cm.; sepals lanceolate, obtuse, margins puberulent, 

3 mm. long; stamens 10, the 5 longer equaling the sepals, the filaments 
glabrous, the very short disk lobes villous as the ovoid ovary that 
is extended by style as long as the longer stamens. Section Pitumba 
(Sleumer). According to the author, Ruiz & Pavon named it under 
another genus in volume 4 of their Flora but he has made the name 
untenable in Casearia by using it for another species; if the plate in 
volume 4 was with analysis the name was not "ined." F.M. Neg. 
28940. 

Junin: Vitoc, Ruiz & Pavon, type. La Merced, 1,000 meters, 
Weberbauer 1862. 



48 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Casearia petiolaris Poeppig ex Eichler in Mart. Fl. Bras. 13, 
pt. 1: 471. 1871. 

Allied to C. obovalis but glabrous or essentially so except the 
ashy-tomentulose pedicels and calyces; leaves oblong, rather long- 
attenuate to the petiole, this 6-8 mm. long, acuminate, the resin 
glands few and little pellucid; pedicels 3-5 mm. long; stamens free, 
the filaments ciliate; style villous at base. F.M. Neg. 24109. 

Loreto: Yurimaguas, Poeppig 2482, type. 

Casearia prunifolia HBK. Nov. Gen. & Sp. 5: 364. 1823. 

Stipules lanceolate, persisting, those on the sterile shoots to 5 mm. 
long; leaves thin, pellucid-punctate in the type (not according to 
original description), oblong-lanceolate, 8-10 cm. long, mostly 2.5- 
3.5 cm. wide, gradually caudate-acuminate, subentire, subsessile at 
the attenuate base, glabrous, the lateral nerves (about 5) with the 
slender veins rather prominent beneath; flowers few, the clusters 
sessile, the pedicels puberulent, to 5 mm. long; sepals appressed, 
pubescent or glabrate without, suberect or perhaps reflexing, acute, 
nearly 3 mm. long; filaments lightly pilose above, exserted, the 
alternate shorter, connate below into a tube, twice as long as the 
densely short-hirsute staminodes; anthers glabrous; style pilose, 
simple, the stigma capitate. In the type at Paris the leaves are 
pellucid-punctate (as to attached leaves) but there may be some 
mixture with Xylosma prunifolia (HBK.) Griseb., which is a true 
Xylosma of Colombia without staminodes. The Colombian Fla- 
courtia Benthami Tul. (F. prunifolia as to Bentham, PL Hartw. 
160, not HBK.), also confused, has leaves 3-4 cm. wide, 5-8 cm. 
long, somewhat callous-toothed; cf. C. combaymensis. F.M. Neg. 
34888. 

Cajamarca: Bacamores near Jae"n, Bonpland, type. 

Casearia resinifera Spruce ex Eichler, I.e. 466. 

Unarmed, glabrous, the dark-colored elenticellate branchlets 
exuding a resin at the leaf nodes; petioles 5-10 mm. long; leaves 
oblong or narrowly oblong, acute or broadly cuneate at base, rather 
caudate-acuminate, 1.5-2.5 dm. long, 4.5-7 cm. wide, subentire or 
remotely serrulate, subcoriaceous, most densely and minutely pel- 
lucid-punctate, pellucid lines none, lateral nerves 11-13, subperpen- 
dicular; capsules ovate-oblong, 14 mm. long, obtuse, glabrous, the 
seeds many, smoothish, 4 mm. long. F.M. Neg. 13688. 

Loreto: Mishuyacu, Klug 307. Brazil. 



FLORA OF PERU 49 

Casearia sylvestris Sw. Fl. Ind. Occ. 2: 752. 1800; 481. 

Shrub or small tree, the growing parts at least more or less rusty- 
puberulent but typically glabrate in age (cf. variety); branches 
gray, lenticellate; stipules cordate-ovate or suborbicular, minute, 
caducous; petioles 2-10 mm. long; leaves oblong or elliptic and lanceo- 
late, little if at all unequal at the acute narrowed or rounded base, 
mostly narrowly acuminate and average size, that is, 6-9 cm. 
long, 2-3 cm. wide, or only somewhat smaller or larger, serrulate 
dentate or subentire, membranous to coriaceous, usually the former, 
usually minutely pellucid-punctate and lineolate, the lateral nerves 
5-8; umbels axillary, sessile, 10-many-flowered; pedicels glabrous or 
puberulent, 2-4 mm. long, subequaled by the glabrate-puberulent 
pale flowers; calyx tube campanulate, the lobes broad, erect-spread- 
ing; stamens 10, free, the filaments sparsely pilose, the subglobose 
anthers glandular, glabrous; disk lobes spathulate, densely tomen- 
tose-barbate, scarcely to much shorter than the filaments; style 
trifid, glabrous or at base pilose; capsule glabrous. Highly variable 
in foliage and pubescence. The var. chlorophoidea (Rusby) Sleumer 
has glabrous flowers, the leaves entire or nearly; var. Lingua (Camb.) 
Eichler, the leaves beneath, at least the younger, the petioles, pedicels 
and calyces finely gray-tomentulose, and the floral parts more pilose. 
This form is called "lingua de Tiu" in Brazil. Variable as is the 
species it is marked by the free insertion of the disk lobes among 
the stamens, the style nevertheless trifid. A few collections of the 
many known are: 

San Martin: San Roque, Williams 7526; Ule 6459 (det. Pilger); 
Spruce 4059. Alto Rio Huallaga, Williams 6831; Weberbauer 6824 
(form with opaque leaves with herb. name). Huanuco: Pillao, 
Pozuzo, Ruiz & Pavdn. Junin: La Merced, Killip & Smith 23809; 
25097. Ayacucho: Near Kimpitiriki, 400 meters, Killip & Smith 
22942. Loreto: Pumayacu, King 3160; 3193 (det. Standl.). Cachi- 
puerto, King 3132 (det. Standl.). Florida, King 2184 (det. Standl.). 
Caballo-Cocha, Williams 2436. Pongo de Manseriche, Tessmann 
4256. Maranon region, Tessmann 4208. Puno: San Gaban, Lechler 
2514 (cf. C. jasciculata). Rio Acre: Ule 9631; Krukoff 5583; 5239; 
5386; 5233. Lower altitudes, warm America. "Sishi-co-ey" (Huitoto). 

Casearia tarapotina Pilger, Verh. Bot. Ver. Brandenb. 47: 161. 
1905. 

Branchlets glabrous; leaves elliptic, shortly acute at base or 
rounded, more or less acuminately and acutely pointed, obsoletely 
denticulate, 8-10 cm. long, 3-4 cm. wide, pellucid-punctate and 



50 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lineolate, the 8-9 lateral nerves prominent beneath; umbels axillary; 
pedicels puberulent, 5-7 mm. long; calyx lobes broadly lanceolate, 
obtuse, 4 mm. long, little puberulent; stamens free, unequal, 5 nearly 
equal, 5 longer than the villous disk lobes; anther ovoid, glabrous; 
ovary glabrous, the style villous below; fruit (as to specimens cited) 
about 6 mm. in diameter, nearly mature. Section Pituma (Pilger) . 
F.M. Neg. 13691. 

San Martin: Tarapoto, Ule 6936, type; Williams 6096 (fr. 
strigose at tip, 5 mm. long, 7 mm. thick); 6723; 6590. Lamas, 
Williams 6403. Loreto: Ucayali region, Tessmann 3061. "Tambor 
huactana." 

17. LAETIALoefl. 

Shrubs or trees with entire or toothed opaque or punctate leaves, 
deciduous stipules and rather large hermaphrodite flowers in terminal 
or axillary cymes or clusters. Bractlets sometimes united. Sepals 
free or nearly so, 4-5, somewhat petaloid. Petals none. Stamens 
10-15 or many, hypogynous or the outer somewhat perigynous, the 
filaments free, the anthers oblong-ovoid, tiny. Disk lacking or not 
marked. Ovary free, 1-celled with 3 placentae. Style lacking, 
simple or 3-parted. Capsule berry-like, dehiscent, the seeds imbedded 
in fleshy arils. Name conserved as of Loefling but original publica- 
tion, Loefling, Iter Hispan. 190. 1758 (German ed. 252) is only by 
inference and the name is scarcely noticeable on the page because 
it is in small type in synonomy under the name Guidonia P. Br., 
this in conspicuous letters. In the German edition both names are 
in the same size of type, but Laetia apetala is in parentheses. 

Flowers glabrous or nearly so; leaves membranous or firm. 
Flowers in clusters on the stems. 

Flower clusters mostly or all axillary L. suaveolens. 

Flower clusters mostly or all extra-axillary L. procera. 

Flowers in dichotomous or simple cymes. 

Leaves short-acuminate or acutish, 6-9 cm. long . .L. corymbulosa. 

Leaves blunt, mostly about 4 cm. long L. apetala. 

Flowers pubescent; leaves heavy coriaceous. 

Leaves oblong; "cup" subsessile, to 5 mm. long L. coriacea. 

Leaves oval; "cup" pedicellate, 2.5-3 mm. long L. ovalifolia. 

Laetia apetala Jacq. Sel. Stirp. Amer. 167. pi. 108. 1763. 
Similar to L. corymbulosa but the leaves more uniformly obovate, 
rounded or blunt at apex and the cymes simple or 2-dichotomous, 



FLORA OF PERU 51 

sometimes puberulent. Probably only one species is concerned that 
is variable in these apparent differences. 

Loreto: Region of the Ucayali, Tessmann 3225; 3405. Brazil to 
Colombia. 

Laetia coriacea Spruce ex Benth. Journ. Linn. Soc. Bot. 5: 
Suppl. 2: 84. 1861. 

Leaves rigid coriaceous and extremely lustrous both sides, oblong- 
elliptic, rounded at base, shortly acuminate, about 14 cm. long, 
5-6.5 cm. wide, probably becoming larger, wavy margined; petioles 

6 mm. long; sepals about 5 mm. long; bractlets united into a deeply 
bilobed cup 5 mm. long, this sessile or subsessile, the pedicels to 

7 mm. long; young fruit globose, densely tomentose. L. cupulata 
Spruce, I.e., Amazonian, is similar but the pedicels are more than 
twice as long as the entire cup. F.M. Neg. 24071. 

Loreto: Iquitos, Kuhlmann 19343 (det. Kuhlm.). Brazil; 
Venezuela. 

Laetia corymbulosa Spruce ex Benth. I.e. 83. 

Tree with oblong-elliptic-obovate leaves and small flowers borne 
laxly in 2-4 dichotomous cymes at the ends of short branchlets; 
petioles 4-7 mm. long; leaves membranous, glabrous, minutely and 
obscurely serrulate, pellucid punctate, slightly cordate and unequal 
at base, rounded or shortly acuminate at apex, 5-9 cm. long, 2.5-4 
cm. wide; cymes to 4 cm. long; pedicels 1 cm. long, the minute 
bractlets more or less puberulent. The variety floribunda (Spruce) 
Eichler has corymbs to 20-flowered, even 4-times dichotomous. 
The following collection (Mexia) was distributed as "Banara laxi- 
flom." The collector noted it as a tree 40 meters high, growing 
higher, common, the wood used for firewood. It is perhaps a variety 
of L. apetala. F.M. Negs. 6511; 34869 (var.). 

Loreto: Left bank of Rio Maranon above Rancho Indiana, 
Mexia 6404- Brazil. "Teareo." 

Laetia ovalifolia Macbr. Candollea 5: 389. 1934. 

Branchlets smooth; petioles strongly sulcate, 13 mm. long; 
leaves broadly elliptic or rotund-elliptic, sometimes 2 dm. long, 
1 dm. wide, but mostly 10-12 cm. long, 6.5-8 cm. wide, rounded at 
base, abruptly and shortly acuminate, coriaceous, lustrous, espe- 
cially above, conspicuously reticulate-veiny on both sides, the 5 
lateral nerves prominent; fascicles sometimes extra-axillary, often 
8-10-flowered; bracteal cup on pedicel 1 mm. long, glabrous, irregu- 



52 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

larly lobed, 2.5-3 mm. long; pedicels as the sepals, silky, ashy- 
pilose without, both 8-9 mm. long; sepals strongly reflexed, puberu- 
lent within, oblong, about 5 mm. long; style trifid at tip, about 
6 mm. long; ovary densely rusty-pilose. Flowers of the type 
noted as white and rose color; of 656 as white, dark green and red; 
a tree to 15 meters high. 

Loreto: Mishuyacu near Iquitos, King 757, type; 656; 814. 

Laetia procera (Poepp. & Endl.) Eichler, Mart. Fl. Bras. 13, 
pt. 1: 453. 1871. Samyda procera Poepp. & Endl. Nov. & Gen. & 
Sp. 3: 67. 1845. 

Sometimes a tall tree; petioles 7-15 mm. long; leaves elliptic- 
oblong, 10 cm. long, 4 cm. wide, sometimes much longer, little 
wider, minutely cordate at base, abruptly short-acuminate, minutely 
appressed denticulate, finely reticulate-veined on both sides, scarcely 
lustrous, pellucid-punctate; fascicles usually 15-30-flowered, the 
pedicels about as long as the petioles; sepals greenish-white, obtuse, 
reflexing, 3-4 mm. long; stamens 12-20; fruit 1.5-2 cm. thick, sub- 
globose, with foveolate seeds about 3 mm. long. To 40 meters high. 
Flowers without odor (Poeppig). F.M. Neg. 24073. 

Loreto: Mishuyacu, Klug 495. Brazil to Guiana and West 
Indies. 

Laetia suaveolens (Poepp. & Endl.) Benth. Journ. Linn. Soc. 5: 
Suppl. 2: 85. 1861. Samyda suaveolens Poepp. & Endl. Nov. Gen. & 
Sp. 3:66. pi. 274. 1845. 

Similar to L. procera but leaves obtuse or acutish at base, heavier, 
densely reticulate-veined, rather lustrous; flowers fewer (4-12), the 
stamens only 10-12 and of diverse lengths; capsules 2 cm. thick or 
larger, the seeds in a red pulp, smooth. Flowers with the fragrance 
of orange blossoms (Poeppig). F.M. Neg. 13636. 

Loreto: Iquitos, Tessmann 3684; Klug 461; 1443. Brazil. 
"Timarehua." 

LACISTEMACEAE 

By Charles Baehni 

A small family of two genera and 20-odd species distributed in 
the New World from Mexico and the West Indies to Peru and 
Paraguay. The flowers are very small, so small that the characters 
used to recognize one species from the next can only be found by 
using a lens. We were of the opinion that good and reliable charac- 



FLORA OF PERU 53 

ters could be found in the flowers alone; study of ample material 
has indeed revealed that there exists a variability in these small 
flowers which could easily justify a splitting of species such as has 
been done in Rosa, Rubus, and other similar genera. Fortunately, 
fairly stable combinations of characters permit a more conservative 
treatment of this family; we cannot help feeling, however, that the 
limits being here difficult to trace, the circumscription of species is 
rather a matter of appreciation. 

There is no known use of any Lacistemaceae. 

Flowers borne on pedicels; inflorescence raceme-like 1. Lozania. 

Flowers sessile; inflorescence catkin-like 2. Lacistema. 

1. LOZANIA [Sinf.] Mutis 
Reference: L. B. Smith, Phytologia 1: 138. 1935. 

Inflorescences raceme-like, each flower borne on a distinct pedicel. 
Basal bract small. Sepals 4, broad, connate at base. Petals absent. 
Stamen 1, 6 ovules in the unilocular ovary. Fruit, the size of a pea, 
opening from the apex in 3 valves. 

Lozania Klugii Mansf. Notizbl. Bot. Gart. Berlin 11: 596. 1932. 
Monandrodendron Klugii Mansf. Repert. Sp. Nov. 30: 178. 1932. 

Small tree (6 meters) ; leaves 6-11 cm. long, 2-3 cm. broad, oblong- 
elliptic, acuminate, cuneate at base, denticulate, covered with 
appressed hairs; petiole 0.2-0.3 cm. long; stipule 0.4 cm. long; 
inflorescence racemose, 2.5 cm. long; flowers yellow-green; stamen 
longer than ovary; anther cells vertically disposed; ovary glabrous; 
style extremely short; capsules pilose, the seed bright orange color. 

San Martin: Pongo de Cainarachi, King 2711; 2681. Loreto: 
Mishuyacu near Iquitos, Klug 1430. Florida, Rio Putumayo, mouth 
of Rio Zubineta, Klug 2297. Pumayacu, Klug 3250. Colombia. 

Lozania Mutisiana Roem. & Schult. Mant. Add. 1: 75. 1822. 
L. nemoralis DC. Prodr. 3: 30. 1828. Monandrodendron Schultzei 
Mansf. Notizbl. Bot. Gart. Berlin 10: 860. 1929. M. peruvianum 
Mansf. Repert. Sp. Nov. 29: 161. 1931. 

Small tree or tree 6-20 meters high; leaves 8-13 cm. long, 2.5-3 
cm. broad, ovate to elongate-elliptic or even lanceolate, acute or 
acuminate, cuneate or rounded at base, glabrous beneath or puberu- 
lous or covered with a felt-like indument; petiole 0.8-1.2 cm. long, 
the rest as in the genus; stamen and ovary subequal; anther cells 
horizontally disposed; indument of ovary variable. The variability 



54 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

of this species is so great that it may be found later to include L. 
Klugii. 

Junin: Pichis Trail, San Nicolas, Killip & Smith 25968. 
Loreto: Pumayacu, King 3204. Florida, King 1 976. San Martin: 
Pongo de Cainarachi, Klug 2694- Venezuela; Colombia. 

2. LAGISTEMA Swartz 

Flowers sessile, disposed in spikes looking like catkins. Basal 
bract large, enclosing the whole flower. Sepals free, often wanting. 
Petals absent. Stamen 1. Ovules 3-6. Fruits like those of Lozania. 

Disk large, not completely hidden by the protective bract. 

Leaves large, 20-35 cm. long L. Macbridii. 

Leaves much smaller. 

Sepal 1 L. rosidiscum. 

Sepals 4. 

Leaves pilose on the middle nerve beneath L. nena. 

Leaves entirely glabrous L. aggregatum. 

Disk narrow, hidden by the protective bract. 

Sepals 2-3 L. purpureum. 

Sepals 4, rarely 3. 
Leaves glabrous below, generally shallowly serrate. 

L. aggregatum. 
Leaves pubescent below, entire. 

Petioles very short (2-2.5 mm.) L. curtum. 

Petioles long (5 mm. and more). 

Sepals broadly ovate L. Poeppigii. 

Sepals oblong-lanceolate L. Weberbaueri. 

Lacistema aggregatum (Berg.) Rusby, Bull. N. Y. Bot. Card. 
4: 447. 1907. Piper aggregatum Berg. Act. Hebr. 7: 131. pi. 10. 
1777. L. myricoides Sw. Prodr. 12. 1788. L. bolivianum Gandoger, 
Bull. Soc. Bot. Fr. 66: 288. 1919. 

Bush or small tree, up to 12 meters high; leaves mostly entire 
or merely undulate but sometimes repando-dentate or even ser- 
rulate, elliptic, acute or acuminate, glabrous, variable in size (7-18 
cm. long, 2.5-7 cm. broad); flowers cream-colored; spikes longer 
than the petioles (up to 12 mm.); sepals 4, lanceolate, serrulate; 
disk variable in size and shape, but generally membranous and 
reddish on dried specimens, more often small and hidden by the 



FLORA OF PERU 55 

bract than large and conspicuous; fruit ellipsoid, glabrous, 6-8 mm. 
long, brownish-red when ripe. 

Loreto: Iquitos, Rio Napo, near Mazan, Mexia 6446; basin of 
Maranon, Tessmann 5340. Florida, Rio Putumayo, at mouth of 
Rio Zubineta, King 2180, 2239. Huanuco: Lower Rio Huallaga, 
Williams 4680; above Cayumba, Mexia 8308. Junin: Hacienda 
Schunke, 5804 (det. Killip). From Mexico to Peru; West Indies. 
"Sishi-co-ey" (Huitoto), "huacapurana," "trompo huayo," "palometa 
huayo." 

Lacistema curtum Macbr. Candollea 5: 393. 1934. 

Tree 5 meters high ; leaves conspicuously pilose on the nerves below 
and on the petioles, glabrous above, entire, caudate-acuminate at 
tip, acute at base, oblong-elliptic, 7-14 cm. long, 2.5-4.5 cm. broad; 
stipule caducous; spikes shorter than the petioles (less than 2.5 mm.) ; 
bracts broadly triangular, slightly acute, finely toothed at margin, 
hirsute-pilose at base alone; sepals 4, oblong-lanceolate, acute, very 
thin, more or less dilacerated. 

Junin: Above San Ramon, 1,400-1,700 meters, Killip & Smith 
24617, type. 

Lacistema Macbridii Baehni, Candollea 8: 42. 1940. 

Leaves remarkably large (to 35 cm. long, 12 cm. broad), entire, 
acuminate, acute at the base, with short petioles (0.5-10 mm. long), 
entirely glabrous; spike (only one seen) 1.5 cm. long; bracts ciliate; 
sepals 4, ovate, denticulate or irregularly lacerate at the tip; disk 
very large, unlobed, uncleft, somewhat triangular; fruit unknown. 

Loreto: Iquitos, Basin of Maranon, between Iquitos and mouth 
of Santiago at Pongo de Manseriche, Tessmann 5282, type. 

Lacistema nena Macbr. Candollea 5: 392. 1934. 

Twigs slender, pilose but becoming glabrous; petiole 4-5 mm. 
long, slightly pilose; leaves entire, acuminate, glabrous beneath 
except on the middle and lateral nerves, 14 cm. long and 4 cm. broad; 
spikes about 1 cm. long, with acute bracts; sepals 4, obscurely 
denticulate; disk conspicuously large, not lobed; fruit sessile, densely 
covered with grayish hairs. 

Loreto: Caballo-Cocha, on the Amazon, Williams 2154; Pebas 
on the Amazon, Williams 1876, type. Amazonas: Basin of Maranon, 
mouth of Santiago at Pongo de Manseriche, Tessmann 4323. "Nena," 
"huacapurana." 



56 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Lacistema Poeppigii A. DC. Prodr. 16, pt. 2: 593. 1868. 

Leaves elliptic or oblong, acuminate, glabrous above, slightly 
appressed-pubescent below, 6-12 cm. long, 2.5-3.5 cm. broad; 
petioles 5 mm. long, spikes about the same length; sepals 4, broadly 
obovate, now and then dentate; disk sublobate, remarkably narrow; 
fruit 8-16 mm. long, shortly stipitate. F.M. Neg. 8075. 

Maynas: Poeppig, type. 

Lacistema purpureum (R. & P.) A. DC. Prodr. 16, pt. 2: 
593. 1868. Synzyganthera purpurea R. & P. Syst. 1: 273. 1798. 

Twigs glabrous; leaves elliptic or oblong-lanceolate, entire or 
remotely subdenticulate, glabrous except below on the nerves, 8-10 
cm. long and 2.5-4 cm. broad; spikes about the same length as the 
petioles (5-7 mm.); sepals 2-3; disk split, indistinctly lobed; fruit 
glabrous, about 12 mm. long. 

Huanuco: "Mufia et Pillao," near Chacahuassi, Ruiz & Pavon 513. 

Lacistema rosidiscum Macbr. Candollea 5: 392. 1934. 

Small tree; leaves oblong-elliptic or even lanceolate, 8 cm. long 
and 2.5-3.5 cm. broad, glabrous above and slightly shining, dull 
below and glabrous also but sometimes pilose, especially on the 
nerves; stipules lanceolate, caducous; spikes 3-5 mm. long, shorter 
than the petioles; disk of flower large, pink, deeply split and irregu- 
larly undulate; 1 sepal narrowly oblong. Williams 6721 has narrower 
leaves than the type (2.5 cm.). 

San Martin: Tarapoto, Williams 7216, type; Williams 6721. 

Lacistema Weberbaueri Baehni, Candollea 8: 46. 1940. 

Leaves much like those of L. Poeppigii, elliptic, shallowly serrate, 
acuminate or even caudate, acute at base, with long appressed hairs 
on the nerves below, 6-8 cm. long, 2.5-3 cm. broad; petiole slender, 
5-7 mm. long; stipules caducous, 6-7 mm. long, covered on the 
middle of the back with more or less stiff yellow hairs; spikes up to 
18 mm. long; flower completely hidden by bract; sepals 3-4, lanceo- 
late or linear, very rarely dentate on one side; disk narrow, slightly 
cleft; fruit ovoid, glabrous, stipitate. 

Loreto: Moyobamba, Weberbauer 44-97, type. 

VIOLACEAE. Violet Family 
By Charles Baehni and R. Weibel 

Reference: Gingins in DC. Prodr. 1: 287-316. 1824. 
Although the family is a very natural one, it is usually easy to 
distinguish one genus from the others, because of striking characters 



FLORA OF PERU 57 

readily discernible in the flowers or in the fruits. There is only one 
genus which has been accepted here with some doubt: the genus 
Rinoreocarpus. The characters indicated by its author, Ducke, are 
beyond doubt of value to the field taxonomist; for the herbarium- 
worker, it seems difficult to separate it from Rinorea. The Andean 
species of Viola need revision. However, the material at our 
disposal was insufficient to alter the accepted limits; it is to be 
hoped that, when foreign exchanges are possible, someone will 
undertake such a revision. 

The fruit of Leonia glycycarpa R. & P. is edible; the roots of 
Corynostylis, Viola, and Hybanthus yield an emetic; those oiAnchietea 
(source of Anchietin) furnish a liniment. Otherwise, besides being 
agreeable as ornamental plants, Violaceae are of little use to man. 

Fruit a berry or a capsule. 

Flowers regular (actinomorphous) or slightly irregular; petals 

not spurred nor saccate. 
Flowers regular. 
Fruit a capsule. 
Connective dilated dorsally into a large appendage. 

1. Rinorea. 
Connective expanded into a small apical scale. 

2. Rinoreocarpus. 

Fruit a berry 3. Gloeospermum. 

Flowers slightly irregular (petals not all same size). 

Filaments completely united 4. Paypayrola. 

Filaments united at the base only 5. Amphirrhox. 

Flowers irregular (zygomorphous) ; 1 petal spurred or saccate. 

Capsule woody 7. Corynostylis. 

Capsule not woody. 

Foremost petal saccate, not spurred 6. Hybanthus. 

Foremost petal spurred. 

Capsule wrapped in the remnants of calyx and corolla; 

half-shrub mostly unramified 10. Noisettia. 

Capsule not as above; habit variable. 
Capsule inflated; seeds winged or ridged (exception: 
A. Raimondi with half-spherical seeds) .8. Anchietea. 
Capsule 3-carinate, not inflated; seeds not winged, ovoid 

or spherical 9. Viola. 

Fruit nut-like . . , . 11. Leonia. 



58 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. RINOREAAubl. 

Reference: Blake, Contr. U. S. Nat. Herb. 20: 491-518. 1924. 

Shrubs or trees with apparently opposite leaves (Peruvian 
species) axillary or sometimes terminal racemes of small white or 
yellowish flowers (exception: R. micrantha). Flowers regular, with 
5 free petals and 3-valved capsules. Stamens entirely free, connec- 
tives dilated dorsally from base to apex into scarious brownish scales 
exceeding the anthers and free from them laterally and apically. 
Each filament (Peruvian species) is adnate outside at the base or 
throughout its length to a gland, which is sometimes longer than 
the filament and is (usually) free at apex. One species, R. paniculata 
(Mart.) Kuntze, 498, with alternate leaves, has been collected in 
the Amazonian region and may be expected. We have found the 
descriptions by Blake concise and complete and they have been 
largely used by us. Williams has noted the name "yurac varilla" 
for some species. 

Style in adult flowers straight or bent at summit; flowers about 

3 mm. long or longer. 

Ovary and style well-defined, the latter exactly straight; flowers 
pedicellate. 

Ovary glabrous or with hairs R. flavescens. 

Ovary densely pubescent. 
Anthers 1-2 cuspidate or 2-4-setose apically. 

Petals pubescent R. Passoura. 

Petals glabrous R. falcata. 

Anthers not appendaged apically. 

Leaves mostly about 5 cm. long; racemes 2-3 cm. long. 

R. juruana. 

Leaves mostly about 8 cm. long; racemes 3.5-6 cm. long. 

R. viridifolia. 

Ovary and style merging, the latter broad and bent to one side 
at tip; flowers subsessile or in cymules. 

Ovary villous; racemes spike-like R. macrocarpa. 

Ovary glabrous; panicles spike-like R. racemosa. 

Style (in adult flowers) sigmoid in form; flowers (Peruvian) scarcely 
2 mm. long (or longer, form) R. Lindeniana. 

Rinorea falcata (Mart.) Kuntze, Rev. Gen. PI. 1, pt. 1: 42. 
1891; 511. Alsodeia falcata Mart, ex Eichler in Mart. Fl. Bras. 13, 
pt. 1: 386. 1871. 



FLORA OF PERU 59 

Similar to R. Passoura; pedicels about 3 mm. long, reflexing; 
petals to 4 mm. long, glabrous; anthers bicuspidate; capsules 2 cm. 
long, glabrate, the seeds glabrous. The Peruvian collection, appar- 
ently referable here, has white flowers only 3.5 mm. long. 

Loreto: San Antonio on Rio Itaya, Killip & Smith 29417 (det. 
F. M.). Brazil. 

Rinorea flavescens (Aubl.) Kuntze, Rev. Gen. PI. 1, pt. 1: 42. 
1891; 507. Conohoria flavescens Aubl. PI. Guian. 1: 239. pi. 95. 1775. 

Shrub or slender tree to 7 meters; petioles 3-7 mm. long, pub- 
escent like the branchlets; blades obovate to oval, 7-20 cm. long, 
2.2-8 cm. broad, shortly acuminate, at base cuneate to rounded, 
obsoletely crenate-serrulate or subentire, glabrous or pubescent on 
the veins above and beneath; racemes about 7 cm. long, finely 
spreading, puberulous; pedicels 2-4 mm. long; sepals oval or ovate, 
rounded or obtuse; petals oblong-lanceolate to elliptic-ovate, yellow, 
4.2-4.5 mm. long, obtusish, with recurved and very sparsely hispid- 
ulous apex, otherwise glabrous; filaments equaling or exceeding the 
deltoid tridenticulate gland, this adnate for half its length or more, 
anthers not appendaged, connective scales oblong-ovate, obtuse; 
ovary most generally glabrous; capsule 2-2.5 cm. long, usually 2- 
seeded; seeds glabrous. Illustrated, Aubl. PI. Guian. pi. 95. 

Loreto: Yurimaguas (Poeppig, 2324 B); 2137. Mishuyacu near 
Iquitos, King 263; 693; 818; 843; 1362; Killip & Smith 29981. 
Brazil to Guianas. 

Rinorea juruana Ule, Verh. Bot. Ver. Brandenb. 47: 158. 
1905; 514. 

Shrub or tree, 3-10 meters high; blades obovate or oblong, 4-6 
cm. long, 2-2.5 cm. broad, acuminate, obtuse at apex, minutely sub- 
spinulose-serrate, short pubescent on the veins, especially above; 
racemes 2-3 cm. long, few-flowered, puberulous, the pedicels 3 mm. 
long; bracts lanceolate, acute; sepals broad-elliptic; petals oblong- 
obovate, whitish yellow, 4-5 mm. long; filaments strongly thickened, 
the connective scales ovate, entire, twice as long as the anthers; 
ovary densely hairy; capsule trigonous, 15 mm. long, acuminate, 
sparsely pilose; seeds solitary in each valve, glabrous, 6 mm. thick. 
R. scandens Ule, I.e. 157, has larger leaves, 10-20 cm. long. With- 
out locality, fide Melchior in Engler & Prantl, Pflanzenfam. ed. 2. 21 : 
352. 1925 (determinations, F. M.). F.M. Neg. 24062. 

Loreto: Santa Rosa, below Yurimaguas, Killip & Smith 8898; 
Williams 4865. Puerto Arturo, Williams 5210; 5088. Recreo, Wil- 



60 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Hams 4127. Alto Rio Itaya, Williams 3244. Brazil; Bolivia. "Virote 
quiro." 

Rinorea Lindeniana (Tul.) Kuntze, Rev. Gen. 1, pt. 1: 42. 
1891; 503. Alsodeia Lindeniana Tul. Ann. Sci. Nat. s^r. 3. 7: 364. 
1847. A. guianensis var. Lindeniana Eichl. in Mart. Fl. Bras. 13, 
pt. 1: 387. 1871. Rinorea micrantha Ule, Verh. Bot. Ver. Brandenb. 
47: 157. 1905; fide Melchior, Notizbl. Bot. Gart. Berlin 10: 546. 1929. 

Shrub or tree, 3-15 meters high; leaves elliptic-oblong, 13-15 
cm. long, 5-7 cm. broad, long-acuminate, usually slightly cordate at 
the oblique base, crenulate or serrulate, glabrous; panicles racemi- 
form, 9 cm. long, the axis puberulent to glabrate, the lower branch- 
lets about 4-flowered; sepals elliptic; petals greenish white (Killip) 
or yellowish (Mexia), ovate-elliptic; filaments bearing a subulate 
gland at base, the connective scales ovate, erose, nearly twice as 
long as the anthers; ovary densely pilose, style S-shaped in adult 
flowers only. The type is Ule 5477 from Rio Jurua, Bocca do Tejo, 
Brazil. F.M. Negs. 24064; 32167; 28286 (R. micrantha}. 

Loreto: Above Pongo de Manseriche, hills on left bank of Rio 
Maranon, in dense forest, alt. 250 meters, Mexia 6346. Florida, 
Tessmann 4630; Rio Putumayo at mouth of Rio Zubineta, forest, 
alt. 180 meters, King 2173. San Antonio, Killip & Smith 29298. 
Soledad, Killip & Smith 29630. San Antonio, Rio Itaya, Killip & 
Smith 29342; 29459. Near Iquitos, King 629. Rio Mazan, Schunke 
148. Rio Acre: Cobija, Ule 9621. Brazil; Bolivia. "Pantui" 
(Aquaruna), "limonsacha," "gopena-go ey" (Huitoto). 

Rinorea macrocarpa Kuntze, Rev. Gen. 1, pt. 1: 42. 1891; 
514. Alsodeia macrocarpa Mart, ex Eichl. in Mart. Fl. Bras. 13, 
pt. 1:385. pi. 78. 1871. 

Tree or shrub 3-9 meters high; leaves oval or obovate-oval, 10-20 
cm. long, 5-8 cm. broad, short-pointed, acute at base, membrana- 
ceous, repand-serrate, at first puberulous, quickly glabrate; racemes 
spiciform, 4-6 cm. long, yellowish-puberulous, the flowers subses- 
sile; sepals ovate-oblong; petals oblong-lanceolate, obtuse, glabrous, 
revolute at apex; stamens scarcely shorter than the petals, the fila- 
ments nearly as long as the anthers, apparently not gland-bearing, 
the dorsal connective scales scarcely once and a half as long as cells; 
ovary villous, merging with the style, the latter broad and bent to 
one side at tip; capsule acute, 4-5 cm. long, at length glabrate, the 
valves 2-seeded, the seeds glabrous. 

Loreto: Leticia, Ule 6192. Surinam; Amazonian Brazil. 



FLORA OF PERU 61 

Rinorea Passoura Kuntze, Rev. Gen. 1, pt. 1: 42. 1891; 507. 
Passoura guianensis Aubl. PI. Guian. 2: Suppl.: 21. pi. 380. 1775. 
Conohoria Passoura DC. Prodr. 1: 312. 1824, in part. Alsodeia 
guianensis Eichl. in Mart. Fl. Bras. 13, pt. 1: 387. 1871. 

Leaves oval to elliptic, 8-17.5 cm. long, 3-7 cm. broad, short- 
pointed to acuminate, at base rounded to cuneate, obscurely crenate- 
serrulate or subentire, pubescent; racemes 3.5 to 10 cm. long, densely 
ferrugineous-puberulous, the pedicels about 5 mm. long; sepals 
broadly ovate, obtusish, strongly several-ribbed; petals oval-oblong, 
white or yellow, obtuse, pilose along midline with erect rufidulous 
hairs; filaments equaling or exceeding the oblong gland, which is free 
above the middle, anthers 1- or 2-cuspidate or 2- to 4-setose at apex, 
connective scales oblong-ovate, obtuse, obscurely erose; ovary 
densely hispid-pilose, style straight; capsule 2 to 3 cm. long, tomen- 
tose, glabrate; seeds 3 to each placenta, velvety-tomentose. R. 
camptoneura (Radlk.) Melchior in Engl. & Prantl, Pflanzenfam. ed. 
2. 21: 352. 1925, Amazonian, has membranous appendices on the 
anthers. R. falcata (Mart.) Kuntze, which would be sought here, 
has glabrous petals. Shrub or tree to 8 meters; wood tough (Mexia) ; 
flowers cream and violet (Klug). 

Junin: Along Rio Perene", Killip & Smith 25154 (det. Standl.). 
San Martin : Juanjui, 400 meters, Klug 3802 (det. Standl.). Chazuta, 
Klug 4081 (det. Standl.). Loreto: Lower basin of Ucayali, Tessmann 
3441,' 3162. Fortaleza, near Yurimaguas, 140 meters, in forest, 
Klug 2787; Mexia 6082; Killip & Smith 28288; 27558; 28825 (only 
Klug spec. det. Baehni). Puerto Arturo, Killip & Smith 27785; 
27727 (det. Standl.). Soledad on Rio Itaya, Killip & Smith 29795 
(det. Standl.). Mishuyacu near Iquitos, Killip & Smith 29930. 
Rio Acre: Seringal Auristella, Ule 9620. Guianas; Amazonian Brazil. 
"Canella de Jacamin," "lluichovaran-dilla." 

Rinorea racemosa (Mart. & Zucc.) Kuntze, Rev. Gen. 1, pt. 1: 
42. 1891; 502. Alsodeia racemosa Mart. Nov. Gen. & Sp. 1: 29. 
pi. 20. 1823. 

Similar to R. macrocarpa, but leaves often merely repand- 
margined, the panicles raceme-like, usually 2 cm. long or longer, the 
cymules with several flowers; petals lightly puberulent, 2.5 mm. 
long or longer, well-exceeding the stamens; capsule about 8 mm. 
long. Flowers greenish yellow (Killip & Smith); orange (Klug). 
Determinations by Melchior. 

Huanuco: Puerto Lira, 300 meters, in dense forest along Rio 
Pachitea, Killip & Smith 26832. Loreto : Rio Itaya, Williams 61. 



62 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Upper Rio Nanay, Williams 8195. Mishuyacu near Iquitos, Klug 504. 
Rio Acre: Near mouth of Rio Macuahan, Krukoff 5391 . Brazil. 
"Majaras caspi" (Inca). 

Rinorea viridifolia Rusby, Mem. Torrey Club 6: 5. 1896; 514. 

Leaves obovate or oblong-obovate, 6-11.5 cm. long, 2-4.5 cm. 
broad, obtusely acuminate or short-pointed at base, cuneate or 
rounded-cuneate, crenate-serrulate, puberulous on main nerve above, 
sparsely strigose at base beneath or glabrous; racemes 3.5-6 cm. 
long (8 cm., Tessmann 5281), sparsely and obscurely puberulous; 
pedicels 2.5-3.5 mm. long; bracts scarious, broadly ovate, keeled; 
sepals oval-ovate, obtuse or acutish; petals elliptic, obtuse, slightly 
reflexed, ciliolate at base and apex; filaments stout, somewhat 
exceeding the emarginate sparsely puberulous gland, nearly as long 
as anther cells, the anthers unappendaged, the connective scales 
oblong-ovate, obtuse, distinctly erose especially below the middle; 
ovary densely hispid-pilose, style straight. R. brevipes Blake, 512 
(R. laxiflora Melchior in Engl. & Prantl, Pflanzenfam. ed. 2. 21: 
352. 1925, Alsodeia brevipes Benth. in Hook. Journ. Bot. 4: 106. 
1842, A. laxiflora Benth. I.e. 107, syn. after Blake) has also nearly 
glabrous petals, the filaments less than one-half as long as the 
anther cells (R. viridifolia filaments as long as anther cells). 

Loreto: Ucayali, Tessmann 3199. La Victoria, Williams 2961; 
2712. Basin of Maranon between Iquitos and mouth of Santiago 
at Pongo de Manseriche, Tessmann 5281. Rio Acre: Cobija, Ule 
9619; Seringal San Francisco, Ule 9613. Bolivia. 

2. RINOREOCARPUS Ducke 

Reference: Ducke, Archiv. Jard. Bot. Rio Jan. 4: 144. 1925. 

Genus related to Rinorea, of which it has the capsules, the free 
petals and the free stamens. However, it can be recognized at once 
owing to the dichasial cymes and its anthers without a dorsal scale 
produced into a short terminal appendix. 

Rinoreocarpus Ulei (Melchior) Ducke, Archiv. Jard. Bot. Rio 
Jan. 5: 173. 1930. R. salmoneus Ducke, I.e. 4: 144. pi. 13. 1925. 
Gloeospermum Ulei Melchior, Notizbl. Bot. Gart. Berlin 9: 164. 1924. 

Shrub or tree to 12 meters high, the young parts covered with an 
evanescent velvety pubescence; leaves obovate, almost glabrous, 
10-16 cm. long, 3-7 cm. broad, shortly acuminate, remotely and 
irregularly serrate, slightly shining above; stipules early caducous; 
flowers salmon-colored, to yellow or orange, about 5 mm. long, almost 



FLORA OF PERU 63 

sessile, with a very short staminal tube (one-fourth of the filament), 
the edge of which is straight; appendix of connective short, foremost 
petal alone plicate; style straight, about twice as long as the glabrous 
ovary; capsules 2-3 cm. long, early orange-yellow later black, rather 
obsoletely transverse-reticulately rugose, the seeds obovate, 6 mm. 
long, lustrous, minutely black-punctate. Cf. Williams, Field Mus. 
Bot. 15: 348. 1936, under Gloeospermum Sprucei Eichl. F.M. Neg. 
24056. 

San Martin: Pongo de Cainarachi, Klug 2727. Loreto: Pebas, 
Williams 1935. Iquitos, Williams 3638. Yurimaguas, Killip & 
Smith 29058. Florida, Klug 2287. Brazil. 

3. GLOEOSPERMUM Trian. & Planch. 
Reference: Melchior, Notizbl. Bot. Gart. Berlin 9: 163-167. 1924. 

Trees and shrubs with distichous, short-petioled and slightly 
toothed leaves. Flowers 2 or 3 at a time, regular, small, greenish 
or reddish, the scorpioid cymes axillary. Filaments united at the 
base with a staminal tube which is in some species as long, in some 
others half as long as the stamens. Tip of connective produced into 
a long membranous appendix sometimes as long as the anthers. 
Fruit a berry. 

Leaves glabrous beneath .G. sphaerocarpum. 

Leaves densely pilose G. pilosum. 

Gloeospermum pilosum Melchior, Notizbl. Bot. Gart. Berlin 
8: 623. 1923. G. peruviana Melchior, nomen nudum in Engl. & 
Prantl, Pflanzenfam. ed. 2. 21: 354. 1925. Achras globosa R. & P. 
Fl. Peruv. 4: pi 425. 1802, fide Melchior. 

Very similar to G. sphaerocarpum, from which it is readily distin- 
guished by the short and dense pubescence of the branchlets, under- 
surface of leaves and sepals. After Melchior, I.e. 9: 164. 1924, G. 
pilosum can be readily distinguished from G. sphaerocarpum by the 
staminal tube which almost equals the filament in the former, and 
is only one-half or three-fourths as long in the latter. From observa- 
tions made in Herb. Delessert, we are inclined to doubt whether 
these proportions hold true; it would seem then that, the pubescence 
being the only certain character to be relied upon, G. pilosum could 
be considered as a mere var. of G. sphaerocarpum. The plate with 
this number in some copies of R. & P., vol. 4 is not this plant. F.M. 
Neg. 24055. 



64 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Uchiza (Tafalla in 1798, type); in Herb. Delessert 
as "Achras globosa sp. n. M. Pavon." 

Gloeospermum sphaerocarpum Triana & Planch. Ann. Sci. 
Nat. se>. 4. 17: 129. 1862; 167. 

Shrub, perhaps tree, entirely glabrous; leaves lanceolate to 
oblong-lanceolate, shortly acuminate at the tip or acute at both 
ends, remotely serrate at the margin, 4-12 cm. long, 1-4 cm. broad, 
papyraceous, paler and conspicuously veined beneath; stipules very 
early caducous (only to be seen in the bud), linear-subulate, almost 
as long as the petiole (0.5 cm.); inflorescences only 0.5 cm. long; 
flowers small (3.5 mm. long), on short pedicels (1-2 mm.); free part 
of the petals short, spreading; tube of corolla thick; stamens about 
as long as the calyx, the margin of the staminal tube irregular; 
appendix of the stamens nearly as long as the lobes, membranaceous; 
ovary attenuated into a style 3 times as long as the stamens; stigma 
punctiform (description after Eichler). The var. latifolium Melch. 
I.e. 8: 622, has broader leaves, 3-5 cm. wide, 5-13 cm. long; the 
flowers are the same. F.M. Negs. 24058; 24057. Illustrated, Mart. 
Fl. Bras. 13, pt. 1: pi. 79. 

Loreto: Pongo de Manseriche, Tessmann 4730. Maynas, woods 
near Yurimaguas, Poeppig 2088 (type of var.); Williams 5256; 4955 
(det. Melchior); 5086. Near Iquitos, Williams 8167 (det. Melchior). 
La Victoria, Williams 3007? (fruit "only). Puerto Arturo, Williams 
5302? (fruit only). Brazil; Colombia. 

4. PAYPAYROLA Aubl. 

Flowers only slightly zygomorphic. Claws of the petals wide, 
cohering into a long tube. Filaments united. Appendage of anthers 
very small. 

Paypayrola grandiflora Tul. Ann. Sci. Nat. se>. 3. 7: 371. 1847. 

Tree 5-7 meters high ; leaves subopposite, borne at the end of the 
twigs, entire, 20-35 cm. long, 7-13 cm. broad; spikes terminal or 
axillary, the few yellow flowers themselves being sessile or shortly 
pedunculate; petals thick, forming an urn-shaped corolla, sometimes 
2 cm. long; capsule shortly acuminate, 4 cm. long, glabrous. 
Determinations (except as noted) by Melchior. Illustrated, Mart. 
Fl. Bras. 13, pt. 1: pi. 76. 

San Martin: Pongo de Cainarachi, King 2717 (det. Standl.). 
Loreto: Pampa del Sacramento, (Huber 1533). Leticia, Ule 6219. 
Basin of Maranon, between Iquitos and mouth of Santiago at Pongo 



FLORA OF PERU 65 

de Manseriche, Tessmann 5543. Iquitos, Williams 8016; Klug 46; 
129; 526; Killip & Smith 27488. Lower Rio Huallaga, Williams 
3970; 4130; 4265; 4847. Santa Rosa, Killip & Smith 28764. La 
Victoria, Williams 2973. Caballo-Cocha, Williams 2012; 2079; 2359. 
Brazil. "Caballo-cocha," "tornilla muena," "pegan-pegan." 

5. AMPHIRRHOX Sprengel 

With the character of Paypayrola but the filaments free except 
at base where united into a low ring and the anther appendage 
elongate. 

Capsules to 3 cm. long, seeds 5-7 mm. long; leaves often obviously 
repand-denticulate. 

Calyx teeth with a mucro below tip A. juruana. 

Calyx teeth without mucro A. latifolia. 

Capsules to 4 cm. long; seeds 10 mm. long or longer; leaves often 
entire or merely repand. 

Leaves broadly obovate A. Sprucei. 

Leaves nearly oblong-subelliptic or narrowly obovate. 

A. surinamensis. 

Amphirrhox juruana Ule, Verh. Bot. Ver. Brandenb. 47: 
156. 1905. 

Shrub to 3 meters; leaves dark green above, 12-18 cm. long, 4-5 
cm. broad, oblong-lanceolate, cuspidate, remotely serrate; flowers 
yellowish-purple, about 1.5 cm. long, slightly zygomorphous in fas- 
cicles at the tip of branchlets, on long pedicels (1-4 cm.); sepals with 
a short point (mucro) on the back; petals free, yellowish-lilac, unguic- 
ulate; filaments united below; connective with appendage; style 
straight. Capsule 3.5 cm. long, half as broad, the light brown seeds 
7 mm. long (Krukoff spec.). F.M. Neg. 24066. 

Rio Acre: Upper Rio Jurupary, Krukoff 5229 ex char. Brazil. 

Amphirrhox latifolia Mart, ex Eichler in Mart. Fl. Bras. 13, 
pt. 1:376. 1871. 

Tree, 15 meters tall or taller, with shortly petioled oblong-elliptic 
leaves acute at both ends or shortly acuminate, firm, membranous, 
veiny, concolor, obscurely or remotely repand-serrulate, mostly 
12-15 cm. long, 5-7 cm. broad, or on young shoots 2 dm. long, half 
as wide; flowers 2-several fasciculate and as in Amphirrhox juruana, 
but roseate or white; calyx 3-4 mm. long, the sepals broadly ovate; 



66 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

capsules about 2.5 cm. long, tuberculate; the obovoid lustrous seeds 
6-7 mm. long. 

Rio Acre: Upper Rio Macauhan, Krukoff 5527. Guianas; 
Brazil. 

Amphirrhox Sprucei (Eichler) Macbr. comb. nov. Gloeosper- 
mum Sprucei Eichler in Mart. Fl. Bras. 13, pt. 1: 390. 1871. 

Leaves obovate, narrowed to the petiole (this about 1 cm. long), 
subacutely and shortly pointed, reticulate-veiny, obsoletely repand 
or denticulate, slightly paler beneath, chartaceo-coriaceous, often 
15 cm. long, half as wide; flowers apparently unknown; capsules 
ovoid-globose, irregularly and bluntly tuberculate, apiculate, at least 
3 cm. in diameter, the light brown, smooth, subovoid seeds with 
yellowish testa, about 1.5 cm. long. Melchior in herb, and in his 
revision ofGloeospermum referred this to Amphirrhox with (!). As I 
know nothing about the relationship, I hesitate to make him respon- 
sible for the transfer which, however, is necessary here because 
Williams 3638 was at one time determined by Melchior as this species; 
cf. Rinoreocarpus (F.M.). Illustrated, I.e. pi. 79. F.M. Neg. 19148. 

Peru? Brazil. 

Amphirrhox surinamensis (Miq.) Eichler in Mart. Fl. Bras. 
13, pt. 1:377. 1871. 

Leaves characteristically oblong-lanceolate, slightly obovate, 
rarely elliptic-membranous or subcoriaceous; flowers about as in 
A. latifolia, 1.5 cm. long, creamy-white or bluish; capsules smooth, 
subglobose, about 4 cm. in diameter; seeds 16-18 mm. long, similar 
to those of A. Sprucei. Miquel gave the name in herb, under another 
genus. 

Loreto: Pebas, Williams 1719 (det. Melchior). Mishuyacu near 
Iquitos, Klug 1360. Venezuela; Guianas. 

6. HYBANTHUS Jacq. 

Reference: G. K. Schulze, Bot. Jahrb. 67: 437-489. 1936. 

Herbs, half-shrubs or rarely shrubs, the leaves alternate or the 
lower sometimes opposite. Flowers either solitary and axillary or 
in racemes, generally white or blue, the sepals free. Of the 5 free 
petals, one is much longer than the others and unguiculate. Stamens 
free. Connectives dorsally dilated from base to apex into scarious 
brownish scales exceeding the anthers and free from them laterally. 
Fruit a 3-valved capsule. 



FLORA OF PERU 67 

Flowers racemose; shrub H. Sprucei. 

Flowers solitary; herbs or half-shrubs. 
Stipules conspicuous, white scarious; leaves pubescent. 

Leaves obovate, rounded at tip H. tarapotinus. 

Leaves elliptic-lanceolate, acute H. albus. 

Stipules minute or inconspicuous; leaves glabrescent. 

Flowers 7-12 mm. long; annual herb H. attenuatus. 

Flowers 3-4 mm. long; half-shrub H. parviflorus. 

Hybanthus albus (St. Hil.) Baill. Bot. Me'dic. Phan. 2: 841. 
1884; 475. lonidium album St. Hil. Me"m. Mus. Hist. Nat. Paris 
11:477. 1824. 

Shrubby, the branches above and the leaves rather densely 
villous; stipules white-scarious in age, the younger scarious-margined; 
leaves alternate, 2.5-4 cm. long (or larger), more or less closely 
serrate; pedicels nearly filiform, 1.5-3 cm. long, mostly or all with- 
out bracts; calyx teeth subequal, entire foremost petal about 1.5 
cm. long, the lateral somewhat, the others much shorter; filaments 
short, the two upper with a short spur; connective scale orange- 
brown; style more or less curved, weakly 2-lobed. The Peruvian 
plant has rather more densely-serrate leaves and the lateral petals 
have not been seen; the determination therefore is not certain 
(F.M.). 

Cuzco: At 650 meters, C. Bues. Southern Brazil; Uruguay; 
Argentina. 

Hybanthus attenuatus G. K. Schulze, Notizbl. Bot. Gart. 
Berlin 12: 114. 1934; 481. lonidium attenuatum Humb. & Bonpl. 
in Roem. & Schult. Syst. 5: 402. 1819. 

Annual herb with four-cornered stems; leaves glabrous, elliptic 
to oblong-lanceolate, 3-7 cm. long, 1-1:5 cm. broad, the margin 
denticulate or remotely serrate or even entire, usually only the lower 
ones opposite; chasmogamic flowers solitary, axillary, about 1 cm. 
long; cleistogamic usually present, smaller; foremost petal clawed, 
twice as long as the others; ovules 3-6. Some of the Peru collec- 
tions were distributed as H. parietariifolius (DC.) Loes. considered 
by Schulze, I.e. 481, as probably a synonym. 

Junin: Chanchamayo, Hacienda de la Esperanza (Raimondi 
8649). Palca Chanchamayo (Raimondi 2773). Montana de Chan- 
chamayo (Raimondi 2435). Chanchamayo Valley (Weberbauer 
1959). Cajamarca: Jae"n (Raimondi 11494). Tumbez: Ricaplaya, 



68 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

100 meters, flowers white, violet spot on petal below, Weberbauer 
7735. Huanuco: Mito, 3339 (det. Johnst.). San Martin: Near 
Moyobamba, King 3483 (det. Standl.). Distributed through 
Central America, from Mexico to Ecuador. 

Hybanthus parviflorus (Mutis) Baill. Bot. Medic. Phan. 2: 
841. 1884; 482. Viola parviflora Mutis in Linn. f. Suppl. 396. 1781. 
lonidium glutinosum Vent. Jard. Malm. 1: no. 27. 1803. 

Half-shrub or herbaceous above the caudex, very variable in 
habit and leaf-form; lower leaves alone or all of them opposite, ovate- 
elongate or obovate or lanceolate, 0.5-3.5 cm. long, generally 
glabrous; flowers at the end of ramifications, forming themselves 
into more or less definite racemes; 2 foremost filaments with an 
upward directed spur at the base; style in S-form. Said to be 
emetic (Baillon). 

Huanuco: Mito, prostrate in old field, flowers light green, 1464; 
shrubby slope, flowers white, 3337 (both det. Johnst.). Cajamarca: 
Camino Magdalena (Raimondi 7150). Cutervo (Raimondi 3032; 
3043; 5708; 7381; 7403) (Jelski 236). South America except in the 
Guianas. "Pachagapichinya." 

Hybanthus Sprucei (Eichler) Hassl. Bull. Soc. Bot. Geneve 
se>. 2. 1: 214. 1909; 456. lonidium Sprucei Eichler in Mart. Fl. 
Bras. 13, pt. 1: 373. 1871. 

Small shrub; leaves oblong-lanceolate, acute or rather acuminate, 
5-8 cm. long, 2-2.5 cm. broad, with a shallowly toothed or entire 
margin, generally quite glabrous; flowers sometimes solitary, more 
generally in axillary panicles composed of opposed pairs of flowers 
with small membranous bracts at the base of peduncles; foremost 
petal 3.5-4 mm. long, the base of the claw being as wide as the 
spatula; other petals a trifle shorter, ovate, dully pointed. F.M. 
Neg. 24013. 

San Martin: Tarapoto, Spruce 3920, type; Ule 6590; Williams 
6899. Brazil. 

Hybanthus tarapotinus Ule, Verh. Bot. Ver. Brandenb. 47: 
158. 1905; 476. 

Easy to recognize because of its white and scarious stipules, 
short internodes and villous leaves; branchlets rather pilose; leaves 
elliptic to obovate, blunt or rounded at tip, 4-6 cm. long, 1.5-2.5 cm. 
broad, denticulate, sparingly above and densely pilose beneath; 
flowers nearly 1 cm. long, solitary on slender pedicels 2-2.5 cm. long; 
no bracts; stigma slightly bifid. F.M. Neg. 24014. 



FLORA OF PERU 69 

San Martin: Tarapoto near Juan Guerra, Ule 6524, type; 
Spruce 4205. Moyobamba, San Miguel between Tabalosos and 
Lamas (Raimondi 1955}. Juanjui, Klug 4389. Chazula, Klug 
3962 (det. Standl.). Cajamarca: Jae"n, Perico (Raimondi 397}. 

7. CORYNOSTYLIS Mart. & Zucc. 

Flowers zygomorph; 2 ulterior petals small, 2 middle ones larger, 
oblique, and a fifth one with a long cylindrical spur. Filaments 
flat, united in pairs, the 2 foremost ones bearing in common one 
villous spur. Anthers with one enlarged connective. Style slightly 
curved. Capsule big, woody. Seeds almost quadrangular. Roots 
used as an emetic. 

Corynostylis arborea (L.) Blake, Contr. U. S. Nat. Herb. 23: 
837. 1923. Viola arborea L. Syst. Nat. ed. 10: 1239. 1759. 

Climbing shrub with alternate oval to elliptic-ovate leaves, 5-9 
cm. long, 3-5 cm. broad, irregularly crenulate, acute or acuminate, 
glabrous; racemes terminal, leafy-bracted ; flowers whitish, very 
fragrant, 2.5-4 cm. long, lip with strongly dilated tip; spur longi- 
tudinally half-twisted. Illustrated, Mart. Fl. Bras. 13, pt. 1: pi. 69. 

Loreto: Creek Itaya, Mexia 6480. Maranon, between Iquitos 
and mouth of the Santiago, at Pongo de Manseriche, Tessmann 
3570; 5123. Florida, Rio Putumayo, at mouth of Rio Zubineta, 
Klug 2097; 2199; 2273. Pebas, Williams 1848. Near Iquitos, Klug 
2518; 1060. Lower Rio Nanay, Williams 528. North to Mexico, 
the Guianas and St. Vincent. "Chinaca-o" or "doira-o," "ampi yacu" 
(Huitoto). 

