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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 latter type, 
C. Poeppigii). Florida, Rio Putumayo, King 2124; 2290; 2235; 
2167 (all det. Standl.). Mishuyacu near Iquitos, King 1219; 
455. Rio Acre: Cobija, Ule 9638 (type, C. Ulei). "Eto sima-ey," 
"jamoro-ey," "farocano-ey" (Huitoto). 

COMBRETACEAE 

Reference: Eichler in Mart. Fl. Bras. 14, pt. 2. 77-127. 1867. 

Trees, shrubs or lianas with usually opposite leaves and no 
stipules. Flowers most often hermaphrodite, racemose or spicate, 
sometimes capitately. Calyx tube adnate, the 4-5 (6-8) lobes 
usually valvate. Petals 4-5 or wanting, often scarcely obvious, 
the stamens as many to twice as many or more, inserted on the calyx, 
the filaments inflexed in bud. Anthers versatile, opening by slits. 
Stigma simple or rarely 4-lobed, the ovary 1-celled, the 2-6 ovules 
funicularly suspended from the top of the cell. Fruit often winged, 
drupe-like. Nearly Rhizophoraceae as to technical character but 
often well-marked in appearance of flowers and inflorescence; the 
ovary in Rhizophoraceae 1-celled only by disappearance of the 
partitions. The expression "calyx tube" or "receptacle" refers here 
to the tube containing the inferior ovary and to its prolongation 
with the calyx lobes, these often forming 2 parts, more or less dis- 
tinctly, the former being the "lower" the latter the "upper." 

Quisqualis indica L., cultivated generally in the tropics, has been 
collected at Iquitos; the style is adnate to the calyx tube, this often 
5 cm. long or longer and topped by the spreading rose or scarlet 
petals that are about 1 cm. long; the appearance of the flower is 
apocynaceous. 

Besides the reference cited above I wish to acknowledge my 
indebtedness to the work of A. W Exell on the family, particularly 
in Pulle's Flora Surinam 3: 164-177. 1935. 



222 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx tube sericeous and with 2 minute bractlets near top. 

1. Laguncularia. 
Calyx tube ebracteolate. 

Leaves opposite; petals present, small 2. Combretum. 

Leaves alternate or verticillately crowded; petals wanting. 

Anthers versatile 3. Terminalia. 

Anthers fixed to the filament tip 4. Buchenavia. 

1. LAGUNCULARIA Gaertn. f. 

Shrub or tree with leathery, opposite oblong-elliptic leaves that 
are scarcely veined. Flowers mostly hermaphrodite in axillary 
spikes. Calyx tube only slightly extended above the ovary, obscurely 
bibracteolate near the tip, 5-parted, persisting. Petals 5, soon 
caducous. Stamens included or only barely exserted. 

Laguncularia racemosa (L.) Gaertn. f. Fruct. 3: 209. pi. 217. 
1805; 102. Conocarpus racemosa L. Syst. ed. 10. 930. 1759. 

Petioles to 2 cm. long, biglandular near the tip; calyx tube 3-6 
mm. long, gray-puberulent, crowned by the persistent calyx. The 
Peruvian collection with longer leaves than usual was described as 
forma longifolia Macbr. The shrub usually is associated with the 
Mangrove but if the latter was with it, unfortunately it was not 
collected; cf. remarks under Rhizophoraceae. 

Tumbez : Tumbez near the sea, Mangrove formation, Weberbauer 
7627, type, forma. Tropical America and Africa. 

2. COMBRETUM L. 

Shrubs, usually scandent, sometimes trees, the leaves entire and 
opposite or rarely whorled, the flowers borne in spikes that are often 
paniculate. Petals small, inserted between the calyx lobes. Style 
free from the upper receptacle. Fruit 4-6-angled or membranous- 
winged and then suggesting that of some Sapindaceae. 

The species, at present at least, seem to be scarcely distinguishable 
in fruit or only by comparison. 

Flowers 5-merous, red, borne in greatly elongating spikes. 

C. Cacoucia. 

Flowers 4-merous. 

Flowers small, the stamens 2-3 mm. long C. laxum. 

Flowers large, spicate, the stamens 10 mm. long or longer. 



FLORA OF PERU 223 

Petals suborbicular; calyx abruptly short-campanulate above, 

densely red-lepidote C. assimile. 

Petals elliptic to linear. 
Calyx abruptly contracted about medially into an upper and 

lower portion C. fruticosum. 

Calyx infundibuliform or subcylindrical, obscurely if at all 
divided, that is, the upper portion rather gradually 
attenuate below. 

Petals narrowly elliptic; calyx lobes little longer, 2-3 (5) 
mm. long. 

Calyx tubular-campanulate C. Basslerianum. 

Calyx infundibuliform C. rotundifolium. 

Petals nearly linear; calyx lobes much longer, 5-6 mm. long. 

C. Llewelyni. 

Combretum assimile Eichler in Mart. Fl. Bras. 14, pt. 2: 
109. 1867. 

Branchlets, calyx tube and rachis densely red-lepidote; leaves 
ovate-elliptic, obtuse or rounded at base, obtusely acuminate, ample, 
subcoriaceous, glabrous and lustrous above, rather sparsely yellow- 
ish-red lepidote beneath; lateral nerves prominent beneath, usually 
7 or 8 and often 2 or 3 of them much closer together than the rest; 
petals orbicular-ovate, acute, shorter than the ovate calyx lobes, 
these yellow-villous within; fruit lightly reddish scurfy, nearly 4 cm. 
wide and 3.5 cm. long. F.M. Neg. 19655. Illustrated (flower), I.e. 
pi 34- 

Loreto: Near Yurimaguas, Klug 2785 (det. Standl.). Mouth of 
Santiago, Tessmann 4567; 3824. Rio Acre: Ule 9663; Krukoff 5698. 
Brazil. 

Combretum Basslerianum Mildbr. Notizbl. Bot. Gart. Berlin 
9: 143. 1924. 

Slender-stemmed liana, the petioles 3-5 mm. long, the leaves 
narrowly ovate or elliptic, obtuse at base, gradually acuminate, 6-11 
cm. long, 2.5-4.5 cm. wide, glabrous above, pale yellow lepidote but 
not densely so beneath, where the 7-10 lateral nerves are prominent; 
terminal spikes secund, dense, 6-8 cm. long, the rachis and ovaries 
densely brown-purple lepidote; receptacles narrowly campanulate, 
manifestly constricted at the insertion of the filaments, 11 mm. long 
with lobes, 7 mm. wide between them, these 3-4 mm. wide, 2 mm. 
long, within (as lobes) densely lanate; petals scarcely 2 mm. long, 



224 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1 mm. wide, lance-elliptic; filaments 2.5 cm. long, the anthers 
scarcely 2 mm. long; immature fruit densely lepidote, undulate- 
winged, 2 cm. long, half as wide. Near C. secundum (C. fruticosum) 
but receptacle longer and with an upper and lower part, i.e. not 
gradually dilated. Named for Dr. Bassler, leader of the Amazonian 
Expedition on which Tessmann made his important Peruvian col- 
lections. F.M. Neg. 23221. 

Loreto: Yarino Cocha, Tessmann 3208, type. 

Combretum Cacoucia (Baillon) Exell, Kew Bull. 469. 1931; 
122. Terminalia Cacoucia Baillon, Hist. PI. 6: 275. 1877. Cacoucia 
coccinea Aubl. PI. Guian. 1: 450. 1775, not Combretum coccinea Lam. 

Shrub or liana with broadly elliptic leaves (to 1 dm. broad), 
little lepidote and usually pentamerous linear-bracteate flowers; 
spikes sometimes even 5 dm. long, the rachis and calyx tube softly 
tomentose-puberulent, the upper part to 10 mm. long and 6-9 mm. 
wide, nearly closed by the annular margin of the pronounced disk; 
fruit 5-ridged. Poisonous seeds used for destroying rats. Illustrated, 
Mart. Fl. Bras. 14, pt. 2: pi. 32. 

Peru (probably, at least in cultivation). Brazil to Central 
America. 

Combretum fruticosum (Loefl.) Stuntz, U. S. Dept. Agric. 
Bur. PL Ind. Seeds & PI. Imp. 31: 86. 1914; 110. Gaura fruticosa 
Loefl. Iter Hispan. 248. 1758; German ed. 320. 1766; C. laxum Loefl. 
I.e. 320, in syn., 396, with description. C. secundum Jacq. Sel. Stirp. 
Amer. Hist. 1: 103. 1763; 110. C. oxypetalum G. Don, Trans. Linn. 
Soc. 15: 420. 1827. C. Loeflingii Eichler in Mart. Fl. Bras. 14, pt. 2: 
110. 1867. 

Scandent or diffuse shrub; leaves beneath and rachis and calyces 
densely yellow lepidote; leaves gradually acuminate, rather char- 
taceous, evenly nerved; upper calyx tube campanulate, 5-7 mm. long, 
the ovate lobes subequaled by or little longer than the lanceolate 
petals, at most 2 mm. long; fruit nearly orbicular, glabrate or some- 
what scurfy lepidote, often red-tinged, about 2 cm. long. The larger 
leaves are about 15 cm. long and half as wide; stamens about 3 cm. 
long. Flowers yellow and red. It is possible that the name of 
Jacquin represents a distinct plant. Some of the specimens cited 
here have fruits 3 cm. wide, 2.5 cm. high. 

Huanuco: Chicoplaya, Ruiz & Pavdn. Shapajillo, 630 meters, 
Woytkowski 39. San Martin: Tarapoto, Spruce 3969 (det. Taub.). 
Pongo de Cainarachi, Klug 2608 (det. Standl.). Chazuta, Klug 



FLORA OF PERU 225 

4118. Juanjui, King 4325. Junin: La Merced, 5318(1) (fruit). 
Loreto: Maynas, Poeppig. Cachipuerto near Balsapuerto, Klug 
3130 (det. Stand!.). Pebas, Isern 2269; 2268. Cuzco(?): Gay. Rio 
Acre: Ule 9664. Mexico to West Indies, Paraguay, and Argentina. 

Combretum laxum Jacq. Enum. PI. Carib. 19. 1760; 115. C. 
Jacquini Griseb. Fl. Brit. W. Ind. 275. 1864. C. obtusifolium Rich. 
Act. Soc. Hist. Nat. Paris 1: 108. 1792, fide Exell. C. puberum Rich. 
I.e., fide Exell. C. odoratum R. & P. ex G. Don, Trans. Linn. Soc. 
15: 430. 1827. 

Highly variable species as to foliage but marked by the small 
fragrant flowers borne in axillary or terminal panicles; leaves gla- 
brous and not lepidote but sometimes with a minute surface indu- 
ment or puncticulation, this most rarely impressed; upper part of 
calyx tube cup-like, 1-2 mm. high, glabrous or pubescent; petals 
broad, exceeding the calyx lobes; fruit oblong to suborbicular, ridged 
to winged. It is possible that with more material the species 
as interpreted by Eichler can be shown to be in reality several. 
Amazonian forms of this affinity include C. Sprucei Eichler, 115, 
completely glabrous and distinctive by the thick quadrate-oblong, 
narrowly winged fruits to about 3 cm. long; C. laurifolium Mart., 117, 
the fruit bipyramidate-tetragonous, attenuate at base, narrowly 
winged, glabrous; C. nitidum Spruce 118, the fruit glabrous, sub- 
rotund, only 7-8 mm. wide, sessile; C. phaeocarpum Mart., 118, the 
fruit sessile, about 16 mm. long, half as wide, rounded at base, rusty- 
tomentose. More distinctive, perhaps, is C. brevistylum Eichler, 118, 
the inflorescence hirsutulous with bright brown, spreading trichomes, 
the style shorter than the stamens. The Ecuadorian C. Pavonii G. 
Don has leaves softly rusty or ashy-pilose beneath, the flowers 
remote. F.M. Negs. 7888 (puberum); 7890 (obtusifolium); 23226 
(odoratum). Illustrated, flower, I.e. pi. 34. 

San Martin: Juanjui, Klug 4183. Loreto: Near Yurimaguas, 
Klug 2780 (det. Standl.). Balsapuerto, Klug 3068 (det. Standl.). 
Florida, Rio Putumayo, Klug 2237 (det. Standl.). Ucayali, Tess- 
mann 3365; 3075(1). Iquitos, Ule 6870; Tessmann 5034; 3698; Wil- 
liams 1379; Klug 1253. Soledad, Tessmann 5205; 5176. Manfinfa, 
Williams 1136. Upper Rio Nanay, Williams 1195. Rio Acre: Ule 
9665; Krukoff 5460; 5727. Argentina to Mexico and the West 
Indies. "Aioho-ayo-o" (Huitoto), "escobilla." 

Combretum Llewelyni Macbr. Field Mus. Bot. 11: 31. 1931. 
Probably scandent shrub, the branchlets, petioles and leaves 



226 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

beneath sparingly rufo-lepidote; petioles to 5 mm. long; leaves gla- 
brous above, lustrous-subchartaceous, broadly elliptic or nearly sub- 
rotund, abruptly caudate-acuminate, 8-10 cm. long, 5 cm. wide; 
lateral nerves 6-7, prominent beneath, curved at the margins and 
with the veins rather reticulate; terminal spikes about 2 dm. long; 
calyx 17 mm. long, reddish-lepidote, narrowly campanulate from 
base, scarcely or not at all constricted, the narrowly triangular lobes 
rarely 3 mm. wide at base, 5-6 mm. long, acuminate, sparsely lanate 
within; petals nearly linear, 2 mm. long; stamens about twice as long 
as the calyx; fruit unknown. Approaching C. fruticosum and C. 
Basslerianum but calyces longer, more narrowly toothed, the petals 
narrower, the leaves broader. Nearly C. rotundifolium but perhaps 
separable on the basis of the narrow petals. 

Loreto: Along the Rio Itaya, Williams 143, type. 

Combretum rotundifolium Rich. Act. Soc. Hist. Nat. Paris 
1: 108. 1792; 109. C. Aubletii DC. Prodr. 3: 19. 1828. 

With the red-lepidote calyces of C. assimile but the calyx tube 
longer and the upper part more nearly cylindrical; leaves more 
densely yellow-red lepidote; lateral nerves apparently always about 
equally remote; disk well developed with pilose border; petals 2 mm. 
long, half as wide, elliptic; fruit oblong to suborbicular, 2-3 cm. long, 
1.5-3 cm. broad, 4-winged, lepidote. Illustrated (flower), I.e. pi. 34. 
F.M. Neg. 7892. 

Loreto: Pampa del Sacramento, Castelnau (det. Exell). To 
Venezuela and Guiana. 

3. TERMINALIA L. 

Shrubs or trees, the leaves usually obovate, long attenuate to the 
petioles and characteristically clustered at the ends of the branchlets. 
Flowers small, usually spicate, petals lacking. Upper part of calyx 
tube deeply 4-5-lobed, soon falling. Stamens 8 or 10 in two series, 
the lower opposite the calyx lobes. Fruit long-ovoid, deeply angled 
or 2-5-winged. Ramatuella HBK. with the flowers and fruits in 
subglobose heads, the fruits trapezoid, rostrate, medially 4-5-winged, 
the wings rigid, may be found either R. argenlea HBK., 99, the 
leaves ashy sericeous beneath, or R. virens Spruce, 100, the leaves 
glabrous. Brandis included the genus in Terminalia but so far as 
known the fruits are distinctive. 

The Asiatic tree T. Catappa L. has been collected in cultivation 
on the lower Rio Huallaga, and at Caballo-Cocha as "Castana" and 



FLORA OF PERU 227 

"Almendra"; the leaves are much larger than those of the native 
species, attaining at least a dm. in width; it is the Indian almond or 
Tropical almond of horticulture; the ovoid angular drupe known as 
"Myrobalans" is edible, has a filbert-like flavor and yields an oil. 
Leaves mostly about 3.5 cm.; fruits about 1 cm. wide with 4-5 wings, 

2 larger than the others T. amazonica. 

Leaves often to 5 or 6 cm. wide; fruits about 3.5 cm. wide, with only 

2 wide-spreading wings T. oblonga, T. tarapotensis. 

Terminalia amazonia (J. F. Gmel.) Exell in Pulle, Fl. Surinam 
3: 173. 1935; 91. Chuncoa amazonica J. F. Gmel. in L. Syst. Nat. ed. 
13. 2: 702. 1791. Gimbernatea obovata R. & P. Prodr. 138. 1794. 
Chuncoa obovata (R. & P.) Pers. Syn. 1: 486. 1805. T. obovata (R. & 
P.) Steud. Nomencl. ed. 2. 2: 668. 1841. 

A tree, the branchlets and inflorescences as the younger leaves 
more or less reddish pilose or strigose; leaves typically obovate with 
a short, broad, very blunt point, glabrate, 6-8 cm. long, 3.5 cm. 
wide, often smaller, tapering at base to the short, slender petiole; 
flowers rather remote, about 2.5 mm. wide, the stamens well- 
exserted; fruit wings 4-5 mm. long or shorter, nearly as wide as 
the fruit, with wings 10-11 mm. wide. To 20 meters high with 
extremely hard wood much valued for construction (Ruiz & Pavon). 
F.M. Neg. 29283. 

Huanuco: Cuchero & Macora, Ruiz & Pavdn, type. San Martin: 
Tarapoto, Spruce 4909. Loreto: Balsapuerto, Klug3107. North to 
Mexico and Trinidad. "Arbol del chunchu," "chuncho." 

Terminalia oblonga (R. & P.) Eichler in Mart. Fl. Bras. 14, 
pt. 2: 90. 1867. Gimbernatia oblonga R. & P. Prodr. 138. 1794. 
Chuncoa oblonga (R. & P.) Pers. Syn. 1: 486. 1805. 

Glabrous tree or the rachis of the spikes finely strigose; leaves 
typically elliptic-oval, apically rounded or shortly and bluntly acute, 
8-9 cm. long, 5-6 cm. wide, sometimes narrower, the stout 
petioles 2-5 mm. long, the blades obscurely biglandular, moder- 
ately veiny, subchartaceous; fruits lustrous, the wings 15 mm. long, 
or longer, about as wide, ovate, rounded, the fruiting rachis typically 
smooth. To 15 meters high, or higher. Illustrated (fruit), Eichler 
I.e. pi. 33. F.M. Neg. 29282. 

Huanuco: Pozuzo, Ruiz & Pavon, type. San Martin: Tarapoto, 
Spruce 4587 (det. Eichler). Rio Acre: Krukoff 5641. 

Terminalia tarapotensis Heurck & M. Arg. Obs. Bot. 213. 
1871. 



228 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branchlet tips and rachi reddish strigose pubescent; petioles to 
1 cm. long; leaves rather few, broadly obovate, shortly and obtusely 
acuminate or acute, acute at base, often 10-12 cm. long, 4-8 cm. wide, 
glabrous at maturity, distinctly veined, somewhat lustrous, eglan- 
dular but with some pellucid dots near the margin, subcoriaceous; 
disk shortly hispid, the villous style and glabrous stamens well- 
exserted, equally long. Doubtfully distinct fromT. oblonga, known in 
fruit. The Tessmann fruiting specimen has fruits 4 cm. wide, 2.5 cm. 
long. In all probability there is only one species. F.M. Neg. 23231. 

San Martin: Tarapoto, Spruce 4570, type. Loreto: Pongo Man- 
seriche, Tessmann 3573; probably also 4014- 

4. BUCHENAVIA Eichler 

With the facies in general of Terminalia. Branchlets often obvi- 
ously enlarged at the tips, the leaves frequently biglandular at base. 
Flowers hermaphrodite and male, the two sorts irregularly mixed on 
the same spike, this sometimes subcapitate. Calyx tube above 
broadly cup-like, subentire or somewhat toothed. Fruit a more or 
less fleshy drupe, pointed, angled. Genus apparently chiefly rep- 
resented in the Amazon Valley; cf. Ducke, Archiv. Jard. Bot. Rio 
Jan. 4: 148-151. 1925 and Archiv. Inst. Biol. Veg. Rio Jan. 2: 63-65. 
1935, for additions to the eight species known to Eichler. 

Leaves sparsely red-villous beneath with long trichomes, pellucid- 
punctate B. punctata. 

Leaves glabrous beneath at least on surface or sparsely puberulent, 
not obviously punctate. 

Leaves about 1.5 dm. long; inflorescence pubescence brownish- 
gray B. macrophylla. 

Leaves much smaller; inflorescence pubescence reddish. 

B. oxycarpa. 

Buchenavia macrophylla [Spruce] Eichler in Mart. Fl. Bras. 
14, pt. 2: 98. 1867. 

Branchlets nodose; petioles to about 1 cm. long; leaves oblong- 
obovate, acute at the rounded tip, biglandular at the shortly atten- 
uate base, slightly lustrous both sides, paler beneath, membranous, 
the venation slender; spikes subsessile, more or less ashy or brownish 
tomentose; ovary neck and calyx glabrous; stamens all exserted an 
equal distance but barely so, the short style scarcely exceeding the 
villous disk. A 7-meter tree, 6 cm. in diameter (Tessmann). Her- 
barium name by Spruce under Terminalia. F.M. Neg. 21768. 



FLORA OF PERU 229 

San Martin: Tarapoto, Spruce 2507, type. Loreto: Soledad, 
Tessmann 5259 (vel aff., Mildbraed). 

Buchenavia oxycarpa [Mart.] Eichler, I.e. 97. 

Tree, glabrous except for the rusty pubescent rachi and sometimes 
also for the ovary, this tomentulose, and the calyces, these pubescent; 
branchlets slender, often verticillate, as the crowded leaves; petioles 
4-12 mm. long, plane above; leaves obovate to oblanceolate, rounded 
or obtusely acuminate, cuneate or attenuate at the somewhat glan- 
dular base, subcoriaceous, rather dull, distinctly venose both sides, 
5-10 cm. long, 2.5-3.5 cm. wide; flowers spicate, 4-6 mm. broad, 
glabrous except the villous disk, the style scarcely exserted from 
this, the alternate stamens included; drupe oblong, acuminate, 
5-angled, shining, under 2.5 cm. long. To 30 meters high, some 
20 meters to the first branch (Tessmann). Name by Martius in 
herbaria under Terminalia. Illustrated, Eichler I.e. pi. 25. 

Loreto: Soledad, Tessmann 5165. Brazil to British Guiana. 

Buchenavia punctata [Spruce] Eichler, I.e. 98. 

More densely pubescent than the somewhat similar B. macro- 
phylla, the branchlets much enlarged at tips and crowned by fascicles 
of obovate leaves, these brown-pilose on the petioles and nerves; 
leaves acute or obtuse, long attenuate to the eglandular petiole, 
opaque, rather firm, 7-10 cm. long, 2.5 cm. wide or wider, the slender 
reticulate venation prominent beneath; drupes about 1.5 cm. thick. 
Name in herbaria by Spruce under Terminalia. F.M. Neg. 14335. 

San Martin: Tarapoto, Spruce 4.945, type. 

LECYTHIDACEAE 

Knuth, Pflanzenreich IV. 219a. 1939. 

Usually tall trees, the leaves often large, ovate or oblong, entire 
or crenulate-serrate, alternate. Stipules none. Flowers sometimes 
solitary, more often racemose or paniculate, the inflorescence axillary, 
terminal or borne directly on the branches. Flowers hermaphrodite. 
Calyx 4- or 6-lobed, the lower part connate with the ovary into a 
receptacle, the lobes imbricate. Petals free, rarely none, 6, 4 or 
sometimes 8, often large. Androphorum colored, frequently extended 
laterally into a long more or less valericuliform appendage. Stamens 
many, frequently borne separately and modified into staminodia. 
Ovary usually 2-6-celled, the ovules 1 to several in each cell. Style 
obsolete or short. Fruit characteristically a pyxidium, the lower 
portion often more or less enlarged by the remains of the sepals, 



230 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the upper provided with an operculum that is often connate with 
the central column and not rarely woody. 

Family noteworthy as the source of Brazil nuts, Bertholletia 
excelsa, native to the Amazon, but scarcely appearing within Peru 
unless cultivated. A number of species, particularly those of 
Gustavia, furnish wood highly prized for construction. 
Androphorum semiglobose; column of the fruit dehiscing with the 
mesocarp; seeds funiculate. 

Anthers linear; petals 6-8 1. Gustavia. 

Anthers short; petals 4 2. Grias. 

Androphorum manifestly bilateral. 

Androphorum urceolate-campanulate, oblique but without tongue; 

operculum joined with the woody column 3. Cariniana. 

Androphorum laterally with an elongate galericulate appendage. 
Processes of the appendage and annulus anther-bearing and 

similar 4. Couroupita. 

Processes of the appendage more or less destitute of anthers, 

those of the annulus much larger. 
Seeds long funiculate; style rather long; fruit 4-celled. 

5. Lecythis. 
Seeds sessile or nearly; style short 6. Eschweilera. 

Androphorum laterally with spirally incurved appendage. 

7. Couratari. 

1. GUSTAVIA L. 

Trees or tall shrubs with ample, crowded, entire or more or less 
serrate, often spathulate leaves and axillary or terminal flowers 
sometimes corymbosely congested, their pedicels medially, basally 
or rarely apically bracteolate. Calyx obovate, the limb 4-6-lobed 
or sometimes nearly truncate. Petals 6-8, about equal. Andro- 
phorum nearly globose, with numerous multiseriate processes, often 
dilated, attached to the inner face, the anthers linear, basifixed, 
finally opening apically by a pore. Ovary inferior, 6- or occasionally 
4-celled. Fruit globose, with 6 seeds in each cell, suspended especially 
from the upper part of the placenta column by long, thick funiculae. 
Species as recognized, very finely drawn, the key, accordingly, only 
suggestive. Genus named in honor of the Swedish king Gustav III. 

Calyx lobes developed, broadly triangular. 
Petals oblong-obovate, 10-15 mm. broad. 

Petals 8, 3.5 cm. long G. iquitosensis. 



FLORA OF PERU 231 

Petals 6 (7), 2-2.5 cm. long G. hexapetala. 

Petals broadly obovate, finally 3 cm. wide or wider. .G. brasiliana. 

Calyx lobes obsolete or depressed; if obvious only 1-2 mm. high, 
the calyx limb often annulate, thick. 

Petals about 8, 2.5 (-3) cm. long, only 10-12 mm. wide. 

Pedicels about 2 cm. long; leaves slightly puberulent. 

G. longifolia. 

Pedicels 3-4 cm. long; leaves glabrous G. mangua. 

Petals 4 (3.5) -6 cm. long, broadly obovate except G. Tessmannii. 
Leaves glabrous. 

Leaves acute at base or decurrent on the more or less developed 
petiole. 

Petals about 6 cm. long. 

Leaves more or less serrulate . .G. augusta. 

Leaves subentire G. Mexiana. 

Petals 3.5-4.5 (5) cm. long. 
Petals broadly obovate. 

Petals about 10, 4-5 cm. long; pedicels 2-3 cm. long. 

G. calycaris. 

Petals about 8, 3.5-4 cm. long; pedicels 2-6 cm. long. 

G. speciosa. 

Petals oblong-spatulate, 1.5 cm. broad. . . .G. Tessmannii. 
Leaves narrowed to a truncate or retuse base. 

G. Poeppigiana. 

Leaves puberulent beneath G. pubescens. 

Gustavia augusta L. Amoen. Acad. 8: 266. pi. 5. 1775; 16. 

Branchlets at their densely leafy tips 5-6 mm. thick; leaves 
oblanceolate, 17 cm. long, 5.5 cm. broad, often much larger, narrowed 
at each end for about a quarter of their length, the base wing-decur- 
rent into a petiole 1 cm. long, the tip acutish, the margin above 
the middle more or less serrulate, firm papyraceous, dull green, 
glabrous, the nerves 1-2 cm. distant; inflorescence terminal, 4- 
flowered, the pedicels 2-4 cm. long, bibracteolate above the middle; 
calyx undulately lobed; petals to 6.5 cm. long, 3.5 cm. broad, white, 
the androphorum 4 cm. across, the anthers 3 mm. long; fruit globose, 
5.5 cm. thick, turbinate at base, truncate; seeds 2.5 cm. long, 1.5 cm. 
broad. In writing augusta (not angusta as often quoted) Linn 



232 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

thoughtfully completed the name for this beautiful plant in honor 
of his king, Gustav III. Illustrated, Knuth, I.e. 17. fig. 6. 

Loreto: Pampa del Sacramento, Castelnau (det. Benoist). Brazil 
to the Guianas. 

Gustavia brasiliana DC. Prodr. 3: 290. 1828; 19. 

Branchlets at the foliose tips 5 mm. thick; petioles obsolete or 
scarcely 1 cm. long, winged; leaves oblong-lanceolate or broader, to 
23 cm. long, 8 cm. broad, contracted toward the acute tip and 
cuneately narrowed about a third from the base, entire or slightly 
undulate margined, firm papyraceous, greenish, glabrous, the nerves 
14-17 mm. distant; inflorescence terminal, 4-5-flowered, the rather 
stout pruinose tomentulose pedicels bibracteolate near the middle; 
calyx lobes 6, broadly triangular, acute, coriaceous; petals 6, white, 
obovate, to (4) 6 cm. long, finally 3-4 cm. broad, retusely rounded; 
androphorum 2.5-3 cm. broad; fruit narrowly 6-winged, 4 cm. 
across. Known to attain 7 meters. G. microcarpa Pilger, I.e. 20, 
probably extending into eastern Peru from Brazil, has the pedicels 
bibracteolate below the tip. Illustrated, Mart. Fl. Bras. 14, pt. 
I: pi. 7. 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5695 (det. Knuth). 
Brazil. 

Gustavia calycaris (Berg) Miers, Trans. Linn. Soc. 30: 185. 
1874; 20. G. augusta L. var. calycaris Berg in Mart. Fl. Bras. 14, 
pt. 1: 471. 1858. 

A shrub-tree 5-6 meters high, the young branchlets brownish- 
gray, rugulose; petioles 8-12 mm. long; leaves oblong-lanceolate, 
2.5-3.5 dm. long, 5-9 cm. broad, constricted at apex into an acute 
acumen, gradually narrowed to base, nearly entire or undulate- 
sinuate, rigid-chartaceous, green and opaque above, slightly paler 
beneath; racemes axillary, 6.5 cm. long, the fragrant roseate flowers 
as many as 10, 9 cm. broad, borne on pedicels 2-3 cm. long, that bear 
2 linear acute bractlets about 7 mm. long a little below their tips; 
calyx limb subquadrate, the broad lobes rotate, as much as 2.25 mm. 
high, pruinose both sides; petals about 10, strongly imbricate, ovate- 
oblong, fleshy, 4-5 cm. long, 2 cm. broad, grayish opaque and pul- 
verulent both sides; androphorum white; ovary turbinate, brown- 
tomentose, 4-celled, the style short, conical. 

San Martin: San Roque, Williams 6549. Loreto: La Victoria, 
Williams 3165. Caballo-Cocha, Williams 2502. Brazil. 



FLORA OF PERU 233 

Gustavia hexapetala (Aubl.) Smith in Rees, Cycl. 17: no. 2. 
1811; 19. Pirigara hexapetala Aubl. PI. Guian. 1: 490. pi. 193. 1775. 

Branchlet-tips slender, 4 mm. thick; leaves mostly 12-14 cm. 
long, 5-6 cm. broad, suddenly contracted to the acute or obtuse 
apex, gradually narrowed from the upper third to the decurrently 
winged petiole, this 1-2 cm. long, entire or serrate-undulate, papy- 
raceous, glaucous-green, the nerves 12-15 mm. remote; inflorescence 
terminal, shortly cymose, 1-4-flowered, the peduncle lacking; 
pedicels to 2.5 cm. long, 2 mm. thick, with 2 broad ovate bractlets 
at the middle; calyx strongly 6-costate, the acutish triangular lobes 
5-6 mm. long, 5 mm. wide at base; petals 2-2.5 cm. long, 10-12 mm. 
broad; androphorum 1.5-2 cm. broad; fruit globose, 6-winged, the 
wings more or less crisped. Said to attain 8 meters. Illustrated, 
Bot. Mag. pi. 5239. 

Rio Acre: Ule 9653. Brazil to the Guianas. 

Gustavia iquitosensis Knuth, Repert. Sp. Nov. 35: 338. 1934; 
24. 

Slender tree with oblong-spatulate leaves, the larger 4-5 dm. 
long, 13-17 cm. broad, obtuse, narrowed to the base from within 
one-fifth of the tip, the base itself subretuse, the margin entire or 
somewhat undulate, papyraceous, glabrous, the nerves nearly 2 to 
2.5 cm. distant; flowers solitary, the 8 spatulate petals 3.5 cm. long, 
12 mm. broad, 6-7 mm. broad near the base, rounded at apex; 
calyx lobes 4, more or less triquetrous, 6 mm. long and broad; 
androphorum yellowish, 3 cm. in diameter, the processes carmine; 
anthers 2.5 mm. long, 1-1.25 mm. broad. The petioles and receptacle 
somewhat tinged with red, the petals pale at base, deep carmine 
above. The type was 2 meters high, not branched, the trunk 
1.5 cm. thick. G. Ulei Pilger, of adjacent Brazil, has glaucous leaves, 
flowers about 3, subsessile, the petals 6, white or rose as the andro- 
phorum. 

Loreto: Iquitos, Tessmann 3645, type. 

Gustavia longifolia Poeppig ex Berg, Linnaea 27: 442. 1854; 24. 

A nearly simple or scarcely branching tree 2 meters high with 
spatulate leaves 3-6 dm. long, 12.5-20 cm. broad at the middle, 
obtusely acute, 2-3.5 cm. broad at the base, coarsely serrulate, firm 
papyraceous, ashy-green above, minutely puberulent beneath, sub- 
sessile; inflorescence 1-4-flowered, borne on the older stems or from 
the lower leaf -axils; pedicels puberulent, 17-24 mm. long, with 
2 minute ovate bractlets below the middle; calyx nearly entire or 



234 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3-4-lobed, the lobes 5 mm. broad, 1.5 mm. high; petals 8, obversely 
oblong, to 2.5 cm. long, 12 mm. broad, greenish-purple; androphorum 
12 mm. long; receptacle 8 mm. long, puberulent, 4-celled, the cells 
many-ovuled; fruit globose, 5 cm. broad, red, with 3-4 seeds. 
F.M. Neg. 31635. 

Loreto: Yurimaguas, Poeppig 2094, type. Lower Rio Huallaga, 
Williams 3926. Balsapuerto, Kittip & Smith 28564- 

Gustavia mangua Macbr. Field Mus. Bot. 11: 29. 1931; 20. 

Petioles 5-10 mm. long; leaves oblong-lanceolate, 3-4 dm. long, 
6-8 cm. broad, narrowly produced at apex into an acute acumen, 
gradually narrowed toward the acute base, subentire or remotely 
crenulate-serrulate, crisply plicate, rigid chartaceous, glabrous, the 
slender straw-colored nerves prominent beneath; flowers many, in 
racemes 2.5 cm. long, bracts numerous, ovate-acute, 3 mm. long; 
pedicels slender, minutely pulverulent, bibracteolate a little below 
the middle, 3-3.5 cm. long; calyx limb entire, obsoletely 4-lobed; 
petals about 8, oblong-obovate, nearly 2.5 cm. long, 7-10 mm. broad, 
pulverulent. 

Loreto: Lower Rio Nanay, Williams 281, type. Mouth of 
Santiago, Mexia 6228a (det. Standl.). "Sacha manga," "mangua." 

Gustavia Mexiana Knuth, Pflanzenreich IV. 219a: 136. 1939. 

A tree 6 meters high, the ultimate branchlets 6 mm. thick, 
glabrous; leaves often more or less verticillately congested, lanceolate, 
to 16 cm. long, 4 cm. broad, gradually narrowed to the acute apex, 
narrowly cuneate at base where passing into the very narrowly 
winged petiole 1 cm. long, firm papyraceous, the margin minutely 
denticulate or subentire; lateral nerves 12-14, slender and with the 
midrib prominent; calyx limb annulate, petals to 6 cm. long, 2.5-3 
cm. broad, rounded, glabrous, whitish; androphorum 4 cm. broad; 
receptacle 1.5 cm. long, 1 cm. broad, turboid. The tough wood 
is used for tool handles and hoops (Mexia). 

Loreto: Iquitos, Mexia 6517, type. "ChopeY' 

Gustavia Poeppigiana Berg, Linnaea 27: 442. 1854; 25. G. 
caballoensis Macbr. Field Mus. Bot. 11: 28. 1931, fide Knuth. 

Young branchlets 7 mm. thick; leaves sessile, oblanceolate, about 
33 cm. long, 11 cm. broad, more or less abruptly contracted at the 
acutish apex, narrowed from the upper third gradually to the retuse 
base, coarsely denticulate-serrate, papyraceous, green, glaucous, 
glabrous, the nerves 2 cm. distant; inflorescence (as to type) terminal, 



FLORA OF PERU 235 

about 3-flowered, the peduncle obsolete, the pedicels to 1.5 cm. long, 
with 1 broad, caducous bract near the base; calyx lobes obsolete, 
the limb annulate; petals 6-8, obovate, somewhat retuse, 4 cm. long, 
2 cm. broad, apparently white; androphorum 3 cm. broad. In 
G. caballoensis the flowers are axillary, solitary, but the character 
may not be significant. Berg recognized 3 forms, their names 
descriptive of the leaf -variation: var. opaca; var. nitens; var. 
rigida. G. conferta (Berg) Miers, Amazonian, is similar but the 
leaves are only 6-7.5 cm. broad, gradually acuminate, and the 
pedicels 2-2.5 cm. long, bibracteolate above the middle. F.M. 
Neg. 31637. 

Loreto: Yurimaguas, Ule 6302; Killip & Smith 28908. Upper 
Rio Itaya, Williams 3406; 3519. Iquitos, King 350. Caballo- 
Cocha, Williams 2152; 2240 (type, G. caballoensis). Brazil. "Chope." 

Gustavia pubescens R. & P. Fl. Peruv. 4: 306. pi. 351. 1802; 24. 

Leaves elongate-oblanceolate, to 8 dm. long, 18 cm. broad, con- 
tracted at the nearly mucronate apex, gradually narrowed from the 
upper third to the more or less rounded base, the margin serrate- 
crenulate or crenulate, papyraceous, glabrous above, softly tomentose 
beneath but at maturity only on the nerves, net-veined, the lateral 
nerves 13-17 mm. distant; inflorescence unknown; petals broadly 
obovate, about 6 cm. long, 3.5 cm. broad; androphorum 5 cm. broad, 
the process 18 mm. long. The type is from Guayaquil. Cf. G. 
Ruiziana, mentioned under G. speciosa. 

Loreto: Lower Rio Huallaga, Williams 3941. Ecuador. "Mem- 
brillo de monte." 

Gustavia speciosa (HBK.) DC. Prodr. 3: 289. 1828; 22. Piri- 
gara speciosa HBK. Nov. Gen. & Sp. 7: 262. 1825. 

A tree, the ultimate branchlet about 7 mm. thick, glabrous 
except for the more or less evanescently tomentulose pedicels and 
calyces; petioles short or to 3 cm. long, 2.25 mm. thick; leaves elon- 
gate-oblanceolate, usually 3-4.5 dm. long, 8.5 cm. broad, acutish, 
gradually narrowed and cuneate into the petiole, entire or remotely 
serrulate, firm or chartaceo-papyraceous, glaucous-green, the nerves 
1.5 cm. distant; inflorescence sessile or nearly, contracted-racemose, 
3-10-flowered, the pedicels 2-6 cm. long, to 3.5 mm. thick, medially 
bracteolate and with 1 deciduous bract at base; calyx subglobose, 
7-8 mm. broad, 3-6 mm. long; petals rotund-obovate, 3.5 cm. long, 
white within, pale rose without; androphorum 3.5 mm. broad, the 
processes yellow. G. Ruiziana Berg, the type from Guayaquil, has 



236 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves densely puberulent beneath, petals to 6 cm. long. F.M. Negs. 
38305; 29401 (Ruiziana). 

San Martin: Tarapoto, Williams 5646; King 3900. Loreto: 
Middle Ucayali, Tessmann 8410. Iquitos, Tessmann 3676; and sev- 
eral other collections. Brazil; Colombia. "Chupa." 

Gustavia Tessmannii Knuth, Repert. Sp. Nov. 35: 338. 
1934; 28. 

A nearly simple tree 1.5-6 meters high, the young branchlets 
1.5-2 cm. thick; petioles stout, often 5 cm. long, 7 mm. across; 
leaves spatulate, to 7.5 dm. long and 22 cm. broad, acutish or obtus- 
ish, from about the upper fourth narrowed to the base into the 
petiole, the margin entire or scarcely undulate, firm papyraceous, 
green, glabrous, the nerves 2-3 cm. remote; inflorescence short, 2-3- 
flowered, borne on the older wood, the pedicels 5-10 cm. long, their 
medial bractlets 2 mm. high; calyx limb annulate; petals 8, spatulate- 
oblong, 4.5 cm. long, 1.5 cm. broad, 6-8 mm. wide at base, white, 
roseate toward the rounded apex; androphorum 3 cm. broad, the 
processes 1.5 cm. long; anthers 2.5 mm. long, 1-1.25 mm. broad. 
G. Duckei Knuth, similar, has serrulate leaves, the inflorescence 
rachis 7 cm. long, with many bracts; G. Spruceana has leaves 10-13 
cm. broad, pedicels 3 cm. long; G. laciniosa Miers is marked by the 
laciniately serrate leaves, nearly those of G. superba (HBK.) Berg, 
but this with puberulent pedicels all species to be expected. 

Loreto: Mouth of Santiago, Tessmann 4574, type. Puerto Melen- 
dez, Tessmann 4728. Umbria, dist. Putumayo, Klug 1880. 

2. GRIASL. 

Tall trees with large leaves crowded together at the tips of the 
branches and shortly pediceled or subsessile flowers borne in close 
corymbs on the old wood. Technically similar to Gustavia but the 
petals always 4, spreading, fleshy, and the small anthers with a sub- 
apical short cleft. 

The fleshy part of the fruit (mesocarp) of G. peruviana and other 
species is said to be eaten, a common name being "membrillo." 
Pedicels slender, 3-4 cm. long, 1.5-2 mm. thick. 

Leaves entire; rachis 1-2 cm. long G. Tessmannii. 

Leaves undulate-serrate; rachis 6-12 cm. long. .G. maranonensis. 
Pedicels shorter, stout. 

Rachis of inflorescence 6-12 cm. long; leaves entire. 

G. Neuberthii. 



FLORA OF PERU 237 

Rachis 1-1.5 cm. long. 

Leaves subentire G. grandifolia. 

Leaves remotely undulate-serrate G. peruviana. 

Grias grandifolia Pilger, Notizbl. Bot. Gart. Berlin 9: 142. 
1924; 31. 

A tree 10-15 meters high, the trunk to 6 dm. thick; leaves sessile, 
lanceolate linear, to 12 dm. long or longer and 2 dm. broad, narrowed 
toward the acute apex and gradually toward the base, entire, gla- 
brous; flowers 6-8, congested on the trunk, to 6 cm. long, the rachis 
1 cm. long; bractlets triangular or broader, acutish or rotund, 4-6 
mm. long, nearly as wide; pedicels 7-17 mm. long, 2 mm. thick; 
receptacle glabrous, 7 mm. long; calyx limb 4-lobed, the lobes finally 
broadly triangular or often none; petals rather broadly ovate, 23 
mm. long, glabrous, pale yellow; processes about 10-14 mm. long; 
anthers rotund, 1 cm. long. The Colombian G. Haughtii Knuth has 
only about 3 flowers, the peduncle obsolete. Illustrated (fruit), 
Knuth, I.e. 6. F.M. Neg. 29165. 

Loreto: Masisea, Tessmann 3110, type. Puerto Limon, Tess- 
mann 3864- Iquitos, Williams 1355. 

Grias maranonensis Knuth, Repert. Sp. Nov. 35: 339. 1934; 30. 

A small tree 4 meters high; leaves narrowly lanceolate, often 8-10 
dm. long, 24 cm. broad, more or less contracted at the acute tip, 
narrowed at base to a petiole often 9 cm. long, 6-8 mm. across, gla- 
brous, the margin remotely undulate-serrulate, the nerves to 4 cm. 
distant, slender but prominent beneath; flowers as many as 15, more 
or less racemose, the inflorescence to 14 cm. long; bractlets to 8 mm. 
long, very acute; pedicels 3-4 cm. long, 1.5-2 mm. thick; calyx limb 
entire or obscurely 4-lobed; petals 2-2.5 cm. long, glabrous, ochro- 
leucous; processes 12-14 mm. long; anthers 1 mm. broad. Allied to 
G. Tessmannii but the petioles much stouter, the inflorescence longer. 

Loreto: Perseveranza, Maranon (Kuhlmann 21501, type). 

Grias Neuberthii Macbr. Field Mus. Bot. 11: 30. 1931; 31. 
G. loretensis Knuth, Repert. Sp. Nov. 35: 339. 1934. 

A tree 13-15 meters high; leaves narrowly lanceolate with the 
petiole often 9 dm. long, 17 cm. broad, rather abruptly contracted 
at the acute apex, gradually narrowed into the slightly winged 
petiole, this to 2 dm. long; leaf blades lightly revolute, entire, gla- 
brous, firm-papyraceous, the nerves often 3 cm. distant, slender, 
distinctly marked below, the veins little conspicuous; flowers about 



238 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5, more or less racemose, the inflorescence to 16 cm. long, the rachis 
sericeous; bractlets to 13 mm. long, acute, chartaceous; pedicels not 
distinct from the receptacle and with it 13 mm. long, 2.5 mm. across 
at base, stout, slenderly sericeous as the calyx, the persistent lobes 
of the latter ovate obtuse, 10 mm. long, 7 mm. broad, thin; petals 
unequal, often 28 mm. long, 15 mm. broad, chartaceous, glabrous, 
yellow; processes 10-12 mm. long; anthers 1 mm. broad. Pulp 
surrounding the seeds is edible (Mexia). Name in memory of Carl 
Neuberth, long custodian of the herbarium, Field Museum. Illus- 
trated, Knuth, I.e. 32. 

Loreto: Caballo-Cocha, Williams 2337, type. Iquitos, Killip & 
Smith 27272, type, G. loretensis; King 252; 680. Mouth of Santiago, 
Mexia 6115 (det. Standl.). "Chope," "mango sacha." 

Grias peruviana Miers, Trans. Linn. Soc. 30: 301. 1874; 31. 

Tree 7-12 meters high, with linear-lanceolate leaves and inflor- 
escences of 1-4 flowers; leaves sessile or decurrent to a broadly winged 
petiole to 8 cm. long, the blades often 6 or 7 dm. long or longer and 
1.5 dm. broad, abruptly contracted to the acuminate tip, the acute 
acumen 1.5-2 cm. long; leaf -margin coarsely and remotely undulate- 
serrate; nerves 1.5 cm. distant; bractlets triangular, 4-6 mm. long; 
pedicels to 2 cm. long, 2 mm. thick; receptacle glabrous, the calyx 
limb often 2-lobed, the lobes becoming 9 mm. long; petals 15 mm. 
long, 10 mm. broad, obtuse; style short. 

San Martin: Spruce 454-1- Rio de la Ventana, Spruce. 

Grias Tessmannii Knuth, Repert. Sp. Nov. 35: 340. 1934; 30. 

Tree 10 meters high, the trunk 15 cm. in diameter; leaves nar- 
rowly lanceolate, often 5 dm. long and 8-10 cm. broad, narrowed to 
the acute apex and to the petioles, these 3-9 cm. long, 2.25 mm. 
across, margin entire, nerves 1.75-2 cm. distant, slender but promi- 
nent beneath; flowers usually 3-4 together, the rachis itself 1-2 cm. 
long; bracts minute, mucronate; pedicels 3.5-4 cm. long, 1.5 mm. 
thick; receptacle 4 mm. long; calyx limb entire or irregularly 4-lobed; 
petals ovate, 2 cm. long, deep yellow; processes 10-12 mm. long. 

Loreto: Mouth of Santiago, Tessmann 4187, type. Rio Huallaga, 
Williams 4181. The lower Yurimaguas, Kittip & Smith 27868. On 
the Amazon, Williams 1609. Ecuador. 

3. CARINIANA Casar. 

Tall-growing trees with small racemose or paniculate flowers. 
Calyx limb 6-lobed. Petals 6. Androphorum urceolate-campanu- 



FLORA OF PERU 239 

late, somewhat oblique but without tongue, the processes all stami- 
niferous. Ovary 3-celled. Fruit woody, the operculum early joined 
to the placentae, finally opening. Seeds with one long lateral wing. 

Gariniana domestica (Mart.) Miers, Trans. Linn. Soc. 30: 286. 
pi. 63. 1874; 36. Couratari domestica Mart. Flora 20: Beibl. 2: 127. 
1837. 

Young branchlets about 2.5 mm. thick, tinged deep purple; 
leaves oblong-elliptic to about 1 dm. long, half as wide, entire or 
undulate margined, barely acute at base, shortly and acutely acumi- 
nate, the 20 pairs of slender nerves subequally prominent on both 
sides; flowers in a thyrse to 11 cm. long, nearly as broad; calyx lobes 
ovate, acute, 2.5 mm. long, the ovate whitish petals twice as long; 
androphorum 4 mm. broad, reddish-yellow; fruit 6 cm. long, obconic, 
3 cm. wide above. Tree to 60 meters high. Illustrated, Mart. 
Fl. Bras. 14, pt. 1. pi. 80. 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5597. Brazil. 

Cariniana estrellensis (Raddi) Kuntze, Rev. Gen. 3, pt. 2: 89. 
1898; 39. Couratari estrellensis Raddi, Mem. Soc. Ital. Modena 18: 
403. 1820. C. excelsa Casar. Nov. Stirp. Bras. 46. 1842, fide Knuth. 

Young branchlets 2.5-3 mm. thick; petioles 7-13 mm. long; 
leaves to 8 cm. long, 5 cm. broad, more or less acutely or obtusely 
acuminate, decurrent into the petiole, serrate-crenulate, with about 
12 pairs of nerves prominent on both sides; spikes mostly solitary or 
binate in the leaf -axils, rarely congested into a thyrse; pedicels 3-5 
mm. long; calyx lobes scarcely 2 mm. high; petals obovate, 7 mm. 
long, yellow; androphorum 2 mm. broad; fruit 5 cm. long, 2.5 cm. 
broad, the operculum 5 mm. high, plane above; seed without wing 
10 mm. long, 8 mm. broad, the wing 3.5 cm. long. A large tree 10-30 
meters high. Illustrated, Mart. Fl. Bras. 14, pt. 1. pi. 79; Knuth, 
I.e. 3 (androphorum), 6 (fruit). 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5568. Brazil; 
Bolivia; Paraguay. "Jequitiba branca." 

4. COUROUPITA Aubl. 

Trees with oblong-cuneate entire or inconspicuously crenate- 
serrate leaves and racemes of large flowers, generally rather terminal, 
rarely from the old wood. Bracts deciduous. Calyx limb 6-lobed. 
Petals 6, somewhat unequal. Androphorum annuliform with a 
lateral tongue-like appendage that is turned over, the processes of 



240 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

both the basal and reversed portions antheriferous. Ovary inferior, 
6-celled; style obsolete, the stigma 6-radiate. Fruit globose, the 
operculum not dehiscing. Seeds long-funiculate. The "Cannon 
Ball Tree," C. guianensis Aubl., to which the Peruvian specimen of 
C. amazonica was at one time referred, may be cultivated; it has 
leaves 1.5-3 dm. long on glabrous petioles, large showy red flowers 
in panicles from the older branches, and globose fruits as large as 
cannon-balls which, when the hard shell is broken, are extremely ill- 
scented as is the wood; the flowers and fruit are borne more or less 
continuously. 

Petals 2 cm. long or shorter; pedicels short C. amazonica. 

Petals 3-5 cm. long. 

Pedicels short, only the lowest rarely 6 mm. long or longer. 

C. subsessilis. 

Pedicels, at least the lower, 2 cm. long or longer. 
Leaves more or less long-cuneate to base, barbate in axils. 

C. peruviana. 

Leaves rounded to the obliquely obtuse base, obscurely if at all 
barbellate C. acreensis. 

Couroupita acreensis Knuth, Pflanzenreich IV. 219a: 46. 1939. 

Tree about 30 meters high; petioles 10-13 mm. long, ciliate on 
the margins; leaves obovate-oblong, 9 cm. long, 4-4.5 cm. broad, 
very obtuse, gradually narrowed, the upper third to the obliquely 
obtuse base, entire, chartaceo-papyraceous, glabrous; raceme 25 cm. 
long, divided below, the rachis 6 mm. thick, glabrous; pedicels 2 cm. 
long, deciduously bracteate at base; calyx lobes suborbicular, purple, 
glabrous, finely ciliolate, 4 mm. long and broad; petals 4-4.5 cm. 
long, 3-3.5 cm. broad, annulus of androphorum 17-19 mm. broad, 
the upper portion 27 mm. broad, the processes 13-14 mm. long. 

Rio Acre: Basin of Rio Purus, Krukoff 5677, type. 

Couroupita amazonica Knuth, Repert. Sp. Nov. 35: 340. 
1934; 45. 

Branchlets puberulent (at tip) as also the petioles, at least in the 
Peruvian plant; petioles 2-2.5 cm. long; leaves elongate-oblanceo- 
late, mostly about 14 cm. long, 4 cm. wide, obtuse or nearly, long- 
attenuate to the acute base, papyraceous, glabrous or somewhat 
barbellate in the nerve-axils, these 16 or more on each side of the 
thick midrib; racemes 10-45 cm. long; pedicels persisting, to 3 mm. 
long; receptacle rugose; calyx lobes only 2-2.5 mm. long and broad; 



FLORA OF PERU 241 

petals 13-20 mm. long, 7-9 mm. broad, ovate-oblong; androphorum 
with tongue 2.5 cm. long; annulus 1 cm. broad. The Peruvian plant 
apparently has somewhat narrower petals but probably belongs here. 
Wide-branching tree 18 meters high; flowers borne on trunks of large 
branches; fruit resembles cannon-ball (Mexia). 

Loreto: Iquitos, Mexia 6502. Brazil. "Haya-uma," "castanha 
de macaco." 

Couroupita peruviana Berg, Linnaea 31: 261. 1862; 47. 

A tree 30 meters high with rusty red branchlets foliose at their 
tips; petioles sparsely puberulent, 2-2.5 cm. long; leaves oblong, 
10-13.5 cm. long, 4.5-5 cm. broad, obtusely and shortly acuminate, 
gradually narrowed to the obtuse base from about the middle, 
obsoletely serrulate, thin-chartaceous; racemes axillary, many- 
flowered, the rachis 4-6 mm. thick, 20-25 cm. long; pedicels 2-2.5 cm. 
long, the small basal bracts ciliate, also with 2 apical bractlets; 
flowers very fragrant, the broadly oblong petals pale flesh-colored 
without and yellowish-punctulate, red within, 4.5 cm. long, 2.75 cm. 
broad; calyx lobes suborbicular, puberulent, the thin margins ciliate; 
androphorum 9 mm. broad, white, the tongue reddish; fruit nearly 
globose, 18 cm. broad, the upper part and operculum small. 
C. surinamensis Mart, is similar but the leaves are decurrent into 
the petioles and sparsely barbate in the axils beneath. Flowers red, 
white and yellow (Klug). 