8. ANCHIETEA St. Hil. 

Reference: Hallier in Meded. Rijks Herb. 19: 64-66. 1913. 

Vines or climbing shrubs with alternate leaves, zygomorph 
flowers in axillary fascicles or sometimes alone, small, whitish, on 
articulated peduncles. Foremost petal big, with a long cylindrical 
spur and a broad asymmetrical limb. Capsule inflated, 3-valved. 
One species (A. salutaria) is said in Brazil to be depurative (St. Hil.). 

Seeds distinctly winged. 

Wing broad, entire or crenate A. parvifolia. 

Wing narrow, more or less cleft A. Selloana. 

Seeds not winged, sometimes with an edge. 

Seeds semi-globular, without edge A. Raimondii. 

Seeds deltoid, with a narrow, thick, more or less entire edge. 

A. peruviana. 



70 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Anchietea parvifolia Hallier, Meded. Rijks Herb. 19: 64. 1913. 

Branches twining; leaves small, 5-8.5 cm. long, 13 mm. broad, 
ovate, crenate-serrulate at the margin; flowers solitary on slender 
pedicels; spur straight, ascending; seeds winged, crenate at the 
margin, orbicular. The type is Herzog 1040 from Bolivia. 

Peru (probably). Bolivia. 

Anchietea peruviana Melchior, Notizbl. Bot. Gart. Berlin 9: 
167. 1924. 

Leaves alternate, ovate or elliptic-orbiculate, acute or acuminate, 
crenate-serrate, glabrescent, 3.5-6 cm. long, 2-3.5 cm. broad; flowers 
yellowish, solitary or fasciculate; spur cylindrical; seeds deltoid, 
sinuate at the margin. 

Lambayeque: Slope above Olmos (Weberbauer 7100). 

Anchietea Raimondii Melchior, Notizbl. Bot. Gart. Berlin 10: 
728. 1929. 

Leaves rather thick, ovate to elliptic, obtuse or acuminate at 
the tip, crenate-serrate, glabrous, 4-7 cm. long, 2-4.5 cm. broad; 
floral pedicels 1 cm. long; flowers small; spur slightly recurved; seeds 
semi-globular, not winged. 

Cajamarca: Prov. Cutervo, Tambillo (Raimondi 3344, type; 
3484; 3538; 3804; 4853). 

Anchietea Selloviana Cham. & Schl. Linnaea 2: 148. 1827; 66. 

Leaves ovate-oblong or even lanceolate, acute or acuminate, 
remotely crenate-dentate or subentire, entirely glabrous, 4-7 cm. 
long, 2.5-4 cm. broad; spur of flower longer than the limb, slender 
and straight; seeds orbicular, 6-7 mm. in diameter, with a very 
narrow and irregularly cleft wing. 

Cajamarca: Tambillo (Jelski 237; det. Szyszy.). Brazil. 

9. VIOLA Tourn. 

Reference: W. Becker, Beih. Bot. Centralbl. 22, pt. 2: 78-96. 
1907. 

Herbs or half shrubs. Leaves of variable shape, alternate or 
disposed in rosettes. Flowers axillary, always solitary, with 2 bracts 
on the petiole, zygomorphous, spurred. Filaments free, extremely 
short; the 2 foremost stamens bear on the back of the connective 
warts or spurs, or appendages. Capsule opening into 3 valves, 
keeled dorsally. Seeds generally smooth. 



FLORA OF PERU 71 

V. capillaris Pers., reported on the basis of a mislabeled Dombey 
specimen, is unknown from Peru. V. odorata L. has been collected 
by Williams, cultivated at Iquitos, the "Violeta." A common name 
for some native species is "Pensamiento." 

Leaves disposed in rosettes. 
Annual plants; roots thin. 

Sepals striate, never trinerved V. micranthella. 

Sepals trinerved V. Weberbaueri. 

Perennial; roots thick. 

Leaf-tip folded back .V. replicata. 

Leaf-tip not folded back. 
Leaves crenate or dentate. 
Leaves oblong or spatulate, obtuse or rounded at tip. 

Margin membranaceous, 2 or 3 teeth only. 

V. membranacea. 

Margin tomentose V. granulosa. 

Leaves linear, acute. 

Margins 4-5, repand crenate V. Hillii. 

Margins ciliate-serrate V. Weibelii. 

Leaves with margin entire or nearly so. 
Leaves linear. 

Stipules present V. pygmaea. 

Stipules absent V. pusillima. 

Leaves not linear. 
Leaves oblong, 2-4 mm. broad. 

Stipules adnate, 1 cm. long V. pallascaensis. 

Stipules none. V, nivalis. 

Leaves spathulate. 

Stipules present, adnate V. nobilis. 

Stipules wanting V. Kermesina. 

Leaves alternate (not merely in rosettes). 
Leaves denticulate or crenate. 
Leaves crenate. 

Leaves 10-20 mm. long, crenate from base to tip. 

y. Lehmannii. 
Leaves 3-4 mm. long, crenate at tip only y. Weddellii. 



72 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves dentate. 

Flowers red, long-spurred V. arguta. 

Flowers whitish, purplish, short-spurred or red-striped. 
Spur strangled (transversely sulcate) at its insertion. 

V. saccata. 
Spur not strangled. 

Leaves rhomboid-ovate, abruptly attenuate at base. 

V. huanucoensis. 

Leaves ovate, subcordate at base. 
Leaf-blades as broad as, or broader than long. 

V. Mandonii. 

Leaf -blades longer than broad. 
Petiole longer than a third of blade's length. 

V. Dombeyana. 
Petiole shorter than a third of blade's length. 

V. Humboldtii. 

Leaf margin entire or glandulo-serrate. 
Leaves acute or acuminate. 
Leaf-base attenuate. 

Petiole much shorter than blade V. stipularis. 

Petiole as long as blade V. fuscifolia. 

Leaf -base asymmetric, cordate. 

Spur's length one-tenth of inferior petal V . Steinbachii. 

Spur's length one-half of inferior petal. 

V. arguta ssp. meridionalis. 
Leaves obtuse, subobtuse and mucronulate at tip.V. Bangiana. 

Viola arguta HBK. Nov. Gen. & Sp. 5: 373. 1823; 92. V. 
corchorifolia Domb. ex DC. Prodr. 1: 304. 1824. 

Twining shrub; leaves denticulate, ovate, acute at tip, cordate 
at base, pilose beneath, mostly glabrous above, 15-45 mm. long, 
7-22 mm. broad, shortly petioled; stipules brown and membranous, 
long ciliate; flowers large, borne on petioles as long as or longer than 
the leaves; petals red, the inferior one carinate; spur obtuse, broad, 
as long as the remaining part of petal; stamens sessile; appendage 
of foremost stamens short and broad, or wanting (Becker); style 
subulate. Var. glaberrima W. Becker, I.e. 92, is like the type, 
but entirely glabrous. The subsp. meridionalis W. Becker, I.e. 
93, is similarly glabrous, but with leaves only indistinctly serrate 



FLORA OF PERU 73 

instead of denticulate. F.M. Negs. 24018; 34830; 34832; 32174 (V. 
corchorifolia) . 

Huanuco: Pavon s.n.; southwest of Panao, alt. 3,300 meters, 
2207. Chinchao, Poeppig (det. Becker). Mito, supported in shrubs, 
the flowers scarlet, 1710. Cajamarca: Near San Pablo (Weberbauer 
3882, the var. glaberrima). Without locality: Pavon 367; Pavon 
s.n. (1827) under V. coccinea; Mathews 1345. Colombia; Ecuador. 

Viola Bangiana W. Becker, Beih. Bot. Centralbl. 22, pt. 2: 
89. 1907. 

Trailing, rhizomatous, the erect parts reaching 10 cm. (Becker), 
much shorter on examined specimens; leaves ovate, subobtuse and 
mucronate at tip, attenuate at base or sometimes subcordate, in- 
distinctly glandulose-serrate, glabrous, 8-22 mm. long and broad; 
petiole one- third of blade's length; stipules lanceolate, brown, with 
long marginal hairs; pedicels longer than the leaves (up to 5.5 cm.); 
petals white, blue-striate, obovate, the inferior one subacuminate; 
appendages of foremost anthers long; ovary ellipsoid; style subulate. 
Cf. V. Dombeyana. 

Puno: SanGaban,Lecfeter 2241, type. Cuzco: Cerro de Cusilluyoc, 
Pennell 13958. Bolivia. 

Viola Dombeyana DC. Prodr. 1: 305. 1824; 90. 

Creeping rhizome bearing stems of 3-10 cm. ; leaves ovate, dentic- 
ulate-crenate, 8-12 mm. long, 7-8 mm. broad, attenuate or subcor- 
date, often blunt; petiole always longer than a third of blade's length; 
pedicels longer than the leaves; petals probably whitish or purplish- 
white, the upper and lateral ones oblong, the inferior carinate, acutish, 
very shortly spurred; connectives of the two foremost stamens pro- 
duced into appendages; capsule globose; style subulate. Species 
closely related to V. Humboldtii (and several other segregates as 
noted under the descriptions). The leaves are here smaller and the 
denticulation less apparent, the petiole is also relatively longer, 
compared with the blade, as in V. Humboldtii. Since there exist 
linking forms between these two species (cf. Jameson 698} it may 
be that V. Humboldtii is a mere variety. 

V. scandens Willd., found as near as Ecuador and apparently in 
Bolivia, has smaller flowers rarely exceeding the leaves and remotely 
dentate-ciliate stipules. F.M. Negs. 24027; 34837. 

Junin: Huassahuassi (Dombey 1785). East of Palca (Weberbauer 
2489). Between Yapas and Enenas, Killip & Smith 25614- Piura: 
East of Huancabamba, 3,400 meters, Weberbauer 6144 (so distrib- 



74 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

uted), petals light violet with dark violet veins. Ayacucho: Choi- 
mocota Valley, petals with veins violet, 2,600 meters, Weberbauer 
7566. Without locality: Pavdn 362, type. 

Viola fuscifolia W. Becker, Beih. Bot. Centralbl. 22, pt. 2: 94. 
1907. 

Rhizome creeping; erect shoot 10-20 cm. long, glabrous, densely 
covered with leaves; leaves ovate, asymmetric, acutish, glandulose- 
serrate, glabrous, thick, subabruptly attenuate into a petiole nearly 
as long as the blade, stipules brown, lanceolate, long-ciliate; pedicels 
longer than the leaves; flowers small, bluish, inferior petal obovate, 
subemarginate, spur extremely short; 2 foremost stamens bearing a 
long dorsal appendage; style subulate. Cf. V. Dombeyana (F.M.). 

Puno: Trail of Sandia to Chunchusmayo, between Tambo Ichu- 
bamba and Tambo Yuncacoya, alt. 1,800-2,600 meters (Weberbauer 
1096}. Prov. Carabaya, on shaded slopes, near San Juan del Oro 
(Weddell s. n., 1847). 

Viola granulosa Wedd. Ann. Sci. Nat. se>. 5. 1: 292. 1864. 

Undoubtedly related to V. membranacea; the leaves are small, 
spathulate, obtuse, finely crenate, tomentose below and at the mar- 
gin, glabrous above, ciliate on the edges of the petiole, with pedicels 
about as long as the leaves; flowers small, with linear sepals; superior 
and lateral petals lanceolate, inferior obovate, emarginate, without 
spur; style S-shaped, bearing at its upper part wing-like appendages. 
-Illustrated, Weddell, Chlor. And. 2: pi. 87 A. F.M. Neg. 38482. 

CuzcoiGay. 

Viola Hillii Becker, Kew Bull. 134. 1928. 

Rhizome vertical, woody, to 10 cm. long, 5 mm. thick, the upper 
portion shortly 2-4-parted and clothed with remains of old leaves; 
rosulae formed of the erect linear-oblong leaves only to 1.5 cm. 
broad; leaves about 8 mm. long, the blade half as long, subfleshy, 
lightly pilose, obviously 4-5 repand-crenate, about 1 mm. wide; 
stipules rudimentary; flowers yellow, scarcely 5 mm. long, the 
peduncle bibracteolate basally, retrorsely hispid, to 6 mm. long; 
sepals oblong, acutish, pilose; upper and lateral petals oblong, the 
lower broad, shortly spurred ; style distinctly geniculate, horizontally 
rostrate, with two rotund lobes on each side at tip. Section Andi- 
nium; remote from all species (Becker). Illustrated, Hook. Icon. 32: 
pi. SI 49. 

Peru: Boundary at Lake Titicaca on red sandstone hills between 
Moho and Vilque Chico, 4,050-4,200 meters, A. W. Hitt 28. 



FLORA OF PERU 75 

Viola huanucoensis W. Becker, Repert. Sp. Nov. 18: 186. 1922. 
y. truncata W. Becker, I.e. 94. 

Different from V. Humboldtii, by the shape of the leaves, which 
are here rhomboid-ovate; petals whitish, the superior and lateral 
oblong, inferior obovate, more or less triangular, plicate, emarginate, 
with very short spur; foremost anthers cuspidate; capsule ellipsoid; 
style subulate. Cf. V. Dombeyana (F.M.). 

Huanuco: Northwest of Monzon (Weberbauer 3715}. 

Viola Humboldtii Triana & Planch. Prodr. Fl. Nov. Granat. 
121. 1862; 90. V. stipularis HBK. Nov. Gen. & Sp. 5: 372. 1823. 
y. capillaris Ging. in DC. Prodr. 1: 304. 1824. 

Thin rhizome prostrate, ramified; erect stalks up to 20 cm. long; 
leaves ovate, densely and sharply serrate, asymmetrically subcordate 
at base, 7-22 mm. long, 6-14 mm. broad; petiole shorter than a third 
of blade's length; stipules brown, membranous, ciliate; flowers 
similar to those of y. Dombeyana (to which it probably goes as a 
var. F.M.). There exists a var. cuneata W. Becker, 91, which is 
said to have cuneate leaves. An examination of a co-type did not 
show any marked difference from the main species. F.M. Neg. 24033. 

Cajamarca: San Miguel, alt. 3,000 meters, Weberbauer 391+7. 
Hualgayoc, alt. 3,100-3,300 meters, Weberbauer ^020 (co-type of 
var.). Colombia; Ecuador. 

Viola Kermesina W. Becker, Repert. Nov. Sp. 7: 124. 1909. 

Species with the same habit as y. nivalis Benth. but the leaves 
here are spathulate, rounded to truncate at tip, ciliate on both faces 
and at margin, attenuate, without stipules; pedicels longer than the 
leaves; sepals broadly ovate, somewhat obtuse; petals scarlet, the 
upper and lateral one oblong-obovate, inferior one carinate, obcordate 
emarginate; spur rather thick; ovary slightly pilose; style S-shaped, 
dilated into two lobes at tip. 

Junin: Near Yauli, along the railroad tracks, between Lima and 
Oroya, alt. 4,600 meters (Weberbauer 330}. 

Viola Lehmannii Becker, subsp. ovalifolia W. Becker, Beih. 
Bot. Centralbl. 22, pt. 2: 91. 1907. 

Resembles y. Humboldtii: same habit and same flowers; leaves 
ovate, crenate, glabrous, 8-20 mm. long, 7-15 mm. broad, rather 
abruptly contracted into a short petiole; stipules as in V. Humboldtii. 
The subsp. cordifolia W. Becker, I.e. 92, has leaves crenate, sub- 
obtuse, cordate; petiole as long as or longer than blade's length. 



76 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Probably not specifically distinct from V. Dombeyana (F.M.). F.M. 
Neg. 24035. 

Amazonas: Between Pucasmajo and Moyobamba, alt. 3,100 
meters (Stuebel 26d). Molinopampa, Osgood & Anderson 43. With- 
out locality, Mathews 3105 (type, subsp. cordifolia). 

Viola Mandonii W. Becker, Beih. Bot. Centralbl. 22, pt. 2: 
95. 1907. 

Species belonging to the complex of V. Humboldtii; the leaves 
here are broadly ovate, often as large as long (10-22 mm.), sub- 
cordate, with spaced sharp teeth; petiole longer than a third of limb's 
length (up to 5 cm.); petals whitish, superior and lateral ones oblong, 
inferior elliptic-obovate, acute or even acuminate at tip; spur short; 
style subulate (cf. V. Dombeyana, F.M.). The type is Mandon 943 
from Bolivia. F.M. Neg. 24036. 

Peru (probably). Bolivia. 

Viola membranacea W. Becker, Repert. Nov. Sp. 7: 123. 1909. 

Related to V. nivalis, showing the same leaves disposed in rosettes 
and the same spindle-like and thick rhizome; leaves oblong, attenuate 
at base, with 3-4 deep and wide crenatures near the tip and a sub- 
membranous edge, 10-20 mm. long, 1.5-4 mm. broad; stipules mem- 
branous, adnate for the most part, ending in a long acumen; flowers 
small, whitish; superior petal erect, larger than the others, emargi- 
nate; inferior petal the smallest, subplicate, ovate. 

Junin: Near Yauli, along the railroad tracks, between Lima and 
Oroya, alt. 4,600 meters (Weberbauer 347). Huanuco: Yanashallas, 
protected rock crevices, flowers pale lavender, 5,000 meters, 2478. 
Ancash: Pelagatos, 4,400 meters, Weberbauer 7228. 

Viola micranthella Wedd. Ann. Sci. Nat. se>. 5, 1: 291. 1864. 

Annual; root thin; leaves disposed in a rosette, almost sessile 
or attenuate into a short petiole, linear or lanceolate-spathulate, 
subobtuse, glabrous, almost entire, 9-20 mm. long, 2-4 mm. broad; 
stipules linear, 2-4 mm. long, membranous; pedicels shorter than 
the leaves; petals white, the superior ones obovate, the inferior 
emarginate; spur at least as long as a third of the inferior petal, 
the 2 foremost stamens with long appendages engaged into the 
spur. F.M. Neg. 34848. 

Junin: Rio Blanco, alt. 4,000 meters, steep grass-shrub hillside, 
775. Above San Bartolome", 1,800 meters, Weberbauer 5211. 
Cuzco: Huarocondo, 3,200 meters, Herrera 3633. Prov. Canas, 



FLORA OF PERU 77 

Vargas 933,Puno: In shade-grasses, 4,100 meters, Araranca, 
Pennell 13463. Pucara, 3,700 meters, Weberbauer 429. Bolivia. 

Viola nivalis Benth. PL Hartw. 160. 1845. 

Rhizome spindle-like, simple or ramified, quite broad in its 
upper portion which is covered with leaf-scars; leaves disposed in 
a rosette, oblong-lanceolate, acute, finely denticulate or almost 
ciliate at the margin, glabrous on both faces, attenuated into a 
membranaceous petiole, 10-20 mm. long, 2-4 mm. broad; stipules 
absent; pedicels much shorter than the leaves; inferior petal broadly 
obovate, emarginate; spur obtuse, short; appendages of the 2 fore- 
most petals filiform and as long as the anther cells; style bill-shaped 
and bearing 2 membranous lobes. F.M. Neg. 24038. 

Peru (probably). Bolivia; Ecuador. 

Viola nobilis W. Becker, Bot. Jahrb. 37: 590. 1906. 

Rhizome woody, spindle-like, ramified ; leaves broadly spathulate, 
obtuse at tip, attenuate below, entire, glabrous, 20-60 mm. long, 
the blade alone measuring only 15-20 mm.; stipule partially adnate, 
ovate-lanceolate, acuminate, entire, glabrous, submembranaceous; 
flowers small, on long pedicels (4-12 cm.) and without bracts; sepals 
trinerved; petals rarely entirely purple, more commonly white and 
purple-veined; inferior petal short, obcordate, emarginate with 
remarkably short spur; style slightly enlarged above with 2 mem- 
branous appendages. 

Junin: West of Huacapistana, alt. 3,200 meters (Weberbauer 
2214). 

Viola pallascaensis W. Becker, Repert. Sp. Nov. 18: 181. 1922. 

Rhizome thick, simple or divided above, as in Viola nivalis 
Benth.; leaves disposed in rosette, oblong repand-margined, sub- 
obtuse, attenuate, hairy above, glabrous below, 7-9 mm. long, 2-3 
mm. broad; petiole 6-8 mm. long, stipules membranous, adnate 
almost on their whole length to the petiole; pedicels little longer 
than the leaves; sepals trinerved; petals 3-4 mm. long, yellow, spur 
very short; style clavate, bent at base, obliquely flattened at the 
tip, papillose, with very short bill. 

Ancash : Prov. Pallasca, below the mines of Huaura, 3,800 meters, 
(Weberbauer 7200). 

Viola pusillima Wedd. Ann. Sci. Nat. se>. 5. 2: 291. 1864. 
Species related to V. pygmaea; same habit and same linear 
leaves, acutish above, densely ciliate up to the base; petiole winged, 



78 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

broader below; no stipules; pedicels shorter than the length of the 
leaf; flowers small, yellowish; inferior petal plicate, broadly obovate, 
emarginate; spur very short; superior petals obovate, not acuminate, 
dorsal appendages of anthers half as long as length of cells; append- 
ages of style erect and short. Similar is V. Orbignyana Remy of 
adjacent Bolivia with retrorsely ciliate leaves equaled by the 
peduncles, the style appendages reflexed. The type is Mandon 941 
from Bolivia. F.M. Neg. 24042. 

Peru (probably). Bolivia. 

Viola pygmaea Juss. ex Poir in Lam. Encycl. 8: 630. 1808. 

Same habit as V. nivalis; rhizome ramified or simple; leaves 
disposed in rosette, linear, sessile, acutish or subobtuse, 1-2 (-5) 
cm. long, 1-2 mm. broad, entire, sometimes with irregularly disposed 
hairs at the margin; scale-like stipules ovate and subacute; pedicels 
as long as or somewhat shorter than the leaves; petals bluish, 
striate- veined, inferior one carinate, broadly obovate, deeply emargi- 
nate with a spur one-sixth of the length of the whole petal; append- 
ages of the 2 foremost stamens filiform, shorter than the cells; style 
S-shaped. F.M. Negs. 24043; 34856. 

Puno: Near Puno, 4,000 meters, Soukup 439. San Gaban, 
Lechler 2218. Limbani near Mount Aricoma in damp turf, 4,400 
meters, Vargas 9650. Cuzco: Gay 2060, fide Gay. Paucartambo, 
3,600 meters, Vargas 19; also 4,000 meters, Vargas 9719 (det. 
Asplund). Without locality, J. de Jussieu; Pavon (1827). Bolivia. 
"Huallpa huallpa." 

Viola replicata W. Becker, Bot. Jahrb. 37: 589. 1906. 

Species with the same habit as V. nivalis; leaves arranged in 
rosette, obovate, deflexed at tip, 10-15 mm. long, 2-3 mm. broad, 
pubescent on both faces, more or less of a grayish-green color;, 
petiole absent; pedicels as long as or longer than the leaves; sepals 
hirsute; petals white, the inferior one obcordate, slightly emarginate, 
plicate, subhirsute below; spur short; ovary fuzzily pilose. It has 
been reported that the natives use a brewage of the whole plant to 
fight against fever and colds. 

Ancash: Near Huaraz, 4,500 meters, Weberbauer 2961, type. 
"Siete sabros." 

Viola saccata Melchior, Repert. Sp. Nov. 28: 96. 1930. 

In habit a species much like V. Humboldtii; erect stems up to 
10 cm.; inferior leaves orbiculate or broadly elliptic; superior leaves 



FLORA OF PERU 79 

larger, asymmetrically ovate or rhomboid-ovate, acute at tip, 
12-23 mm. long, 8-20 mm. broad, glabrous, finely and sharply 
serrate; petiole shorter than one-half or even one-third of blade's 
length; pedicels longer than the leaves; sepals distinctly toothed; 
petals bluish-white, inferior one carinate, ending in a strangled spur 
five times shorter than the petal; stamen sessile; apical appendages 
as long as cells, the 2 foremost anthers bearing at the base of con- 
nective an appendage broad at base, filiform at tip; ovary glabrous, 
long, conical; style straight. The spur, sulcate transversely to the 
petal, and sac-like, has been described as "strangled," appropriately. 
Cuzco: Quispicanchi, Marcapata, alt. 3,100-3,200 meters, Weber- 
bauer 7780, type. 

Viola Steinbachii W. Becker, Notizbl. Bot. Gart. Berlin 9: 
1040. 1926. V. boliviano, W. Becker, Beih. Bot. Centralbl. 22, pt. 2: 
88. 1907, not Britton, 1889. 

Trailing or twining shrub; twigs reaching 40 cm. long; leaves 
ovate, acute at tip, asymmetrically cordate at base, glabrous, 
indistinctly glandulous-serrate, 20-50 mm. long, 11-30 mm. broad; 
stipule as in V. Humboldtii, membranaceous, ciliate; pedicels longer 
than the leaves; flowers white, blue-striate; inferior petal subacumi- 
nate; spur short; appendages of foremost anthers long; style subulate. 

Puno: Sandia, alt, 2,100-2,300 meters (Weberbauer 6^5). Cuzco: 
Cerro de Cusilluyoc, Pennell 13993 (det. Melchior). 

Viola stipulates Swartz, Prodr. 117. 1788; 86. V. begoniaefolia 
Benth. PL Hartw. 161. 1845. lonidium stipulare Roem. & Schult. 
Syst. 5: 392. 1819. 

Species related to V. Steinbachii; creeping rhizome; erect stem 
10-20 cm. long, rarely reaching 30 cm. (Schlim 579) ; leaves generally 
only at the end of ramifications, elliptic-lanceolate, glandulous- 
serrate or slightly and shallowly dentate or. crenate, 23-82 mm. 
long, 9-34 mm. broad; stipules brown, up to 20 mm. long and 6 mm. 
broad; pedicels shorter than length of leaf; petals whitish, blue- veined, 
inferior one obovate, subacuminate; spur very short; appendages of 
foremost anthers long; style subulate. 

Junin: Tarma, west of Huacapistana (Weberbauer 2102). With- 
out locality (Lobb s.n.). West Indies; Venezuela; Colombia; Ecuador. 

Viola Weberbaueri W. Becker, Bot. Jahrb. 37: 588. 1906. 

Related to V. micranthella; diameter of rosette up to 6 cm.; 
annual with thin root; leaves linear-spathulate, subobtuse, entire, 



80 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glabrous but for the ciliate margin, 15-30 (45) mm. long, 1.5-3 mm. 
broad; stipules small, very narrow or wanting; pedicels shorter than 
leaves; sepals linear-oblong, subobtuse, glabrous, trinerved, mem- 
branous at the margin; petals pale yellow, inferior petal carinate, 
emarginate; spur short; style curved, bill-shaped at tip and bearing 
3 membranaceous appendages, the middle one short and directed 
backwards, the lateral ones longer, narrow and protruding forwards; 
ovary glabrous. 

Arequipa: Mollendo, in loma formation, alt. 100 meters (Weber- 
bauer 1541, type); Hill 29 (det. Becker). Arequipa: Tingo, open 
sandy slopes, flowers citron yellow, Pennell 13115 (det. Melchior). 

Viola Weddellii W. Becker, Repert. Sp. Nov. 26: 25. 1929. 

Species with the same aspect as V. Humboldtii; stem 5-6 cm. 
long, densely foliate; leaves orbicular, 3-4 mm. long and broad, 
rounded or almost cuneate at base, obtuse at tip with 2-3 (4) 
crenatures; pedicels 15-20 mm. long; flowers small, 6-7 mm. long; 
spur short; ovary red-spotted. 

Puno: Carabaya, above San Juan del Oro (Weddell 1847). Cuesta 
de Quequenta (Raimondi 9215}. Cuzco: Cerro de Cusilluyoc, 3,400 
meters, petals light violet, Pennell 13890 (det. Melchior). Prov. 
Acanacu, Paucartambo, 3,500 meters, Vargas 318 (det. Standl.). 

Viola Weibelii Macbr., sp. nov. 

Rhizoma verticale, lignosum, plus minusve divisum ; folia lineari- 
oblonga, circa 1.5 cm. longa, 2 mm. lata, inciso acuteque irregulariter 
ciliato-serrata, supra subfoveolata, glabra, vix in petiolum angustata; 
stipulae elongatae, chartaceae, profunde fimbriatae; pedunculi circa 
1 cm. longi; sepala lineari-lanceolata acuminata; petala superiora 
lateraliaque obovata integra; petala infimum late obovata crenata 
valde calcaratum; stylus paullo dilatatus vel geniculatus. Section 
Andinium. Flowers white or faintly lavender, with yellow eye. 

Junin: Cerro de Pasco, limestone slopes, loose soils, Macbride 
3060, type. 

10. NOISETTIA HBK. 

Flowers zygomorphous, the petals being very unequal in size and 
form, the foremost one large with a long and thin spur. Stigma 3- 
lobed. Capsule wrapped in the remains of corolla and calyx. 

Noisettia orchidiflora (Rudge) Ging. in DC. Prodr. 1: 290. 1824. 
N. longifolia HBK. Nov. Gen. & Sp. 5: 384. pi. 499b. 1823. 



FLORA OF PERU 81 

Half-shrub 30-50 cm. high, often unramified; leaves oblong- 
lanceolate, 10-15 cm. long, 2-4 cm. broad, glandulous-serrulate; 
inflorescences few-flowered (2-5) ; flowers yellowish or white; stamens 
and style shorter than the calyx; capsule greenish, 7-10 mm. long; 
seeds becoming black, 2 mm. in diameter. Illustrated, Mart. Fl. 
Bras. 13, pt. 1: pi. 70. 

San Martin: Tarapoto, Spruce 4006; Williams 6013. Pongo de 
Chilcayo, Ule 6342. Loreto: Chinganilla (Huber 1500). Guiana; 
Brazil. 

11. LEONIA Ruiz & Pavon 

Flowers actinomorph. Stamens completely united into a tube, 
the anthers being immersed in its edge and without appendages. 
Fruit a nut with a hard pericarp. Wood esteemed for construction 
(Williams). 

Leonia cymosa Mart. Nov. Gen. & Sp. 2: 88. 1826. 

Tree 3-8 meters high; leaves oblong or elliptic, slightly acumi- 
nate, 10-15 cm. long, 4-6 cm. broad, entire or shallowly dentate, 
a little decurrent on the upper part of the petiole; flowers yellow 
or orange, 3-4 mm. long, arranged irregularly in dichotomous cymes 
3-4 cm. long; fruit berry-like, with woody pericarp. Illustrated, 
Mart. Nov. Gen. & Sp. 2: pi. 169. 

Loreto: Basin of the Maranon, mouth of Santiago at Pongo de 
Manseriche, Tessmann 4343, 4436. Yurimaguas, Poeppig 2369. Near 
Iquitos, King 893; 303; 325; Killip & Smith 26959; Williams 8009; 
1835; 3702. Pebas, Williams 1573. Lower Rio Nanay, Williams 692. 
Brazil. 

Leonia glycycarpa R. & P. Fl. Peruv. 2: 69. pi. 168. 1799. L. 
racemosa Mart. Nov. Gen. & Sp. 2: 86. 1826. 

Shrub 1.5-6 meters or tree up to 35 meters (Krukoff); leaves big 
(12-35 cm. long, 5-12 cm. broad), oblong, shortly acuminate; inflo- 
rescences more or less pendulous, sometimes 40 cm. long; flowers 
whitish; fruit the size of a peach or smaller. The flesh of the fruit 
is soft or jelly-like, edible. Common in forest free from inundations 
(Williams). Illustrated, Mart. Fl. Bras. 13, pt. 1: pi. 80. 

San Martin: Tarapoto, Williams 6565, Spruce 4424, 4613; Ule 
6444- Junin: Rio Perene", 600 meters, Killip & Smith 25101. 
Ayacucho: Rio Apurimac, valley at 400 meters, Killip & Smith 
23049. Loreto: Basin of Maranon, Tessmann 4455; 5136. Ucayali, 
Tessmann 3185; 5428. Balsapuerto, Klug 2958. Near Iquitos, 



82 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

King 1223; Killip & Smith 27144; 27017; 27457; 27130. Williams 
8014; 8166. Florida, Rio Putumayo, at mouth of Rio Zubineta, 
Klug 2313. San Antonia De Playa (Tafalla s.n.). Yurimaguas, 
Williams 5254- La Victoria, Williams 3095; 2946. Rio Itaya, Wil- 
liams 3483; Killip & Smith 29429. Caballo-Cocha, Williams 2195. 
Bolivia; Brazil; Acarai Mountains at the border of British Guiana. 
"Witu-malla," "urca-tamara," "nina caspi," "trapiarana." 

TURNERACEAE 

Reference: Urban, Jahrb. Bot. Gart. Berlin 2: 1-150. 1883. 

Herbs or low shrubs (sometimes tall or trees) with alternate 
leaves, no stipules or these small, regular solitary or few axillary 
hermaphrodite flowers, their 5 petals soon falling, twisted in bud, 
their caducous sepals imbricated, united below. Peduncles sometimes 
joined to the petiole. Stamens 5, inserted at middle or base of the 
calyx tube. Ovary 1-celled, the 3 styles simple or parted, the stigmas 
conspicuously brush-like. Capsules thin or firm with many pitted 
seeds. The style character is distinctive. Urban's work appeared 
later the same year in Mart. Fl. Bras. 13, pt. 3: 86-170, with a num- 
ber of illustrations. 

Calyx tube without a crown; styles simple 1. Turner a. 

Calyx tube with a fringed crown at petal-base; styles parted. 

2. Piriqueta. 

1. TURNERA [Plum.] L. 

Pubescence usually simple or nearly wanting. Leaves various, 
often glandular toward or at the petiole. Flowers sometimes crowded 
or aggregated, always bracteolate, usually with no pedicel, the 
peduncles free or joined to the petioles. Petals generally yellow. 
Since Urban's monograph a number of species have been described 
by him from the Amazonian Valley, notably in Repert. Sp. Nov. 13: 
155-159. 1914, which may of course be found in eastern Peru. 

Peduncles adnate to the petioles. 

Softly villous annual; seeds curved, tuberculate T. Pumilea. 

Strigose perennial or suffrutescent; seeds little curved, striate. 

T. ulmifolia. 
Peduncles free. 

Pedicels none; leaves ovate-lanceolate, rather coarsely serrate. 
Calyx 12-18 mm. long; bracteoles 2.5-8 mm. wide. . . .T. acuta. 
Calyx 8-10 mm. long; bracteoles to 1.5 mm. wide. 

T. macrophylla. 



FLORA OF PERU 83 

Pedicels developed; leaves narrowly lanceolate, obscurely serrate. 

Styles conspicuously flagellate, to 6 mm. long T. luminosa. 

Styles minutely flagellate, to 4 mm. long T. Weddelliana. 

Turnera acuta Willd. ex Roem. & Schult. Syst. 6: 678. 1820; 96. 

Lightly pubescent shrub attaining at least 1 meter; petioles 4-10 
mm. long, glandular beneath the leaf blade, this unequally narrowed 
at base and apex, the larger 5-12 cm. long, 2-4 cm. wide, shortly 
pilose beneath on the nerves; flowers dimorphic, solitary, the fruiting 
peduncles 3-10 mm. long; bracteoles 8-12 mm. long; calyx lobes 
narrow with setaceous tip; petals about 13 mm. long; filaments 
finely pubescent; styles hirsute, the longer 9-11 mm. long, flagellate 
for 1 mm., the shorter half as long; capsules hirtellous, the seeds 
little to manifestly curved, striate. Illustrated, Mart. Fl. Bras. 13, 
pt. 3: pi. 38. 

Loreto: Iquitos, Williams 3661. Rio Nanay, Williams 778. 
Brazil; Venezuela. 

Turnera luminosa Macbr. Candollea 6: 12. 1934. 

Branchlets striate, caducously rusty-appressed-strigose as the 
buds, petioles and young leaves on the midnerves beneath; stipules 
1 mm. long; petioles 2-3.5 mm. long, eglandular; leaves narrowly 
elliptic, cuneately acute at both ends, 4-7 cm. long, 1.5-2 cm. wide, 
medially crenate-serrulate, pale green, membranous, soon glabrous 
except for sparse minute pubescence on the veins; flowers solitary; 
peduncles 5-10 mm. long, the pedicels 4-5 mm. long; calyx appressed- 
pilose, 7-9 mm. long, narrowly campanulate, lobes half as long, 
ovate-acuminate, the tube pilose within,' scarcely 4 mm. wide at 
throat; petals nearly 15 mm. long; filaments pilose below; anthers 
long-apiculate, 4 mm. long; styles glabrous, 6 mm. long, the 5-7 
flagellate stigmas about 1 mm. long; capsules glabrous, verruculose- 
tuberculate, the seeds strongly curved, obovate-oblong, sparsely 
pilose, reticulate, attenuate above. A shrub 1 meter high (Weber- 
bauer), perhaps not separable from T. Weddelliana typically of 
Paraguay but the flowers apparently monomorphic with longer 
styles; stipules also longer. 

Cuzco: Bush wood in savanna near Quillabamba, Urubamba 
Valley, Weberbauer 7952, type. 

Turnera macrophylla Urban, I.e. 95. 

Similar and allied to T. acuta but with rather larger leaves more 
pilose beneath, monomorphic flowers, the fruiting peduncles 4-5 mm. 



84 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, bracteoles 3-6 mm. long, the filaments merely puberulent. 
The Peruvian occurrence is recorded by Urban as "in andibus 
Peruvianis: fide herb. Acad. Petropol." F.M. Neg. 13590. 

Peru (cf. above). Amazonian Brazil. 

Turnera Pumilea L. Amoen. Acad. 5: 395 (reprint, 377) 1760; 
114. 

White-pubescent, estipulate, often bushy branched annual, the 
monomorphic flowers aggregated in terminal clusters; petioles 5-10 
mm. long; leaves oblong-lanceolate or more or less obovate, cuneate 
at base, acute or rather obtuse, 2-6 cm. long, 1-2.5 cm. wide, coarsely 
crenate-serrate, sometimes glandular at base; calyx 6-7 mm. long, 
the petals little longer than the acuminate lobes; filaments glabrous 
as the style or this sparsely pilose, 3.5-5 mm. long, unequally 
flagellate; capsules hirsute; seeds curved, reticulate- tuberculate. 
Similar and to be expected is T. melochioides Camb., 115, perennial 
or shrubby, the leaves sometimes narrower, the flowers dimorphic, 
the seeds not tuberculate. 

Tumbez: Weberbauer 7732. Piura: Serran, Weberbauer 5986; 
59^6 (det. Urban). Tablazo, Haught 149. Cabo Blanco, Haught 
F-53. San Martin: Tarapoto, Spruce 4075 (det. Urban). Warm 
America. 

Turnera ulmifolia L. Sp. PI. 271. 1753; 138. T. peruviana 
Willd. ex Roem. & Schult. Syst. 6: 679. 1820. 

More or less woody perennial with usually narrowly obovate 
short- or long-petioled leaves and short- or well-peduncled homo- or 
heterostylous flowers, the peduncles nearly always totally adnate; 
calyx 11-25 mm. long; petals sometimes reddish; filiform style 
branchlets 1-2 mm. long; seeds obovate to oblong, little if at all 
curved, reticulate striate. Highly variable vegetatively; among the 
many named forms may be mentioned the var. intermedia Urb., 140, 
with yellow heterostylous flowers, calyx 12-20 mm. long, peduncles 
3-7 mm. long, lower leaves ovate. Notwithstanding the name, 
T. peruviana was from Venezuela; cf. HBK. Nov. Gen. & Sp. 6: 
127. 1823. Illustrated, Fawc. & Rendle, Fl. Jam. 5, pt. 3: fig. 91. 

Cajamarca: Jan, Weberbauer 6236 (det. Urban). San Martin: 
Tarapoto, Williams 5540; 5533; 6416; Ule 6460 (var. intermedia, 
det. Urban); Williams 5533. Near Moyobamba, King 3570 (det. 
Standl.). Rio Huallaga, King 4073. Huanuco: Huanuco, 3515; 
Stork & Horton 9386. Warm America; India. 



FLORA OF PERU 85 

Turnera Weddelliana Urb. & Rolfe, ex Urban, I.e. 90. 

Vegetatively like T. luminosa but the stipules minute, rarely 
even 0.5 mm. long; peduncles 2-15 mm. long; pedicels 3-8 mm. long; 
calyx 6-12 mm. long, the lobes a third longer than the tube; flowers 
dimorphic; longer filaments 6-7 mm. long, the shorter 3 mm. long; 
longer styles 3.5-4 mm. long. Very probably the Peruvian collec- 
tion, if Peruvian, is referable to T. luminosa. The closely allied 
T. Hindsiana Benth., 91, from southern Ecuador, may be distin- 
guished by its broader leaves and glabrous filaments. F.M. Negs. 
13612 (var.); 24132. 

Peru(?): Without locality (Lobb 292). Southern Bolivia to 
Paraguay. 

2. PIRIQUETA Aubl. 

Except as noted in the key with the general characters of Turnera 
but often stellate-pubescent and flowers generally solitary in the 
axils, not always bracteolate, and the peduncles always free. In 
some related families the calyx may or may not have a crown in the 
same genus. 

Piriqueta cistoides (L.) Griseb. Fl. Brit. W. Ind. 298. 1860; 
73. Turnera cistoides L. Sp. PI. ed. 2. 387. 1762. 

Annual but often stout and several dm. high; leaves subsessile 
or the petioles to 1 cm. long, ovate or nearly oblong, 3-8 cm. long, 
not glandular; flowers homostylous; bracteoles none or minute; 
pedicels 3-6 mm. long; calyx 4-9 mm. long, the lobes about half as 
long as the tube; capsules pilose but smooth, the slightly curved 
seeds etuberculate. Variable, the Peruvian form being var. latifolia 
Urban, I.e. 74, with mostly ovate leaves 1.5-3 cm. wide, nearly always 
deeply and usually irregularly crenate or erose-dentate. P. Duarteana 
(Camb.) Urban, 66, which has been found in Brazil near southern 
Peru is a perennial with violet or roseate heterostylous flowers, the 
calyces 7-16 mm. long, the capsules tuberculate. Illustrated, Mart. 
Fl. Bras. 13, pt. 3. pi. 35. F.M. Neg. 38468 (var.). 

San Martin: Tarapoto, Williams 7119. Warm America. 

MALESHERBIACEAE 

Pubescent herbs or mostly simple-stemmed half-shrubs with 
densely crowded frequently narrow leaves and more or less elongate 
spike-like racemes of yellowish or red flowers, the leaves gradually 
reduced above and bract-like in the inflorescence. Technically the 
family is similar to Passifloraceae and to Turneraceae but is distin- 



86 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

guished from both by the lack of aril and by the placement of the 
3-4 styles, these arising from below the ovary tip and more or less 
remotely from each other. Ovary stiped. Capsule enclosed in the 
calyx tube. 

MALESHERBIA R. & P. 

Gynopleura Cav. Icon. 4: 51. pi. 376. 1797. 

This, the only genus, was dedicated to Lamoignon de Males- 
herbes, "lover of plants and patron of botanists"; he was a minister 
under Louis XVI and was beheaded, perhaps because of the "great- 
ness of his soul." (Ruiz & Pavon.) Harms, in Notizbl. Bot. Gart. 
Berlin 8: 209-212. 1922, gave a helpful synopsis of the Peruvian 
species, at that time only 6. A common name is "clavelina." 

Calyx elongate, cylindrical or more or less enlarged at the middle. 
Leaves variously crenate or dentate but not pinnately divided. 
Calyx tubular, that is, little if at all inflated medially. 

Flowers yellowish, sometimes reddish tinged; crown more or 

less undulate. 

Calyx 3 cm. long, 5-6 mm. broad; leaves unequally ser- 
rulate M. tubulosa. 

Calyx 3.5-4 cm. long, 6-10 mm. broad; leaves evenly 

crenulate-dentate or subentire. 

Calyx somewhat inflated, 9-10 mm. broad; leaves 

appressed sericeous pilose, eciliate . . M. Weberbaueri. 

Calyx tubular, 6-8 mm. broad; leaves spreading villous, 

viscid-ciliate with crisped hairs M. Galjufii. 

Flowers fiery red ; crown deeply lobed, the lobes dentate. 

M. ardens. 

Calyx definitely inflated medially. 

Leaves sessile, linear-oblong, finely and irregularly toothed. 

M. thyrsiflora. 
Leaves narrowed to petiolar base, elliptic, evenly toothed. 

M. scarlatiflora. 

Leaves pinnately divided M. angustisecta. 

Calyx funnelform, 1.5-2 cm. long. 

Leaves deeply pinnatifid; crown 7 mm. high M. haemantha. 

Leaves not pinnatifid; crown 13 mm. high M. turbinea. 

Malesherbia angustisecta Harms, Notizbl. Bot. Gart. Berlin 
8: 210. 1922. 



FLORA OF PERU 87 

Softly ashy-villous, little if at all branched shrub with greenish- 
white flowers; leaves oblongish, deeply pinnatisect, the 4-5 divisions 
narrowly linear and sometimes with a lateral tooth or lobe; stipules 
linear to 1.5 cm. long; flowers shortly pedicellate often 2-4 on short 
branchlets, these forming an elongate terminal panicle; calyx nearly 
cylindrical, slightly enlarged toward the tip, 2-2.7 cm. long, 5-6 mm. 
broad; sepals oblong-ovate, acutish, sparsely pilose or glabrate, 7-8 
mm. long, about equaled by the obovate broadly based petals; 
crown minute; ovary a little villous only at tip. Nearly M. pulchra 
Phil, but that has a broader calyx and much higher crown; cf. Harms' 
remarks. Similar is M. tenuifolia D. Don, also Chilian apparently 
instead of Peruvian as per Kew Index, but crown of that is deeply 
lobed. Only more collections can determine if the crown difference 
is constant but also the Chilean plants seen are more strigillose than 
loosely villous. To 0.5 meters high (Weberbauer). 

Arequipa: In rocks with other shrubs and cacti, 2,350 meters, 
above Caraveli, Weberbauer 7179, type. 

Malesherbia ardens Macbr. Field Mus. Bot. 4: 117. 1927. 

Softly ashy-villous strict shrub with racemes of tubular fiery red 
flowers; upper stem leaves sessile, oblong-lanceolate, 3-4 cm. long, 
6-12 mm. wide, cuneate at base, obtuse, unequally and crisply 
crenate-dentate or deeply serrate especially at the base, rather sub- 
appressed-hirsute, the margins ciliate with some long yellowish 
hairs; raceme-leaves little reduced, the racemes 1-3 dm. long; pedi- 
cels 1-1.5 cm. long; bracts narrowly oblong-lanceolate, about 1 cm. 
long; calyx little inflated medially, 2.5-3 cm. long, 8-10 mm. broad, 
lightly pilose; sepals ovate, acute, 6 mm. long, 2 mm. wide at base; 
petals narrowly oblong-lanceolate, 5 mm. long; crown about 3 mm. 
high, deeply lobed, the lobes acutely dentate; stamens exserted about 
8 mm.; capsules pilose, long-exserted, the oval seeds foveolate. To 
0.5 meters high (Weberbauer). 

Moquehua: In shrubby dry ravines, southeast of Moquehua, 
Weberbauer 7436a, type; also between Moquehua and Torata, 7436. 

Malesherbia Galjufii Macbr. Field Mus. Bot. 4: 92. 1925. 

Softly villous throughout with a close indument of short spread- 
ing hairs; leaves sessile, acuminate or acute, linear-lanceolate, mi- 
nutely crenate-dentate, conspicuously crisply ciliate on the margins 
with yellow hairs even 1 mm. long; racemes leafy bracted to 3 dm. 
long; pedicels 8 mm. long; flowers greenish-yellow, long-pilose, 
tubular, 3.5-4 cm. long, rarely 9 mm. broad; sepals lanceolate, 



88 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

acuminate, 7 mm. long, the petals similar but shorter; crown irreg- 
ularly and lightly dentate; stamens exserted about 5 mm. With 
flowers similar to M. tubulosa but with entirely different leaves and 
pubescence, this velvet-like except for the strikingly ciliate leaf- 
margins. It is named for Sr. Cristobal Galjuf, proprietor of the coal 
mines at Huaron, who generously contributed mules and arriero to 
a Field Museum expedition on which this handsome plant was found. 
Junin: Huartas, in loose rocky soils of steep canyon side, 1347. 

Malesherbia haemantha Harms, Notizbl. Bot. Gart. Berlin 8: 
211. 1922. 

Hirsute-villous half-shrub with deeply pinnatisect leaves and 
blood-red turbinate flowers borne in long terminal racemes; pinnae 
12-17, narrowly lanceolate, 3-7 mm. long, often coarsely and irregu- 
larly crenate-serrate, the serrations setose-mucronulate; racemes to 
about 2.5 dm. long, bracteate, with reduced lanceolate leaves; calyx 
villous, 1.5-2 cm. long, at top 8-10 mm. broad; sepals lanceolate, 
acute, setulose at tip, 7-8 mm. long, nearly 3 mm. wide at base; 
petals nearly lanceolate, setose tipped, 5-7 mm. long, 1 mm. wide at 
base; crown to 7 mm. high, variously cleft. Calyx and stamens 
blood-red, the crown blood-red with black throat, the anthers black 
(Weberbauer). 

Arequipa: Above Quicacha, prov. Camana, 2,100 meters, xero- 
phytic cactus-shrub formation, Weberbauer 7181, type. 

Malesherbia scarlatiflora Gilg, Bot. Jahrb. 50: Beibl. Ill: 11. 
1913. 

Strict or little branched shrub more or less clothed below by the 
persisting crowded leaf -remains; leaves oblong-obovate, rounded at 
tip, gradually narrowed to a petiolar base, densely silky yellowish 
tomentose on both sides, equally serrate-dentate, 1-10 cm. long, 
2.5-3.5 cm. wide, the nerves and veins prominently reticulate 
beneath ; flowers scarlet, the limb yellow or greenish-yellow in racemes 
to 3.5 dm. long, the bracteate leaves strongly reduced above; calyx 
somewhat inflated, about 4 cm. long, 10 mm. broad at middle, densely 
silky yellow tomentose; sepals lanceolate, very acute, 7-8 mm. long, 
2 mm. wide at base, the petals shorter and narrower; otherwise like 
M. Weberbaueri, fide Gilg. My specimen is from a plant 1 meter 
high, candelabra-like in branching, leafless except at inflorescence, 
the flowers orange red, the corolla tips greenish-yellow. 

Lima: In stony outcrop, above Matucana, 2,400 meters, Weber- 
bauer 5219, second no. cited; in rocks, Savatier 488; rocky out- 



FLORA OF PERU 89 

crop, 1205. Huancavelica: Huaytara, between 13 & 14 above 
Pisco, 2,700 meters, Weberbauer 5411, first no. cited (type?). 

Malesherbia thyrsiflora R. & P. Fl. Peruv. 3: 30. pi. 254- 1802. 

Suffruticose, little branched below, the simple branches to 1 
meter high, rather yellowish villous-hirsute throughout; leaves 
linear- or oblong-lanceolate, more or less irregularly, sometimes 
deeply, sinuate-dentate, sessile, 5-7 cm. long, 3-6 mm. wide, the 
upper somewhat reduced and bract-like in the rather loose racemes, 
these 1-2 dm. long; flowers reddish yellowish-green, densely villous, 
about 3.5 cm. long, 8 mm. wide medially where somewhat inflated; 
sepals narrowly linear-lanceolate, long-acuminate to 6 mm. long, the 
petals about half as long, ovate-acuminate; crown about 3 mm. 
high, lobed. This excludes G. tubulosa Cav. cited by Ruiz & Pavon 
as a synonym possibly in confusing the species but apparently 
rather by assumption, since the generic name was a synonym. 
Pubescence slightly "sticky," the plants somewhat foetid. 

Lima: St. Eulalia, Chosica, Weberbauer 5723 (det. Harms). 
Cheuchin, Dombey; Ruiz & Pavdn (part type). Canta, Huarochiri 
and Caxatambo (Ruiz & Pavon, part type). Rio Chillon, Pennell 
14476. 

Malesherbia tubulosa (Cav.) Macbr. Candollea 5: 393. 1934. 
Gynopleura tubulosa Cav. Icon. 4: 52. 1797. M. cylindrostachya 
Urb. & Gilg, Bot. Jahrb. 37: 592. 1906. 

Stems strict, densely and softly short-ashy-pilose, leafless below 
but marked by the bases or remains of the numerous leaves, these 
oblong-linear, cuneate at base, acute, appressed hirsute-pilose, un- 
equally and crisply serrate, to several cm. long, 4-8 mm. wide, the 
upper gradually reduced where bract-like and partly concealed in 
the elongate spike-like racemes of reddish-tinged ochre colored 
flowers; calyx narrowly cylindrical to 3 cm. long, 5-6 mm. broad, 
pilose, the lobes lanceolate, acute, 4-5 mm. long; petals narrowly 
ovate, scarcely 3 mm. long, the crown erect, deeply undulate, 1.5 
mm. high; filaments to 1 cm. exserted; ovary stiped, the 3-4 styles 
to 2 cm. exserted; tips of capsule valves exserted; seeds oval, striate 
both across and lengthwise. Well-developed plants half-shrubs to 
1 meter high. Part of the material at Madrid labeled recently as 
by Ne'e is referable to M. thyrsiflora but, as indicated by Cavanilles' 
name and by a specimen by Ne in Cav. manuscript at Geneva the 
application of his name is as interpreted here. F.M. Negs. 2450 
(cylindrostachya); 24124. 



90 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: (Haenke). Lima: Above Matucana (Ball). In rock 
detritus, 1,700 meters, above St. Bartholome", Weberbauer 1690, type 
M. cylindrostachya. Purruchucho, Nee (type, as at Geneva); also 
between Obrajillo and San Buenaventura, Nee (Madrid). 

Malesherbia turbinea Macbr. Field Mus. Bot. 4: 118. 1927. 

Apparently allied to M. haemantha (only flowering branches 
known); calyx 12-15 mm. long, 7 mm. broad, sparsely pilose; crown 
13 mm. high, irregularly denticulate-crenate; sepals 8-9 mm. long; 
pedicels nearly 15 mm. long; stamens scarcely exserted; capsules 
pilose, little exserted; seeds obscurely transversely but strongly 
longitudinally striate, about 1 mm. wide, nearly 2.5 mm. long. 
Flowers blood-red, the anthers apparently white or yellowish. The 
upper bract-like oblong-ovate leaves are merely crenate. Each 
leaf-crenation is tipped with one long cilium. 

Tacna: In rainy green shrubs and Cereus, Candarave, Weberbauer 
7364, type. 

Malesherbia Weberbaueri Gilg, Bot. Jahrb. 50: Beibl. Ill: 11. 
1913. 

Densely appressed silky-villous simple-stemmed shrub with 
crowded linear-lanceolate sessile leaves and long terminal racemes 
of leafy bracted greenish yellow or reddish tinged sub tubular flowers; 
leaves subentire or obscurely crenate-serrulate, 8-12 cm. long, about 
1 cm. wide, acute base and apex; calyx 3.5-4 cm. long, 8-10 mm. 
wide, sepals lanceolate, acuminate, 7-8 mm. long, 2 mm. wide at 
base, the petals shorter and narrower; crown irregularly and slightly 
dentate; capsule included or barely exserted; seeds minutely and 
obscurely striate. To about 1 meter high in steep, loose, rocky 
soils. Species distinctive in its closely appressed sericeous pubes- 
cence. F.M. Neg. 2460. 

Huancavelica: Valley of the Huarpa, Weberbauer 5672, type. 

PASSIFLORACEAE. Passionflower Family 
By E. P. Killip, United States National Museum 

References: Masters in Mart. Fl. Bras. 13, pt. 1. 1872; Killip in 
Field Mus. Bot. 19. 1938. 

Passifloraceae consists of 12 genera, four of which are found 
in the New World. By far the largest genus is Passi flora, which 
has about 400 species, nearly 360 of them being endemic to the 
New World. The flowers of most of the species occurring in Peru 



FLORA OF PERU 91 

are very showy, and their structure is highly complicated. The 
names commonly applied to passionflowers in Peru are "granadilla," 
"niorbo," and "tumbo." "Tumbo," "tintin" and "apincoya" are 
Quechua names (Cook). Herrera has recorded the native name 
"inti sisa," but has by error associated it with a Brazilian species. 

Stamens 5, in the Peruvian species borne on a well-developed gyno- 
phore; sepals 5; petals (if present) 5 1. Passi flora. 

Stamens 8 or 10, inserted on the floor of the calyx; sepals and petals 
normally 4 2. Dilkea. 

1. PASSIFLORA L. 

Herbaceous or woody vines or scandent shrubs, usually climbing 
by tendrils, rarely (one Peruvian species) small trees or erect shrubs; 
leaves alternate, petiolate, simple or compound, lobed or unlobed, 
the petiole often glanduliferous; inflorescence axillary, simple or 
occasionally compound; bracts usually present, scattered or forming 
an involucre; calyx tube patelliform, campanulate, funnelform, or 
cylindric; sepals 5; petals 5, alternate with the sepals, sometimes 
wanting; corona of 1 to several series of distinct or more or less 
united filaments; operculum borne within the corona, membranous, 
flat or plicate, entire or filamentose; stamens 5, the filaments mona- 
delphous in a tube closely adnate to a gynophore, distinct above, the 
anthers 2-celled; ovary borne on a gynophore; styles 3, the stigmas 
capitate; fruit indehiscent, containing a mucilaginous pulp; seeds 
borne on 3 parietal placentae, more or less compressed, reticulate, 
punctulate, or transversely grooved. 

Passi flora rosea, probably a hybrid, has an abnormal genital 
structure, and the above description does not apply to it. 

A. Ovary 3-angled, broadly truncate at apex, the styles projecting 
from the top of the angles; trees, shrubs, or woody vines, without 
tendrils or with weak ones; leaves simple, unlobed, entire; 
bracts and stipules setaceous, soon deciduous. 
Calyx tube campanulate or cylindric-campanulate, shorter than 
the sepals; flowers white or greenish. 

Peduncles terminating in a tendril P. cirrhipes. 

Peduncles not terminating in a tendril. 

Erect shrub or a tree; peduncles once-furcate. .P.frutescens. 
Scandent shrubs; peduncles simple. 

Corona filaments in about 6 series; leaves rounded or 
obtuse, coriaceous, strongly nerved P. costata. 



92 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Corona filaments in 2 series; leaves mostly acute, not 
strongly nerved, membranous P. Tessmannii. 

Calyx tube cylindric, longer than the sepals (subequal in P. cauli- 
flora}; flowers red, yellow, or orange, in short fascicles or 
in racemes. 

Inflorescence fasciculate; operculum cleft into 5 linear segments. 

Calyx tube less than 2.5 cm. long; ovary pubescent; leaves 
coriaceous P. cauliflora. 