San Martin: Tarapoto, Spruce 4495, type. Juanjui, Klug 4357 
(det. Standl.). Loreto: Puerto Melendez, Tessmann 4777. Balsa- 
puerto, Klug 2835. "Aya Uma" (i.e. "Totenkopf," fide Tessmann). 

Couroupita subsessilis Pilger, Verhandl. Bot. Ver. Brandenb. 
47: 163. 1905; 45. 

A tall, glabrous tree or the leaves sparsely barbate in the axils 
beneath; branchlets often 1.5 cm. thick, densely marked by the 
leaf scars, the leaves crowded on them apically and these narrowly 
obovate-oblong, 2 dm. long, 8.5 cm. broad, obtuse, gradually cuneate 
from the upper portion into the narrow petiole (2 cm. long, 2 mm. 
thick), firm papyraceous to chartaceous, the nerves 5-7 mm. distant, 
stout and prominent on both sides; racemes terminal, to 7 dm. long, 
the rachis 8 mm. thick; calyx lobes rotund-ovate, 6 mm. long; petals 
3-5 cm. long, 2-2.5 cm. broad, whitish; androphorum yellow, the 
processes roseate-purple; fruit 6 cm. broad. C. amazonica Knuth 
has membranous leaves and petals 13-20 mm. long; C. elata A. C. 
Smith has calyx lobes 3-4 mm. long but Ducke, fide Knuth, thinks 



242 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

it is the same as C. subsessilis. The Indian name, fide Tessmann, 
means "Totenkopf," a skull, from the form and stench of the fruit 
(broken). F.M. Neg. 29157. 

Loreto : Mouth of Santiago, Tessmann 4827. Brazil. "Aya Uma." 

5. LECYTHIS Loefl. 

Tall trees with nearly the character of Eschweilera, which could 
be included as a subgenus but the ovary is 4-celled, the style rather 
elongate, the ovules many in each cell and, especially, the seeds 
long-funiculate in a very large fruit, the top of which is united 
firmly to the woody column of the placentae. The name is doubt- 
fully correct (cf. Knuth) and, as the monographer suggests, should 
be conserved. 

It is possible that one or more species may occur within the 
boundaries of Peru, especially in the region adjacent to the territory 
of the Rio Acre, but the range of the genus is toward the Atlantic 
and to Central America. For Lecythis subbiflora R. & P. see 
Eschweilera. 

Lecythis hians A. C. Smith, Phytol. 1: 123. 1935; 62. 

A glabrous tree known to attain 30 meters; branchlets densely 
lenticellate; petioles 3-7 mm. long, 1.5-1.75 mm. thick; leaves 
oblong, 7-10 cm. long, 3-4 cm. wide, acutely acuminate, obtuse or 
rounded at base, serrulate, papyraceous, the nerves 7 mm. distant; 
racemes mostly corymbose-congested, 5-12 cm. long; rachis rugose, 

2 mm. thick; flowers subsessile, the deltoid calyx lobes 4-5 mm. long, 

3 mm. broad; petals rounded-oblong, 15-17 mm. long, 12-14 mm. 
wide; androphorum 15 mm. broad; processes of annulus scarcely 
1 mm. long; style fleshy, 1 mm. long; fruit subglobose, 14 cm. long, 
16 cm. broad, rugose above calycaric zone, upper zone entire, 11-13 
cm. in diameter. 

Peru (perhaps). Amazonian Brazil. 

6. ESCHWEILERA Mart. 

Chytroma Miers, Trans. Linn. Soc. 30: 164, 229. pi. 34b. 1874. 

Medium to large trees with coriaceous entire to serrate leaves 
and rather showy flowers in axillary or terminal racemes or panicles. 
Calyx lobes and petals 6, the latter often somewhat unequal. Andro- 
phorum pelviform or annuliform, the processes of the extended 
tongue-like portion all or most of them not antheriferous; anthers 
finally opening by a longitudinal chink. Ovary 2-4-celled, the ovules 



FLORA OF PERU 243 

few, sessile in each cell. Style nearly obsolete to rather short. 
Fruit coriaceous, the top finally opening, the placentae deciduous. 
Seeds few, efuniculate or nearly. 

Lecythis subbiflora R. & P. Fl. Peruv. 4: pi. 461. 1802 (G. Don, 
Gen. Syst. 2: 873. 1832) known as "Camaron" is the same as (and 
the earlier name for) E. integrifolia (R. & P.) Knuth, 97; Ecuadorian 
rather than Peruvian, it is omitted here, especially as it suggests 
Colombian species, for instance, E. bogotensis Knuth, 95. 

The fruit of most species is unknown. "Matamata" is the native 
name for the species which furnish hard wood; the inner bark is the 
source of a good fiber. 

Flowers sessile or the pedicels only a few mm. long. 
Leaves 4-8.5 cm. wide. 

Leaves small, to about 1 dm. long; inflorescence glabrous. 

Petioles 5 mm. long; calyx lobes fimbriate E. Klugii. 

Petioles 6-10 mm. long. 

Leaves acute E. iquitosensis. 

Leaves rounded at tip E. ucayalensis. 

Leaves medium to large, mostly or all of them longer than 1 dm. 
or only to 13 cm. in E. Tessmannii with tomentulose inflores- 
cence. 
Leaves broadly cuneate at base; panicles tomentulose. 

E. Tessmannii. 
Leaves rounded at base; inflorescence glabrous. 

Leaves caudate with acumen to 1 cm. long. . .E. itayensis. 
Leaves merely subcuspidate or acute. 

Leaves subcoriaceous; rachis stout E. juruensis. 

Leaves papyraceous; rachis slender E. loretensis. 

Leaves 13-18 cm. wide. 

Petals 2 cm. long; racemes 3 dm. long. .E. Mexiana, E. Knuthii. 

Petals to 4.5 cm. long; racemes 1 dm. long E. gigantea. 

Flowers pedicellate, the pedicels about 10 mm. long. 
Leaves about 5 cm. wide. 

Leaves obovate, rounded at tip '. ?;;' J V ;'!*. . .E. ucayalensis. 

Leaves oblong, pointed at tip .'. '. . !'.' E. timbuchensis. 

Leaves about 10 cm. wide E. fractiflexa. 

Eschweilera fractiflexa Knuth, Pflanzenreich IV. 219a: 110. 
1939. 



244 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles stout, 1.5 cm. long; leaves oblong, to 23 cm. long, 10 cm. 
broad, contracted toward the tips, acute, rounded at base, entire, 
chartaceous-coriaceous or coriaceous, drying brown-olive, glabrous 
or very finely puberulent below on the thick midnerve; veins barely 
manifest below; racemes 10-16 cm. long, panicled, about 15-flowered, 
the somewhat purplish olive rachis 4-5 mm. thick at base, zigzag; 
pedicels about 12 mm. long, 1.5 mm. thick, the base persisting; recep- 
tacles shortly crateriform; lobes broadly ovate, obtuse or nearly 
rounded, 4-4.5 mm. long; petals to 3 cm. long, 2 cm. broad, yellow- 
ish white; androphorum extended, 4 cm. long, annulate portion 
1 cm. broad; anthers subrotund, 1 mm. long; processes of the 
tongue sterile, 6 mm. long; style minute, ovary 2-celled. Tree 12 
meters high. 

Loreto: Mishuyacu, King 353, type. 

Eschweilera gigantea (Knuth) Macbr. Candollea 8: 25. 1940. 
Lecythis gigantea Knuth, Repert. Sp. Nov. 38: 113. 1935. Chytroma 
giganlea Knuth, Pflanzenreich IV. 219a: 85. 1939. 

Tree to 40 meters high with a trunk diameter of 7 dm.; petioles 
woody, to 5 mm. thick, purple tinged as the branchlets; leaves 
oblong, 5 dm. long, 18 cm. broad, rounded but with a slender mucro 
about 5 mm. long, rounded at base, entire or remotely undulate- 
dentate, the teeth large, glabrous, the stronger nerves 1.5-2 cm. 
distant; racemes about 1 dm. long, 14-18-flowered; rachis often 8 mm. 
thick, strongly angled; bracts minute, broadly triangular; flowers 
subsessile, the crateriform receptacle 7 mm. long, the lobes 6, 
rounded-ovate, entire, 14 mm. long, 12 mm. broad; petals various 
in form and size, sometimes 4.5 cm. long, 3 cm. broad, more or less 
erose-undulate, pale yellow; extended androphorum 6 cm. long, the 
processes of the annulae about 5 mm. long, brownish-yellow, those 
of the tongue deep-yellow and 10 mm. long or longer. 

Loreto: Mouth of Santiago, Tessmann 4064, type. 

Eschweilera iquitosensis Knuth, Pflanzenreich IV. 219a: 
111. 1939. 

A tree 15-20 meters high with a trunk diameter of 68 cm.; 
branches squarrose, gray, the younger deep purple; petioles 6-9 mm. 
long, 2 mm. thick; leaves to 7 (10) cm. long, 4 (5) cm. broad, more 
or less abruptly contracted to an acute tip, more or less rounded at 
base, the lightly revolute margin entire, drying chartaceous and 
olive-brown; nerves and veins slender but conspicuous beneath; 
inflorescence paniculate, about 7 cm. broad, the numerous flowers 



FLORA OF PERU 245 

sessile; receptacle 1.5 cm. long; calyx lobes 5 mm. long, ovate, ob- 
tusish, glabrous; petals broad, white, 15-20 mm. long; androphorum 
expanded, 2.5-3 cm. long, the annulus 1 cm. broad, the processes and 
minute anthers yellow, those of the tongue pale yellow. 

Loreto: Iquitos, Tessmann 3606, type; Klug 1180. Lower Itaya, 
Tessmann 5208. Brazil. "Matamata." 

Eschweilera itayensis Knuth, Pflanzenreich IV. 219a: 111. 
1939. 

A tree, the ultimate branches slender; petioles 1 cm. long, 2 mm. 
thick, dark purple; leaves oblong-ovate, 16 cm. long, 8.5 cm. broad, 
rather rounded-acuminate, the acumen itself 1 cm. long, nearly 
rotund at base, entire, drying papyraceous-chartaceous and olive- 
green, the moderately stout nerves 1 cm. distant, the veins very 
conspicuous beneath; panicle about 8 cm. long and broad, the gla- 
brous branchlets deep purple; pedicels persistent, 1 mm. long, 
finally enlarged; receptacle glabrous, 6 mm. long; calyx lobes broadly 
ovate, lacerate-fimbriate, glabrous; petals 1.75 cm. long and broad; 
androphorum extended, 2 cm. long, the sterile galericulate processes 
3 mm. long; annulus 10 mm. broad, the processes and anthers minute. 

Loreto: Upper Rio Itaya, Williams 3260, type. 

Eschweilera juruensis Knuth, Pflanzenreich IV. 219a: 111. 
1939. 

Tree known to attain 40 meters, the ultimate branchlets 4-5 
mm. thick, brown, glabrous, squarrose; petioles 1.5 cm. long, deep 
brown, glabrous, somewhat winged, 2-3 mm. broad; leaves oblong, 
to 22 cm. long, 8.5 cm. broad, often smaller, merely cuspidate at 
the suddenly contracted apex, rounded at base, entire or incon- 
spicuously undulate, drying chartaceo-coriaceous or chartaceous and 
brown, the nerves 7-10 mm. distant, pronounced above but scarcely 
prominent beneath, the intermediary nerves many, also prominent 
above, the veins beneath slender; racemes axillary, 10-13 cm. long, 
about 15-flowered, the stout rachis 4-6 mm. thick, lightly zigzag, 
glabrous, often sparsely branched but the branches very short; 
flowers subsessile; bracts glabrous, 8 mm. long, ovate; receptacle 
stout, 7 mm. long, half as broad or larger; sepals semi-ovate, obtuse, 
glabrous. Petals and androphorum undeveloped. 

Rio Acre: Near mouth of Rio Macauhan, Krukoff 571+1; 5755, 
type. Brazil. 



246 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Eschweilera Klugii Knuth, Pflanzenreich IV. 219a: 111. 1939. 

An 18 meter tree, the ultimate dark gray branchlets 2-3 mm. 
thick; petioles 5 mm. long, 1.5 mm. thick, brown; leaves lanceolate- 
oblong, to 10 cm. long and 4 cm. broad, often narrower, abruptly 
contracted to an acute tip, basally broadly cuneate, drying char- 
taceous, green above, brownish-green beneath, entire, the nerves 
rather irregularly 6-12 mm. distant; racemes 5 cm. long, the many 
flowers 2-6 mm. distant, panicled; pedicels persisting, finally enlarged, 
1 mm. long; calyx lobes broadly ovate, lacerate-fimbriate, glabrous; 
petals broad, to 15 mm. long, yellow; expanded androphorum 15 mm. 
long, the sterile processes of the tongue 3 mm. long, the fertile ones 
of the annulus, this 8 mm. broad, minute, as also the style. 

Loreto: Mishuyacu near Iquitos, Klug 597, type; 386. 

Eschweilera Knuthii Macbr. Candollea 8: 25. 1940; 90. 
Chytroma Tessmannii Knuth, Repert. Sp. Nov. 38: 113. 1935, not 
E. Tessmannii Knuth, Pflanzenreich IV. 219a: 115. 1939. 

A tree known to attain 10 meters, the trunk 9 cm. thick, branch- 
ing at 6 meters, the bark of the branches gray; petioles 1-2 cm. long, 
3-3.5 mm. thick; leaves lanceolate-oblong to 4.5 dm. long and 13 cm. 
broad, abruptly contracted toward the tip and coarsely mucronate, 
cuneate or rounded (the larger leaves) at base but not decurrent 
into the petiole, entire, drying firm papyraceous and deep olivaceous, 
glabrous, the nerves prominent beneath as the veins; racemes simple, 
3 dm. long, about 20-flowered, corymbed, the rachis olive-brown, 

3 mm. thick at base, the nodes 3 mm. long; pedicels obsolete; re- 
ceptacle broadly crateriform, coriaceous; calyx lobes semicircular, 
very obtuse, 4 mm. long; petals about 2 cm. long, 13 mm. broad, 
deep carmine; androphorum carmine to flesh color, 3.5-4 cm. long, 
extended, the processes of the annulus 1-2 mm. long, the anthers 
minute, those of the tongue sterile, broadly linear, 7 mm. long. 
Perhaps the same as E. Mexiana but petioles shorter, leaf veins 
prominent, calyx lobes smaller; E. andina (Rusby) Macbr., comb, 
nov., Lecythis andina Rusby, Mem. Torrey Club 6: 37. 1896, is even 
less clearly distinct. 

Loreto: Mouth of Santiago, in flood-free wood, Tessmann 3935, 
type. 

Eschweilera loretensis Knuth, Pflanzenreich IV. 219a: 112. 
1939. 

Tree 10 meters high, the branchlets ashy brown, rather short, 

4 mm. thick; petioles 1-1.5 cm. long, 2-2.5 mm. broad, glabrous; 



FLORA OF PERU 247 

leaves oblong, usually 2 dm. long and 8 cm. wide, obtuse, rounded 
at base but the petiole more or less winged by the extension of the 
leaves, entire, papyraceous and green, glabrous, the lateral nerves 
about 20, with the veins moderately prominent beneath; racemes 
1-1.5 dm. long, panicled, few, the rachis 2 mm. thick, glabrous, 
armed by the rudiments of the pedicels, these about 1 cm. remote, 
persisting, 2 mm. long, 1.5 mm. thick; pedicels 2 mm. long, subsessile; 
sepals rotund-cordate, 5 mm. broad, 4 mm. long, glabrous, entire; 
petals red, rotund-obovate, 2 cm. long, glabrous; androphorum 
extended, 4.5-5 cm. long, the tongue fleshy, 12-14 mm. broad; 
galericulum 2 cm. wide; anthers of the annulus ovoid; style conoid, 
3 mm. long. 

Loreto: Cachipuerto on the Rio Cachiyaco between Balsapuerto 
and Moyobamba, Klug 3121, type. 

Eschweilera Mexiana (Knuth) Macbr. Candollea 8: 25. 1940. 
Chytroma Mexiana Knuth, Pflanzenreich IV. 219a: 136. 1939. 

Said to be a tree attaining 18 meters, the flowers white but only 
the fruit described, this 5-6 cm. high and broad, the part below the 
operculum turboid, 3 cm. high, the calyculate zone to within 15 mm. 
of the operculum, 5 cm. broad, the persisting sepals subligneous, 
triquetrous, 12 mm. high, 15 mm. broad; interzonal vittae 14 mm. 
high, concave below, convex above; superior zone 5.5 cm. in diameter, 
lightly undulate; operculum conoid, more or less cusped apically, 
5.5 cm. in diameter, more or less radiate, thick woody as the 
pyxidium; cells (in the fruit) 4. In specimen seen the branchlet is 
7 mm. thick, the attached petiole 5 mm. thick, 3.5 cm. long, the 
leaf nearly oblong, 4.5 dm. long, 18 cm. wide, rounded and apiculate 
at apex, broadly rounded or subtruncate at base, chartaceo-coriace- 
ous, nerves mostly about 1.5 cm. distant, rather prominent beneath, 
scarcely apparent above, the veins obscure even beneath; flowers 
detached and broken but seemingly sessile, the petals apparently 
3 cm. long, half as wide, the 6 sepals glabrous, suborbicular, 1 cm. 
long; androphorum 3 cm. long, the annulus about 1.5 cm. broad; 
style short. Hardwood used for beams (Mexia). 

Loreto: Pongo de Manseriche on the right shore of the Rio San- 
tiago, 220 meters, in thick wood, Mexia 6243, type. "Hua capu." 

Eschweilera Tessmannii Knuth, Pflanzenreich IV. 219a: 115. 
1939. 

Known to attain 25 meters with a trunk diameter of 7 dm., the 
ultimate branches densely marked with dark minute lenticels; petioles 



248 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1 cm. long, dark purple; leaves often 3-10 cm. distant, oblong, to 
13 cm. long, 5.5 cm. broad, abruptly contracted to the acute tip, 
broadly cuneate at base, drying chartaceous, gray above, pale brown 
beneath, entire; nerves slender and scarcely obvious above, the 
netted venation slender beneath; inflorescence paniculate, to 2 dm. 
long, the branchlets many, brown- tomentulose, abundantly flowered ; 
true pedicels 1-1.5 mm. long, finally persisting and enlarging to 1.5 
mm. thick; receptacle minutely tomentose, 7 mm. long; calyx lobes 
ovate or ovate-oblong, rounded interiorly, minutely tomentose; 
petals pale rose, whitish at base, broad, 1.5 cm. long; androphorum 
2.5 cm. long, the annulus 1 cm. broad, the brownish anthers minute. 
Loreto: Mouth of Santiago, Tessmann 4691, type. Mishuyacu, 
King 156; 258. Upper Rio Nanay, Williams 3180. 

Eschweilera timbuchensis Knuth, Pflanzenreich IV. 219a: 

115. 1939. 

A tall tree, the ultimate branchlets 2.5-3 mm. thick; petioles 
stout, 2 mm. thick, to 9 mm. long; leaves oblong, usually 13 cm. long, 
5 cm. broad, rather suddenly contracted to an acute tip, nearly rotund 
at base, entire or lightly undulate-margined, the midnerve prominent, 
the nerves irregularly so and 7-15 mm. distant, the venation con- 
spicuous; pedicels 1 cm. long; fruit (perhaps not mature) 4.5 cm. 
in diameter, 5 cm. long, obtuse. Apparently similar to, if indeed 
distinct from E. Krukovii A. C. Smith, 112, of adjacent Brazil. 

Loreto: Timbuchi, on the Upper Rio Nanay, Williams 979, type. 

Eschweilera ucayalensis Knuth, Pflanzenreich IV. 219a: 

116. 1939. 

Trunk of the type with a diameter of 8 dm.; petioles to 1 cm. 
long, 2 mm. thick, more or less brown-colored; leaves obovate, 
to 9.5 cm. long, 5.5 cm. broad, rounded or nearly retuse, broadly 
cuneate at base, entire or obscurely crenate, drying pale brown and 
firm papyraceous, glabrous, the nerves slender but prominent, 8 mm. 
distant; calyx lobes broadly ovate, 4 mm. long; petals pale yellow, 
very broad, 15-20 mm. long; androphorum about 3 cm. long; 
galericulum pale, the annulus deep yellow, the latter 1 cm. broad, 
the processes, anthers and style minute. 

Loreto: Yarina-cocha, on the Middle Ucayali, Tessmann 3218, 
type. 

7. COURATARI Aublet 

Trees in many respects similar to Eschweilera but the lateral 
extension of the androphorum spirally incurved back toward or 



FLORA OF PERU 249 

over the antheriferous processes. Ovary 3-celled, the ovules many 
in each cell. Fruit oblong-cylindrical, finally opening. Seeds few, 
thin, wing-margined. 

Couratari macrosperma A. C. Smith, Bull. Torrey Club 60: 
383. 1933; 131. 

Ultimate branchlets thick; petioles stout, to 1 cm. long; leaves 
oval-oblong or oblong, to 18 cm. long, 8 cm. broad, obtuse, broadly 
cuneate at base, lightly undulate-margined, drying olive-green, the 
lateral nerves about 25; racemes axillary and terminal, 1-2 dm. 
long, nearly simple; rachis stout, densely brown-stellate pubescent, 
15-20-flowered; pedicels thick, densely pubescent, 5-8 mm. long, 
articulate below the middle and minutely 3-bracteolate; calyx lobes 
oblong-rotund, obtuse, to 8 mm. long and broad, pubescent without, 
the margins shortly ciliate; petals white, obovate, puberulent with- 
out, about 2 cm. long and broad; androphorum extended, 3 cm. long, 
the tongue 10 mm. broad; fertile processes 1 mm. long, the oblong 
anthers as long; receptacle turbinate, pubescent, the style stout; 
fruit without the operculum 18 cm. long, gradually narrowed to 
the calycarate base, this portion 16 cm. above the true base, 8.5 cm. 
across; interzonal vitta erect, 2 cm. high, the upper zone 8 cm. broad. 
Ducke, according to Knuth, regards this species as highly variable 
in character of fruit, and includes 2 other species here. Illustrated 
(androphorum), Knuth, I.e. 3. 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5638; 5687. Brazil. 

MELASTOMACEAE 

References: Cogniaux, DC. Monogr. Phan. 7: 1891; Gleason, 
Brittonia 1: 127-184. 1932 and Carnegie Inst. Wash. Publ. 522, 
No. 20: 323-373. 1940. 

The characteristics of this well-known family are given in detail 
in the classical monograph cited, pages 7-8; there too is a key to 
the tribes and the genera which it has not seemed necessary to copy 
here for it would be in any case incomplete. On the other hand, there 
follows a key which perhaps accounts for the Peruvian species. 
Attention may be called to the genera Mouriria and Alloneuron, 
which depart from the familiar family character of several leaf 
nerves from the base to apex of the leaves in having pinnately nerved 
leaves; there are, however, species of Miconia (shrubs) as well as 
an herb with only one conspicuous nerve (midnerve). Unfortunately, 



250 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the first division of the key has to rest on the character of the fruit, 
but often this is evident. 

The generic lines are probably drawn in trivial fashion in many 
cases; in Leandra one finds axillary and terminal (becoming lateral) 
inflorescences and yet elsewhere in the family one must be content 
to consider the position of the inflorescence as of prime importance, 
one or two "genera" having nothing more to recommend them. 
Even the development of the appendages of the anthers has been 
used in delineating genera in an exaggerated way sometimes, as in 
the tribe Bertolonieae, while quite as much variation elsewhere has 
received only sectional (if that) recognition. Apropos are the 
words of Cogniaux himself, I.e. 4: "The delimitation of genera is 
arbitrary enough; one could easily distinguish more of them, at the 
same time one could justify the reunion of many of those which are 
generally admitted today. I believe as does M. Baillon who conserves 
only 54, that one admits generally too many; however, in order not 
to disturb too much the nomenclature ... I have united only those 
which it appeared to me completely irrational to leave separated." 
(The italics are mine.) 

It is with no little diffidence that I, with my imperfect knowledge 
of the family, have ventured in two or three instances to unite 
groups accepted as distinct since the appearance of the Cogniaux 
monograph, but in these cases I have merely followed Baillon's 
suggestions; actually, it is my own feeling that, by and large, his 
proposed classification is sound, serving, that is, the purpose of 
practical taxonomy real usefulness and at the same time presenta- 
tion of the probable evolution of the groups and their apparent 
present relationship. However, compare a noteworthy article by 
Epling, Amer. Nat. 72: 547-561. 1938, as well as the thought-pro- 
voking symposium entitled "The Concept of the Genus," Bull. 
Torrey Club 67: 349-389. 1940. 

Finally, there is more than a hint in the remarks of Cogniaux that 
he viewed Baillon's work with sympathy, but was bound by tradi- 
tion. Of course between certain groups as those surrounding 
Clidemia there is nothing to be contributed, probably, by changes 
in status because the lines of demarcation are arbitrary, in order 
to serve the historical interest as well as for convenience; on the 
other hand there are a number of smaller groups that merit study 
with a view to their union for the purpose of contributing to a 
more practical and uniform or logical classification within the family. 



FLORA OF PERU 251 

With thanks I acknowledge my indebtedness to the publications 
of Gleason, notably in the Flora of Surinam, and in his synopsis 
of the family for British Guiana; besides, I have copied largely from 
his careful and discriminating work as cited. For convenience the 
well-known monograph of Cogniaux has been cited as "Cogn. 
Melast." 

The following key is sometimes only suggestive and sometimes 
applies only to the Peruvian species. 

Fruit a pod (possibly baccate in one herbaceous genus) ; ovary usually 
free from the calyx. 

Woody plants, usually shrubs, trees, or vines. 
Pubescence on stems or branches, usually also in the inflores- 
cence, glandular. 

Calyx teeth one-half the length of the tube or longer; capsules 

short. 

Flowers 5-merous; seeds oblong-ovoid. . .2. Rhynchanthera. 
Flowers 4-merous. 

Leaves about 2 cm. long; seeds pyramidate. 

3. Centradeniastrum. 

Leaves 3 cm. long or longer; seeds cochleate. 

4. Ernestia. 

Calyx teeth much shorter than the tube; capsules 1-2 cm. 
long 5. Arthrostemma. 

Pubescence not glandular or scarcely so, rarely the calyx; 
plants sometimes glutinous-punctate or resinous. 

Connective of at least some of the anthers obviously produced 

below the cells. 

Leaves small, about 1 cm. long or shorter; seeds oblong- 
ovoid 1. Microlicia. 

Leaves usually much larger; seeds cochleate or angled 

(unknown in Sandemania). 
Calyx lobes prominent (sometimes caducous); seeds 

cochleate. 

Flowers tubular-campanulate, the petals strongly 
incurved 8. Brachyotum. 

Flowers open, the petals spreading. 

Calyx setose, with branched or stellate trichomes 
between the lobes 12. Pterolepis. 



252 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx not stellate setulose. 

Petals obtuse or retuse; ovary usually setose at 

tip; flowers usually in terminal panicles. 
Anther connective appendaged anteriorly. 
Stamens subequal; stems not inordinately 

spreading-villous 13. Tibouchina. 

Stamens very unequal; stems inordinately 

villous 7. Desmoscelis. 

Anther connective bilobed or gibbous poster- 
iorly only 9. Macairea. 

Petals acute; ovary glabrous; flowers paniculate. 

10. Sandemania. 
Calyx lobes minute or less than one-half as long as the 

tube; seeds angled except in Aciotis, cochleate. 
Calyx limb not a circumscissile cone. 

Calyx tube globose or ovoid 15. Aciotis. 

Calyx tube oblong-campanulate . . .21. Graff enrieda. 

Calyx limb a closed cone, circumscissile at anthesis. 

22. Calyptrella. 

Anther connective not obviously produced. 
Leaves with 3-9 longitudinal nerves. 
Calyx limb not a circumscissile cone. 

Plants more or less pubescent with long hairs. 

Flowers tubular-campanulate, the petals strongly 

incurved ; seeds cochleate 8. Brachyotum. 

Flowers open, the petals spreading. 
Flowers 4-merous; seeds cochleate. 

Calyx oblong-campanulate. . .16. Monochaetum. 

Calyx globose or ovoid 15. Aciotis. 

Flowers 5-merous; seeds linear or angled or 

spindle-shaped. 

Scandent or sarmentose plants; connective with 
a bidentate erect appendage. 

18. Adelobotrys. 
Erect shrubs or trees. 

Petals 1 cm. long or longer. 
Appendages not inflated or tumid. 

19. Meriania. 



FLORA OF PERU 253 

Appendages much inflated or tumid. 

20. Axinaea. 

Petals shorter than 1 cm.. . .21. Graff enrieda. 
Plants glabrous or the pubescence short (puberulent 
or furfuraceous), never scandent (cf. the scandent 
Adelobotrys)', seeds cuneate-angled or spindle- 
shaped. 

Petals less than 1 cm. long. 
Calyx truncate, very minutely toothed. 

24. Tessmannianthus. 

Calyx lobed or irregularly ruptured. 
Seeds winged; appendage filiform . . 17. Huberia. 

Seeds not winged ; appendage short. 

21. Graffenrieda. 
Petals 1 cm. long or longer. 
Appendages not inflated or tumid ... 19. Meriania. 
Appendages much inflated or tumid . 20. Axinaea. 
Calyx limb a closed cone circumscissile at anthesis. 

Flowers small, to 6 mm. long 22. Calyptrella. 

Flowers large, 1 cm. long or longer 23. Centronia. 

Leaves pinnately nerved 25. Alloneuron. 

Herbs, or half-shrubs, woody only toward base, erect or sprawling 

(Salpinga rarely suffrutescent). 
Calyx and capsule terete, not strongly ribbed-angled or winged; 

seeds cochleate or reniform (in Catocoryne unknown). 
Calyx lobes prominent at least about one- third the length of 
the tube. 

Plants silky villous, stout; calyx hairs simple. 

7. Desmoscelis. 
Plants slender, not villous. 
Plants more or less pubescent. 

Calyx hairs in part branched or stellate. 

12. Pterolepis. 
Calyx hairs if present, simple. 

Filiform plant 35. Catocoryne. 

Slender but robust plants. 

Half-shrub; hairs on leaves above partly adnate 
to leaf -surface . . .13. Tibouchina. 



254 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Annual herb; hairs if present, not adnate. 

14. Acisanthera. 

Plants glabrous 6. Nepsera. 

Calyx lobes minute or at least much shorter than the tube. 

Calyx tube oblong-campanulate, 1-2 cm. long. 

5. Arthrostemma. 
Calyx tube ovoid or globose, small 15. Aciotis. 

Calyx or capsule strongly ribbed-angled or winged; seeds ovoid- 
angled, except in Pterogastra cochleate. 

Leaves 1-7-nerved; stamen-connective not spurred, some- 
times tubercled. 

Leaves 1-5-nerved; connective smooth or tubercled. 

Calyx tube winged, ciliate-echinate 11. Pterogastra. 

Calyx tube 8-10-costate, smooth 26. Macrocentrum. 

Leaves 7-nerved; stamen-connective not spurred. 

-27. Salpinga. 

Leaves 3-7-plinerved; stamen-connective 1-3-spurred. 
Leaves 3-5-plinerved; plants stemless; connective 1-spurred. 

28. Monolena. 
Leaves 5-7-plinerved; plants caulescent. 

Stamen-connective 3-spurred 29. Triolena. 

Stamen-connective 2-spurred 30. Diolena. 

Fruit baccate, berry-like or hard; ovary usually adherent; shrubs or 
trees; seeds ovoid-angled or subrotund. 

Leaves usually with 3-9 longitudinal nerves (exceptions are in 
Miconia) ; seeds not subrotund, many. 

Leaves not striolate between the primary nerves (unless rarely 
in Miconia). 

Inflorescence characteristically terminal. 

Petals acute or acuminate, narrow or if broad very acum- 
inate; anther-connective not or little prolonged, often 
raised ; 31. Leandra. 

Petals obtuse or rarely acute in Miconia but then the anther- 
connective spurred or appendaged. 

Leaves not vesicular; anther-connective not elevated. 

Calyx limb a closed cone circumscissile at anthesis. 

32. Conostegia. 



FLORA OF PERU 255 

Calyx limb not calyptriform or at least not circum- 

scissile (closed in some species of Miconia). 
Stamens minutely if at all appendaged; filaments 

often narrow. 
Calyx lobes unappendaged or minutely. 

33. Miconia. 
Calyx lobes with conspicuous outer teeth. 

36. Heterotrichum. 

Stamens conspicuously appendaged; filaments broad. 

34. Icaria. 

Leaves often vesicular at base; anther-connective raised. 

37. Tococa. 

Inflorescence lateral or axillary. 
Petals obtuse. 

Leaves vesicular at base. 

Anther cells convoluting 38. Maieta. 

Anther cells not convoluting. 

Anthers straight or nearly 39. Clidemia. 

Anthers, at least somewhat, curving apically toward 

flower-center 37. Tococa. 

Leaves not vesicular. 

Anthers narrow or long rostrate, 1-pored; flowers small. 
Flowers borne with the leaves (or rarely below them, 

C. septuplinervid) 39. Clidemia. 

Flowers in small clusters, usually borne below the 

existing leaves 41. Henriettea. 

Anthers oblong, usually short and thick, 1-2 pored; 

flowers usually large 40. Belinda. 

Petals acute or acuminate. 
Flowers borne below the current leaves, clustered or 

solitary 41. Henriettea. 

Flowers axillary (at least some inflorescences subtended 

by leaves) solitary or densely glomerate, cymose or 

paniculate (Leandra rarely might be sought here). 

Calyx limb regularly toothed or truncate. .42. Ossaea. 

Calyx limb irregularly lobed, closed or dehiscing above 

at anthesis 43. Myriospora. 

Leaves finely striolate between the primary nerves (cf. one or 
two Miconias) ; flowers bracted 44. Blakea. 



256 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves pinnately nerved but the lateral nerves often scarcely if at 
all obvious; seeds subrotund, few 45. Mouriria. 

1. MICROLICIA D. Don 

Small often slenderly branched shrubs, usually somewhat eri- 
caceous in general appearance. Leaves frequently shorter than 10 
mm. and wanting toward the base of the branches or stems, some- 
times more or less imbricated. Flowers small, 4-5-merous. Stamens 
unequal; anthers all perfect, rostrate and with connective long-pro- 
duced below. Ovary 3-celled; the capsule apically dehiscent. 
Sometimes, as in some other genera, the connective of larger anthers 
may be extended upward as an appendage. 

Besides the following, M. peruviana Cogn. in Mart. Fl. Bras. 14, 
pt. 3: 83. 1883; 52, might be sought here because of its name although 
it is known only from Cochabamba, Bolivia, and more likely ranges 
into Brazil than into Peru; it has glabrate branches, entire setulose- 
margined leaves 3-4 mm. wide, persisting below. Similarly the 
broom-like M. arenariaefolia [Mart. & Schr.] DC., 44, with awl- 
shaped leaves, reported by Cogniaux from "Cochabamba, Peru," is 
known only from Bolivia and Brazil. 
Leaves (some of them) setose-denticulate; flowers 5-merous. 

M. Weddellii. 

Leaves all entire, the revolute margins eciliate; flowers 4-merous. 

M. sphagnicola. 

Microlicia sphagnicola Gleason, Bull. Torrey Club 58: 215. 
1931. 

Shrubby, about 1 meter high, freely branched, the younger 
branches densely glandular with short stout trichomes and setose 
at the nodes; leaves narrowly elliptic, 2 mm. wide, punctate both 
sides; calyx tube obconic, glandular, with 4 setae; sepals ovate- 
lanceolate, 2 mm. long, ending in an awn 0.7 mm. long; petals 6-7 
mm. long; glandular-margined connective of outer larger anthers 
long-produced, truncate, those of the smaller shortly produced, 
obtuse. Characteristic plant of type-locality (collectors). 

Junin: Enefias, Pichis Trail, in open sphagnum bog or swamp, 
1,700 meters, Killip & Smith 25680, type. 

Microlicia Weddellii Naud. Ann. Sci. Nat. se>. 3. 12: 251. 
1849; 40. 

Low shrub with somewhat flexuose gray-barked stems (the 
younger setose at nodes) that are leafless below; leaves oblong-elliptic, 



FLORA OF PERU 257 

5-7 mm. long, 2-2.5 mm. wide, glabrous but some sparsely long 
setulose-denticulate; calyx tube glandular-punctate, 4 mm. long, 
the shorter subulate lobes long-setose apically; petals roseate, 4-7 
mm. long; anthers ovoid, very shortly beaked. M. paraensis Ducke, 
Archiv. Jard. Bot. Rio Jan. 3: 225. 1922, is marked by the long 
pilose nodes, somewhat larger leaves and flowers, the anthers subu- 
late, beaked. F.M. Neg. 36105. 

Puno: Prov. of Carabaya, now Sandia, Weddell, type. Mountains 
of Yuncacoya, (Weberbauer, 241). 

2. RHYNCHANTHERA DC. 

Shrubby or sometimes herbaceous, usually glandular-pubescent 
plants with 5-9-nerved ovate leaves, paniculate, rather showy, 
5-merous purple flowers and glandular calyces with narrowly subu- 
late lobes that are nearly as long or longer than the tube. Stamens 
very unequal, 5 sterile, the anthers filiform, the 5 larger fertile ones 
with rostrate anthers, their slender-produced connective usually 
tuberculate or bidenticulate at its insertion on the filament. 
Calyx teeth filiform, longer than the tube, glandular-setose. 

R. Haenkeana. 

Calyx teeth shorter than the tube, somewhat hirsute or villous- 
glandular R. Hookeri. 

Rhynchanthera Haenkeana DC. Prodr. 3: 107. 1828; 104. 
R. Matthaei Naud. Ann. Sci. Nat. se>. 3. 12: 209. 1849. 

Calyx teeth glandular-setulose, linear-filiform, 5-7 mm. long, 
longer than the tube; petals to 15 mm. long; branchlets hispidulous 
with gland-tipped hairs of unequal length; petioles mostly 5 mm. 
long; leaves glandular-ciliolate, scattered-hispidulous both sides, 
5 (-7) -nerved, oblong-ovate, acuminate, rather rounded or acute 
at base; style about 15 mm. long; capsule globose, 4 mm. in diameter. 
One of the fertile stamens is larger. The filiform calyx teeth are 
the conspicuous feature of this species, as they are for R. grandiflora 
(Aubl.) DC., 101, distinguished by cordate leaves, long petioles. 
F.M. Neg. 36114. 

Amazonas(?): Mathews 1316 (also 1273; 1274; 1275). Without 
locality (Haenke, type). 

Rhynchanthera Hookeri Naud. Ann. Sci. Nat. se>. 3. 12: 212. 
1849; 109. R. Williamsii Gleason, Bull. Torrey Club 58: 216. 1931(7). 

Calyx teeth slightly glandular-pubescent, subulate, 2-4 mm. long, 
shorter than the more densely glandular villous tube; petals 7-10 mm. 



258 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, rounded and apiculate at tip; fertile stamens about equal; 
connective 2-toothed at base; stems hispid; leaves cordate-ovate, 
7-nerved, acuminate, ciliate-margined, glabrate, about 5 cm. long, 
3 cm. wide, reduced in the open inflorescence; ovary ovoid, the 
style 6.5-15 mm. long; capsule globose to 6 mm. thick, slightly 
puberulent-glandular at tip. The author of R. Williamsii wrote: 
"R. Hookeri has a hispid stem, shorter and spreading sepals, longer 
and narrower petals attenuate at base, shorter anther beak, a slender 
and more elongate connective 2-toothed at base, ovoid ovary and 
much longer style." But in the type (as seen by me) the stems are 
not more hispid or the sepals more spreading, and in both types the 
petals are somewhat attenuate; the other "differences" are clearly 
relative, the connective being 2-toothed in both types; probably the 
material seen by Gleason was less mature, or actually mixed. F.M. 
Neg. 26098. 

San Martin: San Roque, Williams 7283 (det. GL); Williams 7284 
(type, R. Williamsii). Near Moyobamba, Mathews 1276; Klug 
3431. Colombia. "Chichirilli-sacha," "chicitilla sacha." 

3. CENTRADENIASTRUM Cogn. 

Small shrubs or sometimes herbs with petioled leaves and rather 
small 4 (5)-merous cymose flowers. Calyx teeth shorter than the tube. 
Stamens unequal, the connective of the larger erostrate anthers 
strongly produced and more or less extended above the insertion of 
the filament. Ovary glabrous, (3) 4-celled. Cogniaux separated this 
genus from Centradenia G. Don because the connective of the 
smaller anthers is not produced, unappendaged ; also, the leaves in 
Centradenia are disparous. In view of the presence of similar varia- 
tions elsewhere in the family it does not seem to me that it is useful 
or necessary to regard the Peruvian plant as generically distinct but 
of course it is quite possible to do so; the other (known) species are 
Mexican or Central American, with one from Colombia, and the 
recognition of Centradeniastrum as more than a section destroys the 
interesting conception of a range for a group of plants obviously 
closely allied. This is yet one more instance where one interested 
in phytogeography finds his materials obscured by a too fine drawing 
of generic lines. The species, if it is transferred to Centradenia, 
could well be named for Cogniaux, there being a C. rosea. 

Centradeniastrum roseum Cogn. Bot. Jahrb. 42: 131. 1908. 

Branches erect, glandular-puberulent, 4-5 dm. high; petioles 
filiform, 5-15 mm. long; leaves bright green and glabrous above, 



FLORA OF PERU 259 

pale beneath, ovate, about 2 cm. long or shorter, 8-18 mm. wide; 
cymes 3-6 cm. long; pedicels 3-8 mm. long; calyx tube about 3 mm. 
long, the triangular subulate teeth half as long; petals 8 mm. long; 
longer anthers 3 mm. long, the produced part as long, but only 
shortly and obtusely extended above the filament-insertion. F.M. 
Neg. 16684. 

Cajamarca: Chugur near Hualgayoc, in wet shrubby wood, 
2,800 meters, Weberbauer 4086, type. 

4. ERNESTIA DC. 

Shrubby or subherbaceous glandular pubescent plants with rather 
thin 3-7-nerved serrulate leaves and open terminal panicles of 
rather small flowers. Calyx glandular the subulate-triangular lobes 
about equaling the 8-ribbed tube, this membranous in fruit. Sta- 
mens unequal but nearly alike, the filiform connective appendages 
longer than the connective. Capsule 3- or 4-celled, apically glabrous 
or setose. The species could readily be included in Arthrostemma. 

Calyx lobes longer than the tube; stamens alike E. tenella. 

Calyx lobes and tube subequal; stamens dissimilar. . . .E. quadriseta. 

Ernestia quadriseta Berg ex Triana, Trans. Linn. Soc. 28: 35. 
1871. E. quadrisetosa Berg, fide Cogn. Melast. 144. 1891. 

Stems, leaves both sides and panicle branches densely short- 
hispidulous with divaricate trichomes, only those on the leaves not 
gland-tipped, those on the leaves beneath rather finer; petioles to 
1.5 cm. long; leaves 5-7-nerved, 3-5 cm. long, 2-3 cm. wide; inflores- 
cence rather close, leafy; calyx tube about 3 mm. long, the purplish 
petals 8-9 mm. long, the calyx lobes narrowly triangular. Illus- 
trated, Triana, I.e. pi. 2. F.M. Negs. 21166; 36124. 

San Martin: Tarapoto, Spruce 4-584 (4884, fide Triana), type; 
Williams 5989 (det. Gl.). Loreto: Balsapuerto, Killip & Smith 
28600 (det. Gl.); King 3021; 2868. 

Ernestia tenella (Bonpl.) DC. Prodr. 3: 121. 1828; 144. Rhexia 
tenella Bonpl. Rhex. 79. pi. 30. 1823. E. tenella var. Sprucei Cogn. in 
Mart. Fl. Bras. 14, pt. 3: 227. 1885. E. Sprucei (Cogn.) Macbr. Field 
Mus. Bot. 4: 172. 1929. 

Similar to E. quadriseta but the inflorescence very open and the 
pubescence longer and looser; calyx lobes subulate, 4-5 mm. long; 
petals white; ovary scarcely puberulent apically. The var. Sprucei 
is a form with purple flowers and ovary setulose, and is the Peruvian 



260 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

plant, but the difference is hardly constant or noteworthy. F.M. 
Negs. 36125; 21167 (var.). 

Amazonas: Moyobamba, Mathews 1277. Brazil. 

5. ARTHROSTEMMA R. & P. 

Weak-stemmed or semi-scandent sparsely glandular-setose herbs 
with long internodes, sometimes somewhat woody toward the base. 
Leaves petioled, broad, ciliately serrulate. Flowers rather large, 
borne apically, cymose or scorpoid, 4-merous. Calyx tube oblong 
or narrowly campanulate, the much shorter calyx lobes persisting on 
the often conspicuous and lustrous fruit. Stamens dissimilar, the 
connective of the larger with filiform erect appendage that is clavate 
and tridenticulate, that of the smaller biaristate. Capsules elongate, 
the seeds cochleate. 

Arthrostemma ciliatum R. & P. Fl. Peruv. 4. pi. 326. 1802; 
Pav. ex D. Don, Mem. Wern. Soc. 4, pt. 2: 299. 1823; 140. Rhexia 
diversifolia Bonpl. Rhex. 119. pi. 45. 1823. 

Herb trailing over bushes or sprawling on wet banks, smooth or 
nearly, even the younger branches little glandular; petioles 1-2 cm. 
long; leaves ovate, lightly cordate, acuminate, 5-nerved, 4-6 cm. 
long, about half as wide; flowers loosely borne, to 1.5 cm. long; cap- 
sules (as to type) 15 mm. long, 6-7 mm. thick, the brown seeds 
beautifully rugose-tuberculate. The type is in fruit but it is doubt- 
ful if there is more than one species characterized by nearly equal 
stamens. A. macrodesmum Gleason (A. campanula, (Naud.) Tr. 
not DC.) found as near as Ecuador has shortly pediceled flowers and 
stamens very unequal, the connective of the episepalous stamens as 
long or longer than the thecae; the connective of the widely dis- 
tributed and otherwise similar A. fragile Lindl. is much shorter than 
the thecae (Gleason). 

San Martin: Tocache, Poeppig 1690. Near Tarapoto, Williams 
6772; 7143. Huanuco: Chinchao and Cuchero, Ruiz & Pavon. 
Pozuzo, 4681. Without locality, Dombey. Junin: La Merced, 5411. 
Ecuador. 

Arthrostemma grandiflorum Markgr. Notizbl. Bot. Gart. 
Berlin 9: 1139. 1927. 

Suffrutescent, the scandent stems more or less glandular-setose; 
leaves 5-7-nerved; petals 2.5 cm. long; stamens unequal; ovary 
apically glandular-setose. Probably a large-flowered race of the 
preceding but allied by the author to A. latifolium D. Don because 



FLORA OF PERU 261 

of the pubescent ovary; the allied genus Ernestia has in E. tenella a 
species in which the pubescence of the ovary varies. As Gleason, 
who has referred here the K. & S. collections, has suggested, the genus 
needs monographic study. The following material, except as to 
type with larger flowers (!), I should refer to the R. & P. species, 
variable in capsule size and pubescence. F.M. Neg. 16688. 

Junin: Puerto Bermudez, Killip & Smith 26618. Huanuco: 
Region Churubamba, Mexia 8156. Loreto: Pongo de Manseriche, 
Tessmann 3904, type. Mouth of Santiago, Tessmann 4153. Balsa- 
puerto, Klug 2892. Florida, Klug 2319. Ayacucho: Killip & 
Smith 23082; 23114. 

6. NEPSERA Naud. 

Herb or woody below with petioled, cordate-ovate, serrulate, 5-7- 
nerved leaves and extremely lax, trichotomously branched panicles 
of small white 4-merous flowers, these with oblong-lanceolate acute 
petals. Calyx lobes as long as the tube, persistent. Stamens unequal, 
the subulate anthers dissimilar; connective arcuately produced and 
anteriorly with 2 recurved-ascending appendages. Ovary globose, 
glabrous, free, 3-celled. Seeds short, cochleate, coarsely foveolate. 

Nepsera aquatica (Aublet) Naud. Ann. Sci. Nat. se>. 3. 12: pi. 
14; se>. 3. 13: 28. 1849; 146. Melastoma aquatica Aublet, PI. Guian. 
1:430. 1775. 

Stems to one meter long or longer, the thin leaves 4-6 cm. long, 
1.5-2.5 cm. wide, the pyramidal panicle 1-3 dm. long, its branches 
exceptionally slender; calyx tube 2-3 mm. long; petals 5-6 mm. long; 
capsule 2 mm. thick. Widely distributed and found near Peru, it 
is to be expected in the Amazonian region. Illustrated, Mart. Fl. 
Bras. 14, pt. 3: pi. 53. 

Peru (probably). Brazil to Ecuador and the Antilles. 

7. DESMOSCELIS Naud. 

Erect, stout or subligneous virgate-stemmed, silky-villous herb 
with medium-sized 5-merous flowers more or less crowded on short 
axillary branchlets. Calyx lobes persistent, about as long as the 
campanulate tube. Stamens very unequal, the oblong-subulate 
anthers dissimilar, the connective of the larger long-produced and 
provided with two anterior filiform appendages, that of the smaller 
arcuate and merely biauriculate anteriorly. Ovary more or less 
adherent, pubescent apically, 5-celled. Seeds cochleate, granulate. 
The habit and long-villous pubescence which extends even to the 



262 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

calyx identify the plant vegetatively; technically it is perhaps, in 
view of variation in the family, arbitrarily separated from Acisan- 
thera P. Br. with free and glabrous 2-4-celled ovary; the latter, in 
A. quadrata Juss., 130, was accredited to the Peruvian flora by 
Cogniaux on the basis of a Pavon (so labeled) specimen, doubtless 
of Antillean or Central American origin. 

Desmoscelis villosa (Aublet) Naud. Ann. Sci. Nat. se>. 3. 12: 
pi. 14; ser. 3. 13: 30. 1849; 147. Melastoma villosa Aublet, PL Guian. 
1:428. 1775. 

Simple or somewhat branched, the flowers commonly crowded 
at the tips of short axillary branchlets or sometimes thyrsoid or 
paniculate; trichomes on the stems 3-4 mm. long or even longer; 
calyx tube and ovate-subulate lobes 5-6 mm. long; petals 1 cm. long; 
capsule 4-6 mm. long. Leaves usually ovate-lanceolate, 3-5 cm. 
long, about 2 cm. wide or narrower; variable particularly in form of 
inflorescence. Illustrated, Bonpl. Rhex. pis. 31 and 1$. 

San Martin: San Roque, Williams 7118; 7^10 (det. GL). Bolivia 
to Guiana and Colombia. 

8. BRACHYOTUM (DC.) Triana 

Shrubs, often compactly branched, with numerous rather small 
or small usually harshly setose leaves and more or less nodding 
tubular-campanulate 4-5-merous flowers commonly borne in three's 
at the ends of branchlets, sometimes solitary or cymose. Calyx 
lobes persistent. Stamens equal, the anthers similar, narrow, the 
connective not elongated, simple or anteriorly bitubercled or bical- 
loused. Ovary free, apically setose. Seeds cochleate. Approaches 
Tibouchina through some species, but characterized by the shape of 
the flowers. In the following key the character "flowers 4- or 5- 
merous" has been avoided because some species are already known 
to vary in this respect; similarly questionable as fundamental is 
the group-division of the species based on anther character; the 
anthers may be bicalloused at base (this sometimes very obscure) or 
smooth in species otherwise much alike. 
Flowers ebracteate, the bracts early caducous. 

Pubescence, at least that of the branchlets, closely appressed- 
strigose; anthers minutely calloused at base except B. nutans, 
B. alpinum, B. asperum, B. ledifolium. 

Leaves narrow or small, usually revolute, all or most of them 
1-4(5) mm. wide, 3-15 mm. long, oblong-linear or, if ovate, 
inordinately calloused. 



FLORA OF PERU 263 

Leaves ovate, the surface completely covered with tubercles. 
Leaves to about 5 mm. long, 2.5 mm. broad .B. lycopodioides. 
Leaves about 3 mm. long, 2 mm. wide B. minimum. 

Leaves narrowly oblong-lanceolate, more or less calloused- 
setose above. 

Leaves sparsely setose beneath B. Grisebachii. 

Leaves densely setulose beneath B. rosmarinifolium. 

Leaves larger, usually broader, or if only 4-6 mm. wide, not at 
all conspicuously calloused. 

Leaves glabrous or only very sparsely setulose above. 

Calyx lobes narrow, soon longer than the tube; anthers 
bicalcarate B. cernuum. 

Calyx lobes ovate, shorter than or as long as the tube; 
anthers smooth. 

Petals about 2 cm. long B. nutans. 

Petals about 1 cm. long B. alpinum. 

Leaves obviously, often densely setulose above. 

Calyx lobes subulate-linear, as long or longer than the 
tube; leaves 5-nerved. 

Leaves setulose B. quinquenerve. 

Leaves silky-villous B. campanula. 

Calyx lobes ovate to narrowly ovate-lanceolate, sometimes 
as long as the tube; leaves 3-nerved. 

Leaves sparsely setulose on nerves beneath, sometimes 
on veins, not at all granulose. 

Calyx lobes remote at base, much shorter than tube. 

Flowers mostly or all ternate B. Pentlandii. 

Flowers corymbose-paniculate B. floribundum. 

Calyx lobes usually contiguous, often about equaling 
the tube. 

Leaves ovate-elliptic, obtuse, about 1 cm. long, 5-8 
mm. wide B. Naudinii. 

Leaves ovate-oblong, subacute, often longer. 

B. strigosum. 

Leaves tomentose beneath or at least granulose on the 
veins, often also more or less setulose. 



264 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx sericeo-strigose. 

Anthers bituberculate at base B. Radula. 

Anthers etuberculate at base B. asperum. 

Calyx very shortly- and appressed-setulose. 

B. ledifolium. 

Pubescence, at least that of the branchlets, spreading or ascending, 

in any case never appressed strigose-setulose, sometimes 

villous, sometimes setulose; anthers smooth except B. Bar- 

beyanam, B. Figueroae, B. parvifolium. 

Leaves oblong- or lanceolate-ovate, 2-3 times longer than broad, 

acute or subacute, usually revolute. 
Leaves small, to about 15 mm. long. 
Calyx lobes oblong-linear; leaves rarely 10 mm. long. 