Calyx tube about 3 cm. long; ovary glabrous; leaves mem- 
branous P. skiantha. 

Inflorescence racemose; operculum filamentose, at least part way. 

Calyx tube puberulent; leaves ovate or ovate-oblong. 

P. pyrrhantha. 
Calyx tube glabrous; leaves oblong P. spinosa. 

A. Ovary terete or subangular, not definitely 3-angled, acute or 
rounded, the styles projecting from the center of the top of the 
ovary, or the ovary tapering to the styles; herbaceous or some- 
times woody vines, usually with well developed tendrils. 

Tendrils of 2 forms, one axillary on the main stem, the other on 
one of each pair of peduncles P. heterohelix. 

Tendrils, if present, all axillary. 

Petiolar glands globose, 3-4 mm. wide, with a stipe 6-8 mm. 
long; bracts borne at middle of peduncle, lacerate. 

P. adenopoda. 

Petiolar glands smaller or wanting; bracts scattered along 
peduncle or forming an involucre near base of flower, want- 
ing in P. rubra. 

B. Bracts linear-subulate or setaceous, less than 1 mm. wide, 
scattered along the peduncle; flowers usually not more than 
5 cm. wide and not highly colored; operculum plicate. 

Petiolar glands present; seeds reticulate. 
Petals none. 

Leaves prevailingly transverse-oblong, much broader than 
long, the glands sessile, borne at or below the middle 
of the petiole P. coriacea. 

Leaves prevailingly 3-lobed (in Peruvian plants), longer 
than broad, the glands borne above the middle of the 
petiole P. suberosa. 



FLORA OF PERU 93 

Petals present. 

Leaves membranous, pubescent, dentate or denticulate, 3- 
lobed to below the middle, the petioles biglandular 

near the apex; ovary densely pilose P. morifolia. 

Leaves coriaceous, essentially glabrous, entire, 3-lobed not 
more than to the middle, the petioles biglandular near 

the base; ovary glabrous P. Lobbii. 

Petiolar glands none, though in 2 species the petioles with 
auricular appendages near their base; seeds transversely 
sulcate. 

Bracts none; fruit elongate-obovoid ; seeds with smooth ridges. 

P. rubra. 
Bracts present; fruit globose or fusiform; seeds with rugulose 

ridges. 
Leaves narrowly lanceolate to ovate-lanceolate, unlobed or 

with 2 reduced lateral lobes. 

Peduncles 2-flowered; petioles without auricular append- 
ages P. quadriflora. 

Peduncles 1-flowered; petioles with 2 auricular append- 
ages at or below the middle. 

Plant glabrous or cinereous-pubescent; fruit globose, 

not more than 1.5 cm. in diameter. .P. auriculata. 

Plant ferruginous- tomentose; fruit broadly ovoid, more 

than 1.5 cm. in diameter P. ferruginea. 

Leaves 2-3-lobed or, if subentire, broader than long. 
C. Inner corona filaments linear, broadly dilated at the apex 
and often lobulate. 

Ovary densely cano-tomentose P. leptoclada. 

Ovary glabrous. 
Leaves 3-lobed a third to a half their length, the lobes 

ascending, acute or subacute. 
Leaves coriaceous or subcoriaceous, lustrous above, green 

beneath P. tricuspis. 

Leaves membranous, dull above and fasciate along the 

nerves, reddish beneath P. trifasciata. 

Leaves 2-lobed, the lobes widely divergent (hence leaves 
transversely oblong), occasionally with a small inter- 
mediate lobe present. 
Petals none; sepals broadly ovate; ovary ovoid. 

P. Poeppigii. 



94 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petals present; sepals linear-lanceolate; ovary fusiform. 

P. amalocarpa. 

C. Inner corona filaments filiform, capillary. 

Leaves distinctly longer than broad, shallowly bilobate, 
densely pilosulous beneath P. bauhinifolia. 

Leaves broader than long or the length along the midnerve 
subequal to the greatest width, glabrous or pulverulent 
beneath. 

Ovary white-sericeous; leaves finely pulverulent beneath. 

P. Candollei. 

Ovary glabrous; leaves glabrous or very sparingly pubescent 

beneath. 

Peduncles more than 4 cm. long; outer corona filaments 
falcate, conspicuously dilated near the middle. 

P. punctata. 
Peduncles less than 4 cm. long; outer corona filaments 

straight, not dilated. 

Fruit ellipsoidal, 6-ribbed, stipitate; leaves membran- 
ous; sepals thin-transparent P. tenella. 

Fruit globose or subglobose; leaves coriaceous; sepals 

rather fleshy. 

Leaves truncate or subtruncate along the upper 
margin; flowers 4-5 cm. wide; outer corona 

filaments 1 cm. long or more P. vespertilio. 

Leaves distinctly 3-lobed, the lobes subequal; flowers 
1.5-2 cm. wide; outer corona filaments much 
less than 1 cm. long P. obtusiloba. 

B. Bracts usually foliaceous, more than 1 mm. wide, ovate, 
lanceolate, or cordate (occasionally narrowly linear and very 
narrow in P. quadriglandulosa) , forming an involucre near 
base of flower, sometimes coalescent part way; flowers usually 
more than 5 cm. wide and highly colored; operculum not 
plicate. 

D. Calyx tube long-cylindric, longer than the sepals; corona 

1- or 2-ranked, usually reduced to tubercles. 

Leaves not lobed, lanceolate P. lanceolata. 

Leaves 3-lobed or 3-foliolate. 

Leaves 3-foliolate P. trifoliata. 

Leaves 3-lobed. 



FLORA OF PERU 95 

Bracts free to the base. 

Plant glabrous; stipules setaceous; leaves not more 
than 5 cm. long; corona a sinuate ridge. 

P. gracilens. 

Plant pubescent; stipules pinnatisect or deeply cleft; 
leaves fully 5 cm. long; outer rank of corona 

short-filamentose P. pinnatistipula. 

Bracts united part way. 

Stipules narrowly linear P. Matthewsii. 

Stipules oblong, ovate, or subreniform. 
Flowers violet, the tube not more than 3.5 cm. long; 

plant glabrous P. glaberrima. 

Flowers red or pink, the tube more than 3.5 cm. 

long; plants glabrous or pubescent. 
Leaves small, less than 4 cm. long and 6 cm. wide; 

peduncles very slender P. parvifolia. 

Leaves larger; peduncles stout. 

Involucre about two-thirds the length of the 

calyx tube P. macrochlamys. 

Involucre not two-thirds the length of the calyx 

tube, usually less than half. 
Stipules more than 2.5 cm. long; leaves thick- 
coriaceous, lustrous above; plant gla- 
brous P. anastomosans. 

Stipules less than 2.5 cm. long; leaves mem- 
branous or subcoriaceous, rarely coria- 
ceous; plants usually pubescent. 
Petioles 2- or 3-glandular at the middle, the 
glands at least 1.5 mm. long. 

P. mesadenia. 
Petioles 4-12-glandular, the glands less 

than 1.5 mm. long, scattered. 
Stem terete, yellow-pubescent; calyx tube 

glabrous P. mollissima. 

Stem angular, glabrous or grayish pubes- 
cent; calyx tube glabrous or pubes- 
cent P. mixta. 

D. Calyx tube campanulate to short-cylindric, usually much 
shorter than the sepals; corona in 3 or more ranks, or if 
2-ranked, the inner rank tubular. 



96 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves 3-foliolate. 

Plant glabrous; bracts lacerate-serrate P. Weberbaueri. 

Plant densely tomentose; bracts deeply laciniate-fimbriate. 

P. trisecta. 
Leaves simple, unlobed or variously lobed. 

Bracts and stipules deeply pinnatisect into filiform, gland- 
tipped divisions; plant often viscous, ill-scented. 
Seeds more than 6 mm. long and 3 mm. wide; lateral 
leaf lobes ascending; plant densely rufo-hirsute. 

P. vestita. 
Seeds smaller; lateral leaf lobes usually divergent. 

P. foetida. 
Bracts and stipules not pinnatisect. 

E. Operculum dependent, the margin not recurved. 
Stipules semi-ovate or subreniform, serrate, persistent; 
leaves 3-lobed; corona filaments of each rank distinct 
to base. 
Flowers red; peduncles rarely more than 8 cm. long; 

bracts free, or united only at base P. manicata. 

Flowers white; peduncles 10-15 cm. long; bracts united 

halfway P. peduncularis. 

Stipules linear or setaceous, deciduous; leaves lobed or 
unlobed; corona filaments of the inner rank united 
part way into a tube. 

Two outer ranks of corona filaments purple at apex, 
white at base; leaves never lobed; bracts ovate, at 

least 1 cm. wide P. coccinea. 

Two outer ranks of corona filaments red or white; leaves 
often polymorphic, unlobed to 3-lobed ; bracts usually 
narrower. 

Plant essentially glabrous. P. quadriglandulosa. 

Plant ferruginous- tomentose P. vitifolia. 

E. Operculum erect or horizontally spreading, not dependent. 
Calyx tube cylindric, subequal to the sepals; sepals and 
petals red; peduncles 10-16 cm. long, very slender. 

P. tarapotina. 

Calyx tube campanulate or broadly short-tubular, much 
shorter than the sepals; sepals and petals variously 
colored, red only in P. alata; peduncles usually much 
less than 10 cm. long. 



FLORA OF PERU 97 

Stem quadrangular, the angles winged; seeds more than 

5 mm. wide. 

Petioles 6-glandular; leaf nerves 10 or more to a side; 
stipules ovate to lance-ovate ... P. quadrangularis. 
Petioles 2-4-glandular; nerves about 8 to a side; stip- 
ules linear or linear-lanceolate P. alata. 

Stem terete or angular, the angles not winged; seeds not 

more than 5 mm. wide. 
Leaves palmately 5-9-lobed to below the middle (rarely 

a few 3-lobed leaves in P. caerulea). 
Stipules linear-subulate; bracts united in the lower 
part; leaf lobes serrulate .. P. serrato-digilata. 
Stipules semi-ovate; bracts free to the base; leaf 

lobes entire P. caerulea. 

Leaves unlobed or 3-lobed. 

Stipules linear or setaceous, often soon deciduous; 
leaves not lobed. 

Ovary glabrous P. nitida. 

Ovary sericeo-tomentose. 

Outermost rank of corona filaments shorter than 
the second rank; flowers solitary in the axils 

or in pairs P. laurifolia. 

Outermost rank of corona filaments as long as 
the second rank; flowers borne on short, 

axillary branches P. riparia. 

Stipules ovate to oblong. 

Bracts united, at least toward the base, more than 

2 cm. long. 

Petiolar glands liguliform, often appearing fili- 
form when dry, at least 3 mm. long. 

P. ligularis. 
Petiolar glands not liguliform, shorter, sessile or 

short-stipitate. 
Leaves minutely denticulate, usually 3-lobed ; 

corona 3-ranked P. triloba. 

Leaves entire at margin, never lobed; corona 

about 5-ranked P. tiliaefolia. 

Bracts free to the base, usually less than 2 cm. 
long; stipules semi-ovate to semi-oblong, 



98 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

attached on one side above the base, hence 
often appearing reniform. 
Plant glabrous throughout. 
Leaves suborbicular in general outline, 3- 
lobed at the apex; bracts imbricate, un- 
equal, one smaller than the others and 
borne a short distance below them. 

P. cuzcoensis. 
Leaves lanceolate or deeply 3-lobed; bracts 

verticillate. 

Leaves lanceolate, not lobed . . P. loretensis. 
Leaves 3-lobed. 

Leaves peltate at least 5 mm. from the 
lower margin, 10-20 cm. wide. 

P. spectabilis. 
Leaves not peltate. 

Sepals dorsally awned with a folia- 

ceous awn 5 mm. long or more; 

flowers 6-9 cm. wide . P. subulata. 

Sepals merely mucronulate; flowers 

3-5 cm. wide P. aristulala. 

Plant hispid-hirsute throughout. 

Ovary glabrous; bracts narrowly lanceolate or 
elliptic-lanceolate, less than 5 mm. wide, 
minutely glandular-denticulate. 

P. menispermifolia. 

Ovary pubescent; bracts ovate or ovate- 
lanceolate, more than 5 mm. wide, serrate 
or entire. 

Leaves ovate or ovate-lanceolate, entire or 
lobulate near base; petiolar glands not 
more than twice longer than thick. 

P. crassifolia. 

Leaves 3-lobed; petiolar glands more than 
twice longer than thick . . P. nephrodes. 

Passiflora adenopoda DC. Prodr. 3: 330. 1828. P. acerifolia 
Schlecht. & Cham. Linnaea 5: 89. 1830. 

Stem glabrate or hispidulous; stipules semi-orbicular, about 1.5 
cm. wide; petioles conspicuously biglandular toward the base, the 



FLORA OF PERU 99 

glands globose, 2-4 mm. in diameter, with a slender stipe 6-8 mm. 
long; blades 7-12 cm. long, 8-15 cm. wide, 3-5-lobed, deeply cordate, 
entire or denticulate, hispidulous; peduncles solitary or in pairs, 
bearing near the middle 3 lanceolate or oblong, lacerate bracts 7-10 
mm. long, 4-6 mm. wide; flowers up to 7 cm. wide, whitish or yellow- 
ish; sepals oblong-lanceolate, terminating in a conspicuous horn; 
petals linear-lanceolate, much shorter than the sepals; corona fila- 
ments in a single series, filiform, 1.5-2 cm. long, white, purple-banded; 
ovary densely brown-tomentose; fruit globose, 2-2.5 cm. in diameter; 
seeds reticulate. 

Junin: La Merced, 900 meters, 5561. Mexico to Venezuela: 
foothills and lower mountain slopes, 900 to 1,600 meters. 

Passiflora alata Dryand. Bot. Mag. 2: pi. 66. 1788. P. latifolia 
DC. Prodr. 3: 328. 1828. P. alata var. latifolia Mast. Trans. Linn. 
Soc. 27: 635. 1871. 

Plant glabrous throughout; stem stout, quadrangular, the angles 
winged; stipules linear or linear-lanceolate; petioles 2-4-glandular, 
the glands orbicular, sessile; blades ovate or ovate-oblong, 8-15 cm. 
long, 7-10 cm. wide, entire or minutely denticulate, penninerved, 
the principal secondary nerves 7 or 8 to a side, the upper ones distant; 
bracts ovate, about 1.5 cm. long; flowers 10-12 cm. wide, crimson 
or carmine, the corona filaments variegated with red, white, and 
purple; sepals and petals oblong; corona 4-ranked; fruit obovoid or 
pyriform, 8-10 cm. long. 

Loreto: Iquitos, 100 meters, Killip & Smith 27468. Without 
locality: Dombey (or Pavon?) 738, type of P. latifolia. Perhaps intro- 
duced into Peru from eastern Brazil, where apparently indigenous. 

Passiflora amalocarpa Barb. Rodr. Vellosia 1: 29. pi. 12. 1888; 
ed. 2. 1: 25. 1891; 3, pt. I: pi. 12. 1891. 

Plant glabrous throughout; stipules narrowly linear, subfalcate; 
petioles glandless; blades membranous, transversely oblong or trans- 
versely linear-oblong in general outline, 3-lobed or sometimes 
truncate at upper margin, 1-4 cm. along midnerve, 3.5-6.5 cm. along 
lateral nerves, 7-12 cm. wide, the middle lobe usually well developed, 
truncate or rounded; peduncles solitary, up to 5 cm. long; bracts 
linear-setaceous; flowers 3.5-4 cm. wide, apparently greenish white; 
sepals linear-lanceolate; petals linear, about half as long as the 
sepals; corona 2-ranked, the outer filaments filiform, subequal to the 
petals, the inner narrowly linear, 2-3 mm. long, capitate and emargi- 



100 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nate; operculum closely plicate; ovary fusiform; fruit fusiform, 
3.5-4 cm. long, 1-1.5 cm. in diameter; seeds transversely sulcate. 

Loreto: Lower Rio Nanay, Williams 526. Known otherwise 
only from the type locality, Manaos, Brazil. 

Passiflora anastomosans (Lambert) Killip, Journ. Wash. 
Acad. Sci. 17: 428. 1927. Tacsonia anastomosans Lambert ex DC. 
Prodr. 3: 335. 1828. 

Plant essentially glabrous throughout; stipules semi-ovate, about 
3 cm. long, aristulate; petioles 2-4-glandular near the apex, the 
glands sessile; blades 7-9 cm. long, 7-12 cm. wide, 3-lobed about 
to the middle, callous-serrate, conspicuously reticulate-veined, thick- 
coriaceous, very smooth and lustrous above; peduncles 2.5-3 cm. 
long; bracts 6-7 cm. long, connate one-half to two-thirds their length; 
calyx tube cylindric, 8-9 cm. long; sepals and petals oblong-lanceo- 
late; corona minutely tuberculate; fruit ovoid-ellipsoidal. 

Huanuco: Pozuzo, Pavon, type. 

Passiflora aristulata Mast, in Mart. Fl. Bras. 13, pt. 1: 
570. 1872. 

Plant glabrous; stipules cordate or semicordate, aristulate; 
petioles filiform, biglandular near the middle; blades 3-lobed about 
to the middle, 3-4 cm. long, 4-6 cm. wide, the lobes broadly oblong- 
ovate, obtuse, subequal; bracts broadly ovate, about 1 cm. long, 
borne at base of flower; flowers 3-4.5 cm. wide, greenish white; calyx 
tube campanulate; sepals oblong, dorsally mucronulate; petals 
ovate-oblong; corona filaments light purple, in 4 series, the outer 2 
filiform, the inner ones narrowly linear, subclavate; operculum 
fimbriate in the upper half; ovary ovoid; fruit edible. 

San Martin: Tarapoto, Spruce 3988, type. Juanjui, Poeppig in 
1830. Loreto: Lower Rio Huallaga, 200 meters, Williams 5180. 
Rancho Indiana, Rio Marafion, 110 meters, Mexia 6424-- "Sandia." 

Passiflora auriculata HBK. Nov. Gen. & Sp. 2: 131. 1817. 
P. cinerea Poepp. & Endl. Nov. Gen. & Sp. 2: 57. pi. 177. 1838. 

Stem glabrous or finely pubescent; stipules filiform; petioles up 
to 2 cm. long, bearing close to the base 2 auricular appendages 
about 2 mm. wide; blades prevailingly ovate-lanceolate in general 
outline, 5-15 cm. long. 2-10 cm. wide, subentire or undulately or 
angulately 3-lobed, acuminate, glabrous above, glabrate or cinereous- 
pubescent beneath; peduncles in pairs, up to 1 cm. long; bracts 
setaceous; flowers 2-2.5 cm. wide; sepals narrowly oblong-lanceolate, 



FLORA OF PERU 101 

yellowish green or pale greenish; petals linear, about half as long as 
the sepals, white; corona 2-ranked, the outer filaments filiform, 
yellowish green, purple at the base, the inner capitellate, white; 
operculum closely plicate; ovary pilosulous; fruit globose, 1-1.5 cm. 
in diameter; seeds obovate, transversely grooved. 

San Martin: Tarapoto, Ule 9644- Loreto: Mainas, Poeppig 1790 
(type of P. cinerea), 2302. Yurimaguas, 135 meters, Killip & Smith 
27589, 28047, 28311, 29035; Williams 4226, 7873. Iquitos, 100 
meters, Klug 141; Mexia 6388. Rio Maranon, Tessmann 4933. 
Balsapuerto, 220 meters, Klug 2933. Junin: Puerto Yessup, 400 
meters, Killip & Smith 26336. Nicaragua to the Guianas and 
Amazonian Brazil and Bolivia. 

Passiflora bauhinifolia HBK. Nov. Gen. & Sp. 2: 132. 1817. 

Stem softly pubescent; stipules linear-subulate, subfalcate, 
purplish; petioles glandless; blades oblong or ovate-oblong in general 
outline, 5-9 cm. long, 4-6.5 cm. wide, shallowly lobate, glabrescent 
above, sparingly or densely appressed-pilosulous beneath; peduncles 
solitary or in pairs; bracts linear-setaceous, purplish; sepals oblong- 
lanceolate, about 1 cm. long; petals ovate-lanceolate, half as long 
as the sepals, white; corona filaments in 2 poorly marked series, 
the outer narrowly liguliform, almost filiform, subtrigonous, the 
inner capillary; operculum plicate; ovary densely villous; fruit glo- 
bose, about 1 cm. in diameter; seeds transversely 6- or 7-sulcate. 
-F.M. Neg. 16521. 

Cajamarca: Huambo, Raimondi 2233. Ecuador. 

Passiflora caerulea L. Sp. PI. 959. 1753. 

Plant glabrous and often glaucous throughout; stipules semi- 
ovate; petioles 2-6-glandular, the glands stipitate; blades palmately 
5 (rarely 3, 7, or 9) -lobed nearly to the base, the lobes linear-oblong 
to broadly ovate-oblong, up to 10 cm. long, usually obtuse, entire; 
bracts broadly ovate, 1.5-2.5 cm. long, borne close to the flower 
base; flowers up to 10 cm. wide, white or pinkish; calyx tube cup- 
shaped, the sepals and petals oblong; corona filaments in 4 series, 
those of the 2 outer radiate, from a half to as long as the petals, blue 
at the apex, white at the middle, purple at the base, the inner fila- 
ments much shorter; operculum filamentose part way; fruit ovoid 
or subglobose, about 4 cm. in diameter; seeds coarsely reticulate. 

Lima: Lima Botanical Garden, Killip & Smith 21530. Tacna: 
Tacna, cultivated, Rusby 489. Without locality: Dombey 740. A 
native of east-central Brazil to Argentina, widely cultivated. 



102 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Passiflora Candollei Tr. & Planch. Ann. Sci. Nat. se"r. 5. 17: 161, 
footnote. 1873. P. lunata Juss. ex DC. Prodr. 3: 331. 1828, not 
P. lunata J. E. Sm. or Poepp. & Endl. 

Stem usually finely pulverulent; stipules linear-subulate, falcate, 
coriaceous; petioles glandless; blades 5-10 cm. along midnerve, 7-15 
cm. along lateral nerves, 7-15 cm. wide, bilobed (lobes lanceolate, 
acuminate, the sinus broadly lunate, with an intermediate lobe some- 
times present, or the upper margin subtruncate), bright green, lus- 
trous, and glabrous above, dull and finely pulverulent beneath ; bracts 
setaceous; flowers 3-5 cm. wide; sepals broadly oblong, fleshy, white 
within; petals ovate-oblong, white, pink-tinged; corona 2-ranked, 
the outer filaments 8-10 mm. long, sub trigonous, yellow, the inner 
filiform, much shorter, minutely capitellate, light green; operculum 
closely plicate; ovary white-sericeous; fruit globose, 1.5-2 cm. in 
diameter; seeds transversely sulcate. 

Amazonas: Chachapoyas, Malhews. Loreto: Iquitos, 100 meters, 
Killip & Smith 27143, 27438; Williams 8070, 8180. Rio Itaya, 
Kittip & Smith 29375, 29503, 29693, 29734, 29735; Tessmann 5273. 
Mishuyacu, King 1306, 1581, 2521. Mouth of Rio Santiago, Tess- 
mann 4942. Lower Rio Huallaga, Williams 4175; Kittip & Smith 
27825. Fontaleza, King 2788. Rio Paranapura, King 3945. 
Huanuco: Pampayacu, 1,100 meters, 5123. Rio Cayumba, 1,100 
meters, Mexia 8213. Chinchao, Ruiz & Pavon. Junin: La Merced, 
700 meters, Kittip & Smith 23517. Colonia Perene", 680 meters, 
Kittip & Smith 25000, 25423. Puerto Yessup, 400 meters, Killip 
& Smith 26308. Puerto Bermudez, 375 meters, Killip & Smith 
26607. Department uncertain: Hacienda Chalhuapuquio, Stevens 
146. Without locality: Dombey 742, type; Pavon. Known also from 
a single Bolivian collection. 

Passiflora cauliflora Harms, Verh. Bot. Ver. Brandenb. 48: 
185. 1907. 

Scandent shrub, glabrescent throughout except the ovary; leaves 
oblong, 15-18 cm. long, 6-10 cm. wide, obtusely acuminate or acute, 
entire, coriaceous, with a dark, thickened band at the margin; flowers 
"brown-yellow" (Klug), borne in short axillary fascicles; calyx tube 
cylindric, 1.5-2 cm. long; sepals and petals oblong, 1.5-2.5 cm. long; 
corona filaments in 3 series, the outermost subdolabriform, nearly 
1.5 cm. long, the others narrowly linear, much shorter; operculum 
erect, cleft in the upper third into 5 linear segments; ovary narrowly 
obovoid, softly ferruginous-tomentellous. F.M. Neg. 16522. 



FLORA OF PERU 103 

San Martin: Zepelacio, 1,600 meters, Klug 3469. Loreto: Cerro 
de Escalera, near Tarapoto, 1,300 meters, Ule 6679, type. Mouth 
of Rio Santiago, Tessmann 4588. Recently collected also at Manaos, 
Brazil, by A. Ducke. 

Passiflora cirrhipes Killip, Field Mus. Bot. 19: 522. 1938. 

Woody vine, glabrous throughout except the ovary; leaves ovate 
or oblong-ovate, 5-9 cm. long, 4-5.5 cm. wide, sharply acuminate, 
membranous; peduncles slender, bifurcate, terminating in a reduced 
or a well-developed tendril, flowers white; calyx tube cylindric- 
campanulate, up to 1 cm. long; sepals linear-oblong, about 1.5 cm. 
long; petals spatulate; corona filaments in 3 series, erect, the outer- 
most subdolabriform, yellow; operculum fimbriate about halfway; 
ovary rufo-tomentose. 

San Martin: Zepelacio, near Moyobamba, 1,100-1,600 meters, 
Klug 3883, type. 

Passiflora coccinea Aubl. PI. Guian. 2: 828. pi 324. 1775. 

Plant rufo-puberulent or rufo-tomentose nearly throughout; 
stipules narrowly linear, entire or minutely glandular-serrulate; 
petioles glandless, or biglandular at the base; blades oblong, 6-14 
cm. long, 3-7 cm. wide, not lobed, subcordate, duplicate-serrate or 
crenate, glabrous or sparingly pubescent above, ferruginous- or 
cano-tomentose beneath; peduncles up to 8 cm. long; bracts ovate, 
up to 6 cm. long, 1-3.5 cm. wide, free to base, coriaceous, crenate or 
sharply serrate, usually glandular at the margin, reddish; flowers 
scarlet or red; calyx tube short-cylindric-campanulate, up to 2 cm. 
long; sepals linear-lanceolate, 3-5 cm. long; petals linear; corona in 
3 series, the 2 outer consisting of subulate filaments about 1 cm. long, 
purple above, pink or white below, the inner rank tubular, fila- 
mentose only at the margin, white; ovary yellowish-tomentose; fruit 
subglobose or ovoid, about 5 cm. in diameter; seeds minutely 
reticulate. 

San Martin: Lamas, 840 meters, Williams 5069, 6379. Loreto: 
Masisea, 275 meters, Killip & Smith 26851. Yurimaguas, 135 
meters, Killip & Smith 27574, 27995, 29062; Williams 4745, 7821. 
Lower Rio Huallaga, 155-210 meters, Williams 3894, 3917, 4145. Rio 
Maranon Valley, 150 meters, Killip, Smith, & Dennis 29186, 29214. 
Rio Pastaza, Tessmann 3787. Rio Ucayali, Huber 1411, I486; Tess- 
mann 3187. Junin: Puerto Yessup, 400 meters, Killip & Smith 
26290. Puerto Bermudez, 375 meters, Killip & Smith 26673. Cahua- 
panas, 340 meters, Killip & Smith 26820. Ayacucho: Kimpitiriki, 



104 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

400 meters, Killip & Smith 22909. Cuzco: Marcapata Valley, 1,200 
meters, Herrera 1155. Department uncertain: Pampa del Sacra- 
mento, Castelnau in 1847. Without locality: Weberbauer 6761. 
Guianas to Peru, Bolivia, and northern Brazil. "Granadilla." 

Passiflora coriacea Juss. Ann. Mus. Hist. Nat. Paris 6: 109. 
pi. 39, f. 2. 1805. 

Plant essentially glabrous throughout; stipules narrowly linear; 
petioles 2 (rarely 4)-glandular usually below the middle, the glands 
sessile; blades transversely oblong, acute, rarely obtuse at the ends, 
3-7 cm. long (midnerve), 7-25 cm. wide, peltate, coriaceous; upper 
inflorescence a terminal raceme, the lower flowers solitary or in pairs 
in the leaf axils; bracts setaceous; flowers 2.5-3.5 cm. wide, yellowish 
green; sepals oblong-lanceolate; petals none; corona filaments in 
2 series, the outer ones filiform, the inner linear; operculum plicate; 
fruit globose, 1-2 cm. in diameter; seeds coarsely reticulate. 

San Martin: Tarapoto, Spruce 4532; Ule 6461. Chasuta, 260 
meters, King 3963. Alto Rio Huallaga, Williams 5517, 5751 
Junin: San Ramon, 1,000 meters, Killip & Smith 24907. Without 
locality: Haenke 1882. Mexico to Peru and northern Bolivia; 
reported once from British Guiana. "Uchu anquirisi." 

Passiflora costata Mast, in Mart. Fl. Bras. 13, pt. 1: 573. 1872. 

A subscandent shrub with few tendrils, the branches puberulent; 
leaves obovate to narrowly oblong-obovate, up to 25 cm. long and 
16 cm. wide, rounded and often emarginate at the apex, entire, 
penninerved, coriaceous, usually finely puberulent beneath; peduncles 
solitary or in pairs, 1-flowered; bracts narrowly linear; flowers 6-7 cm. 
wide, white, fragrant; calyx tube broadly campanulate or slightly 
funnel-shaped; sepals oblong-lanceolate, 3-4 times longer than the 
tube; petals oblong-spatulate; corona filaments in about 6 series, 
the outermost filaments 1.5-2 cm. long, orange, reddish at the 
middle, the others much shorter; operculum minutely denticulate; 
ovary narrowly ovoid, densely tomentose. 

Loreto: Iquitos, Tessmann 3675. Soledad, Tessmann 5286. 
Guianas and the Amazon Basin. 

Passiflora crassifolia Killip, Journ. Wash. Acad. Sci. 20: 
380. 1930. 

Stem densely villous with spreading, grayish hairs; stipules sub- 
reniform, aristate; petioles 4-6-glandular, scattered ; blades lanceolate 
or ovate-lanceolate, 10-18 cm. long, 6-12 cm. wide, unlobed or rarely 



FLORA OF PERU 105 

slightly lobate below the middle, obtuse, deeply cordate, thick-cori- 
aceous, glabrous, appressed-villous on the nerves and veins beneath ; 
bracts lanceolate, 1.5-2 cm. long, sharp-acuminate, entire; calyx 
tube tubular-campanulate; sepals oblong, dorsally awned; corona 
filaments in 3 or 4 series, the outermost about 7 mm. long; operculum 
filamentose in the upper half; fruit ovoid or ellipsoidal, 4-6 cm. 
long, villosulous; seeds coarsely reticulate. 

Junin: La Merced, 700 meters, 5371; Killip & Smith 23382, 
23434, type. Colonia Perene', 600 meters, Killip & Smith 25181. 

Passiflora cuzcoensis Killip, Journ. Wash. Acad. Sci. 20: 
379. 1930. 

Plant glabrous; stipules semi-oblong, 2-2.5 cm. long; petioles 
biglandular in the upper third, the glands minute, subulate; blades 
suborbicular in general outline, 5-7 cm. long, 5- or 7-nerved, 3-lobed 
at the apex, the lobes rounded; peduncles slender, up to 3.5 cm. long; 
bracts subimbricate, unequal (one larger than the others), cordate, 
2.5-3 cm. long, crenate-serrulate toward the apex; flowers about 
5 cm. wide; calyx tube campanulate; sepals and petals lance-oblong, 
the sepals awned dorsally just below the apex, the awn foliaceous; 
corona filaments in 2 series, the outer radiate, about 2 cm. long, the 
inner filiform, up to 2 mm. long; operculum denticulate. 

Cuzco: Marcapata Valley, near Chilechile, Weberbauer 7872, 
type. 

Passiflora ferruginea Mast, in Mart. Fl. Bras. 13, pt. 1: 
556. 1872. 

Stem rufo-tomentose; stipules narrowly linear; petioles up to 
5 cm. long, bearing near the middle 2 auriculate appendages; blades 
broadly ovate in general outline, 8-20 cm. long, 5-12 cm. wide, 
subangulately 3-lobed, rounded at the apex, sinuate-dentate or 
sinuate-denticulate, rufo-tomentose beneath; bracts setaceous; 
flowers 2.5-3 cm. wide, otherwise similar to those of P. auriculata; 
ovary sericeo-villous; fruit broadly ovoid, about 3.5 cm. long, 2.5-3 
cm. in diameter; seeds broadly obcordate, transversely grooved. 

San Martin: Tarapoto, Spruce 4901, type. Juanjui, Alto Rio 
Huallaga, 400-800 meters, King 4159, 4299. 

Passiflora foetida L. Sp. PL 959. 1753. 

Stipules semi-annular about the stem, deeply cleft into filiform, 
gland-tipped segments; petioles glandless, though often with gland- 
tipped hairs; blades prevailingly 3-lobed; bracts involucrate, 2-4- 



106 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pinnatifid or -pinnatisect, the segments filiform, gland-tipped; 
flowers 2-5 cm. wide, white or pink (in Peruvian plants), the corona 
filaments white or banded with purple and white, in several series, 
those of the 2 outer series filiform, about 1 cm. long, the inner ones 
capillary, 1-2 mm. long; operculum minutely denticulate; fruit 
globose; seeds ovate-cuneiform, obscurely tridentate, coarsely reticu- 
late at the center of each face. 

An herbaceous, ill-scented vine, highly variable in the degree 
and the nature of the indument, the shape of the leaves, the lacinia- 
tion of the bracts, and the size and color of the flowers. The species 
is widely distributed in tropical America and has been introduced 
into many parts of the Old World. Several varieties may be recog- 
nized, of which four, including typical P. foetida, have been reported 
from Peru. These may be keyed as follows: 

Ovary pubescent; fruit sparingly to densely pubescent. 

Stem, petioles, and peduncles hirsute with spreading, rather stiff, 
yellowish or brownish hairs averaging more than 1.5 mm. 
long P. foetida (typical). 

Stem, petioles, and peduncles softly pilosulous, often viscous, the 
hairs averaging less than 1.5 mm. long var. gossypifolia. 

Ovary glabrous. 

Bracts small, bipinnatisect or tripinnatisect, the segments straight, 
or at least not closely interwoven var. hirsuta. 

Bracts very large, tripinnatisect or quadripinnatisect, the segments 
closely interwoven var. hispida. 

The typical form is known in Peru only from Juan Guerra, San 
Martin, 720 meters (Williams 6843). 

Var. gossypifolia (Desv.) Mast. 

Piura: Piura, Spruce 64-58; Gaudichaud. Talara, Haught 69. 
Parinas Valley, Haught 209. Libertad: Pacasmayo, Forbes in 1912. 
Lima: Chosica, 1,000 meters, 523. Lima, Cuming 1046; Mathews 
408. Huanuco : Maria del Valle, 2,100 meters, 4954. Huancavelica : 
Mantaro Valley, below Colcabamba, 1,800-1,900 meters, Weber- 
bauer 6456. Without locality: Dombey 737; Ruiz & Pawn; Maclean; 
Nee: Wilkes Expedition. 

Var. hirsuta Mast. (P. Baraquiniana Lemaire). 

San Martin: Tarapoto, 750 meters, Williams 5413. Loreto: 
Contamana, 150 meters, Killip & Smith 26870. Iquitos, 100 meters, 
Killip & Smith 27093; King 959, 1483. Yurimaguas, 135-200 



FLORA OF PERU 107 

meters, Kittip & Smith 27828; Williams 5069, 7847; Poeppig 2173. 
Lower Rio Huallaga, Williams 3962, 3977, 5112. Rio Ucayali, Tess- 
mann3091, 5464- 

This variety is confined to the Amazon Basin of Peru and Brazil. 

Var. hispida (DC.) Killip. 

Tumbez: Hacienda La Choza, Weberbauer 7690. 

Passiflora foetida is known in Peru as "bedoca" and "purupuru." 

Passiflora frutescens Ruiz & Pavon ex Killip, Field Mus. Bot. 
19: 527. 1938. 

Erect shrub or tree, the branchlets finely puberulous; leaves 
ovate, oblong-ovate, or oblong-obovate, 20-25 cm. long, 12-14 cm. 
wide, obtuse and emarginate at the apex, cordulate, entire, penni- 
nerved, glabrous above, glaucescent and puberulent beneath; 
peduncles once-furcate; calyx tube funnel-shaped, about 1 cm. long; 
sepals and petals linear-oblong, 3-3.5 cm. long, white, densely red- 
spotted; outer corona filaments subdolabriform, about 1.5 cm. long, 
bearing just above the middle a knoblike projection. 

Huanuco: Chinchao, Ruiz & Pavon (No. 247, Fl. Peruv. Chil. 
ined.), type. 

Passiflora glaberrima (Juss.) Poir. in Lam. Encycl. Suppl. 2: 
844. 1812. Tacsonia glaberrima Juss. Ann. Mus. Hist. Nat. Paris 6: 
394. 1805. 

Plant glabrous throughout; stipules semi-ovate, about 1 cm. 
long, callous-serrate; petioles conspicuously 2-4-glandular at the 
apex; blades coriaceous, 3-5 cm. long, 4.5-7 cm. wide, 3-lobed four- 
fifths their length, the lobes ovate, acute, sharply dentate, the basal 
ones horizontally divaricate or slightly reflexed; peduncles about 
1.5 cm. long, slender; bracts about 1.5 cm. long, connate below the 
middle; flowers violet; calyx tube cylindric, 3.3 cm. long; sepals and 
petals oblong, 1-1.5 cm. long; corona reduced to a low, etuberculate 
ring; ovary ellipsoidal. 

Piura: Paramo de Guamani, Humboldt & Bonpland, type. 

This is the earliest described species of a small group of tac- 
sonias with blue, violet, or magenta flowers, and with conspicuous 
petiolar glands. The group is widespread in Colombia and Ecuador. 

Passiflora gracilens (Gray) Harms in Engl. & Prantl, Pflanzen- 
fam. 3. 6a: 91. 1893. Tacsonia gracilens Gray, U. S. Expl. Exped. 
639. 1854. T. boliviana Rusby, Mem. Torrey Club 3, pt. 3: 37. 1893. 
Passiflora boliviana Harms, Repert. Sp. Nov. 19: 28. 1923. 



108 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plant essentially glabrous throughout; stipules setaceous; petioles 
very slender, glandless or with 2 small glands at the apex ; blades 1-4 
cm. long, 2-5 cm. wide, minutely serrulate or subentire, 3-lobed 
nearly to the base, the lobes narrowly ovate-oblong, the basal ones 
divaricate; bracts ovate, 5-8 mm. long, minutely serrulate or entire, 
free to the base, often early deciduous; calyx tube narrowly cylindric, 
1.5-2 cm. long, 3-5 mm. in diameter (small for tacsonias), green at 
base, pink above without, bluish green within; sepals and petals 
oblong-lanceolate, 8-20 mm. long, bright pink; corona reduced to a 
shallowly sinuate, purplish ring; fruit ovoid, 2-3.5 cm. long; seeds 
cuneate, reticulate. 

Huanuco: Huariaca, Mathews 915, type. Junin: Ingahuasi, 
Mantaro Canyon, 3,150 meters, Killip & Smith 22177. Huanca- 
velica: Iscuchaco, Mantaro Valley, Weberbauer 5679; Raimondi 
11585. Cuzco: Herrera 487. Paucartambo, 3,300-3,500 meters, 
Balls 6682; Pennell 14170. Colquipata, 3,300 meters, Pennell 13791 , 
Hacienda Ccapana, 3,450 meters, Herrera 1061. Hacienda Fanccac, 
2,760 meters, Herrera 2108. Oropeza Valley, 3,400 meters, Herrera 
2599. Hacienda Cutija, 2,500 meters, Bues (Herrera 2126). Huasao, 
Herrera 3101. Ollantaitambo, 2,800 meters, Herrera 3342. Uru- 
bamba Valley, Herrera 1660, 2243. Calca, Vargas 154. Also in the 
mountains of Bolivia. "Jukucha-jampajhuai," "pichincho-jampa- 
jhuai" (Inca). 

Passiflora heterohelix Killip, Field Mus. Bot. 19: 563. /. 2a. 
1938. 

Plant glabrous throughout; tendrils slender, of 2 forms, one 
axillary on the main stem, the other on one of each pair of peduncles; 
stipules setaceous; petioles biglandular just below the apex; blades 
oblong or oblong-lanceolate, 4-11 cm. long, 1.5-5 cm. wide, cus- 
pidate-acuminate, entire, penninerved, coriaceous; peduncles 1- 
flowered, in pairs, one simple, ecirrhose, the other bifid, with one 
branch floriferous and the other a tendril; bracts narrowly ovate, 
2-3 cm. long, free to the base; calyx tube barely 1 mm. long; sepals 
elliptic-ovate, about 1.5 cm. long; petals slightly shorter than the 
sepals; corona consisting of a few linear filaments 2-3 mm. long, 
apparently in a single series; ovary narrowly ellipsoidal. 

Loreto: Tarapoto, 750 meters, Williams 5902, type. This may 
represent an undescribed genus; at least, it has no close relative in 
Passiflora. Final decision must await the collecting of better 
developed flowers. 



FLORA OF PERU 109 

Passiflora lanceolata (Mast.) Harms, Bot. Jahrb. 18: Beibl. 
46: 11. 1894. Tacsonia lanceolata Mast, in Mart. Fl. Bras. 13, pt. 1: 
536. 1872. Passiflora acutissima Killip, Journ. Wash. Acad. Sci. 17: 
428. 1927. 

Stem pilosulous; stipules lanceolate, 1-1.5 cm. long, deeply pin- 
natisect; petioles minutely biglandular at the apex; blades lanceolate, 
4-8 cm. long, 1-1.5 cm. wide, attenuate-acuminate, subrevolute, 
coriaceous, glabrous; peduncles about 5 cm. long; bracts lance- 
olate, 1.5-2 cm. long, free to the base, laciniate; calyx tube cylindric, 
7-8 cm. long, glabrous; sepals oblong, 3-3.5 cm. long, dorsally aris- 
tate; petals similar and subequal to the sepals; corona apparently 
tuberculiform ; ovary glabrous. 

Peru: Without locality, Mathews 1252, type. The nearest rela- 
tives of this are Colombian or Venezuelan species. 

Passiflora laurifolia L. Sp. PI. 956. 1753. 

Plant glabrous except the ovary and bracts; stipules narrowly 
linear; petioles biglandular just below the apex; blades oblong or 
ovate-oblong, 6-12 cm. long, 3.5-8 cm. wide, rounded to subacute, 
entire, thick-coriaceous; bracts ovate-oblong, 2.5-4 cm. long, glandu- 
lar-serrate toward the apex, puberulent; flowers 5-7 cm. wide; calyx 
tube cylindric-campanulate, up to 1 cm. long; sepals and petals 
oblong, red or purplish red ; corona filaments banded with red, blue, 
violet, or purple, and white, in 6 series, the outermost about 2 cm. 
long, the second series, 3-4 cm. long, those of the succeeding series 
much shorter; operculum minutely denticulate; ovary sericeo-tomen- 
tose; fruit ovoid, 5-8 cm. long, edible; seeds finely reticulate. 

Loreto: Tarapoto, 360 meters, Ule 6337, 6545. Huanuco: 
Chicoplaya, Ruiz & Pavon. Common in the West Indies and the 
Guianas; rare in Venezuela and northern Brazil. "Granadilla." 

Passiflora leptoclada Harms, Notizbl. Bot. Gart. Berlin 9: 
979. 1926. 

Plant glabrous throughout except the ovary; stipules linear- 
setaceous; petioles glandless; blades membranous, transversely 
elliptic or transversely oblong in general outline, truncate or shallowly 
3-lobed at the apex, 1.5-4 cm. along the midnerve, 2.5-4 cm. along 
the lateral nerves, 4-7 cm. wide, the lobes obtuse or rounded; 
peduncles in pairs; bracts subulate; flowers up to 3 cm. wide, ap- 
parently greenish white; sepals narrowly oblong; petals similar to 
the sepals but shorter; corona 2-ranked, the outer filaments filiform, 
subequal to the sepals, the inner linear, 3-4 mm. long, broadly 



110 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

capitate and emarginate; operculum closely plicate; ovary ovoid, 
densely cano-tomentulous; fruit about 1.5 cm. in diameter; seeds 
transversely sulcate. Close to the widely distributed P. misera, 
but having a dense indument on the ovary. F.M. Neg. 16541. 
Loreto: San Isidro, Tessmann 4969, type. Soledad, Rio Itaya, 
110 meters, Kittip & Smith 29775. Mishuyacu, 100 meters, Kittip 
& Smith 29989. La Victoria, Williams 2737. Puerto Arturo, 
Williams 5252. "Mashu sisa." 

Passiflora ligularis Juss. Ann. Mus. Hist. Nat. Paris 6: 113. 
pi. 40. 1805. 

Plant glabrous throughout; stipules ovate-lanceolate or oblong- 
lanceolate, up to 2.5 cm. long; petioles up to 10 cm. long, bearing 
4-6 scattered, liguliform or filiform glands 3-10 mm. long; blades 
broadly ovate, 8-15 cm. long, 6-13 cm. wide, abruptly acuminate, 
deeply cordate, entire; bracts ovate-lanceolate, connate toward the 
base; flowers 6-9 cm. wide; sepals ovate-oblong, white within; 
petals oblong, white or pinkish white; corona 5-7-ranked, the 2 
outer rows of filaments equaling the petals, radiate, blue at the apex, 
banded with white and reddish purple; fruit ovoid, 6-8 cm. long, 
4-5 cm. in diameter, the pericarp parchment-like, the pulp white, 
edible; seeds narrowly obcordate, reticulate. 

Lima: Lima, cultivated, Killip & Smith 21529; Rose 18776 
Huanuco: Huanuco, Ruiz & Pavon. Cuchero, Poeppig 1695. 
Junin: Huacapistana, 1,800 meters, Killip & Smith 24330. Tarma, 
cultivated, Killip & Smith 21947. Ayacucho: Aina, 800 meters, 
Killip & Smith 22681. Cuzco: Torontoy, Urubamba Valley, 2,400 
meters, Cook & Gilbert 814- Department uncertain: Carabaya, 
Weddell 4777. Without locality: Dombey 739, type; Pavon. The 
common "granadilla" of western South America, extending to 
central Mexico. Its pulp is used for ices and cooling drinks. 

Passiflora Lobbii Mast, in Mart. Fl. Bras. 13, pt. 1: 553. 1872. 
P. obtusiloba var. glandulifera Harms, Repert. Sp. Nov. 19: 25. 1923. 

Stem finely pilosulous or glabrous; stipules linear-setaceous; 
petioles biglandular near the base, the glands minute, sessile; blades 
2-3 cm. long, 4.5-8 cm. wide, 3-lobed one-third to one-half their 
length (lobes lance-ovate, obtuse, subequal), shallowly cordate, 
entire, coriaceous, glabrescent; bracts setaceous; flowers 1.5-2 cm. 
wide; sepals oblong-lanceolate, purplish without, grass-green within; 
petals linear-oblong, grass-green; corona 3-ranked, the outermost 
filaments filiform, purple and light green, the others pink or purple, 



FLORA OF PERU 111 

capitellate, the tip green; ovary glabrous; fruit globose, about 1.5 cm. 
in diameter; seeds closely reticulate. 

Ancash: Between Samanco and Caraz, Weberbauer 3165, type of 
P. obtusiloba var. glandulifera. Huanuco: Ambo, 2,400 meters, 
2415. Ayacucho: Pampalca, 3,200 meters, Killip & Smith 23241- 
Huanta, 2,800 meters, Killip & Smith 23322. Department uncer- 
tain: Caururu, Savatier 1436. Without locality: Lobb, type; Gay; 
MacLean; Mathews. The type bears the inscription "Lobb, Co- 
lumbia." It is certain that many Lobb specimens so labeled actually 
came from Peru, and presumably this one did. 

Passiflora loretensis Killip, Journ. Wash. Acad. Sci. 21: 
349. 1931. 

Plant essentially glabrous; stipules semi-oblong, 1.3-1.8 cm. long; 
petioles up to 2 cm. long, bearing 2 pairs of subulate glands, one near 
the apex, the other near the middle; blades lanceolate, 10-12 cm. 
long, 5.5-6 cm. wide, acuminate, cordulate and subpeltate at the 
base, entire, coriaceous; bracts cordate-ovate, 2-2.5 cm. long; flowers 
about 5 cm. wide, pink(?); calyx tube broadly campanulate, the 
sepals and petals oblong, the sepals awned; corona filaments in 5 
series, filiform, the 2 outer radiate, about 1.5 cm. long, pale pink; 
operculum fimbrillate to the middle; fruit globose, about 5 cm. in 
diameter; seeds coarsely reticulate. 

Loreto: La Victoria, Amazon River, Williams 3086, type. 

Passiflora macrochlamys Harms, Bot. Jahrb. 42: 130. 1908. 

Plant glabrous nearly throughout; petioles 5-glandular; blades 
coriaceous or subcoriaceous, denticulate-serrate, 5-6 cm. long, 
6-9 cm. wide, 3-lobed more than half their length, the middle lobe 
broadly ovate, 3-3.5 cm. wide at the base, abruptly acuminate, the 
lateral lobes widely divaricate; peduncles 3-5.5 cm. long; bracts 
connate two- thirds their length, about 5 cm. long; calyx tube cylin- 
dric, 6-6.5 cm. long, the sepals and petals oblong; corona minutely 
tuberculate. F.M. Neg. 16544. 

Huanuco: Monzon, 2,000-2,500 meters, Weberbauer 3541, type. 
Without locality: Ruiz & Pavon. 

Passiflora manicata (Juss.) Pers. Syn. PI. 2: 221. 1806. Tac- 
sonia manicata Juss. Ann. Mus. Hist. Nat. Paris 6: 393. pi. 59, f. 2. 
1805. 

Stem angulate; stipules semi-ovate, coarsely dentate; petioles 
4-10-glandular; blades 4-8 cm. long, 5-9 cm. wide, rarely larger, 



112 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3-lobed to the middle, callous-serrulate or serrate, tomentose beneath, 
the lobes ovate or ovate-oblong; peduncles up to 8 cm. long; bracts 
free or united toward the base, ovate, 2-3 cm. long; calyx tube 
urceolate-campanulate, 1.5-2 cm. long, green without; sepals oblong- 
lanceolate, green, pink- tinged without, scarlet within; petals scarlet; 
corona in 3 or 4 series, the outer 2 or 3 filamentose, the fila- 
ments 2-4 mm. long, blue; ovary obovoid, glabrous; fruit ovoid or 
subspherical, 3.5-5 cm. long, glabrous and lustrous; seeds finely 
reticulate. 

Cajamarca: Hacienda La Tajona, Weberbauer 4051. Western 
Venezuela and Colombia to Peru, sometimes cultivated. 

Passiflora Matthewsii (Mast.) Killip, Journ. Wash. Acad. Sci. 
17: 428. 1927. Tacsonia Matthewsii Mast, in Mart. Fl. Bras. 13, 
pt. 1:539. 1872. 

Stem tomentellous; stipules narrowly linear, 2.5-3 mm. long; 
petioles 6-glandular; blades 5-6 cm. along the midnerve, 3.5-4 cm. 
along the lateral nerves, 4-6 cm. between the apices of the lateral 
lobes, coriaceous, glabrous above, densely grayish-tomentose beneath, 
3-lobed about to the middle, the lobes ovate-lanceolate or oblong- 
lanceolate, acute, serrulate; peduncles 1-2 cm. long, stout; bracts 
about 2.5 cm. long, connate to the middle; calyx tube cylindric, 
about 4 cm. long, tomentellous without; sepals and petals oblong, 
2-2.5 cm. long, rose; corona minutely tuberculate. 

Amazonas: Chachapoyas, Mathews, type. Libertad: Near Cerro 
Uruchalda, 3,350 meters, West 8175 (doubtfully referred here). 

Passiflora menispermifolia HBK. Nov. Gen. & Sp. 2: 137. 1817. 
P. villosa Dombey ex Tr. & Planch. Ann. Sci. Nat. se"r. 5. 17: 154. 1873, 
as synonym. 

Plant densely hispid-hirsute with light brown hairs nearly 
throughout; stipules subreniform, glandular-denticulate or subentire; 
petioles 2-4-glandular; blades broadly lanceolate or suborbicular in 
general outline, 10-16 cm. long, 8-13 cm. wide, angulately 3-lobed, 
cordate, remotely glandular-denticulate to dentate, the lobes acute 
or rounded, the middle one broadly ovate-deltoid; bracts narrowly 
lanceolate or elliptic-lanceolate, 1-2 cm. long, acuminate, glandular- 
denticulate; flowers up to 6 cm. wide, violet; sepals lance-oblong; 
petals linear-oblong; corona filaments in several series, the outermost 
filiform, about 2 cm. long, the others very dense, shorter; operculum 
filamentose in the upper half; ovary ovoid, glabrous; seeds coarsely 
reticulate. 



FLORA OF PERU 113 

Cajamarca: Between Tomependa and Jae"n de Bracamoras, 
Humboldt & Bonpland, type. Loreto: Yurimaguas, 135 meters, 
Killip & Smith 28705. Canchahuaya, Rio Ucayali, Huber 1408. 
Mishuyacu, 100 meters, Klug 1064. Balsapuerto, 220 meters, Klug 
3091. Huanuco: Chinchao, Ruiz & Pavon. Ayacucho: Aina, 900 
meters, Killip & Smith 22795. Cuzco: Urubamba Valley, 1,800 
meters, Cook & Gilbert 1085. Without locality: Dombey, type of 
P. villosa Dombey; Mathews 2074- Nicaragua to Peru. Also in 
upper Amazonian Brazil. 

Passiflora mesadenia Killip, Journ. Wash. Acad. Sci. 17: 427. 
1927. 

Stem subquadrangular, pilosulous; stipules subreniform, glandu- 
lar-dentate; petioles conspicuously biglandular at the middle, some- 
times with a third gland above or below this pair; blades 4-6 cm. 
long, 6-8 cm. wide, 3-lobed to the middle, subauricular or cordulate 
at the base, shallowly and irregularly dentate-serrate, glabrous above, 
pubescent beneath; peduncles 4-5.5 cm. long, stout; bracts 4-5 cm. 
long, united about half their length, glabrous; calyx tube cylindric, 
7-8 cm. long, pink, glabrous; sepals oblong, pink without, white 
within; petals oblong-spatulate, white; corona minutely tuberculate; 
ovary glabrous, pruinose. 

Huanuco: Vilcabamba, Rio Chinchao, 1,800 meters, 4960, type. 
Pampayacu, Kanehira 26 5a; Sawada P-l. 

Passiflora mixta L. f. Suppl. 408. 1781. P. longiflora Lam. 
Encycl. 3: 39. 1789. P. Tacso Cav. Diss. 10: 451. pi. 277. 1790. 
Tatsonia mixta Juss. Ann. Mus. Hist. Nat. Paris 6: 394. 1805. 
T. mixta var. longiflora DC. Prodr. 3: 335. 1828. 

Plant glabrous throughout or more usually pubescent on the 
under side of the leaves, the involucre, the calyx tube, and the ovary; 
stem angulate; stipules subreniform, callous-dentate or -serrate; 
petioles minutely 4-8-glandular; blades 5-10 cm. long, 6-17 cm. wide, 
3-lobed to or to slightly below the middle, the lobes ovate-oblong, acute 
or abruptly acuminate, coarsely or finely callous-serrate; peduncles 
stout, up to 6 cm. long; bracts united one-half to three-quarters their 
length into a tubulate-campanulate or nearly cylindric involucre 
2-5 cm. long; calyx tube cylindric, 8-11 cm. long, about 1 cm. in 
diameter, yellow-green to orange-red without, cream-color within; 
sepals oblong, 3-4 cm. long; petals pink to orange-red; corona short- 
tuberculate, deep lavender or purple; fruit ovoid, 4.5-6 cm. long, 
2-2.5 cm. in diameter; seeds reticulate. 



114 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Mito, 2,700 meters, 1615. Muna, 2,400 meters, 4315. 
Pillao, Ruiz & Pavdn. Cuzco: Lucumayo Valley, Cook & Gilbert 
1 348. Panticalla Pass, 3,600 meters, Cook & Gilbert 1 824. Urubamba 
Valley, 2,400 meters, Cook & Gilbert 828. Paucartambo, 3,000 meters, 
Herrera 2999. Ayacucho; Tambo, Weberbauer 5621. Puno: Sandia, 
Weberbauer 872. Without locality: Jussieu, type of P. longiflora 
and of P. Tacso; Dombey 746; Nee; Haenke 1957, 2040. Venezuela 
and Colombia to Peru and Bolivia, between 2,500 and 3,600 meters. 
Often cultivated. Fruit edible. "Tacso," "tumbo," "monte- 
turubo"; "xamppajrrai" (Quechua). 

Passiflora mollissima (HBK.) Bailey, Rhodora 18: 156. 1916. 
P. tomentosa Lam. Encycl. 3: 40. 1789(?). Tacsonia mollissima 
HBK. Nov. Gen. & Sp. 2: 144. 1817. Passiflora tomentosa var. 
mollissima Tr. & Planch. Ann. Sci. Nat. seY. 5. 17: 131. 1873. 

Stem terete, densely and softly yellow-villous; stipules subreni- 
form, 7-9 mm. long; petioles bearing 8-12 sessile glands; blades 5-1Q 
cm. long, 6-12 cm. wide, 3-lobed about two-thirds their length (lobes 
ovate or ovate-oblong, acute, the sinuses acute), sharply serrate- 
dentate, softty pubescent above, grayish- or yellowish-tomentose 
beneath; peduncles 2-6 cm. long; bracts 2.5-3 cm. long, united one- 
third to one-half their length; calyx tube 6.5-8 cm. long, olive-green, 
often red-tinged without, white within, glabrous; sepals oblong, 
2.5-3.5 cm. long; petals pink; corona reduced to a purple band with 
a few tubercles or crenulations; ovary sericeo-tomentose; fruit oblong- 
ovoid, 6-7 cm. long, 3-3.5 cm. in diameter, yellowish, softly pubes- 
cent; seeds reticulate. F.M. Neg. 16548. 