B. microphyllum. 

Calyx lobes ovate-oblong; leaves mostly 10-12 mm. long. 

B. Trianaei. 
Leaves mostly longer than 2 cm. 

Pubescence spreading; petioles 5-10 mm. long. 

B. Weberbaueri. 
Pubescence subappressed ; petioles 2-4 mm. long. 

B. ledifolium. 

Leaves broadly ovate-elliptic, rarely twice longer than broad, 

often obtuse, often plane. 
Branchlets setulose, the trichomes firm. 
Pubescence on leaves beneath merely setulose. 
Pubescence sparse, confined to veins; shrub lax. 

B. lutescens. 

Pubescence dense; shrub compact, strict. . .B. Figueroae. 
Pubescence on leaves beneath consisting of two sorts of 

trichomes B. canescens. 

Branchlets villous or villous-hirsute. 

Calyx sparsely setulose; anthers smooth B. rostratum. 

Calyx densely sericeous; anthers minutely bicalcarate. 

Leaves 1.5-3.5 cm. long B. Barbeyanum. 

Leaves 8-15 mm. long B. parvifolium. 

Flowers bracteate. 

Bracts suborbicular; anther connective not calloused. 

Leaves small; flowers solitary B. confertum. 

Leaves ample; flowers racemose B. racemosum. 



FLORA OF PERU 265 

Bracts narrowly ovate or lanceolate; anther connective obscurely 

bicalloused. 
Bracts narrowly ovate; leaves about 1 cm. wide. 

B. Benthamianum. 

Bracts narrowly lanceolate; leaves wider B. quinquenerve. 

Brachyotum alpinum Cogn. Melast. 167. 1891. 

Apparently like B. nutans but the calyx lobes narrowly ovate- 
lanceolate, 4 mm. long, the tube 6 mm. long and the petals only 11- 
12 mm. long. Of the same alliance but the much larger leaves 
slightly setulose above, the flowering peduncle filiform, is B. graci- 
lescens Triana, 167, of Ecuador, to be expected. B. alpinum also 
being Ecuadorian I suspect that at least the Cuzco collection cited 
here belongs to B. nutans. F.M. Neg. 25862. 

Ayacucho: Quinua, 3,300 meters, Weberbauer 5540 (det. Cogn.). 
Totorabamba, 3,600 meters, Weberbauer 5484 (det. Cogn.). 
Cuzco: Valle del Apurimac, Herrera 757 (det. Markgr.). Ecuador. 
"Carhuinchu." 

Brachyotum asperum Cogn. Bot. Jahrb. 42: 132. 1908. 

Young branchlets shortly appressed-setulose; petioles 2-5 mm. 
long; leaves rigid, nearly plane, narrowly ovate-oblong, entire, 3-5- 
nerved, rather densely callous-asperous above but green, the setae 
very short, densely and shortly ashy-setulose beneath, 3-4.5 cm. 
long, 20-27 mm. wide; flowers 5-merous, solitary, ternate, nodding; 
calyx ashy appressed-setulose, the tube 8-9 mm. long, the oblong, 
apiculate, rigid lobes 4-5 mm. long; petals narrowly obovate, setulose 
dorsally toward the acute tip, 17-18 mm. long; anthers 5 mm. long. 
Allied by the author to B. rostratum. Corolla dark gentian blue, 
calyx light rust-color (S. & H.). Shrub 1 meter high. B. gracilescens 
Triana, 167, Ecuadorian, has thin, less pubescent leaves, petioles 
to 10 mm. long. F.M. Neg. 16707. 

Cajamarca: Near Hualgayoc, 3,400 meters, Weberbauer 4013, 
type. In clay-gravel, 3,000 meters, La Tajona, Stork & Horton 
10023. 

Brachyotum Barbeyanum Cogn. Melast. 158. 1891. 

Branchlets appressed-villous-hirsute, the hairs reddish; petioles 
to 6 mm. long; leaves narrowly ovate, densely appressed-setose 
above the surface, soft to the touch but the trichomes minutely 
bulbous-based, the 3 nerves densely red-villous beneath, 1.5-3.5 cm. 
long; calyx lobes ovate-lanceolate, about equaling the softly long- 



266 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

villous tube, this 8 mm. long; flowers 5-merous, the subtruncate 
petals scarcely ciliate, 2 cm. long; anthers 5 mm. long; ovary shortly 
setose; style 2.5 cm. long. In age leaves are probably larger, pubes- 
cence harsher above; the type is only a flowering branchlet. F.M. 
Neg. 36853. 

Amazonas: Chachapoyas, Mathews, type. 

Brachyotum Benthamianum Triana, Trans. Linn. Soc. 28: 
49. 1871; 162. 

Younger branchlets shortly appressed-strigose; leaves densely 
and finely appressed bullate-setulose, the 3 nerves prominent above, 
finely appressed-setulose beneath, many only 2 cm. long, 1 cm. wide 
or smaller; bracts 1-1.5 cm. long, glabrous within; flowers 4- or 
5-merous, subsessile, solitary or ternate; calyx densely sericeous, the 
membranous lobes slightly longer than the tube, this 6 mm. long; 
petals narrowly obovate, subrotund at tip, 1.5 cm. long. Similar 
Ecuadorian species also to be expected areB. Jamesonii Triana, 161, 
and B. campylanthum Triana, 163, the former with smaller, less 
pubescent leaves, the latter with flowers solitary at tips of lateral 
branchlets, the bracts obovate. F.M. Neg. 16708. Illustrated, 
Baill. Hist. PI. 7: 8. fig. 11. 

Peru (?): (Seemann 773 bis, fide Cogn.). Ecuador. 

Brachyotum campanulare (Bonpl.) Triana, Trans. Linn. Soc. 
28: 48. 1871 ; 153. Rhexia campanularis Bonpl. Rhex. 35. pi. 14. 1823. 

Leaves densely sericeous-villous above and below, the trichomes 
above, at least typically, densely appressed but not at all callous- 
based, 2-3 cm. long, 12-17 mm. wide, the upper smaller; flowers 
usually solitary; calyx tube 4-5 mm. long, the lobes 7-9 mm. long, 
lanceolate-linear, acuminate; petals 12-14 mm. long, obovate-oblong, 
the acute tip terminating in a bristle. A Pavon specimen without 
locality, referred here by Cogniaux, is appressed-setulose above, 
appressed-sericeous beneath, the calyx lobes only about equaling 
the tube; there is no proof that it is Peruvian and it seems to me 
that it does not belong here. The type actually came from Loja. 
B. rotundifolium Cogn., 152, Ecuador, has 7-nerved leaves to 2 cm. 
long and nearly as wide. F.M. Neg. 36135. 

Huanuco: Chinchao (MacLean, fide Cogn.). Ecuador. 

Brachyotum canescens (Bonpl.) Triana, Trans. Linn. Soc. 28: 
48. 1871; 165. Rhexia canescens Bonpl. Rhex. 14. pi. 6. 1823. 

Petioles 2-3 mm. long; leaves finely appressed-setulose above, 
the trichomes only minutely calloused or scarcely, the midnerve 



FLORA OF PERU 267 

and nerves beneath setulose hispid, a few minute setae mixed with 
the intervening tomentum, 3-nerved, ovate, 1-2 (3) cm. long, 5-10 
mm. wide; calyx lobes firm, shorter than or nearly equaling the tube, 
this 6 mm. long; flowers 5-merous the scarcely ciliate petals purple 
or yellowish. F.M. Neg. 36136. 

Ancash: Between Huaraz and Yungay, Weberbauer 2928; 179. 
Colombia. "Sarzilejo." 

Brachyotum cernuum (Bonpl.) Triana, Trans. Linn. Soc. 28: 
48. pi. 3. 1871; 157. Rhexia cernua Bonpl. Rhex. 32. pi. 13. 1823. 

Branches 4-angled, the younger sparsely appressed-setulose, 
especially at the nodes; petioles 2-5 mm. long; leaves oblong-ovate, 
1.5-3 cm. long, 6-15 mm. wide, acute, glabrous except the 3 nerves 
lightly setulose beneath; flowers usually 3-5, nodding on pedicels 
to 1 cm. long; calyx sparsely and minutely setose, the tube 5-7 mm. 
long, the erect narrow membranous lobes twice to thrice as long; 
petals obovate, subacute, shortly ciliolate, 2-2.5 cm. long, the style 
as long; anthers 5 mm. long. Description by Cogniaux. B. tri- 
chocalyx Triana, 158, with small, glabrous leaves but long-setose 
calyx may be expected from Ecuador. F.M. Neg. 36137. 

Peru (perhaps). Ecuador. Colombia. 

Brachyotum confertum (Bonpl.) Triana, Trans. Linn. Soc. 
28: 49. pi. 3. 1871; 163. Rhexia conferta Bonpl. Rhex. 53. pi. 
20. 1823. 

Abundantly and slenderly branched shrub to about 1 meter high, 
with numerous almost tiny leaves; branchlets minutely but densely 
strigose; petioles to 2 mm. long; leaves narrowly ovate, 3-nerved, 
rigid, revolute, glabrate beneath except for setulose midnerve and 
margin, finely appressed-setulose above, 3-7 mm. long, 2-3 mm. 
wide; flowers 5-merous subtended by 6 suborbicular rather mem- 
branous appressed-sericeous bracts 8-12 mm. long; calyx sericeous, 
the tube 6 mm. long, the ovate lobes slightly shorter; petals subor- 
bicular, 10-12 mm. long; anthers 3 mm. long. Illustrated, Bot. 
Mag. pi. 6018. B. Andreanum Cogn., 164, Ecuadorian, has larger 
leaves and flowers. 

Amazonas: Chachapoyas, Mathews; Fielding 1225. Above 
Balsas, 3,400-4,000 meters, Weberbauer 4291; 268. Ecuador. 

Brachyotum Figueroae Macbr. Field Mus. Bot. 4: 173. 1929. 
Branchlets crowded, densely subappressed-setulose, tardily gla- 
brate; petioles and leaves beneath densely hirsute, the leaves broadly 



268 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ovate, about 9 mm. long, 5-6 mm. wide, conical-tubercled above, 
the tubercle setiferous; flowers solitary or 3-4, 4-5-merous, whitish, 
the rounded obovate shortly ciliolate petals about 12 mm. long; 
calyx lobes ovate, about 4 mm. long, scarcely half as long as the 
sparsely setulose tube; anther minutely bituberculate. Simulates 
B. lutescens with smooth anther, and is doubtfully distinct. About 
1 meter high. 

Ancash: Catuc, 7 km. east of Huaraz, 2504. "Cotchkis bianco." 

Brachyotum floribundum [Griseb.] Triana ex Cogn. Melast. 
155. 1891. 

Shrub with relatively long and slender appressed-setulose branch- 
lets distinctive by its cymose inflorescence; petioles 2-4 mm. long; 
leaves mostly 3-4 cm. long, 1.5 cm. wide, acutish both ends, ap- 
pressed-setulose above, the setae with very slender callus at base, 
beneath rather sparsely setose-hispid on the veins and 3-5 nerves, 
otherwise glabrous; flowers 4-merous; petals 12-14 mm. long; 
pedicels slender, setulose, articulate at middle, 5-15 mm. long; 
calyx attenuate at base, 5 mm. long, slightly setose, the broad lobes 
scarcely 1 mm. long. Grisebach assigned the name under Chaeto- 
gastra in herbaria. B. sanguinolentum Triana, 154, to be expected 
in southern Peru from Bolivia, is glabrate, the flowers twice as 
large. F.M. Neg. 16709. 

Puno: Tabina, Lechler 1857, type. Sandia Valley, 2,300 meters, 
Weberbauer 626; 237. Cuzco: Near Huallahualla, Marcapata, 3,300 
meters, Vargas 9708. "Tili tili." 

Brachyotum Grisebachii Cogn. Melast. 153. 1891. 

Similar to R. rosmarinifolium; typically leaves 1-1.5 (2) cm. 
long, 4-7 mm. wide, sparsely setulose above, not bullate, glabrous 
beneath except the sparsely setulose nerves; flowers 4-merous; calyx 
sparsely appressed-setose, the lobes about as long as the tube, this 
4-5 (6) mm. long; petals about 2 cm. long, subacute. According to 
Cogniaux the calyx tube is 7-8 mm. long, the lobes 8-10 mm. long. 
F.M. Neg. 16711. 

Puno: Near Agapata, Lechler 1856, type. Cuzco: Prov. of Cuzco, 
Gay; Weddell; Herrera. Ecuador? 

Brachyotum ledifolium (Desr.) Triana, Trans. Linn. Soc. 28: 
48. 1871; 165. Melastoma ledifolia Desr. in Lam. Encycl. 4: 48. 
1797. Chaetogastra sulphured Naud. Ann. Sci. Nat. se"r. 3. 14: 135. 
1850. C. Bonplandiana Naud. I.e. 137. pi. 4(1). 



FLORA OF PERU 269 

Branches glabrate, exfoliating; branchlets appressed-setose, the 
setae rigid; petioles 2-4 mm. long; leaves oblongish, 1.5-2.5 (3) 
cm. long, 5-7 (10) mm. wide, bullate (not calloused) above and 
shortly but sparsely setulose, beneath granulose-setulose, usually 
^revolute at margin; flowers yellowish, often ternate, 5-merous, sub- 
sessile, nodding; calyx lobes oblong-elliptic, 3-4 mm. long, much 
shorter than the campanulate, appressed-setulose tube; petals 
obovate-suborbicular, 1.5 cm. long; style 2 cm. long; anthers 5 mm. 
long. C. Bonplandianum Naud. of Ecuador is similar but is probably 
distinct as the branchlets are singularly ashy scurfy-pubescent. 
F.M. Neg. 36131. 

Peru (?):/. Jussieu, type. Without locality, Ruiz & Pavon (det. 
Markgr.). Ecuador. 

Brachyotum lutescens (R. & P.) Triana, Trans. Linn. Soc. 28: 
48. 1871 ; 166. Rhexia lutescens R. & P. Fl. Peruv. 3: 84. pi. 319. 1802. 

Densely branched shrub, the branches and branchlets spreading 
rusty-setulose; petioles 2-3 mm. long; leaves ovate, blunt, plane, 
1-1.5 cm. long, 7-9 mm. wide, bullate-calloused setulose above, 
sparsely setulose beneath; flowers yellowish or whitish and some- 
times red-edged, often ternate, often nodding, 4- or 5-merous; calyx 
at first somewhat glandular hirsute, the glands evanescent, the 
oblongish lobes about half as long as the tube, this about 8 mm. 
long; petals obovate, somewhat rounded at tip, 15 mm. long; anthers 
7-8 mm. long. Also at Muna and Chaglla (Ruiz & Pavon). Prob- 
ably with more material will be found to include B. Trianae and 
B. Figueroae. F.M. Neg. 16713. 

Huanuco: Pillao, Ruiz & Pavon, type. Mito, 1872. 

Brachyotum lycopodioides Triana, Trans. Linn. Soc. 28: 49. 
1871; 161. 

Closely allied to B. rosmarinifolium but the leaves lineate, tuber- 
cled above, the lustrous tubercles usually in 4 rows and leaving no 
free leaf -surf ace; leaves mostly only 3-4 (6) mm. long, strigose-hir- 
sute beneath, rigid, revolute-margined; flowers nodding, 5-merous; 
oblongish calyx teeth and tube subequal, 3-4 mm. long; petals 
broadly ovate, retuse, 9 mm. long. Illustrated, Weberbauer, page 234. 

Amazonas: Bajasan (Mathews 1254, type). Chachapoyas, 2,400 
meters, Weberbauer 4399; 264 (det. Cogn.); Williams 7587 (det. Gl.). 
Huanuco: Mountains of Pozuzo (Pearce). Junin: West of Hua- 
capistana, Weberbauer 2072; 251, 234. 



270 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Brachyotum Maximowiczii Cogn. Melast. 154. 1891. 

Younger branches densely appressed-setose; petioles 2-3 mm. 
long; leaves ovate, minutely crenulate, blunt, densely calloused 
above with conical tubercles, thickly pilose beneath, 8-12 mm. long, 
5-7 mm. wide; pedicels very short but bibracteolate, the bractlets 
ovate-oblong, 5-7 mm. long; calyx tube 6 mm. long, the narrowly 
ovate lobes somewhat rounded at apex; petals narrowly obovate, 
subrounded at tip, 13 mm. long; anthers 2.5-3 mm. long. The 
flowers were described as 4-merous. 

Amazonas: Bejasan, prov. Chachapoyas (Fielding 1265). 

Brachyotum microphyllum (Naud.) Triana, Trans. Linn. Soc. 
28: 49. 1871; 164. Chaetogastra microphylla Naud. Ann. Sci. Nat. 
ser. 3. 12: 136. 1849. 

Branchlets and leaves beneath densely hispidulous with short, 
spreading trichomes; leaves rather sparsely strigose above, setulose 
beneath, 6-9 mm. long, 3-5 mm. wide, obsoletely 3-nerved; flowers 
5-merous, nodding; calyx sparsely setulose, the membranous subu- 
late-linear lobes subequaling the tube, this 5 mm. long; petals 
broadly obovate. Perhaps a variety of B. rosmarinifolium (Triana). 
In the type the leaves are finely densely callous-setulose above, 
densely setulose beneath. F.M. Neg. 36133. 

Peru (?) : Without locality, Bonpland. 

Brachyotum minimum Markgr. Notizbl. Bot. Gart. Berlin 13: 
459. 1937. 

Nearly B. lycopodioides; leaves ovate, 3 mm. long, 2 mm. broad; 
calyx tube 8 mm. long, densely sericeous-setose, the lanceolate 
teeth half as long, 1 mm. broad; petals orbicular, shortly acuminate 
and glandular-apiculate in bud, only 2 mm. long and broad; fruit 
campanulate, 6 mm. broad. 

Cajamarca: With old fruits and young flowers, May, 1879 
(Raimondi, three sheets, type). 

Brachyotum Naudini Triana, Trans. Linn. Soc. 28 : 48. 1871 ; 159. 

Much branched; petioles 3-4 mm. long; leaves ovate-elliptic, 
about 1 cm. long, 5-8 mm. wide, appressed-setulose above and all 
over beneath, the trichomes on the upper surface with slender cal- 
loused base; flowers nodding, sometimes 5-, sometimes 4-merous; 
calyx lobes broadly triangular, acute, equaling the tube, this about 
5 mm. long; petals ovate, acutish, ciliate, 1.5 cm. long. The leaves 



FLORA OF PERU 271 

are plane, scarcely bullate. The Williams collection has narrower, 
longer calyx lobes than type, slightly larger leaves. F.M. Neg. 
36134. 

Huanuco: Palca, Dombey. Amazonas: Chachapoyas, Williams 

7572. 

Brachyotum nutans Gleason, Bull. Torrey Club 54: 25. 1927. 

Branchlets sparsely appressed-setose with a ring of setae at the 
nodes; petioles 2-3 mm. long; leaves 3-nerved, glabrous above, 
sparsely appressed-setose beneath, ovate-oblong, 8-12 mm. long, 4-6 
mm. wide; flowers 4-merous; peduncle nodding, 5 mm. long; calyx 
glabrous or nearly so, the broadly triangular sepals a little shorter 
than the tube, this 4.5 mm. long; petals 19 mm. long; connective 
with minute dorsal protuberance at base and 2 anterior callosities. 
Contrasted with B. alpinum with shorter petals. 

Cuzco: Paso de Tres Cruces, Cerro de Cusilluyoc, Pennell 138^7, 
type. Paucartambo, 3,700 meters, Vargas 320; Soukup 379. Yucay, 
Soukup 736. Ecuador. 

Brachyotum parvifolium Cogn. Bot. Jahrb. 42: 132. 1908. 

Younger numerous and irregular branchlets with the petioles, 
pedicels and leaves beneath densely villous, with rather short, erect- 
spreading, reddish trichomes; petioles 1-3 mm. long; leaves elliptic- 
ovate, 8-15 mm. long, 6-11 mm. wide, obscurely 3-nerved, pilose 
above; flowers solitary, 5-merous, the petals glabrous or ciliolate, 
truncate or retuse, about 14 mm. long; calyx densely long-appressed- 
villous, the rigid, triangular-linear lobes 6-7 mm. long equaling the 
tube; anthers 6 mm. long, minutely bicalcarate anteriorly; style 16 
mm. long; ovary densely setose above. Near B. Barbeyanum. 
F.M. Neg. 16715. 

Amazonas: Tambo Ventillas, Chachapoyas, Weberbauer 4406, type. 

Brachyotum Pentlandii (Naud.) Triana, Trans. Linn. Soc. 28: 
49. 1871; 156. Chaetogastra Pentlandii Naud. Ann. Sci. Nat. se*r. 3. 
14: 133. 1850. 

Younger branchlets minutely appressed-strigose; petioles 2-4 
mm. long; leaves nearly plane, narrowly ovate, entire, 3-nerved, 
obtuse, above shortly and finely calloused-setulose, beneath minutely 
and sparsely setulose on the nerves, 1.5-2.5 cm. long, 8-15 mm. 
wide; flowers often ternate; pedicels 3-5 mm. long, sometimes brae- 
tea te at base; calyx tube shortly dense-setulose, campanulate-hemi- 
spheric, 8-9 mm. long, the teeth only 2 mm. long, subulate; petals 



272 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

obcordate, scarcely ciliolate, 18 mm. long; style 2-2.5 cm. long. 
One of three similar Bolivian species to be expected in adjacent Peru; 
B. microdon (Naud.) Triana, 155, has petioles 5-10 mm. long, 
oblong-lanceolate acute leaves 4-8 cm. long, the pedicels 5-10 mm. 
long, the calyx pulverulent-setulose; B. hermannioides (Naud.) 
Triana, 156, is scarcely different (fromB. microdon), the leaves ovate, 
the calyx teeth erect, linear lanceolate. F.M. Neg. 36138. 
Peru (probably). Bolivia. 

Brachyotum quinquenerve (R. & P.) Triana, Trans. Linn. 
Soc. 28: 48. pi. 8. 1871; 153. Rhexia quinquenervis R. & P. Fl. Peruv. 
3: 83. pi 321. 1802. 

Branchlets slender, shortly appressed-setulose above; petioles 
5-10 mm. long; leaves ovate-oblong, 4-7 cm. long, 1.5-2.5 cm. wide, 
appressed-setulose above, densely so beneath, there the hairs appres- 
sed on the nerves, subspreading and finer between them; flowers 
nodding, 4-merous, ebracteate but the bracts on the pedicels not 
promptly caducous; calyx lobes narrowly linear, soon 10 mm. long, 
or finally even to half again as long as the tube; flowers several in a 
nodding panicle or 3-5 on short branchlets; pedicels 3-6 mm. long, 
equaled by the narrow bracts; petals obovate, acute, 1.5 cm. long; 
style 2 cm. long. A slender open shrub about 1 meter high, well 
marked by the combination of elongate calyx lobes and large leaves. 
The Pavon specimen mentioned under B. campanulare seems to 
belong here. The Cuzco plant is scarcely typical, the flowers more 
crowded, the bracts larger, more prominent, less promptly caducous. 
F.M. Neg. 16716. 

Huanuco: Without locality, Sawada P89 (det. Gl., B. micro- 
phyllum but connective calloused). Panao, Chaglla and Muna, 
Ruiz & Pavdn. Chinchao and Pillao, Ruiz & Pavon. Tambillo south- 
west of Panao, 3572. Playapampa, 4858. Yanano, 494-0. Junin: 
Kittip & Smith 24131; 24448 (det. GL). Chanchamayo, Isern 2099. 
Huacapistana, Weberbauer. Ayacucho: Killip & Smith 22272; 
22333 (det. Gl.). Amazonas: Chachapoyas, Weberbauer 4426. 
Cuzco: Basin of the Urubamba (Herrera 1964)', 3224; 1559; 3208. 
Convention, 2,400 meters, Vargas 801. Department unknown: 
Andimarca (Mathews 1170). Without locality, Bonpland; River o. 
"Cachiquis." 

Brachyotum racemosum Cogn. Bot. Jahrb. 42: 132: 1908. 
Branchlets shortly and densely sericeous; petioles 6-13 mm. long; 
leaves plane, oblong, 4-7 cm. long, 14-27 mm. wide, 5-nerved, densely 



FLORA OF PERU 273 

serieeous-villous both sides; racemes 3-5 cm. long; flowers greenish 
white, the shortly ciliate roundish petals 16-18 mm. long; bracts 4- 
seriate, suborbicular; calyx lobes triangular, acuminate, much shorter 
than the appressed sericeous tube, this 8-9 mm. long; anthers 5-6 
mm. long; style 2.5 cm. long. Allied to B. confertum. F.M. Neg. 
16714. 

Cajamarca: Prov. of Chota, 3,100 meters, west of Huambos, 
Weberbauer 4170, type. 

Brachyotum Radula Triana, Trans. Linn. Soc. 28: 48. pi. 3. 
1871; 160. 

Branchlets closely appressed-setulose; petioles 5 mm. long, some- 
times much longer; leaves ovate-elliptic or oblong, 2.5 cm. long, 12 
mm. wide or much smaller, often obviously and closely denticulate, 
rather sparsely and minutely callous-tuberculate with short setae 
above, shortly tomentose beneath; flowers pendent, few, 5-merous, 
calyx lobes membranous, rounded at tip, slightly shorter than the 
appressed-setulose tube, this 6-7 mm. long; petals broadly oblong, 
obtuse, more or less hirsutulous or glabrous, ciliolate, 15-18 mm. 
long; anthers 5 mm. long. F.M. Neg. 16717. 

Amazonas: Chachapoyas, Mathews 3210, type; Fielding 1260. 
Cajamarca: Near Hualgayoc, 3,100 meters, Weberbauer 1+130 (det. 
Cogn.);261. 

Brachyotum rosmarinifolium (R. & P.) Triana, Trans. Linn. 
Soc. 28: 49. 1871; 161. Rhexia rosmarinifolia R. & P. Fl. Peruv. 3: 
84. pi. 318. 1802. 

Densely branched shrub the younger branchlets appressed- 
strigose; petioles 1-2 mm. long; leaves narrowly oblong, revolute, 
2-4 mm. wide, 8-12 mm. long, callous-setulose above, appressed 
hirsute-setulose beneath; flowers 5-merous, nodding; calyx lobes 
ovate quite as long (type) or a little shorter than the moderately 
callous-setose tube, this 5 mm. long; petals obovate, scarcely ciliolate; 
anthers 3 mm. long. The Weberbauer 3378 and 2218 were referred 
by Cogniaux to B. Maximowiczii but they have the 5-merous flowers 
and foliage of this species. The flowers are sometimes several. F.M. 
Neg. 16907. 

Amazonas: In the mountains above Chachapoyas (Mathews 1253; 
MacLean; Weberbauer, 264). Ayacucho: Huanto (Pearce). Huan- 
uco: (Ruiz & Pavdn). Mito, 1870. Monzon, 3,300 meters, Weber- 
bauer 3378; 253. Junin: Tarma, Ruiz & Pavdn. Huacapistana, 



274 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3,300 meters, Weberbauer 2218; 251. Altos de Palca, Isern 583. 
"Cachiquis." 

Brachyotum rostratum (Naud.) Triana, Trans. Linn. Soc. 
28: 48. 1871; 165. Chaetogastra rostrata Naud. Ann. Sci. Nat. se>. 
3. 14: 135. 1850. 

Branchlets deciduously but at first densely long-hirsute; petioles 
2-4 mm. long; leaves 10-22 mm. long, 6-14 mm. wide, ovate, densely 
callous setulose in zones above, hispid on the nerves and hirtellous 
beneath; flowers 5-merous; calyx tube sparsely setose, 8 mm. long, 
the membranous oblongish lobes scarcely or nearly half as long; 
petals glabrous, somewhat truncate, 12 mm. long; anthers 7 mm. 
long, beaked. F.M. Negs. 16718; 38256. 

Peru: Without locality, Dombey, type. 

Brachyotum strigosum (L. f.) Triana, Trans. Linn. Soc. 28: 
49. 1871; 159. Melastoma strigosa L. f. Suppl. 236. 1781. Rhexia 
stricta Bonpl. Rhex. 19. pi. 8. 1823. 

Branchlets slender, the younger densely strigose; petioles 1-3 
mm. long; leaves ovate-oblong, revolute, rigid, sparsely and minutely 
hirtellous especially on the nerves beneath, finely setulose above, 
1-2 cm. long, 4-8 mm. wide; flowers nodding, 5- or sometimes 4- 
merous, solitary or ternate; pedicel bracts 1-1.5 cm. long; calyx tube 
broadly campanulate, 5 mm. long, densely long-setose, the mem- 
branous, narrowly oblong, acute lobes 6-8 mm. long; petals rounded 
at apex, ciliolate, 1-1.5 cm. long; style 1.5-2 cm. long, anthers rather 
long-bicalcarate. Very similar if not the same is B. Lindenii Cogn., 
159, Colombian, the calyx shortly strigose, the anthers shortly 
bicalcarate; to it Cogniaux referred Weberbauer 2930. 

Ancash: Pichin, 3,600 meters, Weberbauer 2930. Cuzco: Pau- 
cartambo, 3,550 meters, Vargas 319 (distr. as B. quinquenerve) . 
Ecuador; Colombia. 

Brachyotum Trianaei Cogn. Melast. 167. 1891. B. callosum 
Macbr. Field Mus. Bot. 4: 172. 1929? 

Branchlets densely hispidulous; petioles 2-3 mm. long; leaves 
ovate-oblong, 8-13 mm. long, 4-6 mm. wide, densely tuberculate- 
setose above, hirtellous-setulose beneath; flowers lutescent, ebrac- 
teate, 4- or 5-merous; calyx tube sparsely strigose, eglandular, the 
membranous acute lobes 4-5 mm. long, shorter than the tube; petals 
obovate, about 12 mm. long in the type of B. callosum. Anthers, as 
here interpreted, sometimes obscurely calloused. So far as one can 



FLORA OF PERU 275 

tell from description there seems to be only one species concerned. 
Shrub, 0.5-1.5 meters high. My 1871, in part, has small leaves 
suggestive of B. microphylla but calyx of this species. Perhaps not 
distinct from B. lutescens. F.M. Neg. 16719. 

Huanuco: In scattering hillside thickets northeast of Huanuco, 
2181, type, B. callosum. Mito, 1871 (petals greenish white, red- 
edged). Yanano, 4927 (flowers white). Huancavelica: Puno forma- 
tion near Salcabamba, 3,871 meters, Stork & Horton 10429. Cuzco: 
Paucartambo, Soukup 387. Puno: Tabina, Lechler 2061, type, B. 
Trianaei. 

Brachyotum tyrianthinum Macbr. Field Mus. Bot. 4: 174. 
1929. 

Moderately branched shrub 1-2 meters high, the slender, gla- 
brous branches little or not at all exfoliating; branchlets, petioles 
and leaves above more or less densely appressed-setulose; petioles 
about 3 mm. long; leaves ovate, mostly 3 cm. long, 1 cm. wide, 
sparsely hirtellous beneath; flowers solitary, nodding, 4-merous (as 
to type); calyx tube appressed-setulose, 7 mm. long, the longer 
lanceolate lobes sparsely setulose; petals subrotund, about 15 mm. 
long; connective obscurely tuberculate. The leaf setae above are 
finely calloused at base. Near B. Naudinii and B. Grisebachii. 

Huanuco: Mito, slender rather open shrub, 1438, type. South- 
east of Huanuco, 2092(1}. "Cachis." 

Brachyotum Weberbaueri Cogn. Bot. Jahrb. 42: 133. 1908. 

Branchlets densely pubescent with short spreading trichomes; 
petioles densely hirsutulous, 5-10 mm. long; leaves plane, 2-4.5 
cm. long, 7-14 mm. wide, above green but tuberculate, the setae 
short, beneath densely ashy-sericeous; flowers often ternate, 4- 
merous, subsessile, nodding, the retuse, glabrous but ciliolate petals 
green with violet edges, 13 mm. long; calyx tube 5 mm. long, spread- 
ing-setulose, the lance-linear acute lobes shorter; anthers 3 mm. 
long; style 2 cm. long. Fide the author, allied to B. Trianaei. To 
1 meter high. F.M. Neg. 16720. 

Amazonas: Tambo Ventillas, 2,400 meters, near Chachapoyas, 
Weberbauer 4405. 

9. MACAIREA DC. 

Similar to Tibouchina but the flowers usually in ample, densely 
flowered pyramidal panicles and the base of the anther-connective 
bilobed or gibbous posteriorly but unappendaged anteriorly. Flowers 



276 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

4-merous, in the Peruvian species not longer than 1 cm. In a few 
species the flowers are in axillary cymes; actually a component of 
Tibouchina, the group is nevertheless a convenient segregate. 
Calyces eglandular, the lobes about as long as the tube. . .M. scabra. 
Calyces glandular, the lobes longer than the tube M. sulcata. 

Macairea scabra Cogn. Bot. Jahrb. 42: 133. 1908. 

Branches slender; petioles 5-8 mm. long, with the peduncles and 
calyces shortly setulose, eglandular; leaves ovate-elliptic, entire, 
little narrowed at base, 3-nerved, 4-6 cm. long, 2-2.5 cm. wide, 
minutely tuberculate-setulose above, shortly and densely setulose 
beneath; panicle 10-17 cm. long, narrowly pyramidal; calyx lobes 
linear-triangular, shorter than or about equaling the tube, this 3-4 
mm. long; petals glabrous, 8-9 mm. long, the style about as long. 
A shrub related to M. albiflora Cogn., 177, about 1 meter high. F.M. 
Neg. 16704. 

San Martin: Rioja west of Moyobamba, 800 meters, Weberbauer 
4708, type; 290. 

Macairea sulcata Triana, Trans. Linn. Soc. 28: 38. 1871; 180. 

Branches angled, sulcate, setose, especially at the nodes; leaves 
elliptic-lanceolate, narrowed at base, resinous-punctate beneath and 
on the nerves, setose, smooth above, 5-8 cm. long, 1.5-2.5 cm. wide; 
inflorescence ample, many-flowered; calyx lobes much longer than 
the tube, this to 3 mm. long; petals 8-10 mm. long. Bark of branches 
exfoliates; the inflorescence branches are brown-hirsute. F.M. Neg. 
32324. 

San Martin: Tarapoto, Spruce 4004, type. 

10. SANDEMANIA Gleason 

Shrub with many paniculate 4-merous flowers. Hypanthium 
herbaceous, lightly 8-costate, the calyx tube not produced, its linear 
lobes short and with no outer teeth. Petals acute. Stamens little 
dimorphic, the slender subulate anthers somewhat beaked and 
dehiscent terminally by a pore; connective more or less produced, 
the outer series provided anteriorly with a U-form appendage or 
shortly bilobed (inner series). Ovary free, 2-celled, glabrous. Style 
slender, narrowed to the punctiform stigma. 

Sandemania lilacina Gleason, Kew Bull. 480. 1939. 
A 2-meter shrub, the upper subterete branches and terminal 
panicles densely strigose and furfuraceous; petioles 4-5 cm. long; 



FLORA OF PERU 277 

leaves narrowly oblong, entire, obtuse or subacute, obtuse at base, 
to 5 cm. long, 12 mm. broad, coriaceous, 5-nerved, glabrous above, 
beneath on veins pubescent like the stems, the surface shortly hir- 
sutulous and minutely tomentulose; panicles 1 dm. long; bracts 
broadly ovate, 3 mm. long, the narrower bractlets 1-2 mm. long, 
ciliate; pedicels 1 mm. long, villous; hypanthium shortly cylindrical, 
2.2 mm. long, sparsely hirtellous at base and with some setae between 
the sepals, these 0.7 mm. long; petals ovate, 3.5 mm. long; filaments 
5.2 or 4.1 mm. long, the connective to 1.3 mm. long; style nearly 4 
mm. long. Said to resemble most closely Macairea in general aspect 
but with different connective, glabrous ovary and filaments, acute 
petals. From Tibouchina and Pterolepis it differs in its glabrous 
ovary. But all these characters exist in degree in some genera in 
the same group. 

San Martin: Rioja, 900 meters (Sandeman 170, type). Jesus del 
Monte, near Moyobamba, 900 meters (Sandeman). 

11. PTEROGASTRA Naud. 

A group of weak-stemmed thin-leaved annuals readily recogniz- 
able by the winged calyx and capsule. Flowers rather large, solitary 
or cymose with unequal but similar stamens, the larger with connec- 
tive usually long-produced and bituberculate at the insertion of the 
filament. Ovary 4-5-celled, apically setose. Calyx enlarged in 
fruit and prominently 4-5-winged, the wings ciliate-echinate. 

Pterogastra divaricata (Bonpl.) Naud. Ann. Sci. Nat. se*r. 3. 
13: 33. 1849; 181. Rhexia divaricata Bonpl. Rhex. 59. pi. 22. 1823. 

Similar to P. major; leaves sparsely puberulent-pilose or setose, 
8-16 mm. wide; calyx lobes nearly as long as the tube, this 7-8 mm. 
long; connective of larger stamens shortly produced; petals to 12 
mm. long, rounded at apex; ovary eglandular. 

San Martin: San Roque, Williams 7116; 7643 (both det. GL). 
Near Moyobamba, Klug 3814- Loreto: Balsapuerto, Klug 3049. 
British Guiana; Orinoco. 

Pterogastra major Triana, Trans. Linn. Soc. 28: 40. pi. 2. 
1871; 182. 

Stems to several dm. long; leaves somewhat long-pilose both 
sides (or setose), oblong-lanceolate, 4-8 cm. long, typically 2-3.5 
cm. wide; calyx tube 7-8 mm. long, the lobes about half as long; 
petals 9-11 mm. long, obtuse; connective of larger anthers long-pro- 
duced; ovary glandular-setulose; capsule about 10 mm. long. 



278 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

P. minor Naud., 181, probably to be expected, has much smaller 
puberulent leaves. Illustrated, Mart. Fl. Bras. 14, pt. 3: pi. 60. 

Peru: (Mathews 1278). Colombia. 

12. PTEROLEPIS (DC.) Miq. 

Usually herbaceous plants with essentially the character of 
Tibouchina, but the Peruvian species marked by the presence of 
stellate or penicillate hairs between (alternating with) the promin- 
ent calyx lobes. Stamens equal or unequal. Anthers subulate, the 
connective short or prolonged at base with 2 anterior rounded tuber- 
cles. The group probably really belongs in Tibouchina, following 
Baillon, but it is a convenient segregate from that more typically 
shrubby and showy-flowered genus. 

Calyx lobes longer than the tube; larger stamens with dilated connec- 
tive P. glomerata. 

Calyx lobes shorter than the tube; larger stamens with filiform 
connective P. pumila. 

Pterolepis glomerata (Rottb.) Miq. Comm. Phyt. 78. 1840; 
190. Rhexia glomerata Rottb. Descr. PL Surin. 8. pi. 4- 1798. 

Stems erect, branched above, several-10 dm. tall, 4-angled and 
more or less densely strigose, with geniculate rigid hairs; leaves ses- 
sile or nearly, oblong-lanceolate, usually 2-4 cm. long, 6-15 mm. 
wide, acute, 3-nerved, somewhat strigose on both sides; flowers 
usually in dense, terminal clusters; calyx pubescent with long 
branched hairs, tube 4-5 mm. long, the lobes often a little longer, 
eglandular; stamens equal; connective short, thick. Variable (prob- 
ably). The Peruvian form would be var. peruviana Cogn. (the type 
from Chiquitos, Bolivia) with pubescent calyx and leaves to 8 mm. 
wide. F.M. Neg. 26146. Illustrated, Bot. Mag. pi. 2838 (as Os- 
beckia glomerata). 

Peru (probably). Warm South America and the West Indies. 

Pterolepis pumila (Bonpl.) Cogn. in Mart. Fl. Bras. 14, pt. 3: 
263. 1885; 185. Rhexia pumila Bonpl. Rhex. pi. 35. 1823. 

Similar to P. glomerata but often simple-stemmed or little 
branched, the leaves shortly petioled and 1-4 cm. long, 4-10 mm. 
wide, the flowers axillary or terminal, single or few; calyx tube about 
2.5 mm. long, hirsute-glandular, the lobes 2 mm. long with a long 
bristle at tip; larger anthers to 2 mm. long, the connective scarcely 
half as long. Resembles most P. trichotoma (Rottb.) Cogn., to be 



FLORA OF PERU 279 

expectedvbut that has anthers 3-5 mm. long, the slender round con- 
nective 2-3 mm. long. For that species, fide Cogniaux, Creuger 
employed the name P. pumila DC.; Linnaea 20: 102. 1847. 

Amazonas: Chachapoyas (Mathews 3220). Brazil to Central 
America. 

13. TIBOUCHINA Aubl. 

Shrubs, trees, or less frequently herbs, often with rather large 
leaves and showy or small 4-8-merous flowers, rarely solitary but 
usually borne in terminal panicles. Stamens subequal or often 
alternately unequal, the connective short- or long-produced and 
variously short- tuberculate or spurred anteriorly or unappendaged. 
Ovary free, setose at tip, usually 4-5-celled, the style usually filiform, 
the stigma punctiform. 

All of the Peruvian species except two have subulate pointed 
anthers. These two belong to the section Purpurella, the anther 
dehiscent by a large pore; Gleason has suggested that the section may 
claim generic rank which would be in logical accord with some other 
genera, so called. The majority of the species listed here, however, 
pertain to the section Diotanthera with 5-merous flowers, ebracteate 
but the pedicels often bracteolate; similar are two species with 8- 
merous flowers. A number of species have bracted flowers, a few 
with calyx lobes soon falling, others with calyx lobes persisting. One 
species, T. gracilis, of this alliance is herbaceous, as one other but 
that has truncate anthers. T. ciliaris (Vent.) Cogn. has been 
accredited to Peru, collected by Dombey; actually the specimen in 
Herb. Jussieu is labeled as from Herb. Dombey, which is another 
matter; however, the species, Colombian, if found, may be known 
from T. stenophylla by the spreading glandular-villous pubescence 
and larger leaves. 

Anthers truncate; herbs or half-shrubs, spreading-hirsute below, or 
if shrubs, petals villous without. 

Shrub; petals villous without T. grossa. 

Herbaceous above; petals glabrous T. clinopodifolia. 

Anthers attenuate; petals glabrous; shrubs, or if half-shrubs, pub- 
escence below not spreading-hirsute (except T. gracilis). 

Flowers 8-merous T. octopetala, T. pukherrima. 

Flowers 4-6-merous. 

Flowers bracted at calyx-base, the calyx (7) 9-10 mm. long, 
or if only 5-6 mm. long, equaled by the lobes. 



280 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Shrubs. 

Stamens glabrous; petioles short T. aspera. 

Stamen connective pilose; petioles often 7-10 mm. long. 

T. Mathaei. 
Herbaceous, at least above, spreading-hirsute below. 

T. gracilis. 

Flowers if basally bracted, caducously; pedicel bracts, if present, 
small but sometimes extending to calyx tube, this then often 
only 3-4 mm. long. 

Flower buds involucrately bracted, the bracts caducous, the 

calyx lobes deciduous. 
Pubescence of bracts and calyx appressed. 
Calyx lobes about half as long as tube; flowers purple. 
Inflorescence several-flowered, exceeding the leaves. 

T. stenocarpa. 
Inflorescence few-flowered, exceeded by the leaves. 

T. coronata. 
Calyx lobes and tube subequal; flowers greenish. 

T. virescens. 
Pubescence of calyx arcuate, spreading T. ochypetala. 

Flower buds ebracteate, in any case not involucrately (cf. 
T. stenocarpa and T. ochypetala with bracts promptly 
caducous) ; calyx lobes persistent (cf . the semi-herbaceous 
T. gracilis). 

Stamens very unequal, the connective of the longer pro- 
duced 2.5-several mm.; pubescence of the branchlets 
widely spreading, those of the calyx, if present, often 
glandular. 

Leaves glabrous and smooth above T. laevis. 

Leaves more or less but always distinctly setulose above. 
Calyx lobes short, inconspicuous. 

Pubescence all eglandular, the branches long-villous. 

T. fulvipilis. 
Pubescence at least in part glandular. 

Calyces and peduncles glandular, the branches 

eglandular. 
Calyx glabrate or sparsely setulose. 

T. pleromoides. 



FLORA OF PERU 281 

Calyx densely hirsutulous T. brevisepala. 

Calyces eglandular, peduncles and branches gland- 
ular T. rhynchantherifolia. 

Calyx lobes prominent, more or less foliose. 

Pubescence short, stiff T. cymosa. 

Pubescence long, rather soft T. laxa. 

Stamens subequal or if more or less unequal the connective 
of the larger produced only 1-2 mm.; pubescence of 
the branchlets appressed or subappressed, strigose- 
setulose except the first three species. 

Branchlets shortly and densely villous-hirsutulous, the 

trichomes spreading-ascending. 

Pubescence in part glandular . . . . T. rhynchantherifolia. 
Pubescence eglandular. 
Upper petioles to 1 cm. long; pubescence villous. 

T. mollis. 

Upper petioles to 2 cm. long; pubescence hirsutulous. 

T. dimorphylla. 

Branchlets strigose-setulose, the trichomes closely appres- 
sed or if rather loose relatively few. 

Pubescence of calyces and branchlets subappressed- 
ascending; calyx lobes flexuous, subulate, as long 
or longer than the tube T. longifolia. 

Pubescence at least of the branchlets closely appressed; 

calyx lobes firm or foliose. 

Calyx lobes conspicuous, foliose, the tube abun- 
dantly strigose or setulose; shrubs. 

Leaves several cm. long T. calycina. 

Leaves 2-2.5 cm. long T. echinata. 

Calyx lobes small or minute, usually much shorter 
than the tube, this often lightly pubescent or 
plants half-shrubs. 
Calyx eglandular; pedicels short. 

Calyx appressed-strigose, longer than broad. 

Leaves rather sparsely asperous-setulose on 
both sides. 

Calyx lobes linear; shrub. . . .T. asperifolia. 
Calyx lobes ovate; half-shrub. T. pentamera. 



282 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves pilose or short-villous, at least beneath. 
Flowers crowded; leaves rounded to the 

scarcely acute base. 
Calyx fusiform, constricted above. 

T. decora. 

Calyx campanulate T. Weberbaueri. 

Flowers laxly borne, few; leaves some- 
what narrowed to acute base. 

T. Solmsii. 
Calyx arcuately setulose, as broad or broader 

than long; half-shrub T. Gay ana. 

Calyx somewhat glandular; pedicels, except in T. 

incarum, elongate. 

Leaves sparsely setulose on both sides, narrow. 
Leaves 5-8 mm. wide; anther-connective con- 
spicuous T. stenophylla. 

Leaves 12-25 mm. wide; anther-connective 

short T. stenopetala. 

Leaves villous-strigillose beneath, ample. 

T. incarum. 

Tibouchina aspera Aubl. PI. Guian. 1: 446. pi. 177. 1775; 238. 

Branchlets densely scaly-scabrous (the scales long) ; petioles 2-5 
mm. long; leaves rigid, 5-nerved, usually about 5 cm. long, 2 cm. 
wide, the midnerve deeply impressed above, subcordate at base, 
acuminate, sparsely, minutely and closely adnate, appressed-set- 
ulose but lustrous above, similarly beneath with the trichomes 
coarser, not so close appressed, especially on the nerves; flowers 5- 
merous, congested, conspicuously bracted, the outer pair of bracts 
5-7 mm., the inner 8-10 mm. long; calyx tube 7-9 mm. long, the 
persistent rigid lobes a little shorter; petals 1.5 cm. long; stamens 
glabrous, the anthers long-attenuate, subulate; style 14-16 mm. long. 
T. Spruceana Cogn., 239, from the upper Amazon, has 3-nerved 
narrow leaves and petals 2-2.5 cm. long. Illustrated (flower), 
Triana, Trans. Linn. Soc. 28: pi. 3. 

San Martin: Tarapoto, Spruce 4585; Williams 7648 (det. Gl.). 
Cerro de Moyobamba, Stuebel (det. Cogn.). To British Guiana. 

Tibouchina asperifolia Cogn. Bot. Jahrb. 42: 137. 1908. 
Branches obscurely tetragonous, shortly appressed-setulose; 
petioles 3-6 mm. long; leaves broadly lanceolate, little narrowed at 



FLORA OF PERU 283 

base, acute or shortly acuminate, entire, rather sparsely appressed- 
setulose on both sides, the larger 5-nerved, 4-7 cm. long, 12-18 mm. 
broad; pedicels 2-5 mm. long; calyx long- and densely appressed- 
setose, eglandular, the tube ovoid-campanulate, 3-4 mm. long, the 
linear, subulate lobes 2-2.5 mm. long; petals purplish, ciliate, 7-8 
mm. long; stamens somewhat unequal, glabrous, the connective 
very shortly produced; style filiform, glabrous, 1 cm. long. Shrub 
5 dm. high. Allied to T. lanceolata Cogn., 262, of Bolivia, Sect. 
Diotanthera. F.M. Neg. 16727. 

Huanuco: South-west of Monzon, prov. Huamalies, Weberbauer 
3357, type. 

Tibouchina brevisepala Cogn. Bot. Jahrb. 42: 135. 1908. 

Shrub 2 meters high, the younger branches rather sparsely setu- 
lose, with short, curved trichomes; petioles slender, 1-2 cm. long; 
leaves oblong-lanceolate, abruptly acutish at base, acute or shortly 
acuminate, subentire, subappressed-setulose above, rather sparsely 
shortly pilose beneath, membranous, the larger 5-nerved, 6-10 cm. 
long, 2-3 cm. broad; pedicels, as the calyx at base, shortly glandular 
hirsute, the former 5-7, the latter 6 mm. long, its lobes scarcely 1 
mm. long; petals lilac, very shortly ciliate, about 12 mm. long; sta- 
mens strongly unequal, glabrous, the connective of the larger pro- 
duced 5-6 mm.; style to 14 mm. long. Section Diotanthera. 

Cuzco: Yanamanchi, Cuzco to Santa Ana, Weberbauer 4965, 
type. 

Tibouchina calycina Cogn. Bot. Jahrb. 42: 135. 1908. 

Branchlets densely appressed-setulose; petioles 8-12 mm. long; 
leaves bright green above, densely bullate-strigose, cinereous beneath 
with a dense short pilosity and markedly foveolate, 7-10 cm. long, 
2.5-3.5 cm. wide, the 7 nerves of the larger prominent; pedicels stout, 
1-2 cm. long, solitary or ternate; calyx densely appressed-setulose, 
8-10 mm. long, the coriaceous lobes to twice as long; petals purple, 
glabrous, 3 cm. long; stamens rather unequal, glabrous, the connec- 
tive produced 1 or 2 mm.; style glabrous, 2 cm. long. The var. 
parvifolia Cogn. has leaves 5-7 cm. long, 1.5-2 cm. broad, the flowers 
smaller. Affine T. laxa Cogn. Section Diotanthera. F.M. Neg. 
16731. 

Puno: Sandia, 2,400 meters, Weberbauer 650, type. Cuzco: 
Santa Ana, Weberbauer 5008, the var. 



284 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Tibouchina clinopodifolia (DC.) Cogn. in Mart. Fl. Bras. 14, 
pt. 3: 411. 1885; 274. Chaetogastra clinopodifolia DC. Prodr. 3: 
133. 1928. 

Stems herbaceous, many, decumbent-ascending, long and sparsely 
hispid, few-several dm. long; petioles to 1 cm. long; leaves oblong- 
ovate, thin, serrate, to 4 cm. long, 1.5-2.5 cm. broad; flowers few, 
the glabrous petals 6-7 mm. long, the calyx tube half as long, the 
lobes shorter, persisting; anthers obliquely truncate at tip, dehiscent 
by a large pore. Section Purpurella. Gleason has suggested that 
this may merit generic recognition. F.M. Neg. 6349. 

Huanuco(?): Haenke. Brazil. 

Tibouchina coronata (Triana) Cogn. in Mart. Fl. Bras. 14, 
pt. 3: 307. 1885; 204. Pleroma coronatum Triana, Trans. Linn. 
Soc. 28: 41. 1871. 

Petioles 1-1.5 cm. long; leaves coriaceous, oblong-lanceolate, 
3-5-nerved, hispid-setulose above, densely tomentose beneath, 5-8 
cm. long, 1.5-2.5 cm. broad; pedicels to 1 cm. long, the two bracts 
very thin, broadly ovate, obtusish, 1 cm. long; petals purple, 2-3 cm. 
long; calyx tube 6-7 mm. long, the broadly triangular, deciduous 
calyx lobes 3-4 mm. long. The calyx pubescence is not rigid. 
Section Involucralis. F.M. Neg. 26104. 

Amazonas: Chachapoyas, Mathews 3211. 

Tibouchina cymosa Cogn. Bot. Jahrb. 42: 134. 1908. 

Branchlets, petioles and short cymes densely pubescent, with 
short, spreading eglandular trichomes (typically stiff-barbellate) ; 
petioles 0.5-1 cm. long; leaves submembranous, broadly ovate, 
rounded at base, acute, minutely serrulate, setulose above, densely 
villous beneath, the larger 7-nerved, 3-5 cm. long, 2-3.5 cm. broad; 
calyx tube 8-10 mm. long, the ovate-lanceolate lobes nearly as long; 
petals violet, shortly ciliolate, 2.5 cm. long; stamens very unequal, 
glabrous, the connective produced, 1.5-2 or 3-4 mm. long; style 
glabrous, 2 cm. long. Allied to T. mollis. Weberbauer 6339 referred 
here by Markgraf but with pilose stems, the trichomes ebarbellate, 
probably is rather T. mollis or T. laxa. Section Diotanthera. F.M. 
Neg. 16740. 

Cajamarca : Below San Miguel, 2,500 meters, Weberbauer 391 1 , type. 

Tibouchina decora Gleason, Bull. Torrey Club 58: 218. 1931. 

Branches, leaves above and leaf nerves beneath densely strigose 

with subulate or papillose-based curved trichomes; petioles 7-11 mm. 



FLORA OF PERU 285 

long; leaves ovate-oblong, entire, rounded at base, acute, 5-nerved, 
3.5-5.5 cm. long, about 2.5 cm. broad or narrower, velutinous above 
and softly villous beneath between the nerves; cymes many-flowered, 
the bracts linear-lanceolate, about 1 cm. long; calyx narrowly cam- 
panulate, densely strigose as the longer pedicel, 7 mm. long, the sepals 
little shorter; stamens glabrous, the anthers subequal, the connec- 
tive shortly produced, anteriorly bilobed; style glabrous, about 
15 mm. long. A shrub to about 2 meters high with magenta flowers, 
the petals freely ciliate, 12 mm. long. Junin specimens det. Gleason. 
Section Diotanthera. 

Ayacucho: In thickets at Aina, 700 meters, (Killip & Smith 
22700, type). Junin: Between 680 and 2,400 meters, (Killip & 
Smith 24230; 24791; 24979). Chanchamayo Valley, Schunke 376; 
381; 346; 1516; 1596. La Merced, 5469. 

Tibouchina dimorphylla Gleason, Bull. Torrey Club 54: 27. 
1927. 

Branchlets sharply angled, thinly spreading-pubescent; petioles 
3-4 cm. long; leaves ovate, acute, minutely denticulate, the largest 
18 cm. long, nearly half as wide, 5-nerved or the upper ones more 
cordate at base, 7-nerved, their petioles 8-15 mm. long, becoming 
bullate and sparsely subappressed-setulose above, thinly and finely 
pubescent beneath, the trichomes on the nerves stouter; inflores- 
cence compact; calyx 5 mm. high, subequaled by the lobes, densely 
hispid, with incurved-ascending trichomes; petals 10-12 mm. long, 
ciliate; connective prolonged 1 mm. in the larger stamens, extended 
into 2 appendages; style 10 mm. long. The leaves on the flowering 
branches are 6.5-10 cm. long. Section Diotanthera. 

Cuzco: Pillahuata, Pennell 14109, type; 14092. Without locality 
on Field Museum sheet, Weberbauer 6926 (det. Gl.). 

Tibouchina echinata (R. & P.) Cogn. Melast. 266. 1891. 
Rhexia echinata R. & P. Fl. Peruv. 3: 85. pi. 319. 1802. 

Branchlets closely appressed-setulose; petioles 2-5 mm. long; 
leaves rigid, 5-nerved, mostly 2-3 cm. long, 7-9 mm. wide, becoming 
harshly appressed-callous-echinate above, appressed-setulose on the 
nerves and hirsutulous between the nerves beneath; flowers nodding; 
petals 2 cm. long or longer; calyx tube in fruit densely bullate- 
echinate, the little shorter lanceolate lobes appressed-setulose with- 
out, glabrous within, about 1 cm. long; stamens subequal, glabrous, 
the connective shortly produced. Section Diotanthera. F.M. Neg. 
26105. 



286 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Lima: Nee. Huanuco: Muria, Ruiz & Pavon, type. Huacachi, 
near Mufia, 3891. Between Rio Maranon and Rio Monzon (Weber- 
bauer 3369a; 253). 

Tibouchina fulvipilis Cogn. Bot. Jahrb. 50: Beibl. Ill: 31. 1913. 

Younger branchlets, petioles and peduncles clothed with spread- 
ing, rigid trichomes; petioles 1-2 cm. long; leaves ovate-lanceolate, 
obsoletely crenulate, submembranous, 5-nerved, sparsely hirtellous 
above and pilose beneath, 6-9 cm. long, 2-4 cm. broad; flowers 
cymose on pedicels 5-15 mm. long; petals 1.5 cm. long; calyx 
eglandular, pubescent only at base, 5-6 mm. long, the lobes 1-1.5 
mm. long; stamens unequal, glabrous, the anther-connective 1.5 or 
4-5 mm. produced; style 11 mm. long. Var. scrobiculata has leaves 
bullate above, scrobiculate beneath. Section Diotanthera. 

Ayacucho: Prov. Huanta, 2,800 meters, erect shrub, Weberbauer 
5619, type; 5625 (var.). 

Tibouchina Gayana (Naud.) Cogn. Melast. 253. 1891. Micran- 
thella Gayana Naud. Ann. Sci. Nat. se>. 3. 13: 350. 1849. 

Branches densely or sparsely appressed-setulose; leaves ovate, 
subacute at base, shortly acuminate, 5 (-T)-nerved, thin, finely 
appressed-hispid above, less so beneath, 4-6 cm. long, about 2 cm. 
broad; calyx tube subglobose, 4 mm. long, the lobes 2-3 mm. long, 
densely hirsute; petals nearly 1 cm. long; connective shortly produced, 
style 8-9 mm. long. A slender, branching plant, more or less woody 
at base. Section Diotanthera. Illustrated, Bot. Mag. pi. 6345. 
F.M. Neg. 36163. 

Cuzco: Gay, type. Puno: San Gaban, Lechler 2182. Valley of 
Sandia, 2,300 meters, Weberbauer 512; 237. 

Tibouchina gracilis (Kunth) Cogn. in Mart. Fl. Bras. 14, 
pt. 3: 386. 1885; 243. Rhexia gracilis Bonpl. Rhex. 138. pi. 52. 1823. 

Simple, erect, pilose-setose herb (often lignescent at base) with 
lanceolate or ovate-lanceolate 5-nerved leaves often about 7 cm. 
long, 1 or 2 cm. wide, subsessile or on petioles a few mm. long; 
flowers glomerate, shortly bracteolate, the petals 1-2 cm. long; 
calyx tube 5-6 mm. long, the setulose lobes as long or longer; stamens 
glabrous; style to 2 cm. long. Highly variable in size and character 
of foliage. Section Simplicicaulis. Illustrated, Bot. Mag. pi. 3481. 

San Martin: Tarapoto, Mathews 3218; Spruce. Paraguay to the 
West Indies. 



FLORA OF PERU 287 

Tibouchina grossa (L. f.) Cogn. in Mart. Fl. Bras. 14, pt. 3: 
297. 1885; 274. Melastoma grossa L. f. Suppl. 236. 1781. 

Densely branched shrub 2-4 meters high, with ovate, obtusish 
5-nerved leaves harshly strigose above, villous or hispidulous beneath, 
2-4 cm. long, half as wide, on petioles 4-8 mm. long; cymes few- 
flowered; calyx tube 6-7 mm. long, densely appressed-setose, equaled 
by the triangular-lanceolate, acuminate lobes; petals 2.5-3 cm. long, 
villous without; style stout, to 3.5 cm. long. Section Purpurella. 
Illustrated, Bonpl. Rhex. pi. 1. 

Amazonas: Chachapoyas, Mathews (det. Cogn.). Ecuador; 
Colombia. 

Tibouchina incarum Gleason, Amer. Jour. Bot. 26: 634. 1939. 

Widely branched shrub to 1.5 meters high, the 4-angled stems 
closely strigose; petioles slender, 1-2 cm. long; leaves thin, closely 
substrigose above, paler and softly villous on the veins and veinlets 
beneath, minutely serrulate, ovate-lanceolate to 11 cm. long, 4.5 cm. 
wide, acuminate, abruptly cuneate at base, 7-plinerved; panicles 
ample, to 2.5 dm. long, glandular-hirsute; pedicels 2 mm. long; 
flowers 6-merous, the broadly obovate, bright pink petals 8 mm. 
long and nearly as wide; sepals recurved, 2.6 mm. long; stamens 
nearly isomorphic but the filaments of two lengths, 4.7 or 3.7 mm. ; 
ovary densely bristly apically. Allied to T. Gayana with smaller 
leaves, merely acute at base, larger sepals and the hypanthium not 
glandular. 

Cuzco: Machu Picchu (Balls B6817, type); grassy hillside (Staf- 
ford 772; 791}; open brush formations, 2,200 meters (West 6456). 

Tibouchina laevis Cogn. Bot. Jahrb. 42: 136. 1908. 

Tetragonous branchlets annulate-enlarged at nodes, glabrous as 
the petioles, peduncles, calyces and leaves above; petioles strongly 
flattened, 3-4 mm. broad; leaves rigid, narrowly ovate, rounded at 
base, acute, minutely setulose-dentate, scarcely setulose beneath, 
5-nerved, 4-5 cm. long, half as broad; pedicels 5-10 mm. long; calyx 
tube narrowly campanulate-ovoid, acute at base, 5-6 mm. long, the 
narrowly triangular lobes half as long; petals yellow, glabrous, 8-9 
mm. long; stamens very unequal, glabrous, the connective of the 
larger produced to 3 mm.; style glabrous, 11 mm. long. Allied to 
T. pleromoides. Section Diotanthera. 

Puno: Between Chunchusmayo and Sandia, Weberbauer 1139, 
type. 



288 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Tibouchina laxa (Desr.) Cogn. Bull. Acad. Belg. se>. 3. 14: 933. 
1887; 250. Melastoma laxa Desr. in Lam. Encycl. 4: 41. 1797. 

Branches hirsute, the trichomes spreading; petioles 5-10 mm. 
long; leaves ovate, somewhat cordate, 7-nerved, only sparsely set- 
ulose on either side, foveolate beneath, 3-5 cm. long, 2-3.5 cm. 
broad; flowers few, rather crowded, the purple or white petals 2 cm. 
long; calyx tube 7-8 mm. long, equaled or exceeded by the lobes; 
stamens glabrous; connective moderately produced; style 1.5 cm. 
long. Var. villosissima Cogn. has broadly ovate leaves, densely 
softly pilose both sides, the branches and calyces villous. Section 
Diotanthera. Illustrated, Bot. Mag. pi. 5629. 

Piura: Near San Felipe, Bonpland. Amazonas: Chachapoyas, 
Williams 7573 (det. Gl.). Without locality, Mathews 1275; J. Jus- 
sieu; Ruiz & Pavon, this the var. (det. Markgr.). Cajamarca: 
Shrubby hillsides near Socota, 2,600 meters, Stork & Horton 10083. 
Ecuador. 

Tibouchina longifolia (Vahl) Baill. Adansonia 12: 74. 1877; 
264. Rhexia longifolia Vahl, Eclog. Amer. 1: 39. 1796 et Icon. PL 
Amer. 2: pi. 15. 1799. R. flexuosa R. & P. Fl. Peruv. 3: 85. pi. 
320. 1802. 

More or less suffrutescent, often only a few dm. high, the branches 
more or less densely spreading-setulose; leaves rather narrowly 
lanceolate, usually less than 1 dm. long, 1-2.5 cm. wide, somewhat 
appressed-pilose, panicle leafy; calyx tube 3 mm. long, the flexuous 
lobes as long or longer; petals white, 5-6 mm. long, equaled by the 
style; stamens glabrous, subequal, the connective scarcely produced. 
A form with glandular calyx is var. simulans Macbr. Field Mus. 
Bot. 4: 174. 1929. Section Diotanthera. F.M. Neg. 21206. Illus- 
trated, DC. M6m. Melast. pi. 8. 

Huanuco: Rio Huallaga Canon, 4237. Churubamba region, 
Mexia 8212 (det. GL). Pozuzo, 4556, the var.; Ruiz & Pavdn. 
Piedra Grande', 3683. Pampayacu, Kanehira 81; Poeppig 1002. 
Casapi, Mathews 1721 (Fielding 1721). Cuchero, Dombey; Ruiz & 
Pavdn. Chinchao and Muna, Ruiz & Pavdn. Junin: La Merced, 
5300, the var.; Killip & Smith 23567; 23789; 24968; 25197; 25278 
(all det. Gl.). Ayacucho: Killip & Smith 22499; 23075 (det. GL). 
San Martin: Tarapoto, Spruce; Williams 6364; 7151 (det. GL). 
Loreto: Puerto Arturo, Killip & Smith 27848; 29216 (det. GL). 
Pongo de Manseriche, Mexia 6207. Yurimaguas, Williams 4610. 
Cuzco: Valle de Vilcanota, Mexia 8091 (det. GL). Paucartambo, 
2,700 meters, Vargas 94. To Mexico and the West Indies. "Mullaca." 



FLORA OF PERU 289 

Tibouchina Mathaei Cogn. in Mart. Fl. Bras. 14, pt. 3: 380. 
1885; 241. Lasiandra lepidota Naud. Ann. Sci. Nat. seY. 3. 13: 155. 
1849, not T. lepidota (Bonpl.) Baill. 

Branchlets and leaves beneath clothed with narrowly oblong 
squamate fringed trichomes; petioles 7-10 mm. long; leaves oblong- 
ovate, to 1 dm. long and about half as wide, often smaller, 5-nerved, 
densely appressed and finely strigillose-setulose above, appressed- 
squamulose or lepidote beneath, the margins ciliate, with coarser 
trichomes similar to those on the nerves; calyx tube 8-9 mm. long, 
the lobes half as long, densely squamate with elongate, lanceolate 
scales; petals about 1.5 cm. long; connective barbate at juncture 
with filaments. Simulates T. aspera. Section Barbigeri. F.M. Negs. 
26108; 36167. 

San Martin: Moyobamba, Mathews 1267, type; Weberbauer M93; 
290. Costa Rica. 

Tibouchina mollis (Bonpl.) Cogn. in Mart. Fl. Bras. 14, pt. 3: 
349. 1885; 249. Rhexia mollis Bonpl. Rhex. 50. pi. 19. 1823. 

Branchlets as many as the leaves, densely appressed or sub- 
appressed, pilose-hirsute; petioles to 1 cm. long; leaves rather thin, 
5-nerved with 2 outer, fainter nerves, ovate, 5-9 cm. long, 2.5-4 cm. 
wide; calyx densely hirsute, 5-6 mm. long, equaled by the linear 
subulate lobes; stamens subequal, glabrous, the narrow anthers 
obtuse; petals about 12 mm. long; style 8-10 mm. long. Section 
Diotanthera. The leaves are soft to the touch beneath, the calyces 
soon globose. Cf. T. Rusbyi. F.M. Neg. 36168. 

Peru(?): Without locality, Rivero (MacLean).- Ayacucho: Prov. 
Huanta (Weberbauer 564-2, det. Cogn.). Ecuador; Colombia. 

Tibouchina ochypetala (R. & P.) Baill. Adansonia 12: 75. 
1877; 213. Rhexia ochypetala R. & P. Fl. Peruv. 3: 86. pi. 321. 1802. 
T. Tessmannii Markgr. Notizbl. Bot. Gart. Berlin 9: 1139. 1927. 

Branches more or less acutely tetragonous, closely appressed- 
strigose with rigid trichomes; petioles 0.5-1.5 cm. long; leaves 
rather rigid, oblong-lanceolate, 6-8 cm. long, 1-2 cm. wide, rather 
sparsely appressed-hispidulous above, more densely appressed- 
pubescent beneath; panicles few-flowered, appressed-setulose; bracts 
ovate, 1-1.5 cm. long; petals 2-3 cm. long; calyx tube stiffly setose- 
pubescent, equaling the oblong lobes, these 5-6 mm. long; larger 
anthers 1 cm. long, the smaller about 6 mm. long, the connective 
anteriorly glandular-pilose; filaments more or less setose; style gla- 
brous except at base. T. Tessmannii Markgr., allied by the author 



290 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to T. nervulosa Cogn., a species with glabrous filaments and style, 
does not seem to me to be separable. A showy shrub usually 2-4 
meters high, but often rather open in growth; a 7-meter tree (Mexia). 
Section Involucrales. F.M. Neg. 38258. 

Huanuco: Vilcabamba, Hacienda on Rio Chinchao, 5006; 16800 
(Tessmannii). Puyash, Sawada 95 (det. Gl.). Churubamba, Mexia 
8204 (det. GL). Chinchao, Cuchero, Pillao, Ruiz & Pawn, type; 
Poeppig 100; 1003. Between Rio Maranon and Monzon, (Weber- 
bauer, 256). Junin: La Merced, 5550. Hacienda Schunke, 5726. 
San Martin: Valley of the Monzon, 1,900 meters, (Weberbauer, 284). 
Tarapoto, Spruce 4000; Williams 5953; 6385; 6492 (det. GL). 
Moyobamba, Mathews 1272 (Fielding). Amazonas: Chachapoyas, 
Mathews 3214; Weberbauer 4363 (det. Cogn.). Loreto: Bristol, near 
Mouth of Rio Pastaza, on Rio Maranon, Tessmann 3788, type, T. 
Tessmannii. Cerro de Escaler, 1,300 meters, Vie 6364- Pumayacu, 
King 3137 (det. GL). Also Killip & Smith 28618; 29210 (det. GL, 
T. Tessmannii). Ecuador. "Chinchincca," "pichuquita," "machu- 
sacha pichirina," "Santa Rosa sisa." 

Tibouchina octopetala Cogn. Bull. Torrey Club 17: 57. 
1890; 273. 

Densely but rather shortly hirsute shrub or the leaves shortly 
villous beneath; petioles 2-3 cm. long; leaves 5-nerved with also a 
pair of faint marginal ones, rounded at base, acute, entire, sub- 
membranous, about 1 dm. long or longer, 5-7 cm. broad; pedicels 
bibracteate above the middle, the obovate purplish bracts shortly 
villous, 1 cm. long; calyx tube about 8 mm. long, densely strigillose- 
setose, equaled by the narrow lobes; petals purple-violet, 12-15 mm. 
long; stamens glabrous, the longer filaments and anthers 7.5-8 mm. 
long; style hirsute below, 1 cm. long. Section Octomeris, to which 
also only belongs T. pukherrima. 

Cuzco: Near Rio Yanamayo, Pennell 14039. Puno: Mountains 
of Yuncacoya, Weberbauer 1140; 242. Bolivia. 

Tibouchina pentamera (Ule) Macbr., comb. nov. Comolia 
pentamera Ule, Notizbl. Bot. Gart. Berlin 6: 352. 1915. 

Stems slender, sparsely subappressed-strigose; leaves narrowly 
ovate, long-acuminate, shortly petioled, thin, 3-nerved, sparsely 
appressed-setose, 2.5-3.5 cm. long, 11 mm. wide or narrower; calyx 
tube densely strigose-setose, 4-5 mm. long, 3-4 mm. broad, the ovate 
lobes densely setose on the margins, 3.5 mm. long; petals 5, white, 
narrowed below, retuse, 7 mm. long; stamens subequal, 5 and 7 mm. 



FLORA OF PERU 291 

long, the short connective 2-lobed; ovary 5-celled, glabrous (fide 
Ule), style 8 mm. long; seeds cochleate. A half-shrub 1-4 dm. 
high, little branched, subdecumbent at the woody base. Gleason 
in Herb. Dahlem has written on the type sheet "Why is this a 
Comolia?" but unfortunately he did not answer his query; when 
the type was photographed I noted in my manuscript "probably a 
Tibouchina," but did not study it further. From description and 
photograph it seems to be allied to T. lanceolate, Cogn., 262, Bolivian, 
or to T. stenopetala. The ovary, if actually glabrous, is an aberrant 
character for Tibouchina, but the ovary may be either glabrous or 
setulose in other groups so it is not in itself a generic character. 
In the glabrous ovary T, pentamera could be traced to Comolia, from 
which its facies as well as its 5-merous flowers exclude it; it is distinct 
from Acisanthera and Pterolepis in the character of its pubescence. 
F.M. Neg. 16844. 

Loreto: Cerro de Escaler, Ule 6742, type. 

Tibouchina pleromoides (Naud.) Macbr. Field Mus. Bot. 4: 
174. 1929. Lasiandra pleromoides Naud. Ann. Sci. Nat. se"r. 3. 13: 
131. 1849. Pleroma maurocarpum Triana, Trans. Linn. Soc. 28: 47. 
1871. Tibouchina maurocarpa Cogn. Melast. 260. 1891. 

Younger branches very shortly appressed-setulose; petioles 1-2 
cm. long; leaves ovate-oblong, obscurely crenulate, 5-nerved, sparsely 
asperous-strigillose above, foveolate and sparsely strigillose beneath 
on the nerves, to about 1 dm. long and a third as broad; panicles 
few-flowered; petals red, ciliate, 1 cm. long; calyx tube 5-6 mm. 
long, glabrous, the minutely ciliate teeth 2-2.5 mm. long; stamens 
glabrous, very unequal, the connective of the larger long-produced. 
The leaves often only 5-6 cm. long, 2-2.5 cm. wide, are bullate above, 
correspondingly foveolate beneath. Section Diotanthera. F.M. 
Neg. 32309. 

Cuzco: Gay. Junin: Pariahuanca, Mathews 874- Apurimac: 
Abancay, 3,000 meters, Vargas 420. 

Tibouchina pulcherrima Gleason, Bull. Torrey Club 54: 
26. 1927. 

Closely allied to T. octopetala, but with conspicuously purple 
pubescence, especially on the nerves of the leaves and the more open 
inflorescence; leaves narrowly ovate-oblong, 15-18 cm. long, 6-7.5 
cm. wide, setose-ciliate, the upper surface densely conic-tuberculate, 
setose, the lower ashy with a spreading pubescence; pedicels 10-15 



292 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

mm. long; longer filaments 7.5-8 mm. long, the appendages of the 
anther connective erect or nearly, 1.3 mm. long. Section Octomeris. 
Cuzco: Cerro de Cusilluyoc, Pennell 13971, type. 

Tibouchina rhynchantherifolia Cogn. Bot. Jahrb. 42: 
134. 1908. 

Younger branches, petioles and peduncles shortly glandular- 
pilose with spreading trichomes; petioles 1-2 cm. long; leaves 
membranous, puberulent-pilose on both sides, narrowly ovate, 
rounded at base, shortly acuminate, the larger 7-nerved, 7-11 cm. 
long, 3-4.5 cm. broad; panicles broadly pyramidal; calyx densely 
pilose but eglandular, the tube 4 mm. long, the lobes scarcely half 
as long; petals white, shortly glandular-ciliate, 5 mm. long; stamens 
strongly unequal, glabrous, the anthers 2.5 or about 4 mm. long; 
style 7-8 mm. long. Affine T. Rusbyi, Section Diotanthera. F.M. 
Neg. 16185. 

Puno: Between Sandia and Chunchusmayo, 1,500 meters, Weber- 
bauer 1062, type. 

Tibouchina Solmsii Cogn. Melast. 250. 1891. 

Branches slender, nearly terete, the younger densely appressed- 
setulose; petioles 3-7 mm. long; leaves membranous, ovate-oblong, 
minutely serrulate, pilose on both sides, 2-3.5 cm. long, 9-17 mm. 
broad; flowers few or solitary; calyx tube setulose (the trichomes 
tuberculate at base), 3.5-4 mm. long, the long ciliate lobes 2 mm. 
long; petals ciliolate, 7-8 mm. long; stamens glabrous, subequal, 
the small anthers apically attenuate; style 6-7 mm. long. Two 
collections are referred here with doubt; several Bolivian forms are 
too similar; the genus needs revision. Section Diotanthera. 

Puno: San Gaban and Tatanara, Lechler 2179; 2555, types. 
Huanuco: Villcabamba on Rio Chinchao, 5195(1). Cuzco: Cerro de 
Cusilluyoc, rocky forest, flowers magenta, Pennell 13992(1). 

Tibouchina stenocarpa (DC.) Cogn. in Mart. Fl. Bras. 14, pt. 
3: 344. 1885; 224. Lasiandra stenocarpa DC. Prodr. 3: 130. 1828. 

Branches appressed-setulose; petioles 0.5-2 cm. long; leaves 
rigid-coriaceous, oblong, 8-12 cm. long, 2-4 cm. broad, 5-nerved, 
green but shortly strigose above, densely appressed-sericeous villous 
beneath; panicles ample; pedicel bracts 7-9 mm. long; petals 2-3 cm. 
long; calyx tube 7-9 mm. long, the lobes about half as long; fila- 
ments conspicuously villous-lanate; style glabrous, 2-2.5 cm. long, 
the ovoid capsule 8 mm. long. A variety boliviensis Cogn. has the 
style pilose below. The single collection may be T. coronata, which 



FLORA OF PERU 293 

seems to be similar, although Cogniaux, following Triana, assigned 

the species to different groups. Section Pleroma. F.M. Neg. 26115. 

Puno: Valley of the Sandia (Weberbauer 1123; 278). Brazil. 

Tibouchina stenopetala (Griseb.) Cogn. Melast. 262. 1891. 
Micranthella stenopetala Griseb. in syn. ex Triana, Trans. Linn. 
Soc. 28: 46. 1871. 

Branches slender, elongate, rather acutely angled, very slightly 
setulose; leaves thin, sparsely appressed-setulose above and beneath 
on the nerves only, 5-6 cm. long, half as wide, obscurely undulate- 
crenulate; panicle diffuse, few-flowered; petals 7 mm. long, long- 
cuneate at base; calyx slightly glandular-hirsute, the tube 4 mm. 
long, the teeth 1 mm. long; stamens glabrous, unequal, the connec- 
tive short; style 5-7 mm. long. A fragile membranous-leaved 
species, the fruiting calyx sharply 10-ribbed. Section Diotanthera. 
F.M. Neg. 6357. 

Puno: San Gaban, Lechler 2271, type. 

Tibouchina stenophylla Cogn. Bull. Torrey Club 17: 56. 
1890; 257. 

Slender-stemmed, the branches obtusely angled, weak, finely 
appressed-setulose; petioles 2-7 mm. long; leaves thin, narrowly 
lanceolate, entire, 3-nerved, subacute, 3-6 cm. long, 5-8 mm. broad; 
flowers few on filiform pedicels 5-8 mm. long; calyx tube 4 mm. 
long, the ciliate lobes 1 mm. long; stamens glabrous, strongly un- 
equal, the connective of the larger long-produced; style 8-9 mm. 
long; capsule ovoid, 5 mm. long. Section Diotanthera. 

Loreto: Mouth of Santiago, Puerto Mele"ndez, Tessmann 4795 
(det. Markgr.). Bolivia. 

Tibouchina virescens Cogn. Bot. Jahrb. 42: 133. 1908. 

Allied to T. coronata; branches slender, obtusely angled, the old 
densely verrucose, the younger shortly and densely appressed- 
setulose; petioles 4-6 mm. long; leaves deep green above but ap- 
pressed-strigillose, ashy-villous beneath, 4-6.5 cm. long, 10-17 mm. 
broad; bracts 4, rather coriaceous, acute, 7-8 mm. long; calyx 
densely appressed, long-setulose; lobes equaling the calyx tube, 
7 mm. long. Section Involucrales. F.M. Neg. 16807. 

Junin: East of Palca, 3,100 meters, Weberbauer 2475, type. 

Tibouchina Weberbaueri Cogn. Bot. Jahrb. 42: 136. 1908. 
A 2-meter shrub with slender, glabrous branches; petioles, 
peduncles, calyces and leaves above shortly but densely appressed- 



294 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

setulose; petioles 0.5-1 cm. long; leaves rather firm, green above, 
ashy-appressed-villous beneath, oblong or ovate-oblong, rounded at 
base, acute, 5-nerved, entire, 4-6 cm. long, 12-22 mm. broad; 
panicles short, leafy, rather densely flowered; pedicels 2-5 mm. long; 
calyx tube 4 mm. long, eglandular, the narrow, acute lobes about 
half as long; stamens very unequal, glabrous, the connective produced 
to 1.5 mm. on the larger anthers, these 5.5 mm. long; style filiform, 
glabrous, 12-14 mm. long. Allied to T. Orbignyana. Section 
Diotanthera. F.M. Neg. 16808. 

Cuzco: Near Santa Ana, 1,700 meters, Weberbauer 5002, type. 
Spreading shrub 1.5 meters high, with deep cerise flowers, 1,750 
meters, Rio Sambray, Mexia 8056 (det. Gl.). Rio Chaupimayo, 
Soukup 569. 

14. ACISANTHERA P. Br. 

Herbaceous or nearly, usually branched, the leaves small, the 
flowers 4-5-merous, axillary or terminal, few in short cymes, some- 
times solitary. Stamens dimorphic, the anthers oblong to subulate, 
obtuse to rostrate; the connective of the larger prolonged and with 
2 broad anterior lobes, that of the smaller similar but the lobes smaller 
or reduced to tubercles. Ovary free, 2-4-celled. Style slender, 
often bent. Seeds reniform or cochleate, minutely pitted. 

Acisanthera quadrata Pers. Syn. PI. 1: 477. 1805. 

Usually well-branched annual with narrowly 4-winged or -angled 
smooth stems or these remotely glandular pubescent; leaves sub- 
sessile, ovate-oblong, finely denticulate, glabrous or glandular, 1-2 cm. 
long, half as wide; flowers axillary, solitary, on pedicels 1-3 mm. 
long, 4-merous; calyx campanulate, 2 mm. long, the triangular sepals 
about as long; petals roseate, 5-6 mm. long; connective of larger 
stamens and anthers subequal, the lobes 1 mm. long, the connective 
of the smaller basally tubercled. Semisucculent plant of moist 
places. 

Peru(?): Pav6n(f), without locality (det. Cogn.). West Indies; 
Central America south to Panama. 

15. ACIOTIS D. Don 

Glabrous or sparsely pilose with 4-angled or -winged stems, 
more or less woody and often sprawling or herbaceous, the herbs 
usually delicate or succulent, their leaves membranous. Flowers 
small or nearly minute, sessile or subsessile, bracteolate, 4-merous, 



FLORA OF PERU 295 

borne in terminal panicles. Stamens equal or nearly, the connective 
shortly if at all prolonged, unappendaged or merely obscurely lobed 
or enlarged. Ovary 2-celled, the stigma truncate or capitate, the 
capsule thin-walled. The leaves of some species are used for greens. 

Anthers roundish, the connective little if at all produced; petioles 

rarely 1 cm. long except in A. aequatorialis. 
Petioles 1.5-3.5 cm. long; branches angled, glandular-pilose. 

A. aequatorialis. 
Petioles 3-8 (-10) mm. long. 
Annual herbs. 

Leaves ovate, 1-3 cm. wide A. dichotoma. 

Leaves oblong-lanceolate, rarely 1 cm. wide . . A. sphaerantha. 
Suffrutescent, the upper winged branches reddish-villous. 

A. brachybotrya. 

Anthers oblongish or linear, the connective somewhat produced; 
petioles often about 1 cm. long to much longer except in 
A. cor data. 
Leaves ovate-lanceolate, many more than twice longer than 

wide (sometimes cordate at base). 

Flowers pedicellate; leaves 5-nerved, glabrous. A. purpurascens. 
Flowers sessile; leaves (5) 7-nerved, at least with a few hairs 

above. 

Petals 4-5 mm. long; acute and terminating in a bristle. 
Petioles slender, sparsely pilose; petals shortly setose at tip. 

Stems terete A. longifolia. 

Stems angled A. purpurascens. 

Petioles stout, densely hirsutulous; petals long-setose at 

tip A. rubricaulis. 

Petals 7 mm. long, aristate A. aristata. 

Leaves ovate, somewhat cordate and scarcely twice longer than 

wide. 
Plants suffrutescent with reddish hirsute winged stems. 

A. caulialata. 

Plants annual or woody only below, the stems often angled. 

Leaves 7-nerved, the outer nerves faint; petioles 1-2 cm. long. 

Panicle open, about as long as wide, the branches subsimple. 

A. polystachya. 

Panicle oblongish, dichotomously branched A. laxa. 

Leaves 5-nerved; petioles rarely 1 cm. long A. cordata. 



296 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Aciotis aequatorialis Cogn. in Mart. Fl. Bras. 14, pt. 3: 464. 
pi. 105. 1885; 301. 

Herb 1-4 dm. high with somewhat glandular pilose, acutely 
angled branches and 5-nerved pilose leaves, these ovate, often some- 
what cordate, acute, 2-5 cm. long, 1.5-3.5 cm. wide; panicles small, 
dichotomously branched, subcorymbose; pedicels 0.5-1 mm. long; 
calyx 2 mm. long; petals 3-4 mm. long, acutish, the style 3 mm. 
long; anthers ovoid-oblong. 

Loreto: Sarayacu (Huber 1497). Yurimaguas, Poeppig. Brazil. 

Aciotis aristata Ule, Notizbl. Bot. Gart. Berlin 6: 352. 1915. 

Branches, these 4-sided but scarcely winged, petioles and panicles 
densely long-pilose; leaves oblong-lanceolate, 9-13 cm. long, 3.5- 
4.5 cm. wide, long-acuminate, 5-7-nerved, sparsely setose-pilose on 
both sides; calyx 5 mm. long, smooth; petals long-aristate at tip; 
connective long-produced, the anthers oblong-linear; capsule 4 mm. 
thick. Except for the larger flower, it seems, from character, to 
be nearly A. rubricaulis. F.M. Neg. 16821. 

Rio Acre: Ule 9667, type. 

Aciotis brachybotrya (DC.) Triana, Trans. Linn. Soc. 28: 51. 
1871; 301. Spennera brachybotrya DC. Prodr. 3: 115. 1828. 

Branches distinctly 4-winged and rather densely villous with 
long, usually reddish, crinkly trichomes, the small leaves typically 
glabrous and about 2 cm. long and wide or 3-4 cm. long and some- 
what pilose, subrotund at base, acute; panicles small, dichotomously 
branched; pedicels scarcely 1 mm. long, about equaled by the 
bractlets; calyx and acute petals 2 mm. long, the style 1 mm. longer; 
capsule 2.5 mm. thick. 

Junin: Pichis Trail, Kittip & Smith 25892 (det. Gl.). Ecuador; 
Brazil. 

Aciotis caulialata (R. & P.) Triana, Trans. Linn. Soc. 28: 
52. 1871; 308. Arthrostemma caulialata R. & P. Fl. Peruv. 4: pi. 327. 
1802. Spennera panicularis Naud. Ann. Sci. Nat. se>. 3. 14: 148. 
pi. 5. 1850. S. caulialata Naud. I.e. 149. 

Stems distinctly suffrutescent below, conspicuously 4-winged 
above, hirsutulous-furfuraceous, the trichomes, as those of the long 
(2-3.5 cm.) petioles red, firm; leaves ovate, often 3-5 cm. long, 
2-3 cm. wide, rather evenly and densely pilose on both sides with 
long hairs, especially beneath on the 5 nerves; panicles pyramidal, 
about 1 dm. wide at base and half again as long; petals ovate, 



FLORA OF PERU 297 

acuminate; anther connective long-produced. The branchlets with 
the wings are 4-5 mm. wide. F.M. Neg. 29489. 

Lima: Gay. San Martin: Moyobamba, Mathews 1279; Lechler 
2186. Huanuco: Pueblo Nuevo, Ruiz & Pavon, type. 

Aciotis cordata (Veil.) Macbr., comb. nov. Melastoma cordata 
Veil. Fl. Flum. 178. 1825; Icones 4: pi. 114. 1827. Spennera dyso- 
phylla Benth. in Hook. Journ. Bot. 2: 296. 1840. Aciotis dysophylla 
(Benth.) Triana, Trans. Linn. Soc. 28: 52. 1871; 310. 

Annual herb with glabrate tetragonous stems and sparsely ap- 
pressed, pilose, 5-nerved, ovate leaves, these to 4 cm. long, 2.5 cm. 
wide or smaller, often cordate at base; panicles to about 3 cm. long 
and as wide at base, the minutely glandular, campanulate calyx 
1.5 mm. long, its pedicel scarcely 1 mm. long; petals 2 mm. long, 
oblong-lanceolate; anthers oblong, the connective short; capsule 
ovoid, 2 mm. long. Lacks the glandulosity of the similar and to 
be expected A. viscosa (Bonpl.) Triana, with ovoid calyx and petals 
3 mm. long. Cogniaux questioned the identity of the Vellozo plant 
but apparently knew only the plate; Bentham set the precedent of 
accepting identifiable Vellozo species. Illustrated, Mart. Fl. Bras. 
14, pt. 3: pi. 106. 

Amazonas: Nazareth, Osgood 16. Loreto: Iquitos, Killip & 
Smith 26888; King 1188 (det. Gl.); Williams 1426; 1447. Pebas, 
Williams 1710; 1712. Rio Mazan, Schunke 213. To Ecuador and 
British Guiana. 

Aciotis dichotoma (Benth.) Cogn. in Mart. Fl. Bras. 14, pt. 3: 
460. pi. 104. 1885; 299. Spennera dichotoma Benth. in Hook. Journ. 
Bot. 2: 295. 1840. 

Herbaceous, rarely more than a dm. or two high, the narrowly 
4-winged branches glandular-pilose; petioles typically rarely more 
than 5 mm. long; leaves ovate-oblong, obtusish, soft, finely crenulate- 
serrulate, glabrous both sides, 3-nerved, 2-3 cm. long, 1-1.5 cm. 
wide; panicles small; pedicels scarcely 0.5 mm. long; calyx 1.5 mm. 
long; petals acutish, reddish, 1.5-2 mm. long; anthers orbicular to 
broadly ovate, all alike; style capitate at tip, 1 mm. long; capsule 
1.5 mm. thick. F.M. Neg. 16822. 

Loreto: Upper Rio Blanco, Tessmann 3032 (det. Markgr.). To 
the Guianas. 

Aciotis laxa [Richard] Cogn. in Mart. Fl. Bras. 14, pt. 3: 476. 
pi. 108. 1885; 309. Spennera laxa [Richard] DC. Prodr. 3: 116. 



298 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1828. A. indecora (Bonpl.) Triana, Trans. Linn. Soc. 28: 52. 1871, 
fide Gleason. Rhexia indecora Bonpl. Rhex. 131. pi. 49. 1823. 

Suffrutescent, robust, sulcately stemmed plant, sometimes a 
meter long; petioles 1-2 cm. long or longer; leaves 5-7-nerved, long- 
acuminate, cordate-ovate, thin, glabrate, to 6 cm. long, 4 cm. wide 
or larger; panicles to 8 cm. long or longer, but often only about 
3 cm. wide; pedicels obsolete; bractlets subulate; calyx 2.5 mm. 
long, the tube campanulate-ovoid ; petals 2.5 mm. long, the style 
twice as long; capsule ovoid, 3.5 mm. long. A form with petioles 
to 5 cm. long, the scarcely cordate leaves 9-11 cm. long, 6-7 cm. 
wide, was named var. macrophylla Cogn. 

Loreto: Yurimaguas, Poeppig. Manfinfa, Upper Nanay, Williams 
1179; 1180 (det. Gl.). Puno: San Gaban, Lechler 2404- (the var.). 
Brazil; Guiana. 

Aciotis longifolia (Mart.) Triana, Trans. Linn. Soc. 28: 52. 
1871; 305. Spennera longifolia Mart, ex DC. Prodr. 3: 115. 1828. 

Slender, typically herbaceous, with purplish subterete stem 
several dm. high; leaves narrowly oblong-lanceolate, long-acuminate, 
rounded at base, mostly with a few hairs on the upper surface, 
otherwise glabrous, very thin, 10-13 cm. long, 3-4 cm. wide; petioles 
to 3 cm. long; panicles diffuse; flowers sessile; bractlets 1-2 mm. 
long; calyx finally glabrous, 3 mm. long, the petals 4 mm. long, the 
style as long; capsule 4 mm. thick. The var. glabra Huber, Bol. 
Mus. Goeldi 4 : 595. 1906 is woody at base, the branches glabrous. 
Illustrated, Mart. Fl. Bras. 14, pt. 3: pi. 106. 

Loreto: Cerro de Canchahuaya (Huber 1475, the var.). Brazil. 

Aciotis polystachya (Bonpl.) Triana, Trans. Linn. Soc. 28: 52. 
1871; 303. Rhexia polystachia Bonpl. Rhex. 98. pi. 38. 1823. 

Similar to A. laxa but annual, with merely acute leaves, these 
4-5 cm. long, 2.5-3 cm. wide, the petioles glandular-pilose; pedicels 
and lanceolate bractlets 1 mm. long; calyx 1.5 mm. long, the tube 
narrowly campanulate; petals 2 mm. long, the style as long, the 
capsule as thick. The stems are densely reddish-brown-hispid, the 
trichomes long; the inflorescence open, the branches secundly 
flowered, about 6 cm. long and wide. F.M. Neg. 36186. 

San Martin: Near Tarapoto, Spruce. Lamas, Williams 6362. 
Loreto: La Victoria, Wittiams 2978. Brazil. 

Aciotis purpurascens (Aublet) Triana, Trans. Linn. Soc. 28: 
52. pi. 3. 1871; 306. Melastoma purpurascens Aublet, PI. Guian. 
402. pi. 154. 1775. 



FLORA OF PERU 299 

Shrubby, the stems 4-angled, often strongly; leaves oblong- 
lanceolate, long-acuminate, slightly narrowed to the margined 
petioles, 5-nerved, glabrous on the surfaces, 6-8 cm. long or longer, 
2-5 cm. wide below the middle; panicles elongate; pedicels and bract- 
lets 1-2 mm. long; calyx 3 mm. long, the tube ovoid; petals and style 
4 mm. long; capsule 3-4 mm. thick. The var. pellucida Cogn. has 
petioles to 2 cm. long, leaves to 13 cm. long, 4-5 cm. wide, pellucid, 
the stem scarcely branched; the var. martinensis Macbr., var. nov. 
foliis supra parce pilosulis. This last suggests A. longifolia and 
A. rubricaulis but it has the winged stems of A. purpurascens 
(Williams 7493, type). Many other collections besides the following 
have been made. 

San Martin: San Roque, Williams 7498. Near Moyobamba, 
Klug 3319 (var. martinensis}. Loreto: Yurimaguas, Poeppig 1837, 
(var. pellucida). Mishuyacu, Klug 144,' 1489 (det. Gl.). Yuri- 
maguas, Williams 3813; 4739; 7848. Near Iquitos, Williams 1485; 
1474; Killip & Smith 26958. Rio Mazan, Schunke 146. Pebas, 
Williams 1688; 1930. La Victoria, Williams 2701; 2568. Rio 
Itaya, Williams 38; 3209. Florida, Klug 2174. Rio Nanay, Williams 
334; 360; 406; 463; 803; 883; 1179; 1185; 1276. "Yaco mullaca." 

Aciotis rubricaulis (Mart.) Triana, Trans. Linn. Soc. 28: 52. 
pi. 3. 1871; 305. Spennera rubricaulis Mart, ex DC. Prodr. 3: 115. 
1828. 

Half-shrub with elongate sharply angled stems and oblong- 
lanceolate, acuminate leaves, somewhat pilose on both sides, their 
petioles densely hirsute; panicles elongate, narrowly pyramidal; 
calyx slightly glandular-pilose; petals shortly setose at tip, 4-5 mm. 
long, the style 3 mm. long; otherwise like A. longifolia, so it seems 
probable that there is only one variable species. Illustrated, Mart. 
Nov. Gen. & Sp. 3: pi. 255. 

San Martin: Mount Campana (Spruce). Brazil to Guiana and 
Central America. 

Aciotis sphaeranthera (Naud.) Cogn. in Mart. Fl. Bras. 14, 
pt. 3: 458. 1885; 298. Spennera sphaeranthera Naud. Ann. Sci. Nat. 
se>. 3. 14: 142. 1850. 

Small annual, the ascending-erect stems 1-2 dm. high, reddish 
pilose on the angles; leaves oblong-lanceolate, entire, both ends 
acute, 3-nerved, sparsely ciliate on the margins and pilose on the 
nerves beneath, mostly 3 cm. long, about 5 mm. wide; panicle 



300 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

about 3 cm. wide, scarcely as high; petals retuse, 2-3 mm. long; 
anthers suborbicular. F.M. Neg. 36189. 
Cuzco: Near Cuzco, Gay, type. 

16. MONOCHAETUM (DC.) Naud. 
Reference: Gleason, Amer. Jour. Bot. 16: 502-522. 1929. 
Shrubs with rather few cymose or solitary medium-sized 4-merous 
flowers and ovate or lanceolate leaves, the 3-7 nerves more or less 
strongly impressed above. Calyx tube oblong-campanulate or 
tubular, the lobes persisting in the Peruvian species. Anther 
connective not produced but the larger anthers with a porrect, 
often elongate and conspicuous appendage at base, the shorter, often 
sterile ones with slenderer and longer appendage. Ovary adherent 
below, 4-celled, pubescent at tip. Seeds arcuate or cochleate. 
The related group Pachyloma DC. of the upper Amazon, to be 
expected, has a truncate calyx, the connective anteriorly biauriculate, 
the ovary glabrous; P. coriaceum DC., 390, has glabrous branches, 
the pedicels 1-3 mm.; P. huberioides (Naud.) Triana, 390, has 
branches slightly pilose at the nodes, the pedicels 1-2 cm. long. 
Branchlets conspicuously spreading-hirsute. 

Calyx somewhat glandular M. subditivum. 

Calyx not at all glandular . M. dicranantherum. 

Branchlets glabrate or more or less appressed-setulose. 

Calyx lobes 5-7 mm. long; petals nearly 10 mm. long. 

M. lineatum. 

Calyx lobes 4.5 mm. long; petals 6-7 mm. long. . .M. subglabrum. 

Monochaetum dicranantherum (R. & P.) Naud. Ann. Sci. 
Nat. se>. 3. 4: 52. 1845; 393. Rhexia dicrananthera R. & P. Fl. 
Peruv. 3: 84. pi. 320. 1802. 

Branches loosely hispid with spreading hairs; calyx not at all 
glandular, the pubescent lobes 3-3.5 mm. long; petals to 1 cm. 
long; larger anthers 4 mm. long, the tail recurving, that of the 
smaller filiform, erect. The narrowly ovate leaves are 5-nerved, 
2-3 cm. long, the pediceled flowers rather many in a terminal cyme. 
F.M. Neg. 16825. 

Huanuco: Cuchero and Muiia, Ruiz & Pavdn; 3929. Cani near 
Mito, 3436. Northeast of Huanuco, 2145. Tomaiquichua, 2430. 

Monochaetum lineatum (D. Don) Naud. Ann. Sci. Nat. se"r. 
3. 4: 52. 1845; 399. Arthrostemma lineatum D. Don, Mem. Wern. 
Soc. 4: 299. 1823. 



FLORA OF PERU 301 

Branches very sparsely setulose, the few setae appressed; leaves 
oblong-lanceolate, 5-plinerved, 3-3.5 (5) cm. long, 10-12 mm. wide 
with a few scattered setae above, or these only in one line between 
the nerves, scarcely more pubescent on the nerves beneath; cymes 
axillary, few-flowered, open; calyx lobes 5-7 mm. long, nearly 
glabrous; larger anthers 6 mm. long, the smaller 3-4 mm. long. 
Markgraf, in Herb. Madrid, has referred the Ruiz & Pavon specimen 
from Chinchao to M. Hartwegianum Naud., 394, a species collected 
at Guayaquil by Ruiz & Pavon, which, as to type, has spreading 
setose-villous branches, subglabrous calyx teeth 3 mm. long, and 
leaves glabrate except for some appressed setae. Cf. remark under 
M. subditivum. F.M. Neg. 26086. 

Huanuco: Chinchao, Ruiz & Pavon. San Martin: Moyobamba 
(Mathews). Amazonas: Chachapoyas, Mathews. Without locality, 
Nee. Ecuador. 

Monochaetum subditivum Macbr. Field Mus. Bot. 4: 175. 
1929. 

An open shrub a meter or so high, similar to M. dicranantherum 
but the leaves 5-plinerved, 3-4 cm. long, the cymes 2-4-flowered 
and the calyx setae mostly gland-tipped; appendage of smaller 
anthers linear, little shorter than the anther. Perhaps a var. of 
M. lineatum, as part of type of that species has glandular calyces. 

Huanuco: Hacienda Villcabamba on the Rio Chinchao, 5194, 
type. Rio Chinchao, Ruiz & Pavon (M. lineatum in part, in some 
herbaria). 

Monochaetum subglabrum Gleason, Bull. Torrey Club 58: 
220. 1931. 

Stems very sparsely pilose, the trichomes 2-3 mm. apart; sepals 
triangular-lanceolate 4.5 mm. long, 1.6 mm. wide at base, sharply 
acute, glabrous except for the setulose tip; petals white or pink, 
6-7 mm. long; small anthers with straight oblanceolate appendage, 
that of the larger arcuate, 4 mm. long, dilated toward the tip; 
style 4 mm. long. 

Ayacucho: Ccarrapa, 1,500 meters (Killip & Smith 22325, 
type). Junin: Carpapata, 2,700 meters (Killip & Smith 24423). 

17. HUBERIA DC. 

Glabrous, sometimes viscid shrubs with 3-nerved oblong or 
lanceolate leaves, and long-pediceled 4-merous cymose flowers. 
Calyx tube oblong, with a dilated lobed limb. Stamens nearly 



302 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

equal, similar, the connective of the elongate anthers not produced, 
appendaged only posteriorly, the appendage directed downwards. 
Seeds broadly winged, the ovoid capsule included in the ventricose 
calyx, 4-angled, 4-celled. 

Huberia peruviana Cogn. in Mart. Fl. Bras. 14, pt. 4: 10. 
pi. 1. 1886; 410. 

Glabrous except for the viscous cymes; petioles nearly 1 cm. 
long; leaves crowded, rather rigid, lanceolate, rather closely serrulate 
above the middle, acuminate, 5-6 cm. long, about 2 cm. wide; 
calyx tube 6 mm. long, the linear teeth to 2 mm. long; petals golden 
yellow, very acute, 7-8 mm. long; capsule 8-9 mm. long; seeds 5-6 
mm. long. F.M. Neg. 25892. 

San Martin: Tarapoto, Spruce 4828. Sambasbambe (Mathews 
1283). Near Moyobamba, King 3271. 

18. ADELOBOTRYS DC. 

Scandent shrubs (rarely erect), often more or less herbaceous 
above and, usually at least, somewhat reddish-setose or furfuraceous, 
especially on the younger parts and on the leaves beneath, these 
oblong-elliptic to suborbicular. Flowers in panicled umbels or clusters, 
or the terminal inflorescence rarely reduced to 2-3 or few flowers. 
Calyx conical to campanulate, the limb dilated as a flange, usually 
merely undulate or barely 5-lobed, the outer teeth often merely 
tubercles or setae, sometimes subulate, rarely well-developed. 
Stamens equal in size, or unequal, or slightly dissimilar, the subulate 
anthers usually curved, the connective basally short-spurred and 
dorsally with an elongate appendage, often bifid or tailed, borne 
parallel to the anther. Filaments narrowly winged or flattened but 
usually filiform. Ovary 3-5-celled, the slender or filiform seeds 
tailed at least at one end. As Gleason has shown, the stamens 
are often dimorphic; therefore the chief character upon which 
Marshallfieldia Macbr. (cf. A. Boissieriana) was based is of no 
value here; earlier descriptions of the genus were either not clear 
or inexact; the above character as regards the calyx and filaments, 
inflorescence also, seems to conform to the facts. The species listed 
are doubtfully all valid; two groups could be based on pubescence; 
this is mostly setulose in M. Boissieriana, M. fuscescens, M. macro- 
phylla, M. rotundifolia scurfy or appressed-strigose and scurfy or 
nearly lacking in remaining species. 

Leaves oblong-lanceolate, 1.5-2 cm. wide A. subsessilis. 

Leaves ovate to subrotund, wider. 



FLORA OF PERU 303 

Leaves minutely, closely denticulate above the middle, below 

the entire margins revolute A. Tessmannii. 

Leaves entire or obscurely crenulate-denticulate. 

Calyx teeth 2.5-3 mm. long or longer A. fuscescens. 

Calyx teeth minute or obsolete. 
Petals about 15 mm. long; divisions of dorsal spur of anthers 

only 0.8 mm. long A. macrantha. 

Petals usually much shorter; dorsal spur lobes about 2 mm. 

long or longer. 

Petioles 5-7 mm. long, the oval leaves inordinately red- 
pubescent margined A. praetexta. 

Petioles often longer (sometimes short); leaves in any 

case otherwise. 
Leaves subrotund or only about one-third longer than 

broad; stamens obviously dimorphic. 
Outer calyx teeth evident, marginal, the limb merely 

dilated A. rotundifolia. 

Outer calyx teeth obsolete or the limb obscurely lobed. 

A. Boissieriana. 

Leaves ovate or elliptic (always?), about twice longer 
than broad; calyx teeth below limb-margin, minute 
or obsolete; stamens nearly isomorphic. 
Leaves more or less densely pilose beneath. 

A. macrophylla. 

Leaves glabrate beneath or only lightly pubescent on 
the nerves A. guianensis. 

Adelobotrys Boissieriana Cogn. in Mart. Fl. Bras. 14, pt. 4: 
21. 1886; 420. Marshallfieldia corallina Macbr. Field Mus. Bot. 4: 
176. 1929. 

Branches scandent, densely long-reddish-villous, especially at 
the nodes; petioles to 12 (20) mm. long; leaves slightly obovate, 
more or less rounded at base, shortly acute or cusped, entire, 3-nerved 
with faint marginal nerves, the nerves and veins as the petioles and 
inflorescence, densely reddish-hispidulous, the leaves finally glabrate 
above, to about 11 cm. long, 8 cm. wide; panicle typically many- 
flowered, to 2 dm. long; pedicels 2-3(-5) mm. long; calyx 7 mm. 
long, the limb dilated, the obscure roundish scarious lobes 2 mm. 
high, the outer teeth about 0.3 mm. long or tuberculif orm ; fila- 
ments subfiliform but flattened, stamens very unequal, 7-8 mm. and 



304 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

11 mm. long, nearly equaled by the long-attenuate anthers or the 
anther shorter, the capillary dorsal appendage 4-5 mm. long, deeply 
bifid; petals narrowly obovate, about 10 mm. long. The type 
of Marshallfieldia was a young, undeveloped specimen; this would 
account for the reduced inflorescence of 2-3 flowers; the character 
of unequal stamens, upon which the genus was particularly seg- 
regated, is actually a character of other species of Adelobotrys, although 
Cogniaux described the stamens as equal, the filaments "filiform." 
In the type of A. Boissieriana the rather short, narrow and long fila- 
ments are 7 and 11 mm., their respective anthers 4-5 and 8-9 mm. 
The species could be kept generically distinct on the basis of the 
aberrant calyx and would be as distinct a "genus" as several recog- 
nized traditionally. F.M. Neg. 29490. 

Huanuco: Chinchao, Pav6n, type. Junin: La Merced, Hacienda 
Schunke, climbing a tree-trunk, 5642, type, Marshallfieldia. Chan- 
chamayo Valley, Schunke 408. 

Adelobotrys fuscescens Triana, Trans. Linn. Soc. 28: 68. 
1871; 420. 

Moderately pilose, especially the leaves beneath and the calyces, 
the latter marking the species at once from other Peruvian forms 
by the long outer teeth, these as much as 3 or 4 mm. long; petioles 
1 cm. long or longer; leaves coriaceous, ovate-oblong, 8 cm. long or 
longer, 5-8 cm. broad, entire, rather abruptly acuminate, 5-nerved; 
panicle ample, rather narrow; pedicels 4-8 mm. long; calyx tube 
5-7 mm. long, subcylindrical, the strongly dilated limb obscurely 
lobed, 5-6 mm. broad; petals 6-7 mm. long. F.M. Neg. 25850. 

Loreto: Mouth of Santiago, Tessmann 4506 (det. Markgr. as 
A. macrophylla) . Brazil; Colombia. 

Adelobotrys guianensis (DC.) Gleason, Brittonia 1: 141. 1932. 
A. adscendens Triana, Trans. Linn. Soc. 28: 67. pi. 5. 1871, not 
Melastoma adscendens Sw. Fl. Ind. Occ. 2: 772. 1800. Davya guia- 
nensis DC. Prodr. 3: 105. 1828. Miconia scandens R. & P. Fl. 
Peruv. 4: pi. 395. 1802. Davya peruviana DC. Prodr. 3: 105. 1828. 
A. multiflora Pilger, Verh. Bot. Ver. Brandenb. 47: 166. 1905. 