Huanuco: Huanuco, 3,200 meters, 2074. Pampayacu, Kanehira 
265. Junin: Tarma, 3,000-3,200 meters, Kittip & Smith 21868, 
21 942. Ocopa, 3,300 meters, Kittip & Smith 2201 1 . Carpapata, 3,200 
meters, Killip & Smith 24481. Huancayo, Valleriesta. Ayacucho: 
Ccarrapa, 1,200 meters, Killip & Smith 23236. Cuzco: Ollantai- 
tambo, 3,000 meters, Cook & Gilbert 270. Guispicanchi, Herrera 696. 
Pauccarccoto, 2,700 meters, Bues (Herrera 2127, 2128). Cuzco, 
Herrera 2943; Popenoe 1355. Arequipa: Arequipa, cultivated, Rose 
18979. Venezuela and Colombia to Peru and Bolivia, usually found 
between 2,000 and 3,000 meters; often cultivated. Fruit edible. 
"Tacso," "tintin," "tumbo," "tumbo del monte," "trompos." 

Passiflora morifolia Mast, in Mart. Fl. Bras. 13, pt. 1: 555. 1872. 

Stem sparingly hispidulous above; stipules semi-ovate, about 6 
mm. long and 3 mm. wide, long-acuminate; petioles biglandular near 
the apex, the glands thick-stipitate; blades 4-11 cm. long, 5-15 cm. 



FLORA OF PERU 115 

wide, 3-lobed to below the middle (lobes acute, the middle one ovate 
or ovate-lanceolate, usually narrowed at the base, the lateral ones 
divergent), deeply cordate, repandly dentate or denticulate, or sub- 
entire, membranous, minutely hispidulous above, minutely pilosulous 
beneath; peduncles solitary or in pairs, widely divaricate; bracts 
setaceous; flowers 2-3 cm. wide; sepals linear-oblong, white, red- 
mottled within; petals linear-lanceolate; corona filaments in a single 
series, filiform, 5-6 mm. long, white, banded with blue or violet; 
operculum plicate; ovary densely pilose; fruit globose, 2 cm. in 
diameter; seeds very slightly compressed, coarsely reticulate. 

Cuzco: Urubamba Valley, near Echarate, Weberbauer 7949. 
Without locality: Gay 941; Pavdn. Common in Argentina; rare in 
Bolivia, Guatemala, and Mexico. 

Passiflora nephrodes Mast. Bull. Torrey Club 17: 282. 1890. 

Plant densely hirsute throughout; stipules subreniform, 1.5-2 
cm. long, coarsely serrate-dentate; petioles 2- or 3-glandular near the 
middle; blades 6-18 cm. long, 5-9 cm. wide, 3-lobed, subcordate, 
denticulate, the middle lobe ovate or ovate-lanceolate, much longer 
than the deltoid-ovate lateral lobes; bracts ovate-lanceolate, 1-2 
cm. long, serrate; flowers 6-8 cm. wide; sepals oblong, dorsally 
awned; petals linear-oblong; corona filaments filiform, in several 
series, the outermost radiate, 2.5-3 cm. long, purple in the lower 
half, white in the upper, the succeeding filaments erect, 3-6 mm. 
long; operculum filamentose in the upper half; fruit ovoid, about 4 
cm. long; seeds reticulate. 

Junin: Yapas, Pichis Trail, 1,600 meters, Killip & Smith 25447. 
Cuzco: Lares Valley, Weberbauer 7920. Also in western Bolivia 
and Amazonian Brazil. 

L 

Passiflora nitida HBK. Nov. Gen. & Sp. 2: 130. 1817. 

Plant glabrous throughout; stipules linear-subulate; petioles 
biglandular at the apex; blades ovate-oblong, ovate-elliptic, or 
broadly ovate, 9-17 cm. long, 6-10 cm. wide, acute or acuminate, 
subentire, undulate-denticulate, or serrulate, usually becoming black- 
ish in drying; bracts oblong-ovate, about 3.5 cm. wide; flowers 9-11 
cm. wide; calyx tube campanulate; sepals oblong-lanceolate, white 
within; petals narrowly oblong, white; corona filaments in several 
series, the 2 outer subequal, 2-3.5 cm. long, white and pink-spotted 
at the base, banded with blue and white at the middle, white at the 
apex, the inner filaments shorter; operculum fimbrillate; fruit glo- 
bose, 3-4 cm. in diameter; seeds reticulate. 



116 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Iquitos, 100 meters, Killip & Smith 26909, 27158; King 
256, 828; Williams 1383, 1517; Tessmann 3703; Mexia 6488. Rio 
Nanay, Williams 1130. La Victoria, Williams 2800. Rio Rumiyacu, 
Raimondi 614- Gamitanacocha, Rio Mazan, 100-125 meters, J. M. 
Schunke 51. Colombia to the Guianas, south to Peru and central 
Brazil. "Granadilla," "puru puru." 

Passiflora obtusiloba Mast, in Mart. Fl. Bras. 13, pt. 1: 554. 
1872. P. Niorbo Planch. Ann. Sci. Nat. se>. 5. 17 : 156, footnote. 1873. 

Stem finely pilosulous or glabrescent below, densely pubescent 
toward the end; stipules setaceous; petioles glandless; blades coria- 
ceous, glabrous or nearly so, 1-2.5 cm. long, 2-5 cm. wide, 3-lobed 
about a third their length, the lobes subequal, rounded; peduncles 
in pairs; bracts setaceous; flowers 1.5-2 cm. wide; sepals narrowly 
oblong, yellowish green; petals linear-spatulate, light green; corona 
2-ranked, the outer filaments terete, about 4 mm. long, yellow-green, 
the inner filiform, 1.5-2 mm. long; operculum closely plicate; ovary 
glabrous. 

Junin: Tarma, 2,800-3,100 meters, Weberbauer 1735; Killip & 
Smith 21943. Palca, Stevens 40. Huasahuasi, Dombey 735 (type of 
P. Niorbo). Ayacucho: Quinua, Weberbauer 5 5 45. Without locality: 
Pavdn (type); Gay. "Niorbo." 

Passiflora parvifolia (DC.) Harms in Weberbauer, Pflanzenw. 
Per. And. 253. 1911. Tacsonia parvifolia DC. Prodr. 3: 335. 1828. 
Passiflora brachychlamys Harms, Notizbl. Bot. Gart. Berlin 10: 815. 
1929. 

Plant essentially glabrous throughout; stipules semi-ovate, 6-10 
cm. long, coarsely and remotely serrate; petioles minutely 3-7- 
glandular; blades small for the subgenus Tacsonia, 2-3 cm. long, 3-6 
cm. between the apices of the lateral lobes, 3-lobed to below the 
middle, the lobes ovate to ovate-lanceolate, 7-15 mm. wide, sharply 
serrulate; peduncles 3-6 cm. long, very slender; bracts 2.5-3 cm. 
long, connate at least to the middle, the tube subconical, the free 
portions ovate-lanceolate, acute; calyx tube cylindric, 6-9 cm. long, 
rose-green or rose-red; sepals and petals oblong, 2-2.5 cm. long, bright 
red; corona minutely tuberculate; ovary narrowly ellipsoidal. F.M, 
Neg. 24153. 

Libertad: Eastern base of Cerro Huaylillas, 3,900 meters, West 
8126. Huanuco: Huanuco, 3,500-4,000 meters, 2198; Weberbauer 
3329. Tambo de Vaca, 4,300 meters, 4405. Mito, 1823. Muna, 
Pearce in 1863. Yanano, 1,800 meters, 5743. Ayacucho: Tambo, 



FLORA OF PERU 117 

Pearce 309. Cuzco: Province of Paucartambo, 3,350 meters, Her- 
rera 484, type of P. brachychlamys. Hacienda Ccapana, Herrera 
588. Without locality: Ruiz & Pavon 533, type; Lobb (erroneously 
cited in Mart. Fl. Bras, as from Colombia). Endemic, Pearce's 35, 
from Ecuador, also cited as this, being P. cumbalensis. "Samppa- 
jhuai" (Cuzco), "jamppajhuai." 

Passiflora peduncularis Cav. Icon. 5: 15. pi. 426. 1799. Tac- 
sonia peduncularis Juss. Ann. Mus. Hist. Nat. Paris 6: 395. 1805. 
T. peduncularis var. Dombeyana DC. Prodr. 3: 334. 1828. T. Dom- 
beyana M. Roemer, Fam. Nat. Syn. 2: 197. 1846. 

Stem quadrangular, tomentulous; stipules semi-ovate, coarsely 
and irregularly serrate; petioles 3-4-glandular; blades deeply cordate, 
serrulate, subcoriaceous, glabrous above, tomentulous on the nerves 
and veins beneath, 3-lobed to below the middle, the lobes ovate, sub- 
equal, 3-6 cm. long, the lateral widely divergent; peduncles 10-15 
cm. long, stout; bracts 2-3 cm. long, united to above the middle; 
flowers white; calyx tube short-cylindric, 1.5-2 cm. long; sepals and 
petals oblong, 3-4 cm. long; corona in several series, the outermost 
consisting of very short, liguliform filaments, the others mostly of 
minute, filiform threads; ovary yellowish-tomentose; fruit spherical, 
3-4 cm. in diameter. 

Ancash: Ocros, Cajatambo, Weberbauer 2675. Lima: Rio Chil- 
lon, above Obrajillo, 3,000 meters, Pennell 14393. Moquehua: 
Carumas, 3,300 meters, Weberbauer 7345. Without locality: Nee, 
type; Pavon; Dombey, type of T. peduncularis var. Dombeyana; 
Wilkes Expedition; Mathews 480; Haenke 2118. Probably restricted 
to the higher mountains of western Peru, reports of its occurrence in 
Chile and Ecuador doubtless being erroneous. 

Passiflora pinnatistipula Cav. Icon. 5: 16. pi. 428. 1799. 
Tacsonia pinnatistipula Juss. Ann. Mus. Hist. Nat. Paris 6: 393. 
1805. T. micradena DC. Prodr. 3: 334. 1828. 

A rather stout vine with very showy flowers, the younger portions 
of the stem white- tomentose or white-lanate; stipules purplish, 5-7 
mm. long, 4-5 mm. wide, pinnatisect or palmately cleft into filiform 
divisions; petioles minutely 4-6-glandular; blades coriaceous, rugose 
and glabrous above, densely white- or grayish-lanate beneath, 
5-10 cm. long, 6-13 cm. wide, 3-lobed about four-fifths their length, 
the lobes lanceolate or oblong-lanceolate, acuminate, sharply serrate; 
peduncles up to 7 cm. long, the bracts free, ovate, 1-1.5 cm. long, 
remotely sharp-serrate, usually reddish purple without; calyx tube 



118 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cylindric, 4.5-5 cm. long, bright pink and usually cano-tomentose 
without, white within, blue-tinged toward the throat; sepals oblong, 
3-4 cm. long, greenish or pinkish without, white within; petals white, 
faintly blue-tinged; corona 2-ranked, the outer rank filamentose, 
1.5-2 cm. long, blue and white, the inner a ring of minute, deep purple 
tubercles; ovary white- tomentose; fruit subglobose, about 5 cm. in 
diameter; seeds reticulate. 

Junin: Tarma, 3,100 meters, Ruiz & Pavon, type of Tacsonia 
micradena; Killip & Smith 21938. Between Tarma and Palca, 
Weberbauer 1733. Ocopa, 3,300 meters, Killip & Smith 22012. 
Huancayo, 3,600 meters, Killip & Smith 22034; Ledig 5. Cuzco: 
Sacsaihuaman, Herrera 295. Paucartambo, 3,350 meters, Herrera 
486. Guispicanchi, 3,400 meters, Herrera 2586, 2605. Ollantai- 
tambo, 3,000 meters, Cook & Gilbert 475. Tinta, Cook & Gilbert 228. 
Probably a native of Peru or Chile but cultivated throughout the 
Andes for its edible fruit or as an ornamental. "Tin-tin," "puru- 
puru," "tacso." 

Passiflora Poeppigii Mast. Trans. Linn. Soc. 27: 630. 1871; 
in Mart. Fl. Bras. 13, pt. 1: 546. 1872. P. lunata Juss. sensu Poepp. 
& Endl. Nov. Gen. & Sp. 2: 58. pi. 178. 1838 (not P. lunata Juss.). 

Plant glabrous throughout; stipules subulate-falcate; petioles 
slender, glandless; blades transversely oblong in general outline, 
1.5-5 cm. along the midnerve, 2-6.5 cm. along the lateral nerves, 
3.5-10 cm. wide, repand-truncate at the upper margin, membranous; 
peduncles 4-6 cm. long, very slender; bracts setaceous; flowers 
1.5-2 cm. wide, white; sepals broadly ovate; petals none(?); corona 
2-ranked, the outer filaments filiform, 7-8 mm. long, the inner 
narrowly linear, much shorter; operculum plicate; ovary ovoid. 

Loreto: Yurimaguas, Poeppig D.2170, type. 

If this species is truly apetalous, as described by the authors, 
it is the only South American representative of its subgenus without 
petals. Authors have frequently overlooked very small petals and 
this may be the case here. It is impossible to decide the point 
because of the condition of the flowers of the type specimen. 

Passiflora punctata L. Sp. PI. 957. 1753. 

Plant glabrous throughout; stipules linear-falcate; petioles gland- 
less; blades transversely oblong in general outline, 2-5 cm. along 
the midnerve, 3-7 cm. along lateral nerves, 6-12 cm. wide, truncate 
and very shallowly 3-lobed at the apex or rather conspicuously 
bilobed, thin-membranous, glaucescent beneath; peduncles 5-8 cm. 



FLORA OF PERU 119 

long, very slender; bracts setaceous; flowers 2.5-4 cm. wide; sepals 
oblong-lanceolate, light yellow-green; petals similar, greenish white; 
corona 2-ranked, the outer filaments liguliform, falcate, dilated near 
the middle, yellow-green at the apex, purple to magenta at the 
center, white at the base, the inner filaments filiform, capitellate; 
operculum plicate; fruit ellipsoidal; seeds transversely 6-sulcate. 

Tumbez: Hacienda Chicama, 900 meters, Weberbauer 7637. 
San Martin: Tarapoto, 750 meters, Williams 5922. Libertad: 
Chinchin, Nee. Lima: Lima Botanical Garden, Killip & Smith 
21527; Nee. Barrana, Wawra 527. Cuzco: Santa Ana Valley, 
Herrera 941. Without locality: Dombey 736; Pavon. Probably 
native in Ecuador and northern Peru, this species is frequently 
cultivated in western South America. "Norbo." 

Passiflora pyrrhantha Harms, Notizbl. Bot. Gart. Berlin 9: 
977. 1926. 

Scandent shrub with a few tendrils, the branchlets puberulent; 
leaves ovate or ovate-oblong, 10-16 cm. long, 6-10 cm. wide, 
acuminulate, remotely denticulate in the upper half, subcoriaceous, 
glabrous; flowers fire-red, in racemes at the end of long peduncles 
which occasionally terminate in a tendril ; calyx tube cylindric, about 
5 cm. long, appressed-puberulent; sepals and petals narrowly oblong; 
corona filaments in 2 series, the outer about 6 mm. long, slightly 
falcate-dilated, the inner subulate, very short; operculum filamentose 
nearly to the base; ovary ovoid, velutinous. F.M. Neg. 16555. 

Loreto: Puerto Melendez, Pongo de Manseriche, Tessmann 
4770, type. Gamitanacocha, Rio Mazan, 100-125 meters, J. M. 
Schunke 358. 

Passiflora quadrangularis L. Syst. Nat. ed. 10. 2: 1248. 1759. 
P. macrocarpa Mast. Gard. Chron. 1869: 1012. 1869. 

Plant glabrous throughout; stem stout, quadrangular, the angles 
winged; stipules ovate or ovate-lanceolate, 2-3.5 cm. long; petioles 
6-glandular, the glands subsessile; blades broadly ovate or ovate- 
oblong, 10-20 cm. long, 8-15 cm. wide, abruptly acuminate, entire, 
penninerved, the principal lateral nerves 10-12 to a side, approximate; 
bracts cordate-ovate, 3-5.5 cm. long; flowers up to 12 cm. wide; 
sepals ovate or ovate-oblong, white, violet, or pinkish within; petals 
slightly narrower than the sepals, white, deeply pink-tinged; corona 
5-ranked, the 2 outer ranks subequal, the filaments up to 6 cm. long, 
radiate, multicolored, the other ranks of the corona much shorter, 
mostly purple and white; fruit oblong-ovoid, 20-30 cm. long, 12-15 



120 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cm. in diameter, terete or longitudinally 3-grooved; seeds obcordate 
or suborbicular, up to 1 cm. long. 

San Martin: San Roque, 1,400 meters, Williams 7250. Loreto: 
Iquitos, 100 meters, Killip & Smith 27152; King 733. Rio Itaya, 
Williams 188. Huanuco: Ambo, 2,600 meters, 2422. Chinchao, 
Ruiz & Pavdn. Extensively cultivated in tropical America. The 
pulp is eaten, and the thick rind is often made into sweetmeats or 
preserves. "Granadilla," "tumbo," "tambo." 

Passiflora quadriflora Killip, Journ. Wash. Acad. Sci. 17: 
424. 1927. 

Plant glabrous throughout; stipules setaceous; petioles glandless; 
blades narrowly lanceolate, 5-8 cm. long, 1.5-2 cm. wide, unlobed, 
entire, acute, coriaceous or subcoriaceous; peduncles in pairs, 2- 
flowered, the main portion and the branches nearly equal; bracts 
setaceous; flowers about 5 cm. wide, greenish white; sepals lanceolate; 
petals linear-lanceolate, about a third as long as the sepals; corona 
2-ranked, the outer filaments filiform, subequal to the petals, the 
inner capillary, much shorter; operculum slightly plicate, finely 
fimbriate; fruit globose-ovoid, subtrigonous, about 2.2 cm. in 
diameter. 

San Martin: Tarapoto, 1,200 meters, Vie 6464; Williams 5514. 
Huanuco: Rio Chinchao, 1,900 meters, 5189, type. 

Passiflora quadriglandulosa Rodschied, Med. & Chir. Bemerk. 
Esseq. 77. 1796. Tacsonia quadriglandulosa DC. Prodr. 3: 335. 1828. 

Stem terete, glabrous or the younger parts pilosulous; stipules 
setaceous, soon deciduous; petioles obscurely biglandular at the 
base; blades polymorphic, unlobed and oblong or oblong-lanceolate, 
8-15 cm. long, 3-8 cm. wide, or asymmetrically 2- or 3-lobed (lobes 
acuminate, the middle one longer than the lateral ones), irregularly 
repand-dentate, glabrous, or finely puberulent on the nerves above, 
finely puberulent or tomentellous beneath; bracts narrowly linear 
to narrowly oblong-lanceolate, 8-15 mm. long, 1-5 mm. wide, 
glandular-serrulate, or (in var. involucrata (Mast.) Killip) broadly 
ovate, up to 5 cm. long and 3 cm. wide, doubly serrate; flowers pink, 
red, or scarlet; calyx tube cylindric, 1-1.5 cm. long; sepals and petals 
similar and subequal, 6-8 cm. long; corona 3-ranked, the 2 outer 
ranks filamentose to the base, the filaments bright red or scarlet, 
the third coronal rank tubular, filamentose in upper third only; ovary 
ferruginous- tomentulous; fruit ovoid, about 3.5 cm. long; seeds 
reticulate. 



FLORA OF PERU 121 

Loreto: Fox 89. Rio Itaya, 110 meters, Kittip & Smith 29408; 
King 1069, 1226. Mishuyacu, 100 meters, Kittip & Smith 29974- 
Lower Rio Nanay, Williams 562. La Victoria, Williams 3118. 
Caballo-Cocha, Williams 2458. Iquitos, Williams 8052. Rancho 
Indiana, Rio Marafion, 110 meters, Mexia 6405. Gamitanacocha, 
Rio Mazan, 100-125 meters, J. M. Schunke 47, 52. Chasuta, 
Raimondi 581. Peras, Fox 119. Rio Ucayali, Tessmann 3309. 
Guianas to Peru, Bolivia, and northern Brazil. "Granadilla acida." 
The variety is represented in Peru by the following collections, all 
from Loreto: Mishuyacu, Klug 342, 417. Rio Nanay, Williams 780, 
1104. Iquitos, Ducke 21230. "Estrella," "granadilla." 

Passiflora riparia Mart, ex Mast, in Mart. Fl. Bras. 13, pt. 1: 
599. pi. 116. 1872. 

Plant glabrous except the ovary; stipules linear, soon deciduous; 
petioles biglandular at the middle; blades oblong or oblanceolate, 
10-15 cm. long, 4.5-8 cm. wide, abruptly acuminate, entire or 
minutely serrulate, subcoriaceous; peduncles 4-5 cm. long, borne on 
short, axillary, leafless (or with much reduced leaves) branches; 
bracts ovate-oblong, 3-4 cm. long, reddish; calyx tube cylindric- 
campanulate, about 1 cm. long; sepals oblong, 4-5 cm. long, white, 
green-tinged; petals oblong-linear, white; corona in several series, 
the filaments of the outer series subequal, 4-5 cm. long, banded with 
blue or violet and white, those of the innermost series about 2 mm. 
long, the intervening ones an irregular mass of tubercles; operculum 
crenulate; ovary rufo-sericeo-tomentose; fruit ovoid or globose, 3-4 
cm. long, apparently not edible; seeds reticulate at the center, striate 
at the margin. 

San Martin: Tarapoto, Williams 5637, 5848. Alto Rio Huallaga, 
Williams 6300. Lamas, 840 meters, Wittiams 6378. Juanjui, 400 
meters, Klug 3897. Chasuta, 260 meters, Klug 4037. Loreto: 
Yurimaguas, 135 meters, Kittip & Smith 28214, 28940; Williams 
7876. Lower Rio Huallaga, 135 meters, Killip & Smith 29012. 
Iquitos, 100 meters, Williams 1392, 1440, 7996. La Victoria, Rio 
Amazonas, Williams 3126. Junin: Puerto Bermudez, 375 meters, 
Kittip & Smith 26683; Puerto Yessup, 400 meters, Kittip & Smith 
26307. Also in Amazonian Brazil and southernmost British Guiana. 
"Purupuru," "chinchorcon," "granadilla." 

Passiflora rosea (Karst.) Killip, Field Mus. Bot. 19: 278. 1938 
(P. pinnatistipula X P. mollissima). Poggendorffia rosea Karst. 



122 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Linnaea 28: 438. 1856; Fl. Columb. 1: pi. 15. 1858. Tacsonia rosea 
Sodiro, Anal. Univ. Quito 18: 343. 1903. 

Plant tomentose; stipules linear-lanceolate, laciniate-dentate 
above the middle; petioles 6-8-glandular; blades 3-lobed to slightly 
below the middle, sharply serrate, the lobes ovate or ovate-oblong; 
bracts cordate-ovate, free to the base; calyx tube 4.5-5 cm. long, 
enlarged near the middle; sepals and petals oblong-lanceolate, 2.5-3 
cm. long, bright pink; corona filamentose, in 2 series; gynophore 
3-4.5 cm. long, the filaments of the stamens arising below its middle, 
the anthers basifixed; ovary densely white-sericeo- tomentose; fruit 
ovoid, about 8 cm. long; seeds broadly obovate, closely reticulate. 

Junin: Tarma, 3,100 meters, Killip & Smith 21885. Huancayo, 
CJiavez 129 Cuzco: Paucartambo, 3,000 meters, Herrera 2999a. 
Also in Colombia and Ecuador. 

This curious plant is the basis of the monotypic genus Poggen- 
dorffia. There are, however, good reasons for believing it a natural 
hybrid between P. pinnatistipula and P. mollissima. In the vegetative 
parts it rather closely resembles P. mollissima, whereas in the length 
and indument of the calyx tube and the filamentose outer series of 
corona filaments it is suggestive of P. pinnatistipula. "Purupuru." 

Passiflora rubra L. Sp. PI. 956. 1753. 

Stem densely grayish pubescent; stipules setaceous; petioles 
glandless; blades membranous, finely pubescent or rarely softly 
hirsute, 2-8 cm. along the midnerve, 4-11 cm. along the lateral nerves, 
3-10 cm. wide, bilobed, the lobes widely divergent, or shallowly and 
subequally 3-lobed; peduncles solitary; bracts none; flowers up to 
5 cm. wide; sepals linear-lanceolate, reddish or greenish without, 
white within; petals half as long as the sepals, white; corona pre- 
vailingly 1-ranked, the filaments narrowly liguliform or nearly fili- 
form, purplish at the base, green or white above; operculum slightly 
plicate; ovary subglobose, densely hirsute with white or brownish 
hairs; fruit ovoid or obovoid, 2-2.5 cm. long, 1.5-1.8 cm. wide, 
rarely larger, tapering at the base, 10-angled; seeds black, trans- 
versely sulcate, the ridges smooth. 

Tumbez: Hacienda Chicama, 900-1,000 meters, Weberbauer 7653. 
San Martin: San Roque, 1,350-1,500 meters, Williams 7088. 
Tarapoto, Ule 6546. Huanuco: Muna, 2,100 meters, 4162. Junin: 
La Merced, 700 meters, Killip & Smith 23400, 23432, 25371 . Yapas, 
Pichis Trail, 1,500 meters, Killip & Smith 25441. Ayacucho: Aina, 
800 meters, Killip & Smith 22800. Cuzco: Uchumayo, Bues 



FLORA OF PERU 123 

(Herrera 2129). Echarate, 900 meters, Bues in 1928. Without 
locality: Ruiz & Pavon. "Mazo-manchachi" (Cuzco). Common 
in the West Indies, and from Colombia to Peru; rarer elsewhere in 
South America. 

Passiflora serrato-digitata L. Sp. PI. 960. 1753. P. serrata L. 
Syst. Nat. ed. 10. 2: 1248. 1759. P. digitata L. Sp. PL ed. 2. 1360. 
1763. P. serrata var. digitata R. & P. ex DC. Prodr. 3: 330. 1828. 
P. digitata R. & P. ex M. Roemer, Fam. Nat. Syn. 2: 183. 1846. 

Plant essentially glabrous throughout; stipules linear-subulate, 
about 1.5 cm. long; petioles biglandular at the apex and at the middle, 
the glands up to 3 mm. long; blades up to 15 cm. long and 18 cm. 
wide, palmately 5-7-lobed to below the middle, the lobes oblong to 
oblanceolate, acuminate, finely serrulate; bracts ovate-lanceolate, 
3-5 cm. long, united a quarter their length, slightly pubescent; 
flowers 6-8 cm. wide; sepals and petals oblong, blue-tinged to pinkish 
blue; corona in several series, the 2 outer filamentose, radiate, banded 
with blue, white, and purple; fruit globose, 4-5 cm. in diameter, the 
pericarp brittle, the pulp edible; seeds reticulate. 

San Martin: Juanjui, 800 meters, Klug 4278. Loreto: Iquitos, 
100 meters, Killip & Smith 27233, 27422; Williams 3751. Yuri- 
maguas, 135 meters, Killip & Smith 27849. Lower Rio Huallaga, 
155-210 meters, Williams 5153. Rio Ucayali, Tessmann 3373. Pongo 
de Manseriche, Tessmann 4955. Junin: La Merced, 700 meters, 
Killip & Smith 23931, 24052. Paucartambo Valley, 700 meters, Killip 
& Smith 25422. Ayacucho : Aina, 900 meters, Killip & Smith 22825. 
Cuzco: Chancamayo, Lares Valley, Weberbauer 7939. Arequipa: 
Vito, Ruiz & Pawn. Department uncertain: Hacienda Chalhua- 
puquio, Stevens 154- "Ccoto-gguantte," "ckoto huanthi." West 
Indies, Guianas, and Amazonian Brazil to southern Bolivia and Peru. 

Passiflora skiantha Huber, Bol. Mus. Goeldi 4: 591. /. 5. 1906. 

Scandent shrub, glabrous throughout; leaves elliptic, up to 17 
cm. long and 10 cm. wide, abruptly acute or acuminate, membran- 
ous; flowers in short, dense fascicles; calyx tube cylindric, 3-3.5 cm. 
long; sepals and petals oblong, 2.5-3 cm. long; corona filaments in 
4 series, the outermost spatulate, about 1.5 cm. long, the others much 
shorter; ovary ellipsoidal. 

Loreto: Cerro de Canchahuaya, Huber 1424, type. 

Passiflora spectabilis Killip, Journ. Wash. Acad. Sci. 20: 
379. 1930. 



124 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plant glabrous; stipules semi-ovate or semi-oblong, up to 6 cm. 
long; petioles 2-3-glandular, the glands sessile; blades 6-12 cm. long, 
10-20 cm. wide, 3-lobed not more than to the middle, peltate near 
the lower margin, glaucescent, the lobes broadly triangular, obtuse or 
subacute; bracts ovate, 6-7 mm. long, borne 6-12 mm. from the base 
of the flower; flowers 5-7 cm. wide, pink or white; calyx tube cam- 
panulate; sepals oblong, short-awned; petals linear; corona filaments 
very slender, in 4 series, the 2 outer subequal to the petals, blue, 
white at the tips; operculum filamentose nearly to the base; fruit 
globose, 5 cm. in diameter, edible; seeds coarsely reticulate. 

Loreto: Mishuyacu, 100 meters, Killip & Smith 29884, type; 
King 242. Iquitos, 100 meters, Killip & Smith 27165, 29846 
Junin: Enenas, 1,600 meters, Killip & Smith 25764. 

Passiflora spinosa (Poepp. & Endl.) Mast. Trans. Linn. Soc. 
27: 630. 1871; in Mart. Fl. Bras. 13, pt. 1: 576. 1872. Tacsonia 
spinosa Poepp. & Endl. Nov. Gen. & Sp. 2: 59. pi. 181. 1838. 

Woody vine, glabrous throughout except the ovary, the tendrils 
often reduced to stout spines; leaves oblong, 10-17 cm. long, 3-9 
cm. wide, acuminate, entire, coriaceous, lustrous; flowers bright red, 
in slender racemes up to 25 cm. long, these sometimes bearing 
reduced leaves; calyx tube cylindric, 4-5 cm. long, glabrous; sepals 
and petals narrowly oblong; corona filaments in 2 series, yellow, the 
outer dolabriform, the inner filiform; operculum filamentose in the 
upper third; ovary narrowly oblong, puberulent. 

Loreto: Yurimaguas, 135 meters, Poeppig D. 2187, type; Killip 
& Smith 28147. Caballo-Cocha, Rio Amazonas, Williams 2458. 
Upper Rio Nanay, Williams 1134. Also in Colombia and Amazonian 
Brazil. 

Passiflora suberosa L. Sp. PI. 958. 1753. 

Plant glabrous to densely pubescent; stipules linear-subulate; 
petioles minutely biglandular above the middle, the glands normally 
stipitate; blades membranous or subcoriaceous, highly variable in 
outline, entire or usually (in Peruvian material) 3-lobed, the middle 
lobe the largest; bracts setaceous; flowers 8-13 mm. wide, solitary 
or in pairs; sepals ovate-lanceolate, greenish yellow; petals none; 
corona filaments in 2 series, filiform, the outer ones purple, white, 
and yellow, the inner green; operculum plicate; fruit globose or 
ovoid, 6-15 mm. in diameter, glabrous; seeds coarsely reticulate. 

Tumbez: Hacienda Chicama, 900 meters, Weberbauer 7638. 
Lima: Callao, Sargent 36; Gaudichaud 154, 154bis. Lima, Ball in 



FLORA OF PERU 125 

1882; Kittip & Smith 21524; Rose 18773; Ruiz & Pavdn; Savatier 
1438; Wawra 2651. Lurin, Pennell 12207. Atocongo, Pennell 14772. 
Ancon, 120 meters, Mexia 8101. Chosica, 1,000 meters, 2855. 
Miraflores, Savatier 11+39. Department uncertain: La Convencion, 
Bues in 1928. Without locality: Dombey 734; Nee; Gay. Common 
throughout tropical America. "Noxbe cimarron." 

Passiflora subulata Mast, in Mart. Fl. Bras. 13, pt. 1: 567. 
1872. P. platyceras Harms, Repert. Sp. Nov. 18: 298. 1922. 

Plant glabrous; stipules semi-oblong-lanceolate, 2-3.5 cm. long; 
petioles biglandular at or above the middle; blades 4-7 cm. long, 5-8 
cm. wide, 3-lobed to the middle, the lobes ovate or ovate-oblong, 
obtuse, ascending; bracts ovate or ovate-oblong, 1.5-2.5 cm. long, 
cordate, borne close to the flower base; flowers 6-8 cm. wide, white; 
sepals oblong-lanceolate, keeled, the keel terminating in a green, 
foliaceous awn 8-12 mm. long; petals lanceolate; corona filaments 
in 3 series, the outermost capillary, about 2 cm. long, the others 2-3 
mm. long; operculum fimbrillate to the middle. 

Junin: Huacapistana, 1,800-2,500 meters, Weberbauer 2171, type 
of P. platyceras; Killip & Smith 24311, 24323. Pariahuanca Valley, 
between Panti and Rocchac, 2,400 meters, Weberbauer 6542. With- 
out locality: MacLean, type. 

Passiflora tarapotina Harms, Verh. Bot. Ver. Brandenb. 48: 
185. 1907. Tacsonia glauca Poepp. ex Mast, in Mart. Fl. Bras. 
13, pt. 1: 569. 1872, as synonym? 

Plant glabrous throughout; stipules semi-ovate, up to 2.5 cm. 
long; petioles up to 4 cm. long, biglandular near the middle; blades 
6-8 cm. long, 9-10 cm. wide, 3-lobed to below the middle, subpeltate, 
glaucous beneath; peduncles 10-16 cm. long, very slender; bracts 
narrowly oblong to ovate-oblong, 10-15 mm. long, serrulate; flowers 
blood-red; calyx tube cylindric, 2-2.5 cm. long; sepals and petals 
narrowly oblong, subequal to the tube, the sepals bearing a large, 
foliaceous awn; corona in 3 or 4 closely approximate ranks of fila- 
ments barely 1 mm. long; operculum filamentose half its length. 
F.M. Neg. 16559. 

San Martin: Tarapoto, about 750 meters, Ule 6462, type; 
Spruce 3923; Williams 5418, 5555, 5709, 5776. Loreto: Juanjui, 
Province of Mainas, Poeppig in 1830 ("Tacsonia glauca?"). Very 
similar in leaf shape to P. aristulata and P. subulata but readily 
distinguished by the flowers. 



126 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Passiflora tenella Killip, Journ. Wash. Acad. Sci. 20: 375. 1930. 

A very slender vine, essentially glabrous throughout; stipules 
setaceous; petioles glandless; blades transversely oblong in general 
outline, 1.5-3 cm. long, 4-7 cm. wide, shallowly 3-lobed at the 
truncate apex (lobes obtuse or truncate; rarely the leaves distinctly 
3-lobed about a third their length), thin-membranous, glaucous 
beneath; peduncles solitary, 2-3 cm. long, very slender; bracts 
setaceous; flowers about 1.5 cm. wide, greenish white; sepals and 
petals thin-transparent, the petals only 2-3 mm. long; corona fila- 
ments filiform, in 2 series, the outer equaling the sepals, the inner 
very short; operculum only slightly plicate; fruit ellipsoidal, about 
3 cm. long and 8 mm. in diameter, 6-ribbed, stipitate; seeds trans- 
versely 4-5-sulcate. 

Tumbez: Hacienda La Choza, 100-200 meters, Weberbauer 7704, 
type. Lima: Lima Botanical Garden, said to have come from forests 
of eastern Peru, Pennell 14801. Ecuador. 

Passiflora Tessmannii Harms, Notizbl. Bot. Gart. Berlin 9: 
978. 1926. 

Scandent shrub; leaves ovate, broadly oblong, or obovate, 5-9 cm. 
long, 2.5-6.5 cm. wide, acute or subacute, entire, membranous, 
minutely puberulous beneath; peduncles solitary or in pairs; calyx 
tube cylindric-campanulate, about 1 cm. long; flowers white; sepals 
narrowly oblong, 1.5 to 2 cm. long, the petals similar but shorter; 
corona filaments in 2 series, the outer ones falcate-dilated above the 
middle, attenuate at the tip, about 7 mm. long, yellow, the inner 
subulate, much shorter; operculum fimbrillate; ovary ovoid, velu- 
tinous. F.M. Neg. 16560. 

Loreto: Rio Maranon, at mouth of Rio Santiago, Tessmann 
4385, type. 

Passiflora tiliaefolia L. Sp. PI. 956. 1753. 

Plant glabrous throughout; stipules ovate-lanceolate or oblong- 
lanceolate, 1-2 cm. long, the upper ones rarely narrowly linear-lanceo- 
late; petioles 2-4-glandular, the glands saucer-shaped, subsessile or 
short-stipitate; blades cordate-ovate, 10-25 cm. long, 8-18 cm. wide, 
abruptly acuminate, cordate, entire; bracts ovate, about 2 cm. long, 
connate part way; flowers about 8 cm. wide, the sepals and petals 
oblong; corona 5-ranked, the 2 outer rows of filaments radiate, 
about half as long as the petals, the 3 inner rows much shorter; 
fruit subglobose, about 5 cm. in diameter. 

Peru: Without locality, Pavon. 



FLORA OF PERU 127 

This is a doubtful species, based originally upon a figure given 
by Feuille'e (1714) of a plant growing at Lima. Specimens from the 
mountains of Colombia (between 1,500 and 2,500 meters) apparently 
represent this species. 

Passiflora tricuspis Mast, in Mart. Fl. Bras. 13, pt. 1: 587. 1872. 

Stem glabrous, the ridges strongly scabrellous; stipules setaceous; 
petioles finely pilosulous or nearly glabrous, glandless; blades variable 
but in Peruvian plants usually 3-lobed from a third to two-thirds 
their length (lobes lanceolate to linear-oblong, subequal or the 
middle one the longer, the lateral lobes ascending), 5-13 cm. along 
the midnerve, 4.5-11 cm. along the lateral nerves, 3-7 cm. wide, 
glabrous or minutely puberulent above, finely pilosulous beneath; 
bracts setaceous; flowers 3-4.5 cm. wide; sepals and petals oblong 
or lance-oblong, white; corona 2-ranked, the outer filaments narrowly 
liguliform, about 1.5 cm. long, the inner narrowly linear, 2-2.5 mm. 
long, capitate; operculum plicate; ovary glabrous; fruit globose, 
about 1.5 cm. in diameter; seeds transversely 7-sulcate. 

Loreto: Rio Ucayali, Tessmanni 3194. Junin: La Merced, 600- 
700 meters, 5436; Killip & Smith 23474. Bolivia to eastern Brazil 
and Paraguay. 

Passiflora trifasciata Lemaire, 111. Hort. 15: pi. 544- 1868. 

Plant glabrous throughout; stipules subulate; petioles glandless; 
blades 5-10 cm. along midnerve, 4-10 cm. along lateral nerves, 4-10 
cm. wide, 3-lobed about a third their length (lobes deltoid, acute 
or subobtuse, the lateral ascending), membranous, dark green, 
mottled with white or yellowish green along the nerves above, 
reddish or violet beneath; bracts setaceous; flowers 2.5-3.5 cm. wide; 
sepals oblong, light green; petals linear, light green; corona 2-ranked, 
the outer filaments terete, 8-10 mm. long, the inner linear-clavate, 
much shorter; operculum plicate; fruit globose, 1.5-2.5 cm. in 
diameter; seeds transversely 6-sulcate. 

San Martin: Lamas, 840 meters, ^illiams 6327. Loreto: 
Yurimaguas, 135 meters, Killip & Smith 27819, 28069, 28297, 
28318; Williams 5075, 5213. Balsapuerto, 200 meters, Killip & 
Smith 28412. Frequently cultivated; probably endemic to northern 
Peru. "Millua caspi." 

Passiflora trifoliata Cav. Icon. 5: 16. pi. 427. 1799. Tacsonia 
trifoliata Juss. Ann. Mus. Hist. Nat. Paris 6: 393. 1805. T. trigona 
DC. Prodr. 3: 334. 1828. 



128 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A rather stout vine, densely short-villous-hirsute throughout; 
stipules oblong or oblong-lanceolate, 1.5-3 cm. long, glandular-fimbri- 
olate; leaves trifoliolate, the petioles obscurely glandular, the leaflets 
ovate-oblong or linear-oblong, 3-5 cm. long, 1.5-2.5 cm. wide, acute, 
sessile, entire or serrulate; peduncles up to 9 cm. long, very stout; 
bracts free to the base, ovate-oblong, 2-3.5 cm. long, closely fimbriate 
or laciniate-fimbriate; flowers pendent, the calyx tube cylindric, 
3-4 cm. long, green, purple-tinged without; sepals ovate-oblong, 
2-3 cm. long, deep pink, white toward the base; petals ovate- 
oblong; corona minutely tuberculate, 2-ranked; fruit ovoid, 4-6 
cm. long, 2.5-3.5 cm. in diameter, longitudinally 3-grooved; seeds 
reticulate. 

Ancash: Tallenga, Weberbauer 2878a. Province of Cajatambo, 
Weberbauer 2654- Lima: "Lima," Weberbauer 261. Rio Blanco, 
3,200 meters, Kittip & Smith 21636. Viso, 2,700 meters, 579. 
Huaros, 3,500 meters, Pennell 14723. Rio Rimac, Forster in 1905. 
Huanuco: Huariaca, Sawada P97. Junin: Tarma, Mathews 674- 
Huancayo, 4,000 meters, Ledig 4- Oroya, Kalenborn 184. Cuzco: 
Paucartambo, Herrera 485. Department uncertain: Guamantanga, 
Nee, type; Cajavilca Valley, Savatier 1631. Without locality: Ruiz 
& Pavdn (type of Tacsonia trigona) ; Lobb 20; Savatier 490; Gay 542; 
Dombey 744; Haenke 1847; Raimondi 8597; Wilkes Expedition. 
"Naupa-mchu-jamppajhuai" (Inca). 

Passiflora triloba R. & P. ex DC. Prodr. 3: 330. 1828. P. colu- 
brina Poepp. & Endl. Nov. Gen. & Sp. 2: 58. 1838. 

Plant essentially glabrous throughout; stipules ovate-lanceolate, 
2-3 cm. long; petioles biglandular at the apex, the glands saucer- 
shaped; blades 10-15 cm. long, 10-18 cm. wide, cordate-ovate or 
usually 3-lobed to about the middle (lobes ovate or oblong-ovate, 
subequal, obtuse, minutely denticulate), deeply cordate, glaucescent 
beneath; bracts ovate, 4.5-7 cm. long, connate only at the base; 
flowers up to 10 cm. wide; sepals oblong-lanceolate, purple-spotted 
or purple-tinged, the petals longer than the sepals; corona 3-ranked, 
the 2 outer rows of filaments banded with purple and white below 
and with blue and white above. F.M. Neg. 16562. 

San Martin: Tarapoto, Spruce 4052. Loreto: Yurimaguas, 135 
meters, Kittip & Smith 27664, 27834, 28716; Williams 4980; Poeppig 
2171, type of P. colubrina. Rio Ucayali, Tessmann 3155. Junin: 
San Ramon, 1,000 meters, cultivated, Killip & Smith 24092. Also 
in Bolivia. 



FLORA OF PERU 129 

Passiflora trisecta Mast, in Mart. Fl. Bras. 13, pt. 1: 564. 1872. 
P. thaumasiantha Harms, Notizbl. Bot. Gart. Berlin 10: 813. 1929; 
Rev. Chil. Hist. Nat. 34. /. 47. 1930. 

Plant ferruginous-villous or -tomentose; stipules ovate-lanceolate, 
1-2.5 cm. long, laciniate-serrate, the serrations glandular; petioles 
5-7-glandular, the glands filiform; blades trifoliolate, the leaflets 
oblong-lanceolate, 4-9 cm. long, 1.5-3 cm. wide, acuminate, petiolu- 
late, serrate; peduncles stout, 8-15 cm. long; bracts ovate, 2.5-3.5 
cm. long, free to base, deeply fimbriate-laciniate; flowers white; 
calyx tube tubular-campanulate, 1-1.5 cm. long; sepals narrowly 
oblong, 2-4 cm. long; petals linear-lanceolate; corona in 3 series, the 
outer 2 filamentose, the innermost dentiform; fruit globose, about 
5 cm. in diameter; seeds reticulate. 

Junin: La Mejorada, 2,600 meters, Kittip & Smith 23345. 
Ayacucho: Anco, Rio Mantaro Valley, 2,800 meters, Kittip & Smith 
22178. Huanta, 2,800 meters, Kittip & Smith 23332. Cuzco: 
Urubamba Valley, Cook & Gilbert 824; Herrera 1673, type of P. 
thaumasiantha. Mollepata, Herrera 1223. Arequipa: Huaspa, 
Raimondi 11479. Also in Bolivia, where known only from the type 
specimen, collected by Pentland. "Tumbo tumbo," "kita tumbo." 

Passiflora vespertilio L. Sp. PI. 957. 1753. 

Stem glabrous or finely puberulent; stipules narrowly linear- 
setaceous; petioles short, glandless; blades coriaceous, glabrous and 
lustrous above, glabrous or slightly puberulent beneath, 2-3-lobed, 
the lobes divaricate, acuminate, the upper margin usually truncate, 
often also undulate; bracts setaceous; flowers 4-5 cm. wide, yellowish 
green; sepals broadly lance-oblong, obtuse; petals oblong, shorter 
than the sepals; corona 2-ranked, the outer filaments narrowly 
liguliform, united at the base into a broad membrane, the free fila- 
ments 1-1.5 cm. long, the inner filaments capillary, much shorter; 
operculum closely plicate; ovary ovoid, glabrous; fruit subglobose, 
1-1.5 cm. in diameter; seeds transversely sulcate. 

Loreto : Rio Nanay, Williams 1201 . Iquitos, 100 meters, Williams 
8180. A common plant of the Guianas and Amazonian Brazil, 
barely entering northern Peru and Bolivia. 

Passiflora vestita Killip, Field Mus. Bot. 19: 473. 1938. 

Plant densely rufo-hirsute nearly throughout; stipules semi- 
annular about the stem, laciniate nearly to the base, the segments 
filiform, gland-tipped; petioles glandless but bearing numerous gland- 
tipped hairs; blades 7-15 cm. long, 6-12 cm. wide, 3-lobed, truncate 



130 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or cordate, densely glandular-ciliate, the lobes ovate-lanceolate, 
acute or acuminate, the lateral lobes suberect; bracts 4-5 cm. long, 
pinnatisect or obscurely twice-pinnatisect, the segments filiform, 
gland-tipped; fruit globose, about 4 cm. in diameter; seeds obcuneate, 
about 8 mm. long and 5 mm. wide, tridentate, reticulate at the mid- 
dle of each face, smooth toward the margin. 

Loreto: Between Yurimaguas and Balsapuerto, 135-150 meters, 
Killip & Smith 28126, type. 

Passiflora vitifolia HBK. Nov. Gen. & Sp. 2: 138. 1817. 
P. punicea R. & P. ex DC. Prodr. 3: 329. 1828. 

Stem, petioles, and peduncles densely ferruginous- tomentose; 
stipules setaceous, deciduous; petioles biglandular at the base, rarely 
with additional glands; blades up to 15 cm. long and 18 cm. wide, 
3-lobed to below the middle, irregularly repand-dentate or crenate, 
tomentulous on the nerves above, densely puberulent or tomentulous 
beneath; peduncles up to 9 cm. long; bracts oblong to oblong-lanceo- 
late, 1.5-2.5 cm. long, 4-8 mm. wide, acuminate, glandular-serrate, 
reddish; flowers scarlet; calyx tube cylindric, 1-1.8 cm. long; sepals 
lanceolate, 6-8 cm. long; petals linear-lanceolate, slightly shorter 
than the sepals; corona 3-ranked, the 2 outer ranks filamentose, the 
outermost filaments 1.5-2 cm. long, red or yellow, the next ones 
somewhat shorter, pale red, the third rank tubular, pale red, short- 
filamentose; ovary densely tomentulous; fruit ovoid, about 5 cm. 
long, puberulent; seeds reticulate. 

San Martin: Juanjui, Alto Rio Huallaga, 400-800 meters, Klug 
4190. Without definite locality: Ruiz & Pavon, type of P. punicea. 
Common from Nicaragua to Amazonian Venezuela; rare in Ecuador. 

Passiflora Weberbaueri Harms, Bot. Jahrb. 54: Beibl. 117: 
79. 1916. 

Plant essentially glabrous; stipules broadly semi-ovate or semi- 
cordate, up to 1.7 cm. long, lacerate; petioles bearing 2 pairs of stipi- 
tate glands; blades trifoliolate, the leaflets oblong to ovate-lanceolate, 
6-14 cm. long, 2-5 cm. wide, subsessile or short-petiolulate, acumi- 
nate, serrate, coriaceous; peduncles 7-12 cm. long; bracts oblong 
or ovate-oblong, 4-4.5 cm. long, free to the base, lacerate-serrate; 
flowers white; calyx tube urceolate-campanulate, up to 1.5 cm. long; 
sepals narrowly lanceolate, 5-6 cm. long; petals broadly linear, 
slightly shorter than the sepals; corona in 3 series, the outer 2 
ranks filamentose, the innermost membranous, short-filamentose; 
ovary velutinous. F.M. Neg. 3285. 



FLORA OF PERU 131 

Cuzco: Cosfiipata, 2,700-2,800 meters, Weberbauer 6933, type. 
Yanamayo, 2,200-2,400 meters, Pennell 14040. 

2. DILKEA Mast. 

Woody vines (the Peruvian species), glabrous throughout, with- 
out tendrils or rarely with a few poorly developed ones; leaves 
alternate, petiolate, simple, entire; bracts minute, subulate; flowers 
(in the Peruvian species) in sessile or subsessile glomerules, white, 
the sepals 4, oblong, 2 broader than the other 2, united below the 
middle when young to form a cylindric or funnel-shaped tube, at 
length separating to the base and readily deciduous, the petals 4 or 5, 
free; corona 2-ranked, the outer rank consisting of slender, free or 
nearly free filaments, the inner tubular in the lower part, cleft above 
into floccose, crispate threads or into segments margined with such 
threads; operculum none; stamens 8, hypogynous; ovary subsessile 
or borne on a short gynophore; fruit globose or ovoid. 

Dilkea is confined to the middle and upper Amazon Basin, and 
is known from only a few collections. Five species are at present 
recognized but additional material may well show that the genus is 
monotypic. 

Sepals not more than 1.5 cm. long; ovary subsessile; leaves obovate. 

D. parviflora. 
Sepals 2-3 cm. long; ovary borne on a short gynophore. 

Leaves cuneiform, truncate at the upper margin except for a 

short lobe at the center D. retusa. 

Leaves broadly ovate to oblong-oblanceolate, rounded or acumi- 
nate D. Wallisii. 

Dilkea parviflora Killip, Field Mus. Bot. 19: 575. 1938. 

Leaves obovate, 15-18 cm. long, 8.5-10.5 cm. wide, rounded at 
the apex, coriaceous; flowers white; sepals and petals oblong, 1.2- 
1.5 cm. long, obtuse; outer corona filaments narrowly liguliform, 
about 1.3 cm. long, the inner ones narrowly flabellate, about 1 cm. 
long, densely crispate-floccose at the margin in the upper half. 

Loreto: Mishuyacu, 100 meters, Klug 1158, type. 

Dilkea retusa Mast. Trans. Linn. Soc. 27: 628. 1871. 

Leaves cuneiform, 15-20 cm. long, 7-11 cm. wide, coriaceous, 
the lateral nerves divaricate at nearly a right angle from the mid- 
nerve, slightly ascending; sepals and petals oblong, 1.5-2.5 cm. long; 
outer corona filaments liguliform, 2-2.5 cm. long, the inner filiform 



132 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

in the lower half, spatulate-dilated in the upper half and margined 
with floccose, crispate threads. 

Loreto: Mishuyacu, 100 meters, Klug 1017. 

Dilkea Wallisii Mast, in Mart. Fl. Bras. 13, pt. 1: 622. pi. 106, 
f. 3. 1872. 

Leaves ovate to oblong-oblanceolate, 12-15 cm. long, 5-10 cm. 
wide, abruptly acuminate at the rounded apex, rounded or cuneate 
at the base, coriaceous, the lateral nerves ascending; sepals and 
petals oblong, 2.5-3 cm. long; outer corona filaments narrowly 
ligulate, about 2 cm. long, the inner filiform below the middle, 
spatulate-dilated in the upper half and margined with floccose, 
crispate threads; fruit depressed-spherical, up to 2.5 cm. long and 
4.5 cm. wide. 

Loreto: Florida, Rio Putumayo, at mouth of Rio Zubineta, 200 
meters, Klug 2100. Gamitanacocha, Rio Mazan, 100-125 meters, 
J. M. Schunke 233. 

CARICACEAE. Papaya Family 

Low or often tall and simple-trunked trees usually characteristic 
in appearance by virtue of the ample, somewhat peltate, palmately 
lobed or 7-9-foliolate leaves crowded and spreading or drooping from 
near the tips of the branchlets and often overhanging the melon- 
like fruits. Flowers usually unisexual, the male and female corollas 
dissimilar, the former panicled, the latter solitary or, if panicled, 
usually few and crowded. Corolla with a long slender tube, the 
limb horizontally divaricate, the 5 petals soon caducous. The sap 
of the leaves is milky, sometimes orange-colored. 
Calyx and corolla segments alternate with each other; stamens 

rarely coalescent below 1. Carica. 

Calyx and corolla segments, at least the male, opposite; stamens 

usually coalescent below 2. Jacaratia. 

1. CARICA L. 

Reference: Solms-Laubach in Mart. Fl. Bras. 13, pt. 3: 174- 
195. 1889. 

Stamens 10, 5 sessile or subsessile, alternating with the corolla 
lobes, 5 inserted in the throat on short filaments. Style wanting 
or short, rarely elongate, the stigmas 5, entire to variously divided. 

The Peruvian species all belong to the section Vasconcellea (St. 
Hil.) Solms-Laubach, characterized by 5-celled fruits (unless when 



FLORA OF PERU 133 

young 1 -celled above) and simple or only 2-lobed stigmas. Kuntze 
referred all the species to Papaya (Tourn.) Adans. Heilborn has 
recently, Sv. Bot. Tidskr. 30: 224. 1936, described C. pinnatifida 
from northwestern Bolivia with ovate-lanceolate subpinnatifid leaves 
but with 1 -celled fruit and branched stigmas. 

The sweetly delicious at its best "papaya," C. papaya L., the 
well-developed tree as beautiful as the fruit is attractive, is some- 
times cultivated in Peru as in all warm countries but is unknown any- 
where in a wild state; it may well have originated from one or the 
other of the following, several of which are more or less satisfactory 
as food and which have long been planted by the inhabitants in the 
mountain villages. Among the Peruvian forms it is recognizable by 
its 1-celled fruits, mostly divided 5-7 leaf-lobes and irregularly 
branched stigmas. It was found cultivated (and collected) by 
Williams at Caballo-Cocha and La Victoria. 

Apart from its fruit the papaya is valuable because the sap yields 
an enzyme, "papain," with strongly digestive properties on pro- 
teins; it is therefore a commercial product in medicine. It is com- 
monly reputed in the tropics that meat may be made tender by 
washing it in the diluted juice, or by cooking it with the leaves, or 
even by enveloping it with them for some hours. 

Besides the above reference I am indebted to Harms' review of 
some South American species with special attention to the Peruvian, 
Notizbl. Bot. Gart. Berlin 8: 91-100. 1922, from which careful synop- 
sis I have freely drawn. As he remarks, it is necessary for an ade- 
quate study that collections be made showing female flowers and 
also fruits, both of which for a number of described species are 
partly or quite unknown. Determinations, unless otherwise men- 
tioned, are mostly by Harms. 

The following key, purely artificial, applies to the more usual 
leaf -forms; the constant characters are found in the inflorescence 
(flowers, particularly style) and in the fruit, but herbarium material 
at present shows these characters only partially. The flowers except 
in two species, C. monoica and C. erythrocarpa, are dioecious; and in 
the dioecious species the male and female inflorescence so far as 
known is subequal and elongate only in C. gossypiifolia and C. 
platanifolia; the male inflorescence is contracted in C. glandulosa, 
C. lanceolata and C. candicans, and laxly paniculate in the remaining 
species (so far as known). Besides the following there is a specimen 
by Klug (3687 from Zepalacio near Moyobamba distributed as C. 



134 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

platanifolia) , but its male flowers are too long and borne in a greatly 
elongate very lax panicle; it is, perhaps, allied to C. paniculata. 
Leaves not lobed, sometimes more or less dentate. 
Margin of leaves entire. 

Leaves white pubescent beneath C. candicans. 

Leaves glabrous, often white-glaucous beneath. 
Leaves oblong-lanceolate, more or less glaucous beneath or 

narrowly long-acuminate. 
Leaves narrowly long-acuminate; male corolla tube 8-10 

mm. long C. acuta. 

Leaves acute or acuminate at tip. 

Male corolla tube 6-8 mm. long; leaves mostly about 

1 dm. long C. lanceolata. 

Male corolla tube 11-12 mm. long; leaves mostly 1.5 

dm. long or longer C. glandulosa. 

Leaves ovate-elliptic to broadly ovate, rather abruptly acu- 
minate, concolor or only lighter green beneath. 

C. heterophylla. 

Margin of leaves coarsely dentate C. Augusti. 

Leaves all or most of them deeply lobed or divided, the lobes entire 
or lobed or dentate. 

Leaf lobes 3, more or less coarsely serrate C. Weberbaueri. 

Leaf lobes entire or more or less lobed or parted or with a few 

remote teeth. 

Leaves repandly or shallowly 3-5-lobed, the margins repandly 

and remotely dentate, usually some lobes entire; male and 

female peduncles both elongate, subequal. . .C. platanifolia. 

Leaves 3-7-lobed, the lobes entire or lobed; male inflorescences 

generally much longer. 
Leaves deeply 3-parted, the divisions narrowly linear. 

C. glandulosa. 

Leaves variously lobed, the lobes broader and not linear. 
Leaves of an ovate or ovate-lanceolate type, or if about as 
broad as long the lobes ovate-lanceolate, and, espe- 
cially, the male inflorescence long, narrow, lax. 

C. heterophylla. 

Leaves typically palmate-suborbicular or when rarely 
simulating C. heterophylla, the male inflorescence 
relatively short, congested. 



FLORA OF PERU 135 

Male and female peduncles subequal; leaves 5-lobed. 

C. gossypiifolia. 

Male and female peduncles very unequal. 
Leaves glabrous or glabrate. 
Leaf lobes 7, at least the basal overlapping. 

C. parviflora. 

Leaf lobes not overlapping at the open base. 
Leaf lobes 3, entire; flowers monoecious; fruits 

scarlet C. erythrocarpa. 

Leaf lobes 5-7, or 3 but at least 1 lobe then more 

or less lobed. 
Leaf lobes oblong-lanceolate, acute; flowers 

monoecious; fruits yellow C. monoica. 