Elongate branches, slender petioles and generally membranous 
leaves sparsely reddish-setulose, becoming glabrate or glabrous, or 
the leaves beneath more or less prominently setulose on the nerves, 
these 5, basal or sometimes partly originating above the base; leaves 
ovate-oblong, rounded at base, long-acuminate, entire, or rarely 
toward the tip remotely serrulate, the lower 1 dm. long, 6-8 cm. 



FLORA OF PERU 305 

wide or larger, the upper often much smaller; panicle much-branched, 
sometimes 2 dm. long or longer, the umbellate slender pedicels 
4-7 mm. long; calyx tube 6-7 mm. long, 4-5 mm. broad at tip, the 
outer teeth 0.3-0.5 mm. long, barely projecting above the truncate 
calyx tube; filaments filiform but compressed, about 7 mm. long, 
the anthers nearly as long. Stamens somewhat dimorphic, but not 
distinctly so as in the true adscendens of Jamaica (Gleason). 

Huanuco: Near Cuchero, Dombey. San Martin: Moyobamba, 
1,200 meters, Weberbauer 4685; 290, det. Cogn. Pongo de Caina- 
rachi, Klug 2603 (det. Gl. as A. rotundifolia) . Without locality, 
Ruiz & Pav6n. Loreto: Mouth of Santiago, Tessmann 1*370; 4120 
(det. Markgr. as A. multiflora); Tessmann 406 (det. Markgr. as A. 
rotundifolia). Balsapuerto, Klug 2948 (det. Gl.) Pampas de Ponasa, 
Ule 6749, type A. multiflora. Bolivia to Mexico and the West 
Indies. 

Adelobotrys macrantha Gleason, Bull. Torrey Club 58: 220. 
1931. 

Younger branches, leaves beneath and panicles moderately brown- 
strigose; petioles stout, 3-5 cm. long, closely and completely pube- 
scent; leaves elliptic, acute at each end, 5-nerved, soon glabrescent 
above; pedicels about 5 mm. long, the obconic-strigose calyx 9 mm. 
long, with broadly rounded sepals, the outer teeth conic-tuberculi- 
form; petals pink, 15-19 mm. long; stamens nearly isomorphic, the 
filaments flat, 12 mm. long, the anthers nearly straight, the longer 
nearly 13.5 mm., the shorter 8.5 mm., the lobes of the dorsal spur 
only 0.8 mm. long; style slender, 16 mm. long, not tapering, the 
stigma truncate. Shrub or small tree, the type noted as having 
rich pink petals and yellow anthers; Killip & Smith 29616, sterile, 
was a "subligneous vine." The largest leaves are 2.5 dm. long, 
14 cm. wide. 

Junin: Cahaupanas, on Rio Pichis, 340 meters, Killip & Smith 
26745, type. Loreto: Rio Itaya, Killip & Smith 29496. Soledad, 
Killip & Smith 29616. Mouth of Santiago, Tessmann 4181 (det. 
Markgr. as A. fuscescens). 

Adelobotrys macrophylla Pilger, Verh. Bot. Ver. Prov. Bran- 
denb. 47: 165. 1905. 

Younger branches and leaves beneath, especially on nerves and 
at margins, densely reddish-setose-villous; leaves elliptic, rounded 
or lightly cordate at base, gradually narrowed above, acute, mem- 
branous, becoming glabrescent above and as large as 18 cm. long, 



306 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

8 cm. wide, 5-nerved; petioles stout, typically only 5-7 mm. long; 
flowers shortly pediceled, the calyx tube densely appressed, ashy- 
setose, 5 mm. long, the dilated limb not obviously lobed; petals 
white or roseate, broadly obovate, 8-9 mm. long; stamens equal, 
the filaments dilated, 7 mm. long, the anther slightly longer; style 
3 mm. exserted. F.M. Neg. 16849. 

Loreto: Leticia, Ule 6199, type. La Victoria, Williams 2914; 
2915 (det. Gl.). 

Adelobotrys praetexta Pilger, Verh. Bot. Ver. Prov. Brandenb. 
47: 167. 1905. 

Species striking by the broadly oval, abruptly caudate leaves 
which are soon glabrous or glabrate except for border of densely 
matted red hairs; petioles 7-15 mm. long; leaves typically about 
6 cm. long, 3.5-4 cm. broad, 5-nerved, the veins obscure; fruiting 
pedicels 4 mm. long; calyces costate, sparsely setose, 6 mm. long, 
the limb not lobed, the outer teeth obsolete or setulose. F.M. 
Neg. 16851. 

Loreto: Pongo de Cainarachi, Ule 6368, type. Santa Ana, Upper 
Rio Nanay, Williams 1231 (det. Gl.). 

Adelobotrys rotundifolia Triana, Trans. Linn. Soc. 28: 68. 
1871; 420. 

Subterete branchlets, petioles, leaves beneath and inflorescence 
moderately reddish-pilose; leaves typically suborbicular, apparently 
sometimes elliptic, about 9 cm. long, 7 cm. wide or smaller, often 
broader in proportion to width, shortly acuminate, rounded or 
emarginate at base, distinctly 5-nerved or the outer nerves only 
partial; pedicels 5-8 mm. long; calyx limb spreading, 5 mm. broad, 
inner lobes obvious, the outer teeth minute, triangular-subulate; 
petals 6-7 mm. long; stamens a little dimorphic, anthers 5 and 7 mm. 
long, the dorsal spur 3 mm. long, bifid for less than 1 mm. Material 
referred here from Peru seems to be referable elsewhere. The species, 
however, has been collected rather near Peru in Brazil, Krukoff 
7277 (det. GL). Illustrated, Mart. Fl. Bras. 14, pt. 4: pi. 5. 

San Martin: San Roque, Williams 7108(1}. Brazil. 

Adelobotrys subsessilis Gleason, Bull. Torrey Club 58: 222. 
1931. 

Slender branches, leaves beneath and compact panicles densely 
strigose with malpighian trichomes; petioles merely 2 mm. long; 
leaves narrowly oblong-lanceolate, rounded at base, acute, entire, 



FLORA OF PERU 307 

3-nerved, with 2 submarginals, 5-7 cm. long, 1-2 cm. wide; calyx 
obconic, each broadly oblong sepal medially elevated into a 
spreading, stoutly conic outer tooth, 0.8 mm. long; petals pink, 
9 mm. long; stamens nearly isomorphic, the anthers 6 and 7 mm. 
long; style slender, 9 mm. long, the stigma punctiform. Type 
from a slender shrub 4 meters high; the leaves bear longitudinal 
strips of deep brown hairs and the panicles are racemiform. 

Loreto: Iquitos, Killip & Smith 27147, type. Caballo-Cocha, 
Williams 2170. 

Adelobotrys Tessmannii Markgr. Notizbl. Bot. Gart. Berlin 
9: 1140. 1927. 

Pale-green and glabrous except early, then slightly scurfy- 
pilose-puberulent on the petioles, leaf nerves beneath, younger 
stems and inflorescence; petioles sometimes 3 cm. long; leaves 
broadly ovate, rounded at base, acuminate, 5-nerved, to 15 cm. long, 
9 cm. wide; pedicels 2-4 mm. long; calyx tube 3.5 mm. long, the 
dilated, merely undulate limb 4 mm. broad, the nearly marginal 
teeth minute but subulate and projected about 0.5 mm.; petals 
white, rose-tinted, 8 mm. long; stamens slightly dimorphic, the 
longer 7, the shorter 4 mm. long, the appendage 3 mm. long; style 
3 mm. long; fruit globose, immature. 

Loreto: Below Itaya, in flooded woods, Tessmann 3572, type. 
Colombia. 

19. MERIANIA Swartz 

Similar to Axinaea but dorsal anther appendage acute or knobbed 
or sometimes rudimentary, not at all inflated. Calyx lobes often 
with exterior teeth. Petals never emarginate. Filaments often 
linear or filiform. Ovary 3-6-celled, the style filiform. Capsule 
always included. Calyx truncate and obscurely if at all lobed. 
Leaves obviously serrulate nearly or quite to the base, ample. 
Leaves cordate-based. 

Pubescence sparse or nearly wanting M. Raimondii. 

Pubescence abundant on leaves beneath M. Radula. 

Leaves rounded or acute at base. 

Calyx glabrous; outer teeth none M. tetraquetra. 

Calyx scurfy; outer teeth developed but promptly caducous. 

M. Weberbaueri. 

Leaves obscurely if at all serrulate or only above the middle and 
then, at least, small or medium in size. 



308 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branches deeply sulcate-angled M. nobilis. 

Branches terete or obtusely quadrate. 

Leaves conspicuously hirsute-pilose M. intonsa. 

Leaves glabrous or scurfy beneath. 
Flowers solitary or rarely ternate. 

Leaves ovate or elliptic; calyx tube grooved. M. prunifolia. 
Leaves oblong-lanceolate; calyx tube smooth. M. longifolia. 
Flowers paniculate, sometimes few. 

Leaves oblong-lanceolate, serrulate M. oblongifolia. 

Leaves ovate-elliptic, entire. 

Calyx densely reddish-puberulent M. urceolata. 

Calyx glabrous. 

Flowers 5-merous M. rugosa. 

Flowers 6-merous M. hexamera. 

Meriania hexamera Sprague, Trans. Bot. Soc. Edinb. 22: 
431. 1905. 

Similar to M. rugosa; petioles 1.5-2.5 cm. long; leaves somewhat 
smaller; calyx merely undulate, outer lobes obsolete; petals 6, 
obovate. The Peruvian collection is nearly glabrous; there are 
essentially 3 principal nerves a little above the base and a very faint 
marginal pair. The material is incomplete as seen by me but 
appears, from photograph, to be Sprague's species. 

Amazonas: Rare tree, trail between Chachapoyas and Moyo- 
bamba, Williams 7598. Colombia. 

Meriania intonsa Gleason, Bull. Torrey Club 58: 223. 1931. 

Scandent, the hirsute nodes 4-8 cm. long; petioles stout, 6-13 
mm. long; leaves elliptic, subcordate, abruptly acuminate, 5-nerved, 
densely long-pilose above, hirsute on the veins, pilose on the face 
beneath, membranous, to 8.5 cm. long, 5.5 cm. wide; inflorescence 
terminal, few-flowered, sessile with sessile cordate-ovate bracts; 
flowers subsessile, the calyx narrowly obconic, 4 mm. long, the 
limb undulate, the hirsute subulate teeth 3.5 mm. long and terminat- 
ing in a longer bristle; petals about 5 mm. long, broadly rounded; 
connective bispurred at base, the basal spur conic, the dorsal elongate; 
ovary 5-celled. The yellowish calyx-bristles are striking, the petals 
lilac or white. 

Loreto: Mishuyacu, Kittip & Smith 29969, type; King 1334; 
2541; 90. Colombia. 



FLORA OF PERU 309 

Meriania longifolia (Naud.) Cogn. Melast. 426. 1891. Chas- 
tenaea longifolia Naud. Ann. Sci. Nat. se>. 3. 4: 55. 1845. 

Nearly glabrous; petioles 4-8 mm. long, minutely pilose above; 
leaves lanceolate, mostly 6 cm. long, 1.5-2 cm. wide, glabrous, long- 
acuminate, minutely serrulate toward the tips, firm; solitary flowers 
axillary and terminal on long slender peduncles but with two leaf- 
like bracts below the calyx, this with rounded tube 7-8 mm. long, 
broadly triangular lobes 3 mm. long, minute outer teeth; petals to 
2 cm. long; appendages rudimentary. The Peruvian plant is the 
var. Spruceana (Cogn.) Macbr., comb. nov. (M. Spruceana Cogn. 
Melast. 426. 1891) distinguished from the typical form by shorter 
petioles, leaves, and flowers. F.M. Neg. 16876. 

San Martin: In the mountains of Campana, Tarapoto, Spruce 
4204, type. San Roque, Williams 7010 (det. Gl.). Near Moyo- 
bamba, Klug 8642. Also in Colombia and Venezuela. "Cruz 
chillca." 

Meriania nobilis Triana, Trans. Linn. Soc. 28: 66. 1871; 436. 

Branches and leaves lustrous, glabrous, the former deeply 
sulcate-angled or somewhat winged and membranously stipulate at 
the enlarged nodes; petioles 4-8 cm. long; leaves cordate-ovate, 
dilated-flanged at base, apically subrounded or apiculate, stoutly 
5-nerved and with another pair of faint marginal nerves, 2 dm. long, 
1.5 dm. wide or larger; pedicels to about 3 cm. long; calyx tube 
campanulate, the cupulate limb obscurely 5-lobed, the rather 
prominent outer teeth confluent with the inner; connective of the 
larger anthers long-clawed, the claw subulate, ascending, the 
appendage of the smaller anthers confluent with the connective. 
It seems possible that the specimen cited here was mislabeled. 
F.M. Neg. 26054. 

Peru(?): Without locality, Ruiz & Pavon. Colombia. "Panda." 

Meriania oblongifolia Cogn. Melast. 429. 1891. 

Glabrous, the branchlets slender and leafy only toward the 
tips; leaves oblong-elliptic, 8 cm. long, 2.5 cm. wide, thin, decurrent 
on the slender petiole, this about 2 cm. long, finely serrulate toward 
the acutish tip, 3-nerved or slightly plinerved ; panicle loose, nodding, 
the pedicels 5 mm. long or longer; calyx obscurely lobed, the tubercle- 
like teeth obvious; petals in herbarium pale yellow; anthers shortly 
and thickly beaked at base. The type may have been found in 
Ecuador. F.M. Neg. 36200. 

Peru(?): Without locality, Grisar, type. 



310 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Meriania prunifolia D. Don, Mem. Wern. Soc. 4: 323. 
1823; 438. 

Branchlets slender, furfuraceous; petioles nearly filiform, 4-7 mm. 
long; leaves ovate, bluntly acute, to 4 or 5 cm. long, 2.5 cm. wide, 
membranous but firm, green and glabrous above, lighter beneath, 
where a little scurfy on the 3-5 nerves; flowers terminal or axillary, 
solitary, subsessile; calyx tube scurfy, campanulate, 4 mm. long, 
the broadly rounded lobes 2 mm. long, the spreading teeth to 4 mm. 
long; appendages elongate. The fruiting calyx is strikingly grooved, 
the outer calyx teeth about a third as long. The K. & S. specimen, 
in fruit with glabrous calyx, elliptic leaves 14 by 5 cm. may not 
belong here. F.M. Neg. 26055. 

Peru(?): Without locality, Ruiz & Pavon, type. Junin: tree, 
8 meters, Santa Rosa, Pichis Trail, Killip & Smith 26184(1}. 

Meriania Radula (Benth.) Triana, Trans. Linn. Soc. 28: 66. 
1871; 431. Axinaea Radula Benth. PI. Hartw. 130. 1844. A. 
purpurea R. & P. Syst. 1 : 122. 1798, not M. purpurea Sw. Miconia 
incarum Macbr. Trop. Woods 17: 12. 1929. 

Stout tree-shrub, the thick branchlets obtusely quadrate; petioles 
2-3 cm. long; leaves cordate-ovate, to 2 dm. long, 1.5 dm. wide or 
larger, crenulate-denticulate, dark green and bullate-asperous above, 
paler and crisply tomentulose beneath, 7-9-nerved; calyx tube some- 
what furfuraceous, subhemispheric, to nearly 1 cm. long, the gla- 
brate limb truncate-dilated; petals about 1.5 cm. long; appendages 
rudimentary. Sometimes a tree 17 meters high. F.M. Neg. 16873. 
Illustrated, R. & P. Fl. Peruv. 4: pi. 410. 

Huanuco: Muna, Ruiz & Pavon, type. Mito, 1867, type, M. 
incarum; 3547. Libertad: Prov. Pataz, Weberbauer 7032. Ecuador. 
"Mogo-mogo." 

Meriania Raimondii Markgr. Notizbl. Bot. Gart. Berlin 13: 
461. 1937. 

Comparable to M. Radula; petioles 3-5 cm. long; leaves some- 
times somewhat plinerved, chartaceous, cordate, shortly acuminate, 
broadly serrate, 12-15 cm. long, 8-9 cm. wide, obtusely bullate, 
glabrous except the sparsely furfuraceous nerves beneath; pedicels 
4-6 mm. long; calyx glabrous with sinuate triangular lobes 1 mm. 
high, 2.5 mm. broad; petals 10 mm. long and broad, glabrous; 
filaments and anthers 7 mm. long, the connective with acute spur 
2 mm. long; fruit 7 mm. thick, globose, angled. 



FLORA OF PERU 311 

Cajamarca: Cutervo, in flower and fruit in April (Raimondi, 
type). San Martin: Moyobamba, Williams 7579. 

Meriania rugosa Markgr. Notizbl. Bot. Gart. Berlin 13: 460. 
1937. 

A tall shrub, glabrous (or essentially) with obtusely quadrangular 
glabrate branchlets, quadrately rugose, subchartaceous, lanceolate- 
elliptic leaves that are oblique at base, minutely undulate-margined, 
5-plinerved, 15-17 cm. long, nearly half as wide; petioles to 4 cm. 
long; panicles lax, terminal, to 2.5 dm. long, the slender pedicels 
5-10 mm. long; calyx tube 5 mm. high, 6 mm. broad, the 5 obtuse 
lobes 2 mm. long, 4 mm. broad, tubercled posteriorly; petals orbicular, 
1 cm. broad; filaments 7 mm. long, 2 mm. broad, the acuminate 
anthers 10 mm. long; connective spur 2 mm. long, unappendaged ; 
style 12 mm. long; ovary costate. Apparently very nearly M. 
hexamera. 

Amazonas: Chachapoyas, Valle de Huayabamba (Raimondi, 
type). Between Cochamal and Santa Rosa (Raimondi). 

Meriania tetraquetra Triana, Trans. Linn. Soc. 28: 66. 
1871; 431. 

Nearly the same as M. Radula and perhaps not specifically 
separable; branches acutely quadrate; petioles longer and leaves 
larger, membranous, more remotely denticulate and merely scurfy 
puberulent on the nerves beneath.- F.M. Neg. 26058. 

Amazonas: Chachapoyas, Mathews, type. Libertad: Prov. Pataz, 
2,000 meters, Weberbauer 7048 (det. Markgr.). 

Meriania urceolata Triana, Trans. Linn. Soc. 28: 67. 1871; 435. 

Branches and petioles terete or nearly, the latter 2.5 cm. long; 
leaves broadly ovate, minutely auriculate at base, to about 15 cm. 
long and half as wide, 5-plinerved, glabrous above, minutely brown- 
stellate beneath, the surface nearly concealed; panicles terminal, 
erect, many-flowered; calyx about 1 cm. long, reddish-scurfy, the 
exactly truncate limb minutely 5-tubercled above the margin; 
appendages elongate. Marked by the red-puberulent tubular calyx. 
A tree to 7 meters (Triana). F.M. Neg. 16877. 

San Martin: Tarapoto, Spruce 4439, type. Moyobamba, 800 
meters, Weberbauer 4570; 4541; 290; King 3451. Brazil; British 
Guiana. 



312 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Meriania Weberbaueri Macbr. Field Mus. Bot. 4: 176. 1929. 

Branchlets somewhat enlarged at the nodes, obscurely quadrate, 
scurfy, finally glabrous; leaves broadly elliptic-ovate, slightly at- 
tenuate at base, repandly serrulate toward the obtuse tip, the larger 
about 2 dm. long and half as wide, firm, glabrous above, minutely 
and densely pubescent beneath, with stellate and pilose hairs inter- 
mixed, 7-plinerved; cymes few-flowered; calyx lobes triangular, 
equaling or longer than the tube, this campanulate, 6 mm. long, the 
outer teeth scarcely 3 mm. long, the limb not at all dilated; petals 
rose-colored, about 2 cm. long; connective strongly enlarged at base, 
produced into a short, minutely bilobed appendage. Said to be a 
member of the section Umbellatae Cogn., 428, but simulates M. 
tetraquetra closely. 

Junin: Valley of the Rio Masamerich, branch of the Rio Pangao, 
Weberbauer 6659, type. 

20. AXINAEA R. & P. 

Chastenaea DC. Prodr. 3: 102. 1828. 

Reference: Donald S. Eves, Bull. Torrey Club 63: 211-226. 
1936. 

Glabrous or pubescent shrubs or trees with rather showy, 4-5- 
merous flowers borne in broad lateral and terminal panicles. Branches 
sometimes conspicuously enlarged at the nodes or stipule-scarred, 
the leaves sometimes blunt, and in many species with the lobes or 
auricles at the base drawn together on the upper side into a small 
tubercle, this structure more or less developed. Calyx limb dilated, 
truncate or very shortly and obscurely lobed. Stamens 10, some- 
times 8, equal or nearly, glabrous, the anther connective inflated 
or fleshy, dorsally appendiculate. Ovary free, 4-5-celled. Seeds 
many, oblong-pyramidal. A. purpurea R. & P. Syst. 1: 122. 1798, 
not mentioned or not identified by Eves, is Meriania Radula. 

Leaves basally tubercled by the confluence of the dilated petiole-tip 
with adjacent leaf-margin, the resultant thickening sometimes 
wart-like, sometimes obscure. 
Leaves with a more or less expanded and recurved auricle on 

petiole-tip A. grandifolia. 

Leaves not so auricled, if at all. 

Leaves villous beneath, with plumose trichomes. . . A. tomentosa. 
Leaves glabrous or more or less scurfy beneath, the trichomes 
not distinctly if at all plumose. 



FLORA OF PERU 313 

Leaves broadly ovate, about twice as long as broad. 

Leaves tomentose beneath A. glandulosa. 

Leaves glabrate beneath A. tetragona. 

Leaves lance-ovate, about thrice as long as broad. 
Branchlets obscurely angled; petioles 1-1.5 cm. long. 

A. nitida. 

Branchlets (the youngest) angled; petioles 2-3 cm. long. 
Leaf auricles reflexed, the leaf base thus appearing acute 

from above A. Pennellii. 

Leaf auricles flat or curved, the leaf base definitely 

cordulate A. Weberbaueri. 

Leaves plane at base, but in A. lanceolata auricled. 

Leaves sessile, or nearly, and cordulate A. sessilifolia. 

Leaves petioled, long-petioled if cordate. 
Leaves long-petioled, definitely cordate; petals 4 (5?). 

Nodes with flange-like annulus A. tetragona. 

Nodes merely enlarged A. crassinoda. 

Leaves short-petioled, 'obscurely if at all cordate; petals 5. 
Leaves not entire. 

Stipules persisting . A. dependens. 

Stipules deciduous. 

Leaves with reflexed basal auricles A. lanceolata. 

Leaves without auricles, the leaf base rarely revolute. 

A. Merianiae. 
Leaves entire A. pauciflora. 

Axinaea crassinoda Triana, Trans. Linn. Soc. 28: 69. 1871; 214. 

A sprawling or straggling shrub or small tree, well marked by 
the strikingly enlarged nodes of the somewhat tetragonous branches, 
the prominent stipule scars and, especially, by the 4-merous flowers, 
in which character the species is unique; petioles stout, 4-7 cm. long, 
densely scurfy-hirsute as the leaves beneath; leaves broadly ovate, 
13-19 cm. long, 12-16 cm. wide, cordate basally, rounded and 
obtuse apically, conspicuously crenate-dentate, subcoriaceous, 7-9- 
nerved; panicles pendulous, about 1.5 dm. long, the branches 
spreading, the pedicels and calyces scurfy, the former 4-10 mm. 
long; calyx limb 4-lobed to truncate; petals pink, retuse, 8-10 mm. 
long; stamens 8, the conic acuminate connective 4 mm. long; style 
slightly curved, 9-12 mm. long. F.M. Neg. 25854. 



314 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Amazonas: Chachapoyas, Mathews 3213, type. Playapampa, 
4873. 

Axinaea dependents R. & P. Fl. Peruv. 4. pi 411. 1802; 218. 
Meriania dependens (R. & P.) Naud. Ann. Sci. Nat. se>. 3. 18: 124, 
129. 1852. 

Branches obscurely tetragonal, little if at all enlarged at the 
nodes, early finely scurfy, the stipules persistent, semi-circular; 
petioles to 4.5 cm. long, scurf y-puberulent, as the 5-nerved or 
shortly plinerved leaves beneath, especially on the nerves and 
veins; leaf -blades ovate- or oblong-elliptic, bluntly acute, acute 
to rounded at the base, 11-16 cm. long, 3-5 cm. broad, submem- 
branous; panicles pendulous, to 2.5 dm. long, the puberulent pedicels 
10-12 mm. long; calyx scurfy-puberulent, 5-7 mm. wide, the limb 
obscurely 5-lobed; petals obtuse, 10-12 mm. long; filaments very 
flat, equaled by the narrowly conic, long-acuminate connectives, 
these 5 mm. long; style attenuate, 12-18 mm. long. F.M. Negs. 
16897; 16899 (as A. glandulosa). 

Huanuco: Chinchao, Ruiz & Pavdn, type. Junin: Vitoc, Ruiz 
& Pavon. 

Axinaea glandulosa R. & P. Fl. Peruv. 4: pi. 412. 1802; 214. 
Chastenaea glandulosa (R. & P.) Naud. Ann. Sci. Nat. se"r. 3. 18: 
123, 124. 1852. 

Similar to A. dependens, but the stipule scars not prominent, the 
leaves 5-7-nerved, auriculate and tubercled at base, the auricles 
reflexed, densely scurf y-tomentose beneath, coriaceous, 5-7 cm. 
wide; panicles erect, the peduncles stout, the thick, scurfy pedicels 
5-8 mm. long; calyx glabrous, rugose, 7 mm. wide; petals apiculate, 
13 mm. long; connective broadly ovate, flattened, obtuse or acute, 
shorter than the curved filaments, these 4-5 mm. long; style nearly 
filiform, 15 mm. long. The flowers are white, the petals apparently 
sometimes 17 mm. long. Eves' application of this name and A. 
dependens is followed, although in some herbaria at least the Ruiz 
& Pavon collections are not so named. 

Huanuco: Chinchao, Ruiz & Pavdn, type. Pampayacu, 
Kanehira 11. 

Axinaea grandifolia (Naud.) Triana, Trans. Linn. Soc. 28: 69. 
1871; 222. Chastenaea grandifolia Naud. Ann. Sci. Nat. se>. 3. 18: 
122. 1852. 

Upper branches obtusely tetragonal, pulverulent near the tip, 
as also the slender petioles, these 3-4 cm. long and transversely 



FLORA OF PERU 315 

tubercled apically; leaves narrowly ovate, 7-16 cm. long, often 6-7 
cm. wide, rounded but apiculate apically, shortly decurrent basally 
and the decurrent portion recurved or expanded and crisp, sinuate- 
dentate, 5-7-nerved, glabrous above, sometimes somewhat scurfy 
beneath, coriaceous; panicles erect, many-flowered, the pulverulent 
pedicels 8-12 mm. long; hypanthium broadly hemispheric, 7-8 mm. 
wide, glabrous or nearly, the limb subentire; petals retuse to apicu- 
late, 10-12 mm. long, purple or white; anther connective ovoid to 
rotund, apiculate to blunt, 2 mm. long; style slightly arcuate, 
7-8 mm. long. To this Eves refers Sawada 68 (here placed in A. 
nitida) which, though it has smaller (7x3 cm.), less pubescent 
leaves, lacks the recurved decurrent auricles. F.M. Neg. 16900. 

Peru: (fide Eves). Colombia; Venezuela. 

Axinaea lanceolata R. & P. Syst. 1: 122. 1798; Fl. Peruv. 4: 
pi. 409. 1802; 216. Chastenaea lanceolata (R. & P.) Naud. Ann. 
Sci. Nat. s<r. 3. 18: 123. 1852. 

Glabrous or nearly so, the upper branches acutely tetragonous, 
the slender petioles 5-6 cm. long, without the foliar tubercle, the 
leaves ovate-lanceolate, 1-2 dm. long, 3-10 cm. wide, bluntly acumi- 
nate, at base auriculate, the auricles conspicuously folded under, 
remotely and coarsely serrate, shortly 5-plinerved, coriaceous; 
panicles pendulous, few-flowered; pedicels sometimes scurfy, 5-10 
mm. long; calyx broadly campanulate, granular-puberulent or gla- 
brous, 6-11 mm. wide, the limb subtruncate; petals rounded or 
retuse, pink to white, 10-18 mm. long; anther sacs subulate, 5-8 mm. 
long, the ovoid connective obtuse or acute; style filiform, 5 mm. 
long. In the type the leaves are scurfy-puberulent and yellowish 
green beneath, becoming nearly glabrous; the pedicels are stout. 
Eves refers this negative to A. glandulosa; it seems to match cotype 
material from Madrid. F.M. Neg. 16901. 

Huanuco: Muiia and Panao, Ruiz & Pavon, type. San Dillani, 
Pearce. Bolivia. 

Axinaea Merianiae (DC.) Triana, Trans. Linn. Soc. 28: 69. 
1871; 224. Chastenaea Merianiae DC. Prodr. 3: 102. 1828. 

Glabrous shrub to 5 meters tall, with white or rose-colored flowers, 
the filaments white, the anthers yellow, the connectives dark red- 
violet; petioles slender, 1.5-2 cm. long, without foliar tubercle; 
leaves elliptic, 4-6 cm. long, 2.5-3 cm. wide, acute or obtuse, basally 
revolute-decurrent and this portion sometimes notably expanded and 
reflexed, sinuate-serrate, 3-5-plinerved, subcoriaceous; panicle some- 



316 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

what nodding, few-flowered; pedicel 10 mm. long; calyx shallowly 
campanulate or saucer-shaped, 7-10 mm. wide, the dilated, often 
reflexed limb truncate or slightly 5-sinuate; petals retuse, white to 
rose-colored, 12-15 mm. long; anther sacs 4 mm. long, the large 
blunt connectives nearly or quite as long; style attenuate, 9-15 mm. 
long. The Peruvian specimen was referred by Markgraf to A. 
affinis (Naud.) Cogn. of Ecuador and Colombia, with 5-nerved leaves, 
a broadly hemispheric calyx and connectives acuminate at one end 
(according to Eves). 

A. sclerophylla Triana, 216, of Ecuador, similar, has the leaves 
pubescent in the manner of A. tomentosa; A. macrophylla (Naud.) 
Triana, 223, found as near as Ecuador, differs in its 7-nerved leaves; 
the submarginal nerves are the sixth and seventh, in A. affinis, the 
fourth and fifth (Eves). 

Piura: Ayavaca, Weberbauer 6371. Libertad: Prov. Otuzco, 
Weberbauer 6997. 

Axinaea nitida Cogn. Bot. Jahrb. 42: 138. 1908; 220. 

Glabrous except for a slight scurfy puberulence on the younger 
petioles, peduncles and leaves beneath, the obscurely tetragonous 
branches becoming lustrous as the coriaceous leaves above, these 
narrowly ovate, 5-9 cm. long, 3.5-5 cm. broad, rounded at base and 
tubercled, shortly acuminate, undulate-denticulate, 5-nerved; pani- 
cles short, densely many-flowered, about 6 cm. long; pedicels 3-8 mm. 
long; calyx broadly hemispheric, 6-7 mm. broad, the limb truncate 
or undulately lobed; petals reddish, rounded to very slightly retuse, 
10-14 mm. long; anthers 5 mm. long, the ellipsoid connectives 
blunt, the nearly filiform style 11-13 mm. long. Distinguished from 
A. grandifolia (which compare) by the lustrous branchlets and upper 
leaf surfaces and by its never decurrent leaf bases. The Sawada 
shrub may be a new species; it is more tardily glabrate and less 
lustrous. F.M. Neg. 16904. 

Amazonas: Molinopampa, east of Chachapoyas, 2,000 meters, 
Weberbauer 4355, type; (Mathews 1287). Huanuco: Pan de Azucar, 
Sawada 68(!). 

Axinaea pauciflora Cogn. Melast. 448. 1891; 219. 

Branches rather slender, the upper obtusely tetragonous and 
with the petioles, cymes, calyces, and leaves beneath lightly scurfy- 
puberulent; petioles stout, 1 cm. long; leaves ovate, subacute at 
base, nearly obtuse, 6-10 cm. long, 4-5 cm. broad, 5-plinerved, entire 
except for a few minute calluses, rigid coriaceous, slightly yellowish- 



FLORA OF PERU 317 

scurfy beneath, the nerves extremely pronounced; cymes terminal, 
the 5 pedicels 1 cm. long in fruit, hispidulous-scurfy, the peduncle 
itself 15 mm. long; anthers linear, 5 mm. long, the anther sac narrowly 
ovoid, acute; style 1.5 cm. long. Possibly collected in Ecuador. 
A. Drakei Cogn., 218, of Ecuador, similar, has much longer glabrous 
leaves; A. lepidota (Benth.) Triana, 225, of the same country, is 
strongly marked by a pale arachnoid pubescence. F.M. Neg. 
36210. 

Peru: (?), Grisar, type. 

Axinaea Pennellii Gleason, Bull. Torrey Club 54: 28. 1927; 217. 

Branches subterete, obscurely 4-sided and rusty-scurfy toward the 
tip, somewhat enlarged at the nodes; petioles slender, 1.5-2 cm. long, 
with a more or less prominent but minute tubercle at the junction 
with the reflexed and overlapping leaf-auricles; leaves long-ovate, 
10-16 cm. long, 3.5-6 cm. wide, blunt-acuminate, conspicuously 
sinuate-serrate, 5-7-nerved, glabrous above, thinly yellowish-scurfy 
beneath, the auricles small; panicles erect, about 10-flowered; 
pedicels thinly scurfy, 5-10 mm. long; calyx broadly hemispheric, 
often glabrous, 10 mm. wide, the limb truncate; petals purple, 
retuse, 16 mm. long; filaments 7-10 mm. long; anther connective 
ovoijl, blunt, 5-6 mm. long; style 14 mm. long. 

Cuzco: Forest and cliff above Rio Pillahuata, Pennell 14003, type. 

Axinaea sessilifolia Triana, Trans. Linn. Soc. 28: 68. 1871; 213. 

In general similar to A. crassinoda but the leaves sessile or nearly, 
their cordate bases clasping the thick quadrate branches, conspicu- 
ously plinerved, to at least 1.5 dm. long, 7 cm. wide, green in color, 
but often lightly scurfy on the nerves beneath or glabrous; pedicels 
6-10 mm. long, glabrous; petals rounded or barely retuse, 13-15 mm. 
long; connectives with a conspicuous dorsal lobe, apiculate, 4 mm. 
long; style abruptly curved near the tip, 8-9 mm. long. A tree 
about 12 meters tall, the petals blood-red, the anthers yellow (Eves). 
F.M. Neg. 32326. 

Junin: Vitoc, Ruiz & Pavon (det. Markgr.). Ecuador. 

Axinaea tetragona Cogn. Bot. Jahrb. 42: 137. 1908. 

Shrub 2 meters high, with stout, acutely 4-angled, or nearly 
winged, glabrous branches, the nodes annulately enlarged, the annulus 
thick, wing-like; petioles slender, lightly scurfy, 3-6 cm. long; 
leaves subcoriaceous, ovate-cordate, acutish, 9-nerved, undulate- 



318 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

denticulate, 7-12 cm. long, 5-9 cm. wide, glabrous and somewhat 
bullate above, sparsely or scarcely scurfy-puberulent and reticulate- 
veined beneath; panicles 2.5 dm. long with few short, suberect 
branches; pedicels 6-10 mm. long; calyx glabrous or sparsely scurfy, 
7 mm. broad, obscurely lobed; petals obovate, broadly rounded, 
eciliate, 10-11 mm. long; anthers linear, 6 mm. long; style filiform, 
11-12 mm. long. Affine A. crassinoda Triana. Eves suggested 
that this might be a species of Meriania, but it seems clearly to 
belong here; the leaves are obscurely tubercled at base. F.M. 
Neg. 16905. 

Amazonas: East of Chachapoyas, near Tambo Bagazan, 2,700 
meters, Weberbauer 4430, type. 

Axinaea tomentosa Cogn. Melast. 451. 1891; 215. 

Branches conspicuously tetragonous, only slightly enlarged 
nodally, or stipule-scarred, furfuraceous as also the stout petioles, 
these 2-3.5 cm. long, with an apical tubercle, but this sometimes 
obscure; leaves broadly elliptic, 12-15 cm. long, 7.5-8 cm. wide, 
apiculate to obtusely acuminate, acute and slightly revolute at 
base, finely and remotely denticulate, 5-7-nerved or shortly plinerved, 
glabrous above, densely rusty-tomentose-villous beneath, with 
crinkly plumose trichomes; panicles erect, compact, many-flowered; 
pedicels densely scurfy, puberulent, thick, 2-5 mm. long; calyx 
broadly campanulate, 7-8 mm. wide, the limb truncate to shallowly 
5-lobed; petals obtuse to subretuse, 12-15 mm. long; filaments and 
anther sacs 6-8 mm. long, the ovoid connective with 2 large, some- 
what flattened lateral lobes; style slender, 17 mm. long. Perhaps 
a var. of A. glandulosa, more obviously pubescent, the trichomes 
longer. 

Amazonas: Chachapoyas, Mathews, type. Prov. Bongara, Weber- 
bauer 7150 (det. Eves). 

Axinaea Weberbaueri Cogn. Bot. Jahrb. 50: Beibl. Ill: 32. 
1913; 215. 

Nearly A. glandulosa, but the upper branches acutely 4-sided, 
the petioles slender, the basal leaf auricles small and somewhat 
recurved but not definitely reflexed; leaves very shortly and rather 
densely stellate- tomentose beneath; pedicels 2-6 mm. long; petals 
violet or purple, 10-16 mm. long, retuse; anther sacs flattened, with 
2 lateral wings extending downward from the connective; style 
slender, 6-15 mm. long. 

Ayacucho: Prov. of Huanta, 2,700 meters, Weberbauer 5648, type. 



FLORA OF PERU 319 

21. GRAFFENRIEDA DC. 

Glabrous, pulverulent, or rarely pubescent shrubs or trees, 
usually with large leaves and small 4-5-merous flowers, these often 
borne in ample terminal panicles. Calyx tube oblong-campanulate, 
the limb regularly lobed or the calyx irregularly splitting at anthesis. 
Stamens equal or subequal, the anthers subulate, the connective 
basally prolonged into a short sharp spur, erect or nearly erect by 
the incurving of the anther. Miconia flaviflora Gleason and M. 
firma Macbr. might be sought here, but both species lack the anther 
spur. Two keys are appended. 

Flowers 5-merous. 
Calyx splitting irregularly. 

Leaves merely 3-nerved G. limbata. 

Leaves 5-nerved or 5-plinerved. 

Leaves stellate-hirsute beneath G. acida. 

Leaves nearly glabrous G. foliosa. 

Calyx regularly lobed. 

Pedicels 1-4 mm. long; petals and calyx lobes rounded. 

G. Candelabrum. 
Pedicels obsolete; petals and calyx lobes acute. 

Leaves tomentose beneath G. stenopetala. 

Leaves hispid beneath G. cinna. 

Flowers 4-merous. 
Principal leaf nerves 5 or the leaves plinerved (cf. G. trichanthera). 

Leaves oblong-lanceolate, acuminate, often plinerved. 

G. Moritziana. 

Leaves ovate, acute or shortly acuminate, shortly if at all 

plinerved. 

Branches and petioles stout; ovary 4-celled. . . .G. emarginata. 
Branches and petioles slender; ovary 3-celled. .G. intermedia. 
Principal leaf nerves 3, basal. 

Leaves ample, 8 cm. wide or wider. 

Calyx tube about 3 mm. long, the lobes a third as long. 

G. floribunda. 

Calyx tube about 2 mm. long, the lobes half as long. 

G. miconioides. 

Leaves ovate-lanceolate, 5 cm. wide or narrower. .G. trichanthera. 



320 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Vegetative key 

Principal leaf nerves 3 (midrib and 1 pair, this at or near margin). 
Calyx tube 4-6 mm. long; inflorescence narrow. 

Inflorescence cinereous; calyx 5-6 mm. long G. limbata. 

Inflorescence glabrate; calyx about 4 mm. long. .G. trichanthera. 
Calyx tube 2-3 mm. long; inflorescence ample. 

Outer pair of nerves marginal G. floribunda- 

Outer pair of nerves submarginal G. miconioides. 

Principal leaf nerves 5 (1 pair about medial, another submarginal, 

often fainter) or the leaves plinerved. 
Leaves oblong-lanceolate. 

Leaves coriaceous, shortly 3-plinerved, acute. . . .G. trichanthera. 
Leaves membranous, often shortly 5-plinerved, acuminate. 

G. Moritziana. 

Leaves broader, ovate to elliptic or if oblongish only twice longer 
than broad. 

Flowers solitary on divaricate branchlets G. Candelabrum. 

Flowers if terminal crowded on short ascending branchlets. 
Leaves oblongish, 3-4 (4.5) cm. wide. 

Shrub nearly glabrous G. foliosa. 

Shrub markedly pubescent G. cinna. 

Leaves broadly ovate to elliptic. 
Leaves acute or acuminate. 

Pubescence stellate-hirsute G. acida. 

Pubescence compact, scurfy or scaly, often nearly lacking. 

Leaves glabrate G. intermedia. 

Leaves rusty scaly-tomentose G. stenopetala. 

Leaves rounded at tip and base G. emarginata. 

Graff enrieda acida Markgr. Notizbl. Bot. Gart. Berlin 13: 
462. 1937. 

Tree with greenish astringent-acidulous bark, obtusely tetragon- 
ous branchlets and short, more or less sparse, stellate-hirsute pub- 
escence extending to the calyces, the leaves only above soon glabrate; 
petioles to 3 cm. long; leaves elliptic, acute at both ends, 12-18 cm. 
long, 5-8 cm. wide, 5-plinerved, minutely dentate above the entire 
base; flowers 5-merous, cymose-corymbose, the pedicels 1-2 mm. 
long; calyx oblong, 4 mm. long, after anthesis the conical limb 



FLORA OF PERU 321 

irregularly lacerate; petals 5, fleshy, obovate, 10 mm. long; fila- 
ments rose-colored, 6 mm. long, nearly equaled by the dark-violet 
anthers; connective produced 1 mm. and shortly spurred posteri- 
orly, red; style 1.5 cm. long, ovary glabrous, 5-celled. Near G. 
foliosa. 

Cajamarca: Tambillo, Cutervo (Raimondi, 5 sheets, type). 
Colombia. 

Graffenrieda Candelabrum Macbr., spec. nov. 

Liana; caulibus petiolisque teretibus minute parceque cum pilis 
paullo stellulatis vel plumosis hispidulis mox glabratis; petiolis 1 cm. 
longis; foliis ovatis, basi rotundatis vel subacutis, acute acuminatis, 
circa 14 cm. longis, 6 cm. latis, 3-nerviis, nervis lateralibus ad mar- 
ginem proximis, nervis cum nervulis transversalibus et venulis retic- 
ulatis subtus praecipue prominentibus, utrinque glabris, nitidulis, 
chartaceo-coriaceis; paniculis amplis, candelabriformis, ramulis gra- 
cilibus; floribus solitariis, ut videtur lateralibus terminalibusque, 
sessilibus, 5-meris; calycis tubo oblongo-tubuloso fere 3 mm. longo, 
glabro, minute setoso-denticulato; petalis distincte acutis circa 3 mm. 
longis; antheris subulatis, connectivo basi calcarato; ovario villoso. 
By Gleason determined as G. patens Triana, 441, which Brazilian 
species, similar in foliage, has the flowers crowded at the tips of the 
panicle-branches. 

Loreto: Mishuyacu near Iquitos, Klug 413, type. 

Graffenrieda cinna Macbr., spec. nov. 

Arbuscula 2 m. alta; ramis petiolis paniculisque dense plumoso- 
furfuraceis et puberulentis; petiolis 1-2 cm. longis; foliis ovato- vel 
oblongo-ellipticis, basi acutis, apice conspicue et anguste caudatis, 
5-9 cm. longis, 3-4.5 cm. latis, supra glabris, nitidulis, subtus laxe 
nervis furfuraceis exceptis hispidis, breviter 5-plinervii; capitulis 
florum mediocriter confertis; floribus sessilibus 5-meris; calyce sub- 
obconico, 2.5 mm. longo, minute furfuraceo, lobis ovatis acutis; 
petalis lineari-lanceolatis, acuminatis, 3 mm. longis; antheris sub- 
ulatis, basi breviter calcaratis; ovario costato, furfuraceo, libero. 
Apparently referable to this genus but remarkable in the variety of 
character the partly simple pubescence, the 5-merous flowers, the 
acuminate petals. 

San Martin: Zepelacio, near Moyobamba, 100 meters, Klug 
3318, type. 



322 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Graffenrieda emarginata (R. & P.) Triana, Trans. Linn. Soc. 
28: 71. 1871; 442. Miconia emarginata R. & P. Syst. 1: 106. 1798; 
Fl. Peruv. 4: pi. 394. 1802. 

Branches and petioles robust, the latter 2-3.5 cm. long; leaves 
broadly ovate, basally rounded and emarginate, acutish, strikingly 
5-nerved, dark brown-scurfy beneath, glabrous above; panicle 
ample, many-flowered; calyces sessile, about 3 mm. long, the broadly 
ovate lobes 1 mm. long; petals 4, oblong-spathulate, 2 mm. long, 
white; ovary 4-celled, puberulent at tip. In the type, the brown- 
scurfy compound inflorescence is 2 dm. wide with a peduncle more 
than 3 dm. long, the leaves 15 cm. wide, 18 cm. long. F.M. Neg. 
16885. 

Huanuco: Chinchao, Ruiz & Pawn, type. San Martin: Tara- 
poto, Spruce 4566. Piura: Ayavaca, 2,600 meters, Weberbauer 6389 
(det. Markgr.). Bolivia; Colombia. 

Graffenrieda floribunda Triana, Trans. Linn. Soc. 28: 71. pi. 5. 
1871; 444. 

Similar to G. miconioides; leaves rigid-coriaceous, to 2.5 dm. long, 
18 cm. broad; flowers umbellate or verticillate, the pedicels 2-4 mm. 
long; calyx tube about 3 mm. long, the ovate lobes 1-1.5 mm. long; 
petals white, rose- tinted, 4-5 mm. long. The leaves are typically 
bluntly rounded at apex, but the species seems doubtfully distinct. 
F.M. Neg. 32325. Illustrated, Mart. Fl. Bras. 14, pt. 4: pi. 10. 

San Martin: Tarapoto, Spruce 4885, type. Near Moyobamba, 
Weberbauer 4631 (det. Cogn.), 290. Loreto: A 5-meter tree, Pum- 
ayacu, Klug 3139 (det. Gl.). 

Graffenrieda foliosa Cogn. Bot. Jahrb. 42: 137. 1908. 

Nearly glabrous, densely leafy shrub 6 meters high with stout, 
obtusely angled branches; leaves coriaceous on petioles to 2 cm. long, 
ovate-oblong, subrotund at base, obtuse, entire, 5-nerved, 6-9 cm. 
long, 3-4 cm. broad; panicles 8-12 cm. long, the many 5-merous 
flowers sessile; calyx lightly ashy-scurfy, the tube 4 mm. long, the 
limb rupturing irregularly into short persistent lobes; petals yellowish 
white, 2-3 mm. long; style to 8 mm. long. F.M. Neg. 16886. 

Huanuco: Prov. Huamalies, 2,000 meters, Weberbauer 3544, type. 

Graffenrieda intermedia Triana, Trans. Linn. Soc. 28: 71. 
1871; 442. 

Glabrate, except the scurfy inflorescence, this strict, the branches 
slender; petioles 2-4 cm. long; leaves firm, ovate-elliptic, shortly 



FLORA OF PERU 323 

and rather abruptly acuminate, acutish at the rounded base, slightly 
5-plinerved, to 15 cm. long, half as wide, often smaller; panicle 
many-flowered, scarcely 1 dm. long; calyx tube 2-2.5 mm. long, the 
4 broadly rounded membranous lobes hardly 1 mm. long; petals 3 
mm. long; ovary 3-celled; style 4-5 mm. long. The leaves are very 
shortly plinerved with a few scattered scales, dark green above, light 
colored beneath. F.M. Neg. 21168. 

San Martin : Tarapoto, Spruce 4258, type. 

Graff enrieda limbata Triana, Trans. Linn. Soc. 28: 70. 1871; 
440. 

Smooth, except for the ashy-scaly ample inflorescences; petioles 
4-5 cm. long; leaves 3-nerved, the lateral near the margin, the hori- 
zontal veins prominent, abruptly and shortly acute, narrowed to 
base, somewhat obovate, lighter colored beneath, 1.5-2 dm. long, 
6-10 cm. broad, sometimes smaller; panicles long-peduncled, with 
many flowers, the 5 obliquely-obovate petals 7 mm. long; calyx tube 
10-costate, 5-6 mm. long, the limb rupturing irregularly in 2-4 lobes, 
2 mm. long; ovary 4-5-celled, the capsule with as many ribs. An 
open shrub or small tree about 4 meters high, with narrow panicles 
of white flowers, large for the genus. Flowers pinkish-lavender with 
touch of white (Woytkowski) ; pale rose (Klug). 

Junin: La Merced, 5568. Colonia Perene", Killip & Smith 24953 
(det. Gl.). San Martin: Tarapoto, Ule 6456 (det. Pilger); sandy 
woods, Spruce 4261, type; Williams 5893 (det. Gl.); 5631. Near 
Moyobamba, 800 meters, Weberbauer 4569 (det. Cogn.); 290; Klug 
3252 (det. GL). Rioja, Woytkowski 6. "Nispero sacha," "dispera 
sacha." 

Graffenrieda miconioides Naud. Ann. Sci. Nat. se*r. 3. 18: 
118. 1852; 444. 

Terete branches as the ample panicles, slightly ashy-puberulent; 
petioles 2-3 cm. long; leaves obovate, oblong, acute or shortly acum- 
inate, acutish at base, moderately coriaceous, 8-18 cm. wide, some- 
what lustrous above; flowers subfasciculate or aggregate, the pedicels 
1-2 mm. long; calyx tube 2-2.5 mm. long, the broadly ovate lobes 
1 mm. long; petals 4, obovate, 6 mm. long, nearly as broad at the 
top; anthers strongly recurved, very acutely produced basally; 
ovary partly free, 3-celled. Trunk 9 meters high, 15 cm. in diameter 
(Tessmann). Compare G. floribunda. F.M. Neg. 36204. 

Loreto: Iquitos, Tessmann 5586 (det. Markgr.). Brazil. 



324 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Graff enrieda Moritziana Triana, Trans. Linn. Soc. 28: 70. 
1871; 443. 

Slenderly branched, somewhat scurfy-puberulent, especially the 
inflorescence; petioles 1-3 cm. long; leaves membranous, green both 
sides, but loosely scurfy on the nerves beneath, oblong-lanceolate, 
long-acuminate, acute or slightly rounded at base, distinctly retic- 
ulate, shortly 5-plinerved, about 12 cm. long, 5 cm. broad; panicles 
8-12 cm. long, the subverticillate pedicels filiform, 4-5 mm. long; 
calyx tube narrowly campanulate, 3-4 mm. long, the broadly tri- 
angular lobes 1 mm. long; petals 4, yellowish, broadly oblong, 4 mm. 
long; capsule lightly 8-costate. Apparently the younger branchlets 
are straggling. F.M. Neg. 16888. 

San Martin: Tocache, Poeppig 2045 (det. Krasser). Venezuela; 
Colombia. 

Graff enrieda stenopetala Ule, Notizbl. Bot. Gart. Berlin 6: 
353. 1915. 

Young branches, petioles, peduncles and calyces rusty-scurfy- 
tomentulose; petioles 4-5.5 cm. long; leaves subcoriaceous, ovate- 
elliptic, rounded at base, obtusely acuminate, about 15 cm. long, 7-9 
cm. wide, 7-plinerved and reticulate beneath, closely rusty- tomen- 
tose; flowers in panicled fascicles, 5-merous, sessile, bracteolate; 
calyx tube 3.5 mm. long, the acute teeth 1.5 mm. long; petals 
whitish, narrowly lanceolate-linear, acute, 5 mm. long, less than 1 
mm. broad; ovary 3-celled, costate, slightly scurfy; style 7 mm. 
long. The Peruvian collection differs from type in having only 
5 nerves, more loosely borne flowers and less pubescent inflorescence 
(Gleason). F.M. Neg. 16894. 

Loreto: Killip & Smith 24950 (det. Gl.). British Guiana. 

Graffenrieda trichanthera Gleason, Bull. Torrey Club 63: 
533. 1936. 

Small tree, glabrous except the minutely puberulent calyx; 
petioles to 10 mm. long; leaves coriaceous, slightly 3-plinerved, 
lanceolate, rounded at base, acute, entire, to 13 cm. long, 3.5 cm. 
wide or the smaller less than half as large; flowers 4-merous, sessile, 
crowded in dense heads; calyx tubular-campanulate, about 4 mm. 
long, at an thesis with 2-4 scarious irregular lobes; petals 3.2 mm. 
long; anther connective and short spur slightly setose; ovary free, 
2.3 mm. long, glabrous, 4-celled. 

San Martin: Zepelacio, near Moyobamba, 1,100 meters, Klug 
3298, type. 



FLORA OF PERU 325 

22. GALYPTRELLA Naud. 

Glabrous trees or shrubs, or the younger parts pulverulent, 
with entire, 3-7-nerved, or very shortly plinerved leaves and small 
flowers often subumbellate at the ends of the peduncles. Calyx 
tube pubescent, before anthesis crowned with the closed top-shaped 
limb. Connective somewhat produced, bearing a porrect appendage 
at its juncture with the filaments. Seeds needle-shaped. The genus 
may be distinguished in flower from Conostegia, with similar calyx, 
by the appendaged stamen and different seeds. 

Flowers 5-9-merous; leaves and inflorescences ample. . .C. cucullata. 
Flowers 4-merous. 

Leaves ample, stellate-puberulent beneath C. robusta. 

Leaves mostly narrower than 7 cm., glabrous or glabrate. 
Leaves 1 dm. long or longer, membranous; calyx scarcely costate 

or ecostate C. gracilis. 

Leaves 8 cm. long or shorter, coriaceous; calyx 8-costate. 

C. tristis. 

Calyptrella cucullata (Pav.) Triana, Trans. Linn. Soc. 28: 72. 
pi. 5. 1871; 453. Melastoma cucullata Pav. ex D. Don, Mem. Wern. 
Soc. 4: 317. 1823. Conostegia cucullata D. Don, I.e. 

Shrub or tree, the thick, somewhat fistulose branches obscurely 
tetragonous; petioles elongate, the leaf blades broadly ovate, dis- 
tinctly 5-7-nerved, often 2 dm. long or longer and fully half as 
wide to nearly as wide; panicles many-flowered, 1-3 dm. long, the 
pedicels 2-8 mm. long; calyx tube 2.5-3 mm. long, the caducous 
pointed limb 5 mm. long; petals 5-6 mm. long. A scaly puberulence 
is mostly confined to the calyx, the nodes and the petioles. Ants 
are sometimes found in the hollow branchlets. 