Leaf lobes ovate; flowers dioecious. 
Leaf divisions broadly ovate; style short or 

obsolete C. paniculata. 

Leaf divisions narrowly acute; style 5-6 mm. 

long C. stylosa. 

Leaves minutely but densely pilose beneath, especially 
on the nerves C. pubescens. 

Carica acuta 0. Heilb. Sv. Bot. Tidskr. 30: 220. 1936. 

Dioecious-flowered tree similar to C. lanceolate, but the narrower 
leaves long- and acutely narrow-acuminate; male flowers 12-14 mm. 
long, the rather rounded-triangular sepals to 1 mm. long, the corolla 
tube 8-10 mm. long; anther connective of short stamens slightly 
protruded; female flowers 12-16 mm. long otherwise with the sepals 
similar to the male; style indistinct, the 5 stigmas linear and entire. 
To 10 meters high, flowering at leafing. Female flowers illustrated, 
Heilb. I.e. 219. Harms I.e. 100, referred these collections to C. lan- 
ceolata. Sap whitish, flowers greenish. 

Apurimac: Among Cacti and shrubs, Hacienda Cuycuhua, 2,800 
meters, Weberbauer 7170; 7171 (types). "Jalasacha." 

Carica August! Harms, Notizbl. Bot. Gart. Berlin 11: 280. 1931. 

Simple-stemmed glabrous shrub with long-petioled oblong-elliptic 
or lanceolate, sometimes rather obovate, irregularly dentate leaves 
that are white-glaucous beneath and sparsely provided with coarse 
setae or gland-like processes on the nerves and veins, a number of 
them also congested at the juncture with the petiole to form a ring; 
leaves 2 dm. long, 8 cm. wide, often larger, rounded at base, acumi- 



136 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nate; male racemes 6-11 cm. long, the sparsely pilose pedicels 3-4 
mm. long; calyx barely 2 mm. long, 5-6-parted nearly to base; 
corolla 15-19 mm. long, the tube 9-12 mm. long, the 5 lanceolate 
segments acute, 6-8 mm. long; longer filaments sparsely pilose, the 
anther connective shortly produced as also for the shorter stamens; 
female peduncles 14-16 cm. long; young fruit lance-ovoid, narrowly 
winged. 

Recalls in its gray-backed toothed leaves C. Weberbaueri but 
that species has 3-lobed leaves; it also suggests C. glandulosa as 
regards the glaucous leaves and the glands but the leaves of the 
latter are entire margined; so, with the related C. Weberbaueri, 
it may meritoriously, as Harms has appropriately decided, per- 
petuate the name of the great student of the Peruvian flora. To 
3 meters high, the flowers greenish. 

Ayacucho: In shady forest among shrubs, Choimacota Valley, 
2,700 meters, Weberbauer 7563, type. "Monte papaya." 

Carica candicans Gray, U. S. Expl. Exped. Bot. 640. 1854; 
177. Vasconcellea candicans (Gray) A. DC. Prodr. 15, pt. 1: 
417. 1864. C. integrifolia Raimondi, Elem. Bot. 2: 230. 1857, fide 
Weberbauer. 

Sparingly branched, foliate only during the winter or wet season, 
with broadly ovate, subcordate palmately 3 (-5) -nerved entire or 
sinuate margined leaves, green above, thinly white-tomentose- 
floccose beneath; leaves sometimes with 3-4 large lateral teeth, 
attaining 2 dm. in length; male peduncles 2.5-5 cm. long, shorter 
than the petioles, the many greenish-white flowers shortly pedicelled; 
calyx 5-7-lobed; corolla tube 12 mm. long, the 5-7 lobes half as long; 
female flowers solitary, at least usually on pedicels 2-2.5 cm. long, 
the lanceolate sharp calyx teeth about 3 mm. long, the corolla lobes 
2-2.5 cm. long, narrowly lanceolate; fruiting pedicel 5-7 cm. long, 
the fruit cucumber-shaped, blunt at the smaller base and at tip, 
obscurely 5-angled, 10-13 cm. long, 3-4 cm. thick, the many seeds 
imbedded in the fleshy pulp; sarcotesta (outer husk) of the seeds 
slimy within, the seeds ovoid, about 8 mm. long without the sarco- 
testa, the inner seed coat brown and nearly smooth when dry 
(Harms). 

Low tree (to about 3 meters high) with thick, often sprawling 
or irregularly growing trunks, characteristic for the lomas and the 
western Andean slopes to around 3,000 meters. On the coast it 
blooms during the dry period or summer season and many examples 
in this leafless state are nearly bizarre, especially if standing alone 



FLORA OF PERU 137 

on some stony outcrop; it grows during the moist or rainy season 
which on the coast corresponds to winter. The fruits are called 
"Mito" and have a good odor and not unpleasant taste (Ruiz & 
Pa von); when ripe the taste is sweet and the aroma delightful 
(Raimondi). Weberbauer, opposite page 143, shows a photograph 
of a tree in leaf, referring to it frequently in the text, 117, 118, 143, 
158, 161, 163, 164, 173, 174, 176. I found it commonly eaten but 
dry, 'fibrous and seedy, the flavor, however, pleasing. 

Lima: Between Lima and Obrajillo, U. S. Exped., type. Obra- 
jillo, Ruiz & Pavon. Viso, among rocks, 770. Matucana, rocky 
place, 2,370 meters, Weberbauer 267; 1693. Lomas, Mount Amancaes, 
300 meters, Weberbauer 5717. Arequipa: Prov. La Union, Cota- 
huasi, 2,700 meters, open shrub, cactus, herb formation, Weberbauer 
6865 (det. Krause). Moquehua, Weberbauer 7354. "Ulicana," 
"papayo," "mito," "jerju." 

Carica erythrocarpa Linden & Andre, 111. Hort. 18: 31. pi. 51. 
1871; 184. 

Trunk simple; petioles grooved above, 10-20 cm. long; leaves 
3-lobed, the lobes ovate-lanceolate, entire, acuminate, green above, 
paler beneath and with some evanescent hyaline trichomes; calyx 
of masculine flowers minute or obsolete; fruit shortly peduncled, 
scarlet, subcostate, lustrous, the size and shape of a hen egg but 
shortly cusped at tip, with many large black suberose seeds. 
Fruit at first taste sweet, but later disagreeable (Andre"). 

Peruvian locality according to Wallis, "Reiseerinnerungen," in 
Gartenflora 25: 301. 1876, but described from plants cultivated by 
Linden, the seed supposed to be from region of Guayaquil, collected 
by Wallis, apparently otherwise unknown. 

Carica heterophylla Poepp. & Endl. Nov. Gen. & Sp. 2: 60. 
pi. 182. 1838; 180. Vasconcellea heterophylla A. DC. Prodr. 15, pt. 1: 
417. 1864. 

Often low or to 2 meters high, with ample membranous glabrous 
leaves, ovate acute or acuminate, mostly entire or more or less 
hastate and 3-lobed, the middle segment broad and largest or now 
and then shortly lobed, the main division sometimes short, sometimes 
extending to below the middle, rarely even pinnately lobed; petioles 
4-6 cm. long or longer; male racemes very lax, greatly elongate, 
narrow and little branched, the female shortly peduncled, 1-3- 
flowered; male flowers slender, 16 mm. long, the tube scarcely more 
than 0.5 mm. wide, the lobes 5 mm. long, very narrow; fruit the size 



138 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and form of a walnut, the seeds 7 mm. long with tubercles nearly 
2 mm. long. Sometimes to 5 meters high, the male flowers yellowish 
(me). Highly variable in leaf -character but the oblong, loose 
male inflorescence distinctive. C. baccata Heilb. Acta Hort. Berg. 9: 
105. 1928, Ecuadorian, has ovate subpinnatifid leaves, 3 acute lobes 
widely spreading on each side, the male inflorescence short. Flowers 
white (Klug). 

San Martin: San Roque, Klug 7249 (det. Standl.). On Rio 
Mayo near Tarapoto, Spruce 4345 (det. Solms-Laubach) ; Williams 
5237; 5261; Killip & Smith 27627 (all det. Harms). Loreto: 
Yurimaguas, Poeppig, type. Pumayacu, Klug 3162 (det. Standl.). 
Near Iquitos, Klug 648 (det. Harms). Cerro de Cumbaso, Ule 
6758. Rio Itaya, Killip & Smith 29424; 29793 (det. Harms). 
Brazil. "Mamorana." 

Carica lanceolata (A. DC.) Solms-Laubach, I.e. 179. Vascon- 
cellea lanceolata A. DC. Prodr. 15, pt. 1: 416. 1864. 

Fascicles of leaves and flowers sometimes from the older branches; 
petioles to 2.5 cm. long; leaves glabrous, unparted and entire- 
margined, lanceolate, obtuse or nearly cordate at base, acute or 
obtusely acuminate, to 8 cm. long, 2.5 cm. wide, glaucous beneath 
and sometimes with a few tubercles near the petiole; male peduncles 
often twice as long as the petiole, many-flowered, the corolla tube 
6-8 mm. long; anther connective shortly extended above the cells. 
Doubtless, if Peruvian (!), from southern Peru. The sap is orange- 
colored, the plant poisonous (Kuntze). Female flowers 10-12 mm. 
long; ovary with truncate top, the style 2 mm. long, the stigma 
linear, entire (Heilborn). F.M. Neg. 38430. Illustrated, female 
flowers, Heilborn, Sv. Bot. Tidskr. 30: 219. 

Cuzcof?): Gay, type. Bolivia; Argentina. 

Carica monoica Desf., Ann. Mus. Hist. Nat. Paris 1: 273. 
pi. 18. 1802; 183. Vasconcellea monoica (Desf.) A. DC. Prodr. 15, 
pt. 1:418. 1864. 

Petioles 12-16 cm. long; leaves suborbicular, only shallowly cord- 
ate, ample, glabrous or sometimes granulate on the nerves near the 
petiole, often 3-lobed with oblong-lanceolate pointed lobes, only 
4-6 cm. wide, the middle lobes mostly 3-lobed, or 5-7-lobed, the 
margins entire or lobulate at base; flowers monoecious, the inflores- 
cence short, thick, with few blossoms; tube of male corolla 14-16 mm. 
long, villous within, the lobes oblong; connective of lower anthers 
produced, ovate, subcrenate; upper anthers glandular only apically; 



FLORA OF PERU 139 

female petals 18 mm. long, linear-oblong. Described from cultivated 
specimens, Paris. Fruit insipid but the leaves a substitute for 
cabbage; the Indians of Pampa Hermosa collect the seeds in the 
quebrada of Chinchao where the plant grows abundantly (Ruiz & 
Pavon). F.M. Neg. 13701. Illustrated, Bot. Mag. pi. 3633. 
Huanuco: Pozuzo, Ruiz & Pavon. "Col de montana." 

Carica paniculata Spruce ex Solms-Laubach, I.e. 177. C. lep- 
tantha Harms, Notizbl. Bot. Gart. Berlin 8: 95. 1922. 

Glabrous dioecious shrub; petioles 12 dm. long or longer; leaves 
broadly ovate or even suborbicular, cordate at base, membranous, 
above the middle 3-5-lobed, the middle lobe ovate or obovate, the 
lateral longer, all entire or often with broad, short, lateral lobes or 
the lateral rarely trilobed, obtuse, acute or all acuminate; male in- 
florescence many-flowered, divaricately branched at apex, sometimes 
sparsely puberulent, the pedicels obscure or lacking; flowers rose- 
colored; calyx 1-1.5 mm. long, shortly 5-denticulate; corolla tube 
10-15 mm. long, the lobes 4-6 mm. long; filaments shortly coales- 
cent at base, the anther connective of the outer scarcely or not pro- 
duced, that of the sessile inner long-extended; peduncle of fruiting 
inflorescence 2-4 cm. long with 2-5 pedicels 4-7 mm. long; fruit 
ovate-lanceolate, scarlet, pointed, 2-2.5 cm. long, the obtusely ver- 
rucose seeds oblong-ovoid, 5-6 mm. long. Shrub 0.5-2 meters high, 
leafless at flowering. The leaves (6009) with 3-lobed middle lobe, 
sometimes with small, broad, lower lobes (Weberbauer) . Descrip- 
tion after Harms. 

Nearly as variable as C. heterophylla in leaves but all sinuses, 
basal as well as lateral, very open and rounded, the lobes rounded to 
a short point or even obtuse. Allied to C. parviflora, 177, with deeply 
7-lobed leaves, the lobes mostly 3-lobed, these narrower, the more 
sharply acute divisions overlapping, especially the basal ones, as not 
at all in C. paniculata. C. stenocarpa Heilb. Sv. Bot. Tidskr. 30: 221. 
1936 of northern Bolivia appears to be related but the leaf sinuses 
are extremely sharp. Perhaps Weberbauer 7080 at 2,300 meters, be- 
low Hacienda Soquian, Libertad, a 5-meter shrub with tiny greenish 
male flowers, no leaves, belongs to C. paniculata or to a new related 
species (Harms). Cf. also the species mentioned under C. pubes- 
cens which, being glabrous, would be sought here. From the material 
seen Harms seems to have relied on a drawing in Herb. Berlin in 
separating his species. F.M. Negs. 13700 (leptantha); 13702. 

Tumbez: Hacienda Chicana, Weberbauer 7659b; 7659c. Caja- 
marca: Below San Pablo, 1,000 meters, Weberbauer 3803. Los Reyes, 



140 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

near Cajamarca, Weberbauer 4806. Piura: Above Piura at Chanro, 
Weberbauer 6004. Between Chanro and the Haciendo San Antonio, 
500 meters, Weberbauer 6009 (see remarks above). Ecuador. 

Carica parviflora (A. DC.) Solms-Laubach, I.e. 177. Vasconcellea 
parviflora A. DC. Prodr. 15, pt. 1: 417. 1864. 

Large-leaved species with small and delicate male flowers; 
petioles 1-2 dm. long; leaves lighter green beneath, appearing pel- 
tate by the overlapping sinuate lower lobes, deeply 7-parted, the 
obovate lobes mostly 3-lobed, acute or acuminate; male inflorescence 
long-peduncled, many-flowered, the pedicels 2 mm. long; calyx 
minute; corolla tube 8 mm. long, villous within, the lobes 3-4 mm. 
long, ovate-oblong; upper stamens apically 1-glandular, the con- 
nective of the anthers of the lower shortly produced. C. paniculata 
Spruce is distinguishable, according to Solms-Laubach, by the diver- 
gent lobes of the leaves. 

In all probability this or C. pubescens is the species referred to by 
Ruiz and Pavon as the "seven-lobed Carica" which they found at 
Pozuzo: the fruit, called "papaya," is the size of a small melon with 
a taste somewhat similar, but to improve it longitudinal incisions 
must be made in it some days before eating so that the milky sap 
that is somewhat bitter may drain (Ruiz & Pavon). 

Huanuco: Pozuzo (cf. above), Ruiz & Pavon. 

Carica platanifolia Solms-Laubach, I.e. 182. 

Stem succulent, the branches elongate; petioles 3 dm. long, the 
large chartaceous glabrous leaf blades 3-5-lobed, sometimes to the 
middle, the broadly acuminate divisions with a few sharp teeth; 
both male and female inflorescences densely flowered and long- 
peduncled, the peduncle even to 2.5 dm. long; male flowers about 10 
mm. long, the calyx small, the linear acute lobes and slender corolla 
tube subequal, only the lower anthers with a short process; female 
flowers in pseudo-umbels of 5-6, 2-5 cm. long, 5 mm. broad, the petals 
extremely narrow, the calyx extremely minute. Solms-Laubach 
suggested the original locality as Cuchero; probably Peru or Ecuador. 

Peru(?): Without locality, Ruiz & Pavdn. 

Carica pubescens (A. DC.) Solms-Laubach, I.e. 185. Vascon- 
cellea pubescens A. DC. Prodr. 15, pt. 1: 419. 1864. 

Branches, petioles, leaves beneath and inflorescences lightly 
pubescent, the pubescence persisting, especially on the leaf nerves 
beneath; petioles greatly elongate, the large leaves broadly cordate, 



FLORA OF PERU 141 

deeply 5-7-parted, the middle lobe 3-5-lobed, the oblong lower and 
lateral lobes oblong, mostly 1-lobed; the lobes not rarely have 1 or 2 
broad teeth and the middle lobe is usually 3-lobed with elongate 
pointed lobules, the lateral lobes descending and mostly with a 
lobule on the outer lower side; male peduncles 2.5-6 cm. long; calyx 
deeply 5-parted, the lobes less than 2 mm. long; corolla glabrous 
within, the tube about 8 mm. long, the linear-lanceolate lobes as 
long; connective of all anthers shortly produced, the filaments of 
the upper glabrous; female inflorescence (Weberbauer 6997) very 
short, few-flowered, the corolla segments 2.5 to nearly 3 cm. long, 
the young fruit with five 2-branched styles. Description compiled 
from the five original sheets, and Harms, I.e. 97, as other descriptions 
herewith; Harms saw only a scrap of the type but identity with the 
Weberbauer plant seems certain. Much planted in mountain villages 
in the east, the fruit about the size of a hen's egg and eaten when 
cooked with sugar, but not raw. Flowers greenish, mostly among 
the leaves but also below on the old wood; sometimes only male 
sometimes only female on the same shrub (Weberbauer). 

The cultivated "Chamburu" of Ecuador, C. candamarcensis Hook, 
f. Bot. Mag. 101: pi. 6198. 1875, has pleasant-tasting yellow fruits, 
5-celled stigmas (bifid) similar to cacao pods in shape and size. 
Heilborn in Arkiv Bot. 17, pt. 12: 1, 2. 1921 has described two 
related, but glabrous species or races, viz., C. chrysopetala (flowers 
reddish-yellow) and C. pentagona, this known as "babaco" it has 
a distinctly sympetalous female corolla and has shown that the 
Hooker species is Ecuadorian and, in fact, dioecious. F.M. Neg. 
8512. 

Huanuco: Cultivated at San Rafael, near Ambo, 2,700 meters, 
Weberbauer 6685; also 6997, fide Harms, but locality not given by 
him. Pozuzo, Ruiz & Pawn, type. "Chamburu," "chiluacan." 

Carica stylosa O. Heilb. Sv. Bot. Tidskr. 30: 218. 1936. 

Tall glabrous herb or with a few trichomes on the principal nerves 
and petioles; leaves palmate, cordate at base, even to 5 dm. long, 
deeply 5-parted, the narrow acute lobes deeply pinnatifid; male 
inflorescence 1.5-2 dm. long, the flowering branchlets short, many 
flowered; sepals about 1 mm. long; corolla tube 18 mm. long, the 
segments 7 mm. long; connective of outer stamen anthers scarcely 
produced, that of the inner long-extended; female inflorescence to 
7 cm. long, the flowering branchlets to 5 cm. long; flowers 2-2.5 cm. 
long, the petals linear; ovary 5-celled, the style 5-6 mm. long, the 



142 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5 stigmas 10-15 mm. long, linear, entire. Illustrated, Heilborn, I.e. 
219 and 221. 

San Martin: San Roque, southeast of Moyobamba at edge of 
manioc and maize plantations (Douglas Melin 175; 279, types). 

Carica Weberbaueri Harms, Notizbl. Bot. Gart. Berlin 8: 
94. 1922. 

Shrub or tree, the branches probably as the elongate petioles 
sparsely hispidulous or scabrous; leaves ovate or oblong-ovate, 
rounded or emarginate at base, often acuminately 3-lobed above 
the middle, large (the middle lobe often 2.5-4 dm. long), the lobes 
more or less dentate-margined or even incised, puberulent-pilose on 
the nerves and veins beneath as well as glaucous; male inflorescence 
3-20 cm. long, sometimes hispidulous-pilose, the branchlets 1-2 cm. 
long or shorter, the pedicels 2-5 mm. long; calyx barely 2 mm. 
long; corolla tube 2 cm. long or longer, the width (pressed) 4 mm., 
little pilose within, the divisions 12-15 mm. long; longer filaments 
sparsely pilose, the connective shortly produced, the connective of 
the subsessile inner stamens sparsely pilose dorsally and produced; 
apical gland none or minimal. No other species except C. Augusti 
has such strongly toothed leaves. To 9 meters high, the blossoms 
brownish-green. 

Libertad: In evergreen bush, 2,000 meters, valley system of the 
Mixiollo, Weberbauer 7072, type. Ecuador. 

JACARATIA [Marcgr.] Endl. 

Branching shrubs or tall trees, aculeate at least below, with 
palmately divided leaves, the 5-12 leaflets sessile or petiolate. 
Calyx and corolla divisions more or less clearly opposite at least in 
the male flowers. Filaments usually somewhat coalescent. Other- 
wise like Carica, all species considered, and only to be regarded as 
a genus for convenience. 

Jacaratia digitata (Poepp. & Endl.) Solms-Laubach, Mart. Fl. 
Bras. 13, pt. 3: 191. 1889. Carica digitata Poepp. & Endl. Nov. 
Gen. & Sp. 2: 60. 1838. J. spinosa (Aublet) A. DC. var. digitata 
(Poepp. & Endl.) A. DC. Prodr. 15, pt. 1: 419. 1864. 

Tall tree with white, soft, spongy bark, the trunk and branches 
armed with stout broad-based prickles, these, reduced in size, some- 
times extending even to the branchlets and petioles; petioles 15-20 
cm. long, the leaves 5-7-foliate, the shortly to markedly petioled 
leaflets becoming obovate-lanceolate, abruptly acuminate, entire, 



FLORA OF PERU 143 

glabrous, 1-1.5 dm. or so long, about half as wide; male peduncle 
as long or longer than the petioles, the flowers subsessile, 1.5 cm. 
long, their stamens unequal; fruit oblong, obscurely costate, the 
seeds with a fragile brown coat. To 30 meters high. The male 
flowers are extraordinarily malodorous (Poeppig). The bark yields 
an astringent sap (Williams). 

The tree, according to Poeppig, is notorious among the Indians 
for its poisonous qualities; they even fear to sleep beneath it lest 
they become afflicted with sores which become gangrenous and which 
the sap is said to cause. Poeppig describes the fruits, however, as 
insipid but not poisonous, inodorous, yellow. Klug, on the other 
hand, observed that the inhabitants make holes in the soft bark; 
these holes then become filled with maggots (Coleoptera) the size 
of a fat finger, which are collected for food. Illustrated, Mart. Fl. 
Bras. 13, pt. 3: pi. 51. 

San Martin: Juanjui, 400 meters, Klug 3816. Loreto: Rio 
Cochiquinas, Yurimaguas, Poeppig, type. Rio Itaya, Williams 3342. 
Puerto Limon, Tessmann 3858. Rio Acre: Ule 9646; Krukoff 5446. 
Near Iquitos, Klug 1222. Brazil. "Shamb-uru," "chamburu," 
"papaya-haspi," "mamao brabo." 

LOASACEAE 

Reference: Urban & Gilg, Nova Acta Acad. Leop.-Carol. 76, 
pt. 1. 1900. 

Usually erect or climbing herbs characteristically more or less 
provided with barbed and often stinging hairs, the plants not infre- 
quently "sticky" to touch. Leaves opposite or alternate. Flowers 
hermaphrodite, the 4-7 calyx lobes persistent, the petals as many, 
generally 5. Stamens usually indefinite, staminodes often present, 
and often nectariferous scales, various in size and form. Ovary 
usually inferior, 1-5-celled, the style entire or 2-3 parted, the fruit 
a 1-3-celled capsule. For convenience the above work is hereafter 
referred to as Monogr. Loas. It is detailed and painstaking, and the 
following synopsis of the many Peruvian forms is compiled from it. 

Stamens 5; vine, cucurbitaceous in aspect 1. Gronovia. 

Stamens 10-many. 

Stamens all fertile or the outer petaloid or filiform . . 2. Menlzelia. 
Stamens in fascicles opposite the petals and fertile, others alternate, 

sterile, mostly transformed into scales. 
Petals 4; staminodia free. 



144 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Fertile stamens 4-14, 1-4 before the petals, 6-10 sterile, 1-3 
before the sepals; capsule spiraled 3. Sclerothrix. 

Fertile stamens 12-28, 3-7 before the petals, 16-20 sterile, 
4-5 before the sepals; capsule straight. . . .4. Klaprothia. 

Petals 5-7; outer staminodia united into a nectariferous scale. 

Capsule straight, clavate or obconic, dehiscent apically be- 
tween the lobes; Peruvian species not scandent.5. Loasa. 

Capsule usually spirally distorted, dehiscing laterally from 
below; plants often vines 6. Cajophora. 

1. GRONOVIA L. 

Vine closely simulating some Cucurbitaceae by reason of the 
cordate leaves with lateral, divaricate, acuminate lobes and the 
peduncled cymose cylindrical flowers. Flowers subtended by fili- 
form bracts. Sepals more or less connate. Staminodia none. 

Gronovia scandens L. Sp. PI. 202. 1753; 11. 

Stems sparsely spreading hispid ulous and also provided with 
conical hooked processes by which the plant "holds"; leaves very 
sparsely asperous, the sinus broad, the lobes entire or with 1-3 
teeth; inflorescence 1-4 cm. long, 10-18-flowered; petals 3-4 mm. 
long, shorter than the sepals. Illustrated, Baillon, Hist. PI. 8: 463. 

Piura: Ayavaca, 1,000 meters, Weberbauer 6394. Near Serran, 
250 meters, Weberbauer 5995. Tumbez : Plain southeast of Hacienda 
La Choza, Weberbauer 7698. North to Mexico. 

2. MENTZELIA L. 

Reference: Darlington, Ann. Mo. Bot. Card. 21: 103-226. 1934. 

Variable as Loasa in vegetative character but the flowers often 
leafy-bracted ; otherwise marked as indicated in the generic key. 
The following is compiled from Miss Darlington's monograph, which 
is so satisfactory as regards the species of North America that her 
treatment is accepted; however, on the basis of material seen, the 
validity of the South American forms in several instances seems open 
to question. In any case more material is needed to prove or dis- 
prove the soundness of the characters here used as indicating specific 
lines; accordingly the monographer's key is appended. 

Calyx tube and capsule cylindrical, somewhat attenuate at base. 

M . aspera. 



FLORA OF PERU 145 

Calyx tube obconical or turbinate, distinctly attenuate below, 

usually pedicellate. 
Leaves sessile; stamens in 2-3 series. 

Stamens 40-50, in 2 (rarely 3) series M. ignea. 

Stamens 50-60, in 3 series M. cordifolia. 

Leaves petiolate; stamens in 4 series M. Fendleri. 

Mentzelia cordifolia Dombey ex Urb. & Gilg, Monogr. Loas. 
74. 1900; 121. 

Often bushy spreading, the papery bark of the older branches 
defoliating; leaves sessile, the upper more or less cordate clasping 
usually with a short lobe at base each side and a few coarse teeth, 
to 6 cm. long, 3.5 cm. wide, scabrous; flowers orange, terminally 
crowded; petals 10-20 mm. long, half as wide, glabrous; calyx tube 
with stipe 7-8 mm. long; capsule subcylindrical attenuate at base to 
stipe, 2-8 mm. long, scabrous, the seeds apparently not margined. 
But the seeds as to type of M. cordifolia are immature! M. sora- 
tensis Urb. & Gilg, 68, from northern Bolivia seems to be only a form 
with somewhat narrower leaves, capsules subsessile; its seeds are 
apparently margined and to it Miss Darlington has referred Pennell 
13069 from Tiabaya, Arequipa, considered by Killip as M. ignea, 
which see. Possibly the specimen from southern Peru will prove to 
be a variety of one of the older species. The petals of the material 
from Chancay are about 15 mm. long. Common according to 
Weberbauer, 100, 158, 163. A specimen by Ward from Matucana 
was det. by Ball, M. hispida Willd., a similar but larger-flowered 
Mexican species. F.M. Neg. 24161. 

Lima: Chancay, Ruiz & Pavon; Dombey, type. San Buena- 
ventura, Pennell 14568 (det. Darlington). Near Viscas, Pennell 
14472 (det. Darlington). Quive, Pennell 14313 (det. Darlington). 
Matucana, 130 (det. Killip); Weberbauer 89; 160. Cajamarca: San 
Miguel, Weberbauer 3923; 188, 189. Huanuco : Stony slopes, 3215 
(det. Killip). Arequipa: Raimondi. Mejia and Pasco ((Zunther & 
Buchtien, det. Bruns). Cuzco: Apurimac, Herrera 1220. Valle del 
Urubamba, Weberbauer, 175. Ecuador to Bolivia. "Manca-r'ajra," 
' 'manca-ppaqui. ' ' 

Mentzelia Fendleriana Urb. & Gilg, Mem. Torrey Club 3, pt. 
3: 34. 1893; 117. 

With the habit and characters of M. cordifolia but the middle and 
lower leaves at least minutely petiolate and narrowly ovate-lance- 



146 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

olate or oblong-lanceolate, green; petals 17-22 mm. long; stamens 
70-80 in four series; capsules subsessile to well-pedicelled; seeds of 
M. cordifolia. M. scabra HBK., 116, credited to "Colombia, 
Ecuador and Peru" by the monographer, who, however, cites speci- 
mens only from Colombia, has 100-120 stamens and ovate leaves. 

Piura: Talara, Haught 5. Lima: (Cuming 1011}. Junin: Uspa- 
chaca, 1296 (det. Killip). Cuzco: Ollantaytambo (Cook & Gilbert 
280; 54-5, fide Darlington); Hen era 3413. Near Urubamba, H err era 
1529; 1336 (det. Killip). Yucay, Soukup 575. Anta, Vargas 299 (det. 
Standl.). Ecuador; Bolivia; Venezuela; Chile? "Manca-ppaqui." 

Mentzelia ignea (Phil.) Urb. & Gilg in Engler & Prantl, Pflan- 
zenfam. 3, 6a: 110. 1894; 122. Loasa ignea Phil. Anal. Mus. Nac. 
Chile, Bot. 1891:24. 1891. 

Similar to M. cordifolia but with only 40-50 stamens in 2 or 3 
series; seeds ovate-oblong, grayish or brownish-yellow, irregularly 
striate-verruculose, not winged, thus as in M. cordifolia and probably 
the earliest name for the species sens lat. F.M. Negs. 10127; 38488. 

Lima: (Gay, fide Urban & Gilg). Amazonas: Chachapoyas, 
Mathews 596; Weberbauer 4360 (det. Urb. & Gilg). Arequipa: 
(Cockerell, fide Darlington). Tiabaya, Pennell 13069 (det. Killip). 

3. SCLEROTHRIX Presl 

Ancyrostemma Poepp. & Endl. Nov. Gen. & Sp. 3: 65. pi. 272. 1845. 
Essential characters as in key. 

Sclerothrix fasciculata Presl, Symb. Bot. 2, pt. 6: 4. pi. 53. 
1833. Ancyrostemma micranthum Poepp. & Endl. I.e. Blumenbachia 
micrantha [R. & P.] G. Don, Gard. Diet. 3: 62. 1834. 

Annual with opposite, thin, minutely dentate, petioled leaves 
about 3 cm. long, 1.5 cm. wide, often much larger, ovate-oblong- 
elliptic, acute both ends, sparsely short-pilose both sides, as the 
stem; flowers few, terminal on short axillary branchlets, the petals 
about 1.6 mm. long; capsule tardily spiraled, 4-9 mm. long. Illus- 
trated, Mart. Fl. Bras. 13, pt. 3. pi. 53. 

San Martin: Tarapoto, Ule 6415. San Roque, Williams 7625; 
7017. Amazonas: Chachapoyas, Mathews. Huanuco: Cuchero, 
Poeppig (type, A. micranthum}. Pozuzo, 4545. Cuzco :Gay. Rosa- 
lina, 650 meters, Bues. Junin: Pichis Trail near San Nicolas, 
Killip & Smith 26124. Brazil to Mexico. 



FLORA OF PERU 147 

4. KLAPROTHIA HBK. 

Sprawling herb with opposite, ovate, dentate leaves and few- 
flowered cymes of rather small white flowers. Essential character 
as in key. 

Klaprothia mentzelioides HBK. Nov. Gen. & Sp. 6: 123. pi. 
537. 1823. 

Leaves ovate, rounded at base or little contracted to the petiole, 
acuminate, 5-8 cm. long, 2-4 cm. wide or larger, somewhat scabrous- 
pilose, bullate-rugose in age above; petals 4, 6-8 mm. long; capsules 
obovate. 

Junin: Pichis Trail, Dos de Mayos, 1,700 meters, Killip & Smith 
25878. Puno: Region of Sandia, 1,500 meters, Weberbauer 1070 
(det. Urban). In the mountains, Bolivia to Colombia and Venezuela. 

5. LOASAAdans. 

Annual or perennial herbs, usually with stinging hairs, sometimes 
half shrubs, rarely vines, the leaves alternate or opposite, the flowers 
generally yellow, less frequently white or red. Flowers 5-7-merous, 
petals plane, concave, naviculiform or cucullate. Nectar-scales 
from 3 (4-7) staminodia, connate, concave or sac-like, usually with 
3 (2-7) threads from the back or below the tip, the 2 (-4) interior 
staminodia linear or subulate. Stamens many, the filiform filaments 
in fascicles opposite the petals. Ovary rarely somewhat superior, 
always 1-celled. Capsule straight, clavate or obconic, rarely sub- 
globose, dehiscent between the calyx lobes, the seeds rarely few. 

In all the Peruvian species the scales are 3-nerved and the stam- 
inodia 2, and none truly scandent have yet been found. The char- 
acters said to separate L. picta from similar species may not be 
specific. 

Capsules 4-5-valved the valves placed before the calyx lobes; leaves 
rather ashy-puberulent both sides; branchlets sublignescent with 
whitish lustrous defoliating thin bark L. incana. 

Capsules 3-5-valved the valves alternating with the calyx lobes. 
Leaves, unless the uppermost, pinnate or trifoliate. 

Leaves mostly trifoliate L. tripkylla. 

Leaves pinnate or bipinnate L. urens. 

Leaves variously lobed, sometimes pinnately, or toothed to 
subentire. 



148 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels more or less adnate to the axis of the pseudoraceme; 

petals apparently 5-6 mm. long L. inconspicua. 

Pedicels not at all adnate to inflorescence axis; flowers longer. 
Scales not bisaccate at base, sometimes more or less inflated. 
Perennial with cordate-orbicular leaves; scales with threads. 

L. heucheraefolia. 
Annuals or if perennial the leaves not orbicular; scales 

naked. 

Petals 16-25 mm. long; scales more or less saccate. 
Petioles short, the upper leaves sessile; sepals ovate. 
Leaves ashy tomentose and esetulose beneath; inflo- 
rescence simple L. carunculata. 

Leaves setose on nerves; inflorescence pseudopa- 

niculate L. macrothyrsa. 

Petioles elongate, even the upper; sepals linear .L. laxa. 
Petals 7-13(15) mm. long; scales not at all inflated. 
Leaves opposite; scales appendaged below; seeds few, 

large L. nitida. 

Leaves opposite; scales smooth; seeds many .L. solaria. 
Leaves alternate; seeds many, small. 
Petals broadly clawed; leaves subentire, stipulate. 

L. Stuebeliana. 

Petals not clawed; leaves dentate, never stipulate. 
Scales with a horizontal fold or band; leaves 

deeply sinuate-dentate L. leiolepis. 

Scales with two horizontal bands; leaves unequally 

dentate L. chenopodifolia. 

Scales at or near base definitely bisaccate. 
Flowers small, the petals, especially if 12-25 mm. long, 

obviously clawed at base and cymbiform. 
Leaves not subpinnately lobed. 

Scales with several transverse plicae above the sacs. 

L. picta. 

Scales with 1-2 ridges between sacs and tip. 
Leaves esetulose or nearly. 

Petals entire; leaves sessile unless the lowest. 

L. kastata. 



FLORA OF PERU 149 

Petals toothed above the broadly clawed base; 

leaves petioled, serrate L. fulva. 

Leaves evidently setulose. 

Leaves subequally serrate, rounded or subcordate. 

L. Poissoniana. 
Leaves irregularly serrate, scarcely rounded at 

base L. ferruginea. 

Leaves subpinnately lobed, the toothed lobes extending 
about half way to the midrib. 

Petals about 15 mm. long; staminodia and scales sub- 
equal L. cuzcoensis. 

Petals about 25 mm. long; staminodia much longer 

than scales L. Vargasii. 

Flowers showy, the petals usually 2-5 cm. long or in any 

case not at all or very shortly clawed, sometimes 

cymbiform. 

Petals oblong or nearly, acute or obtusish, in any case 

not obovate and broadly rounded at apex. 
Leaves serrate or repandly lobulate. 
Subscandent plant with small (3 cm.) leaves. L. lenta. 
Erect or suberect plants, the leaves larger. 

Petals about 3 cm. long; petioles 1-3 cm. long. 

Upper petioles about 1 cm. long; scales efila- 

mentose L. solaria. 

Upper petioles 2.5-3 cm. long; scale filamentose. 

L. loxensis. 
Petals about 2 cm. long; petioles elongate. 

Leaves narrowed below L. olmosiana. 

Leaves rounded at base L. tingomariensis. 

Leaves irregularly lobed or coarsely serrate, often 
jaggedly. 

Petals somewhat cymbiform; scale neck entire or 
minutely bilobed. 

Petals 3-4.3 cm. long; leaves subequally lobed. 

L. grandiflora. 
Petals 5 cm. long; leaves unequally lobed. 

L. macrantha. 
Petals plane; scale neck 3-toothed L. magnifica. 



150 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petals obovate, broadly rounded at apex. 
Leaves suborbicular or reniform, often cordate. 

Leaves peltate L. peltata. 

Leaves not peltate. 

Petals 4 cm. long; leaves fleshy L. rugosa. 

Petals 2.5-3 (-4) cm. long; leaves thin or firm. 
Leaves opposite. 

Petals esetulose, 1 cm. wide L. Aspiazui. 

Petals slightly setulose, 2 cm. wide. 

L. limata. 
Leaves alternate. 
Leaves, even the lowest, repandly serrate. 

L. macrorrhiza. 

Leaves coarsely lobed-dentate. 
Petals 2.5 cm. long; scale neck 2-3-lobed. 

L. ranunculi/olid. 
Petals 3-4 cm. long; scale neck 3-lobed. 

L. magnified. 

Leaves ovate or triangular-ovate. 
Petals 4-5 cm. long. 
Petals 5 cm. long; leaves herbaceous. 

L. macrophylla. 
Petals 4 cm. long; leaves coriaceous. 

L. Weberbaueri. 

Petals about 2 (3) cm. long L. carnea. 

Loasa Aspiazui Macbr., spec. nov. 

Herba perennans; caulibus petiolisque conspicue bruneo-setosis 
paullo vel vix strigillosis; foliis superioribus suboppositis, fere orbicu- 
latis basi distincte cordatis, obscure lobulatis et sparse calloso- 
denticulatis circa 1 dm. longis et latis, supra sparsissime minuteque 
setulosis, subtus praesertim ad nervos longe setosis; floribus ut videtur 
in apice caulis circa 5, cincinnose collectis; pedicellis circa 1 cm. 
longis; calycis lobis adpresse bruneo-setulosis et minute strigillosis 
12 mm. longis, basi 4 mm. latis, acuminatis, tubus ovoideis 7 mm. 
crassus densissime rufo-setosus; petalis firmo-chartaceis puberulentis 
planis apice rotundatis 25-28 mm. longis, 10-11 mm. latis; squamis 
8 mm. longis, dorso sub medio bicalcaratis (calcaribus 5 mm. longis), 
filis nullis, alis ventralibus supra collum (collo 3-lobulato) fere 3 mm. 



FLORA OF PERU 151 

oblongo-productis; staminodiis pilosis 12 mm. longis; staminibus 
circa 20 mm. longis. 

Apparently related to L. alpina Urb. & Gilg, and with similar 
scales but that Ecuadorian species has deeply lobed leaves. Named 
for Dr. R. Aspiazu, Lima physician; cf. note under Fuchsia Aspiazui. 

Junin : Rio Masamerich, 3,200 meters, Weberbauer 6669, type. 

Loasa carnea Urb. & Gilg, Bot. Jahrb. 45: 469. 1911. 

Several dm. tall with sparsely setose herbaceous stem, but 
perhaps perennial; lower petioles to 12 cm. long, the upper leaves 
ovate-oblong, cordulate or truncate at base, acute, more or less 
deeply lobed, the ovate-triangular acute lobes obsoletely serrate, 
sparsely and shortly asperous both sides with a few setae intermixed 
on the veins and nerves, the larger to 8 cm. wide; inflorescence lax, 
few to 7-flowered, the pedicels to 1.5 cm. long; calyx tube about 
5 mm. long, to 5 mm. wide, densely setose, the ovate lobes 6-7 mm. 
long, 4 mm. wide at base; petals plane, obovate-oblong, rounded 
at apex, scarcely 2 cm. long, half as wide; scales 6-7 mm. long. 
Allied to C. aurantiaca Urb. & Gilg, 209, Ecuadorian, with denticulate 
leaf lobes, fewer larger flowers, the calyx lobes narrow and twice 
as long. Flowers flesh-colored; common shade plant (Weberbauer). 
F.M. Neg. 10182. 

Cajamarca: Woods by Chugur, 2,700 meters, Weberbauer 4082. 

Loasa carunculata Urb. & Gilg, Monogr. Loas. 243. 1900. 
L. vestita Killip, Journ. Wash. Acad. Sci. 19: 194. 1929. 

Stems erect, straight, to several dm. high, clothed with reddish- 
yellow smooth trichomes and smooth setae, both 1-1.5 mm. long, 
intermixed; leaves alternate, the lower shortly petioled, oblong- 
lanceolate, rounded at base, acute, deeply 8-12 lobed and dentate- 
serrate, to 17 cm. long, 7 cm. wide, laxly asperous above with 
bulbous based hairs, beneath gray-tomentose, the trichomes slender 
and smooth, setae completely lacking; calyx tube 7-8 mm. long, 
reddish-setose, the oblong-lanceolate lobes about 1 cm. long; petals 
apparently orange, cymbiform, 17-19 mm. long, contracted for 
two-fifths of their length into a claw 2-2.5 mm. broad at base; 
scales at the base about 1.5 mm. broad, at one-fifth their height 
more or less obscurely saccate-inflated, sometimes scarcely so, then 
narrowed to the top, this portion conspicuously calloused ; staminodia 
2, 9-10 mm. long, dilated at base, then suddenly geniculate-incurved. 
Illustrated, scales and staminodium, Urb. & Gilg, I.e. pi. 6. 



152 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Peru(?): Without locality, Lobb 358, type. Huancavelica : 
Gravelly hillsides, 2,400 meters, Mejorada, Stork & Horton 10905. 
Ayacucho: North of Huanta, grassy steppe, 3,500 meters, Weber- 
bauer 7591 (type, L. vestita). 

Loasa chenopodifolia Desr. in Lam. Encycl. 3: 580. 1791; 231. 
L. xanthiifolia Juss. Ann. Mus. Hist. Nat. Paris 5: 26. 1804. 

Erect annual to 4 dm. high, with erect spreading branches more 
or less densely pubescent with small retrorse trichomes; lower leaves 
often opposite, the lax upper alternate, the petioles to 3 cm. long, 
sometimes half as long as the ovate leaf blade, this to 5 cm. broad, 
unequally serrate and crenate-dentate, not at all setose, loosely 
asperous above, rather densely ashy puberulent-pilose beneath; 
flowers in pseudo-racemes, to 1.5 dm. long; calyx tube 7-8 mm. long, 
2.5-3 mm. broad above, pubescent with an ashy puberulence and 
brown setae intermixed, the lobes 4-5 mm. long, 2-2.5 mm. broad 
at base; petals white, 9-10 mm. long, narrowed but not clawed at 
base; scale 3.5-4 mm. long, 2.5-3 mm. broad at base, strongly 
saccate or inflated-convexed only in the upper part with 2 thickened 
parallel lines or folds, apically strongly callous-thickened, the neck 
recurved-crenulate, or 3-lobed, the 3 nerves ascending to the tip, 
the margin itself strongly produced into teeth 1.5 mm. long; 
staminodia about 5 mm. long, inflated basally. Illustrated, Tratt. 
Archiv Gewachsk. 1: pi. 26. F.M. Neg. 10183. 

Lima: Near Huamatanga, Dombey; Jos. Jussieu, type. Viso, 
577. Amazonas: Chachapoyas, Mathews 3002. 

Loasa cuzcoensis Killip, Journ. Wash. Acad. Sci. 18: 91. 1928. 

Stout stemmed annual several dm. high, the stem pubescent 
with small whitish hairs and brown setae intermixed; petioles 0.5-3.5 
cm. long, alternate or subopposite, the leaves ovate to lance-ovate, 
5-12 cm. long, 3-10 cm. wide, subpinnately lobed, the 7-9 acute 
lobes sharply serrate, hispidulous and sparingly setose; flowers 
5-merous, the peduncles to 3 cm. long; calyx densely and divaricately 
brown setose, the ovate lobes 5-6 mm. long; petals obovate, about 
15 mm. long, 6 mm. wide, cucullate distally, clawed toward base, 
slightly pilose without, apparently light yellow; scales triangular- 
ovate, about 5 mm. long, bisaccate toward base, the sacs much 
inflated, callous, thickened toward apex, otherwise smooth, the 
marginal teeth lanceolate, scarcely 1 mm. long; staminodia 2 in 
each scale, subulate, 4-5 mm. long; anthers purplish; capsule 2 cm. 



FLORA OF PERU 153 

long, nearly 1 cm. wide. Series Saccatae, apparently nearest L. fer- 
ruginea and L. Poissoniana. Is employed in popular medicine as a 
diuretic and cultivated for its flowers (Herrera). 

Cuzco: San Sebastian, 3,200 meters, Herrera 1465, type; also 
Cuzco in March, Herrera 2363, and at 3,000 to 3,600 meters, July, 
1923, Herrera; Soukup 41- Sacsahuamdn above Cuzco, Pennell 
13571 (probably, fide Killip). Valle del Urubamba, Herrera 3380. 
' ' Iscai-ckorota, ' ' ' 'china-quisa. ' ' 

Loasa ferruginea Urb. & Gilg, Monogr. Loas. 225. 1900. 

Erect, the slender (about 4 mm. thick) stems densely clothed 
with long brown setae and small white trichomes intermixed; lower 
leaves opposite, on petioles to 1 cm. long, the oval or oval-oblong 
blades to 2.5 cm. long, 17 mm. wide, unequally serrate-dentate, 
subrounded or subacuminate at base, acute, fulvous setose above 
and beneath on the nerves; calyx tube 4-5 mm. long, 3-4 mm. thick 
above, pubescent like the stem, the narrowly ovate lobes about 4 mm. 
long; petals 11-12 mm. long, narrowly clawed below one-third to 
one-fourth of their length, densely pilose without; scales about 4 mm. 
long, bisaccate above the base, strongly calloused and also thickened 
apically, and with a horizontal fold or band, the marginal teeth 
1.2-1.3 mm. long; staminodia 7 mm. long; capsule about 2 cm. 
long, 5-6 mm. wide. Probably should be included inL. Poissoniana. 
F.M. Neg. 10187. 

Cuzco: San Sebastian, 3,200 meters, Herrera 522. Puno: Lechler 
1897, type. Bolivia? "China quisa." 

Loasa fulva Urb. & Gilg, Monogr. Loas. 224. 1900. 

Closely allied to L. ferruginea but more branched and only 
sparsely setose; leaves esetose or rarely a few setae obvious on the 
midnerve beneath; petals 8-10 mm. long, broadly clawed one-fourth 
to one-fifth their length; staminodia about 5 mm. long, little exceed- 
ing the scales. Otherwise apparently like L. ferruginea. The 
indument on leaves beneath and on calyx is especially fine and 
short. There is an herbarium name by Ruiz & Pavon referring to 
the asperity. F.M. Neg. 10191. 

Peru: Without locality, the type cultivated in 1846 at Berlin. 
Lima: Santa Clara, Weberbauer 1672 (det. Urb. & Gilg); 145. Lima 
& Chancay, Ruiz & Pavon (det. Werderm.). 

Loasa grandiflora Desr. in Lam. Encycl. 3: 580. 1791; 202. 
L. cymbopetala Urb. & Gilg, Bot. Jahrb. 45: 468. 1911. 



154 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loosely bushy-branching when well-developed, the sparsely 
puberulent stems as well as the abundant rufous bristles lustrous; 
petioles, leaves both sides and calyx lobes puberulent-pilose, rather 
more densely pilose and cinereous beneath, scattered setose above 
and on the nerves especially on the under surface; lower leaves 
suborbicular and 5-lobed, the upper more ovate, deeply pinnately 
lobed, the lobes more or less lobed or unequally toothed; lower 
pedicels sometimes several cm. long; calyx lobes gradually acuminate, 
13-18 mm. long, 5-7 mm. wide; petals navicular-concave or probably 
when living less so, 3-4 cm. long, 10-15 mm. wide, medially setose 
without, thin; scales 13-17 mm. long, the 2 incurved subglobose 
calluses a fourth as long, the minutely notched front of the scale 
a little shorter than the triangular wings; staminodia narrowly subu- 
late, pilose on the middle part; filaments about 100, 2-2.5 cm. long, 
the anthers 3-3.5 mm. long; capsule 15-17 mm. long, pilose and 
setose, 3-valved; seeds 1.5-1.7 mm. long, elevated reticulate, black- 
ish. Petals and saccate base of green scales light red (K. & S.). 
There is sometimes, if indeed not always, more or less glandulosity. 
The species is the type of several segregated variants which may 
prove not to be distinct, and the key characters need proving by 
more collections. The long hairs are extremely stiff, easily pene- 
trating thick gloves; the flowers are deep orange (Ball). F.M. 
Negs. 10185; 24171. Illustrated, Urb. & Gilg, pi. 5 (flower parts); 
Juss. Ann. Mus. Hist. Nat. Paris 5: pi. 4- 

Lima: Chicla (Ball) ; Weberbauer 250; 169. San Mateo, Raimondi; 
Isern. Near Lima, Jos. de Jussieu, type; Dombey; Ruiz & Pawn; 
Mathews; Abadia. Rio Blanco, 2977; Killip & Smith 21710 (my 
collection det. Killip as L. macrorrhiza). Junin: Tarma, Ruiz 
& Pavdn (locality mistake?). Ancash: grass-shrub formation, 3,300 
meters, above Ocros, Weberbauer 2758 (type, L. cymbopetala). 

Loasa hastata Killip, Journ. Wash. Acad. Sci. 18: 92. 1928. 

Erect, somewhat succulent annual, the stem 12 mm. thick below, 
densely setose with pale setae about 0.5 mm. long, and toward the 
top, with many longer setae, brown and 2 mm. long; lower, nearly 
opposite, ovate-lanceolate leaves with petioles to 1.5 cm. long, the 
alternate upper sessile, more hastate, 3-10 cm. long, 1.5-7 cm. wide, 
acuminate to a subacute or obtuse tip, subcordate or subtruncate 
at base, irregularly sinuate-lobed, hispidulous but glossy above, 
finely pilose beneath; flowers 5-merous; calyx 3-4 mm. long, the 
lobes 6-7 mm. long; petals 13-15 mm. long, clawed below, green 
tinted, sparingly setiferous on nerves without; strongly inflated 



FLORA OF PERU 155 

scales about 6 mm. long, bright yellow and red shading into rose- 
pink, the neck with 2 thick horizontal bands, the lanceolate teeth 
1 mm. long; staminodia 8 mm. long; capsule 2 cm. long, 5 mm. broad, 
pilose and densely long-setose. Series Saccatae. 

Lima: Wet, sunny places, 2,500 meters above Matucana, 416, type. 

Loasa heucheraefolia Killip, Journ. Wash. Acad. Sci. 18: 
90. 1928. 

Perennial, the subligneous tomentulous and more or less setose 
stem to 1 cm. thick; setae very slender; leaves alternate or sub- 
opposite, the petioles 2-4 cm. long, the orbicular blades 3-6 cm. long, 
shallowly and irregularly 7-lobed, the lobes undulate-crenate, cor- 
date with narrow sinus, tomentulous, pilosulous and sparingly setose 
both sides; flowers white, 5-merous, axillary, solitary (?), the pedun- 
cles 1-1.5 cm. long; calyx 8-10 mm. long, 10 mm. wide, pilosulous- 
setulose, the lobes 8 mm. long, half as wide; petals 15 mm. long, 8 
mm. wide, narrowed at base; scales rectangular, 5 mm. long, 4 mm. 
wide, the 3 nerves slightly inflated and terminating in a filament 

3 mm. long; staminodia subulate, 12-14 mm. long; stamens about 
50; anthers oval, 1.2 mm. long. 

Ancash: Moist cliff pockets, Tambo de Pariocota, 2543, type. 

Loasa incana Grah. Edinb. Phil. Journ. 11: 169. 1830; 260. 
L. atriplicifolia Presl, Symb. Bot. 1: 61. pi. 39. 1832. 

Often bushy-branched and about 3 dm. (-2 m.) high, the branches 
characteristically marked by the white paper-like outer bark that is 
more or less breaking off; leaves alternate, ovate, often about 3.5 
cm. long, 3 cm. wide, truncate or slightly subcordate, acute, more or 
less deeply dentate on the sides or even lobulate; calyx tube about 

4 mm. broad, the entire ovate lobes 6 mm. long, petals 13-15 mm. 
long, cochleariform, rather narrowly clawed, glabrous within, green- 
ish-white; scales 3-nerved; capsule nearly subglobose. Plants more 
or less grayish puberulent-tomentose. Illustrated, Urb. & Gilg, I.e. 
pi. 7; Bot. Mag. pi. 3048. 

Piura: Parinas Valley, Haught 111; 186. Lima: Pavon; River o; 
Mathews; MacLean. Matucana, 257; Isern 2245. Quive, 1,200 
meters, Pennell 14309. Obrajillo, Ruiz & Pavon. Arequipa: Near 
Islay, Orbigny. 

Loasa inconspicua Urb. & Gilg, Monogr. Loas. 221. 1900. 
Annual, the slender stems about 2.5 dm. high, puberulent and 
above somewhat setulose; petioles 1 cm. long; leaves alternate, 



156 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ovate-oblong, equally dentate or with few large teeth alternating 
with the smaller, sparsely setose as well as scabrous above, densely 
pilose beneath, to 3 cm. long, 17 mm. broad; flowers in pseudo- 
racemes, the pedicels 13 mm. long, or shorter, always more or less 
adnate to the axis; calyx tube 7-8 mm. long, 2-3 mm. thick, the lobes 
3-4 mm. long, 1.5 mm. broad, rounded or scarcely acute; petals 
apparently only 5 or 6 mm. long; capsules about 14 mm. long, 3 mm. 
thick, scabrous. 

Peru(?): Eastern part (Cuming 1051}. 

Loasa laxa Macbr., spec. nov. 

Herba ut videtur perennans; caulibus plus minusve tortuosis 
minute strigillosis haud setulosis; petiolis elongatis, ad 8 cm. longis 
setulosis; foliis ovatis vel oblongo-ovatis, basi subtruncatis utrinque 
2-3-lobulatis, margine calloso-denticulatis, superioribus alternis, ad 

15 cm. longis, 10 cm. latis, membranaceis, ubique minute hispidulis; 
floribus ut videtur in cymam dispositis, cymae ramis inflorescentiisque 
ex axillis foliorum superiorum laxe cincinnoseque evolutis, conspicue 
setulosis et minute strigillosis; calycis segmentis fere linearibus 8 mm. 
longis, 1-1.5 mm. latis; petalis 6 subplanis, oblongis, acutiusculis, 

16 mm. longis, 4 mm. latis, longe setulosis; squamis 4 mm. longis, 
manifeste saccato-inflatis, ultra saccum in collum contractis, collo 
supra saccum callis incrassatis ornato, marginibus superne ad apicem 
dilatatis; staminodiis 7 mm. longis; staminibus vix 6 mm. longis. 

Apparently most closely allied toL. carunculata Urb. & Gilg, 243. 
The collector's data for the collection in Field Museum is wanting. 

Peru: Weberbauer 6993, type. 

Loasa leiolepis Urb. & Gilg, Monogr. Loas. 230. 1900. 

Puberulent-stemmed, short-setose, alternate-leaved annual; peti- 
oles to 1 cm. long; leaves ovate-oblong, rounded or acutish at base, 
shortly decurrent on the petiole, with 2-5 serrate lobules or large 
teeth, sparsely asperous or nearly glabrate above, densely pilose- 
puberulent beneath and esetose, unless a few setae on the nerves; 
calyx tube 4-5 mm. long, to 3 mm. broad, the narrow lobes to 6 mm. 
long, sometimes minutely denticulate, in fruit becoming 8 mm. long; 
petals 7-8 mm. long, scarcely clawed; scales dorsally smooth but 
with a thick horizontal bend above, the tip itself callous- thickened, 
the marginal teeth about 1 mm. long. Puberulence rusty and 
plant with aspect of L. fulva. Type cultivated 1843 at Berlin, said 
probably to be of Peruvian origin. F.M. Neg. 10200. 



FLORA OF PERU 157 

Puno: Sandia, Weberbauer 562. Amazonas: Chachapoyas, 
Raimondi. 

Loasa lenta Macbr., spec. nov. 

Planta verosimiliter perennans; caulibus adscendentibus vel sub- 
scandentibus breve setosis; petiolis 2 cm. longis, dense setulosis et 
hispidulis; foliis oppositis, ovatis, serratis, basi subtruncatis, acumin- 
atis, plerumque 3 cm. longis, 2 cm. latis, supra sparse setulosis, sub- 
tus plus minusve hispidulis et setulosis; floribus in apice caulis 
ramorumve in monochasia 2-4-flora dispositis; pedicellis 5-10 mm. 
longis; calycis lobis oblongo-lanceolatis, acutis, leviter setulosis 10 
mm. longis, 2.5 mm. latis, tubus 5 mm. crassus dense setosis; petalis 
17 mm. longis, fere 10 mm. latis, planis, glabris, vix acutis; squamis 
7 mm. longis, calcaribus squamae duplo brevioribus, filis nullis, collo 
manifeste 3-lobo, marginibus subito aliformi-dilatatis; staminodiis 
(pilosis) staminibusque 10 mm. longis. 

Seems from the herbarium material to be a clambering plant with 
tough stems; flowers scarlet. Series Alatae. 

Huanuco: Near Muna, Weberbauer 6722, type. 

Loasa limata Macbr., spec. nov. 

Ut videtur herba annua ad 0.8 m. alta; caulibus petiolisque nitidis 
parce setosis; petiolis (supremis) oppositis circa 5 cm. longis; foliis 
fere rotundato-ovatis circa 1.5 dm. longis, basi inaequaliter cordatis, 
irregulariter serrato-lobulatis, lobulis acutis, denticulis calloso- vel 
glanduloso-apiculatis, ubique parcissime setulosis, subtus minutissime 
pilosis, praesertim supra intense viridibus, membranaceis; floribus in 
axillis foliorum superiorum 1-2 dispositis; calycis segmentis 10 mm. 
longis, basi 3.5 mm. latis, acuminatis, parce setosis, tubus densissime 
brunneo-setosus, ovoideus, 5 mm. crassus; petalis planis, membrana- 
ceis, solum ad nervum medium parcissime setulosis, apice rotundatis, 
3 cm. longis, fere 2 cm. latis; squamis 15 mm. longis, calcaribus circa 
squama 3-plo brevioribus, filis nullis, alis ventralibus 2-2.5 mm. latis 
supra collum parte circa 4 mm. triangulari-productis, collo fere 
truncatis; staminibus quam petala manifeste brevioribus; staminodiis 
glabris circa 15 mm. longis. 

"Flowers salmon-orange, the glands dark red." Series Alatae; near 
the Ecuadorian L. aurantiaca Urb. & Gilg, 209, with smaller leaves, 
setosely ciliate, twice as long calyx lobes, papillose staminodia; sug- 
gests also L. acuminata Wedd., 198, of Ecuador, with much longer 
calyx lobes, pilose staminodia. 



158 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Apurimac: Ampuy, in black loam, 3,200 meters, Stork, Horton 
& Vargas 10624, type. 

Loasa loxensis HBK. Nov. Gen. & Sp. 6: 116. 1823; 216. 

More or less suffrutescent perennial, the leaves except the lowest 
alternate; petioles 2.5-3 cm. long, the ovate leaf blade somewhat 
decurrent or truncate, acuminate, with 2-3 broadly triangular lobes 
above the middle, the margins minutely glandular-dentate, scabrous 
above, puberulent-tomentose beneath and setose on the nerves both 
sides or esetose beneath; pedicels 1-1.5 cm. long; calyx tube 4-5 mm. 
broad, densely hirsute, the lobes 8-12 mm. long; petals flat or nearly 
so, about 3 cm. long, 7-8 mm. broad, shortly reddish hirsute dorsally; 
scales 11 mm. long with 3 very short threads below the neck or these 
obsolete, the scale wing dilated, the wings 5 mm. long; filaments as 
long as the petals; capsule to 17 mm. long, 10 mm. broad at top. 
The Peruvian specimen is smoother than the typical form, the leaves 
dark green above with a few asperous trichomes and short setae on 
nerves, only pilose beneath, the margins repand and subentire to 
shallowly lobed. Related species found as near as Ecuador include 
L. campaniflora Tr. & Planch., 218, similar, but the lanceolate calyx 
lobes 15 to 23 mm. long, and scale wings shorter; and L. aurantiaca 
Urb. & Gilg, 209, L. calycina Benth., 210, both distinguished by the 
stamens being evidently shorter than the petals, the calyx lobes 
about 17 mm. long, and from each other by the size of the flowers, 
the petals of the former being at least 3 cm. long, those of the latter 
scarcely 2. F.M. Neg. 10202. 

Amazonas: Chachapoyas, Mathews. Ecuador. 

Loasa macrantha Urb. & Gilg, Bot. Jahrb. 45: 468. 1911. 

Perennial herb, the stems and branches densely setose; upper 
leaves with petioles to 1 cm. long, the blades broadly ovate to sub- 
orbicular, to 9 cm. long, 8 cm. wide, unequally 5-lobed, the middle 
lateral lobes larger, all acute, unequally serrate-lobed, above sparsely, 
underneath densely pilose and both sides especially on nerves some- 
what setulose; pedicels 5-6 cm. long, setose, the 3-4 flowers igneous; 
calyx 7-8 mm. long, as broad, setose as the lanceolate very acute 
lobes, these also pilose, 2 cm. long, 7 mm. wide; petals somewhat 
concave, about 5 cm. long, 1.5 cm. wide, acute; scales about 15 mm. 
long. Allied to L. grandiflora Desr., 202. F.M. Neg. 10203. 

Junin: Grass steppe with scattered shrubs, 3,400 meters, moun- 
tains east of Palca, Weberbauer 2494, type. 



FLORA OF PERU 159 

Loasa macrophylla Urb. & Gilg, Bot. Jahrb. 45: 467. 1911. 

Erect perennial to 1 meter high with ovate-rotund leaves to 2.5 
dm. long and broad and large 5-merous salmon-rose colored flowers 
in terminal and upper-axillary cymes; petioles 5-6 cm. long; leaves 
unequally and deeply 5-lobed, the larger middle one unequally lobed- 
serrate, the others unequally serrate, all acute, both sides densely 
short pilose, marginally setulose; pedicels 5 cm. long; calyx densely 
long-setose; petals nearly plane, rounded base and apex, about 5 cm. 
long, 22-25 mm. wide; scales nearly 2.5 cm. long. Allied to L. 
argemonoides Juss., 195, Colombian, with different leaves and flowers 
(Urban & Gilg). F.M. Neg. 10204. 

Ancash: Along trail, above Ocros, 2,900 meters, Weberbauer 
2718, type. 

Loasa macrorrhiza Urb. & Gilg, Bot. Jahrb. 45: 467. 1911. 

Perennial apparently from a much-divided horizontal rhizome, 
the densely setose flowering stems to 4 dm. high; pubescent and 
sparsely setose petioles of the densely crowded basal leaves 6-8 cm. 
long, the leaf blades suborbicular, 4 cm. long to 6 cm. wide, broadly 
cordate at base, unequally and more or less deeply lobulate-serrate 
but not lobed; stem leaves much smaller, shortly petioled, similar in 
form, all of both sides short-pilose, with no setae or very few; flowers 
2-3, igneous color, the densely setose pedicels 5-8 cm. long; calyx 
7-8 mm. long and as broad at top, setose, as slightly also the very 
acute lanceolate lobes, these 15 mm. long, 5-6 mm. wide; petals obo- 
vate, rounded at tip, dorsally setose, 25-27 mm. long, 15 mm. wide; 
scales about 16 mm. long. Allied to L. ranunculifolia Humb. & 
Bonpl., 201, with lobed leaves. F.M. Neg. 10206. 

Ancash: In moist rocky ravines, 4,100 meters, above Huaraz, 
Weberbauer 3080, type. 

Loasa macrothyrsa Urb. & Gilg, Bot. Jahrb. 45: 469. 1911. 

Suffrutescent, strict, to 2 meters high, the terete lignescent stems 
densely long-setose; leaves alternate, the lower subsessile, all oblong, 
subcordate, acute, to 14 cm. long, half as wide, base to apex unequally 
and deeply dentate-lobed, the 5-8 lobes unequally serrate, gray- 
tomentose beneath, above laxly, beneath densely, on nerves with 
setae intermixed; flowers greenish-yellow, subnutant, in false panicles, 
pedicels to 1.5 cm. long; calyx tube 1 cm. long, 6 mm. wide, densely 
setose, the ovate-oblong lobes as long, 5-6 mm. wide; petals strongly 
cymbiform, 2-2.5 cm. long, 8-9 mm. wide; scales 8-9 mm. long, yel- 



160 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lowish with 2 horizontal reddish-brown lines. Related to L. carun- 
culata Urb. & Gilg. F.M. Neg. 10205. 

Cajamarca: In open or closed formation at 2,400 meters, below 
San Miguel, Weberbauer 3907, type. 

Loasa magnifica Urb. & Gilg, Monogr. Loas. 204. 1900. 

Apparently in general like L. grandiflora but all or all but the 
lowest leaves alternate and the upper more broadly cordate; calyx 
lobes 17-23 mm. long, 7-12 mm. wide above the base; petals plane 
or nearly, 30-43 mm. long, 12-22 mm. wide, with a short claw 3-4 
mm. wide; scales 18-22 mm. long, 4-6 mm. wide, the neck with 3 
lobes 2-3 mm. long. Petals orange-pink, filaments and scales 
(except pink sacs) white (K. & S.). In the type the leaves are very 
setose, sharply lobed and toothed; the following collections may 
represent a distinct species but only uppermost leaves are present, 
these clasping-sessile, repandly lobed or sub-entire. Illustrated, 
Urb. & Gilg (flower parts), pi. 5. F.M. Neg. 10207. 

Peru: Without locality, MacLean, type. Lima: Rio Blanco, 
Kittip & Smith 21625. Matucana, 369 (det. Killip). Above Obra- 
jillo, Pennell 24392? (det. Killip). 

Loasa nitida Desr. in Lam. Encycl. 3: 581. 1791; 132. 

Pale-puberulent or glabrate, rather succulent, thin-leaved annual, 
essentially esetose except for a few pale bristles on the campanulate 
calyx; lower leaves decurrent on the petiole, the upper subamplexi- 
caul, all more or less shallowly and repandly pinnate, the lobes with 
a few teeth or crenate, ovate, nearly glabrous above, rather ashy 
puberulent beneath, sometimes a dm. or more long; calyx tube 4-5 
mm. long, the lobes 5-7 mm. long, 2-2.5 mm. wide; petals yellow, 
7-8 mm. long, a little cucullate at tip, plane but not clawed at base; 
scales about 4 mm. long, with 3 lamelliform dorsal appendages; sta- 
mens to about 50; capsule thicker than long, scabrous, the valves 
3. Petals yellow, the nectaries blood-red on white back (Weber- 
bauer). L. triloba Dombey with mostly trilobed leaves and capsules 
longer than thick, sometimes accredited to Peru, is in all probability 
Chilean, this surely the origin of the Dombey specimen. F.M. Negs. 
24173; 38481. Here, because of its opposite leaves, would be sought 
the Bolivian L. Herzogii Urb. & Gilg, Meded. Rijks Herb. Leiden 
29: 54. 1916 related to L. chenopodifolia but leaves opposite, on 
petioles 3-7 cm. long; petals 11-12 mm. long. Illustrated, Juss., 
Ann. Mus. Hist. Nat. Paris 5: pi. 2; Tratt. Arch. Gewachsk. 1: 
pi. 23. 



FLORA OF PERU 161 

Lima: Chancay, in sand and stone, Ruiz & Pavon. Lurin, 5938 
(det. Killip). Near Lima, Dombey, type; Abadia. Amancaes Hill, 
Isern; Weberbauer 1620; 145. Arequipa: Raimondi. 

Loasa olmosiana Gilg, spec. nov. in herb. 

Herba robusta perennans; caulibus petiolisque (petiolis 3-6 cm. 
longis) mediocriter setosis et hispidulosis; foliis alternis ut videtur 
late vel elliptico-lanceolatis basi apiceque sensim angustatis superiori- 
bus circa 2 dm. longis, 1 dm. latis, plus minusve obscure lobulatis, 
toto margine minute calloso-denticulatis, membranaceis, supra 
minute setulosis, subtus puberulis vel ad nervos sparse setulosis; 
calycis floribus laxe dispositis, pseudopaniculatis; lobis 17 mm. longis, 
8-9 mm. latis, acuminatis, strigillosis, tubus vix 4 mm. crassus per- 
dense setosis; petalis subplanis sparse setulosis et pilosis membran- 
aceis acutis 22 mm. longis, 8 mm. latis; squamis 10-11 mm. longis, 
saccis paullo inflatis, filis nullis, collo integro marginibus superne 
sensim dilatatis supra collum 2.5 mm. productis atque 5.5 mm. ali- 
formi-dilatatis; staminodiis 16 mm. longis; staminibus vix 15 mm. 
longis. 

From the scales evidently a species of the Alatae but apparently 
undescribed. 

Lambayeque: Olmos, Weberbauer 7097, type. 

Loasa peltata Spruce ex Urban & Gilg, I.e. 205. 

Lowest leaves densely crowded, the lower always peltate, sub- 
orbicular, the petiole joined at the lower third of the palmately 5-7- 
lobed blade, this about 5 cm. wide, the lobes ovate, obsoletely 
dentate; decreasing upper leaves not at all peltate, alternate as 
the lower, oblong-lanceolate, denticulate, finally sessile, all sparsely 
asperous with few short setae on nerves; flowers 5-merous, yellow, 
the pedicels to 13 mm. long; calyx 4-5 mm. long, 3-4 mm. thick, 
the lobes 12-13 mm. long, 3-4 mm. wide; petals slightly concave 
above, to about 22 mm. long and half as wide, densely pilose with- 
out but esetose, broadly clawed or narrowed at base; scales rec- 
tangular from the bicalcarate base, deeply bilobed by the extended 
"wings" (cf. pi. 5, Urban & Gilg). Species unique in the peltate 
leaves, but related is L. rugosa. 

Peru(?)r Mount Mulmul (Spruce, type). 

Loasa picta Hook. f. Bot. Mag. 75: pi. W28. 1849; 222. 

More or less puberulent-pilose as well as rusty-setulose annual, 
1-several dm. high, often branching from the base; leaves rather few, 



162 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

alternate, the petioles to 1.5 cm. long, the broadly oval to ovate 
blades to 6 cm. long and nearly as wide, cuneate at base, acute, 
coarsely serrate, the teeth often of two sizes, sparsely setulose as 
well as asperous above, puberulent beneath, both sides; calyx 6-7 
mm. long, 3-4 mm. wide, the lobes about 4 mm. long, 2.5-3 mm. 
wide; petals 11-12 mm. long, narrowed to base, not clawed; scales 
from side saccate-convex, from back ovate, above the base with 2 
dorsal strongly inflated calluses and with fold or callus above these 
and irregularly dotted or blotched, the tip thickened, the 3 nerves 
extended about 1 mm.; staminodia and the 50 or so filaments about 
7 mm. long, the former pilose, strongly thickened at base; capsule 
more than 2 cm. long, 3-4 mm. thick. The var. Stuebeliana Urb. 
& Gilg, 223, has oblong-lanceolate leaves to 10 cm. long, 4.5 cm. wide, 
long-cuneate at base, the longer petioles 5 cm. long, the flowers more 
than 3 cm. wide. Probably should include L. Poissoniana and L. 
ferruginea, which see. Type material was cultivated at Kew, and, 
according to Herrera also in Peru for its flowers, their scales yellow 
with transverse red stripes. 

Amazonas: Chachapoyas, Mathews. Between Pacasmayo and 
Moyobamba at Tambo Viejo, 3,100 meters, Stuebel (the var.). 
Above Samanco (Weberbauer, 170) (at least this collection is typical). 
Cajamarca: San Miguel, Weberbauer 3887(1}. Ancash: Region 
Huaraz, Weberbauer 3146(1). Cuzco: Santa Rita, Valle del Urubamba, 
2,300 meters, Herrera 1550. Huaillai, Marcapata, Vargas 9709(1). 
"Angel-tuana," "chino-quisa." 

Loasa Poissoniana Urb. & Gilg, I.e. 226. 

Like L. ferruginea and probably only a variety but the leaves 
ovate-oblong, the lower to 4 cm. long, 3-3.5 cm. wide, rounded or 
subcordate at base, the lower more or less deeply 3-5-sinuate-den- 
tate or serrate-dentate, the teeth unequal; marginal teeth of scale 
rounded at tip, 1-1.2 mm. long. Type probably, if Peruvian, from 
Cuzco region. Some material cited under L. picta may belong here 
with L. ferruginea. F.M. Neg. 38482. 

Peru: Gay 1975, type. Lima: Matucana, 179. Ambo, 3154. 
Moquehua: Weberbauer 7394- 

Loasa ranunculifolia Humb. & Bonpl. PI. Aequin. 1: 50. pi. 14- 
1806; 201. 

Densely setose perennial with the general characters of L. grandi- 
flora but well-marked by its closely appressed hirsute and scattered 
setose 7-9-lobed lower leaves, the lobes extending less than half 



FLORA OF PERU 163 

way and by its comparatively small flowers, the oval petals only 25 
mm. long, 13-15 mm. wide; scales 12-13 mm. long, the neck only 
twice as long as the calluses and the wings more extended (cf . illus- 
tration, Urb. & Gilg, I.e. pi. 5). The Ecuadorian L. alpina Urb. & 
Gilg, 197, has 5-7-lobed leaves and the ventral wings produced above 
the scale-neck are as long as the scale-body. My 1934 from Mito, 
Huanuco, referred by Killip to L. ranunculifolia, may or may not be 
this species as it is in fruit. Ecuadorian in distribution, a collection 
given by Herrera from southern Peru is referred here to L. cuzcoensis, 
which see. F.M. Neg. 38483. 

Cajamarca: 3,000 meters, Bonpland, type. Ecuador. 

Loasa rugosa Killip, Journ. Wash. Acad. Sci. 18: 91. 1928. 

Perennial to 3.5 dm. high with a stout minutely pilose and densely 
red-setose stem nearly 1 cm. thick; basal leaves numerous, the cauline 
few; petioles 2-6 cm. long; blades reniform 2.5-3.5 cm. long, 4.7 cm. 
wide, shallowly and irregularly 5-9-lobed, deeply cordate, coriaceous, 
strongly rugose, nearly glabrous above, rufo-tomentose on nerves 
and veins beneath; peduncles stout, to 4 cm. long; calyx densely 
setose, the lobes to 2 cm. long, 1 cm. wide; petals 4 cm. long, 2.5 cm. 
wide, orange; scales petaloid, rectangular, 12 mm. long, erectly 
bilobed to below middle with 2 roundish appendages without near 
base; staminodia setaceous, 15 mm. long, densely pilose; stamens to 
2.5 cm. long, the linear-oblong anthers to 2 mm. long; capsule 2-2.5 
cm. long, 2 cm. wide, densely brown-setose. Series Grandiflorae; 
differs from L. pellata Spruce in non-peltate thicker rugose leaves 
and much larger flowers. The pubescence was extremely stinging. 

Junin: Tambo de Vaca, rocky uplands, 4,000 meters, 4350, type. 

c 

Loasa solaria Macbr., spec. nov. 

Herba annua; caulibus pedicellisque plus minusve setulosis; 
petiolis usque 10 mm. longis; foliis superioribus sessilibus oppositis 
oblongo-ovatis, acuminatis, fere aequaliter denticulatis, denticulis 
conspicue setosis, supra adpresse hispidulosis, subtus glabratis, 5-8 
cm. longis, circa 3.5 cm. latis; pedicellis circa 1 cm. lorgis; petalis 
12 mm. longis manifeste unguiculatis ad nervum medium setulosis, 
squamis dorso calcaribus vel alis nullis in parte superiore plica inte- 
gro horizontali notatis, apice ipso calloso-incrassatis marginibus 
superne in dentes supra collum productis. 

Referred at one time to L. chenopodifolia from which its opposite 
regularly toothed and setose leaves distinguish it. In its clawed 



164 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

petals it approaches L. Stuebeliana. The collectors noted the flowers 
as bright yellow with scarlet centers. 

Lima: Viso, wet shrubby slope, 3,000 meters, 577, type. 

Loasa solata Macbr., spec. nov. 

Manifeste perennans; caulibus petiolisque breviter setulosis et 
plus minusve puberulis; petiolis oppositis 10-15 mm. longis; foliis 
fere oblongo-lanceolatis circa 7 cm. longis, 3 cm. latis obscure 
ubique 2-3-lobatis et crenulato-denticulatis, acuminatis supra minute 
strigillosis et sparse setulosis, subtus puberulis haud setulosis; 
floribus in apice caulis circa 7 in cincinnum dispositis; pedicellis 
ad 3.5 cm. longis, calycis lobis paullo vel haud setulosis, puberulis, 
11 mm. longis, basi 4 mm. latis, acuminatis, tubus densissime bruneo- 
setosis; petalis 27 mm. longis vix 5 mm. latis, acutis, subplanis, apice 
paullo cucullatis, puberulis et sparse setosis; squamis papilloso- 
scabridis, 15 mm. longis, filis nullis, saccis 3 mm. longis, collo anguste 
oblongo 6 mm. longo, alis ventralibus supra collum 6 mm. productis. 

With nearly the foliage of L. calycina Benth. and of L. campani- 
flora Tr. & PL, 218, but the scales suggestive of (though different 
from) those of L. acuminate, Wedd., 198, Ecuadorian and Colombian 
species. The data for the type sheet is lacking in Field Museum. 
Urban and Gilg gave the plant a new name but failed to publish it, 
perhaps deciding that it was referable to a species already described. 
Because of this uncertainty I have not taken up their name in this 
as in several other instances. 

Peru: Weberbauer 6408, type, Field Museum. 

Loasa Stuebeliana Urb. & Gilg, Monogr. Loas. 228. 1900. 

Similar and closely allied to L. leiolepis but the lower petioles 
to 8 cm. long, the leaf blades well-decurrent and subentire or merely 
repand, esetose, or essentially, and at base with 2 reniform sub- 
amplexicaul stipules; petals 12-13 mm. long, broadly clawed; other- 
wise not marked. F.M. Neg. 10219. 

Amazonas: Chachapoyas, Mathews. Between Pacasmayo and 
Moyobamba at 2,000 meters, Stuebel 52d, type. 

Loasa tingomariensis Macbr., spec. nov. 

Herba an perennans(?) 0.5 m. alta; caulibus petiolisque sparse 
setulosis et minute strigillosis; petiolis alternis vel per paria ap- 
proximatis ad 8 cm. longis; foliis viridibus suborbiculatis a basi (paullo 
cordatis) usque ad apicem leviter et inaequaliter lobulatis, lobulis 
utrinque circa 5 ovato-triangularibus, margine toto dense minuteque 



FLORA OF PERU 165 

denticulatis, supra mediocriter sed laxe setulosis subtus imprimis 
venis cum pilis brevibus setuloso-villosulis, membranaceis, viridibus, 
superioribus 15 cm. longis, 13 cm. latis; floribus in apice caulis vel 
in axillis foliorum superiorum circa 6 cincinnose dispositis; calycis 
lobis oblongo-ovatis, acutis fere 1 cm. longis, 6-7 mm. inferne latis, 
tubus circa 4 mm. longus superne circa 6 mm. latis; petalis planis 
vel subplanis oblongis acutis 20-22 mm. longis, 12 mm. latis, dorso 
baud setulosis leviter pilosis, intus glabris, membranaceis; squamis 
8 mm. longis a dorso oblongis, collo obscure 3-crenulato-lobatis lobis 
recurvatis, lateralibus obscure papillosa emittentibus, marginibus 
superne sensim dilatatis supra collum valde aliformi-dilatatis 5-6 
mm. latis; staminodiis 2 interioribus squamam superantibus; 
staminibus quam petala evidente brevioribus. 

With the floral structure of the Alatae, the scales similar to those 
of L. calycina, 210, Ecuadorian, its leaves oblong-lanceolate, the 
leaves nearly those of L. loxensis, but broader, larger, the petals 
esetulose and the scales differently proportioned. L. aurantiaca 
Urb. & Gilg, 209, of Ecuador has narrower oblong-lanceolate petals, 
much longer scales. "Flowers orange-red to tomato red." 

Huanuco: Sandy bank 10 km. south of Tingo Maria, 700 meters, 
Stork & Horton 9512, type. Junin: Yapas, Killip & Smith 25469. 

Loasa triphylla Juss. Ann. Mus. Hist. Nat. Paris 5: 27. pi. 5. 
1804; 235. 

Variable annual but well-marked by virtue of its trifoliate lower 
leaves, the ovate leaflets always petiolulate and often with small 
leaflets or lobes between them; upper leaves lobed, the lobes ovate; 
petals white, 11-22 mm. long; scales obsoletely, bluntly and shortly 
bihorned at the base, banded purple and white, thickened above, 
verruculose beneath the emarginate apex; capsule to 2 cm. long. 
Several names have been given to variations apparently of no taxo- 
nomic value. Often more or less setose and 1 meter high. L. Hum- 
boldtiana Urb. & Gilg, 240, Ecuadorian, has 3 enlarged nerves from 
the bicalcarate base instead of the 2 blunt horns that characterize 
L. triphylla. Type by Jos. Jussieu, "Peru." 

Piura: Ayavaca, Weberbauer 6424- Lambayeque: Weberbauer 
7096. Lima: Matucana, Weberbauer 56. Widely distributed in 
Warm America. 

Loasa urens Jacq. Obs. 2: 15. pi. 38. 1767; 232. L. ambrosiae- 
folia Juss. Ann. Mus. Hist. Nat. Paris 5: 26. pi. 4- 1804. 



166 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Annual, the stems abundantly yellow-brown setose, and with 
oblong-pinnate or bipinnate leaves, well-petioled, the larger to 1 or 
2 dm. long, pilose-puberulent especially beneath as well as setose 
on the nerves to glabrate; petals yellow, about 2 cm. long, narrowly 
clawed; calyx tube 4-8 mm. long, 4-6 mm. across, the lobes 7-10 
mm. long, 3-4 mm. wide; scales 6-7 mm. long, the two calluses 
somewhat spreading and with 2-5 green or roseate folds above them ; 
capsule 16-20 mm. long, 6-9 mm. thick. Leaf-pinnae more or less 
lobed, narrowly oblong; setae lustrous, many. Illustrated (flowers), 
Urban & Gilg, I.e. pi. 5. 

Lima: Amancaes, Weberbauer 1623; 142, 143, 146; Soukup 1306; 
Mathews 729. Near Lima, Raimondi; Caslelnau; Abadia. Chorillos, 
5875. Arequipa: Pasco, Mejia and Cochendo (fide Bruns). Are- 
quipa, Raimondi. Mollendo, Weberbauer 1471; Johnston 3554- 
"Inea." 

Loasa Vargasii Macbr., spec. nov. 

Herba annua 0.7 m. alta; caulibus petiolisque mediocriter strigil- 
losis et setulosis; petiolis alternis circa 5 cm. longis; foliis late oblongo- 
ovatis basi circa 14 cm. longis, 12 cm. latis (superioribus paullo 
reductis?) fere truncatis subpinnatim lobatis, lobis acutis utrinque 
2-3 inferioribus majoribus fere oblongis 3-4 cm. longis, 2.5-3 cm. 
latis, grosse vel repando-serratis, membranaceis, ubique scabro- 
hispidulis haud setulosis; floribus cernuis in apice caulis in pseudo- 
racemos usque ad 7 cm. longis dispositis; calycis lobis late ovatis 
acutis 9 mm. longis, 5 mm. latis; petalis 2.5 cm. longis, 7 mm. pro- 
fundis margine setulosis in parte % m ^- subito in unguiculum angus- 
tatis; squamis 8 mm. longis distincte supra basin in calcaria bina 
valde inflata protractis, filis nullis, supra incrassato notatis, apice 
ipso 4-crenato-calloso-incrassatis marginibus ipsis superne in dentes 
1 mm. longos supra collum productis; staminodiis 2 squamam multo 
superantibus. 

Flowers white, the glands patterned in red. Clearly allied to 
L. Schlimiana Planch. & Lind., 227, of Colombia but differs in 
foliage and scales. 

Apurimac: Rio Pachachaca, in gravelly rainy-green shrubland, 
2,000 meters, Stork, Horton & Vargas 10531, type (University of 
California; scrap and photo, Field Museum). 

Loasa Weberbaueri Urb. & Gilg, Bot. Jahrb. 45: 467. 1911. 

Perennial herb to 2 meters high, the sublignescent stems densely 

setose; petioles 8-9 cm. long; leaves nearly suborbicular, cordate at 



FLORA OF PERU 167 

the subequal base, to 22 mm. long, nearly as broad, subpinnately 
lobed, the 5 ovate lobes about one-third as deep as the leaf-breadth, 
acute, unequally dentate, densely asperous above, ashy or brown- 
tomentulose beneath; flowers borne in twos, the very densely setose 
pedicels to 3.5 cm. long; calyx tube about 1 cm. long and broad, 
densely setose as the acute lobes dorsally, these to 2.5 cm. long, 
1 cm. wide at base; petals plane, 4 cm. long or longer, 2.5 cm. wide, 
rounded at tip; scales about 2 cm. long. Related to L. acuminata 
Wedd., 198, Ecuadorian, leaf lobes often 7, flowers smaller. F.M. 
Neg. 10225. 

Cajamarca: In half shade by deciduous wood near Huambos, 
3,100 meters, Weberbauer 4183, type. 

6. CAJOPHORA Presl 

Round-stemmed vines or annual or perennial herbs mostly abun- 
dantly provided with white stinging bristles, the leaves opposite, 
the flowers white, yellow or red, axillary or in pseudoracemes or 
panicles. Capsules rarely straight, always dehiscing from base up- 
wards, remaining closed at the top. Otherwise like Loasa. The 
similar genus Blumenbachia Schrad. may be found in southern Peru; 
it has a sharply 4-angled stem, the capsule wall thick instead of thin 
and different (unparted) placentae. A number of too closely related 
Bolivian forms, doubtfully distinct, have been included here, for 
they are to be expected in the same mountain range running into 
southern Peru. But in the known Peruvian species the seeds are 
merely foveolate-reticulate, never bi winged or irregularly angled. 
It seems necessary to follow the monographers and separate the 
species on habit but several forms, as remarked in the descriptions, 
are not clearly defined as to this as well as other characters. Some 
common names are "Ortiga macho," "ortiga colorada." 

Stems never scandent, sometimes decumbent or weak. 
Flowers 5-merous. 
Stems weak, or if erect, the leaves rosulate below. 

Leaves rosulate, on the short suberect stems crowded. 

C. rosulata. 

Leaves not rosulate, on the elongate, weak, subdecumbent 
stems remote. 

Scales with 3 threads; capsule (so far as known) straight. 

Plants abundantly setose C. coronata. 

Plants sparsely setose C. andina. 



168 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Scales without threads; capsules often somewhat contorted. 

Scales about 5 mm. long, entire or subentire (cf. also C. 
carduifolia). 

Leaves narrowly lanceolate; front edge of scales erect, 
thin, entire C. cirsiifolia. 

Leaves ovate-oblong; front edge of scales slightly 
incurved and toothed C. cymbifera. 

Scales about 10 mm. long, bidentate at tip. .C. pauciseta. 

Stems erect or suberect, the leaves never rosulate and the stems 
short. 

Scales provided with 3 threads C. andina. 

Scales without threads. 

Stems 2.5 mm. thick; scales apically thickened; petals 14 
mm. long C. pachylepis. 

Stems stout; scales apically thin; petals 17-25 mm. long. 

Petals 25 mm. long or longer C. carduifolia. 

Petals 17-18 mm. long C. cymbifera. 

Flowers 6-7-merous. 
Flowers sessile or subsessile (always?). 

Scales with a small appendage at the front C. superba. 

Scales obsoletely 3-lobed at the front C. heptamera. 

Flowers well-peduncled. 
Scales neither dorsally keeled nor apically thickened. 

C. chuquitensis. 

Scales keeled and apically thicker. 

Capsules subglobose; leaves with 2-3 pairs pinnae. 

C. sphaerocarpa. 

Capsules oblong-ovoid; pinnae often several. 
Leaves slightly ovate; pedicels to 1 cm. long. .C. horrida. 
Leaves oblong-lanceolate; pedicels 3-7 cm. long. 

C. Mandoniana. 

Stems truly scandent (cf. C. pachylepis). 
Petals 6. 

Leaf lobes entire to dentate; flowers orange-red. .C. madrequisa. 

Leaf lobes pinnate-serrate; flowers scarlet C. scarlatina. 

Petals 5. 



FLORA OF PERU 169 

Petals flat, ecalloused. 

Petals 3-5 cm. long; scales with 3 threads C. canarinoides. 

Petals 2-2.5 cm. long; scales without threads. . . .C. Buraeavi. 
Petals evidently cymbiform. 

Scales not calloused dorsally, sometimes apically. 
Scales dorsally provided with 3 threads or 3 appendages. 
Scales thin at front, not thickened. 
Leaves ovate-oblong; petals about 1.5 cm. long. 

C. contorta. 
Leaves lanceolate; petals about 2 cm. long. 

C. Pentlandii. 

Scales thickened apically or squamose-appendaged. 
Scales with 3 threads. 

Capsules narrowly oblong, about 5 times longer than 
broad; leaves lanceolate C. tennis. 

Capsules oblong, 2-4 times longer than broad; leaves 
ovate C. aequatoreana. 

Scales with 3 thickish appendages. 

Petals entire C. Smithii. 

Petals serrulate C. serropetala. 

Scales without threads. 

Interior staminodia without appendage C. cinerea. 

Interior staminodia appendaged. 
Seeds not winged; petals 2-3 cm. long. 

Leaf lobes unequally serrate; petals to 2.5 cm. long. 

C. sepiaria. 

Leaf lobes equally dentate; petals about 3 cm. long. 

C. macrantha. 
Seeds winged; petals about 15 mm. long. 

C. pterosperma. 

Scales bicalloused below the neck at insertion of 3 threads. 

C. stenocarpa. 

Cajophora aequatoreana Urb. & Gilg, Monogr. Loas. 313. 
1900. 

Climbing, the terete stems to 2.5 mm. thick; petioles to 27 mm. 
long; leaves ovate, pinnately lobed about half the breadth or nearly 
pinnate, the ovate divisions or lobes repand-dentate or serrate, 



170 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lightly asperous above, pilose beneath, slightly setose both sides, 
5-8 cm. long, about 5 cm. wide; calyx tube 7-8 mm. long, 5-6 mm. 
across, the lobes about 1 cm. long, regularly serrate; petals about 16 
mm. long, 8-9 mm. wide, deeply cymbiform-concave; scales 4-5 mm. 
long, ecarinate but with 3 longer threads and evidently thickened 
marginally; capsules 2.5 cm. long or a little longer, 2-3 times longer 
than broad. Illustrated, Bot. Mag. pi. 6134 as B. contorta. C. 
Kuntzei Urb. & Gilg, 314, Bolivian, has leaves half as wide, capsules 
only 2 cm. long. F.M. Neg. 24164. 

Huanuco: Pacasmayo to Moyobamba near Cumallca, 3,600 
meters, Stuebel 35a. Ancash: Raimondi. Huancabamba, Raimondi. 
Ecuador. 

Cajophora andina Urb. & Gilg, Monogr. Loas. 287. 1900. 

Stems erect(?), or ascending from a fusiform root; flowering 
stems several to many, a dm. long or longer, densely puberulent- 
pilose but esetose; leaves opposite, oblong, to 9 cm. long, nearly 
3 cm. wide, pinnate, the pinnae about 6 pairs, oblong, acute, indis- 
tinctly dentate, densely tomentulose beneath, not at all setose but 
with a few setae above; flowers terminal, solitary; calyx tube 6 mm. 
long, the linear subentire lobes 8-9 mm. long, setae lacking; petals 
to 2 cm. long, not more, 8 mm. wide, deeply cymbiform; scales 
about 4 mm. long, dorsally 3-keeled and with 3 threads; stamens 
about 100; capsule unknown. To be expected on extension of the 
Bolivian ranges. Described as "erect, rigid" but material seen and 
referred here by Urban & Gilg has weak ascending stems. F.M. 
Neg. 24165. 

Peru (probably). Bolivia. 

Cajophora Buraeavi Urb. & Gilg, Mem. Torrey Club 3, pt. 3: 
37. 1893; 328. 

Slender vine, the stems about 2 mm. thick; petioles 1-3 cm. long; 
leaves ovate, acuminate, deeply serrate or subpinnate, the lobes 
sharply serrate, minutely asperulous above, pilose-tomentulose 
beneath and there with a few scattered setae, if any, to 11 cm. long, 
and about half as wide; calyx tube 8 mm. long, the regularly dentate 
lanceolate lobes little longer; petals 22 mm. long, 13 mm. wide, 
acute, little concave; scales 6 mm. long, the dorsal nerves not 
prominent, threads lacking, the lobes rounded and very densely 
papillose. 

Junin: Cerro de Pasco (Mathews). Huanuco: 3,600 meters, 
Cumallca, Stuebel 35a. Puno: Carabayo, Weddell 4735. Bolivia. 



FLORA OF PERU 171 

Cajophora canarinoides (Lenne* & C. Koch) Urb. & Gilg, in 
Pflanzenfam. 3, 6a: 119. 1894; 326. Illairea canarinoides Lenne" & 
C. Koch, Verh. Ver. Bef. Gartenbaus, N. R. 1: 397. 1853. 

Closely allied to C. Buraeavi but leaves sagittate from a truncate 
or emarginate base, long-acuminate, sometimes pinnate at base; 
calyx tube 5-9 mm. long, the ovate lobes equally or unequally 
dentate, 8-15 mm. long; petals 3-5 cm. long, 10-18 mm. wide, 
variable in form: scales with 3 threads as long or a little longer- 
Illustrated, Bot. Mag. 83: pi. 5022. 

Puno: Cuyocuyo, Sandia, 3,180 meters, Weberbauer 862; 184. 
Bolivia. 

Cajophora carduifolia Presl, Rel. Haenk. 2: 42. 1831; 290. 

Stems "ascending" or perhaps erect, stiff, 3-4 mm. thick below, 
more or less densely tomentose with some setae intermixed especially 
above, to 2.5 dm. high or higher; petioles to 2.5 cm. long; leaves 
ovate to broadly ovate, to 11 cm. long, 4.5 cm. wide, pinnate or 
nearly, the ovate always acute lobes distinctly but not deeply serrate, 
abundantly setose above, little beneath but densely tomentose; 
flowers solitary, terminal; calyx lobes dentate, 12-13 mm. long; 
petals yellow, 26-28 mm. long, 14-15 mm. wide, deeply cymbiform- 
cucullate; scales 6-7 mm. long, deeply saccate-convex, the back with- 
out keel, no threads, the margin always evidently thinner apically, 
not notched before. The original description calls for a weak- 
stemmed plant and indeed here Ball referred material from Chicla 
that was sprawling without support; cf. C. contorla. 

Huanuco: (Haenke). 

Cajophora chuquitensis (Meyen) Urb. & Gilg, I.e. 301. Loasa 
chuquitensis Meyen, Reise 1: 483. 1834; Walp., Nov. Act. Acad. 
Leop.-Carol. 19: Suppl. 1: 339. 1843. 

Stems "stiffly erect" or ascending, 2.5-3 mm. thick, tomentose 
and laxly setose; petioles about 1 cm. long; leaves lanceolate, 
rarely somewhat ovate, to 10 cm. long, 22 mm. wide, densely long- 
setose above, usually only tomentose beneath or setae very few, 
pinnate, or pinnately divided, the divisions subpinnately dentate; 
flowers 2-3, 7-merous, erect; pedicels to 2.5 cm. long, not lengthening 
in fruit; calyx lobes entire or nearly, linear-lanceolate, about 10 mm. 
long; petals 13-14 mm. long, 7-8 mm. wide, deeply cymbiform- 
cucullate; scales about 5 mm. long, ecarinate, apically thin, the 
dorsal threads much longer; placentae plane, dichotomous; capsule 
slightly longer than 2 cm., 8-9 mm. across. F.M. Neg. 10145. 



172 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Puno: Lake Titicaca, Meyen, type. San Antonio, Lechler. 
Ayacucho: One day from Ayacucho, 3,500 meters, Weberbauer 5486. 

Cajophora cinerea Urb. & Gilg, I.e. 321. 

A vine, the stems to 3 mm. thick, the petioles 8 mm. long, the 
leaves oblong or ovate-oblong, regularly serrate-dentate, stoutly 
setose above, tomentose beneath; calyx tube 6-7 mm. long, 3-4 mm. 
across; the lobes as long, deeply dentate; petals nearly or quite 
15 mm. long, only slightly concave; scales about 4.5 mm. long, 
ecarinate, efilamentose, manifestly thickened above, not winged 
below but edge a little inflexed; inner staminodes not appendaged 
dorsally above the base. Type without locality, but possibly from 
northern Bolivia and therefore to be expected in southern Peru. 
F.M. Neg. 32587. 

Peru (possibly). Bolivia. 

Cajophora cirsiifolia Presl, Rel. Haenk, 2: 42. pi. 56. 1831; 
280. C. Lechleri Urb. & Gilg, Pflanzenfam. 3, 6a: 119. 1894, fide 
Urb. & Gilg. 

Stems to several dm. long, weak and decumbent-ascending, about 
3 mm. thick, the lowest leaves rosulate; petioles to 7.5 cm. long; 
leaves oblong-lanceolate, pinnate, to nearly 2 dm. long, 2.5-3.5 cm. 
wide, the usually broadly ovate pinnae deeply dentate or lobulate, 
tomentulose beneath and setose especially above and marginally, 
the white bristles scattered but prominent as also on the stems 
and petioles; flowers pseudolateral ; calyx tube 5-6 mm. long, 3-4 
mm. across, the narrow lobes 8-9 mm. long, obsoletely and remotely 
dentate; petals about 16 mm. long, 11 mm. wide, saccate-cymbi- 
form; scales 4.5-5 mm. long, efilamentose, entire at front and 
neither thickened nor winged; capsule subglobose to 22 mm. long, 
somewhat spiraled below, or scarcely. Flowers brick-red (Mexia). 
F.M. Neg. 24167 (Lechleri). 

Huanuco(?): (Haenke, type). Puno: Near Acanzaro and San 
Antonio, Lechler 1768 (C. Lechleri). Pucara, 3,700 meters, Weber- 
bauer 401 ; 185. Sandia, 4,400 meters, Weberbauer 962; 218, 219, 220. 
Crevices in crags, Granja Salcedo, 3,835 meters, Mexia 7778 (det. 
Johnst.); also Soukup 9, distr. as C. horrida. Araranca, wide clumps 
on rocky siliceous slopes, 4,100 meters, Pennell 3458 (det. Killip). 
Bolivia. "Llungo-llungo." 

Cajophora contorta Presl, Rel. Haenk. 2: 42. 1831; 306. 
C. Preslii Urb. & Gilg, I.e. 306. Blumenbachia grandiflora R. & P. 
ex G. Don, Gard. Diet. 3: 62. 1834, probably. 



FLORA OF PERU t 173 

Stems only about 2 mm. thick, scandent; petioles to 12 mm. long; 
leaves lanceolate, to 6 cm. long, 4.5 cm. wide with as many as 12 
pairs oblong to ovate pinnae, these usually regularly serrate-dentate, 
little setose, not at all beneath where tomentulose; calyx tube 
7-8 mm. long, 4-5 mm. across, the ovate deeply dentate lobes nearly 
as long; petals 15 mm. long or scarcely longer (to 20 mm.), about 
half as wide, deeply concave; scales 5 mm. long, only 2 mm. wide, 
with 3 nearly filiform and greatly elongate threads, medially keeled, 
the margin not thickened nor winged below nor involute; filaments 
about 60; capsules unknown. Ball, Journ. Linn. Soc. 22: 39. 1885, 
referred his material to C. carduifolia with the remark: the slender 
but rather stiff stems twine, although in this district the plant 
ordinarily grows in positions where it can obtain no support. F.M. 
Neg. 10161. 

Lima: Chicla, 3,720 meters, Weberbauer 56; (Ball?). Puente 
Infernillo (Ward, fide Ball). Huanuco(?): Andean valleys, Haenke, 
type. Cuzco: H err era 523. Urubamba, Herrera 686; 34.12; Vargas 
330. Prov. Calca, Vargas 329. Arequipa: Isern 2019. Bolivia. 
"Huasjia-quisa," "quisa." 

Cajophora coronata H. & A. in Hook. Bot. Misc. 3: 327. 
1833; 278. 

Closely allied to and in habit and pubescence resembling C. cirsii- 
folia but the leaves more ovate, to 7 cm. wide, the calyx tube 9-25 
mm. long, about 7 mm. across, the lobes pinnate, the petals 2-5.5 
cm. long, 17-30 mm. wide and, more particularly, the scales provided 
with 3 filaments and 3-lobed before and the capsule always straight, 
to 4 cm. long, 2 cm. thick. Highly variable in foliage and flowers, 
nevertheless well-marked by the scales and capsules. Localities not 
surely Peruvian. There is a fine habitat photograph in Ostenia 
opposite page 240. 1933. Illustrated, Bot. Mag. pi. 8125. 

Moquehua(?): Tacora-Sajama, Stuebel 102. Cuzco(?): Gay. 
Bolivia; Argentina; Chile. 

Cajophora cymbifera Urb. & Gilg, I.e. 281. 

Stems "stiffly erect," but rather suberect or even scandent, about 
3 mm. thick; petioles to 4.5 cm. long; leaves ovate or ovate-oblong, 
to 12 cm. long and half as wide, pinnatifid or pinnately parted, the 
ovate divisions usually 5-7 on each side, sparsely serrate, the teeth 
often minutely dentate, densely setose above, tomentose beneath 
and setose on the nerves; flowers in few-flowered cymes, the pedicels 
5-6 cm. long; calyx tube 7-8 mm. long, half as wide, the lobes about 



174 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

15 mm. long, remotely but deeply dentate; petals 17-18 mm. long, 
deeply cymbiform, narrowed to base, a little above it 7-8 mm. wide; 
scales 4-5 mm. long and nearly as broad, ecarinate, efilamentose 
and thin apically, the interior staminodes bearing above the base a 
cymbiform appendage 5-6 mm. long. A photo by Killip of type 
shows a weak-stemmed plant, perhaps even scandent. Although 
allied by Urban and Gilg to C. cirsiifolia in their key, 271, it is 
described as erect and its alliance seems to be with C. carduifolia, 
the staminodia with shorter appendage. The locality is given as 
Colombia but the probability is that the plant was found in central 
Peru. Illustrated (flower parts), Urb. & Gilg, I.e. pi. 7. 

Peru(?): Without locality (Lobb, type). Ecuador. 

Gajophora heptamera (Wedd.) Urb. & Gilg, Pflanzenfam. 3, 
6a: 119. 1894; 295. Loasa heptamera Wedd. Chlor. And. 2: 
218. 1857. 

Similar to C. superba; lower pinnae of the leaves often sub- 
pinnately lobed; calyx lobes 16 mm. long, 6-7 mm. wide at base, 
more or less deeply sinuately lobed; scales obsoletely 3-lobed before, 
the middle lobe emarginate, the lower margin broadly incurved. 

Peru (probably). Bolivia. 

Gajophora horrida (Britton) Urb. & Gilg, Mem. Torrey Club 3, 
pt. 3: 36. 1893; 297. Loasa horrida Britton, Bull. Torrey Club 17: 
281. 1890. 

A coarse, strict, erect, and abundantly white-bristly-pubescent 
plant with pinnate or subpinnate leaves, their divisions even to 10 
pairs and deeply dentate with acute, entire teeth ; leaves ovate, often 
broadly so, to 1.5 dm. long, usually at least a third as wide, sub- 
tomentose beneath; flowers solitary, terminal, the pedicels elongating 
to as much as 4 cm.; calyx tube 8-9 mm. long, 7-8 mm. across, 
densely setose, the ovate-lanceolate, regularly serrate dentate lobes 
about 14 mm. long; petals 26 mm. long, 12-14 mm. wide, deeply 
cymbiform-cucullate, broadly winged below, glabrous within; scales 
about 7 mm. long, 4 mm. wide, well exceeded by the dorsal threads, 
prominently 3-keeled above the base, trilobed before, the upper 
margin thickened, and broadly winged below; inner staminodia 
about 13 mm. long, dorsally appendaged; placentae 4, dichotomous, 
plane; capsule 2.5 cm. long, 2 cm. thick, densely setose on the 
12-14 prominent nerves; seeds oblong, 1.1 mm. long. As known 
from northern Bolivia it is to be expected in southern Peru. F.M. 
Neg. 38493. 



FLORA OF PERU 175 

Arequipa: Sumbay, 4,200 meters, Karl P. Schmidt (det. Standl.). 
Bolivia. 

Cajophora macrantha Killip, Journ. Wash. Acad. Sci. 18: 
94. 1928. 

Scandent terete stem 2.5 mm. thick, sparingly setose with slender 
retrorse bristles; leaves opposite, on petioles 1-2.5 cm. long, the 
lanceolate blades 5-8 cm. long, 3-5 cm. wide, acuminate, cordate, 
pinnatifid about half, the ovate-lanceolate segments regularly 
dentate, setulose above, appressed-hirsutulous on nerves and veins 
beneath; flowers 5-merous, subterminal, the peduncles about 5 cm. 
long; ovary 1 cm. long, 12 mm. wide at throat, densely setose, the 
linear-oblong lobes 10-12 mm. long, irregularly toothed; petals 
white, about 3 cm. long and 22 mm. wide, slightly narrowed at base; 
scales deeply concave, 9-10 mm. long, nearly as wide, papillose, 
green, truncate, dorsal thread none; staminodia linear, about 10 mm. 
long, the slightly longer stamens with linear-oblong anthers 1.5 mm. 
long. Section Dolichocarpae; with the scales of C. sepiaria but leaves 
less deeply divided, flowers larger and anthers longer and narrower 
(Killip), but it seems to be very near. 

Junin: Tambo de Vaca, 3,600 meters, 4468, type. 

Cajophora madrequisa Killip, Journ. Wash. Acad. Sci. 18: 
94. 1928. 

Herbaceous vine, the sparingly appressed setose terete stem 1.5-2 
cm. thick; petioles 1-2 cm. long; leaves opposite, lanceolate-oblong, 
3-6 cm. long, 1.5-3 cm. wide, subtruncate or cordulate, at base below 
the acuminate apex pinnately lobed, the lobes 5-6 each side, even 
suborbicular, dentate, finely appressed-hispidulous above, rufo- 
tomentose beneath; flowers solitary or in 2-3-flowered cymes, the 
densely retrorse-hirtellous peduncles to 8 cm. long; calyx densely 
setose, about 1 cm. wide at throat, the lobes to 2 cm. long, 3 mm. 
wide, subulate-dentate; petals 6, scarcely narrowed at base, tomen- 
tulous-hirtellous without, apparently light yellow, 17-19 mm. long, 
10-12 mm. wide; scales convex, 5-6 mm. long, 3-4 mm. wide, 
shallowly bidentate, with 3 slender threads 1.5 mm. long; staminodia 
12 mm. long, 2 mm. wide at base with a dorsal roundish appendage 
near base; stamens about 100, ovate-orbicular anthers 1.2 mm. long; 
capsule clavate, 5 cm. long, 12 mm. wide at top, tapering to short 
stipe, spiraled to right. Apparently nearest the much more densely 
setose C. scarlatina with more deeply divided leaves, larger scarlet 
flowers. 



176 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cuzco: Lucumayo Valley, 1,800-3,600 meters, (Cook & Gilbert 
294, type). Cerro de Cusilluyoc, Pennell 13829 (det. Killip). "Mad- 
requisa," "marrequisca." 

Cajophora Mandoniana Urb. & Gilg, Monogr. Loas. 299. 1900. 

Similar to C. horrida but the oblong-lanceolate leaves with fewer 
(to 6) pinnae, these sparsely and coarsely toothed, the stems usually 
2-flowered, the pedicels 3-7 cm. long, but little if at all lengthening 
in fruit; petals not recurved or winged; scales 5 mm. long, and nearly 
as broad, the middle keel little marked; capsules about 3 cm. long, 
16 mm. thick; otherwise scarcely differs and, indeed, it may be 
remarked here that more collections are needed to prove the value 
of these characters as indicating species, in this case as well as in 
several other closely related forms as indicated in the key. As the 
type is from northern Bolivia, its occurrence in Peru is probable. 
Leaves sparsely setose both sides but otherwise nearly glabrate, 
dark green above. F.M. Neg. 24168. 

Peru (probably). Bolivia. 

Cajophora pachylepis Urb. & Gilg, Monogr. Loas. 288. 1900. 

Allied to and with many of the characters of C. andina except 
that the stems are densely setose especially above, as well as the 
leaves to some extent on both sides; leaves ovate or the upper lanceo- 
late, to 7 cm. long, 2.2 cm. wide the pinnae ovate; calyx tube about 
8 mm. long, the lobes pubescent as the leaves; petals about 14 mm. 
long, 9 mm. wide; scales 4.5 mm. wide and long, with a dorsal keel 
but no threads; stamens 50-60; capsules to 3.3 cm. long, 17 mm. 
thick, scabrous; seeds 0.8 mm. long. The scant type has a facies 
that suggests that the plant may be scandent. F.M. Neg. 38495. 

Peru: (Presumably) quebrada de Bilcacota, Castelnau. Without 
locality, Mathews. 

Cajophora pauciseta Killip, Journ. Wash. Acad. Sci. 18: 
93. 1928. 

Lax, apparently decumbent herb 2 dm. long or longer, the slender 
stem finely pilose and with many stiffer hairs and a few slender 
white setae intermixed; leaves opposite; petioles to 5 cm. long, 
about half as long as the lanceolate-ovate blade, this 2-5 cm. wide, 
deeply and regularly pinnate-lobed, the 6 pairs of lobes dentate, 
thin, densely cpvered with stiff, appressed, hyaline hairs, glabrous 
on nerves, densely white-tomentose beneath; peduncles subterminal, 
erect, 4-5 cm. long; calyx densely yellow-brown setose, the lanceolate 



FLORA OF PERU 177 

lobes 1 cm. long, 2 mm. wide, remotely denticulate; petals 5, 15-18 
mm. long, 10-13 mm. wide, clawed; scales sac-like, 10-12 mm. 
long, 7 mm. wide, the 3 nerves not terminating in threads, bidentate, 
the teeth 2 mm. long; staminodia linear, marginally papillose, 12 mm. 
long; anthers broadly ovate. Scales of C. cymbifera but not with a 
triangular lobe. 

Junin: La Oroya, 3,300 meters (Kalenborn 48, type). 

Gajophora Pentlandii (Paxt.) G. Don ex Loud. Encycl. PL 
Suppl. 2: 1438. 1855; 307. Loasa Pentlandii Paxt. Mag. Bot. 9: 
7. 1842. 

Resembling and allied to C. contorta but stems to 3 mm. thick; 
petioles to 6 cm. long, leaves to 19 cm. long, 4.7 cm. wide, calyx 
lobes linear-lanceolate, subentire, to 9 mm. long, petals 21-24 mm. 
long, nearly 2 cm. wide, nearly plane; scales 6 mm. long or longer, 
4 mm. wide, with 2 threads, long-beaked before and somewhat 
winged below; stamens 80-100; otherwise similar but capsules un- 
known. Perhaps the Cuzco specimen cited under C. contorta 
belongs here. The type from Peruvian seeds (Loudon) and cultivated 
1840-46. Employed as an infusion in the treatment of sciatica 
(Herrera). Illustrated, Hook. Bot. Mag. 70: pi. 4095. F.M. Neg. 
10158. 

Cuzco: Saxaihuaman, Herrera 1268. Huasco, Herrera 3026. 
Puno: Raimondi. "Huajia-quisa," "huascjia-quisa," "orcco-quisa," 
"ckora-quisa." 

Cajophora pterosperma (R. & P.) Urb. & Gilg, Monogr. 
Loas. 324. 1900. Blumenbachia pterosperma G. Don, Gard. Diet. 
3: 62. 1834. Loasa pterosperma R. & P. in herb. 

A slender vine, the stems even below less than 2 mm. thick; 
lower leaves broadly ovate, gradually upwards narrowing to oblong- 
lanceolate above, pinnate or pinnately parted, the divisions ovate 
and more or less deeply sinuately incised, above asperous but with 
few setae and none on the pilose lower surface, to 14 cm. long, half 
as wide, the petioles to 2 cm. long; flowers 5-merous, 1-4, on pedicels 
to 7 cm. long; calyx tube about 7 mm. long, 4 mm. across, the some- 
what longer linear-lanceolate lobes distinctly and regularly dentate; 
petals to 18 mm. long, 12-14 mm. wide, deeply cymbiform and 
broadly winged below; scales 7-8 mm. long, 5-6 mm. wide, ecarinate, 
efilamentose, distinctly tricuspidate with subequal acute teeth; 
staminodia appendaged; stamens 50 or so; capsules 26-29 mm. long, 
nearly or quite 10 mm. thick, often contorted below, the placentae 3; 



178 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

seeds oval-orbicular, broadly winged. Perhaps Ecuadorian. F.M. 
Neg. 10162. 

Peru: Without locality, Ruiz & Pavdn. 

Cajophora rosulata (Wedd.) Urb. & Gilg, Monogr. Loas. 276. 
1900. Loasa rosulata Wedd. Chlor. And. 2: 219. 1857. 

Perennial with densely crowded leaves; flowering stems apparently 
erect or ascending, only 1 mm. thick; petioles to 7 mm. long; leaves 
about oblong, pinnate with regularly dentate ovate pinnae, 3.5 cm. 
long, 15-18 mm. wide, sparsely puberulent above, densely so beneath, 
esetose or nearly; flowers 5-merous, solitary, erect, reddish-orange; 
calyx tube about 3 mm. long and nearly as much across, the narrow 
lobes twice as long and deeply but remotely and subciliately dentate; 
petals 14 mm. long or slightly shorter, about 4 mm. wide, cymbi- 
form-cucullate but not winged; scales deeply saccate-convex, 3-3.5 
mm. long, 2-2.5 mm. broad, glabrous, ecarinate, efilamentose, obtuse 
before, neither incised nor thickened; stamens 30-40; placentae 3; 
capsules unknown to the authors. Flowers fiery red (Weberbauer), 
cadmium-orange (Pennell). F.M. Neg. 38497. 

Arequipa: Nevado de Chachani, Pennell 13320. Cotahuasi, 45 
meters, Weberbauer 6881. Tacna: Arica, 4,000 meters, Woitschach. 
Moquehua: Tacora, Weddell, type. 

Cajophora scarlatina Urb. & Gilg, Bot. Jahrb. 45: 470. 1911. 

Stems scandent, with long internodes, densely pilose, sparsely 
setose; petioles 7-8 cm. long; leaves oblong or lanceolate, to 16 cm. 
long, 7 cm. wide, pinnate, the pinnae extending nearly to the midrib, 
acute, shortly gray-pilose on both sides with a few setae intermixed 
only above; flowers scarlet, apparently 6-merous, solitary, the pedicels 
2-4 cm. long; calyx tube about 1 cm. long, nearly as wide, the 
regularly dentate lobes loosely setose, 15 mm. long, a third as wide; 
petals about 2.5 cm. long, nearly as wide, deeply cymbiform, laxly 
setose. Allied to C. mollis Urb. & Gilg, 291, Argentinian. F.M. 
Neg. 10167. 

Puno: Cuyocuyo, in rocks, 3,400 meters, Weberbauer 392, type. 
Arequipa: Nevado de Chachani, Pennell 13262 (det. Killip). 

Cajophora sepiaria (R. & P.) Macbr., comb. nov. Loasa 
sepiaria R. & P. in herb. Blumenbachia sepiaria R. & P. ex G. Don, 
Gard. Diet. 3: 62. 1834. L. contorta Desr. in Lam. Encycl. 3: 579. 
1791. C. contorta Urb. & Gilg, 323, not Presl, as to the plant 
described. 