Huanuco: Pozuzo, 4701. Pampayacu, Kanehira 33. Casupi, 
Sawada 102. Valley of the Monzon, Weberbauer 8503; 284. Chin- 
chao, Ruiz & Pavdn, type. Junin: La Merced, 5596. Chanchamayo 
Valley, Schunke 266. San Martin: Tarapoto: Spruce 4329; 4899; 
Williams 6459; 7109 (det. Gl.). Ecuador. "Nispero," "dispero." 

Calyptrella gracilis Triana, Trans. Linn. Soc. 28: 72. 1871; 454. 

A small tree with lightly pulverulent, slender, 4-angled branchlets 
and small (1-2 dm. long), membranous, lanceolate-acuminate leaves, 
minutely and sparsely brown-scurfy beneath; three leaf nerves 
prominent, the 2 outer fainter ones joining the middle-laterals above 
the shortly attenuate leaf base; petioles 2 cm. long; petals yellow, 



326 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to 2 mm. long, ovate; anther connective shortly spurred dorsally. 
Var. ovata Huber, Bol. Mus. Goeldi 4: 595. 1906, has ovate leaves, 
truncate at base. F.M. Neg. 25867. 

Junin: Chanchamayo Valley, Schunke 440; 441. San Martin: 
Tarapoto, Spruce 4896, type. Rio Tocachi, Poeppig 2045. Loreto: 
Cerro de Canchahuaya (Huber 1427, type var.). 

Calyptrella robusta Cogn. Bot. Jahrb. 42: 138. 1908. 

Leaves densely stellate-puberulent beneath; calyces sessile, 
densely furfuraceous, 4-merous, the tube 2 mm. long; petals 1.5-2 
mm. long; capsule subglobose, 3 mm. thick. Otherwise similar to 
C. cucullata. F.M. Neg. 16882. 

Huanuco: In shrub-tree formation, 1,600 meters, Weberbauer 
3572, type. 

Calyptrella tristis Triana, Trans. Linn. Soc. 28: 72. 1871; 454. 

Shrub or small tree with rounded, ovate, bluntly acute, glabrous 
but distinctly papillose leaves more or less rounded at base, 3-5- 
nerved or very shortly plinerved, rather conspicuously ribbed, often 
8 cm. long and half as broad, sometimes much smaller; petioles about 
12 mm. long; inflorescence 4-6 cm. long, narrow in flower; petals 
white, 2-2.5 mm. long; calyx, especially in fruit, conspicuously 
ribbed. F.M. Neg. 16883. 

San Martin: Tarapoto, Spruce 4823, type; Williams 5987 (det. 
Gl.). Loreto: Pumayucu, 2-meter shrub, 600-1,200 meters, Klug 
3213 (det. Gl.). 

23. CENTRONIA D. Don 

Similar to Calyptrella but the flowers large, showy, the anther 
connective not produced, the seeds pyramidal. Leaves entire or 
crenulate-dentate, often thick or fleshy, finely reticulate-veined 
beneath, glabrous or rusty-tomentose or pilose on the under surface, 
as sometimes also the branches. 

Leaves 5-7-nerved, tomentose or pilose beneath. 

Pubescence dense, tomentose C. excelsa. 

Pubescence loose, pilose C. peruviana. 

Leaves 3-nerved, glabrous. 

Calyx scurfy-tomentose C. reticulata. 

Calyx densely setose C. laurifolia. 



FLORA OF PERU 327 

Centronia excelsa (Bonpl.) Triana, Trans. Linn. Soc. 28: 72. 
pi. 5. 1871; 460. Rhexia excelsa Bonpl. Rhex. 90. pi. 34- 1823. 

Stems herbaceous, often rooting, 2-3 meters high, the branches 
tetragonous, brown-scurfy or glabrate in age; petioles 2-5 cm. long; 
leaves ample, often somewhat plinerved, broadly ovate, crenulate, 
glabrous and more or less bullate-reticulate-veined above, at first 
densely and softly tomentose beneath; calyx densely tomentose, 
1.5 cm. long and broad in fruit; flowers 5-6-merous; anther connec- 
tive appendaged posteriorly; ovary smooth. 

Amazonas: Chachapoyas, Mathews. Ecuador. 

Centronia laurifolia D. Don, Mem. Wern. Soc. 4: 315. 
1823; 455. 

A tree with stout, glabrous branches and petioles, and coriaceous, 
glabrous, elliptic-caudate, acuminate, entire, 3-plinerved leaves, 
to 2.5 dm. long, 10 cm. wide; flowers subsessile, the broadly ovate, 
subacute petals 1 cm. long; anthers 7-8 mm. long, with a very sharp 
spur; style nearly filiform, 12 mm. long; ovary dissected apically 
into 5-6 spui^-like projections. The calyx is conspicuously setose, 
the flower bud 18 mm. long. F.M. Neg. 29495. 

Huanuco: Chicoplaya, Ruiz & Pawn, type. 

Centronia peruviana Macbr., spec. nov. 

Arbuscula 3 m. alta; ramulis petiolis paniculisque laxe denseque 
rufo-villosis; petiolis 2-5 cm. longis; foliis ellipticis basi rotundatis, 
apice acutis vel paullo acuminatis, manifeste calloso-crenulatis, 
breviter 5-plinerviis, 18 cm. longis, 10 cm. latis, supra glabris, obscure 
vel leviter et grosse reticulato-venosis, subtus longe fulvo-pilosis 
praecipue ad nervos nervulosque et stellato-pubescentibus, haud 
bullatis; pedicellis 7 mm. longis; alabastris solum apiculatis; calycis 
tubo 12 mm. longo haud sulcato, limbo circa 15 mm. longo; petalis 
2.4 cm. longis; filamentis antheraeque subaequalibus 10 et 12 mm. 
longis, connectivo appendice obtuso instructo, appendice superne 
leviter tuberculato; ovario vertice obscure 5-lobato; stylo 2 cm. 
longo. Nearly C. grandiflora Standl. of Panama with more acumi- 
nate bullate-reticulate leaves; both species seem to be closely allied 
to C. tomentosa Cogn., 460, of Ecuador and C. excelsa, species with 
short, crinkly-tomentose pubescence. C. peruviana, further, has a 
different connective appendage than the latter but apparently the 
same as that of the former; C. tomentosa, however, has a beaked bud, 
the beak 5-parted. Flowers between pink and rose (Stork & Horton). 



328 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Carpish, bank of mountain stream, 2,850 meters, 
Stork & Horton 9928, type. 

Centronia reticulata Triana, Trans. Linn. Soc. 28: 71. pi. 5. 
1871; 456. 

Similar to C. laurifolia but the leaves nearly basally 3-nerved, 
the calyx densely furfuraceous and not at all or only sparsely setose; 
pedicels 1-3 mm. long; anthers 5-6 mm. long, the cells crisply un- 
dulate, the connective shortly spurred; ovary 5-celled, apically 
dentate; style 9 mm. long. 

San Martin: Mountains of La Campana, Spruce 4840, type. 
Loreto: Cerro de Escaler, Ule 6747 (det. Pilger). Huanuco: Chico- 
playo, Ruiz & Pavdn (det. Markgr.). 

24. TESSMANNIANTHUS Markgr. 

Tree with large 3-nerved glabrous leaves, the lateral nerves near 
the margin and small 5-merous flowers with obtusely acuminate 
petals, borne in large terminal panicles. Calyx campanulate, 
truncate. Stamens very unequal and dissimilarly appendaged (con- 
nective not produced), the larger with 2 acuminate dorsal appendages, 
the smaller obtuse, parted, dehiscent by one broad pore, the connec- 
tive indistinctly biauricled posteriorly. Allied to Graffenrieda 
(Markgr.). 

Tessmannianthus heterostemon Markgr. Notizbl. Bot. Gart. 
Berlin 9: 1141. 1927. 

Tree 30 meters high, with obovate-obtuse leaves to 1.5 dm. 
long, 8 cm. wide; inflorescence paniculate, 12 cm. long, 8 cm. wide, 
ebracteate; pedicels 3 mm. long; calyx glabrous, 4 mm. long, 4 mm. 
across, the lobes scarcely marked; petals 4 mm. long; longer filaments 
4 mm., anthers 7 mm., the appendages 2 mm. long; shorter filaments 
2.5 mm., anthers 3 mm. long, divergent 1 mm. at apex, auricles 0.5 
mm. long; style 8 mm. long; ovary 3-celled, glabrous. F.M. Neg. 
17313. 

Loreto: Mouth of Santiago, Tessmann 4585, type. 

25. ALLONEURON Pilger 
Meiandra Markgr. Notizbl. Bot. Gart. Berlin 9: 1142. 1927. 

Shrubs or trees with pinnately veined leaves like Mouriria but 
with capsular fruit. Leaves crowded at the ends of the branches, 
long-elliptical-lanceolate or narrowly cuneiform, ciliate, narrowed to 



FLORA OF PERU 329 

the rather abruptly dilated base. Flowers small, 4- or 5-merous, 
in axillary panicles with few long, slender, or filiform branches. 
Anther connective shortly spurred. Ovary inferior, 2-celled. 

Alloneuron ma jus Markgr. ex Macbr. Trop. Woods 17: 13. 
1929. Meiandra maior Markgr. Notizbl. Bot. Gart. Berlin 9: 1142. 
1927. 

A tree to 30 meters high with few, many-flowered panicles of 
small white blossoms; leaves chartaceous, glabrous except the 
ciliate margins, 6-8 cm. wide; inflorescence 15 cm. long, about 
6 cm. wide, scurf y-puberulent, the flowers secundly crowded, the 
4 petals 3 mm. long. The specific name inadvertently written 
maior by me in Tropical Woods. Illustrated, Markgr. I.e., page 
1143. F.M. Neg. 17395. 

Loreto: Mouth of Rio Santiago, Tessmann 4625, type. 

Alloneuron Ulei Pilger, Verh. Bot. Ver. Brandenb. 47: 185. 
1905. Meiandra minor Markgr. Notizbl. Bot. Gart. Berlin 9: 1142. 
1927. 

Similar to A. majus but leaves smaller; inflorescence to about 
9 cm. long, a third as broad, the flowers scattered on the filiform 
branches; petals 5, 2.5 mm. long, yellowish. F.M. Neg. 17396. 

Loreto: In rocks at the Pongo de Cainarachi, Vie 6417, type. 
Pongo de Manseriche, Tessmann 4853. 

26. MACROCENTRUM Hook f. 

Lax or weak-stemmed, usually branching, nearly glabrous herb 
with ciliate, oblong-ovate, 3-5-nerved leaves, small 4-5-merous 
flowers borne in secund cymes or solitary on long peduncles. Sta- 
mens alike or nearly, the anther connective with 1 slender spur at 
base directed backward. Ovary free, 3-celled. Capsule 3-cornered, 
each enclosed in a persistent deeply 8-10-costate calyx. Cf . Salpinga, 
with which Baillon merged it. 

Macrocentrum fasciculatum (L. C. Rich.) Triana, Trans. 
Linn. Soc. 28: 79. 1871; 534. Aulacidium fasciculatum L. C. Rich, 
ex DC. Prodr. 3: 113. 1828. M. peruviana (Cogn.) Macbr. Field 
Mus. Bot. 4: 177. 1929. Salpinga fasciculata DC. I.e. 

Stems simple, 1-2 dm. long; petioles 5-13 mm. long; leaves 2-3 
cm. long, 10-17 mm. broad; calyx in fruit 13-14 mm. long; flowers 
pale lilac. This is the var. peruviana Cogn. Bot. Jahrb. 42: 138. 
1908; the typical form is often much-branched, the petioles longer, 



330 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the leaves larger, but the fruiting calyx only 8-9 mm. long, and may 
be specifically distinct. The similar M. cristatum (DC.) Triana has 
4-merous flowers, 8-costate calyx. F.M. Neg. 16916 (var.). Illus- 
trated, Mart. Fl. Bras. 14, pt. 4: pi. 14. 

San Martin: North of Moyobamba, 1,500 meters, Weberbauer 
4645. French Guiana. 

27. SALPINGA Mart. 

More or less suffrutescent and branched or herbaceous, simple- 
stemmed herbs with ovate 5-7-nerved thin leaves and medium-sized 
white or roseate flowers, sessile or subsessile, secund. Calyx tube 
thick-ribbed, enclosing the 3-angled capsule, the sepals obscure. 
Stamens 10, isomorphic or nearly, the connective of the subulate 
anthers with an elongate linear appendage at the base directed for- 
ward and a minute basal spur. 

Salpinga cilia ta Pilger, Verh. Bot. Ver. Brandenb. 47: 167. 1905. 

Ascending or decumbent, about 1 dm. high; leaves 4.5-8 cm. 
long, 3-5 cm. broad, densely long-fimbriate-ciliate on the margin. 
F.M. Neg. 16913. 

Loreto: Cerro de Cumbasso, 800 meters, Ule 6743, type. 

Salpinga secunda Schrank & Mart, ex DC. Prodr. 3: 113. 
1828; 536. 

Stems erect, often suffrutescent below and branched or simple, 
fleshy above; leaves 6-10 cm. long, 5-7 cm. broad, sparsely or 
minutely and inconspicuously ciliate on the margins, sometimes a 
little hirsute above, thin, crowded, cuneate at base; fruiting calyx 
elongating to 16 mm., deeply 10-sulcate. F.M. Neg. 16914. 

Loreto: Sarayacu to Catalina, Pampa del Sacramento (Huber 
1509). Iquitos, Ule 6744 (also many collections by Killip & Smith, 
as 29921, and King 517, all det. Gl.). Timbuchi on Rio Nanay. 
Williams 944; 945. To Guiana. "Puca mullaca," "mullaca azul." 

28. MONOLENA Triana ex Benth. & Hook. 

Stemless, succulent, glabrous herbs with large flowers borne in 
scorpoid cymes on short slender scapes. Calyx 3-angled, glabrous, 
the lobes simple. Stamen connective anteriorly spurred and poster- 
iorly tubercled. Capsule topped with the persistent 3-lobed base of 
the style. 



FLORA OF PERU 331 

Monolena primulaeflora Hook. f. Bot. Mag. pi. 5818. 1870; 538. 

Leaves firm but drying thin, broadly elliptic, 3-5-plinerved, 
often red-purple beneath. 

Loreto: Mouth of Santiago, Tessmann 4300 (det. Markgr.). 
San Martin : Moyobamba, Weberbauer 4731 ; 288. Without locality 
(Spruce; Lechler 2370). Rio Acre: Ule 9668. Junin: Pichis Trail, 
Killip & Smith 26131; 26505 (det. Gl.). Colombia. 

29. TRIOLENA Naud. 

More or less pubescent herbs with large, 5-7-plinerved leaves 
and small flowers borne in scorpoid spikes or racemes. Connective 
of the larger anthers anteriorly 3-winged. Calyx finally 3- winged, 
enclosing the capsule, widely curved at summit. This genus and 
Monolena and Diolena are only academic segregates of Salpinga; 
the aspect is exactly that of Diolena. Rationally, similar segregation 
is required in Bother tribes. 

Triolena Barbeyana Cogn. Melast. 542. 1891. 

Leaves narrowly ovate, distinctly unequal in each pair, glabrous 
above, puberulent on the nerves beneath; calyx densely hirsute. 
The similar T. hirsuta Triana has been collected in adjacent lands; 
its nearly equal leaves are very sparsely setulose or glabrous above 
but appressed hirsutulous on the nerves beneath. T. pustulata 
Triana, of Ecuador, is similar but the oblongish leaves are cordate 
at base, bullate above. 

Peru(?): Without locality, Pavon, type. 

30. DIOLENA Naud. 

Herbs, usually simple with often unequal ovate-oblong or lanceo- 
late, acute or acuminate leaves and white flowers secundly borne in 
scorpoid racemes or spikes. Stamen connective anteriorly 2-spurred. 
With Triolena, very similar to Salpinga. 

Diolena amazonica Pilger, Verh. Bot. Ver. Brandenb. 47: 168. 
1905. 

Stems creeping and rooting at base, subligneous, 1.5-3 dm. high, 
above with the petioles densely villous-setose; leaves unequal in 
each pair, the larger irregularly subelliptic, acuminate, shortly 
villous-setose beneath on the reticulate nerves, the margin irregularly 
serrulate and setose, 11-12 cm. long, about half as wide; smaller leaves 
lanceolate-elliptic, 4-6 cm. long; rachis setose-villous, 5-6 cm. long; 



332 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

petals white, 6-7 mm. long; stamens unequal, 5 with connective 
merely tuberculate not long produced; fruit 3-winged, excavated 
apically, 9 mm. broad. F.M. Neg. 16906. 

Junin: Puerto Bermudez, Killip & Smith 26660 (det. GL). Pebas, 
Killip & Smith 25385; Rio Pichis, Killip & Smith 26800. Loreto: 
Williams 1804; 3421. Soledad, Killip & Smith 29568. Brazil. 

31. LEANDRA Raddi 

Herbs or low shrubs conveniently grouped apart from Miconia 
by their ovate to linear, always acute or aristate petals, the bristle 
if present borne below the petal tip. Anther connective often ele- 
vated near base, rarely obscurely or minutely prolonged. Even 
though there are now known some species of Miconia with pointed 
petals, these are typical of that genus in other respects, that is, 
especially the anthers are somewhat appendaged or the connective 
spurred; there is a certain aspect, too, to most species of Leandra 
distinct from that of Miconia, yet the genus, all species considered, 
is scarcely a natural concept. Inflorescence really terminal but 
some species sometimes suggest Ossaea. Besides the following, 
the wood of Williams 45 from lower Rio Itaya has been described 
by the collector, Field Mus. Bot. 15: 381. 1936, under the name 
"yutobanco"; the collection is in fruit and may not be a Leandra. 

Flowers not borne secundly. 
Flowers fasciculate, sessile. 
Leaves oblong or nearly, to about 2 cm. wide. 

Pubescence stellate-scurfy L. nervosa. 

Pubescence mostly hirsutulous L. Cogniauxii. 

Leaves broader and longer. 

Branches setose; calyx 4-5 mm. long. 
Leaves 3-5 (-7) cm. wide; calyx nearly 3 mm. wide. 

L. crenata. 
Leaves 6.5-9.5 cm. wide; calyx about 2 mm. wide. 

L. Raimondiana. 
Branches appressed-sericeous-pubescent; calyx about 2 mm. 

long L. Francavillana. 

Flowers solitary or in any case pedicellate and not secund. 

Leaves 3-nerved L. cuspidata. 

Leaves 5-7-nerved or plinerved. 

Leaves rounded-cordate or unequal at base. 



FLORA OF PERU 333 

Branches glabrous; leaves basally inequilateral. 

L. Boissieriana. 
Branches hirtellous; leaves equally rounded at base. 

L. aristigera. 

Leaves attenuate to base; the branches glabrous or hirsute. 

L. chaetodon. 
Flowers borne secundly on the spreading inflorescence branches, 

sessile. 
Pubescence of stems and petioles ascending-appressed, sometimes 

becoming loosely crisped. 

Leaves densely appressed-pilose beneath; inflorescence a narrow, 
elongate thyrse, ample, the aggregate flowers not obviously 

secund L. Francavillana. 

Leaves lightly pilose beneath; inflorescence small, about as 

broad as long, open, the flowers markedly secund. 
Flowers 4-merous; anthers 1 mm. long; ovary 3-celled, gla- 
brous; leaves densely minutely denticulate. 

L. secundiflora. 
Flowers 5-merous; anthers, except in L. divaricata with 

subentire leaves, about 1.5 mm. long. 
Leaves subentire; ovary 3-celled, glabrous; calyx eglandular. 

L. divaricata. 
Leaves densely denticulate; ovary 5-celled, setulose; calyx 

usually somewhat glandular L. secunda. 

Pubescence of stems and petioles definitely divaricate, usually 

more or less reflexed or arcuately reversing. 
Pubescence of branchlets glandular, divaricate, short; ovary 

6-celled L. solenifera. 

Pubescence of branchlets eglandular, often more or less reflexed, 

long; ovary 3-5-celled. 
Leaves at least twice longer than broad; ovary 5-celled or, 

if 3-celled, pubescence closely reflexed. 
Calyx trichomes about 2.5 mm. long or longer; ovary 

setulose. 
Leaves 5(-7)-nerved; petals 2 mm. long. . . .L. longicoma. 

Leaves 7-9-nerved; petals 3-4 mm. long L. reversa. 

Calyx trichomes about 2 mm. long or shorter; ovary 

glabrous or minutely glandular. 
Pubescence dense, reflexed-arcuate; ovary glandular. 

L. dichotoma. 



334 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence reflexed, subappressed; ovary glabrous. 

L. retropila. 

Leaves rounded-ovate, less than twice longer than broad; 
ovary 3-celled; pubescence divaricate L. rotundifolia. 

Leandra aristigera (Naud.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
185. 1886; 683. Clidemia aristigera Naud. Ann. Sci. Nat. se"r. 3. 
17: 366. 1851. 

Young branches, petioles and inflorescence densely hirtellous 
with long-spreading, mostly minutely gland-tipped trichomes; leaves 
very shortly 5-7-plinerved, rounded at base, caudate-acuminate, 
loosely long-hirtellous beneath, sparsely so above, about 15 cm. long, 
9 cm. wide, thin; calyx densely glandular-hispid, the tube 3 mm. 
long, the filiform bristle- tipped teeth 1.5 mm. long; petals ovate- 
oblong, stellate-scurfy without, 2 mm. long, bearing on the back 
below the tip a glandular setulose bristle 3-4 mm. long; style 5 mm. 
long. F.M. Neg. 36222. 

Lima: Near Lima, Gay. Brazil. 

Leandra Boissieriana Cogn. in Mart. Fl. Bras. 14, pt. 4: 184. 
1886; 683. 

Branches glabrous; petioles and leaf margins ciliate-hirsute, the 
leaves ovate, scarcely attenuate to the rounded base, slenderly 
acuminate, 10-13 cm. long, 6 cm. wide, 5-7-plinerved; pedicels 
bibracteolate, 4-8 mm. long; calyx densely glandular-hirsute, 3.5-4 
mm. long, the teeth 1.5 mm. long; petals triangular subulate, long- 
acuminate-aristate, glandular-pilose dorsally, 3-4 mm. long; style 
4 mm. long. F.M. Neg. 34166. 

Huanuco(?): Ruiz & Pavon, type. 

Leandra chaetodon [Schrank & Mart.J Cogn. in Mart. Fl. 
Bras. 14, pt. 4: 178. 1886; 678. Melastoma chaetodon Schrank & 
Mart, ex DC. Prodr. 3 : 117. 1828, in syn. Spennera chaetodon DC. I.e. 

Branches slender, glabrous; petioles 5 mm. long or longer, sparsely 
glandular-ciliate as the young leaves, these soon glabrous, lanceolate- 
ovate, attenuate at base, long and narrowly acuminate, to 16 cm. 
long, 4-6 cm. wide, 5-plinerved, the 2 basal nerves faint; flowers 
in glabrous or glabrate corymbs or racemes, 2-6 cm. long, the white 
lanceolate-subulate petals 2-3 mm. long; calyx tube 2-3 mm. long, 
the teeth 1-2 mm. long; fruit blue, subglobose, 4 mm. thick. F.M. 
Neg. 6414. 



FLORA OF PERU 335 

Huanuco: Pueblo Nuevo, Ruiz & Pavon. San Martin: Tocachi, 
Poeppig 1869.Jumn: Pichis Trail, Killip & Smith 26211 (det. Gl.). 
Loreto: Killip & Smith 26928. Iquitos, Killip & Smith 26971; 
Williams 1328; 3762 (det. GL). La Victoria, Williams 2687; Rio 
Nanay, Williams 835. Without locality: Gay. Venezuela; Brazil; 
Colombia. "Sinchi mullaca." 

Leandra Cogniauxii Ule, Notizbl. Bot. Gart. Berlin 6: 367. 
1915. L. purpurascens Cogn. Bot. Jahrb. 42: 139. 1908, not 
(DC.) Cogn. 

Branches, petioles and peduncles densely setulose with ascending 
reddish trichomes; petioles 2-5 mm. long; leaves rigid, drying black, 
and minutely scabrous above, slightly stellate and setulose on the 
veins and 5 nerves beneath, narrowly oblong, little narrowed basally, 
acutish, 3-5.5 cm. long, 6-15 mm. broad; calyx sparsely scurfy and 
setulose, sessile, the narrowly campanulate tube 2 mm. long, the 
outer teeth 0.5 mm. long; petals pale purple, narrowly triangular, 
about 3 mm. long; style 5-6 mm. long, the ovary glabrate. Allied 
to L. nervosa, the inflorescence a narrow thyrse of glomerate flowers. 
F.M. Neg. 16926. 

San Martin: North of Moyobamba, 1,100 meters, Weberbauer 
4604, type. 

Leandra crenata (Pavon) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
137. 1886; 649. Melastoma crenata Pav. ex D. Don, Mem. Wern. 
Soc. 4: 308. 1823, in syn. Clidemia crenata D. Don, I.e. 

Branches more or less harshly hispid, the younger densely so; 
leaves 5- or partly 7-nerved (the outer 2 nerves faint and only basal), 
densely hirsute-strigillose above, villous-hirtellous beneath; petioles 
1-2 cm. long; panicle thyrsoid; calyx densely long-hirsutulous, the 
tube 4-5 mm. long, the outer teeth half as long; petals yellowish, 
oblong-lanceolate, 4 mm. long; style about 10 mm. long. Flowers 
white to pink (Stork & Horton). The leaves are often 8-10 (12) cm. 
long, 3.5-4 (5) cm. wide, gradually acuminate, minutely cordate at 
base, the surface generally concealed by the fulvous stiff pubescence; 
the aspect of the shrub, which is sometimes 2 meters high, suggests 
Clidemia. F.M. Neg. 16929. 

Huanuco: Cuchero, Poeppig 1397; Dombey. Casapi, Poeppig 83. 
Puno: Valley of Rio Sandia, Weberbauer 1103 (det. Cogn.; 278). 
Huancavelica: East of Surcubamba, in bushwood, 2,700 meters, 
Stork & Horton 10349. Without locality, Pavon, type. Paraguay 
to Bolivia and Brazil. 



336 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leandra cuspidata (Triana) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
175. 1886; 675. Oxymeris cuspidata Triana, Trans. Linn. Soc. 28: 
93. 1871. 

Younger branches more or less densely stellate-furfuraceous, the 
oblong-lanceolate, long-caudate-acuminate leaves scarcely so beneath, 
glabrous and lustrous above, subcoriaceous, unequal, the larger 
7-10 cm. long, 2.5-4 cm. wide, the smaller hardly half as long; 
panicles many-flowered, often nodding, 6-10 cm. long, sometimes 
very diffuse; pedicels 1-3 mm. long; calyx tube 2 mm. long, obtusely 
5-lobed; petals linear-subulate, 2 mm. long; style 3 mm. long; 
fruit black, globose, 4 mm. thick. Species well-marked by the 
3-nerved leaves and the open panicles. F.M. Negs. 36225; 21173. 

Loreto: Mouth of Santiago, Tessmann 5519 (det. Markgr.). 
Brazil. 

Leandra dichotoma (Pav.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
200. 1886; 692. Melastoma dichotoma Pav. ex D. Don, Mem. Wern. 
Soc. 4: 307. 1823, in syn. Clidemia dichotoma D. Don, I.e. 

Pubescence of branches more or less spreading and crisply 
hirsutulous, red or reddish-brown, drying black at times, eglandular; 
petioles elongate; leaves ovate, acuminate, 7-9-nerved, membranous, 
rather densely villous, especially beneath; panicle-branches secund; 
inflorescence and calyx pubescence dense and spreading, eglandular; 
calyx tube 2.5 mm. long, the teeth scarcely 1 mm. long; petals 
lanceolate-acuminate, about 2.5 mm. long, style 4 mm. long, the 
fruit 4-5 mm. thick, purple. The pubescence, at least sometimes, 
is notably soft, the inflorescence branches deep red. F.M. Neg. 
16935. 

Junin: Chanchamayo Valley, Schunke 324- San Ramon, Killip 
& Smith 24736 (fide Gleason, L. reversa}. La Merced, 5509; Schunke 
98; 149; 101; 324; 1484; 1456. Huanuco: Pampayacu, 5041; 
Kanehira 39. Chicoplaya, Ruiz & Pawn, type. Cuchero, Poeppig 
1448. San Martin: Tocachi, Poeppig 1822. Tarapoto, Spruce 4520; 
Williams 7186 (det. Gl.). Near Moyobamba, Klug 3485. San 
Roque, Williams 6934. Loreto: Rio Mazan, Schunke 173; Killip 
& Smith 24803. Rio Acre: Ule 9680. Bolivia to Central America. 

Leandra divaricata (Naud.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
196. 1886; 689. Clidemia divaricata Naud. Ann. Sci. Nat. se>. 3. 17: 
373. 1851. 

Often marked by the pubescence of the branches, this rigid, 
erect, appressed; petioles 1-4 cm. long; leaves ovate, acuminate, 



FLORA OF PERU 337 

shortly 5-plinerved, sparsely setulose above, pilose beneath, (5) 
8-9 cm. long, about 3 cm. broad; panicle small, few-flowered, the 
divaricate branches secund; flowers 5-merous, remote, the linear- 
subulate petals 1.5-2 mm. long, the calyx tube as long, its teeth 
nearly 1 mm. long; anthers about 1 mm. long; style only 2.5 mm. 
long; ovary glabrous, 3-celled. The leaves of the Guiana form may 
be 3-6 cm. wide, and the Williams and Schunke specimens with leaves 
3 cm. wide may belong elsewhere. 

Loreto: Yurimaguas, Poeppig; Nauta, Spruce 3858 (det. K. 
Rechinger); Rio Mazan, Schunke 252; Rio Nanay, Williams 783. 
Brazil to Costa Rica and British Guiana. 

Leandra Francavillana Cogn. in Mart. Fl. Bras. 14, pt. 4: 
197. pi. 43. 1886; 690. 

A tall shrub with ovate, beautifully acuminate, 7-plinerved or 
7-nerved leaves, densely appressed-pilose beneath with long-rusty 
silky trichomes, sparsely and minutely setulose above, about 1 dm. 
long and half as broad; inflorescence a narrow thyrse, the 5-merous 
flowers subglomerate, the linear subulate petals 2 mm. long, the 
calyx tube and calyx teeth as long; ovary 3-celled, setulose at apex, 
the style 4 mm. long; anthers 1-1.5 mm. long. The following collec- 
tions were referred here by Gleason but seem to be L. secunda or 
new; the inflorescence is diffuse, the flowers secund, the leaves 
sparsely hairy, shortly acuminate. F.M. Neg. 36230. 

Junin: Pichis Trail, Killip & Smith 26223. Loreto: Near Iquitos, 
(^Williams 8238; Mexia 6437. La Victoria, Williams 2686; 2555. 
Rio Mazan, Schunke 70. 

Leandra longicoma Cogn. in Mart. Fl. Bras. 14, pt. 4: 202. 

pi. 43. 1886; 693. 

Similar to L. dichotoma but the 5-nerved leaves only sparsely pi- 
lose beneath and the calyx pubescence very long, the petals some- 
what shorter, 2 mm. long. Calyx typically shaggy-villous, the outer 
lobes little shorter than the tube. The leaves are slightly plinerved 
in the type; apparently common in department of Loreto, many 
specimens of recent collectors being so determined by Gleason. 
The calyces vary in this material. The flowers are yellow, the 
fruit blue (Schunke). F.M. Neg. 32348. 

Loreto: Yurimaguas, Poeppig, type; Williams 7838. Pampas del 
Ponasa, Ule 6740 (det. Pilger). Lower Rio Nanay, Williams 600; 
631; 377; 704- Pebas, Williams 1693. La Victoria, Williams 2708. 
Caballo-Cocha, Williams 2327. San Antonio, Rio Itaya, Killip & 



338 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Smith 29453. Near Iquitos, King 1150. Rio Mazan, Schunke 387. 
-Rio Acre: Ule 9670. Costa Rica. "Muyaca," "mullaca." 

Leandra nervosa (Naud.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
73. 1886; 657. Clidemia nervosa Naud. Ann. Sci. Nat. se>. 3. 17: 
341. 1851. 

Branches, petioles and peduncles scurfy-stellate, some setae 
intermixed ; petioles 3-7 mm. long; leaves 5-nerved, oblong-lanceolate, 
acuminate, only 1.5-2 cm. wide, 8-10 cm. long, glabrous above, a 
little stellate-furfuraceous beneath or villous; panicles compact, 5-7 
cm. long; calyx sessile, puberulent-scurfy, the tube 2.5 mm. long, the 
outer teeth scarcely 0.5 mm. long; petals white, oblong, shortly 
acuminate, 3 mm. long; ovary scarcely pilose apically. F.M. Neg. 
25908. 

Amazonas: Chachapoyas, Mathews 1265. Ayacucho: Ccarrapa, 
Killip & Smith 22370 (det. Gl.). To Colombia. 

Leandra Raimondiana Markgr. Notizbl. Bot. Gart. Berlin 13: 
464. 1937. 

Similar to L. crenata; branchlets quadrangular; petioles, inflo- 
rescence and leaves beneath densely villous, the latter broadly ovate, 
9.5-13 cm. long, 6.5-9.5 cm. wide, 7-plinerved, papillose-hispid above; 
outer calyx teeth less than 1 mm. long; ovary apically setose; style 
5 mm. long; filaments 2 mm. long. Calyx 2 mm. across, about half 
as wide as in the related species. 

Puno: Isilluma (Raimondi, type). 

Leandra retropila Cogn. in Mart. Fl. Bras. 14, pt. 4: 198. 
1886; 690. 

Branches hirsute-setulose with more or less deflexed trichomes; 
leaves very membranous, 5-nerved, nearly oblong, to about 1 dm. 
long, 3.5-5 cm. broad, rounded or obtuse at base, acuminate, sparsely 
asperous above, pilose-puberulent beneath; calyx tube 2.5 mm. long, 
the teeth about half as long; petals triangular-lanceolate, 2 mm. long; 
ovary glabrous. F.M. Neg. 16970. 

San Martin: Rio Tocachi, Poeppig 2044- Ayacucho: Estrella, 
Killip & Smith 22672 (det. Gl.). Junin: Pichis Trail, Killip & 
Smith 26134 (det. Gl.). Brazil; Central America. 

Leandra reversa [Ser.] Cogn. in Mart. Fl. Bras. 14, pt. 4: 198. 
1886; 691. Melastoma reversa Ser. ex DC. Prodr. 3: 161. 1828, in 
syn. Clidemia reversa DC. I.e. 



FLORA OF PERU 339 

Closely allied to L. longicoma and to L. retropila; leaves 7-9- 
nerved, rather densely pilose beneath, rather unequal in size, 1-2.5 
dm. long, 5-15 cm. broad; calyx teeth 2-3 times shorter than the tube, 
this to 3 mm. long; petals linear-subulate, 3-4 mm. long; ovary 
apically setulose. Cf . a specimen under L. dichotoma, perhaps refer- 
able here. F.M. Neg. 25915. 

Peru (perhaps). Bolivia; Brazil. 

Leandra rotundifolia Macbr., spec. nov. 

Fruticosa, 6 dm. alta; ramis teretiusculis junioribus dense cum 
setulis valde divaricatis haud velpaullo reflexis rufo-hirsutis; petiolis 
dense hirsutis 2-3 cm. longis; foliis late ovatis vel fere rotundatis, 
basi rotundato-truncatis plerumque haud acutis, apice subabrupte 
breviterque acuminatis, 5-7 cm. longis, 3.5-5 cm. latis, membra- 
naceis, minute ciliolato-denticulatis supra paullo setulosis vel gla- 
bratis, subtus mediocriter setulosis et obscure sparseque pilosis, 5-7- 
nerviis vel interdum subplinerviis; paniculis circa 5 cm. longis et 
latis, ramis secundifloris; floribus 5-meris; calyce setulis adpressis 
dense hirtello obscure glanduloso praecipue ad apicem, tubo cam- 
panulato, 2.5 mm. longo, segmentis exterioribus 3-4-plo brevioribus; 
petalis subulato-acuminatis 2.5 mm. longis; antheris vix 1.25 mm. 
longis; ovario 3-loculari, apice parce setuloso, stylo 4 mm. longo. 
Suggests L. retropila Cogn. but differs in foliage, pubescence and 
floral character. 

Loreto: Bush at Nacimiento, Rio Mazan, Jose M. Schunke 176, 
type. 

Leandra secunda Pav. ex Cogn. in Mart. Fl. Bras. 14, pt. 4: 
199. 1886; 691. Melastoma secunda Pav. ex D. Don, Mem. Wern. 
Soc. 4: 308. 1823, in syn. Clidemia secunda D. Don, I.e. 

Near L. dichotoma but distinguishable by the appressed and 
glandular pubescence of the calyx (this often obscure) and by the 
less pubescent leaves; inflorescence branches open and spreading in 
fruit, the secund flowers on each few; anthers 1-1.25 mm. long; ovary 
as to type 5-celled, setulose. The Loreto collections determined by 
Gleason, except Williams 46; 153; 3441, all from Rio Itaya, seem 
referable here, at least at present. 

San Martin: Tocachi, Poeppig 2037. Huanuco: Pueblo Nuevo, 
Ruiz & Pavon, type. Loreto: Near Iquitos, King 1262 (det. Gl.). 
Santa Rosa, Killip & Smith 28774. La Victoria, Williams 2554. 
Pebas, Williams 1601 ; 1658; 1670; 1908. Rio Nanay, Williams 1291. 



340 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Palta-Cocha, Williams 1259; 1260. Caballo-Cocha, Williams 2080; 
2207. Pinto-Cocha, Williams 786; 788; 797. "Santa mullaca." 

Leandra secundiflora (DC.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 
195. 1886; 689. Clidemia secundiflora DC. Prodr. 3: 164. 1828. 

Similar to L. divaricata but leaves sparsely hirsute with long, 
closely appressed trichomes above, setulose beneath, the panicles 
many-flowered and the flowers 4-merous; anthers only about 1 mm. 
long. The branches are crisp-hirsute, with the trichomes more or 
less spreading. Cf. L. secunda; elsewhere in this family (as in 
Tococa) the same species may have 3- or 5-celled ovaries. F.M. 
Negs. 6424; 36243. 

Lima: Near Lima, Gay. Loreto: Leticia, Ule 6205 (det. Pilger). 
Brazil; Panama. 

Leandra solenifera [Schrank] Cogn. in Mart. Fl. Bras. 14, pt. 4: 
192. pi. 42. 1886; 687. Melastoma soleniferum Schrank ex DC. 
Prodr. 3: 164. 1828, in syn. 

Branches, petioles, peduncles and calyces densely glandular- 
pilose; leaves slightly emarginate at base, obliquely acuminate, 
unequally dentate; panicle-branches secund; flowers 6-7-merous; 
otherwise rather similar to L. divaricata. The following, as to Field 
Museum specimen, is Heterotrichum polyandrum (Benth.) Gleason. 

Loreto: Pebas, Williams 1905 (det. GL). Without locality: 
Pavdn. Brazil to the Guianas. 

32. CONOSTEGIA D. Don 

The Peruvian species are shrubs with slender, slightly furfura- 
ceous-pubescent branches, membranous 3-nerved or 5-plinerved 
leaves and ample, terminal panicles of rather small, 6-8-merous 
flowers. Calyx limb a closed cone or top-like, circumscissile at base 
in anthesis. Otherwise similar to Miconia but, on account of the calyx 
character, simulating Calyptrella from which it may be distinguished 
in flower by the unappendaged stamens. 

Conostegia attenuata Triana, Trans. Linn. Soc. 28: 98. 
1871; 699. 

Branchlets slender, nearly terete, minutely scurf y-puberulent; 
petioles 5-10 mm. long; leaves narrowly lanceolate, acute at base, 
long-acuminate, 5-10 cm. long, 1-2 cm. wide, obscurely granu- 
late-puberulent, 3-nerved; flowers 5-merous, few in a subcorym- 
bose panicle, the pedicels about 1 cm. long, articulate near the middle; 



FLORA OF PERU 341 

hypanthium ovoid-conic, scurfy, 6-7 mm. long, 2.5-3 mm. thick; 
petals narrowly obovate, acute, 4-5 mm. long; anthers oblong, 2 mm. 
long; style filiform, 4 mm. long, the stigma capitate. Description 
as to foliage applies only to the Peruvian and Ecuadorian shrub; 
the Colombian type was described as glabrous, branches tetragonous, 
leaves attenuate at both ends; the southern form may therefore be 
noticed as var. peruviana Macbr., var. nov., ramulis teretiusculis 
minute pulverulentis; foliis anguste lanceolatis, basi acutis, apice 
longe acuminatis. The Peruvian collection being sterile, the type 
of the variety is Mexia 8485, Prov. Esmeraldas, Ecuador. F.M. 
Neg. 36358. 

Loreto: Along the Rio Nanay, Williams 1080. Ecuador; 
Colombia. 

Conostegia Poeppigii Cogn. in Mart. Fl. Bras. 14, pt. 4: 211. 
1887; 703. 

Leaves obscurely scurfy-puberulent on the nerves beneath, 
oblong-ovate, acuminate, 2-3 dm. long, 7-11 cm. broad; panicles 
about 1 dm. long; flower-bud 6 mm. long, half as thick, glabrous, 
shortly and acutely acuminate; petals subtruncate, 3 mm. long. 
The widely distributed C. icosandra (Sw.) Urban, 706 (C. subhirsuta 
DC.), has densely scurfy branchlets, flower bud subglobose, 8-9 mm. 
long, the flowers 8-10-merous, with petals 7-8 mm. long. F.M. 
Neg. 16998. 

Huanuco: Pampayacu, Poeppig. Brazil. 

33. MICONIA R. & P. 

For the region comprised this group practically includes all those 
species with baccate or coriaceous, irregularly rupturing fruit, 
terminal inflorescence, this often much branched, and leaves that 
are rarely striolate-veined between the longitudinal nerves. Generally 
the petals are obtuse or retuse or rounded, the connective is little 
if at all produced and not appendaged, at least dorsally. For 
reasons of tradition or convenience a number of segregates, for 
which the key provides (cf. also Cogniaux, I.e. 613-614), are main- 
tained. If a specimen does not key out to one of these offshoot 
groups but sometimes the distinctions are not well-sustained it 
is a Miconia, if the fruit is as indicated above. 

Gleason has written that the "range of structure in the stamens, 
the calyx and the inflorescence strongly suggests that the genus, 
as now accepted, is a heterogeneous assemblage greatly in need of 
segregation." This "range of structure" is a fact but it is not clear 



342 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to me how segregation would serve the purpose of showing the 
probable evolution and present relationship of the species; sections 
under one group-name satisfy better, or at least satisfactorily, the 
endeavor to show the historical picture, and certainly the mechanics 
of the taxonomy are much simplified. Logically then, other groups 
now segregated could be reunited (or united) to conform with the 
range of structure admitted in Miconia. It is an error to insist 
that segregation necessarily serves sound taxonomy; actually the 
drawing of generic boundaries too finely, breaks, even if it does not 
in many instances lose, the lines of development for anyone except 
perhaps the specialist, who overlooks the fact that he himself (by 
much familiarity) may retain in his own mind his segregates within 
the larger units in which they really belong. 

Technical key by sections, after Cogniaux and Gleason. 
On page 360 is an artificial key to all the species. The sections 

are not always well marked. The number following the section 

name refers to the page for the key to its species. 

Anthers subulate or extremely narrow and the sides parallel, dehiscent 
by a single minute terminal pore, the connective simple, pro- 
longed, or sometimes even appendaged at base (Jucunda, 
Tamonea, Adenodesma, Octomeris, Laceraria are sections included 
here) 1. Eumiconia (p. 343). 

Anthers somewhat or scarcely enlarged toward the truncate terminal 
pore, this usually as wide as the anther connective, evidently 
extended (rarely the anther is sterile below, the upper portion 
then obovate) 2. Glossocentrum (p. 351). 

Anthers oblong, dehiscent by clefts that typically extend nearly 
or quite to base of cells, the short anther thus soon expanded 
and open on one side; connective stout, conspicuously extended 
but unappendaged 3. Chaenanthera (p. 352). 

Anthers semiellipsoid, blunt, curved on the outer sides, with 1 
minute pore, the connective neither appendaged nor prolonged 
(M. lasiocalyx with 2-pored anthers) .4. Amblyarrhena (p. 353). 

Anthers obovate or cuneate, with 2-4 broad, truncate pores; fila- 
ments usually flat or geniculate 5. Cremanium (p. 355). 

Anthers similar to Cremanium, but the pore 1, this partly 2-cleft 
and produced anteriorly 6. Chaenopleura (p. 358). 

Section EUMICONIA 

Note: This key and those to other sections are surely largely 
artificial but so also are those of Cogniaux. A natural arrangement, 



FLORA OF PERU 343 

as already suggested by Gleason, will probably be based at least in 
part on the nature of the connective and its appendages. However, 
these characters alone are doubtfully satisfactory, as sometimes 
species otherwise similar are thereby separated widely; and though 
the character of appendages may be constant, the degree of pro- 
longation of the connective does not seem to be. Probably in this 
genus only consideration, of a combination of floral and vegetative 
characters will result in a fairly natural alignment. Here, for 
convenience, the nature of the pubescence has been used more than 
the number of leaf nerves (this is often subject to various interpre- 
tations) as by Cogniaux. 

1. EUMICONIA 

Calyces oblong-cylindrical, at least by prolongation of tube, the 
limb often dilated, characteristically longer than broad, not or 
little narrowed at base, often 4 mm. long or longer, never 
secund; anthers often subulate, falcate, elongate, frequently 
4-8 mm. long (cf. M. calvescens, flowers verticillate; M. capitata, 
flowers capitate). 

Leaves at least shortly plinerved, sometimes only 1 nerve origi- 
nating above the base, pubescent or sessile, not lepidote. 

Flowers not subtended by conspicuous bracts; pubescence not 

extremely compact. 
Leaves attenuate at base; branches appressed-strigose. 

Leaves strikingly plinerved M. nervosa. 

Leaves shortly plinerved M. radulaefolia. 

Leaves amplexicaul or auricled at base; branches not ap- 
pressed-strigose. 
Leaves shortly plinerved. 

Branchlets densely hispid M. impetiolaris. 

Branchlets puberulent or tomentulose. 

Leaves elliptic M. grandifolia. 

Leaves obovate M. nobilis. 

Leaves strikingly plinerved. 
Calyces 5-8 mm. long. 

Leaves broad, ample M. amplexans, M. tomentosa. 

Leaves oblongish, elongate M. biglandulosa. 

Calyces about 2.5 mm. long M. pandurata. 



344 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers subtended by conspicuous bracts but these more or 
less promptly caducous; pubescence extremely compact on 
leaves beneath. 

Leaves sessile or subsessile M. megaphylla. 

Leaves distinctly petioled. 

Leaves strikingly plinerved M. pachydonta. 

Leaves shortly plinerved M. mucronata. 

Leaves basally nerved or essentially, or glabrous or petioled or 

lepidote-pubescent. 
Bracts subtending the large flowers at least in bud, conspicuous 

but caducous; calyces 4-6 mm. long, pubescent. 
Leaves minutely denticulate (3 doubtfully distinct species, 

the "differences" apparently variable). 
Filaments hirsutulous; flowers often sessile. 

Leaves medium size M. bubalina. 

Leaves ample M. dodecandra. 

Filaments glabrous or lightly pilose; flowers pedicellate. 

M. serrulata. 

Leaves entire or essentially so. 
Flowers pedicellate; connective minutely prolonged into 

2 spurs M. guianensis. 

Flowers sessile; connective scarcely prolonged. 

M. amazonica. 

Bracts not conspicuous, minute or wanting, the flowers small, 
the calyces rarely if ever more than 4 mm. long or the 
plants glabrous (cf. M. Tessmannii). 
Leaves more or less pubescent but not silvery-lepidote or 

plinerved. 

Flowers in sessile verticils arranged in a spike or spike-like 
panicle or panicled; leaves sessile or subsessile or linear, 
entire. 

Leaves narrowly linear-lanceolate M. riparia. 

Leaves broad. 

Leaves shortly petioled or not amplexicaul. 

Leaves 3-nerved, smooth M. triplinervis. 

Leaves 5-7-nerved, bullate M. rufescens. 

Leaves amplexicaul M. amplexicaulis. 

Flowers, if more or less verticillate, borne at the ends of 
branchlets; leaves well-petioled or serrate. 



FLORA OF PERU 345 

Leaves well-petioled, entire or nearly. 

Calyces ashy-tomentose, as often the leaves beneath. 

Connective simple; panicle narrow M. Traillii. 

Connective appendaged; panicle branched. 

M. pubipetala. 

Calyces glabrate or scurfy as also the leaves beneath. 
Leaves glabrous above. 
Panicles not at all setose. 

Calyx tube pubescent M. Donaeana. 

Calyx tube glabrous M. vittata. 

Panicles or peduncles setose M. mazanana. 

Leaves sparsely setose above M. ciliaris. 

Leaves shortly petioled, coarsely serrate M. mollis. 

Leaves glabrous or plinerved or the pubescence lepidote. 
Leaves plinerved (cf. M. triplinervis). 
Branches terete. 
Leaves not decurrent. 

Calyx scurfy M. flaviflora. 

Calyx glabrous M. juruensis. 

Leaves well decurrent M. pteropoda. 

Branches sulcate-winged M. flaccida. 

Leaves basally nerved. 

Leaves glabrous; flowers 4-merous M. Tessmannii. 

Leaves densely lepidote beneath; flowers 5-merous. 

M. aulocalyx, M. abbreviata. 

Calyces campanulate, urceolate, sometimes narrowed to base, thus 
obconic or turbinate, rarely a little longer than broad, often 
small, usually paniculate, sometimes verticillately spicate, 
sometimes secund; anthers various (cf. M. flaviflora). 
Flowers sessile, in sessile or shortly peduncled verticils arranged in 
a spike or panicle, but not restricted to branchlet tips; pubes- 
cence never simple (cf. M. Duckei). 

Leaves sessile M. dipsacea. 

Leaves well petioled. 

Leaves several times longer than broad. 

Leaves stellate-puberulent beneath M. Martiniana. 

Leaves scurfy on the nerves beneath M. glomerata. 



346 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves 2-3 times longer than broad. 
Panicle branched. 

Inflorescence puberulent. 
Leaves not decurrent at base. 

M. calvescens, M. Urbaniana. 
Leaves more or less decurrent on petioles. 

M. pteropoda, M. prasina. 

Inflorescence tomentose M. eriocalyx. 

Panicle simple M. compacta. 

Flowers if sessile not borne as above, sometimes verticillate but 
then at the ends of the branchlets, sometimes aggregate or 
even congested into a more or less capitate inflorescence or 
the pubescence in some part simple. 

Anthers noticeably subulate, often elongate, often with a cluster 
of glands at the broader base, usually falcate at the nar- 
rowed tip (cf. M. glomerata). 
Anthers more or less conspicuously glandular laterally at 

base, at least some of them (the episepalous). 
Inflorescence about 1.5 cm. wide, raceme-like. 

M. compacta. 
Inflorescence pyramidal. 

Calyx truncate or regularly lobed or not closed in bud 

by a scarious membrane. 
Leaves minutely stellate all over beneath. 

M. pubipetala. 

Leaves scurfy on the nerves beneath or glabrate. 
Leaves entire; panicles glabrate or puberulent. 

Calyx campanulate, pedicellate M. aurea. 

Calyx obconic, sessile or nearly. 

Stems terete M. juruensis. 

Stems winged M. flaccida. 

Leaves undulate-denticulate; panicles scurfy. 

M. glandulifera. 

Calyx closed in bud, the membrane splitting into irregu- 
lar lobes at an thesis M. aureoides. 

Anthers all eglandular at base (but connective itself glandular 

in M. Donaeana et rel.). 

Calyx closed in bud, the membrane splitting into irregular 
lobes; style and filaments glabrous. 



FLORA OF PERU 347 

Leaves plinerved M. decurrens. 

Leaves basally nerved. 

Petals white, 5 M. Duckei, M. capitata. 

Petals green, 4 M. Wagneri. 

Calyx regularly lobed; sometimes shallowly; style and fila- 
ments hirtellous. 

Leaves sparsely setulose above, ciliate M. ciliaris. 

Leaves glabrous above. 
Panicles esetulose. 

Calyx scurfy M. Donaeana. 

Calyx tube glabrous M. vittata. 

Panicles more or less setulose M. mazanana. 

Anthers linear, rarely a little attenuate apically, rarely sterile 
below and obovately enlarged apically (division not clearly 
marked; cf. especially M. compacta and the 2 preceding 
species as well as those with closed calyces, and M. juruen- 
sis, the calyx obconic). 

Flowers variously disposed in a panicle or thyrse but never 

secund. 

Pubescence if present various, except never clearly simple 
in considerable part or entirely. 

Leaves at least shortly plinerved or sessile or subsessile; 
or lightly pubescent species, the pubescence minute 
or scurfy, usually without sharp definition of the 
trichome character (cf. M. grandiflora, M. pandurata 
with large sessile leaves). 

Leaves sessile or decurrent on the short petiole to the 
subsessile base. 

Leaves clasping at base M. membranacea. 

Leaves decurrent to base M. glaucescens. 

Leaves distinctly petioled, even if long-decurrent. 

Leaves long-attenuate at base, usually decurrent on 
the petiole, the free portion of the latter then 
shorter than the leaf-edged part. 

Flowers pedicellate at least in part. 

Primary branches of inflorescence simple or 

shortly branched M. longiracemosa. 

Primary branches of inflorescence well branched. 



348 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branches acutely angled; leaves puberulent. 

M. Poeppigii. 
Branches subterete; leaves glabrous. 

M. pyrifolia. 
Flowers sessile. 

M. pteropoda, M. prasina, M. longiracemosa. 
Leaves rounded to acute at base, shortly if at all 

decurrent on the petiole. 
Pubescence obvious, at least on the calyx. 
Leaves slightly auricled at the rounded base. 

M. Chamissois. 
Leaves not at all auriculate. 

Leaves ample, mostly about 1.5 dm. long or 

longer M. Poeppigii. 

Leaves medium, mostly about 10 cm. long or 

shorter. 
Leaves entire. 

Leaves caudate M. cuspidata. 

Leaves obtusely acuminate . M. loretensis. 
Leaves at least lightly undulate-crenulate. 

M. filamentosa. 
Pubescence lacking or essentially. 

Connective glandular M. pauciglandulosa. 

Connective eglandular M . spennerostachya. 