FLORA OF PERU 179 

Scandent stems about 2.5 mm. thick below, where densely leafy; 
petioles to 4 cm. long; leaves oblong-ovate, often broadly so, pinnate 
or pinnately parted, the oblong or ovate divisions to 7 pairs, un- 
equally and more or less bluntly serrate, the serrations too some- 
times denticulate, sparsely asperous-setulose above, tomentulose 
beneath; flowers 2-4, remote, orange, usually pendulous; pedicels 
4.5-6 cm. long; calyx tube 7-8 mm. long, 4-5 mm. across, the linear 
lobes about 12 mm. long, remotely subulate-dentate; petals to 24 mm. 
long, scarcely wider than 15 mm., nearly plane, not winged; scales 
8-9 mm. long, half as wide, efilamentose, ecarinate, quadrangularly 
excised in front, the lobes forming a rounded tip, below narrowly 
winged and long-ciliate; staminodia conspicuously appendaged; 
stamens 60-70; capsule turbinate-oblong, scabrous and setose, 2 cm. 
long, 8 mm. thick; seeds oblong, nervose. Leaves nearly esetulose 
or with a few setae above; flowers yellow-green to orange. Illus- 
trated, Lam. 111. pi. lf.26; Juss. Ann. Mus. Hist. Nat. Paris 5: 25. 
pi. 3. F.M. Negs. 10147; 29438. 

Lima: Cheuchin, Ruiz & Pavon, type; Jos. Jussieu. Rio Blanco, 
4,000 meters, prostrate on stony talus or supported in low shrubs, 
680 (det. Killip). Canta, Pennell 14616. Obrajillo, Nee. Caja- 
marca: Ocros, Weberbauer 2684- Tacna: Weddell. 

Cajophora serropetala Macbr., spec. nov. 

Planta scandens; caulibus subadpresse sparseque setulosis haud 
vel paullo villosulis petiolis circa 2.5 cm. longis; foliis ovatis circa 
8 cm. longis, solum inferne pinnatipartitis supra pinnatilobatis 
pinnis inferioribus utrinque 1-2, profunde et irregulariter lobatis, 
supra dissite setulosis et mediocriter scabridis, subtus minutissime 
villosulis et ad nervos setulosis; pedunculis circa 1 cm. longis; flo- 
ribus 5-meris; calycis segmentis 1 (1.5) mm. latis, 1 cm. longis parce 
denticulatis; petalis paullo cymbiformis, 2 cm. longis, circa 1 cm. 
latis, margine infra medium plus minusve serratis; indumento ut 
in caule; squamis 5 mm. longis, oblongis, apice incrassatis, bicornis, 
3-nerviis dorso prope medium appendicis 3 oblongis acutis ad basin 
attenuatis, usque 3.5 mm. longis, 1 mm. latis; staminodia ad quamque 
squamam 2 papillosa apice longe filiformi-attenuata circa 5 mm. 
longa; stamina circa 60; capsula immatura dense setulosa paullo spi- 
raliter nervosa, obconica nutante, 1.5 cm. longa, superne 7 mm. lata. 

An uncommon creeper around a dwarf shrub, the flowers delicately 
transparent with a tint of pronounced green (Woy tkowski) . Ap- 
parently nearly C. Smithii but remarkable in the serrulate petals; 



180 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the scale appendages in the dried specimen are conspicuous and bright 
white except for the yellow narrowed lower half. 

Junin: Huassahuassi, Woytkowski 39, type. 

Cajophora Smithii Killip, Journ. Wash. Acad. Sci. 24: 51. 1934. 

Slightly puberulent and sparsely setulose vine; petioles 2-3.5 cm. 
long; leaves lanceolate, 5-7 cm. long, 3.5-5 cm. wide, pinnately 
lobed above the middle, divided below the middle, the segments 
dentate, appressed-scabrous above, puberulent beneath; peduncles 
1 cm. long; calyx about 7 mm. long, 5 mm. across, densely setose, the 
linear lobes to 1 cm. long, to 1 mm. wide; petals cymbiform, 1 cm. 
long, 7 mm. wide, villosulous and sparingly and weakly setulose with- 
out, orange; scales green, saccate-convex, oblong, 4 mm. long, 1.5 mm. 
wide, strongly thickened at apex, 2-horned, 3-nerved, bearing dor- 
sally near middle 3 oblanceolate acute white appendages about 2 mm. 
long, 0.8 mm. wide; staminodia falciform, filiform at tip, 5 mm. 
long; stamens about 80. Compared by the author with C. clavata of 
Argentina with different scales. 

Junin: Carpapata above Huacapistana, 3,000 meters (Killip & 
Smith 2441 9). 

Cajophora sphaerocarpa Urb. & Gilg, Monogr. Loas. 296. 
1900. 

Similar to C. horrida but the upper leaves at least with only 
2-3 pairs of leaflets; flowers several, the pedicels to 3 cm. long, 
elongating to 7 cm. in fruit; petals about 32 mm. long; capsule 24 mm. 
long and nearly as thick, the seeds 1.2-1.3 mm. long; otherwise like 
the allied species with which it was found in northern Bolivia, so 
to be expected in adjacent Peru. F.M. Neg. 38498. 

Peru (probably). Bolivia. 

Cajophora stenocarpa Urb. & Gilg, Monogr. Loas. 330. 1900. 

Scandent species marked by the presence of 2 calluses on the 
back of the floral scales below the neck; petioles to 7 mm. long; 
leaves oblong or oval-oblong, to 2.5 cm. long, 1.5 cm. wide, pinnately 
parted, the uppermost divisions entire, the lower even sometimes 
pinnate, sparsely asperous above, merely tomentulose beneath and 
without setae; flowers 5-merous, about 3, remote, the pedicels to 
3 cm. long, scarcely longer in fruit; calyx tube 5-6 mm. long, about 
3 mm. across, the narrow entire lobes nearly as long; petals little 
if at all longer than 1.5 cm., cymbiform, shortly clawed; scales 7 mm. 
long, 2-2.5 mm. wide, recurved at the calloused apex; capsule narrow, 



FLORA OF PERU 181 

about 27 mm. long, 6-7 mm. thick, sparsely setose, scarcely or not 
contorted, the strongly angled seeds about 1.2 mm. broad. Illus- 
trated, Urb. & Gilg, I.e. pi. 8 (flower parts). 

Arequipa: 4,000 meters, Hopp. Without locality, Maclean, type. 

Cajophora superba R. A. Phil. Anal. Mus. Nac. Chile, Bot. 
1891: 23. 1891; 292. Loasa heptamera Wedd., var. chelidonifolia 
Wedd. Chlor. And. 2: 218. 1857. 

Stout, erect, with lanceolate pinnate leaves that are white- 
setose above and subtomentose beneath with a few setae on the 
nerves; pinnae about oblong, with entire rounded teeth; petioles to 
4.5 cm. long, the blades to 18 cm. long, 5.5 cm. wide; flowers mostly 
solitary the pedicel scarcely 2 mm. long; calyx tube about 11 mm. 
long, nearly as wide above, the twice as long lobes obsoletely dentate; 
petals 27 mm. long, 12 mm. wide, deeply cymbiform, not winged; 
scales 8-9 mm. long, 6 mm. wide, with a triangular appendage in 
front and lateral wings; stamens very many; capsule 23 mm. long, 
nearly as thick. The following collection with flesh-colored flowers 
not surely correctly named. Illustrated, Bot. Mag. pi. 6143. 

Moquehua: Cuajones Mine, Torata, 3,200 meters, Weberbauer 
7468 (det. Killip). Bolivia; Chile. 

Cajophora tenuis Killip, Journ. Wash. Acad. Sci. 18: 93. 1928. 

Slender, scandent herb, the sparsely pilose stem less than 1 mm. 
thick; petioles 2-3.5 cm. long; leaves opposite, lanceolate, cordulate, 
acute, 3-5 cm. long, 1.5-3 cm. wide, pinnately 6-lobed, the ovate 
lobes sinuate-denticulate, thin, densely appressed-pilose above, 
hirsutulous beneath; flowers solitary, the very slender peduncles to 
5 cm. long, toward tip with many short retrorse bristles; calyx 
densely yellowish setose, the lobes narrowly linear, entire, 6-7 mm. 
long; petals 16-18 mm. long, 10-12 mm. wide, narrowed at base, 
finely hirsutulous, lightly setose, pale cream-colored; scales saccate- 
convex, 8 mm. long, 3 mm. wide, puberulous, shallowly bidentate, 
with 3 threads 3 mm. long; staminodia about 10 mm. long, thickened 
above; capsule 2 cm. long, 4 mm. wide (young). Dolichocarpae, 
and, by shape of capsules, near the different Argentinian C. cernua. 

Huanuco: Maria del Valle, 2,200 meters, 3560, type. "Ortiga." 

BEGONIACEAE Lindl. 
By L/yman B. Smith and Bernice G. Schubert 

References: A. DeCandolle in DC. Prodr. 15, pt. 1: 266. 1864; 
Irmscher in Engl. & Prantl, Pflanzenfam. ed. 2. 21: 548. 1925. 



182 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Characters largely those of Begonia, the one large genus, but the 
tepals may be joined in either or both sexes, while the ovary is some- 
times half superior. Five genera, all but Begonia of very restricted 
range. 

1. BEGONIA L. 

Herbs, erect or scandent shrubs, or rarely small trees, caulescent 
or stemless. Base fibrous or tuberous. Leaves alternate, simple in 
the Peruvian species, more or less asymmetric, stipules free, decidu- 
ous. Inflorescence usually cymose, its bracts mostly deciduous. 
Flowers monoecious. Perianth not clearly divided into 2 series, its 
tepals free, 2-11 in the Peruvian species. Staminate tepals usually 
2 or 4, when 4 the outer pair often distinctly larger. Stamens many 
or rarely few, free or more or less united in a column, anthers various 
in form, generally with a lateral dehiscence. Rudiment of the ovary 
lacking in the staminate flowers. Pistillate tepals 2 or 5 or rarely 
more. Ovary wholly inferior, usually 3-celled. Placentae usually 
axillary and bilamellate. Styles various but most often 2-branched 
with the stigmatic papillae forming a spiral down the branches and 
joined between them. Fruit usually an appendaged capsule. 

More than 700 species in the tropical and subtropical zones of 
both hemispheres. 

Weak, slender, branching annuals; placentae simple and 2-parted in 
a single ovary; staminate tepals 2 or 4; pistillate tepals 5; 
neither over 4 mm. long. 

Capsule-wings unequal, subdeltoid; stipules serrate; leaves mostly 
oblique or transverse, pilose above. 

Stem glabrous; capsule 6-8 mm. long B. humilis. 

Stem villous; capsule 10-14 mm. long B. hirtella. 

Capsule-wings subequal, semicircular; leaves mostly straight, gla- 
brous or subglabrous B. Spruceana. 

Perennial herbs or shrubs; placentae consistently simple or divided 

in each species. 
Placentae simple; leaves straight; plants caulescent; staminate 

tepals 4; pistillate tepals 5. 
Leaves broadly ovate to palmate; inflorescence a many-flowered 

diffuse cyme. 
Shrubby; leaves palmate, lobed, 2-6 dm. wide; capsule- wings 

subequal, cuneate at base B. parviflora. 

Scandent; leaves broadly ovate; capsule- wings very unequal. 

B. glabra. 



FLORA OF PERU 183 

Leaves elliptic or oblong; inflorescence racemose or of 1-2 flowers. 
Inflorescences terminal, laxly racemose; filaments free. 

B. buddleiaefolia. 
Inflorescences axillary, 1-2-flowered; filaments connate near 

base B. Poeppigiana. 

Placentae bilamellate or further divided. 
Filaments united into an elongate column; tepals narrow, acute, 

the staminate 4, the pistillate 6 B. monadelpha. 

Filaments free or united only near base. 

Capsule with narrow equal horns and a columnar apex; styles 
much branched, covered with stigmatic papillae; stam- 
inate tepals 4; pistillate tepals 5. 
Staminate tepals enlarging at anthesis, up to 18 mm. long; 

leaves broad, cordate at base B. hirta. 

Staminate tepals not enlarging at anthesis, 4 mm. long; 
leaves narrow, acute on one side of base, obtuse on 

other B. columnaris. 

Capsule with flat, generally unequal wings which extend to 

its apex. 

Staminate tepals 2; pistillate tepals 2 or 5. 
Tepals thick and fleshy, papillose-hirsute; leaves very 

broadly ovate B. lophoptera. 

Tepals membranaceous. 
Plant densely pubescent; stem geniculate. 

B. gesnerioides. 
Plant glabrous. 

Bracts of the pistillate flowers accrescent, exceeding 
the ovary; largest capsule-wing 20-36 mm. 
long; leaves nearly symmetric, elliptic. 

B. Rossmanniae. 

Bracts of the pistillate flowers inconspicuous. 
Leaves nearly or quite straight, penninerved, 
short-ciliate; style-branches 2, again divided. 

B. peruviana. 
Leaves strongly oblique or transverse, more or 

less cordate at the palminerved base. 
Style-branches again divided; leaves eciliate. 

Capsule-wings subequal B. glauca. 

Capsule- wings strongly unequal. 

B. viridiflora. 



184 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Style-branches 2, simple. 
Pistillate tepals 2; leaf-margins undulate, 

ciliate. 
Stipules ovate with a narrow apex. 

B. bracteosa. 
Stipules ample, very broadly obtuse or 

truncate B. cyathophora. 

Pistillate tepals 5 B. altoperuviana. 

Staminate tepals 4 or more. 
Bracts of the inflorescence petaloid, glandular, very large, 

covering the flowers B. velata. 

Bracts of the inflorescence not petaloid nor covering the 

flowers. 

Staminate tepals 5-11; roots tuberous. 
Plant white-lanate; tepals acute, 20-30 mm. long. 

B. polypetala. 
Plant not lanate; tepals obtuse, 4-30 mm. long; 

styles with 4 or more branches. 
Tepals red or purplish, narrow, 4-14 mm. long; 

anthers oblong. 
Tepals 4-6 mm. long; styles 4-branched. 

B. gracillima. 
Tepals 6-14 mm. long; styles many-branched. 

B. pleiopetala. 
Tepals white, broad, 2-3 cm. long; anthers broadly 

obovoid B. octopetala. 

Staminate tepals 4. 

Leaves peltate; tepals white B. Pennellii. 

Leaves basifixed. 

Roots tuberous; tepals red. 
Styles more than 2-branched, papillose all over; 
tepals 8-12 mm. long; leaves sharply lobu- 

late B. geraniifolia. 

Styles 2-branched, stigmatic tissue linear, spiral; 

tepals 25-30 mm. long. 
Stamens 8-10; outer tepals ovate. .B. Davisii. 
Stamens very numerous. 
Stem elongate; leaves broader than long. 

B. cinnabarina. 



FLORA OF PERU 185 

Stem very short; leaves mostly longer than 

broad B. Veitchii. 

Roots not tuberous; stem elongate. 
Leaves straight. 

Leaves broadly ovate, obtuse, truncate at 

base B. cucullata. 

Leaves lanceolate, acuminate, cuneate. 

B. maynensis. 

Leaves oblique or transverse. 
Capsule- wings very unequal. 
Pistillate tepals obovate, 3-4 mm. long. 

B. tovarensis. 
Pistillate tepals linear-lanceolate, to 14 mm. 

long B. stenotepala. 

Capsule- wings subequal B. piurensis. 

Begonia altoperuviana A. DC. in Ann. Sci. Nat. se*r. 4. 11: 
123. 1859. 

Suffrutescent, glabrous. Branches elongate, herbaceous, purplish. 
Leaves obliquely ovate, acute, cordate toward the middle with the 
lower lobe amply rounded, 75 mm. long, 25-40 mm. wide, crenate- 
serrate, 10-12 nerved from the base, petiole 25-50 mm. long, stipules 
deciduous, ovate-oblong, subacute. Inflorescence dichotomous, 
much-branched, peduncle straight, 75 mm. long, exceeding the 
leaves. Bracts deciduous, the upper ones oblong, 2 mm. long, 
those beneath the pistillate flowers ciliate-dentate. Staminate tepals 
2, obtuse, 6-8 mm. long. Stamens on a low torus, anthers elliptic, 
about equaling the filaments, the connective produced, ovate. 
Pistillate tepals 5 or 4(?), ovate, acute, 1-2 mm. long. Styles 3, 
bifid, the stigmatic tissue linear, spiral, continuous. Capsule obtuse 
at base, 8-10 mm. long, 16 mm. wide, the styles tardily deciduous, 
wings very unequal, the largest ovate, obtuse. 

From the boundary region of Peru and Bolivia north of Lake 
Titicaca, Weddell 4556, type. "Caupolipan" not located, but the 
species undoubtedly occurs in Peru. 

Begonia bracteosa A. DC. in Ann. Sci. Nat. se>. 4. 11: 132. 
1859, ex char. 

Erect caulescent herb, 3-25 dm. high, glabrous, very variable. 
Leaves strongly oblique or transverse, broadly ovate, abruptly 
acuminate, cordate at base, rounded elsewhere, 7-13 cm. long, 



186 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

margin undulate, crenate-serrate, short-ciliate, petioles 3-9 cm. long, 
stipules deciduous, ovate with a narrow mucronate apex. Cymes 
mostly terminal, densely or laxly many-flowered, peduncles 9-17 
cm. long. Bracts soon deciduous, the lowest broadly elliptic, 15 mm. 
long, entire, membranaceous. Pedicels very slender, 5-25 mm. long. 
Flowers rose. Staminate tepals 2, suborbicular, 8-13 mm. long. 
Stamens numerous, free, anthers oblong, equaling or exceeding the 
filaments, the connective produced. Pistillate tepals 2, 4-6 mm. 
long. Pistils 3, deeply 2-parted, the stigmas linear, spiral, continuous, 
placentae bilamellate, ovuliferous on all sides. Capsule 7-14 mm. 
long, very unequally 3-winged, the largest wing ascending, ovate or 
subtriangular, obtuse, to 2 cm. wide. 

Mountain forests. Cuzco: Machu Picchu, 2,400 meters, Vargas 
3169; West 6416; Balls 6818. Prov. Paucartambo, between Tambo- 
mayo and Pillahuata, 1,800-2,200 meters, West 7115. Paucartambo 
Valley, 2,400 meters, Vargas 83. Indefinite: Herb. Hooker (type). 

Begonia buddleiaefolia A. DC. in Ann. Sci. Nat. se"r. 4. 11: 
141. 1859. 

Plant 6-9 dm. high, succulent, fulvous-pubescent. Stem erect, 
branching. Leaves distinctly asymmetric, oblong, acuminate, acute 
at one side of base and obtuse at the other, 9-18 cm. long, 20-35 mm. 
wide, irregularly dentate, ciliate, on the under side the nerves promi- 
nent and hispid, petioles 6-12 mm. long, stipules lanceolate, 8 mm. 
long. Inflorescences terminal, laxly racemose, few-flowered, 7-12 cm. 
long. Bracts much shorter than the pedicels, lanceolate to ovate. 
Staminate tepals 4, elliptic, obtuse, sparsely pubescent, the inner 
pair much the smaller. Stamens on a low torus, anthers elliptic, 
shorter than the filaments. Pistillate tepals 5, elliptic, 4 mm. long, 
glabrous. Styles 3, slenderly 2-parted, stigmatic tissue linear, spiral, 
placentae simple. Capsule cordate at base, its wings unequal, ovate, 
obtuse. F.M. Neg. 7331. 

Humid slopes. San Martin: Tarapoto, Spruce 3998, type. 
Endemic. 

Begonia cinnabarina Hook, in Bot. Mag. pi. 4483. 1849. 
B. Clarkei Hook. f. in Bot. Mag. sub pi. 5663, pi. 5675. 1867. 

Long-caulescent herb with tuberous base, to 6 dm. high. Stem 
erect, few-branched, flexuous, stout, puberulent. Leaves obliquely 
ovate or subreniform, palminerved, 1-2 dm. in diameter, obtuse, 
lobulate, crenate-serrate, pubescent, petioles 8-17 cm. long, stipules 
ovate, acuminate. Peduncles axillary, stout, 2 dm. long, red or green, 



FLORA OF PERU 187 

2-6-flowered. Bracts elliptic, acute, 15-18 mm. long. Pedicels 
slender, decurved, 1-4 cm. long. Staminate tepals 4, broadly 
obovate, 25 mm. long. Stamens very numerous, anthers short 
and broad. Pistillate tepals 5, 25-30 mm. long. Styles 3, 2-parted, 
the stigmatic tissue linear, spiral, continuous, placentae bilamellate. 
Capsule very unequally 3-winged, 2 wings narrowly marginiform, 
the third subdeltoid. 
Peru: Clarke. Bolivia. 

Begonia columnaris Benth. PI. Hartw. 131. 1839. Sassea 
columnaris Kl. Begon. 134. 1855. Casparya columnaris A. DC. in 
DC. Prodr. 15, pt. 1: 274. 1864. 

Herbaceous, 2-4 dm. high. Branches fuscous-puberulous, becom- 
ing glabrous. Leaves asymmetric especially toward base, mostly 
straight, ovate or elliptic, acute or acuminate, base acute on one 
side, obtuse on other, 3-7 cm. long, shallowly lobed, serrate, ciliate, 
puberulent especially beneath on the nerves, petiole 2-10 mm. long, 
stipules deciduous, ovate, exceeding the petiole. Peduncles erect, 
finally exceeding the leaves, 1-4-flowered. Bracts deciduous, elliptic, 
setaceous-dentate at apex. Pedicels 6-16 mm. long. Flowers red. 
Staminate tepals 4, subequal, 4-8 mm. long, elliptic. Stamens on 
a low torus, anthers linear, the connective produced, subacute, 
filaments short. Pistillate tepals 5, elliptic. Styles 3, much-branched, 
wholly covered with stigmatic papillae, placentae bilamellate. 
Capsule 20-25 mm. long, obtuse or subcordate at base, the 3 tri- 
angular acute horns shorter than the cylindric apex. F.M. Neg. 
7313 (var. glabra. F.M. Neg. 20855). 

Huanuco: Huallaga y Mufia, 3,100 meters, Weberbauer 6723. 
Ecuador; Colombia. 

Begonia columnaris var. glabra (A. DC.) Smith & Schubert, 
comb. nov. Casparya coccinea Kl. Begon. 128. 1855. Sassea glabra 
Kl. I.e. 134, pi. 12, fig. C. 1855. Casparya columnaris A. DC. /3 gla- 
bra A. DC. in DC. Prodr. 15, pt. 1: 274. 1864. 

Completely glabrous. 

Huanuco: Muna, Ruiz. Endemic. 

Begonia cucullata Willd. Sp. PI. 4: 414. 1805. B. spatulata 
Lodd. Bot. Cab. 2: pi. 107. 1817. B. paludicola C. DC. in Bull. 
Soc. Bot. Geneve, se>. 2. 6: 125. fig. 7. 1914. 

Perennial herb, caulescent, stoloniferous, glabrous, 1-10 dm. 
high. Leaves slightly asymmetric, broadly ovate with the base 



188 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

truncate and usually inrolled, obtuse, palminerved, to 8 cm. long, 
7 cm. wide, crenate-serrate, ciliate, petioles 25 mm. long, stipules 
persistent, oblong, obtuse, 2-3 cm. long. Cymes axillary, few- 
flowered, peduncle 3-5 cm. long. Bracts persistent, ovate, serrulate, 
5 mm. long. Pedicels slender. Staminate tepals 4, 8-13 mm. long, 
the outer ones suborbicular, the inner smaller and narrowly obovate. 
Stamens free, numerous, filaments short, anthers linear. Pistillate 
tepals 4-5, obovate. Styles 3, 2-parted, the stigmatic tissue linear, 
spiral, continuous, placentae bilamellate. Capsule 24-30 mm. long, 
unequally 3-winged, the largest wing triangular, subacute, seeds 
acute in the typical variety. 

Valley forests. Cuzco: Prov. Convencion, 900 meters, Vargas 
739 (no mature seeds, variety uncertain: a form with large, dark 
red petals). To Brazil. 

Begonia cyathophora Poepp. & Endl. Nov. Gen. & Sp. 1: 7, 
pi. 11. 1835. Cyathocnemis obliqua Kl. Begon. 101. 1855. Begonia 
subciliata A. DC. in Ann. Sci. Nat. se>. 4. 11: 132. 1859. 

Succulent herb, 6-10 dm. high, glabrous. Stem subsimple, strict. 
Leaves obliquely broad-ovate, palminerved, short-acuminate, cor- 
date at base, 15-22 cm. long, to 16 cm. wide, shallowly angular- 
lobate, crenate-serrulate, pale green, petioles 5-10 cm. long, stipules 
deciduous, very broadly elliptic or obovate, distinct but simulating 
a continuous cyathium, 2-4 cm. long, entire, thin. Peduncle elongate, 
with 2 'large bracts at summit, cyme dichotomous, many-flowered. 
Upper bracts deciduous, broadly ovate. Pedicels capillary, 12-24 
mm. long. Staminate tepals 2, broadly ovate, cordate, obtuse, 
9-11 mm. long. Stamens on a low torus, filaments short, anthers 
oblong, the connective produced and dilated. Pistillate tepals 2. 
Styles 3, short, 2-parted, placentae bilamellate. Capsule 8 mm. 
long, the largest wing oblong, 2 cm. wide, the other 2 very small. 
F.M. Neg. 20861 (subciliata 8518). 

Forests and thickets. Huanuco: Cuchero, Poeppig, type. Dist. 
Churubamba, Hacienda San Carlos, waterfall Rio Ysabel, 1125 
meters, Mexia 8128. Junin: La Merced, 700 meters, Killip & 
Smith 23761. Colonia Perene", 680 meters, Killip & Smith 24988 
Cuzco: Prov. Convencion, Chaupimayo, Soukup 815. Indefinite: 
Pav6n (type of B. subciliata). Endemic. 

Begonia Davisii Veitch ex Hook. f. in Bot. Mag. pi. 6252. 1876. 

Subacaulescent herb with tuberous base. Leaves spreading, 

palminerved, obliquely ovate-cordate to suborbicular, 8 cm. long, 



FLORA OF PERU 189 

subacute, green and glossy above with scattered stiff hairs, more 
glabrous and red beneath, margins shallowly lobulate and crenulate, 
ciliate, petiole 4-5 cm. long, stout, hirsute. Peduncles simple, erect, 
12 cm. long, bright red, glabrous, 3-4-flowered. Bracts 2, broadly 
ovate or oblong, 15-18 mm. long, serrate, ciliate, red. Pedicels 2-3 
cm. long. Flowers bright red. Staminate tepals 4, subequal, 3 cm. 
long, the outer broadly ovate, subacute, the inner elliptic, obtuse. 
Stamens free, 8-10, anthers oblong, as long as the filaments. Pistillate 
tepals 5, elliptic. Styles 3, 2-parted, the stigmatic tissue linear, 
spiral, continuous, placentae bifid. Capsule-wings subdeltoid, one 
3-4 times larger than the others. 

Indefinite: Near Chupe, 3,000 meters, Davis. Endemic. 

Begonia geraniifolia Hook, in Bot. Mag. pi. 3387. 1835. B. 
petalodes Lindl. in Bot. Reg. 21: pi. 1757. 1835 (pistillate flowers 
freak or drawing error?). Eupetalum Lindleyanum Gaud. Atl. 
Bonite pi. 50. 1846. Begonia Gaudichaudii Walp. Rep. 5: 769. 1846. 
Eupetalum Kunthidnum Kl. Begon. 21. 1855. E. tuberosum Kl. I.e. 
22. 1855. 

Herb, 15-30 cm. high from a large tuberous base, glabrous, 
succulent. Stem erect, branching well above the base with leaves 
and branches usually whorled. Leaves reniform or suborbicular, 
cordate at base, palminerved with 5-9 acute lobes, to 6 cm. long and 
8 cm. wide, coarsely dentate, green, often with a narrow red margin, 
petiole 1-8 cm. long, stipules persistent, broadly ovate, entire or 
sparsely and coarsely crenate-dentate, 5-9 mm. long. Peduncles 
erect, 1-2-flowered, much exceeding the leaves. Bracts persistent, 
like the stipules. Pedicels 15-25 mm. long. Staminate tepals 4, 
subequal, 8-12 mm. long, suborbicular. Stamens numerous on a 
short but distinct column, radially seriate(?), anthers oblong, about 
as long as the filaments. Pistillate tepals 5, suborbicular. Styles 3, 
bifid with the branches 2-3-parted, each division with a tight spiral 
of stigmatic tissue at apex, placentae 2-parted with the parts usually 
again divided, ovuliferous on all sides. Capsule 6-8 mm. long, acute 
at base, the 3 wings subequal, narrowly triangular, often ascending, 
up to 25 mm. wide. F.M. Neg. 20862 (Gaudichaudii 7328). 

Clefts of limestone rocks. Lima: Amancaes, Ruiz & Pavon; 
Andre 4088. Lima, Wilkes Expedition; Mathews 734, type. San 
Lorenzo, Gaudichaud. Atocongo, 250-500 meters, Pennell 14756. 
Prov. Chancay, 5 km. north of Barranca, 200-600 meters, Worth 
9115. Endemic. "Flor de San Juan." 



190 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Begonia gesnerioides Smith & Schubert, sp. nov. 

Herba, 5 dm. alta, ferrugineo-pubescens; caule geniculata; foliis 
ellipticis, acutis, basi inaequaliter cuneatis, penninerviis, ad 11 cm. 
longis, 4 cm. latis, serratis, sparse hirsutis, petiolis 7 mm. longis, 
stipulis ellipticis, 6-8 mm. longis, brunneis, membranaceis; cymis 
laxe subpaucifloris, pedunculis 33 mm. longis; bracteis lanceolatis, 
ad 9 mm. longis, brunneis, membranaceis; pedicellis gracilibus 8-9 
mm. longis; tepalis masculinis 2, late ovato-cordatis, subacutis, 9 mm. 
longis, albis; staminibus liberis, antheris ellipticis, filamenta subae- 
quantibus; floribus femineis capsulisque ignotis. 

Forest. San Martin: Juanjui, Alto Rio Huallaga, 400-800 
meters, King 4296 (type in Gray Herb.). 

Begonia glabra Aubl. PI. Guian. 2: 916. pi. 349. 1775. B. scan- 
dens Sw. Prodr. 86. 1788. B. elliptica HBK. Nov. Gen. & Sp. 7: 180, 
pi. 641. 1825. B. lucida Otto & Dietr. in Allg. Gartenz. 16: 162. 1848. 
B. Moritziana Kunth & Bouch6 in Ind. Sem. Hort. Berol. 16. 1848. 
B. physalifolia Liebm. in Kjoeb. Vid. Medd. 19. 1853. Wageneria 
deflexa Kl. Begon. 113. 1855. W. lucida Kl. I.e. 114. 1855. W. mon- 
tana and W. glabra I.e. 115. 1855. 

Stem scandent, rooting at the nodes, glabrous. Leaves variable, 
nearly symmetrical, broadly ovate, 4-12 cm. long, short-acuminate, 
rounded or barely cordate at base, sparsely serrate and ciliate to 
entire, glabrous, petioles 1-8 cm. long, stipules persistent, ovate- 
oblong, mucronate, entire, 10-22 mm. long. Cymes axillary, many- 
flowered, diffuse, peduncle 6-12 cm. long. Bracts persistent, minute. 
Pedicels 6-16 mm. long, fine. Staminate tepals 4, the outer broadly 
obovate, 3-6 mm. long, the inner narrower. Stamens free, few, 
anthers oblong. Pistillate tepals 5, elliptic, 4-6 mm. long. Styles 
3, 2-parted, completely covered by stigmatic papillae, placentae 
simple. Capsule 6-9 mm. long, the largest wing oblong, spreading 
or barely ascending, 10-14 mm. wide, the other two marginiform, 
very narrow. 

Dense humid forests. Loreto: Alto Rio Itaya, 145 meters, 
Williams 3497. Lower Rio Huallaga, 155-210 meters, Williams 
3911. Soledad, on Rio Itaya, 110 meters, Killip & Smith 29717. 
Cachipuerto, Rio Cachiyacu, between Balsapuerto and Moyobamba, 
250 meters, Klug 3129. Cajamarca: Jae"n, Humboldt & Bonpland. 
San Martin: Tarapoto, Spruce 2846. Huanuco: Mufia, Ruiz. 
Junin: Puerto Bermudez, 375 meters, Killip & Smith 26642. 
Indefinite: Chacahuassi, Pavdn. Eastern Peru, Spruce 3960. Cen- 
tral America; West Indies; Guiana to Colombia to Bolivia. 



FLORA OF PERU 191 

Begonia glabra var. amplifolia (A. DC.) Smith & Schubert, 
comb. nov. B. scandens /3 amplifolia A. DC. in DC. Prodr. 15, pt. 1: 
362. 1864. 

Leaves up to 20 cm. long and 15 cm. wide, entire. Largest wing 
of the ovary ascending. 

San Martin: Near Tocache, Poeppig 1928. Huanuco: Pam- 
payacu, Poeppig 1063. Colombia. 

Begonia glauca (Kl.) R. & P. ex A. DC. in DC. Prodr. 15, pt. 
1: 330. 1864. Pritzelia glauca Kl. Begon. 109. 1855. 

Shrubby, glabrous. Branches subsimple, woody at base, sub- 
flexuous, even. Leaves oblique or transverse, ovate, acuminate, 
deeply cordate at base with one side broadly rounded, palminerved 
with the longest side penninerved, pale beneath, margin undulate, 
remotely denticulate, eciliate, petioles 2.5-5 cm. long, stipules 
deciduous, ovate-oblong, obtuse or mucronate, 16-20 mm. long. 
Cymes dichotomous, 4-5 times branched, diffuse, peduncle axillary, 
5 cm. long, exceeded by the leaves. Bracts deciduous, the lower 
ones not known, the upper 2 mm. long, elliptic. Pedicels 10-13 mm. 
long. Staminate tepals 2, suborbicular, 6 mm. long. Stamens on 
a torus. Pistillate tepals 2, ovate. Capsule orbicular with sub- 
equal rounded wings, 8-10 mm. long. F.M. Neg. 7341. 

Huanuco: Near Muna, Ruiz & Pavdn. Endemic. 

Begonia gracillima A. DC. in Ann. Sci. Nat. se> 4. 11: 120. 
1859. 

Low stemless herb, presumably with a tuberous base. Leaves 
broadly ovate, obtuse, more or less cordate at base, palminerved, 
12-24 mm. long, 16-24 mm. wide, irregularly crenate-dentate, 
puberulent above and on the nerves beneath, purplish beneath, 
petiole 12-24 mm. long, subglabrous. Peduncle flexuous, 75 mm. 
long, 2-4-flowered. Bracts 2, persistent, elliptic-ovate, entire, 3 mm. 
long, glabrous. Pedicels 10-12 mm. long. Staminate tepals 5-8, 
subequal, elliptic or narrowly obovate, 6 mm. long, puberulent at 
base, purplish. Stamens free, anthers oblong, the connective not 
produced. Pistillate tepals 9, 4 mm. long. Ovary puberulent, styles 
3, twice forked, the stigmatic tissue in separate spirals at the ends 
of the divisions. Capsule winged. 

Indefinite, Gay. Description compiled. This may be only a 
depauperate form of B. pleiopetala. 



192 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Begonia hirta (Kl.) Smith & Schubert, comb. nov. Casparya 
hirta Kl. Begon. 127, pi. 11, fig. C. 1855. C. cordifolia A. DC. ft 
hirta A. DC. in DC. Prodr. 15, pt. 1: 273. 1864. 

Suffrutescent, 3-6 dm. high, branching, ascending. Branches 
densely ferruginous-hirsute. Leaves obliquely ovate, acute or acu- 
minate, unequally cordate at base with one side very broadly rounded, 
5-20 cm. long, unevenly dentate, ciliate, pubescent especially beneath 
on the nerves, petioles 12-30 mm. long, stipules ovate-oblong, 6-20 
mm. long. Cymes axillary, laxly 4-8-flowered, peduncles shorter 
than the leaves. Bracts deciduous, oblong or obovate, subentire. 
Staminate pedicels 2-4 cm. long. Staminate tepals 4, accrescent, 
elliptic, becoming 18-35 mm. long. Stamens on a low torus, fila- 
ments shorter than the linear anthers. Pistillate pedicels 5-10 mm. 
long. Pistillate tepals 5, like the Staminate. Styles 3, much divided, 
the elongate linear branches covered with stigmatic papillae, pla- 
centae 2-parted. Capsule 18-22 mm. long and broad, obconic at 
base, the 3 conical lateral horns about as long as the columnar apex. 
F.M. Negs. (cordifolia 20853, var. hirta 20854). 

Huanuco: Near Muna, Ruiz. Junin: Rio Masamerich, 3,100 
meters, Weberbauer 6639. Endemic. 

Begonia hirta var. cordifolia (A. DC.) Smith & Schubert, 
comb. nov. Casparya cordifolia A. DC. in DC. Prodr. 15, pt. 1: 273. 
1864. Begonia cordifolia Warb. in Engl. & Prantl, Pflanzenfam. 3, 
6a: 146. 1894, not A. DC. 1864. 

Stem and branches soon glabrous. F.M. Negs. 7315, 20853. 

Huanuco: Muna, Ruiz & Pavon. 

Begonia hirtella Link, Enum. Hort. Berol. 2: 396. 1822. B. 
ciliata HBK. Nov. Gen. & Sp. 7: 178. 1825. B. villosa Lindl. in Bot. 
Reg. 15: pi. 1252. 1829. B. diversifolia Grah. nana Walp. in Nov. 
Act. Acad. Leop.-Carol. 19: Suppl. 1: 408. 1843. B. hirtella Link. 
var. nana A. DC. in Mart. Fl. Bras. 4, pt. 1: 345. 1861. 

Herb, 2-9 dm. high. Stem branching, villous with brownish 
hairs 2 mm. long. Leaves strongly asymmetric, transversely ovate, 
acute, very shallowly cordate at base, 3-11 cm. long, broad, palmate- 
nerved, shallowly or not lobed, crenate-serrate, sparsely pilose above, 
petioles 2-7 cm. long, stipules persistent, ovate-oblong, lacerate- 
ciliate. Cymes axillary, usually few-flowered, peduncle 3-5 cm. long. 
Bracts persistent, small, lacerate. Pedicels 4-12 mm. long. Stami- 
nate tepals 4, the outer 2-4 mm. long, suborbicular, the inner oblong, 
much smaller. Stamens free, 9-22. Pistillate tepals 5, obovate. 



FLORA OF PERU . 193 

Styles 3, 2-parted, the stigmatic tissue linear, spiral, continuous, 
placentae variable. Capsule 10-14 mm. long, the wings subdeltoid, 
unequal, the largest 9-12 mm. wide. F.M. Neg. 20892. 

Humid forests. San Martin: Tarapoto, Spruce 3982. Arequipa 
or Puno: Altos de Toledo, Meyen. 

Begonia humilis Ait. Hort. Kew. ed. 1. 3: 353. 1789. B. lucida 
Haworth, Saxifrag. Enum. 197. 1821. B. Meyeniana Walp. in Nov. 
Act. Leop.-Carol. 19: Suppl. 1: 409. 1843. B. Pavoniana A. DC. in 
Ann. Sci. Nat. se> 4. 11: 142. 1859. 

Slender herb, 3-6 dm. high, branching at base, green. Leaves 
strongly asymmetric, transversely ovate, cordate at base, acuminate 
at one side, 5-11 cm. long, subpinnate-nerved, shallowly lobed, 
serrate, ciliate, subdensely pilose above, petioles 1-4 cm. long, 
stipules persistent, narrowly ovate, serrulate, ciliate, 5-7 mm. long. 
Cymes axillary, laxly 2-5-flowered, peduncle 2-3 cm. long. Bracts 
minute, ovate, lacerate. Pedicels 5-10 mm. long. Staminate tepals 
2 or 4, the outer ones suborbicular, 3-4 mm. long. Stamens free, 
8-20, anthers elliptic, the connective produced. Pistillate tepals 5, 
2-4 mm. long. Styles 3, 2-parted, the stigmatic tissue linear, spiral, 
continuous, placentae very variable even in the same ovary, simple 
or unevenly 2-parted. Capsule 6-8 mm. long, emarginate at base, 
wings subdeltoid, obtuse, unequal, the largest 7-9 mm. wide. 
(B. Pavoniana. F.M. Neg. 20808). 

Moist shaded banks. Junin: Colonia Perene", 680 meters, Killip 
& Smith 25003. Indefinite: Mountains, Pavon. Tobago; Trinidad; 
Guiana; Venezuela; Brazil. 

Begonia lophoptera Rolfe in Kew Bull. 28. 1914. 

Erect herb, over 3 dm. high. Stems rather stout, pilose. Leaves 
oblique, very broadly ovate, acute, subtruncate at base, palminerved 
with very short angular lobes, densely serrate, 6-10 cm. long, 5-11 
cm. wide, lustrous above, pubescent on the nerves beneath, thin, 
petioles 3-12 cm. long, villous, stipules narrowly ovate, 15-25 mm. 
long, denticulate. Peduncles suberect, 6-8 cm. long, hirsute-villous, 
scarlet, 8-12-flowered. Bracts elliptic, denticulate. Flowers scarlet 
or white, hirsute-villous to nearly papillose outside. Staminate 
tepals 2, spreading, broadly elliptic, obtuse, 10-15 mm. long. Sta- 
mens on a globose torus, anthers linear, exceeding the filaments, 
the connective conspicuously produced. Pistillate tepals 2, broadly 
ovate, obtuse. Styles 3, deciduous, 2-parted, the stigmatic tissue 
linear, spiral, continuous, placentae bilamellate. Capsule pendulous, 



194 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

hirsute-tuberculate, unequally 3-winged, the smaller wings triangular, 
subobtuse, 8 mm. long, the largest wing broadly oblong, 13 mm. 
long, thickish, its apex truncate, thickened, toothed and pilose. 

Forests. San Martin, Moyobamba District, Forget. Huanuco: 
Pozuzo, Pearce 556 (type). Cuzco: near Rio Yanamayo, below 
"Pillahuata," 2,000-2,300 meters, Pennell 14069. Marcapata 
Valley, 1,200 meters, Herrera 1173. Endemic. 

Begonia maynensis A. DC. in Ann. Sci. Nat. seY. 4. 11: 
126. 1859. 

Herbaceous, 3-8 dm. high, glabrous. Stem simple, straight, 
succulent, decumbent and stoloniferous at base. Leaves erect, 
crowded at top of stem with lower ones deciduous, strongly asym- 
metric but straight, penninerved, oblanceolate, acuminate, cuneate, 
10-28 cm. long, 3-8 cm. wide, serrulate, petioles 16-36 mm. long, 
stipules subpersistent, narrowly lanceolate, acuminate, mucronate, 
entire, 1-2 cm. long, dark brown. Cymes axillary, much exceeded 
by the leaves, diffuse, few-flowered, peduncles 25-40 mm. long. 
Bracts persistent, small, narrowly lanceolate. Pedicels 10-25 mm. 
long. Staminate tepals 4, acute, the outer ovate, 6-7 mm. long, the 
inner smaller, elliptic. Stamens numerous, subfree, anthers small, 
broadly obovoid, much shorter than the filaments. Pistillate tepals 
5, lanceolate, acute, 4-7 mm. long. Styles 3, 2-parted, the stigmatic 
tissue linear, spiral, continuous, placentae 2-parted. Capsule 14-23 
mm. long, rounded at base, truncate at apex, wings subequal, rounded 
or acute, 8-12 mm. wide. F.M. Neg. 7345. 

Dense humid forests. Loreto: Maynas, Spruce 4859, type. 
Soledad, on Rio Itaya, 110 meters, Killip & Smith 29553. Mouth 
of Rio Santiago above Pongo de Manseriche, 200 meters, Mexia 6130. 
Hills to left of Rio Maranon above Pongo de Manseriche, 250 meters, 
Mexia 6348. San Martin: Zepelacio, near Moyobamba, 1,100 
meters, Klug 3716 (sterile, doubtful; leaves variegated). 

Begonia monadelpha (Kl.) R. & P. ex A. DC. in DC. Prodr. 
15, pt. 1: 286, 1864. Barya monadelpha Kl. Begon. 23, pi. 2, fig. B. 
1855. 

Herbaceous or suffrutescent, much branched, up to 2 meters 
high. Branches puberulent. Leaves very asymmetric, obliquely 
broad-ovate, acuminate, cordate at base, palmate-nerved, 5-18 cm. 
long, short-lobed, serrate, puberulent especially beneath on the 
nerves, petioles 1-11 cm. long, puberulent, stipules deciduous, 
elliptic, acuminate, entire. Cymes axillary, dichotomous, diffuse, 



FLORA OF PERU 195 

many-flowered, red, peduncle stout, 5-14 cm. long. Bracts decid- 
uous, elliptic, large. Pedicels slender, 10-35 mm. long. Staminate 
tepals 4, lanceolate, acute, 18-24 mm. long. Stamens numerous on 
an elongate column, anthers broadly elliptic, the connective scarcely 
produced. Pistillate tepals 6. Styles 3, 1 cm. long, short-connate, 
deeply bifid, placentae bilamellate. Capsule ovoid, 10-14 mm. long, 
the largest wing narrowly triangular, 25-30 mm. wide, the others 
narrowly marginiform. P.M. Neg. 20852. 

Forests. Amazonas: Chachapoyas, Mathews. Libertad: Prov. 
Pataz, valley of Rio Mishiollo, 2,700-2,800 meters, Weberbauer 7035 
(young, flowers atypical?). Huanuco: Muna, Ruiz & Pavdn, type. 
Huallaga and Mufia, Weberbauer 6714- Near Carpish, 2,800 meters, 
Stork & Horton 9895. Cuzco: near Rio Yanamayo, below "Pilla- 
huata," 2,000-2,300 meters, Pennell 14050. 

Begonia octopetala L'He"r. Stirp. 101. 1788. B. grandiflora 
Knowles & Westc. Floral Cab. 1: pi. 25. 1837. Huszia octopetala 
Kl. Begon. 18, pi. 1, fig. A. 1855. 

Perennial stemless herb with tuberous base. Leaves very broadly 
cordate-ovate or suborbicular, shallowly palmatifid with 7-9 obtuse 
lobes, 6-20 cm. long, serrate, sparsely puberulent, petioles 10-45 cm. 
long, puberulent, stipules very broadly ovate, obtuse, entire, drying 
dark brown, glabrous. Peduncles erect, 3-4 cm. long, puberulent, 
3-10-flowered. Bracts like the stipules, 7-11 mm. long. Flowers 
white. Pedicels 15-70 mm. long, puberulent. Staminate tepals 8, 
elliptic or obovate, obtuse, 2-3 cm. long. Stamens numerous, free, 
filaments elongate, anthers short, broadly obovoid. Pistillate tepals 
6. Styles 3, much-branched with the stigmatic tissue in separate 
spirals at the end of each division, placentae bilamellate. Capsule 
10-16 mm. long, its largest wing suboblong, erose-truncate, 25 mm. 
wide, several times larger than the other two. 

Thickets and rock-crevices. Lima: Near Lima, Dombey; Pavdn; 
MacLean; Mathews. San Buenaventura, 2,700-2,900 meters, Pennell 
14531. Valley of Almancaes, 305 meters, Mexia 04020. Dist. 
Pachacamac, Atacongo, 360 meters, Mexia 04051. Loma de Ata- 
congo, 600-700 meters, West 3617. Amancay, 5 km. southeast of 
Lima, 300 meters, Stork & Horton 9269. Prov. Huarochiri, quebrada 
southwest of Matucana, 2,900 meters, Goodspeed 11329. Indefinite: 
Soukup 1302. Endemic. 

Begonia parviflora Poepp. & Endl. Nov. Gen. & Sp. 1: 7, 
pi. 12. 1835. B. micrantha Steud. Nomencl. ed. 2. 1: 194. 1840. 



196 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Scheidweileria parviflora Kl. Begon. 59. 1855. Begonia myriantha 
Britton in Bull. Torrey Club 18: 35. 1891. 

Shrubby, 2-4 meters high. Trunk simple, slightly branched near 
top, 20-25 cm. in diameter. Branches spreading, 6 dm. long. 
Leaves palmate, subsymmetrical, shallowly or deeply 5-7-lobed with 
lobes sometimes subdivided, cordate at base, 2-6 dm. wide, finely 
serrate, subglabrous above, ferruginous-tomentose beneath, espe- 
cially on the nerves, bearing cystoliths, petioles up to 6 dm. long, 
ferruginous-tomentose, stipules lanceolate, deciduous. Inflores- 
cences axillary and terminal, solitary, dioecious, cymose, many- 
flowered, 2-6 dm. broad, peduncle 3-6 dm. long. Bracts minute, 
deciduous. Staminate tepals 4, elliptic or obovate, subequal, 3-4 
mm. long. Stamens on a low torus, anthers elliptic. Pistillate tepals 
5, elliptic, 4 mm. long. Styles 3, 2-branched, the stigmatic tissue 
linear, spiral, continuous, placentae simple. Capsule 8 mm. long, 
10 mm. wide, wings obcordate, cuneate-decurrent at base. 

Forests. Loreto: Balsapuerto, 220 meters, Klug 3073. Puma- 
yacu, between Balsapuerto and Moyobamba, 600-1,200 meters, 
Klug 3225. San Martin: Moyobamba, Mathews. Tarapoto, Spruce 
3945. Huanuco: Pampayaco (Cuchero), Poeppig, type. Junin: 
Pangoa, Mathews 1211. Colombia; Bolivia. 

Begonia Pennellii Smith & Schubert, sp. nov. 

E fragmentis solum cognita, verisimiliter tuberosa et acaulis, 
perennis; foliis peltatis, 7-nerviis, palmatifidis, ad 2 dm. longis et 
aequilatis, utrinque puberulis, dense serratis, lobis brevibus sed 
acuminatis, sinibus late rotundatis, petiolis ad 28 cm. longis; pedicellis 
gracilibus, glabris, 2-4 cm. longis; tepalis masculinis 4, subaequalibus, 
ellipticis, obtusis, ca. 25 mm. longis, glabris, exterioribus roseis, 
interioribus albis vel extus roseis; staminibus numerosis, liberis, 
antheris late obovoideis, brevissimis, filamentis elongatis. 

Thickets. Cuzco: above "Pillahuata," Cerro de Cusilluyoc, 
2,800-3,100 meters, Pennell 14126 (type in Gray Herb.). 

Begonia peruviana A. DC. in Ann. Sci. Nat. ser. 4. 11: 133. 1859. 

Perennial herb 1.5-2.5 meters high, glabrous. Stems subscan- 
dent. Leaves straight or nearly so, asymmetric, elliptic, acuminate, 
5-10 cm. long, 18-36 mm. wide, acute at one side of base and the 
other obtuse, penninerved, thin, serrate, short-ciliate, petioles 6-15 
mm. long, stipules oblong, 16-30 mm. long. Cymes axillary, laxly 
many-flowered, peduncles 2-5 cm. long. Bracts deciduous, minute. 
Staminate tepals 2, suborbicular, 4-5 mm. long. Stamens free, 



FLORA OF PERU 197 

anthers oblong, the connective produced. Pistillate tepals 2, like 
the staminate. Styles 3, 2-branched with the branches again 
divided, placentae bilamellate, ovule-bearing on all sides. Capsule 
unequally 3-winged, 2 costiform, the third ovate. F.M. Neg. 24199. 

Woods and thickets. Huanuco: Huallaga and Muna, 2,500- 
2,600 meters, Weberbauer 6718Junin: Huacapistana, 1,800-2,400 
meters, Killip & Smith 24514- Pichis Trail, Yapas, 1,350- 
1,600 meters, Killip & Smith 25595. Cuzco: Near Rio Yanamayo, 
below "Pillahuata," 2,000-2,300 meters, Pennell 14054- Southwest 
slope of Huayna Picchu, 2,700 meters, West 6440. Indefinite: 
near Palca, Pavdn. Near Jambrasbamba, Mathews. 

Begonia ( Begoniastrum) piurensis Smith & Schubert, sp. nov. 

Herba perennis, ad 1 m. alta, glabra, caulescens; foliis valde 
asymmetricis, transverse ovatis, acuminatis, 6.5-14.5 cm. longis, 
ad 6 cm. latis, basi late cordatis, 4-5-nervis, marginibus undulatis, 
crenato-serratis, petiolis ad 45 mm. longis, stipulis ignotis; cymis 
laxe dichotomis, submultifloris, axillaribus, pedunculis 65 mm. longis; 
bracteis deciduis, late ovatis, integris, 4 mm. longis; pedicellis graci- 
libus, 15-24 mm. longis; floribus roseis vel fere albis; tepalis mascu- 
linis 4, exterioribus late ovatis, subacutis, 9 mm. longis, interioribus 
anguste obovatis, 8 mm. longis; staminibus liberis, antheris ellipticis, 
quam filamenta multo brevioribus; tepalis femineis 5, subaequalibus, 
obovatis, subacutis, 9 mm. longis; stylis 3, profunde bifidis, stig- 
matibus spiraliter cinctis; placentis bipartitis, undique ovuliferis; 
capsulis 11 mm. longis (alis inclusis), alis subaequalibus, late sub- 
triangularibus, obtusis, basi decurrentibus, ad 8 mm. latis. 

Piura: Prov. Huancabamba, on clay banks, quebradas immedi- 
ately northwest of Canchaque, 1,200 meters, Stork 11393 (type in 
Herb. Univ. California). 

Begonia pleiopetala A. DC. in Ann. Sci. Nat. se"r. 4. 11: 
121. 1859. 

Stemless herb. Caudex globose or short-cylindric, 20-45 mm. 
long, covered with roots, gemmae and stipules. Leaves obliquely 
cordate-ovate, palminerved, shallowly and bluntly lobed, crenate- 
serrate, 3-8 cm. long, sparsely puberulent above, petioles 3-6 cm. 
long, tomentose, stipules oblong, obtuse, entire, eciliate, 6 mm. long. 
Peduncles slender, 11-15 cm. long, tomentose, 1-2-flowered. Bracts 
2, persistent, elliptic, obtuse, entire, 5-8 mm. long, drying red-brown. 
Pedicels 10-48 mm. long, tomentose. Flowers rose. Staminate 
tepals 9-11, oblong or narrowly obovate, obtuse, 8-14 mm. long, the 



198 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

outer ones pubescent at base. Stamens numerous, free, anthers 
oblong, the connective produced. Pistillate tepals 8, elliptic. Styles 
3, much-branched, with the stigmatic tissue in separate spirals at 
the end of each division, placentae bilamellate. Capsule 10 mm. 
long, one wing deltoid, 7-11 mm. wide, the others narrowly costi- 
form, inconspicuous. 

Piura: Prov. Huancabamba, above Palambla, 2,700-2,800 meters, 
Weberbauer 6026. Puno: Prov. Sandia, Limbani, 3,200 meters, 
Vargas 1317. Indefinite, MacLean, type; Soukup 1226. Bolivia. 

Begonia Poeppigiana (Kl.) A. DC. in DC. Prodr. 15, pt. 1: 
376. 1864. Lepsia Poeppigiana Kl. Begon. 63. 1855. 

Much-branched shrub or perennial herb, erect, glabrous. 
Branches slender, spreading. Leaves pinnate-nerved, elliptic, sub- 
acute, unequally obtuse at base, 8-18 mm. long, setose-serrate, thin, 
white-spotted beneath, petioles 1-2 mm. long, stipules persistent, 
lanceolate, acuminate, entire except for the setose apex, brown, 
membranaceous, 2-4 mm. long. Inflorescences axillary, the stami- 
nate 2-flowered, the pistillate 1-flowered, peduncle very slender, 
5-12 mm. long. Bracts minute, persistent. Pedicels 5-9 mm. long. 
Staminate tepals 4, 4 mm. long, the outer obovate, the inner oblong. 
Stamens borne on a column, anthers elliptic, the connective produced. 
Pistillate tepals 5, elliptic, subacute. Styles 3, 2-parted, the stigmatic 
tissue linear, spiral, continuous, placentae simple. Capsule 6-8 mm. 
long, the wings unequal, the largest deltoid, acute or obtuse, 7-8 mm. 
wide. F.M. Neg. 20823. 

On rocks in forests. Huanuco: Cuchero, Poeppig, type. Trail 
along Rio Huallaga, near Riachuela Chontalagua, 815 meters, 
Mexia 8298. Endemic. 

Begonia polypetala A. DC. in The Garden, 14: 531. 1878. 

Herb, stemless with a tuberous base. Leaves very broadly ovate 
or suborbicular, acute, deeply cordate, to 25 cm. long, 20 cm. broad, 
palmately 7-nerved with very short obtuse lobes, irregularly serrate, 
pubescent above, white-lanate beneath, petioles 18-20 cm. long, 
white-lanate. Peduncle 40-42 cm. long, white-lanate, 2-flowered. 
Bracts persistent, in pairs, elliptic, acute, 18-20 mm. long, entire. 
Pedicels 20-22 mm. long. Staminate tepals 9-10, narrowly elliptic, 
acute, 25-30 mm. long, red. Stamens numerous, anthers oblong, 
shorter than the filaments, the connective produced, acute. Pistillate 
tepals 9, 20 mm. long, deep red, tomentose outside. Styles 3, 
branched, placentae bilamellate, ovuliferous on all sides. Capsule 



FLORA OF PERU 199 

tomentose, unequally 3-winged with one very large and ascending. 
F.M. Neg. 24202. 

Andes of Peru, Froebel. Endemic. 

Begonia Rossmanniae A. DC. in DC. Prodr. 15, pt. 1: 333. 
1864. Rossmannia repens Kl. Begon. 99, pi. 9, fig. A. 1855, not 
Begonia repens Lam. 1785. 

Scandent(?) shrub, glabrous. Leaves but slightly asymmetric, 
elliptic, acuminate, rounded at base, penninerved, serrate, ciliate, 
6-9 cm. long, petioles 4-15 mm. long, stipules deciduous, oblong, 
acuminate, 3-10 mm. long. Cymes terminal, 4-12-flowered, pedun- 
cle 2 cm. long. Lower bracts large, membranaceous, red, fugacious. 
Flowers rose to brick red. Pedicels 15 mm. long. Staminate tepals 
2, broadly ovate, 6-8 mm. long. Stamens free, anthers elliptic, 
obtuse, shorter than the filaments, the connective produced. Bracts 
below the pistillate flowers persistent, accrescent, suborbicular, 
cordate, exceeding the ovary. Pistillate tepals 2, broadly ovate. 
Styles 3, 2-parted, the stigmatic tissue linear, spiral, continuous, 
placentae bilamellate. Capsule with 2 very small wings, the third 
subascending, oblong, 20-36 mm. long. F.M. Neg. 20860. 

Huanuco: Pueblo Nuevo, Ruiz & Pavon, type. Cuzco: Prov. 
Paucartambo, Cosnipata, 800 meters, Weberbauer 6939. Colombia; 
Ecuador. 

Begonia Spruceana A. DC. in Ann. Sci. Nat. se>. 4. 11 : 142. 1859. 

Slender herb, 2-6 dm. high, branching at base, green, leaves 
strongly asymmetric, straight or rarely transverse-ovate, unequally 
cuneate to cordate at base, acuminate, subpinnate-nerved, shallowly 
lobed, serrate, glabrous or nearly so, petioles 1-2 cm. long, stipules 
persistent, narrowly lanceolate, acuminate, entire, glabrous. Cymes 
axillary, laxly few-flowered, peduncle 15-30 mm. long. Bracts per- 
sistent, minute, ovate, entire. Pedicels 5-10 mm. long. Staminate 
tepals 2, suborbicular, 2 mm. long. Stamens free, few, anthers 
elliptic or ovate, the connective produced. Pistillate tepals 5, 
elliptic, 2 mm. long. Styles 3, 2-parted, the stigmatic tissue linear, 
spiral, continuous, placentae very variable even in the same ovary. 
Capsule 6 mm. long, subelliptic, the wings equal forming an even 
rounded margin 2-3 mm. wide. F.M. Neg. 7330. 

Forests. Loreto: Caballo-Cocha on the Amazon River, Osgood 5. 
Between Yurimaguas and Balsapuerto, 135-150 meters, Killip & 
Smith 28091. Balsapuerto, 220 meters, King 2840, 2923. Garni- 



200 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

tanacocha, Rio Mazan, 100-125 meters, Schunke 323. San Martin: 
Tarapoto, Spruce 4211, type. Colombia; Ecuador. 

Begonia ( Begoniastrum) stenotepala Smith & Schubert, 
sp. nov. 

Herba perennis, 2-5 dm. alta, glabra, basi fibrosa, caule elata; 
foliis transverse ovatis, abrupte acuminatis, basi truncatis, alibi 
rotundatis, palminerviis, 12-13 cm. longis, atro-viridibus, lucidis, 
marginibus integris, eciliatis, leviter undulatis, petiolis 7-9 cm. longis, 
stipulis ellipticis, rotundatis apiculatisque, 2 cm. longis, integris, 
viridibus, tenuibus; cymis axillaribus, solitariis, laxe 4-8-floris, 
pedunculis 8-10 cm. longis; bracteis inferioribus late ovatis, 2 mm. 
longis; floribus masculinis ignotis; pedicellis gracilibus, 1-2 cm. longis; 
bracteis floris feminei late ovato-cordatis, serratis, ovarium fere 
aequantibus, atro-rubris; tepalis femineis 5, subaequalibus, ad 14 
mm. longis, anguste ellipticis, rubris; stylis 3, profunde bifidis, stig- 
matibus spiraliter cincta; placentis bipartitis undique ovuliferis; 
capsulis 9-10 mm. longis, rubris, alis duabus parvis, costiformibus, 
tertia anguste subtriangulari, obtusa, ad 27 mm. lata. 

Cuzco: Forest of middle and lower montana zone, along Rio 
Tambomayo, between Tambomayo and Asuncion, 900-1,400 meters, 
West 7114 (type in Herb. Univ. California). 

Begonia tovarensis Kl. Begon. 31. 1855. B. Moritziana Kl. 
I.e. 1855. 

Herb, 2-12 dm. high, very sparsely brown-pilose to glabrous. 
Stem erect, red. Branches short. Leaves more or less asymmetric, 
broadly ovate or suborbicular, acute or rounded, cordate at base, 
2-9 cm. long, palminerved, crenate-serrate, ciliate, glabrous above, 
petioles 5-45 mm. long, stipules ovate-oblong, to 10 mm. long, 
ciliate-serrulate. Cymes axillary, few-flowered, peduncle 15-45 mm. 
long. Bracts persistent, ovate, 2-4 mm. long, fimbriate. Pedicels 
5-17 mm. long. Staminate tepals 4, the outer orbicular, 8 mm. 
long, the inner smaller, narrowly obovate. Stamens free, numerous, 
filaments short, anthers oblong. Pistillate tepals 5, obovate, 3-4 
mm. long. Styles 3, 2-parted, the stigmatic tissue linear, spiral, con- 
tinuous, placentae bilamellate. Capsule 10-15 mm. long, its wings 
decurrent, very unequal, the largest usually ascending, tapering and 
often hooked, to 23 mm. wide, seeds curved with attenuate apices. 
-F.M. Neg. 20901. 

Forests. San Martin: Zepelacio, near Moyobamba, 1,100-1,200 
meters, Klug 3389. Cuba; Mexico; Central America; Colombia; 
Venezuela. 



FLORA OF PERU .201 

Begonia Veitchii Hook. f. in Bot. Mag. pi. 5668. Sept. 1867. 
B. rosaeflora Hook. f. in Bot. Mag. pi. 5680. Dec. 1867. 

Succulent herb, 30-75 cm. high, with tuberous base, quite vari- 
able. Stem very short. Leaves very broadly cordate-ovate to sub- 
orbicular or reniform, obtuse, 5-10 cm. long, lobulate, crenate-ser- 
rate, dark green and sparsely pubescent to glabrous above, glaucous 
and often densely pubescent beneath, often bullate, petioles stout, 
3-12 cm. long, stipules triangular, red. Peduncles stout, 2-3 dm. 
long, overtopping the leaves, red or green, pubescent or glabrous, 
2-4-flowered. Bracts paired, broadly elliptic, entire or crenulate. 
Pedicels 1-3 cm. long. Staminate tepals 4, broadly obovate or 
elliptic, obtuse or emarginate, 25-30 mm. long, red. Stamens very 
numerous, free, anthers broadly elliptic, shorter than the filaments. 
Pistillate tepals 5, like the staminate. Styles 3, 2-parted, stigmatic 
tissue linear, spiral, continuous, placentae bilamellate. Capsule 
obcuneate, 2 wings very narrow, the third 2-3 times larger, sub- 
deltoid. 

Cliffs and open slopes. Cuzco: Near Cuzco, 3,700-3,800 meters, 
Pearce, type. Machu Picchu, 2,300 meters, West 8025; Vargas 536; 
Hunnewell 16057. Lares Valley, Soukup 24. Apurimac: Soccllac- 
casa Pass, Abancay-Cuzco trail, 3,800 meters, West 3818. Bolivia? 

Begonia velata Smith & Schubert, sp. nov. 

Herba ramosa, glabra sed utrinque sparse glandulosa; caule 
robusta, geniculata; foliis valde asymmetricis, palmatis, profunde 
5-fidis, ad 16 cm. latis, laxe dentatis, basi late cordatis, lobis sub- 
triangularibus, late acutis, petiolis ad 9 cm. longis, stipulis membran- 
aceis, amplis, mox deciduis; pedunculis axillaribus, ad 9 cm. longis, 
paucifloris; bracteis maximis, amplis, flores juniores occultantibus, 
pulchre rubris, glandulosis, integris, petaloideis; pedicellis ad 2 cm. 
longis; floribus fulgide rubris; tepalis masculinis 4, late ovatis, 
exterioribus ad 2 cm. longis, interioribus subduplo brevioribus; 
staminibus numerosis, liberis, antheris crasse obovoideis, quam fila- 
menta bene brevioribus; tepalis femineis verisimiliter 5, late ovatis, 
ad 25 mm. longis; stylis 3, breviter bis bipartitis, stigmatibus spir- 
aliter cinctis; placentis bipartitis undique ovuliferis; capsulis imma- 
turis ad 14 mm. longis, alis duabus parvis, tertia triangulari, ascen- 
dente, ad 11 mm. lata. 

Piura: Prov. Huancabamba, above Palambla, 2,700-2,800 meters, 
Weberbauer 6021 (type in Gray Herb.). 

Begonia viridiflora A. DC. in Ann. Sci. Nat. ser. 4. 11: 132. 
1859. 



202 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Shrubby, glabrous. Branches even. Leaves obliquely ovate, 
acuminate, cordate at base with one lobe broadly rounded and 
undulate, remotely denticulate, stipules deciduous, lance-ovate, 
mucronulate, 10 mm. long and 5 mm. wide in the single deformed 
one known. Peduncle about equaling the leaves, inflorescence 
dichotomously much-branched. Bracts deciduous, the upper ones 
elliptic, 3 mm. long. Pedicels 18-24 mm. long. Staminate tepals 2, 
broadly elliptic, 2 mm. long and 3 mm. wide in the bud. Pistillate 
tepals 2 (?), narrowly obovate, cuneate. Styles 3, deciduous only 
at maturity, short-connate, 4-5 mm. long, deeply bifid, the branches 
subdivided and verrucose toward apex, bearing stigmatic papillae 
throughout, placentae bilamellate, ovuliferous on all sides. Capsule 
12 mm. long, 16-18 mm. wide, subtriangular, obtuse at base, truncate 
at apex, the largest wing ovate, the other two marginiform, much 
smaller. 

Huanuco: Cuchero, Poeppig 1063. 

Begonia viridiflora var. parviflora Smith & Schubert, var. nov. 
Tepalis femineis 4-5 mm. longis, roseis; capsula 6-7 mm. longa, alis 
valde inaequalibus. 

Huanuco: Prov. Huanuco, Mirador-Chinchao road, 2,300 meters, 
Mexia 04152 (type in Gray Herb.). 

EXCLUDED SPECIES 

Begonia albomaculata C. DC. in Bol. Mus. Goeldi, 4: 593. 1906, 
nomen subnudum. From region of Rio Ucayali, Loreto, but descrip- 
tion insufficient for identification. 

Begonia anemonoides Azara, Voy. 2: 503. 1809, nomen subnudum. 
Large-flowered species of high altitudes, possibly related to B. 
Veitchii. 

Begonia Boissieri A. DC. Cited by the author as: "In Mexico? 
vel Peruvia?" The specimen is evidently one of the Sess and 
Mocino Mexican specimens acquired by Pavon, and is undoubtedly 
a form of B. gracilis HBK. 

Begonia Maurandiae A. DC. in DC. Prodr. 15, pt. 1: 279. 1864. 
Cited from: "Peruvia prope Lloa" from Jameson material. Actually, 
Lloa is near Quito in Ecuador. 

Begonia rubricaulis Hook. Cited by A. DC. as "E Peruvia," but 
the original description says the origin is unknown and the species 
now appears to be native of Argentina. 

Begonia serotina A. DC. in Ann. Sci. Nat. se"r. 4. 11: 121. 1859. 
Cited from "Guayaquil, Peruviae," which is actually Ecuador. 



FLORA OF PERU 203 

THYMELAEACEAE 

Rather similar to the Lythraceae but the Peruvian species 
definitely woody and always apetalous or the petals reduced to scales. 
Leaves usually alternate, entire, estipulate. Flowers, at least those 
of the Peruvian shrubs, crowded in umbels or heads, these some- 
times panicled and few-flowered. Flowers usually hermaphrodite or 
dioecious, the male at least tubular, with 4-5 imbricate lobes. Sta- 
mens in 1 or 2 series, in the former case the same number as the 
sepals and opposite them, in the latter the second series attached 
lower and alternate to the others. Petaliferous scales when present 
are between or just below the upper series of anthers. Disk if pres- 
ent variously developed or of separate scales. Ovary superior, the 
style often excentric, the cells 1 or 2, 1-ovuled, the ovule pendulous. 
Seed with or without endosperm. Domke, Bibl. Bot. Ill, 1934, has 
given a complete and scholarly treatment of the family and the 
genera. 

To this group, largely Mediterranean and African, belongs the 
delightfully fragrant, often cultivated "Daphne," Daphne Mez- 
ereum L., source of "Mezereon Bark." 

Flowers usually in simple umbels or heads; stamens in 2 series. 

1. Daphnopsis. 
Flowers panicled; stamens 4 in 1 row 2. Schoenobiblus. 

1. DAPHNOPSIS Mart. & Zucc. 

Trees or shrubs with alternate leaves and dioecious flowers borne 
in heads or umbels, the peduncles sometimes branched, usually 
terminal. Male flowers slender, with 4 spreading lobes. Anthers 
sessile or nearly so. Ovary minute with 2-4 small scales beneath. 
Female flowers smaller, often with minute staminodes, the ovary 
sessile with short thick style and capitate stigma. Seeds without 
endosperm. Lasiadenia Benth. and Lophostoma Meisn., the latter 
particularly of wide distribution in Brazil and therefore to be expected 
in Peru, have hermaphodite 5-merous flowers with 10 stamens, the 
anthers of the former (L. rupestris Benth.) subsessile, of the latter 
with filiform exserted filaments ;L. calophylloides Meisn. has branched 
peduncles, the calyx glabrous, L. ovatum Meisn., the peduncle bifid, 
the calyx puberulent. 

Peduncles simple. 

Pubescence of flowers tomentose, more or less persisting on leaves 
beneath . . . . D. Pavonii. 



204 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence of flowers sericeous, finally absent from leaves. 

D. Weberbaueri. 
Peduncles more or less dichotomously branching D. peruviensis. 

Daphnopsis Pavonii Meisn. in DC. Prodr. 14: 522. 1857. 

Branchlets, petioles and leaves beneath (especially the midnerve) 
and inflorescences softly pubescent with spreading or loosely matted 
coarse brown trichomes; petioles 2-5 mm. long, some subopposite; 
leaves oval to elliptic-obovate, rounded at base and apex or slightly 
emarginate, slightly lustrous but sparsely appressed-strigillose above, 
3-7 cm. long, 3-4 cm. wide; peduncles axillary about 3 cm. long, the 
pedicels 2-4 mm. long; male flowers infundibuliform to nearly 8 mm. 
long, the ovate obtuse lobes half as long as the tube and tomentulose 
within; female flowers unknown. The name "emarginata R. & P.," 
introduced into literature by Gilg, is invalid, as it is based merely 
on a herbarium name. From as near as Guayaquil is D. Humboldtii 
Meisn. var. (1}Boissieriana Meisn., I.e. 521, with oblong leaves 4-4.5 
cm. long, 2-2.5 cm. wide, the flower-tube and lobes subequal, the 
flower sessile. D. bogotensis Meisn., I.e., of Colombia to which 
Peruvian plants have sometimes been referred was described from 
female flowers only, but these are on peduncles only 6-12 mm. 
long, the pedicels scarcely 2 mm. long. F.M. Negs. 29665; 35034; 
36829 (var.). 

Junin: Above Palca, Ruiz & Pavdn, type. 

Daphnopsis peruviensis (Domke) Macbr., comb. nov. D. 
caribaea Griseb. var. peruviensis Domke, Notizbl. Bot. Gart. Berlin 
12: 727. 1935. 

Petioles 5-10 mm. long; leaves obovate-lanceolate, 12-15 cm. 
long, 3.5-5 cm. wide, long-caudate with acumen 1-1.5 cm. long; 
lateral nerves at angle of 70-80; male flowers sessile, the tube 7 
mm. long, the lobes 2 mm. long; female calyx tube 2 mm. long, the 
lobes half as long; fruit about 6 mm. long. Description compiled 
in part from D. caribaea of the Lesser Antilles and Margarita species 
with lateral nerves at angle of 40. Domke describes, I.e., var. 
ecuadoriensis which I have not seen. 

Junin: La Merced, Weberbauer 1813, type. 

Daphnopsis Weberbaueri Domke, Notizbl. Bot. Gart. Berlin 
12: 722. 1935. 

Branchlets glabrescent with roundish lenticels, yellowish puber- 
ulent toward the tips; leaves oblong-spatulate or obovate or some- 



FLORA OF PERU 205 

times oblong-elliptic, 3-7 cm. long, 1-3 cm. wide, attenuate to the 
short (1-2 mm.) petiole, broadly rounded or rarely slightly acute at 
tip, glabrous at maturity but beneath progressively sericeous-pub- 
escent, puberulent, glabrous; secondary nerves prominent on both 
sides, the veins reticulate; heads or umbels 10-30-flowered, the 
peduncles 1-2 cm. long, appressed-subsericeous; male flowers infun- 
dibuliform, 6-7 mm. long, 2-3 mm. broad at throat, the pedicels 
about 1 mm. long, subsericeous without, glabrous within; calyx lobes 
reflexed(?), triangular ovate, about 2 mm. long, rounded at apex, 
tomentose as the tube; anthers sessile, broadly elliptic or suborbic- 
ular, 0.5-0.7 mm. long, the lower series included; female flowers 
somewhat urceolate, about 3 mm. long, 2-2.5 mm. broad at the 
middle, the scales rudimentary; ovary glabrous, rather ovoid, 3 mm. 
high, 2 mm. broad, rounded at tip, the subglobose stigma exserted; 
disk nearly lacking. Differs in character and abundance of pubes- 
cence from D. Pavonii (Domke). A shrub 2 meters high. 

Cajamarca: In evergreen shrubs of 3,200 meters above Namas 
(Weberbauer 7217, male, and 7219, female, types). Without locality, 



2. SCHOENOBIBLUS Mart. & Zucc. 

Similar to Daphnopsis and with the 4-merous flowers of that group 
but the calyx tube broadly funnelform, petal-scales lacking and fila- 
ments elongate, a portion of them adnate to the calyx. Rudimentary 
pistil none. 

Schoenobiblus peruvianus Standl. Field Mus. Bot. 11: 
169. 1936. 

Older reddish branches glabrate, the younger hirsute or some- 
times nearly glabrous; petiole stout, to 1 cm. long; leaves oblanceo- 
late-oblong or narrowly oblanceolate, 16-30 cm. long, 5-12 cm. wide, 
abruptly long-acuminate, the acumen narrowly attenuate, gradu- 
ally attenuate to base and sometimes decurrent nearly to base of 
petiole, membranous, glabrous and prominently veined above, nearly 
concolor beneath, sometimes densely hirsute with pale rigid tri- 
chomes, these persistent or deciduous; inflorescence to 12 cm. long, 
long-peduncled, many-flowered, the thick branchlets appressed- 
pilose or subhirsute as the pedicels, these subumbellate and to 1 cm. 
long; stamens half as long as the sepals, these white, somewhat 
unequal, narrowly oblong, obtuse, 5-7 mm. long, dorsally unicostate; 
fruit suboblique, ovoid, glabrous, 15-18 mm. long, obtusely apiculate. 
Probably most nearly related to the Amazonian S. daphnoides 
Mart. & Zucc. with glabrous leaves only to 2.5 cm. wide (Standley). 



206 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Small tree to 8 meters high, the slender trunk unbranched for 
nearly 2 meters, the crown irregular, the bark dark chocolate brown. 
Uncommon in dense forest often near streams (Williams). 

San Martin: Pongo de Cainarachi, Klug 2692. Loreto: Tim- 
buchi, Alto Rio Nanay, Williams 1162, type. Paraiso, Alto Itaya 
(Williams 3354). Pebas, Williams 1707; (1878; 1595). Florida, 
Klug 2213 (det. Standl.). Rio Puranapura, Klug 3958. "Barbasco- 
caspi," "difarog-gu" (Huitoto). 

LYTHRACEAE 

Reference: Koehne, Pflanzenreich IV. 216. 1903. 

Plants various in habit and duration from annual herbs to trees 
but always with entire leaves that are usually decussate and flowers 
that are ordinarily axillary or extra-axillary, rarely racemose or 
paniculate, the pedicels often bibracteolate. Flowers usually actino- 
morphic, hermaphrodite, 4-6 (3-16)-merous. Calyx flat to tubular, 
the lobes mostly valvate, sometimes with alternate appendages. 
Petals, if present, inserted between the lobes in the calyx throat. 
Stamens inserted variously in the calyx tube, few (1-6) or many 
(even to 200 or more), the anthers generally dorsal. Ovary free, 
incompletely 2-6-celled or rarely 1-celled, the placentae then parietal. 
Style simple or none, capsule indehiscent or variously dehiscent. 
Embryo without endosperm. 

The Crape Myrtle, "Locura," Lagerstroemia flos-reginae Retz. was 
found cultivated in Iquitos by Williams; it suggests Physocalymma 
in its panicles of showy flowers with conspicuously erose-crisped 
petals but these are usually six, and the rather small, subsessile 
leaves are not at all scabrous. Likewise he collected in cultivation 
the "Amorfino," related to the Locura, Lawsonia inermis L., also 
probably native to Asia or Africa; it has an indehiscent fruit, white 
or yellowish petals only about 5 mm. long. 

Leaves conspicuously black-dotted 1. Adenaria. 

Leaves not black-dotted. 

Calyx tubular, often narrowly. 
Calyx gibbous at base, usually characteristically curved ; branches 

terete 2. Cuphea. 

Calyx straight, not at all gibbous; branches quadrate. 

^ , , . , . 3. Lythrum. 

Calyx campanulate or turbmate. 

Herbs, sometimes small. 
Small plants, the flowers as capsules scarcely 2 mm. long. 

5. Rotala. 



FLORA OF PERU 207 

Erect, often tall, the flowers larger 4. Ammannia. 

Shrubs or trees. 

Calyx persistent, inflated in fruit, the lobes not at all plicate 
in bud 6. Physocalymma. 

Calyx finally deciduous, strongly plicate and the folds fleshy. 

7. Lafoensia. 

1. ADENARIA HBK. 

A shrub or small tree with numerous black dot-like glands on the 
foliage, calyx and capsule. Leaves opposite, subsessile, oblong- 
lanceolate, more or less pubescent, especially beneath. Flowers small, 
yellowish, in axillary, shortly peduncled or sessile pseudo-umbels. 
Ovary placentae complete, continuous with the style, the calyx per- 
sistent about the indehiscent fruit, which, as the ovary, is black- 
glandular. 

Adenaria floribunda HBK. Nov. Gen. & Sp. 6: 188. 1824; 246. 

The only species, 2-6 meters high, the wood pale yellow, mod- 
erately soft, light and fine-grained. Calyx is 2.5-5 mm. long, the 
petioles 1-6 mm. long, the leaves 1.5-4 cm. long, 3-9 mm. broad, 
narrowed at both ends or obtuse at base; peduncles obsolete to 7 
mm. long, as also the pedicels. Several forms of doubtful taxonomic 
value have been recognized by Koehne. 

Piura: River banks, Weberbauer 3788; 150. Cajamarca: Are- 
nales, in clay, 3,000 meters, Stork & Norton. Huanuco: Piedra 
Grande, 3695, Pampayacu, Kanehira 246. Junin: Below Hua- 
capistana (Weberbauer, 252). Ayacucho: Ccarrapa, Killip & Smith 
22373. San Martin: Tarapoto, Spruce 2428; Lamas, Williams 6440. 
Pongo de Cainarachi, Klug 2677. Loreto: Brea Parina, Tessmann 
5536. Pongo de Manseriche, Tessmann 3810. Florida, Klug 2224. 
Yurimaguas, Williams 4169. Pebas, Williams 1855. Puno: Valley 
of Sandia, Weberbauer 572; 237. Bolivia to Brazil and Mexico. 
"Rumo-caspi," "gurima-ey" (Huitoto), "puca varilla." 

2. CUPHEAAdans. 

Peruvian species low, slenderly branched shrubs or more or less 
suffrutescent perennials with brown exfoliating bark, small to med- 
ium-sized leaves, and axillary or interpetiolar, shortly pedicelled, red 
or purplish zygomorphic flowers. Petals usually 6, infrequently 4, 
2 or none. Flowers 6-merous, but the dorsal stamens lacking. Pla- 
centae interrupted, not continuous with the style, the fruit dehiscing. 



208 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

At the base of the ovary a disk or gland is often present and 
usually easy to see if the boiled calyx is opened from the spur; this 
disk is reflexed or refracted if it appears as a pedicel to the ovary; 
again it is horizontal and flange-like, less frequently erect or even 
appressed against the ovary. The best descriptions by Koehne are 
to be found in Bot. Jahrb. 1 and 2, or, for species occurring in Brazil, 
in Mart. Fl. Bras. 13, pt. 2. 

Besides the following, Koehne proposed a herbarium name for 
Weberbauer 5860, allied to C. calophylla C. & S., 115. I did not note 
the locality, presuming the name was published. 
Calyx 15 (13)-20 (30) mm. long. 

Calyx conspicuously colored red and green; petals none. 

C. speciosa. 
Calyx green or reddish, not bicolored ; petals usually present, very 

unequal. 

Stamens somewhat exserted. 
Leaves slightly reduced into the inflorescence. . . .C. dipetala. 

Leaves greatly reduced above, bract-like C. cor data. 

Stamens scarcely if at all exserted C. ianthina. 

Calyx rarely 10 (11) mm. long; petals never strikingly unequal. 
Flowers axillary or extra-axillary. 

Flowers evidently pedicellate, the pedicels, at least some of them, 
5 mm. long or longer; disk horizontal; leaves sometimes 
oblong-linear. 
Leaves narrowly oblong-lanceolate-linear, several times longer 

than broad. 

Leaves acute both ends, especially at base. . .C. Spruceana. 
Leaves obtuse both ends, even rounded at base. 

C. hyssopifolia. 

Leaves broader, or much shorter in proportion to width. 
Stamens nearly equaling lobes; calyx 10 mm. long. 

C. Vargasii. 

Stamens not equaling the tube; calyx 5.5-8 mm. long. 
Leaves often glandular-ciliate, narrow; disk 3-lobed. 

C. ciliata. 
Leaves merely scabrous; disk narrower, not 3-lobed. 

C. serpyllifolia. 

Flowers sessile or apparently, few if any of the pedicels 3 mm. 
long; disk erect or deflexed or if horizontal the leaves never 
oblong-linear. 



FLORA OF PERU 209 

Disk horizontal or tardily refracting, plane above, subglobose 
beneath, leaves all or many of them 15 (10) mm. long or 
longer; ovules 6 (4)-12. 
Stamens shorter than tube; disk small, spreading. 

C. carthagenensis. 
Stamens equaling the tube; disk prominent, refracting. 

C. strigulosa. 

Disk erect or if deflexed strongly so and more or less excavated 
beneath; leaves mostly or all of them shorter than 10 mm. 
(4-13) or very broadly ovate; ovules 3 (-5). 
Disk deflexed; leaves lance-ovate rarely more than a few 

mm. wide C. gracilis, C. antisyphilitica. 

Disk erect; leaves mostly broadly ovate C. pustulata. 

Flowers in racemes borne above the leaves. 
Petals 6; leaves oblong-ovate or ovate. 
Pedicels (3) 4-6 mm. long; calyx usually glandular. 

C. racemosa. 

Pedicels 1-2 mm. long; calyx rarely glandular C. setosa. 

Petals 2; leaves narrowly oblong. 

Petals rounded apically ; disk refracted C. bombonasae. 

Petals acuminate; disk erect C. tarapotensis. 

Cuphea antisyphilitica HBK. Nov. Gen. & Sp. 6: 202. 
1824; 141. 

Small-leaved, sprawling or suberect shrub, the slender stems 2-7 
dm. long, strigose-hirtellous and scabrous but not viscid; leaves very 
acute or acuminate, subcordate or rotundate at base, 7-30 mm. long, 
3-4 mm. broad; pedicels 3 mm. long or shorter, bracteolate apically; 
calyx 5.5-7.5 mm. long, villous within below the stamens but gla- 
brate ventrally; petals purple, the dorsal 2 narrowly cuneate-lan- 
ceolate, the 4 ventral narrowly cuneate-oblong; style glabrous or 
villous below, sometimes finally slightly exserted. The Peruvian 
plant has been described as forma gracillima Koehne, the calyx 4-4.5 
mm. long. F.M. Neg. 17912 (var. ined. C. serpyllifolia) . Illustrated, 
Mart. Fl. Bras. 13, pt. 2: pi. 51. 

San Martin: San Roque, Williams 76^6; 7659 (both distributed as 
C. ciliata). Sandy pampas near Tarapoto, Ule 6372. Locality not 
noted by me, Weberbauer 5004. Brazil; Colombia; Venezuela. 
"Chiagari." 

Cuphea bombonasae Sprague, Ann. Bot. 17: 161. pi. 11. 
1903; 114. 



210 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Sprawling-ascending, shrubby, the branchlets toward the tips 
and the midnerve on both sides somewhat appressed-hispid with 
dark-colored trichomes; leaves green, paler beneath, minutely and 
sparsely appressed-strigose, especially above, shortly petioled, acute 
both ends, lanceolate, 4-7 cm. long, (2) 5-10 mm., usually 7 mm. 
broad; racemes simple, terminal, ashy-strigillose, at first dense, the 
caducous bracts rhombic, about 3 mm. long, densely and shortly 
hispid, minutely round-bracteolate near the base or above the mid- 
dle; calyx 6 mm. long (pilose within below stamens) the spur scarcely 
1 mm. long; petals 2, purple or white, ovate, rounded at tip, half as 
long as the calyx, the episepalous stamens exserted about one- third, 
the epipetalous ventral 4 less exserted, the 2 dorsal somewhat shorter 
than the tube; style pilose, or scarcely so, exserted; disk finally 
refracted; ovules 6 or 7. Simulates the normally 6-petaled C. 
epilobiifolia Koehne, 112, of Panama and Central America and 
probably only a variety with 2 petals, this variation already in the 
species. 

Junin: Pichis Trail, Killip & Smith 26209. Loreto: Pongo de 
Manseriche, Tessmann 4302. Pumayacu, King 3216. 

Cuphea carthagenensis (Jacq.) Macbr. Field Mus. Bot. 8: 124. 
1930; 122. L/ythrum carthagenense Jacq. Stirp. Amer. Hist. 148. 
1763. C. Balsamona Cham. & Schlecht. Linnaea 2: 363. 1827. 

Scabrous-leaved annual or half-shrub, the stems sparsely hispid 
and minutely strigillose, the former trichomes dark-colored and more 
or less glandular; leaves shortly petioled, 3-3.5 cm. long, sometimes 
smaller, usually attenuate at base, obovate to lanceolate-oblong, 
acute, more or less scabrous or early somewhat strigose, rarely gla- 
brous; pedicels scarcely 1 mm. long, bracteolate apically; calyx 4.5-6 
mm. long, sparsely hirsute; petals rose-violet or purple; stamens 11, 
shorter than the tube, the ventral 9 unequal; disk small, plane above, 
semiglobose beneath; style glabrous, always included; ovules 4-8; 
seeds smooth, narrowly winged. Illustrated, Mart. Fl. Bras. 13, pt. 
2: pi. 45. 

Peru (surely). Widely distributed. 

Cuphea ciliata R. & P. Prodr. 66. 1794; Syst. 120. 1798; 118. 
C. microphylla HBK. Nov. Gen. & Sp. 6: 201. 1824. C. loxensis 
HBK. I.e. 200. 

Subdecumbent or sprawling shrub resembling vegetatively C. 
antisyphilitica and C. gracilis but usually minutely viscid-hirtellous 
and with fruiting pedicels mostly 5 mm. long or longer; leaves sub- 



FLORA OF PERU 211 

sessile, oblong to linear-lanceolate, 5-13 (16) mm. long, 1.5-5 mm. 
broad, often glandular-ciliate and viscid-puberulent or glabrous, 
rigid-coriaceous; pedicels 3-9 mm. long, bracteolate apically; calyx 
6-8 mm. long, dark violet, densely hirtellous or viscid-hispidulous; 
petals deep purple or violet; disk horizontal, broader than ovary, 3- 
lobed; style glabrous or slightly villous above, always included; 
ovules 8-10. The 11 stamens are shorter than the tube. Often 
common on grassy or partly shrubby slopes. Illustrated, Koehne, 
I.e. 119. 

Piura: Ayavaca, Bonpland. Cajamarca: Cutervo, Jelski 311 (det. 
Szyszyl.). Huanuco: Southeast of Huanuco, 2094. Mito, 1414- 
Between Huanuco and Pampayacu, Kanehira 283. Huanuco, Dombey 
792; 793. Junin : Pillao, Ruiz & Pavdn, type. Uspachaca, 1308. San 
Martin: Between Pacasmayo and Moyobamba, Stuebel 42d, 44, 49b. 
Cuzco: Convention, Vargas 828 (det. Standl.). Yucay, Soukup 
904- Amazonas: Chachapoyas, Williams 7559. Ecuador. "Hierba 
de la culebra," "culibrilla," "yerba de culebra." 

Cuphea cordata R. & P. Prodr. 66. pi. 11. 1794; Syst. 119. 1798; 
Fl. Peruv. 4: pi. 404. 1802; 177. 

Shrub, the bark exfoliating on the older stems; leaves ovate- 
lanceolate, often acuminate, 1-6.5 cm. long, 3-40 mm. broad, 
typically lustrous above, usually acuminate and subcoriaceous; stems 
toward the tips more or less glandular pubescent; flowers conspicuous, 
bright red, the reddish calyces 17-23 mm. long, dorsally biwinged 
within, densely hirtellous and viscid-puberulent, on pedicels 4-10 
mm. long, interpetiolar, bracteolate below the apex; petals 6, the 
2 dorsal much larger, rounded or ovate-elliptic; stamens semi- 
exserted, 11; disk ovate, cordate, fleshy, deflexed; ovules 10-30, 
usually 14; style finally long-exserted. Weberbauer, 101, remarks 
that this may be the most frequent shrub of the eastern Andes 
between 1,800-2,500 meters. It usually grows in sunny thickets 
where its lax branches find partial support. Illustrated, Bot. Mag. 
pi. 4208. 

Huanuco: Chaglla, 3653. Pampayacu, Kanehira, 279; Sawada 
342. Acomayo, Ruiz & Pavon. On the Rio Huallaga, Spruce 4591, 
part. Junin : Pillao, Ruiz & Pavon. Huacapistana, Killip & Smith 
24178; 24451. Huasahuasi, Dombey. Tarma, 2,500 meters, Weber- 
bauer 1 780; 247. Ayacucho : Ccarrapa, Killip & Smith 22413. Pam- 
palca, 3,200 meters, Killip & Smith 22223. San Martin: Moyo- 
bamba, Raimondi. Cuzco: Cerro de Cusilluyoc, Pennell 14087. 
Without locality, Gay. Machupinchu, 2,400 meters, Herrera 1970. 



212 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Puno: Valley of Sandia, 2,300 meters, Weberbauer 522; 238. 
Colombia; Bolivia. 

Cuphea dipetala (L.f.) Koehne, Bot. Jahrb. 2: 422. 1882; 177. 
Lythrum dipetalum L. f. Suppl. 250. 1781. C. verticillata HBK. Nov. 
Gen. & Sp. 6: 207. pi. 552. 1824. 

Similar to C. cordata but the inflorescence very leafy, the pedicels 
sometimes extra-axillary, the stems often densely glandular-pubes- 
cent, the flowers and calyces violet and, especially, the two dorsal 
petals large, nearly equaling or slightly shorter than the tube, the 
4 ventral 1.5-3.5 mm. long, sometimes lacking; ovules 12-22, usually 
17-19. Illustrated, Fl. Serres 6: pi. 540. 

Puno: Tabina, Lechler 1 865; Soukup 523. Cuzco: Prov. Con- 
vencion, bush 1 meter high among shrubs and grasses, 1,750 meters, 
Mexia 8054 (det. Bacig.). Paucartambo, Vargas 4% (distr. as C. 
cordata). Without locality, Bonpland. "Yahuar-choncca." 

Cuphea gracilis HBK. Nov. Gen. & Sp. 6: 199. 1824; 140. 

Distinguished from C. antisyphilitica by the linear or narrowly 
lanceolate merely acutish leaves, and the reddish violet or white 
obovate or obovate-oblong petals. The Peruvian form is var. minor 
Koehne, the leaves 4-15 mm. long, 1-3 mm. wide, the calyx 4-4.5 
mm. long, ashy-strigose like the stems and branches. Doubtfully 
distinct from C. antisyphilitica unless in the shape of the petals and 
this character alone is scarcely more than a varietal difference; cf. 
the drawings in Fl. Bras. I.e. Illustrated, Mart. Fl. Bras. 13, pt. 2: 
pi. 51. 

San Martin: Moyobamba, 1,000 meters (Weberbauer 4622*). Bra- 
zil to Colombia and British Guiana. 

Cuphea hyssopifolia HBK. Nov. Gen. & Sp. 6: 199. 1824; 127. 

Similar to C. Spruceana but the leaves obtuse at both ends; calyx 
glabrous or with a few stiff trichomes; style sometimes slightly villous; 
ovules 5-8, mostly 6. Illustrated, Koehne, I.e. 127 (flowers and 
leaves). 

Peru (probably). Chile; Ecuador; Colombia; North America. 

Cuphea ianthina Koehne in Mart. Fl. Bras. 13, pt. 2: 238. 
1877; 177. 

Half shrub more or less glandular pubescent, the ascending stems 
several dm. long; leaves ovate, often subcordate at base, subsessile, 
usually about 2 cm. long, half as broad, finely setulose and above 
with some scattered longer setae; pedicels 3-15 mm. long, bracte- 



FLORA OF PERU 213 

olate medially or higher; calyx about 15 mm. long; petals usually 6, 
the dorsal sometimes even as long as the tube; style finally exserted; 
ovules 9-14 (-18). Variable; cf. Koehne for named forms. 

Puno: Slopes of Limbani, 3,400 meters, Vargas 9660. Cuzco: 
Urubamba Valley, Herrera. Bolivia. "Llinqui-llinqui." 

Cuphea pus tula ta Koehne in Mart. Fl. Bras. 13, pt. 2: 224. 
1877; 118. 

Related to C. ciliata but leaves ovate or ovate-oblong, 10-18 mm. 
long, 7-10 mm. wide, calyx only 5 mm. long, pedicels 1-2 mm. long, 
petals violet, vesicles obvious below the stamens and, especially, the 
disk erect or suberect and nearly half as long as the glabrous ovary, 
this at least equaled by the glabrous style. 

San Martin: Lamas Mountains near Tarapoto, Spruce. Without 
locality, (Mathews 1984)- Tarapoto, Williams 5393; 5492. Lamas, 
Williams 6479. 

Cuphea racemosa (L.f.) Spreng. Syst. 2: 455. 1825; 104. 
Lythrum racemosum L. f. Suppl. 250. 1781. C. spicata Cav. Icon. 4: 
5Q.pl. 381. 1797. 

A slender herbaceous or suffrutescent viscid-hirsute species with 
ovate-elliptic leaves, these usually 2-5 cm. long, 1-3.5 cm. broad, 
acute at each end, glabrous or nearly so except for the scabrous mar- 
gins; flowers in elongate racemes, on slender pedicels 3-6 mm. long, 
the pink petals about half as long as the usually glandular hirtellous 
calyx, this 5.5-10 mm. long; pedicels usually with a roundish bract 
about 1.5 mm. long at base; disk horizontal, ovate or narrowly oblong; 
style villous, finally exserted. Species unmistakable except for C. 
origanifolia C. & S., possibly occurring, but which has an erect or 
semi-erect gland at the base of the ovary (Bacigalupi). Illustrated, 
Mart. Fl. Bras. 13, pt. 2: pi. 41. 

Lima: Canta, Nee. San Martin: Near Moyobamba, Klug 3487 
(det. Standl.). San Roque, Williams 7231; 7163Loreto: Pongo 
de Manseriche, Tessmann 4398. Rio Santiago, Mexia 6160 (det. 
Standl.). South America, to Mexico and Cuba. "Hierba del fraile." 

Cuphea serpyllifolia HBK. Nov. Gen. & Sp. 6: 201. pi. 550. 
1824; 118. 

Closely allied to C. ciliata from which it differs chiefly in its 
much broader leaves with merely scabrous margins, the leaves 4-13 
mm. long, 3-6 mm. broad, thus broadly ovate or ovate-oblong, 
rounded or subcordate at base, acute; pedicels to 13 mm. long; 



214 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

calyx 5.5-7 mm. long, cylindrical in fruit, shortly glandular-his- 
pidulous; disk ovate or orbicular, carinate-convex beneath; ovules 
6-10. Used as a remedy for diarrhoea. 

Cajamarca: San Pablo, 2,400 meters, Weberbauer 3843; 257. 
Cuzco: Gay, 656; 665; 1046. Urubamba Valley, Herrera. Colom- 
bia; Venezuela. "Hierba del toro." 

Cuphea setosa Koehne in Mart. Fl. Bras. 13, pt. 2: 223. 
1877; 110. 

A perennial herb in general resembling C. racemosa, but the 
pedicels much shorter, these only 1-2 mm. long, bibracteolate at 
base, the bractlets minute; leaves to 7.5 cm. long, appressed-hirsute, 
sometimes glabrate; racemes bracted, the ciliate bracts 1-5 mm. 
long; calyx 4-6 mm. long, more or less hirtellous or hispid, rather 
long-spurred, the spur subascending; petals six, violet; disk lanceo- 
late, base ascending, but acutely refracted at the middle. Illus- 
trated (flower), Koehne, I.e. 111. 

Puno: San Gaban, Lechler 2322. Cuzco: Gay 1087; 669 (det. 
Bacigalupi). Bolivia to Mexico and Tobago. 

Cuphea speciosa (Anders.) Kuntze, Rev. Gen. 3, pt. 2: 96. 
1898; 159. Melvilla speciosa Anders. Journ. Sci. 25: 207. 1807. 

A shrub 1-2 meters high, the slender virgate or subscandent 
branches terminating in dense bracted racemes of showy red and 
green calyces, these 2-3 cm. long, strigose puberulent and usually 
glandular hirtellous, borne on pedicels 3-11 mm. long, bracteolate 
below the middle or near the base; petals none; stamens longer or 
little exserted, 11. If the name C. speciosa Hort. (cf. Koehne) is 
accepted as in conflict this plant must be called C. Melvilla Lindl. 
Bot. Reg. 10: pi. 852. 1824. Illustrated (flower), Koehne, I.e. 159. 

Amazonas: Nazareth, Osgood 11. Loreto: Puerto de Neuvo York, 
(Huber 1582). Iquitos, Williams 8027; Mexia 6390. Near Yuri- 
maguas, Klug 2803. Between Urarinas and San Regis on the 
Maranon, Poeppig. Pro, Williams 1976. Caballo-Cocha, Williams 
2302. Rio Itaya, Williams 48. Rio Nanay, Williams 467. Pebas, 
Williams 1873. Widely distributed, South America. "Bosquesito." 

Cuphea Spruceana Koehne in Mart. Fl. Bras. 13, pt. 2: 226. 
1877; 128. 

A low shrub with sessile or subsessile, glabrous or nearly glabrous, 
oblong-linear or linear-lanceolate leaves, to 7 mm. wide, 1-3 cm. 
long; flowers pink, axillary, on apically bracteolate pedicels 5-9 mm. 
long; calyx minutely and sparsely hirtellous, 7-8 mm. long; stamens 



FLORA OF PERU 215 

and tube subequal; style glabrous; ovules 10-14; seeds obtusely 
or retusely margined. Flowers white and yellow (Klug). F.M. 
Neg. 21926. 

San Martin: Juanjui, Klug 4375. Chazutu, Klug 4819. Junin: 
Rio Perene", Killip & Smith 25147. Loreto: Yurimaguas, Poeppig; 
Spruce 4591, type. Pongo de Manseriche, Mexia 6156; 6155 (det. 
Standl.); Tessmann 4839. Bolivia. 

Cuphea strigulosa HBK. Nov. Gen. & Sp. 6: 204. 1824; 123. 
C. strigulosa HBK. subsp. opaca Koehne in Mart. Fl. Bras. 13, pt. 
2: 257. 1877. 

A small shrub with medium-sized scabrous leaves (1) 2-3 (4) cm. 
long and axillary subsessile pink flowers; stems usually more or less 
hirsute; leaves somewhat petiolate; pedicels 1-2 mm. long, bracteolate 
toward the tip; calyx 6-8 mm. long, strigillose or very shortly hirtel- 
lous and hispidulous, villous within below the stamens, the oblong 
roseate or violet petals half as long; infrastaminal vesicles none; 
stamens usually shorter than the tube; disk plane above, semiglo- 
bose beneath; ovules 6-8 (-12); seeds margined. Var. Sinclairii 
Koehne, pedicels 2.5-4 mm. long, is Ecuadorian. Nearly C. cartha- 
genensis, to which some of the following collections could perhaps 
be referred, but apparently they all have the slightly different disk 
and longer stamens of the HBK. species. 

San Martin: Moyobamba, Mathews 1632; Klug 3274; 3444 (both 
distr. as C. carthagenensis) . Chazuta, Klug 4072 (distr. as C. cartha- 
genensis). San Roque, Williams 6967; 7062; 7690. Lamas, Williams 
6478. Tarapoto, Williams 6774. To Trinidad and Colombia. 

Guphea tarapotensis Sprague, Ann. Bot. 17: 160. pi. 11. 
1903; 114. 

A little shrub 2-5 dm. high, closely allied to C. bombonasae but 
the disk erect, cylindrical; bracts broadly ovate, long and densely 
setose-ciliate, densely strigose beneath; pedicels axillary; petals 2 
or in some flowers 3 or 4; ovary villous dorsally, otherwise glabrate; 
disk about one-half as long as the ovary, oblong, subcomplanate ; 
ovules 6-8; seeds 1.3 mm. long and broad. Description from 
Koehne based on the Ule specimen. 

San Martin: Tarapoto (Spruce, type). Loreto: San Antonio, 
Ule 6755 (det. Koehne). 

Cuphea Vargasii Macbr., spec. nov. 

Fruticulosa 3-10 dm. alta; caulibus base procumbento-adscen- 
dentibus demum ut videtur suberectis, flexuosis, parce ramosis 



216 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

mediocriter foliosis plus minusve glanduloso-setulosis et minute 
strigillosis; petiolis vix 1 mm. longis; foliis ovatis vel ovato-lanceo- 
latis base subcordatis, apice acutis vel breviter acuminatis; plerum- 
que 17-20 mm. longis, circa 10 mm. latis, superioribus paullo reductis, 
supra sparse setulosis, nervis vix notatis, subtus minute scabrido- 
hispidulis, nervis prominentibus; pedicellis ad 6 mm. longis, apice 
bibracteolatis; calycibus 10 mm. longis, fere cylindraceis baud 
bilabiatis, obscure vel vix calcaratis minutissime strigillosis et sparse 
glanduloso-hispidulis, intus infra staminibus glabris; staminibus 11, 
9 aequalibus fere exsertis; petalis subaequalibus late obovatis 3.5- 
4.5 mm. longis; ovario 2.5 mm. longo, stylo 3 mm. longo glabro 
incluso; ovulis 8 (9); disco suberecto subgloboso distincte 3-lobato 
vel 3 rotundato-carinato; seminibus fere orbiculatis, 2 mm. longis, 
1.75 mm. latis, fusco-marginatis, exalatis. In some important 
characters this species approaches C. calophylla C. & S. of eastern 
Brazil; its disk is somewhat similar to that of C. ciliata, to which 
it is probably allied. Flowers red-lavender or violet-purple; old 
plants with long, slender, naked woody stems (collectors). 

Apurimac: Between Abancay and Aurpury, 2,600 meters, Stork, 
Norton & Vargas, 10589, type. Sunny hillside among boulders, 
1 km. east of Abancay, 2,400 meters, Stork, Norton & Vargas 10558. 

3. LYTHRUM L. 

Smooth-barked slender herbs or sometimes somewhat suffrutes- 
cent toward the base. Stems or branches often wand-like, the leaves 
ascending, the calyces closely appressed. Flowers solitary, axillary 
or in terminal spikes. 

Ly thrum maritimum HBK. Nov. Gen. & Sp. 6: 194. 1824; 69. 

More or less suffrutescent plant with rather small axillary roseate 
purple flowers; leaves 7-33 mm. long, 2-14 mm. broad, obtuse or 
rounded at base; calyx 5-8 mm. long, the petals somewhat shorter; 
stamens 6, subequaling the style at anthesis, the latter 2 times 
shorter than or about equaling the ovary that is narrowly annulate 
at base. The annual or short-lived L. hyssopifolia L., 65, the ovary 
eannulate, the stamens usually 4-6, called "Romerillo" in Chile, 
has been reported but probably by error; the similar but perennial 
European species, L. flexuosum Lag., 68, with brachystylous flowers, 
12 stamens, may be found as an introduction. Illustrated, Mart. 
Fl. Bras. 13, pt. 2: pi. 39. 



FLORA OF PERU 217 

Lima: In the river bed at Callao, and Lima (Wilkes Exped.). 
Chanchai, Dombey. Without locality, Ruiz & Pavon. Tacna: A 
shrub more than 1 meter high, Shepard 280 (det. Johnston) ; Werder- 
man 710. Apurimac: Curahuasi, Vargas 9610; 1211. Uruguay and 
Argentina to Chile and Colombia. 

Ly thrum Salicaria L. Sp. PI. 446. 1753; 73. 

Inflorescence a long, interrupted spike, the clustered flowers 
large, magenta-colored; calyx 4-8 mm. long, the lobes usually half 
as long as the tube, the petals much longer than the calyx. The 
single record known for South America is based on a poor specimen 
and doubtless represents a casual introduction. 

Piura: Pisacoma (according to Koehne). Widely introduced 
from Europe. 

4. AMMANNIA [Houston] L. 

The Peruvian species are smooth, erect herbs 2 dm. to 1 meter 
high, with linear-lanceolate, oblong, closely sessile leaves and small, 
green or red, sessile flowers that are solitary or clustered in the leaf- 
axis. Petals none. Ovary 2-4-celled. Capsule globose or ellipsoid, 
papyraceous, irregularly dehiscing transversally, the septae not at 
all striate. Bractlets whitish, fertile. 

Ammannia latifolia L. Sp. PI. 119. 1753; 50. 

Leaves 2-8 cm. long, 2-15 mm. broad; calyx lobes obsolete; 
petals none; style very short. Wet or cultivated ground. 

Lima: Callao, Gaudichaud 141- In the river bed at Callao 
(Wilkes Exped.). Chancai, Ruiz & Pavon. Brazil to southeastern 
United States. "Crab-weed," "yerba de cancer." 

Ammannia coccinea Rottb. PI. Hort. Univ. Havn. Progr. 
Descr. 7. 1773; 49. 

Similar to the preceding but, especially, the leaves always 
auricled at base, the calyx lobes obvious and the style one-half to 
two times as long as the ovary. A. auriculata Willd. found as near 
as Guayaquil by Ruiz & Pavon, is similar, but the flowers are mostly 
long pedicellate. 

Cajamarca: Raimondi. Nearly cosmopolitan. 

5. ROTALA L. 

Glabrous herbs closely allied to Ammannia, but the partitions of 
the septicidal capsules densely and elegantly horizontally striate 
and the bractlets sterile. 



218 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rotala ramosior (L.) Koehne in Mart. Fl. Bras. 13, pt. 2: 194. 
pi. 39. 1877; 32. Ammannia ramosior L. Sp. PI. 120. 1753. 

Stems 1-several dm. long, quadrate, often ascending and branched ; 
leaves 1-4 cm. long, 2-11 mm. broad, somewhat attenuate to the 
short petiole; bractlets usually equaling or half as long as the calyx, 
this becoming 4-5 mm. long, the lobes equaled or slightly exceeded 
by the white or roseate petals. 

Loreto: Caballo-Cocha, Williams 2299. North and South Amer- 
ica; Philippine Islands. 

6. PHYSOGALYMMA Pohl 

A tree with widely spreading branches, decussate leaves and 
showy flowers borne in panicled racemes. Flowers 8-9-merous; 
calyx subglobose-turbinate, about 16-striate, vesicular and persistent 
in fruit. Petals crenate-undulate. Stamens 24-28. The tree was 
long supposed to be the source of tulip-wood, or rose-wood, of Brazil, 
which now is known to be obtained from a leguminous tree; com- 
pare Record & Mell, Timbers of Tropical America 288, 455. 1924. 
According to these authors, the somewhat pinkish wood suggests 
that of maple or birch, but is not suitable for cabinet work. 

Physocalymma scaberrimum Pohl, Flora 10: 153. 1827; 210. 
P. floridum Pohl, PI. Bras. Ic. 1: 100. pis. 82-83. 1827. 

A beautiful tree 5-25 meters high, with scabrous leaves, cori- 
aceous, lustrous, several to 10 cm. long, 3-6 cm. wide, and bright 
purple-red flowers, the rhombic petals about 16 mm. long and broad. 

Loreto: Near Yurimaguas, Poeppig 2077; Williams 3882; 4025; 
Killip & Smith 27551 . Junin : On river bluff, La Merced, 5398. 
Colonia Perene", Killip & Smith 24975. Chanchamayo Valley, 
Schunke 432. Bolivia; Brazil. "Huianava," "huainuma." 

7. LAFOENSIA Vand. 
Calyplectus R. & P. Prodr. 73. 1794. 

Very much like Physocalymma but glabrous, the few large flowers 
borne in dense subpaniculate racemes, the campanulate or semi- 
globose calyx coriaceous, its limb remarkably plicate, finally decid- 
uous. Stamens 16-32. Capsule indurate. Named in honor of 
Joannis de Braganza, Duke of Lafoens in the early eighteenth 
century. A native name is "chusma." 

A beautiful yellow dye is obtained from two South-American 
species, L. pacari St. Hil. and L. punicaefolia DC. 



FLORA OF PERU 219 

Lafoensia acuminata (R. & P.) DC. Me*m. Soc. Phys. Hist. 
Nat. Geneve 3, pt. 2: 73. 1826; 218. Calyplectus acuminatus R. & P. 
Syst. 129. 1798. L. speciosa (HBK.) DC. I.e., at least as to Peru. 

Branches terete; leaves 5-10 cm. long, nearly half as broad, 
oblong or obovate-oblong with many subhorizontal nerves that are 
equally prominent both sides, 3-4 mm. apart; leaf -pore apparently 
lacking (in true L. speciosa of Colombia it is present); pedicels 
18-23 mm. long, terete or subcompressed dorsally, the axis raceme 
4-angled; calyx 2 cm. long; flowers 14-merous; pod spherical, accord- 
ing to Ruiz & Pavon radiately stellate from the middle to the apex. 
The sap stains the skin (Rimbach). F.M. Neg. 17918. 

Huanuco: Pozuzo, Ruiz & Pavdn. Amazonas: Chachapoyas, 
Mathews (referred toL. speciosa by Koehne). Cajamarca: Cutervo, 
Weberbauer 7132. Bolivia? Ecuador. "Amarillo," "pocol," "chuspo," 
"cabeza de monge." 

Lafoensia punicaefolia [Bert.] DC. Me"m. Soc. Phys. Hist. Nat. 
Geneve 3, pt. 2: 86. pi. 1. 1826; 218. 

Branches early angulate; leaves rather light coriaceous, oblong- 
lanceolate, 5-8 cm. long, 2-2.5 cm. wide, acute at slender petiole, 
more or less abruptly and bluntly acuminate-caudate; leaf -pore 
small; pedicels 2-4 cm. long, the inflorescence sometimes a simple 
raceme; flowers 12-16-merous; calyx 3-4 cm. long, campanulate but 
pyriformly contracted below the middle; petals about 3 cm. long, 
the filaments 12 cm. long; capsule apiculate-ovoid. A tree about 
15 meters high; flowers flesh-red and yellow (Klug), this collection 
apparently somewhat larger than typical; it was given an herbarium 
name by Standley and may indeed be distinct from the species of 
DeCandolle, originally from Central America but more material 
is needed upon which to base it. The name has been written 
"punicifolia." 

San Martin: Juanjui, 400-800 meters, Klug 4327. Bolivia (fide 
Koehne) to Venezuela and Guatemala. 

RHIZOPHORACEAE. Mangrove Family 

Trees or shrubs with usually opposite, stipulate, more or less 
coriaceous leaves and hermaphrodite flowers borne in axillary 
inflorescences. Stipules united between the petioles, soon deciduous. 
Calyx tube adnate in Rhizophora, free in Cassipourea, the persistent 
lobes valvate. Petals often lobed or fringed, convolute or inflexed 



220 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

in bud, the stamens usually many, often opposite to the petals in 
pairs, inserted on a lobed disk. Ovules usually 2 pendent from near 
the tip of the 2-5(6) cells. 

Strangely enough the Mangrove so conspicuously fringing most 
tropical shores that are muddy seems to be unrecorded from Peru; 
Weberbauer does not mention it, but for that matter the related 
family Combretaceae is omitted by him. However the Mangrove 
seems really to be absent, probably from the lack along the Peruvian 
coast of the muddy flats and estuaries that elsewhere it so often 
characteristically fills with its curved prop-roots. Apart from its 
habit, Rhizophora Mangle L. may be known, if found, by the forked 
flower-peduncles, the cupulate-bracted calyx, the inferior ovary, not 
to mention the remarkable perforation of the fruit that persists 
on the tree by the radicle of the embryo, the elongate club-like 
radicle then descending into the mud. 

CASSIPOUREA Aublet 
Reference: Alston, Kew Bull. 241-276. 1925. 

Smooth with entire or somewhat toothed leaves and white or 
yellowish flowers usually clustered and more or less pediceled, the 
pedicels with minute bractlets at their base. Petals 4-5, thickly 
fringed. Stamens 10-40 on the disk margin, this with as many 
lobes. Fruit somewhat fleshy, tardily dehiscent septicidally with 
1 ariled seed in each of the 3-4 cells. 

Cassipourea peruviana Alston, I.e. 268. C. Ulei Briq. Candollea 
4: 347. 1931. C. Poeppigii Briq. I.e. 349. C. elliptica var. dentata 
Engler in Mart. Fl. Bras. 12, pt. 2: 430. 1876. 

A tree, the slender branchlets slightly pubescent toward the tips 
or glabrate; petioles 3-8 mm. long; leaves lanceolate to broadly 
elliptic, somewhat rounded to the shortly cuneate base, more or 
less acuminate, entire or obscurely and remotely denticulate, often 
1-1.5 dm. long, 4-6 cm. wide, often smaller, glabrous or a little 
pilose at first on the midrib; stipules 3-5 mm. long; flowers few to 
several; pedicels 6 (5) mm. long, apically articulate, distinctly but 
sometimes rather sparsely short-hirsutulous or pilose; calyx broadly 
campanulate, glabrous or with a few to a number of appressed 
trichomes, the 5 lobes densely sericeous within; petals to about 10 
mm. long, densely fringed; stamens about 20, exserted; ovary 3- 
celled, with the style appressed strigose. G. guianensis Aubl., 269, 
and C. Spruceana Benth., 270, both known from the Amazon Valley, 



FLORA OF PERU 221 

have sessile or subsessile flowers, the former with petioled leaves 
and the calyx without somewhat pilose, the latter with subsessile, 
slightly cordate leaves, the calyx without glabrous. Although at 
one time I thought to distinguish C. Poeppigii and so named two of 
the collections cited, it seems to me now, with a larger series of 
specimens at hand, that there is no fundamental distinction and that 
there is only one species concerned, somewhat variable in leaf size 
and form and in pubescence and length of pedicels. C. Ulei may be 
a local variety, the leaves to 3 cm. wide. F.M. Negs. 23233; 23235. 

San Martin: Along the Rio Mayo near Tarapoto, Spruce 4005, 
type. Loreto: Yurimaguas, Poeppig 2088; 2234 (the latt