Leaves basally nerved, or in any case, especially if some- 
what plinerved, definitely puberulent with clearly 
defined trichomes, never sessile (cf. M. calvescens). 
Inflorescence coarsely scurfy or stellate, or capitate, or 
if merely puberulent, the trichomes distinctly 
reddish. 

Trichomes markedly stiped, stellate-branched api- 
cally, especially on upper branchlets. 

Petals glabrous; leaves hispid above. 

M. barbinervis. 
Petals puberulent M. stelligera, M. clavistila. 

Trichomes sessile or subsessile, sometimes barbellate- 
plumose, if stiped so dense that stipe is not 
readily obvious (cf. M. mollis, leaves serrate). 
Pubescence ashy-fulvous. 



FLORA OF PERU 349 

Leaves ample, often membranous. 
Calyx limb lacerate. 

Petals 5, whitish M. Duckei. 

Petals 4, green M. Wagneri. 

Calyx limb lobed or truncate. 

Calyx limb undulate-truncate . M. capitata. 
Calyx limb distinctly 5-lobed. 
Leaves compactly tomentulose beneath. 

M. crassipes. 
Leaves minutely scurfy on nerves be- 

j neath M. zubenetana. 

Leaves small, heavy M. floccosa. 

Pubescence reddish. 

Leaves shortly acuminate M. rubiginosa. 

Leaves caudately acuminate. . .M. phaeophylla. 
Inflorescence glabrous, glabrate or finely puberulent. 
Branches slender; calyx 2 mm. long or shorter. 
Leaves elliptic, about twice longer than broad. 

Sepals regular M. eugenioides. 

Sepals irregular M. tetrasperma. 

Leaves oblong, 3-4 times longer than broad. 

M. cecidophora. 

Branches stout; calyx 4 mm. long M. Malatestae. 

Pubescence at least in considerable part clearly simple. 
Pubescence of branches stellately branched at tip. 

Calyx very pubescent M. barbinervis. 

Calyx glabrous or nearly, but densely hispid about the 
base M. cuneata. 

Pubescence of branches simple. 

Flowers in obviously bracted clusters, the bracts ovate. 
Leaves plane, basally nerved M. bracteata. 

Leaves minutely bullate, shortly plinerved. 

M. radulaefolia. 

Flowers if in clusters ebracteate or obscurely, the 
bracts small. 

Leaves evidently but minutely crenulate; calyx 
glabrous or glandular. 



350 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves oblong-lanceolate M. acreana. 

Leaves elliptic M. erioclada. 

Leaves entire or essentially so. 

Leaves closely bullate above M. Ruizii. 

Leaves plane. 

Leaves basally nerved. 
Leaves longer than 1 dm., glabrous above. 

M. Matthaei. 
Leaves shorter than 1 dm., often pubescent 

above M. heteromera. 

Leaves shortly plinerved. 

Calyx minutely pubescent; leaves broadly 

elliptic M. brachybotrya. 

Calyx hirsute; leaves oblongish. 

M. ibaguensis. 
Flowers secund but the branches sometimes short and the 

character then at least early obscure. 
Principal leaf nerves 5, or if only 3, the outer pair not 
submarginal, but well within a faint submarginal pair. 
Leaves basally nerved. 

Pubescence lanate or very fine and compact or minutely 

stellate-scurfy. 
Leaves to 3 dm. long, petioled; connective somewhat 

produced. 

Petals glabrous; pubescence fulvous. . .M. dispar. 
Petals glandular; pubescence ashy. 

M. argyrophylla. 
Leaves medium, usually subsessile. 

Leaves emarginate at base M. albicans. 

Leaves rounded to acute at base. 
Leaves acutely acuminate, acute at base. 

M. serialis. 

Leaves obtuse or obtusely pointed. 
Leaves softly short-lanate beneath. 

M. fallax. 
Leaves compactly tomentulose beneath. 

M. stenostachys. 
Pubescence lepidote. 
Leaves acute at base M. lepidota. 



FLORA OF PERU 351 

Leaves deeply cordate, subrotund M. tiliaefolia. 

Leaves plinerved M . elaeagnoides. 

Principal leaf nerves 3, the outer pair submarginal, rarely 

also a faint marginal pair. 
Leaves densely and compactly pubescent beneath. 

Pubescence red-punctate lepidote M. punctata. 

Pubescence compactly tomentulose .... M. pterocaulon. 
Leaves glabrate beneath. 

Branchlets compressed, wing-margined . M. scorpioides. 

Branchlets subterete '.M. cannabina. 

j 

2. GLOSSOCENTRUM 

(Cf. also M. dolichorrhyncha, etc.) 

Pubescence densely lepidote. 

Anthers fertile to base; leaves abruptly acuminate ..... M. Klugii. 
Anthers sterile below middle; leaves gradually acuminate, opaque 

above M. semisterilis. 

Pubescence not densely if at all lepidote, sometimes lacking. 
Anthers sterile below, expanded at fertile tip. 

Leaves rounded or acutish at base; petals ovate, 1 mm. long. 

M. stellipilis. 

Leaves acute at base; petals elliptic, 1.5 mm. long . M. Pilgeriana. 
Anthers fertile their entire length. 
Stamens 8-10. 

Petals 4; petioles 2 cm. long or longer. 

Leaves basally nerved M. consimilis. 

Leaves shortly plinerved M. centrandra. 

Petals 5, petioles often shorter. 
Petioles 3-6 (8) mm. long; stamens at least slightly 

dimorphic. 
Leaves basally nerved, opposite. 

Leaves rounded or minutely cordate at base. 

Leaves minutely cordate-ovate M. puberula. 

Leaves rounded at base. 

Sepals scarcely 0.3 mm. long M. minutiflora. 

Sepals nearly 1 mm. long M. myriantha. 

Leaves obviously acute at base. 



352 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx lobes broadly rounded, regular. 

M . cayumbensis. 
Calyx lobes 2-4, irregular, deciduous. 

M. tetrasperma. 

Leaves shortly plinerved M. longifolia. 

Petioles all or mostly longer than 8 mm.; stamens usually 

isomorphic. 

Leaves 2-5 (6) cm. wide, most or all of them narrower 
than 4 cm. and wider above the middle than below. 
Leaves shortly plinerved; stamens dimorphic. 

M. longifolia. 
Leaves basally nerved. 
Leaves verticillate, at least the upper, glabrate. 

M. ternatifolia. 

Leaves all opposite, scurfy beneath . . M . sclerophylla. 
Leaves all or most of them wider than 4 cm. and wider 

below the middle than above. 
Petioles 1.5-5 cm. long; connective not simple. 
Branches glabrate; connective spurred. 

M. centrodesma. 
Branches scurfy; connective bituberculate. 

M . zubenetana. 
Petioles 1-1.5 cm. long; connective spur minute or 

lacking. 
Leaves rounded or abruptly acute at base; flowers 

subsessile M. Martiusiana. 

Leaves narrowed at base; flowers sessile. .M. egensis. 
Stamens about 20. 

Leaves shortly plinerved, green beneath M. icosandra. 

Leaves basally nerved, ashy-pubescent beneath. M. Herrerae. 

3. CHAENANTHERA 

Leaves brown-silvery lepidote beneath M. chrysophylla. 

Leaves granular or stellate-puberulent beneath or glabrate. 

Leaves long-caudate-acuminate, stellate-puberulent beneath. 

M. dolichorrhyncha. 

Leaves cuspidate or shortly caudate, somewhat granular beneath. 

M. Regellii. 



FLORA OF PERU 353 

4. AMBLYARRHENA 

(Cf . also M. Ruizii, M. heteromera in Eumiconia) 

Flowers secund (but in M. comosa congested) on divaricate branches; 
branches or leaves or both often conspicuously ciliate; style 
bent horizontally near tip (Section Hartigea). 
Calyx lobed, the depressed sepals long laciniate-setose . .M. lacera. 
Calyx lobes not obvious or much shorter than the tube, entire. 
Branches glabrous or ciliate with long trichomes. 

Branches at first pubescent M. pileata. 

Branches glabrous / M. ciliata. 

Branches as the leaves beneath, shortly papillose-hirtellous. 

M. comosa. 

Flowers paniculate, not secund (cf. M. comosa, prominently bracted). 
Branches glabrous or minutely scurfy apically, the leaves glabrous 

or glabrate. 

Leaves basally nerved, obovate or cordate-ovate. 
Leaves glabrous; branches sub terete. 

Leaves obovate or elliptic M. Malatestae, M. alypifolia. 

Leaves cordate-ovate M. retusa. 

Leaves tomentose beneath; branches winged M. glutinosa. 

Leaves plinerved, often very shortly, or the leaves lanceolate. 
Branches or inflorescence branches acutely or narrowly wing- 
angled. 

Inflorescence open, ample. 
Inflorescence branches wing-angled, glabrous. 

Calyx terete M. expansa. 

Calyx winged at base M. Baittoniana. 

Inflorescence branches acutely angled, lightly scurfy. 

M. Adrieni. 

Inflorescence congested M. monzoniensis. 

Branches terete or somewhat angled-sulcate but bluntly, 

never sharply (to the touch). 
Leaves completely glabrous. 
Leaves 5 (4) -8 cm. wide. 

Flowers 5-merous; stigma little dilated. 

Leaves acuminate M. Terera. 

Leaves abruptly cuspidate-acuminate. . . .M. Cookii. 



354 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers 4-merous; stigma subpeltate. .M. Barbeyana. 
Leaves 1.5-3 (5) cm. wide. 

Filaments glandular; calyx lobes long. .M. longisepala. 
Filaments glabrous; calyx lobes short. 

Flowers 6-merous; petals 2 mm. long. .M. Grayana. 
Flowers 5-merous. 
Petals 2.2-3 mm. long. 

Leaves gradually acuminate M. pedicellata. 

Leaves abruptly acuminate M. Cookii. 

Petals 1-1.5 mm. long. 
Stigma clavate; pedicels 2-3.5 mm. long. 

M. livida. 
Stigma peltate; pedicels 0.5-2 mm. long. 

M. elongata. 
Leaves minutely and sparsely pubescent on the nerves. 

M. Pulgari. 

Branches strongly pubescent as often also the leaves beneath. 
Leaves asperous above or in any case distinctly bullate. 
Leaves shortly plinerved. 

Leaves acuminate M. hamata. 

Leaves obtusish or rounded at tip M. Radula. 

Leaves basally nerved. 

Leaves acuminate or acute or cuspidate. 
Inflorescence pubescent. 

Leaves lanceolate, glabrous or nearly M. rugifolia. 

Leaves oblong-obovate or ovate-elliptic. 

Leaves small. M. centrophora, M. pseudo-centrophora. 

Leaves 1 dm. long or longer M. obscura. 

Inflorescence glabrous M. Lechleri. 

Leaves somewhat narrowed to apex but this itself blunt. 

Leaves glabrate above M. papillosa. 

Leaves densely appressed-setulose above . . . M. trichrona. 
Leaves plane above, glabrous or pubescent. 

Leaves small, rarely if ever 8 cm. long, nerved basally. 

Leaves fleshy-coriaceous; stigma peltate M. crassifolia. 

Leaves sometimes rigid but not fleshy; stigma clavate or 
subcapitate. 



FLORA OF PERU 355 

Leaves only 2-3 cm. long M. aspergillaris. 

Leaves distinctly larger. 

Flowers all pedicellate M. chrysanthera. 

Flowers sessile or partly pedicellate. 

M. centrophora, M. pseudo-centrophora. 

Leaves medium, regularly about 1 dm. long or longer. 

Pubescence strikingly and loosely long-hispid. 

M. sanguined. 
Pubescence short or dense. 

Leaves basally nerved. 

Nerves 5; filaments glabrous M. obscura. 

Nerves 7; filaments pilose M. inamoena. 

Leaves plinerved, at least shortly. 

Petioles 1-1.5 cm. long. 

Leaves setulose on the nerves beneath. 

M. subglabra. 

Leaves puberulent beneath M. vismioides. 

Petioles elongate. 

Flowers subsessile, densely glomerate. 

M. lasiostyla. 

Flowers well-pedicellate, loosely fascicled. 

M. modica. 

5. CREMANIUM (cf. also Chaenopleura) 

Pubescence at least in part, usually on the leaves, simple, sometimes 
minute, sometimes confined to the nodes of the upper branches. 

Small-leaved species, the leaves characteristically ovate, rarely 
oblong-elliptic, in any case much shorter than 1 dm., or the 
branches setose at the nodes. 

Flowers 7-8-merous M . crocea. 

Flowers 4-5 (6)-merous. 

Branches glabrous, sometimes setose at the nodes. 

Nodes of the branches shaggy-setose M. trichogona. 

Nodes of the branches smooth. 

Pubescence on the leaves beneath very sparse or obscure 
in the axils. 



356 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves about three times longer than broad, basally 

nerved M. media. 

Leaves about two times longer than broad, plinerved. 

M. brews. 
Pubescence conspicuous on the leaf nerves and on the 

petioles M. Ottikeri, M. latifolia. 

Branches more or less pubescent M . laurina. 

Medium-leaved species, or in any case the leaves not ovate, 
especially if shorter than 1 dm.; branches never nodosely 
setose. 
Leaves oblong-elliptic, 1.5-3 (3.5) cm. wide, 3-nerved, a 

marginal pair obscure if at all obvious. 
Pubescence dense. 

Branchlets closely scurfy-stellate M. lugubris. 

Branchlets shortly plumose-hirtellous M. atrofusca. 

Pubescence sparse, shortly setulose M. setinervia. 

Leaves ovate-elliptic or broadly elliptic, generally 4-9 cm. wide, 
often 3-nerved but an additional marginal pair obvious. 

Leaves petioled. 
Leaves hirsute beneath; stigma peltate. 

Leaves 3-nerved with a fainter pair M. caerulea. 

Leaves 5-nerved with 1-2 fainter pairs . . . M. cyanocarpa. 

Leaves papillose-hirtellous to glabrate beneath; stigma 

truncate. 

Leaves sparsely and obscurely scabrous above . M. aprica. 
Leaves densely, if shortly, setulose above. 

Leaves shortly plinerved M. br achy anther a. 

Leaves basally nerved M. peruviana. 

Leaves sessile M. opacifolia. 

Pubescence lacking or in no part obviously simple, often stellate or 

scurfy (incl. the merely scabrous M. aprica). 
Pubescence lacking unless on the calyx or rarely at the nodes. 
Leaves sessile, cordate. 

Leaves glabrous M. lilacina. 

Leaves scabrous above (to the touch) M. opacifolia. 

Leaves petioled. 

Leaves small, 5-12 mm. long. 



FLORA OF PERU 357 

Leaves obovate; flowers 5-merous M. buxifolia. 

Leaves ovate; flowers 4-merous M. vaccinioides. 

Leaves much larger. 

Leaves fleshy, closely and transversely striate- veined. 

M. calophylla. 

Leaves merely coriaceous, not striately veined. 
Anthers 2-pored. 

Petioles short, few if any as long as 2 cm.; small or 

medium-leaved species. 
Branchlets terete or bluntly quadrate. 
Stigma evidently peltate. 

Leaves ovate-oblong, 5-8 cm. long. 

Leaves thin ; petals minute M. rubens. 

Leaves firm; petals 1.5-2 mm. long.M. media. 
Leaves oblong-lanceolate, longer . M. glaberrima. 
Stigma obtuse or minutely capitate. 
Style obsolete; leaves oblong-lanceolate. 

M. brevistylis. 
Style obvious; leaves ovate-elliptic. 

M. galactantha. 
Branchlets conspicuously and acutely angled or 

winged M. densifolia. 

Petioles at least in part longer than 2 cm., or in any 
case the leaves considerably longer than 1 dm. 

Branches acutely sulcate M. sulcata. 

Branches terete or obtusely quadrate. 
Leaves attenuate at base or at least acute. 

Stigma obtuse; leaves thin M. clathrantha. 

Stigma peltate; leaves rather firm. 

Leaves 4-5 cm. wide M. cremophylla. 

Leaves 6-11 cm. wide M. coelestis. 

Leaves cordate at base M . Weberbaueri. 

Anthers 4-pored M. theaezans. 

Pubescence obvious, at least on the younger parts. 

Leaves oblong-linear or narrowly lanceolate, about five times 
longer than broad. 

Flowers 4-merous; leaves 2-3.5 cm. long M. salicifolia. 

Flowers 5-merous; leaves 3.5-11 cm. long M. neriifolia. 



358 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves much broader in proportion to length. 
Leaves small to medium, not much longer than 1 dm., often 
shorter. 

Leaves verticillate, sessile or subsessile M . quadrifolia. 

Leaves opposite, petioled. 

Stigma peltate or capitellate; flowers often distinctly 

pedicellate. 

Leaves obovate or oblong-elliptic, acute or obtuse. 
Leaves about 3 cm. wide or wider. . .M. polyneuron. 
Leaves about 2 cm. wide or narrower. 

M. crassistigma, M. dumetosa. 

Leaves rounded-ovate, obtuse M. Miles-Morgani. 

Style enlarged apically, the stigma obtuse; flowers sessile 

or subsessile. 
Leaves not at all scabrous above. 

Leaves glabrate M. galactantha. 

Leaves compactly tomentulose beneath . . . . M. Tiri. 

Leaves scabrous above M. aprica. 

Leaves ample, mostly or all considerably longer than 1 dm. 

Leaves rotund-cordate M. viti flora. 

Leaves oblong-elliptic. 

Calyx about 2 mm. long or shorter. 

Flowers glomerate M. pulverulenta. 

Flowers not distinctly glomerate. 

Pubescence reddish-plumose M. hygrophila. 

Pubescence cinereous, stellate or nearly lacking. 
Leaves 6-11 cm. wide. 

Leaves 3-nerved M. coelestis. 

Leaves plinerved M. saxatilis. 

Leaves 3-5 cm. wide M. cremophylla. 

Calyx 2.5-3 mm. long. 

Leaves brown granulose-stellate beneath . M. granulosa. 
Leaves reddish scurfy beneath M. Pavoniana. 

6. CHAENOPLEURA 

Pubescent shrubs with subrotund leaves, or at least many of the 
leaves less than twice as long as broad. 



FLORA OF PERU 359 

Pubescence in part simple or nearly lacking. 
Low, prostrate-ascending shrubs, the branches short. 

Leaves suborbicular M. rotundifolia. 

Leaves much longer than broad M. chionophila. 

Erect or suberect, the panicles elongate. 
Leaves strongly bullate above, pilose beneath. . . .M. bullata. 
Leaves merely with impressed veins above, glabrate beneath. 

M. trichocaula. 
Pubescence scurfy-stellate, j 

Leaves to 2 cm. long M. grisea. 

Leaves about 5 cm. long M. Miles-Morgani. 

Glabrous or usually lightly pubescent shrubs, or in any case the 

leaves distinctly longer than broad, if pubescent. 
Panicles narrow, thyrsiform, the pedicels, at least in part, more or 
less nodding. 

Leaves shortly plinerved M. integrifolia. 

Leaves basally nerved. 

Leaves less than twice longer than broad M. nitida. 

Leaves at least twice as long as broad, at least most of them. 

Branches densely hirsute-setose M. Griffisii. 

Branches glabrous or scattered-setulose. 
Branches quadrate, often acutely; flowers many. 

M . alpina, M. andina. 
Branches terete or nearly; flowers few. 

Leaves entire; branches scurfy M. thyrsoidea. 

Leaves setose-serrulate or crenate; branches glabrous. 

Leaves sharply acuminate M. fruticulosa. 

Leaves obtuse M. cauingia. 

Panicles pyramidal, the pedicels rigid, erect or spreading. 
Leaves petioled. 
Branchlets sharply sulcate, at least the upper. 

Leaves glabrous M. corymbiformis. 

Leaves densely pubescent beneath M. ftrma. 

Branchlets terete. 

Leaves ovate-cordate M. secundifolia. 

Leaves oblong-lanceolate M. nigricans. 

Leaves sessile . . . M. retusa. 



360 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Artificial key to Miconia 

Leaves definitely bicolor, that is, the upper surface glabrous (or 

essentially), the lower densely pubescent, or if not, or in any 

case, the flowers secund or in sessile verticils at intervals on one 

elongate axis. Contrast, 364. 

Pubescence on the leaves beneath obviously lepidote or lepidote- 

stellate. 
Flowers ternate, glomerulate or spicate, not secund. 

Inflorescence or its few branches simple or little branched. 

M. aulocalyx, M. abbreviates. 

Inflorescence branches much-branched M . Klugii. 

Flowers secund, the inflorescence branches sometimes short. 
Leaves acute at base or in any case not cordate. 

Leaves shortly plinerved, silvery beneath ; branchlets acutely 

angled M. elaeagnoides. 

Leaves basally nerved, the trichomes more or less colored. 
Leaves oblong-lanceolate, at least about three times longer 

than broad. 
Uppermost leaves verticillate; anthers opening by 

slits M. chrysophylla. 

Uppermost leaves opposite; anthers opening by a pore. 

M. punctata. 
Leaves usually elliptic, typically scarcely or rarely three 

times longer than broad. 

Nerves of leaves basal; anthers linear, fertile to base. 

M . lepidota. 
Nerves of leaves partly arising just above base; anthers 

obovate above, sterile below M. semisterilis. 

Leaves cordate at base M. tiliaefolia. 

Pubescence on the leaves beneath not definitely lepidote, sometimes 
lepidote-stellate but then the stellate character the more ob- 
vious, sometimes an extremely compact indument, the char- 
acter of the trichomes not marked, sometimes merely punctate, 
sometimes lacking. 
Calyces oblong, limb often dilated, usually about 5 mm. long, 

bracted but the bracts caducous, not borne secundly. 
Leaves strikingly plinerved. 

Leaves sessile or subsessile, the short petiole if developed 
nearly concealed by the leaf base. 



FLORA OF PERU 361 

Pubescence on the leaves beneath obviously stellate. 
Leaves about half as long as broad; style pubescent. 

M. amplexans, M. tomentosa. 

Leaves about a third as broad as long; style glabrous. 

M. biglandulosa. 

Pubescence on the leaves beneath compactly tomentulose. 

M. megaphylla. 

Leaves obviously petioled M. pachydonta. 

Leaves basally nerved or merely shortly plinerved. 
Leaves usually less than twice longer than broad, entire 

or merely calloused denticulate. 
Leaves broadly ovate or subrotund, distinctly cordate 

at base M. amazonica. 

Leaves acute, rounded or merely emarginate at base. 
Flowers pedicellate, not obviously bracted. 

M. pubipetala. 
Flowers sessile, early bracted. 

Leaves firm; petals 7-9 mm. long. . . .M. mucronata. 
Leaves membranous; petals 5 mm. long.M. Traillii. 
Leaves mostly at least twice longer than broad, rather 
oblong, often not entire. 

Calyx glabrate or glabrous M. guianensis. 

Calyx permanently tomentose. 

Style glabrous below M. dodecandra. 

Style pilose below. 

Principal leaf nerves 3 M. bubalina. 

Principal leaf nerves 5 M. serrulata. 

Calyces campanulate, usually much shorter than 5 mm. or 

borne secundly. 
Flowers secund but the branches sometimes short. 

Leaves not strikingly ciliate nor flowers ciliate-bracted. 
Pubescence on the leaves beneath minutely stellate or 
sparse and obscure. 

Petioles elongate M. dispar. 

Petioles short or nearly lacking. 

Leaves 5-9 cm. wide; branchlets compressed. 

M. scorpioides. 

Leaves rarely wider than 4 cm. ; branchlets grooved. 

M. cannabina. 



362 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence on the leaves beneath lanate or extremely 

compact, the trichomes not defined. 
Leaves only 3-nerved, the outer pair submarginal; 

branches 4-winged M. pterocaulon. 

Leaves with only 3 principal nerves, but also with a 

faint marginal pair. 
Leaves minutely but distinctly emarginate-cordate 

at base M. albicans. 

Leaves acute to rounded at base. 
Leaves barely acute apically; anther connective 

obscurely if at all produced. 
Leaves more or less distinctly petioled. 

M. stenostachya. 
Leaves all or many of them subsessile. 

M. fallax. 
Leaves acutely acuminate; anther connective 

basally produced. 
Branchlets acutely 4-sided; pubescence ashy or 

sordid M. argyrophylla. 

Branchlets subterete; pubescence fulvous. 

M. serialis. 
Leaves strikingly ciliate or at least the flowers ciliate- 

bracted. 

Branches glabrous or ciliate with long trichomes. 
Calyx lobed, the depressed sepals long laciniate-setose. 

M. lacera. 

Calyx lobes not obvious or much shorter than the tube, 
entire. 

Branches at first pubescent M. pileata. 

Branches glabrous M. ciliata. 

Branches, as the leaves beneath, shortly papillose-hirtel- 

lous M. comosa. 

Flowers not secund. 
Leaves sessile or nearly. 

Flowers sessile in verticils that are spicately arranged on 
an elongate axis. 

Leaves basally nerved M. triplinervis. 

Leaves plinerved M. nervosa. 

Flowers paniculate. 



FLORA OF PERU 363 

Leaves nearly oblong M . fakata, M. dipsacea. 

Leaves obovate M. nobilis. 

Leaves well-petioled. 

Leaves large, plinerved, nearly oblong . . . M. Martiniana. 
Leaves medium, usually 10-15 cm. long, sometimes small. 
Leaves minutely auriculate-cordate or subcordate. 

Leaves 3-nerved M . crassipes. 

Leaves 5-plinerved M. Martiniana. 

Leaves rounded or acute at base. 

Leaves about a third longer than broad; pubescence 

obviously reddish; anthers slender. 
Leaves obtusely caudate M. phaeophylla. 

Leaves acutely or shortly acuminate. 

M. rubiginosa. 

Leaves distinctly more than a third longer than 
broad; pubescence fulvous-stellate or scurfy, 
rarely rufescent; anthers short, cuneiform except 
M. Herrerae, M. sclerophylla. 

Leaves shortly plinerved, long acuminate. 

M . vismioides. 

Leaves basally nerved, acute, obtusely acuminate 

or rounded at tip. 

Leaves oblong or nearly, 1-3.5 (5) cm. wide. 
Leaves firm or coriaceous but fragile, oblong- 
elliptic; petioles 1 cm. long or longer. 
Leaves acute, firm, ashy-pubescent beneath. 

M. Tiri. 

Leaves acuminate, fragile, brown or rufes- 
cent beneath. 
Pubescence reddish; anthers linear. 

M. sclerophylla. 

Pubescence bright brown; anthers broad. 

M. granulosa. 
Leaves heavy coriaceous; petioles often shorter 

than 1 cm. 

Flowers 5-merous; leaves 3.5-11 cm. long. 
Leaves obtusish; calyx lobed, subsessile. 

M. neriifolia. 



364 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves acuminate or acute; calyx 
obscurely lobed, sessile. .M. floccosa, 
Flowers 4-merous; leaves 2-3.5 cm. long. 

M. salicifolia. 
Leaves elliptic or obovate, mostly 4 cm. wide or 

wider. 

Branches obtusely or acutely angled or com- 
pressed. 
Pubescence stellate. 

Stamens 10; stigma clavate. 

M. Pavoniana. 
Stamens 20; stigma capitate. 

M. Herrerae. 
Pubescence on leaves beneath hirsute. 

M. firma. 

Branches wing-angled M. glutinosa. 

Leaves often not exactly concolor but not bicolor as above, that is, 
the surfaces not in contrast by virtue of the pubescence or the 
lack of it; flowers never secund. 

Pubescence in some part simple, never entirely lacking, present 
at least in leaf-nerve axils and on petioles, often with more 
or less branched or stellate pubescence intermixed, this 
sometimes dominate, usually not; simple pubescence rarely 
minute or only a few trichomes in the inflorescence (cf. 
M. impetiolaris, M. rufescens, the trichomes actually pseudo- 
simple). This division does not include scurfy species rarely 
with 1-several simple trichomes in the inflorescence. Cf. 368. 
Leaves uniformly pubescent at least on one side, the trichomes 
there usually simple, not confined to the nerves and veins 
but the leaves sometimes merely scabrous above, the 
pubescence sometimes sparse, minute or lacking on one 
or even both surfaces. 

Inflorescence simple or nearly, spiciform (rarely lower verticils 
on short branches). 

Leaves strikingly plinerved M . nervosa. 

Leaves shortly plinerved or scarcely M. radulaefolia. 

Inflorescence more or less branched. 
Leaves glabrous above in age. 

Petioles 1-2 cm. long M. Matthaei. 



FLORA OF PERU 365 

Petioles all or most of them shorter than 1 cm. 

Pubescence mostly simple M. ibaguensis. 

Pubescence mostly scurfy-stellate . . . . M. chrysanthera. 
Leaves pubescent above, sometimes merely scabrous. 
Pubescence of the branches spreading, rather loose, fine 
or soft, sometimes glandular. 

Trichomes, at least many of them, stiped-stellate. 

Calyx glabrous or sparsely setulose M. cuneata. 

Calyx densely setulose M. barbinervis. 

Trichomes simple. 

Flowers well-pediceled M. sanguinea. 

Flowers sessile, glomerulate. 

Flowers minute, many, the calyx about 1.5 mm. long. 
Branchlets terete; petioles often elongate. 

Branchlets hirsute M. cyanocarpa. 

Branchlets substrigose M. br achy anther a. 

Branchlets angled or lightly sulcate; petioles 

about 1 cm. long M. caerulea. 

Flowers medium, the calyx about 3 mm. long. 

Petioles to 2 cm. long; branchlets long-pubes- 
cent M. erioclada. 

Petioles to 1 cm. long; branchlets rather shortly 
hirsute M. ibaguensis. 

Pubescence of the branches crisped, strigose or more or 
less appressed, often tomentulose or coarse, some- 
times lacking or nearly, if spreading very short. 

Principal leaf nerves 3, and 1 pair fainter submarginal 
ones. 

Leaves sessile M. opacifolia. 

Leaves petioled. 

Leaves sparsely scurfy-stellate or hirtellous be- 
\ neath, ovate-elliptic. 

Leaves sparsely scabrous above M. aprica. 

Leaves setulose above M. peruviana. 

Leaves densely pubescent beneath, oblong. 
Upper leaf surface distinctly bullate, lower 
foveolate M. lugubris. 



366 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Upper leaf surface setulose but not bullate. 

M. caelata, M. atrofusca. 

Principal leaf nerves 3-7, if only 3 equally strong, 
there are then 2 pairs of fainter submarginal ones. 
Nerves all basal. 

Leaf surface smooth, sparsely setulose or glabrous 

above. 
Leaves sparsely setulose above, serrulate. 

M. inamoena. 

Leaves glabrous above, entire . . M. brachybotrya. 
Leaf surface bullate, sometimes minutely. 
Leaves sparsely setulose above; petioles elon- 
gate M. papillosa. 

Leaves densely setose or strigose above; petioles 

short (rarely 2-3 cm. long). 
Leaves rounded at base, 7-nerved. 

Minutely bullate upper leaf surface con- 
cealed by pubescence M. Ruizii. 

Coarsely bullate upper leaf surface con- 
spicuous M. trichrona. 

Leaves cordate at base, 9-nerved . M. Lechleri. 
Nerves in part arising shortly above leaf base. 
Calyx pubescent. 

Leaves densely pubescent beneath. 
Leaves villous beneath; calyx 2 mm. long. 

M. hamata. 

Leaves setulose beneath; calyx 4 mm. long. 

M. Radula. 
Leaves sparsely and minutely pubescent on 

both sides M. lasiocalyx. 

Calyx tube glabrous M. br achy anther a. 

Leaves more or less pubescent marginally or beneath but at 
least chiefly only on the nerves or transverse veins or in 
the nerve axils; in any case the pubescence not uniformly 
distributed over the leaf surface or on the veinlets, rarely 
mostly marginal (cf. M. brachybotrya, M. brachyanthera) . 
Leaves subrotund, 1-1.5 cm. wide. 

Branches conspicuously setulose and scurfy; leaves glabrous 
above, the veins impressed M. trichocaula. 



FLORA OF PERU 367 

Branches merely or chiefly scurfy; leaves setulose or strongly 

bullate. 
Leaves smooth but sparsely setulose above. 

M. chionophila, M. rotundifolia. 

Leaves strongly bullate but glabrous above. .M. bullata. 
Leaves distinctly longer than broad. 

Nodes at least in panicle setose, the branches (below at 

least) glabrous or puberulent. 
Leaves ample. 

Leaves cordate M. Weberbaueri. 

Leaves rounded or subacute at base .... M. mazanana. 
Leaves about 1 dm. long or smaller. 

Nodes shaggy-setose M . trichogona. 

Nodes sparsely setulose M. andina. 

Nodes glabrous or pubescent like the stems. 
Leaves oblong-lanceolate, at least many more than three 
times longer than broad (M. comosa, with ciliate 
bracts, might be sought here). 

Leaves densely bullate-roughened above. .M. rugifolia. 
Leaves smooth above or the veins merely impressed. 
Panicle not racemiform. 

Calyces densely pubescent M. subglabra. 

Calyces glabrous or obscurely puberulent. 
Leaves rugose beneath by the many transverse 

nerves M. setinervia. 

Leaves smooth or in any case the veins few. 
Branchlets densely villous to glabrate; leaves 

acuminate. 
Branchlets villous; leaf nerves basal. 

M. acreana. 
Branchlets glabrate; leaves shortly pli- 

nerved M. Pulgari. 

Branchlets long setose-hirsute; leaves obtus- 

ish M. setulosa. 

Panicle racemiform M. heteromera. 

Leaves ovate-oblong to elliptic, two to three, mostly 

about two, times longer than broad. 
Flowers 7-8-merous . . . . M. crocea. 



368 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers 4-5 (G)-merous. 
Flowers 5-6-merous. 

Branchlets pilose- or setulose-scurfy or scurfy- 
stellate. 
Pubescence mostly scurfy-stellate. 

M. centrophora, M. chrysanthera. 

Pubescence simple M. laurina. 

Branchlets glabrous or sparsely or long-setulose. 

Setae few, scattered or lacking M. media. 

Setae abundant M. setulosa. 

Flowers 4-merous. 
Branches densely but shortly hispid-setose. 

M. Griffisii. 

Branches glabrous or glabrate, the branchlets 
sometimes obscurely pilose or sparsely setu- 
lose, rarely tomentose. 

Leaves pilose on veins beneath; anthers trun- 
cate M. latifolia. 

Leaves glabrous beneath or pubescent on 

nerves or in nerve axils. 
Leaves shortly plinerved, entire. 

M. integrifolia. 
Leaves basally nerved. 

M. andina, M. Ottikeri, M. alpina. 

Pubescence in no part simple, sometimes entirely wanting, often 

a sparse and scurfy puberulence, often conspicuously stellate 

or branched (generally merely scabrous species have been 

included in the above division with simple trichomes). 

Leaves sessile or subsessile, the short petiole, if obvious, more or 

less concealed by the leaf base. 

Leaves more or less stellate pubescent above, sometimes gla- 
brate in age. 

Leaves subentire, the surface bullate M. rufescens. 

Leaves coarsely toothed, the surface nearly plane. 

M. mollis. 

Leaves glabrous above, at least at maturity, unless the mid- 
nerve, or merely scabrous. 

Leaves glabrous beneath or merely minutely puberulent or 
lepidote-punctate at maturity. 



FLORA OF PERU 369 

Leaves narrowed and not clasping at base, this rarely 

auricled. 
Verticils of flowers arranged in a spike . . . M. triplinervis. 

Verticils of flowers panicled M. glaucescens. 

Leaves even if narrowed below, sessile, clasping at base. 
Leaves basally nerved or obscurely plinerved. 

Leaves narrowed at base M. membranacea. 

Leaves cordate at base. 
Leaves 7-17 cm. long. 

Leaves thin, scabrous above M. opacifolia. 

Leaves coriaceous, glabrous M. lilacina. 

Leaves 3-6 cm. long M. retusa. 

Leaves distinctly plinerved, the upper nerves arising 
1 to several cm. above the leaf base. 

Leaves strongly pandurate M. pandurata. 

Leaves not at all pandurate M. grandifolia* 

Leaves obviously if sometimes finely, sparsely or com- 
pactly pubescent beneath, the pubescence if scurfy 
sometimes confined to the nerves. 

Leaves small, verticillate M. quadrifolia. 

Leaves ample, opposite. 

Branches densely hirsute with barbellate trichomes. 

M. impetiolaris. 

Branches scurfy-puberulent or tomentulose. 
Pubescence of leaves beneath a compact indument. 

M. megaphylla. 
Pubescence of leaves beneath scurfy-stellate. 

Calyces 2-3.5 mm. long, usually companulate- 

turbinate. 
Calyces pubescent; leaves long-oblong. 

M. falcata, M. dipsacea. 

Calyces soon glabrous; leaves obovate or 
elliptic. 

Leaves obovate, tapering to base. .M. nobilis. 

Leaves ovate-elliptic M. amplexicaulis. 

Calyces about 5 mm. long, oblong-cylindric. 
Style and filaments glabrous . . . M. biglandulosa. 



370 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Style and filaments somewhat pubescent. 

M . amplexans. 

Leaves petioled, sometimes shortly but the petiole obvious, the 

leaf base usually merely rounded to acute or decurrent. 

Small-leaved species, the leaves, at least many of them, only 

about 8 cm. long, usually much shorter or less than 1 cm. 

wide. Contrast, 373. 

Panicles and branchlets definitely pubescent or puberulent. 
Leaves mostly 2-3 (4) cm. long. 

Flowers nodding, well-pediceled M. grisea. 

Flowers sessile or nearly, crowded. 
Leaves early densely stellate-scurfy beneath; petals 

2 mm. long M. aspergillaris. 

Leaves lightly scurfy beneath; petals about 0.5 mm. 

long M. dumetosa. 

Leaves mostly 4-8 cm. long. 

Leaves more or less caudately or long-acuminate, if 
the latter, linear-oblong. 

Leaves linear-oblong M. riparia. 

Leaves ovate to elliptic, more or less conspicuously 

caudate. 

Anthers laterally dehiscent; leaves shortly caudate. 

M. Regelii. 
Anthers terminally dehiscent. 

Leaves minutely cordate at base . . . M. puberula. 
Leaves acutish at base. 
Calyx glabrate or glabrous. 
Calyx 2.5-3 mm. long; petioles slender, 
7-11 mm. long M. heteromera. 

Calyx about 1.5 mm. long; petioles stouter, 
shorter M. minutiflora. 

Calyx fulvous puberulent M. cuspidata. 

Leaves obtuse, acute or very shortly acuminate, al- 
ways broader than oblong-linear. 
Leaves oblong-lanceolate or oblong-obovate, 1.5-2 

(2.5) cm. wide. 
Petioles 1 cm. long or longer; flowers minute. 

M. crassistigma. 



FLORA OF PERU 371 

Petioles 4-7 mm. long; petals 2 mm. long. 

M. nigricans. 
Leaves ovate-elliptic, rounded to an acutish or 

cordate base, (2) 2.5-4 cm. wide. 
Leaves not heavy-coriaceous. 
Leaves truncate-cordulate at base. 
Calyx glabrate; leaf nerves basal. 

M. Miles-Morgani. 
Calyx scurfy; leaves shortly plinerved. 

M. secundifolia. 

Leaves acutish or merely rounded at base; 
calyx at least sparsely stellate-pubescent. 

Leaves smooth (or nearly) above, even in age. 

Flowers on pedicels 1-2 mm. long. 

M. chrysanthera. 

Flowers sessile or subsessile. 

Anthers open at tip; stigma capitate. 

M. centrophora. 

Anthers open on the side; stigma clavate. 

M . Regelii. 

Leaves bullate above, foveolate beneath. 

M. pseudocentrophora. 
Leaves exceptionally heavy-coriaceous. 

M. crassifolia. 
Panicles and branchlets glabrous or nearly, the pubescence 

if present evanescent or very sparse. 
Leaves 3.5 cm. long or longer. 

Branches acutely angled or winged. 

Leaves tomentose beneath M. glutinosa. 

Leaves glabrous beneath. 

Leaves narrowly oblong, 2.5 (3) cm. wide or 
narrower. 

Leaves 3-nerved, the one pair submarginal. 

M. densifolia. 

Leaves 3-nerved with also a faint submarginal 
pair M. andina, M. alpina. 

Leaves obovate to elliptic, 3 (2.5) cm. wide or 
wider. 



372 FIELD MUSEUM OF NATURAL HISTORYBOTANY, VOL. XIII 

Panicles open, ample M. expansa. 

Panicles dense M. monzoniensis. 

Branches terete or bluntly quadrate. 
Leaves about three times longer than broad, minutely 
denticulate at least toward tip, rarely entire 
(the oblong-lanceolate glabrous M. livida and 
M. Gray ana might be sought here). 
Leaves usually ciliate-serrulate; pubescence often 

early scurfy; calyx minute. 
Leaves acute or acuminate. 

Leaves firm, scurfy on midnerve. 

Flowers all pedicellate; petals 1.5 mm. 

long M. polyneura. 

Flowers sessile or nearly; petals scarcely 

1 mm. long M. galactantha. 

Leaves thin, glabrous M. rubens. 

Leaves caudate M. minutiflora. 

Leaves calloused-dentate or spinulose-serrulate or 
entire; pubescence lacking or not scurfy; 
calyx 2.5-4 mm. long. 
Leaves sharply or even spinulose-serrulate; 

flowers 4-merous. 
Calyx turbinate, narrowed to base. 

M. fruticulosa. 
Calyx campanulate, rounded at base. 

M. andina. 

Leaves calloused-denticulate, rarely some of 
them entire; flowers 5-merous. . .M. media. 
Leaves about twice longer than broad, remotely or 
obscurely, bluntly or crenately toothed. 

Leaves shortly plinerved; flowers numerous. 

M . brevis. 

Leaves basally nerved ; flowers few . M. thyrsoidea. 
Leaves 3 cm. long or shorter. 
Leaves rounded at apex, entire. 
Leaves obovate or oblanceolate. 

Leaves 5-8 mm. broad M. buxifolia. 

Leaves 10-18 mm. broad M. alypifolia. 



FLORA OF PERU 373 

Leaves oblong-elliptic M. vaccinioides. 

Leaves ovate, narrowed to obtuse apex, obscurely 

crenate M. cauingia. 

Medium to large-leaved species, the leaves, at least most of 
them, 9 cm. long to much longer (M. pachydonta and 
M. Regelii might be sought here). 

Pubescence obvious, at least on some part, this usually the 
panicle, and, if even only minutely scurfy, uniformly 
spread where occurring or conspicuous, not early 
deciduous, often permanent (see also some specimens 
of M. nobilis, M. dipsacea). Contrast, 377. 
Large-flowered species, the petals 4-6 mm. long, some- 
times exserted from the calyx only about 2 mm. 
Flowers ternate or in any case subsessile. 

Leaves conspicuously toothed M . mollis. 

Leaves entire or essentially. 

Leaves acute at base M. aureoides. 

Leaves rounded at base. 
Minutely pubescent species. 

Calyx tube glabrous M. vittata. 

Calyx tube densely tomentulose. 

M. Donaeana. 

Conspicuously pubescent species . . M. rufescens. 
Flowers obviously pedicellate, at least in part. 
Petals glabrous or nearly; anthers eglandular. 

M. ciliaris. 

Petals puberulent; anthers or filaments glandular. 
Leaves more or less repand-crenulate. 

M. glandulifera, M. muricata. 
Leaves entire or remotely calloused. 

Leaves glabrate; anthers subulate. . .M. aurea. 
Leaves stellate beneath; anthers blunt. 

M. modica. 
Small-flowered species, the petals rarely about 3 mm. 

long. 

Leaves cordate to rounded at base, sometimes narrowed 
toward the base but base itself cordate-rounded 
or obtuse, never decurrent, and in any case the 
pubescence marked, at least in the inflorescence. 



374 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves more or less cordate at base, sometimes 

minutely but definitely. 

Pubescence in inflorescence a fine ashy puberu- 
lence; anthers linear-subulate, conspicuous. 

M. calvescens. 
Pubescence and anthers not concomitantly as 

above. 

Inflorescence thickly lanate with ashy trichomes. 

M. eriocalyx. 
Inflorescence variously pubescent with rusty 

trichomes. 
Leaves at the least three times longer than 

broad M. Martiniana, M. glomerata. 

Leaves about twice longer than broad. 

Leaves strikingly plinerved . . M. lasiostyla. 
Leaves basally nerved or only shortly 

plinerved. 

Pubescence in part lanate with barbate 
trichomes; calyces pedicellate. 

M. vitiflora. 
Pubescence not lanate. 
Calyces sessile, glomerate. 

Flower verticils spicately arranged. 
M. pulverulenta. 

Flower verticils terminal on the 
branchlets of the open or con- 
gested inflorescence. 

M. Duckei, M. Wagneri. 

Calyces pediceled M. secundifolia. 

Leaves merely rounded or obtuse (rarely acutish) 
at the base (exceptional specimens of the 
big-leaved M. calvescens might be sought here). 
Panicle narrow, subcapitate or racemiform. 

Panicle racemiform M, compacta. 

Panicle compact M. Duckei, M. capitata. 

Panicle more or less branched. 

Pubescence on branchlets distinctly stiped- 

stellate, or calyx lacerately lobed. 
Calyx regularly lobed; stiped-stellate tri- 
chomes conspicuous. 



FLORA OF PERU 375 

Leaves glabrous above M. clavistyla. 

Leaves sparsely stiped-stellate above. 

M. stelligera. 

Calyx irregular ; stiped-stellate trichomes short. 

M. Duckei. 

Pubescence on branchlets compact, scurfy and 

not, at least obviously, stiped-stellate. 
Flowers almost minute, the calyx only 1-2 

mm. long. 
Leaves more or less conspicuously caudate, 

entire. 

Caudation greatly prolonged ; pubescence 
loose M. dolichorrhyncha. 

Caudation short; pubescence minute. 

M. stellipilis. 

Leaves acute or acuminate or if a little 
caudate, not entire. 

Flowers densely glomerulate. 

M. pulverulenta. 

Flowers ternate or few or not in separated 

glomerules. 
Leaves entire. 

Leaves firm, the trichomes barbel- 
late M. hygrophila. 

Leaves medium, the trichomes stel- 
lulate M. saxatilis. 

Leaves denticulate-crenulate. 

M. zubenetana. 

Flowers medium, the calyx 2.5-3 mm. long. 
Pubescence gray. 

Leaves 3-4 cm. wide M. Tiri. 

Leaves mostly 4-8 cm. wide .M. Herreme. 

Pubescence bright brown. . .M. granulosa. 

Leaves more or less decurrent on the petiole, even if 

rounded to base at least minutely decurrent, in 

any case not markedly pubescent, rarely evanes- 

cently along the principal nerves, sometimes 

densely pubescent in the inflorescence. 



376 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves long-decurrent (even to base) on the thereby 

winged petiole, plinerved. 
Leaves glabrous; calyx irregularly lacerate. 

M. decurrens. 

Leaves obscurely puberulent on the nerves be- 
neath; calyx minutely toothed. 
Vein areoles more than 1 mm. wide, oblongish. 

M. pteropoda. 
Vein areoles 0.3 mm. wide, mostly 5-6-sided. 

M. longiracemosa. 
Leaves decurrent only at tip of petiole. 

Leaves always drying black, often caudate. 
Petals subacute; calyx opening irregularly; 
puberulence extremely fine, dense. 

M. tetrasperma. 

Petals obtuse; calyx regularly toothed; puberu- 
lence uneven or coarser. 
Leaves about twice longer than broad. 

Leaves to 12 cm. long. . . .M. Martiusiana. 
Leaves to 2.5 dm. long. 

M. egensis, M. Chamissois. 

Leaves about three times longer than broad. 

Leaves 2-3 cm. wide; calyx about 3 mm. 

long M. cuspidata. 

Leaves (or some of them) 4-8 cm. wide; 

calyx 1.5-2 mm. long. 
Leaves shortly plinerved; anthers terete. 

M. Poeppigii. 
Leaves basally nerved. 
Anthers obovate above . M. Pilgeriana. 
Anthers slender throughout. 

M. minutiflora. 

Leaves always drying brownish-green or yellowish, 
usually only acuminate or obtuse (cf. M. 
Chamissois'). 
Leaves basally nerved. 

Leaves in part ternate; flowers minute. 

M. ternatifolia. 
Leaves all opposite; flowers medium. 

M. cayumbensis. 



FLORA OF PERU 377 

Leaves shortly plinerved. 

Leaves distinctly crenate-serrulate. 

M. filamentosa. 
Leaves entire or essentially. 

Pubescence a puberulence; calyx open. 

M. flaviflora, M. acuminifera. 
Pubescence scurfy; calyx closed in bud. 

M. aureoides. 

Pubescence lacking or evanescent (calyx only sometimes 
scurfy) or merely a sparse pulverulence even on the 
leaf nerves beneath, the upper branches and panicles 
(some species, variable in pubescence, are repeated 
under the contrast, except M. minutiflora and M. 
ternatifolia, the latter with ternate leaves). 
Calyx 2-3.5 mm. long or longer, campanulate; anthers 
short, blunt; branchlets and peduncles usually stout, 
often sulcate or angled. 
Branches obtusely quadrate or sulcate. 
Leaves mostly 5-8 cm. wide, acute or shortly 
acuminate. 

Petioles stout, short M. Malatestae. 

Petioles slender, elongate. 
Calyx obscurely pulverulent, the lobes acute. 
Inflorescence peduncled, leafy, the leaves 

acuminate M. Adrieni. 

Inflorescence sessile; leaves cuspidately acu- 
minate M. Cookii. 

Calyx glabrous, the lobes obtuse . . . . M. Terera. 
Leaves small, 2-3 (4.5) cm. wide, long-acuminate or 

caudate. 

Filaments glandular; calyx lobes and tube sub- 
equal M. longisepala. 

Filaments glabrous; calyx lobes short. 

Petals 3 mm. long M. pedicellata. 

Petals 1-1.5 mm. long. 

Stigma clavate M. livida. 

Stigma peltate M. Grayana, M. elongata. 

Branches, or at least inflorescence branchlets, acutely 
or wing-angled. 



378 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Panicles ample, open. 

Leaves glandular-ciliate M. expansa. 

Leaves minutely setose-serrulate. .M. Bailloniana. 
Panicles congested. 

Flowers sessile or subsessile M. monzoniensis. 

Flowers well-pediceled M. corymbiformis. 

Calyx minute or sometimes 2-3 mm. long, rarely longer 
and often constricted, at least not campanulate or 
the anthers narrowed and not blunt; branchlets 
usually slender, sometimes compressed or bluntly 
angled. 

Leaves shortly plinerved (cf. also M. theaezans with 4- 

pored anthers). 
Calyx minute or small, campanulate or subglobose, 

1-2.5 (3) mm. long, never densely scurfy. 
Flowers 4-merous; anthers obovate. 

Leaves ciliate M. centrandra. 

Leaves eciliate M. Barbeyana. 

Flowers 5-merous; anthers narrow. 

Stamens about 20 M. icosandra. 

Stamens usually 10. 

Leaves usually ternate, oblong-lanceolate, 
decurrent on petioles 5-10 (15) mm. long. 

M. longifolia. 

Leaves opposite, usually elliptic, the petioles 
mostly elongate or the leaves well- 
decurrent. 

Leaves not or scarcely decurrent, about 

twice longer than broad. 
Petioles usually appendaged; flowers 
sessile or subsessile; anthers tapering. 
Anther connective obsolete, eglandular. 
M. spennerostachya. 
Anther connective prolonged, glan- 
dular M. pauciglandulosa. 

Petioles unappendaged; flowers sessile or 
on pedicels 0.5-3 mm. long; anthers 
truncate or linear. 



FLORA OF PERU 379 

Flowers mostly pedicellate. 

M. centrodesma. 
Flowers sessile or subsessile. 
Anthers narrow but truncate; con- 
nective produced.. M. egensis. 
Anthers linear; connective not obvi- 
ous M . Chamissois. 

Leaves usually decurrent, more than twice 

longer than broad M . prasina, 

M. pteropoda, M. longiracemosa. 

Calyx 3-5 mm. long or in any case scurfy or not 

globose-campanulate, often narrowed at base or 

constricted at limb. 

Petioles margined nearly to base by the decurrent 

leaves. 
Calyx lacerately opening; stigma punctiform. 

M. decurrens. 
Calyx evenly lobed ; stigma peltate. 

M. longiracemosa. 
Petioles emargined, the leaves little if at all 

decurrent. 

Branches and panicle-branches sharply mar- 
gined M. flaccida. 

Branches terete or bluntly tetragonous. 
Calyx densely scurfy. 

M. flaviflora, M. cremophylla. 
Calyx lightly scurfy, soon glabrate or glabrous. 
Flowers distinctly pedicel ed. 

Flowers 5-merous M. aurea. 

Flowers 6-merous M. Grayana. 

Flowers subsessile M . juruensis. 

Leaves not at all plinerved (unless in M. theaezans, 

the anthers 4-pored). 

Upper petioles or some of them longer than 2 cm. 
Branchlets sharply quadrate-sulcate . . .M. sukata. 
Branchlets obtusely tetragonous. 

Leaves 9-11 cm. wide, ciliate. .M. Weberbaueri. 
Leaves 5-10 cm. wide, calloused-ciliate. 

M. clathrantha. 



380 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Upper petioles rarely 2 cm. long or few of them so 

long. 

Transverse leaf veins fine, numerous, approximate. 

M. calophylla. 

Transverse leaf veins several mm. distant. 
Anthers short, cuneiform or obovate. 

Calyces minute, about 1 mm. long, sessile or 

nearly. 
Style obsolete; leaves lanceolate, 3-5 cm. 

wide M. brevistylis. 

Style obvious, soon exserted; leaves oblong- 
elliptic. 

Anthers 4-pored; calyx lobed; style 2-3 
mm. long M . theaezans. 

Anthers 2-pored; calyx obscurely lobed; 
style 1 mm. long. . . .M. galactantha. 

Calyces about 1.5-2 mm. long, pedicellate or 
sessile. 

Calyces sessile, about 2 mm. long; leaves 
ample . . . . M. coelestis, M . cremophylla. 

Calyces pedicellate, about 1.5 mm. long; 

leaves narrow. 

Anthers 1-pored; leaves narrowed at 
base M. elongata, 

Anthers 2-pored ; leaves scarcely acute at 
base M . glaberrima. 

Anthers narrow or linear. 

Leaves oblong, caudate M. cecidophora. 

Leaves broadly elliptic, acuminate. M. egensis. 

Miconia abbreviata Markgr. Notizbl. Bot. Gart. Berlin 9: 
1145. 1927. 

Compressed younger branchlets compactly stellate-scurfy and 
somewhat minutely hirsute; leaves dark green and glabrate above, 
silvery brown beneath with a dense stellate-scurfy puberulence, 
obovate, narrowed at base, abruptly long-caudate, 3-nerved, about 
8 cm. long and half as wide; flowers spicate, the inflorescence only 
3 cm. long; calyx oblong, sessile, white-scurfy, 4 mm. long, 1.5 mm. 
across, with 5 broadly triangular small lobes; petals white, orbicular, 



FLORA OF PERU 381 

scurfy below; anthers acuminate; style 6 mm. long; ovary villous 
apically, 3-celled. Section Eumiconia, vicinity of M. aulocalyx 
Mart. F.M. Neg. 17015. 

Loreto: Mouth of Santiago, 2-meter tree, Tessmann 4548, type. 

Miconia acreana Ule, Notizbl. Bot. Gart. Berlin 6: 360. 1915. 

Younger parts, including inflorescence and leaf nerves beneath, 
rather conspicuously hirsute-villous; petioles 4-10 mm. long; leaves 
oblong-lanceolate, narrowed at base, long and acutely acuminate, 
notably thin, denticulate-ciliate, nearly glabrous above, minutely 
stellulate and long-hispidulous on the 5 nerves beneath, more shortly 
so on the veins, these reticulate, 10-18 cm. long, 3-5 cm. broad; 
panicles shorter than the leaves, few-flowered, the flowers 5-merous; 
calyx campanulate, glabrous, 4.5 mm. long, the 3-cornered acute 
teeth less than 1 mm. long; petals 3 mm. long; filaments 2.5 mm. long, 
the anthers as long, their cells a little narrowed above, the tip blunt; 
connective weakly 2-lobed; style 5 mm. long. A slender-branched 
shrub. Probably a variety of M. erioclada. Section Eumiconia. 
F.M. Neg. 16999. 

Rio Acre: Seringal San Francisco, Ule 9676, type. 

Miconia acuminifera Triana, Trans. Linn. Soc. 28: 105. 1871; 
744. 

Younger parts, as the peduncles and leaves beneath, very finely 
furfuraceous; petioles stout, 2-3 cm. long; leaves oblong-lanceolate, 
shortly decurrent into the petiole, abruptly caudate-acuminate, 
distinctly 5-plinerved, pale green, coriaceous, 2-2.5 dm. long, 7-10 
cm. broad; flowers ternate at the tips of the branchlets of the pyra- 
midal panicles; pedicels 1-2 mm. long; calyx densely scurfy, slightly 
constricted medially, 4 mm. long; filaments glabrous; connective 
simple. Cf. M. flaviflora. Section Tamonea. F.M. Neg. 25921. 

Peru (possibly). Colombia. 

Miconia Adrieni Macbr. Field Mus. Bot. 4: 180. 1929. 

Essentially glabrous except for a slight scaly scurfiness, especially 
at the nodes and on the calyx; petioles 1-2.5 cm. long; leaves elliptic, 
narrowed at both ends, 14-18 cm. long, 5.5-7.7 cm. broad, definitely 
3-plinerved; panicles somewhat leafy, to nearly 1.5 dm. long, the 
branches acutely tetragonous; flowers 5-merous, subsessile, the 
calyx 4 mm. long, acutely lobed, the white acutish petals about as 
long; anthers oblong-linear, 1-pored, scarcely 3 mm. long, little 
attenuate, minutely bituberculate; style about 5 mm. long. 



382 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Clump shrub, hillside thicket. Apparently near M. elongata. Named 
for M. Paul Adrien of the Huaron Mining Company. Section 
Amblyarrhena. 

Huanuco: Pampayacu, 5077, type. 

Miconia albicans (Sw.) Triana, Trans. Linn. Soc. 28: 116. 1871; 
785. Melastoma albicans Sw. Prodr. 70. 1788. 

Branchlets and leaves beneath densely white-tomentose, in strik- 
ing contrast to the glabrous dark-green upper surfaces of the latter, 
these subcoriaceous, ovate, subcordate at base, obtuse or acute, 
3-5-nerved or slightly 3-plinerved, usually 6-14 cm. long; petioles 
5-15 mm. long; calyx 2.5-3 mm. long; panicle branches secund, 
erect; petals 2.5 mm. long, anthers isomorphic, the connective some- 
what produced, broadly lobed basally; style strongly dilated and 
truncate at the tip, 4-5 mm. long. A compact shrub, or tree-like, 
1-3 meters high. A Poeppig specimen in bud from Maynas, referred 
in herbaria to M . holosericea (L.) Triana, seems rather to belong here; 
the Linne" plant has calyces 6-7 mm. long and linear anthers. The 
pubescence of both these species has been used as tinder. Section 
Eumiconia. 

Huanuco: Pampayacu, 5103; Kanehira 22. Chinchao, Ruiz & 
Pavon. Junin: La Merced, 5520. San Ramon, 1,000 meters, Killip 
& Smith 24777. Chanchamayo Valley, Schunke 333. San Martin: 
Moyobamba, Mathews 1296; Klug 2365. Tarapoto, Spruce 4265. 
San Roque, Williams 7437. Cuzco: Paucartambo, Vargas 104. 
Without locality, Gay 2140. Brazil to Mexico and the West Indies. 
"Yurac mullaca." 

Miconia alpina Cogn. Bot. Jahrb. 42: 146. 1908. 

Glabrous; branchlets acutely 4-angled; leaves minutely spinulose- 
serrulate, coriaceous, oblong, rounded at base, acute, 3-nerved, 4-5 
cm. long, 1.5-2.5 cm. broad; petioles 1-2.5 cm. long; panicle many- 
flowered, 5-10 cm. long, the 4-merous flowers on pedicels 2-5 mm. 
long; calyx broadly campanulate, drying black, the dilated limb ob- 
scurely 4-lobed, 4-5 mm. broad; petals broadly obovate, 2.5 mm. 
long; filaments obscurely glandular-puberulent, to 3 mm. long, the 
arcuate anthers shorter, the style 4-6 mm. long. Near M. thyrsoidea; 
cf. M. andina which as interpreted in this work should include the 
following collections. 

Huanuco: Prov. Huamalies, 3,500 meters, Weberbauer 3313 
(cited first as type). Cuzco: Above Yanamanchi, 3,900 meters, 
Weberbauer 4593 (cited second as type). 



FLORA OF PERU 383 

Miconia alypifolia Naud. Ann. Sci. Nat. se*r. 3. 16: 215. 1851; 
878. 

Glabrous, the slender branches acutely 4-angled; leaves obovate, 
attenuate at base, obtuse or rounded, 3-nerved, coriaceous, 1.5-2.5 
cm. long, 10-18 mm. broad; petioles 2-4 mm. long; panicle thyrsoid, 
2-3 cm. long; calyx ovoid, shortly and acutely dentate, 2.5 mm. long; 
petals broadly obovate, emarginate, 3 mm. long. Shrub with pale 
green entire leaves. Section Amblyarrhena. F.M. Neg. 25924. 

Cajamarca: West of Huambos, Weberbauer 4168 (det. Cogn.); 
260. Ecuador. 

Miconia amazonica Triana, Trans. Linn. Soc. 28: 103. 1871; 
735. 

Branchlets and leaves beneath densely subscurfy- tomentose; 
petioles 7-15 cm. long; leaves broadly ovate-cordate, entire, shortly 
acuminate, glabrous above, subcoriaceous, 7-9-nerved, 2-3 dm. long, 
1.5-2.5 dm. broad; panicle ample, with bracts 4-6 mm. long; 
calyx 5-6 mm. long; flowers 5-6-merous, the petals white- tomentose 
without, 5-6 mm. long; filaments hirtellous, not glandular; connec- 
tive simple; style slightly pilose below, 12-14 mm. long; stigma 
capitate. Section Tamonea. F.M. Neg. 17005. 

San Martin: Tocachi, Poeppig 2003. Tarapoto, Williams 6076 
(det. GL). Loreto: Yurimaguas, Poeppig. Puerto Melendez, Tess- 
mann 4780 (det. Markgr.). San Ramon, Yurimaguas, Williams 
4588. Brazil. "Dispera sacha blanca," "dispero bianco," "nispero 
sacha bianco." 

Miconia amplexans (Crueg.) Cogn. in Mart. Fl. Bras. 14, pt. 
4. 256. 1887; 749. Pogonorhynchus amplexans Crueg. Linnaea 20: 
107. 1847. M. umbrifera Naud. Ann. Sci. Nat. se>. 3. 16: 116. 1851. 
M. symplectocaulos Pilger, Verh. Bot. Ver. Brandenb. 47: 175. 1905, 
fide Gl. 

Branchlets densely scurfy- tomentose; leaves sessile, obovate, 
abruptly acuminate and abruptly attenuate below, where typically 
auriculate, prominently 3-plinerved, sparsely or rarely densely stel- 
late-puberulent beneath, entire or sinuate-denticulate, submembra- 
naceous, often 3-4 dm. long and half as broad, the lateral nerves 
originating several to 12 cm. above the base; panicle ample, the 
flowers somewhat congested, minutely bracteolate, 5-merous; calyx 
oblong, truncate or obscurely 5-lobed, 7 mm. long; petals white, 
retuse, glabrous, 6-8 mm. long; connective curved into a half-circle 
(Gleason) pedicellate-glandular at base anteriorly; style 10-12 mm. 



384 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long; fruit blue. Leaves typically amplexicaul with 2 prominent 
ears. Gleason refers here Klug 88, Mishuyacu, the uppermost 
leaves cuneate at base and not amplexicaul, Williams 1577, Loreto, 
with cuneate base, petioled leaves, and Williams 1473, Klug 1419, 
the stellate pubescence denser, longer and softer. These are probably 
referable to M. tomentosa (which cf.). Flowers variously reported as 
lilac to red. Section Adenodesma. F.M. Neg. 36304. 

San Martin: Tocache, Poeppig 1915. San Roque, Williams 7005. 
Huanuco: Pueblo Nuevo and Pillao, Ruiz & Pavdn. Loreto: 
Yurimaguas, Poeppig 2408; Williams 4259. Quebrada de Cancha- 
huaya (Huber 1423). Iquitos, Tessmann 3599 (det. Markgr.); 
Williams 3782; Klug 88; Klug 1419; Klug 288; Killip & Smith 
26197. Pebas, Williams 1633; Williams 1577; Williams 1698. Rio 
Nanay, Williams 677. Caballo-Cocha, Williams 2049. Florida, 
Klug 2278. Pumayacu, Klug 2325. Rio Acre: Krukoff 5489. 
Without locality, Gay 1447; 1524. Bolivia to Trinidad and Central 
America. "Carache caspi," "muringa" (Yahua), "pichirina," 
"chaita ida" (Huitoto), "muringa." 

Miconia amplexicaulis Naud. Ann. Sci. Nat. se>. 3. 16: 136. 
1851; 776. 

Branchlets densely tangled-hirsute, the longer trichomes barbel- 
late; leaves ovate or elliptic, cordate-clasping, narrowly acuminate, 
coriaceous, soon glabrous above, a little hirtellous on the veins 
beneath, with sessile, stellate trichomes, 3-nerved and with 2 faint 
outer incomplete nerves, 1.5-3 dm. long, 7-12 cm. broad; panicle 
branches spicate, mostly simple, scurfy, the trichomes obscurely 
stellate, the stipe fuzzy as the trichomes on the leaf nerves beneath; 
calyx subtruncate, in age glabrate, sessile, costate, 2.5-3 mm. long; 
petals retuse, faintly puberulent without, about 2 mm. long; style 
4-5 mm. long. Var. parvifolia Cogn. has leaves somewhat triangular, 
acute, only 5-8 cm. broad. In the type there are no trichomes that 
are well-marked stellate above the stipe. The Poeppig specimen from 
Pampayacu has extremely caudate-acuminate leaves. Section 
Eumiconia. 

Huanuco : Cuchero and Pampayacu, Poeppig 11 ; 78 ; 1 424; Mathews 
1719; 1728. Chinchao, Ruiz & Pawn. San Martin: San Roque, 
Williams 7005 (det. Gl.). Mount Campana, Spruce 4373, type var. 

Miconia andina Naud. Ann. Sci. Nat. se>. 3. 16: 236. 1851; 935. 
Chiloporus andinus Naud. I.e. se>. 3. 4: 57. pi. 3. 1845. 

Branchlets quadrate, glabrous or with a few setae only at the 
nodes; petioles glabrate or somewhat setose, 7-20 mm. long; leaves 



FLORA OF PERU 385 

remotely denticulate, oblong-ovate, rounded at base, acuminate, 
yellowish, glabrous above, rarely a little pilose beneath, 3-nerved, 
3-4.5 cm. long, or sometimes twice as long, 10-17 mm. wide or wider; 
panicle dense, narrow, glabrous, 5-12 cm. long; pedicels 1-5 mm. long; 
calyx 3-4 mm. long, the minute teeth ovate, glabrous; petals 4 
(typically), 2.5-3 mm. long; style 3 mm. long, broadly capitate-pel- 
tate. In the Gay specimen the leaves are 8 cm. long, 3.5 cm. wide, 
glabrous; apparently the leaves may be entire except for the stout 
remote cilia; there are really only 3 principal nerves with 2 outer 
very faint ones. The species appears to be variable in pubescence 
development; cf. Naudin, I.e.; probably M. Ottikeri ought to be in- 
cluded, as well as M. alpina, M. latifolia, M. laurina. Section 
Chaenopleura. F.M. Neg. 25926. 

Piura: Huancabamba, Weberbauer 6083. Huanuco: Yanano, 
4930. Cuzco: Marcapata, 3,300 meters, Weberbauer 7792. Near 
Cuzco, Gay. Cerro de Cusilluyoc, 3,800 meters, Pennell 13859 
Puno: Tabina, Lechler 2062; 2067. Bolivia to Colombia and Vene- 
zuela. "Mote mote." 

Miconia aprica Gleason, Bull. Torrey Club 58: 244. 1931. 

Similar to M. peruviana; branches above and petioles (10-15 
mm. long) pilose with long branched trichomes; leaves yellowish- 
green, obovate to ovate-oblong, acuminate, spinulose-denticulate, 
scabrous above, 7-10 cm. long, half as wide; panicle freely branched, 
5-12 cm. long, densely pilose-tomentose-stellate; calyx glabrous, red- 
punctate; sepals acute, 0.5 mm. long; petals obovate, 1.1 mm. long, 
not retuse; anthers 2.5 mm. long, the connective slightly prolonged, 
somewhat elevated; style stout, glabrous, about 1.5 mm. long, the 
stigma truncate. A tree or shrub sometimes 8 meters high, the 
leaves mealy, stellate on the nerves and yellowish-green beneath, 
sparsely scabro-setulose above. Section Cremanium. 

Junin: In thickets and open woods, 2,000 meters, Huacapistana 
(Killip & Smith 242^6), type; 243 Ib. Carpapata, 2,700 meters, 
Killip & Smith 24381. Ayacucho: Ccarapa (Killip & Smith 22311). 

Miconia argyrophylla [Schrank & Mart.] DC. Prodr. 3: 181. 
1828; 789. Melastoma argyrophyllum Schrank & Mart, ex DC. I.e., 
in syn. 

Very similar to M. albicans but the younger branchlets evidently 
quadrate, the angles sharp and the petals glandular-ciliate; petioles 
sometimes longer; leaves membranous, to 2.5 dm. long, 10 cm. wide, 
long-acuminate; flowers 5-merous; calyx about 3 mm. long, the lobes 



386 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

0.5 mm. long; petals 2.5 mm. long, the glandular trichomes conspic- 
uous; style 6.5 mm. long; connective obviously produced, nearly 
bilobed in front. The Peruvian collection by Ruiz & Pavon has not 
been verified. Section Eumiconia. Illustrated, Mart. Nov. Gen. & 
Sp. 3: pi. 284. 

Peru: Without locality (Ruiz & Pavon). Widely distributed, 
South America. 

Miconia aspergillaris (Bonpl.) Naud. Ann. Sci. Nat. ser. 3. 16: 
214. 1851; 883. Melastoma aspergillaris Bonpl. Melast. 44. pi. 
19. 1816. 

Branchlets and leaves beneath densely stellate- tomentose; petioles 
2-6 mm. long; leaves minutely denticulate, ovate-oblong, obtuse, 
often rounded at base, glabrous above, 3-nerved, 2-3 cm. long, 10-13 
mm. wide; flowers crowded, sessile or pedicellate, the inflorescence 
2-3 cm. long; calyx 2 mm. long; petals white, suborbicular, 2 mm. 
long; style 4 mm. long, the stigma slightly enlarged. Section 
Amblyarrhena. F.M. Neg. 25931. 

Cajamarca: West of Huambos, 3,000 meters, Weberbauer 1+162 
(det. Cogn.); 260. Piura: Prov. Huancabamba, Weberbauer 6052. 
Colombia; Ecuador. 

Miconia atrofusca Cogn. Bot. Jahrb. 42: 142. 1908. 

Branchlets and leaves beneath shortly and densely papillose- 
pilose; petioles 4-7 mm. long; leaves rigid, oblong, acutish, narrowed 
to the obtuse base, very shortly and densely setulose above, not 
foveolate beneath, 3-nerved, 5-8 cm. long, 2-3 cm. broad; panicle 
5-8 cm. long, the minute flowers crowded, sessile; calyx minutely 
5-dentate, 2 mm. long; petals subrotund, about 1 mm. long; style 
3 mm. long, the stigma capitate. A specimen by Ruiz & Pavon 
without locality, Markgraf as M. caelata, rather belongs here. Sec- 
tion Cremanium. F.M. Neg. 17017. 

Huanuco: Southwest of Monzon, Weberbauer 3726, type. Playa- 
pampa, 4875. Villcabamba, 5188. 

Miconia aulocalyx Mart, ex Triana, Trans. Linn. Soc. 28: 115. 
1871; 771. 

Branchlets under a lens densely covered with a stellate lepidote 
tomentum; petioles 1.5-4 cm. long; leaves thin, elliptic-ovate, 
abruptly acuminate, acutish at base, green above, silvery beneath 
with appressed stellate-pilose pubescence, 3-nerved or obscurely 
3-plinerved, 1-2 dm. long, 5-12 cm. broad; spikes paniculate; calyx 



FLORA OF PERU 387 

stellate-scurfy, 10-sulcate, truncate or obscurely 5-lobed, 2.5-3 mm. 
long; petals 1.5 mm. long; style 4-5 mm. long. Section Eumiconia. 
F.M. Negs. 25933; 17013. 

Loreto: Yurimaguas, Poeppig 2308, type; Williams 487; 5282 
(det. GL). Rio Nanay, Williams 485; 487. Iquitos, Killip & Smith 
27406. Colombia; Brazil. 

Miconia aurea (D. Don) Naud. Ann. Sci. Nat. sr. 3. 16: 244. 
1851; 745. Chitonia aurea D. Don, Mem. Wern. Soc. 4: 319. 1823. 

Branchlets glabrous or slightly scurfy; petioles 1-4 cm. long; 
leaves oblong-elliptic, long-acuminate, glabrous or glabrate except 
sometimes minutely scurfy beneath on the nerves, 1-2 dm. long, 
3-7 cm. broad, 3-plinerved; panicle thyrsoid, 1 dm. long or longer, 
the pedicels 2-3 mm. long; calyx reddish, slightly stellate-scurfy, 
to 3 mm. long; petals 4-5 mm. long, scurfy without; filaments gla- 
brous or the connective glandular; style 7-8 mm. long. The var. 
latifolia Markgr. has leaves 2 dm. long, half as wide, conspicuously 
sinuate-margined. Section Tamonea. F.M. Neg. 17014. 

Junin: Chanchamayo Valley, Schunke 334; 276. San Martin: 
Tarapoto, Spruce 4%44- Vchiza, Ruiz & Pavon. Huanuco(?): Rio 
Huallaga (Mathews 1302). Loreto : Yurimaguas, Poeppig 2213; 2319; 
Williams 7858 (det. Gl.). Mouth of Apaga, Tessmann 4829 (var. lati- 
folia). Maucallacta, Rio Paranapura, Klug 3930 (det. Standl.). Brazil. 

Miconia aureoides Cogn. in Mart. Fl. Bras. 14, pt. 4: 260. 

pi. 52. 1887; 762. 

Branchlets densely scurfy; petioles 1-2 cm. long; leaves subcoria- 
ceous, obovate-elliptic, abruptly acuminate, slightly stellate-scurfy, 
becoming glabrous, 3-5-plinerved, the larger 10-15 cm. long, 4-7 cm. 
broad, the smaller scarcely half as long; panicle somewhat nodding, 
5-10 cm. long, the flowers 5-merous, sessile or subsessile, minutely 
bracteolate; calyx to 4 mm. long, scurfy-stellate, campanulate, the 
thin limb closed and obtuse, finally lacerating into irregular persistent 
lobes; petals 5-6 mm. long; anthers elongate, minutely 1-pored, 
the connective eglandular; style 6-7 mm. long. Section Laceraria. 
F.M. Neg. 21183. 

San Martin: Tarapoto, Spruce 4519, type; (Killip & Smith 
29528, "tentatively," Gleason). Loreto: Cerro de Ponasa, 1,100 
meters, Ule 6748 (distr. as M. coronata (Bonpl.) DC., Colombian). 

Miconia Barbeyana Cogn. Melast. 872. 1891. 

Glabrous or nearly so; leaves thin, lustrous, oblong-elliptic, acute 
at base, shortly acuminate, 3-plinerved, with 2 very faint outer 



388 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nerves, 1-2 dm. long, 4-7 cm. broad, entire, the petioles to 3 cm. 
long; panicle many-flowered; pedicels about 1 mm. long; calyx 1.5 
mm. long, broadly urceolate, obscurely lobed; petals ovate, 1 mm. 
long; anthers obovate, 1 mm. long; style 4 mm. long, the stigma 
subpeltate. Perhaps the 4-merous variant of M. Terera. Section 
Amblyarrhena. F.M. Neg. 34153. 

Huanuco: Chicoplaya, Ruiz & Pawn, type. 

Miconia Bailloniana Macbr., nom. nov. Pterocladon Sprucei 
Hook. f. ex Cogn. in Mart. Fl. Bras. 14, pt. 4: 209. pi. 45. 1886; 
697, not M. Sprucei Triana. 

Glabrous shrub with acutely angled sulcate branches (wing- 
angled in the inflorescence), thin, slender petioles, 5-plinerved leaves 
and small flesh-colored flowers borne in short terminal panicles; 
leaves ovate-lanceolate, shortly acuminate, usually unequal, the 
larger to 2.5 dm. long, 10 cm. broad, the smaller half the size; calyx 
slightly 4-winged below, the wings alternate with the ovate obtuse 
lobes, dark purple, the tube about 4 mm. long, the lobes scarcely 
half as long; petals fleshy, 4-5 mm. long. The ebracteolate pedicels 
are 4-winged. According to Weberbauer, ants inhabit the stems. 

Although Baillon, Hist. PI. 7: 17, 54. 1880, reduced Pterocladon 
to Miconia (without, however, transferring the single species) it 
has been maintained as a genus chiefly on the basis of the winged 
inflorescence including the calyx base, the ovary rough. These char- 
acters in some degree occur also in Tococa, and in Miconia in different 
sections, as in M. flaccida and in M. expansa; the anthers are nearly 
those of Section Amblyarrhena that contains species with anther con- 
nective enlarged, more than in Tococa with different ovary and 
aspect. 

San Martin: San Roque, Williams 7050 (det. Gl.). Campana, 
Spruce 4312, type. Valley of the Monzon, Weberbauer 4764 (det. 
Cogn.); 291. Junin: Pichis Trail, Killip & Smith 26102, "a terato- 
logical form with abortive flowers" (Gleason). 

Miconia barbinervis (Benth.) Triana, Trans. Linn. Soc. 28: 113. 
1871; 778. Clidemia barbinervis Benth. Bot. Voy. Sulphur 95. 1844. 

Branchlets, leaves beneath on the nerves and inflorescence, even 
to the calyces, densely long-hirsute with stiped-stellate trichomes; 
petioles 3-7 (-10) mm. long; leaves elliptic, acute at base, acuminate, 
obscurely repand-denticulate, sparsely pilose-setulose above, the 
trichomes simple, prominently 3-nerved with a marginal pair of 
faint nerves, usually 1-2 dm. long and about half as broad; panicles 



FLORA OF PERU 389 

few-flowered, short-pyramidal, the branches spreading, the flowers 
verticillate at or toward their tips; calyx 3-4 mm. long; petals retuse, 
3 mm. long; connective scarcely produced, minutely biauricled; 
style strongly enlarged apically, 4-6 mm. long. One of those 
species marked by the long trichomes that are stellately branched 
at their tips. Cf. also M. stelligera, M. clavistila. Secti<mEumiconia. 
Huanuco: Cuchero, Poeppig 1009; 1451; 1627. San Martin: 
Rio Tocachi, Poeppig 1864- Pongo de Cainarachi, tree 4 meters, 
flowers rose, in clearing, Klug 2619 (det. Standl.). Loreto: Near 
Iquitos, Klug 927. Pebas, Williams 1795 (det. Gl.). Florida, 
Klug 2218 (det. Standl.). Rio Acre: Krukoff 5347 (det. Gl.). 
Brazil to Nicaragua and Trinidad. "Acano-fue," "pero-ey" (Huitoto). 

Miconia biglandulosa Gleason, Bull. Torrey Club 59: 367. 1932. 

Tree 2-5 meters high, the younger obtusely 4-angled and 4- 
sulcate branches very stellate-scurfy with bright trichomes, the 
firm leaves beneath similarly but less densely pubescent, glabrous 
above, oblong-obovate, acuminate, subcordate at base, subentire, 
5-plinerved, 2.5-3.5 dm. long, 1-1.5 dm. wide; panicle narrow, 
densely brown-scurfy like the stem, freely branched; flowers sessile, 
5-merous; calyx cylindric, tomentulose, nearly 6 mm. long with 
ovate-obtuse lobes 0.8 mm. long; petals oblong-obovate, 5 mm. long, 
retuse, essentially glabrous; stamens nearly isomorphic, the filaments 
glabrous, 4 or 5 mm. long, equaled by the subulate, slightly arcuate 
anthers, the slender connective elevated near the base; the sides 
extending into 2 lobes marginally biglandular, the glands stalked. 
Petals greenish-white, filaments and style white, anthers purple 
(K. & S.). 

Loreto : Iquitos, Killip & Smith 26933, type. 

Miconia brachyanthera Triana, Trans. Linn. Soc. 28: 129. 
1871; 900. 

Branchlets densely substrigose-hirsute, obtusely tetragonous; 
leaves ovate-oblong, obtuse, rounded at base, sparsely short-setulose 
above, densely papillose-hirtellous beneath, at least on the nerves, 
1.5-2 dm. long, 6-9 cm. broad, 5-plinerved, on petioles 2-3 cm. long; 
panicle broadly pyramidal, 7-8 cm. long, the minute flowers glomer- 
ate; calyx sessile, tube glabrous, 2 mm. long, the 5 lobes short, the 
petals scarcely 1 mm. long; style 1 mm. long, the stigma truncate. 
The broadly pyriform or subglobose anthers are only 0.5 mm. long. 
Nearly M. caerulea unless separable by shorter pubescence and 
character of stigma. Section Cremanium. 

Huanuco: Casapi (Mathews 1726). Without locality (Haenke). 



390 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia brachybotrya Triana, Trans. Linn. Soc. 28: 111. 
1871; 818. 

A small tree, the branches, petioles and leaves beneath densely 
long-hirtellous (the trichomes subappressed) but the leaves com- 
pletely glabrous and plane above, elliptic-obovate, shortly and 
obtusely acuminate, acutish at base, entire, nearly basally nerved 
or barely 5-plinerved, membranous, rather unequal, the larger about 
2 dm. long and half as broad; panicles congested, 4-5 cm. long; 
calyces sessile, minutely and sparsely pilose, about 3 mm. long, the 
white petals as long, the style twice as long; anthers broadly subu- 
late, obscurely 2-lobed at base; ovary truncate, glabrous. The 
glabrate obconic calyces are densely hairy about their bases. Section 
Eumiconia. F.M. Neg. 17022. 

Loreto: Mouth of Santiago, Tessmann 4452. Brazil. 

Miconia bracteata (DC.) Triana, Trans. Linn. Soc. 28: 111. 
1871; 780. Clidemia bracteata DC. Prodr. 3: 162. 1828. 

Stems, petioles, leaves beneath and inflorescence except the 
bracts densely long-hirsute, the trichomes subappressed, rather stiff, 
especially those of the branches; leaves oblong-ovate, serrulate, 
cuneately subrotund at base, sparsely setulose above, 1-2.5 dm. long, 
shortly 3-(5) plinerved; petioles 1-5 cm. long; flowers rotundly 
bracteate in small glomerules at tips of the panicle branches; calyx 
2-3 mm. long, the lobes erect, the outer teeth shorter; petals 4-5 
mm. long; stamens alike, slender, the filaments and anthers subequal, 
2.5-3 (4) mm. long; connective prolonged backward; ovary two- 
thirds free, pubescent above. It is not certain that the Peruvian 
collection, given a name in syn. by Triana, is the same as the plant 
from the Guianas. M. papillosa (Desr.) Naud., 892, from Bolivia 
and Ecuador, with very obtuse strongly bullate leaves, might be 
sought here. Section Eumiconia. F.M. Neg. 36255. 

Loreto: Prov. Minas (Poeppig 1865). Guianas. 

Miconia brevis Macbr. Field Mus. Bot. 4: 188. 1929. 

Small, entirely glabrous shrub with terete branches, obscurely 
crenulate-denticulate, subcoriaceous, ovate-elliptic leaves rounded or 
nearly rounded at both ends, and nodding panicles of white flowers; 
petioles 1-2 cm. long; leaf nerves 5, only 3 prominent; pedicels 
scarcely 1 mm. long; calyx obscurely 5-denticulate, 2 mm. long; 
petals irregularly suborbicular; filaments triangular, 1.5 mm. long; 
anthers broadly obovoid, scarcely 1 mm. long; style glabrous, nearly 



FLORA OF PERU 391 

5 mm. long; stigma capitellate. Nearly M. media, but leaves shortly 
plinerved, mostly 4 cm. long, usually at least half as broad; com- 
pletely glabrous as also the petioles. 

Piura: Between Huancabamba and Ayavaca, Weberbauer 6334, 
type. 

Miconia brevistylis Cogn. Bot. Jahrb. 42: 144. 1908. 

Entirely glabrous shrub 3 meters high, with stout, obtusely angled 
branches; petioles 6-12 mm. long; leaves rather coriaceous, lanceolate, 
long-attenuate at base, entire or remotely and finely denticulate, 3- 
nerved, to 2 dm. long, 4-5 cm. broad; panicle broad, the minute 
5-merous sessile flowers in small glomerules; calyx 1 mm. long, the 
teeth broadly triangular; petals white, scarcely 1 mm. long; style 
nearly obsolete, the stigma punctiform. Section Cremanium. F.M. 
Neg. 17024. 

Huanuco: Prov. Huamalies, 1,600 meters, Weberbauer 3553, type. 

Miconia bubalina [Pav.] Naud. Ann. Sci. Nat. se>. 3. 16: 244. 
1851; 733. Melastoma bubalina Pav. ex D. Don, Mem. Wern. Soc. 4: 
319. 1823, in syn. Chitonia bubalina D. Don, I.e. 

Branches, leaves beneath and inflorescence densely scurfy- 
tomentose with minute, often reddish trichomes; petioles 1-4 cm. 
long; leaves closely but rather prominently crenulate-serrulate (or 
apparently or sometimes entire), elliptic-oblong, acuminate, the 
stellate pubescence beneath extremely compact, glabrous above, 5- 
nerved, 1-3 dm. long, 5-10 cm. broad; panicle about 1 dm. long, not 
open, the bracts 5-7 mm. long; calyx 6-7 mm. long, as also the petals, 
these tomentose without; filaments hirtellous, not glandular; style 
pilose below, 1 cm. long. Killip & Smith made 3 collections in 
Loreto (Gleason). With M. serrulata, probably not distinct from M. 
dodecandra. Section Tamonea. F.M. Neg. 17025. 

Loreto: Iquitos, Williams 3666; 3707 (det. Gl.); Killip & Smith 
27216. Lower Rio Nanay, Williams 678. Without locality, Ruiz 

6 Pavon. Brazil to Trinidad. 

Miconia bullata (Turcz.) Triana, Trans. Linn. Soc. 28: 130. 
1871; 935. Schizanthera bullata Turcz. Bull. Soc. Nat. Mosc. 1862, 
pt. 2: 322. 1862. 

Branchlets densely clothed with red scales and trichomes; leaves 
subrotund, small, petioled, glabrous but bullate-tuberculate above, 
pilose on the conspicuous veins beneath, 3-nerved; flowers bracteate 
at base in axillary 2-flowered racemes; petals 4, obovate-rotund, 



392 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

small; calyx lobes short, the tube campanulate; style exserted, the 
stigma capitate. Section Chaenopleura. 
Peru: (Mathews 873, type). 

Miconia buxifolia Naud. Ann. Sci. Nat. se>. 3. 16: 208. 1851; 
917. 

Much-branched shrub with slender, obtusely quadrate branch- 
lets; petioles 2-3 mm. long; leaves coriaceous, entire, obovate, gla- 
brous beneath, slightly punctate above, 3-nerved, 8-12 mm. long, 
5-8 mm. broad; panicle 1-3 cm. long, closely flowered; calyx 2 mm. 
long, pedicellate, the pedicel 1-1.5 mm. long; petals white, 1.5 mm. 
long; style 3-4 mm. long. The leaves typically are rounded at apex; 
in the Weberbauer specimen they are acutish; the calyx, too, is a 
little narrower. Section Cremanium. F.M. Neg. 25936. 

Huanuco: Huambos, Weberbauer 4163 (det. Cogn.); 260. Colom- 
bia; Venezuela. 

Miconia caerulea [Pav.] Naud. Ann. Sci. Nat. se"r. 3. 16: 245. 
1851 (name); 899. Melastoma caerulea Pav. ex D. Don, Mem. 
Wern. Soc. 4: 313, 1823, in syn. Cremanium caeruleum D. Don, I.e. 
Miconia agyna Naud. I.e. 232. 

Branchlets quadrisulcate and, as the leaves beneath, rather 
densely pilose with simple trichomes; petioles about 1 cm. long; 
leaves membranous, elliptic-oblong, subrounded at base, acuminate, 
sparsely setulose-asperous above, long-pilose on the 3-5 nerves 
beneath, 8-15 cm. long, 2.5-5 cm. broad, entire; panicles broad, 
about 1 dm. long, the branches densely hispid; calyx sessile, urceolate, 
tube glabrous, dentate, 1.5 mm. long; petals scarcely 1 mm. long; 
style 2.5 mm. long, the stigma broadly peltate. The leaves in the 
type are very shortly 3-plinerved, but with 2 outer faint marginal 
nerves; young parts reddish-purple pubescent. Section Cremanium. 
F.M. Neg. 17027. 

Huanuco: Chinchao, Ruiz & Pavon, type; (Haenke; Mathews 
1177)J\min: San Ramon, Killip & Smith 24758 (det. Gl.). San 
Martin: San Roque, Williams 7757; 7419 (distr. as M. ibaguensis). 
Ecuador. 

Miconia calophylla (D. Don) Triana, Trans. Linn. Soc. 28: 
129. 1871; 918. Cremanium calophyllum D. Don, Mem. Wern. Soc. 
4: 313. 1823. M.(?) myrtiformis Naud. Ann. Sci. Nat. s<r. 3. 16: 
244. 1851. 

Related to M. buxifolia Naud. but the branchlets robust, nodose, 
4-sided, and the leaves cuneately oblong-obovate, ample, even to 



FLORA OF PERU 393 

1.5 dm. long and 6 cm. wide, entire, coriaceous, with many slender 
transverse veins; petioles 1.5-2 cm. long; panicle to 1.5 dm. long; 
calyx 1.5 mm. long; petals white, less than 1 mm. long. The leaves 
simulate those of some Guttiferae or Blakea. The mid-nerve is pro- 
nounced, with 2 faint submarginal nerves. Section Cremanium. 
F.M. Neg. 17029. 

Peru: Patasania, in 1787, Ruiz & Pavdn, type. 

Miconia calvescens [Schrank & Mart.] DC. Prodr. 3: 185. 
1828; 799. Melastoma calvescens Schrank & Mart, ex DC. I.e., in syn. 

Branchlets, these often sulcate, inflorescences and sometimes the 
5 leaf nerves beneath more or less mealy-pubescent with minute 
stellate trichomes; petioles 2-6 cm. long; leaves somewhat undulate 
or repand-denticulate, ovate or oblong-elliptic, acute or acuminate, 
rounded or subcordate at base, to narrowed (rarely), in age glabrous 
or nearly, sometimes 4 dm. long; flowers mostly congested at the 
ends of the spreading-ascending panicle branchlets; calyx about 3 mm. 
long, nearly entire; petals 2-3 mm. long; connective minutely bilobed 
anteriorly; style 5-7 mm. long, the stigma subpeltate. Tree some- 
times 10 meters high, but often bushy. Very common in the low- 
lands of Peru and down the Amazon to its mouth (Gleason, who 
lists many collections). Section Eumiconia. 

Cajamarca: Prov. Jae"n, 1,500 meters, Weberbauer 6199. 
Huanuco: Pozuzo, 4653 (?). Cuchero, Ruiz & Pawn; Dombey. 
San Martin: Tarapoto, Spruce 4551. Valley of the Monzon, Weber- 
bauer 3576; 287. Moyobamba, MathewslSOl. Pongo de Cainarachi, 
King 2633(1). Junin: La Merced, 5262. Puerto Yessup, Killip & 
Smith 26258. Colonia Perene", Killip & Smith 25028. Ayacucho : 
Estrella, Killip & Smith 22653. Loreto: Prov. Minas, Poeppig 
2914- Iquitos, Tessmann 3551 (det. Markgr.). Pebas, Williams 
1618; 1619; 1742; 1637. Caballo-Cocha, Williams 2457; 2116. 
Pongo de Manseriche, Mexia 6264; 6345. Florida, King 2207. 
La Victoria, Williams 2703; 2876; 2936. Rio Nanay, Williams 297. 
Rio Itaya, Williams 3367; 3314; Killip & Smith 29560. Near 
Iquitos, King 1428; Williams 1386; 1479. Puno: San Gaban, 
Lechler 2394; 2394a. Widely distributed, South America. "Juze- 
goro-ey," "misha" (Yahua), "huisha" (Huitoto). 

Miconia cannabina Markgr. Notizbl. Bot. Gart. Berlin 9: 
1145. 1927. 

Branchlets compactly white-scurfy; leaves verticillate, narrowly 
lanceolate, sharply acuminate, decurrent on the short petioles, 



394 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

densely but minutely scurfy on the veins beneath and the 3 nerves, 
glabrous above, about 1 (-2) dm. long, only 3.5 (-4.5) cm. wide; 
panicles to 2 dm. long, 8 cm. wide, the alternate branches secund; 
calyx sessile, densely scurfy, about 2 mm. high, minutely 5-lobed, 
the limb persistent in fruit; petals 2 mm. long, glabrous; style 
glabrous, 5 mm. long. The type was about 2.5 meters high. Said 
to be common near Iquitos. Section Eumiconia. 

Loreto: Marsh at Iquitos, Tessmann 5056, type; 3519; Klug 
1076; Williams 297; 2116; 3367, and other collections det. Gl. 
Rio Itaya, Williams 65. "Mullaca." 

Miconia capitata Ule, Notizbl. Bot. Gart. Berlin 6: 360. 1915. 

Branchlets and petioles densely clothed, almost velvety, with 
shortly stiped or subsessile stellate trichomes; petioles 2-3.5 cm. 
long; leaves membranous, broadly obovate-elliptic, shortly acumi- 
nate, cuneate at base, 1.5-2.5 dm. long, 1 dm. wide or wider, soon 
glabrous above, lightly pubescent beneath with loosely stellate 
branched trichomes, especially on the 5-7 (2 faint) nerves and at 
margins, the trichomes sessile or shortly stiped; flowers 5-merous, 
sessile in a small capitately congested panicle, 2-3 cm. long and 
broad; calyx 5.5 mm. long, the dilated limb with 5 separate fascicles 
of stellate trichomes or glabrate; petals 5 mm. long; anthers linear- 
ellipsoid, attenuate, 5 mm. long, the connective anteriorly gibbous 
at base; style 6 mm. long. Section Eumiconia. 

Loreto: Pongo de Cainarachi, Ule 6367, type. Balsapuerto, 
1 meter shrub with white and rose flowers, Klug 2873 (det. Gl.). 

Miconia cauingia Macbr., spec. nov. 

Arbuscula glaberrima 2-4 dm. alta; petiolis usque 10 mm. longis; 
f oliis ovatis vel ovato-ellipticis basi rotundatis et minute emarginatis, 
obtusis, usque 2.5 cm. longis, 13 mm. latis, supra nitidulis fere planis 
vel paullo bullatis, subtus pallidioribus nervis (3) et venis reticulatis 
paullo notatis, rigide coriaceis, margine minute vel obscure calloso- 
crenulatis; paniculis paucifloris circa 3 cm. longis; floribus 4-meris, 
pedicellis 2-4 mm. longis; calyce late campanulato undulato obscure 
lobato, apice 5 mm. lato; petalis late obovato-rotundatis 2.5 mm. 
longis; staminibus glabris, filamentis 2.5, antheris 1.5 mm. longis; 
stylo leviter piloso circa 3 mm. longis, stigmate vix dilatato. 
Handsome shrub with glossy leaves and waxy white flowers some- 
times faintly rose-colored within; fruit said to be edible (Stork & 
Horton). Section Chaenopleura. 



FLORA OF PERU 395 

La Libertad: Above Cachicadan, against boulders, open areas, 
3,500 meters, Stork & Norton 9992, type. "Caningia." 

Miconia cayumbensis Gleason, Bull. Torrey Club 66: 416. 
1939. 

Slender terete younger branches, leaves beneath and calyces 
minutely ashy-stellate-pubescent; petioles 5-8 mm. long; leaves 
firm, glabrous above, 3-nerved, elliptic or oblong, acute at base, 
shortly acuminate, obscurely and remotely denticulate, to 16 cm. 
long, 6 cm. wide; inflorescence paniculate, about 10 cm. long; 
flowers sessile, 5-merous, the campanulate calyx 2 mm. long, the 
broadly rounded sepals 0.3-0.4 mm. long, the outer teeth reduced 
to a callus; petals 2.8 mm. long, narrow; stamens dimorphic, the 
filaments 3.3 and 2.8 mm. long; anthers thick-linear; connective 
in inner series with one dorsal lobe and 2 lateral. Width of anther 
pore suggests Eumiconia; with more natural classification will 
probably be placed near M. longifolia and M. minutiflora now 
assigned to section Glossocentrum (Gleason). These also have 
dimorphic stamens but the former has plinerved leaves and in the 
latter as in M. puberula the leaves are rounded at base. A tree 
10 meters high, or a clump shrub, the flowers fragrant, white. 

Huanuco: Near mouth of Rio Cayumba, 880 meters (Mexia 
8285, type). Junin: La Merced, 5476. Loreto: Balsapuerto, Klug 
3036. 

Miconia cecidophora Naud. Ann. Sci. Nat. se>. 3. 16: 166. 
1851; 796. 

Branchlets pilose(?); petioles 1-3 cm. long; leaves chartaceous, 
repand-denticulate, glabrous above, obscurely and most minutely 
scurfy (or pilose?) beneath on the 3-5 nerves, stellate-punctate, 
oblong-lanceolate, long-caudate-acuminate, 1-2 dm. long, 3-7 cm. 
broad; flowers not secund, the corymbs crowded; calyx campanulate, 
minutely and acutely 5-lobed, 2.5 mm. long; petals 2.5-3 mm. 
long, narrowly obovate; style 5-6 mm. long, the stigma clavate- 
punctate. The mature leaves show only some scattered minute 
scales beneath, are often scarcely 10 cm. long, 3 cm. wide and with 
no pilose or lepidote pubescence. Section Eumiconia. F.M. Negs. 
17033; 36285. 

Loreto: Pampa del Sacramento, Castelnau. Brazil. 

Miconia centrandra Ule, Notizbl. Bot. Gart. Berlin 6: 361. 1915. 

Simulates closely M. centrodesma but the flowers are sessile and 
4-merous; calyx scarcely 1.5 mm. long, the limb truncate, mucronu- 



396 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

late by the outer teeth, completely smooth; petals bright yellow, 
round-elliptic; style 5 mm. long; anthers ellipsoid, the connective 
spurred posteriorly; ovary 4-celled. Placed by the author in section 
Glossocentrum but apparently rather belongs to section Amblyarrhena. 
F.M. Neg. 17034. 

Rio Acre: Ule 9675, type. 

Miconia centrodesma Naud. Ann. Sci. Nat. se>. 3. 16: 164. 
1851; 850. 

A glabrous slender-branched shrub or the branchlet tips and the 
lax panicles minutely and sparsely scurfy; petioles 1.5-5 cm. long; 
leaves chartaceous, 5-plinerved, usually broadly elliptic, rounded 
to the acute base, sharply acuminate, obscurely appressed-ciliate 
on the subentire or repand-crenulate margins, 6-12 cm. broad, often 
twice as long; calyces about 2 mm. long, pedicellate or subsessile; 
petals white, to 1.5 mm. long; connective conspicuously spurred. 
Section Glossocentrum. F.M. Neg. 17035. 

Loreto: Mouth of Rio Blanco, Tessmann 3034 (det. Markgr.). Rio 
Nanay, Williams 1127 (det. Gl.). Brazil; Colombia; Nicaragua. 

Miconia centrophora Naud. Ann. Sci. Nat. se"r. 3. 16: 209. 
1851; 883. 

Similar to M. aspergillaris but leaves becoming twice as large, 
minutely scurfy-stellate only on the nerves and veins beneath, the 
petioles 1-2 cm. long, the petals yellow, 1.5 mm. long, and the 
stigma subcapitate. There are a few stiffer trichomes mixed with 
the dense scurfy-stellate pubescence in panicles and on branchlets. 
M. pichinchensis Benth., 881, was found by Ruiz & Pavon as near 
as Guayaquil; its thin, subsessile leaves are subcordate at base, the 
flowers openly borne on pedicels 2-7 mm. long. Section Amblyar- 
rhena. F.M. Neg. 25938 (as to Mathews plant). 

Amazonas: Chachapoyas, Mathews 1263. Cajamarca: Pass south 
of Conchon, prov. Chota, 2,500 meters, Stork & Horton 10079. 
Ecuador? 

Miconia Chamissois Naud. Ann. Sci. Nat. seY. 3. 16: 179. 
1851; 800. 

Glabrous shrub (or inflorescence reddish-puberulent) with ob- 
tusely angled branches or these lightly compressed above, and firm 
3-5-plinerved leaves; petioles to 2 cm. long; leaves ovate, more or 
less acuminate, sometimes scarcely decurrent at base, to 2 dm. long, 
about half as broad; panicle ample, the sessile flowers crowded at 



FLORA OF PERU 397 

the tips of the branches; calyx campanulate, 2.5-3 mm. long, the 
limb sub truncate; petals white or pale rose, somewhat truncate, 
about 2 mm. long; stamens dimorphic, the connective of the larger 
prolonged and broadly dilated, that of the smaller shorter with 2 
deflexed lateral lobes (Gleason); style 5 mm. long. The Peruvian 
collections are reddish-puberulent in inflorescence and perhaps should 
be studied further, especially since the flowers are not aggregated. 
Section Eumiconia. F.M. Neg. 17037. 

Loreto: Iquitos, Williams 7982; Killip & Smith 27178; King 339 
(det. GL). Widely distributed, South America. 

Miconia chionophila Naud. Ann. Sci. Nat. se>. 3. 16: 236. 
1851; 935. 

Branches crisply hirsutulous, spreading or possibly scandent; 
petioles hirtellous, 0.5-2 cm. long; leaves narrowly ovate or sub- 
rotund, more or less rounded at base, obtuse, sparsely setulose or 
glabrate above and beneath, the margins undulate and ciliate, 
1-2.5 cm. long, 6-15 mm. broad, 3-nerved; flowers usually ternate; 
calyx 3-4 mm. long; petals retuse, 3 mm. long; style 3 mm. long, 
the stigma broadly peltate. The type from Colombia is rather more 
pubescent than the Peruvian form; Triana wrote the name chiono- 
phylla. Doubtfully distinct from M. rotundifolia. Section Chaeno- 
pleura. F.M. Neg. 17038; 36286. 

Puno: Tabina, Lechler 2063; 2065. Bolivia; Colombia. 

Miconia chrysanthera Cogn. Bot. Jahrb. 42: 141. 1908. 

Shrub 3 meters high, the obtusely angled younger branches as 
the petioles and leaves beneath, these more especially on the 5 
nerves, stellate-puberulent; petioles 8-25 mm. long; leaves nearly 
membranous, plane, narrowly ovate, rounded at base, entire or sub- 
entire, very shortly acuminate, glabrous above in age, sometimes 
sub-5-plinerved, 4-8 cm. long, 2-3.5 cm. broad; panicle peduncles 
stellate and sparsely setulose, the many 5-merous flowers on pedicels 
1-2 mm. long; calyx campanulate-ovoid, slightly stellate-scurfy, 
about 2 mm. long; style filiform, 6-7 mm. long, scarcely dilated 
apically. Not clearly distinct from M. centrophora according to 
Cogn. Near M. pichinchensis Benth. Section Amblyarrhena. F.M. 
Neg. 17040. 

Cajamarca: Below San Pablo, 2,200 meters, Weberbauer 3847, 
type. 

Miconia chrysophylla (L. C. Rich.) Urb. Symb. Ant. 4: 459. 
1910; 857. Melastoma chrysophylla L. C. Rich. Act. Soc. Hist. Nat. 



398 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Paris 109, 1792. Melastoma fulva L. C. Rich, in Bonpl. Melast. 23. 
pi. 11. 1816. Miconiafulva DC. Prodr. 3: 180. 1828. 

Shrub or tree well marked by the ashy or reddish-lepidote indu- 
ment that clothes the younger acutely angled branches and the 
leaves beneath, these green and glabrous above, narrowed both 
ends, sharply acuminate, 3-nerved with also a faint pair of sub- 
marginal ones; panicle branches secundly flowered; calyx 1.5 mm. 
long, subtruncate; petals yellowish, about 1.5 mm. long; connective 
prolonged, unappendaged ; style 3 mm. long. The upper leaves are 
usually verticillate, 3-5 cm. broad, 3 to several times longer. Sec- 
tion Chaenanthera. 

Loreto: Iquitos, Williams 3660. Rio Nanay, Williams 1151. 
Rio Huallaga, Williams 4809. Rio Maranon, Killip & Smith 
27530. Rio Acre: 25-meter tree, Krukoff 5268. Brazil to Guianas. 
"Puca mullaca." 

Miconia ciliaris Triana, Trans. Linn. Soc. 27: 105. 1871; 737. 

Branches slightly scurfy; petioles 1.5-4 cm. long; leaves ovate, 
caudately acuminate, sparsely setulose above, a little scurfy on 
nerves and veins beneath, the minutely denticulate margins hispid- 
ciliate, 5-7-nerved, or slightly 5-plinerved, about 9 cm. wide, often 
1.5-2 dm. long; inflorescence with a compact, reddish puberulence; 
pedicels 2-7 mm. long; calyx campanulate, 3.5-4 mm. long; petals 
4-5 mm. long, glabrate; filaments glandular-pilose, the anthers 
eglandular, the connective simple; ovary 5-celled, glabrous; style 
glandular pilose, 6-7 mm. long, the stigma subpeltate. The leaves 
are slightly cordate at base. Cf. M. muricata. Section Tamonea. 
F.M. Neg. 29499. 

Junin: Vitoc, Ruiz & Pawn, type. 

Miconia ciliata (L. C. Rich.) DC. Prodr. 3: 179. 1828; 867. 
Melastoma ciliata L. C. Rich. Act. Soc. Hist. Nat. Paris 109. 1792. 

Nearly glabrous shrub usually marked by the slightly serrulate 
but rather conspicuously ciliate leaf margins, oblong or obovate- 
oblong leaves, these 3-nerved with an additional pair of faint ones, 
mostly 8-12 cm. long, 3-7 cm. broad; petioles 1-3 cm. long, often 
slightly pilose above; calyces sessile, secund, glabrous, 2-2.5 mm. 
long; petals roseate, about 2.5 mm. long; connective not prolonged; 
style 2 mm. long. Var. congestiflora (DC.) Cogn. has the flowers 
clustered on reduced branches. Section Amblyarrhena. 



FLORA OF PERU 399 

San Martin: Tarapoto, Spruce 4257 (the var.). Loreto: Con- 
chamaya (Huber 1451). Canela Ucsha, Ule 6737, var. (det. Pilger). 
Brazil to Mexico and the West Indies. 

Miconia clathrantha [Mart.] Triana, Trans. Linn. Soc. 28: 129. 
1871 (name); 918. Cremanium clathranthum Mart, ex Triana, 
I.e., in syn. 

Glabrous, with slender, obtusely 4-sided branchlets; petioles 
1-3 cm. long; leaves membranous, obovate-oblong, scarcely denticu- 
late, shortly acuminate, very unequal, the larger to about 2 dm. 
long, 5-10 cm. broad, 3-nerved with an additional pair of faint 
marginal ones; flowers paniculate, sessile, minute, the obscurely 
denticulate calyx scarcely 1 mm. long, the petals 0.5 mm. long; 
style 1 mm. long, the stigma obtuse. The extremely thin leaves 
show prominently many veins; the panicles are 5-8 cm. long. Sec- 
tion Cremanium. F.M. Neg. 17041. 

Huanuco: Cuchero, Poeppig 110; 1719, type. Junin: Pichis 
Trail, 1,100 meters, Kittip & Smith 26051; 26187 (det. Gl.). 

Miconia clavistila Gleason, Bull. Torrey Club 58: 229. 1931. 

Similar to M. stelligera Cogn.; leaves broadly oblong, narrowed 
to the rounded or obscurely subcordate base, subentire, 3-nerved 
but with a submarginal additional pair, the upper surface opaque 
and glabrous except the minutely stellate-tomentose mid-vein, to 
16 cm. long and half as wide; filaments filiform, anthers 4.5 mm. 
long, connective 2-lobed at base and curved anteriorly into 2 lateral 
lobes; style straight and slender, 8 mm. long, after anthesis conspicu- 
ously enlarged above, the truncate stigma 0.7 mm. in diameter. 
Said by author to differ from both M. barbinervis and M. stelligera 
"in leaf -form and elongated style" but these differences seem to be 
scarcely apparent. However in the Mexia collection the trichome- 
stipes are much shorter, the pubescence thus more compact than 
in the other species. Section Eumiconia. 

Loreto: Rio Nanay, Williams 373, type. Above Pongo de 
Manseriche, Rio Santiago, Mexia 6240 (det. Gl.). San Martin: 
Pongo de Cainarachi, Klug 2629. 

Miconia coelestis [Pav.j Naud. Ann. Sci. Nat. se>. 3. 16: 245. 
1851; 919. Melastoma coelestis Pav. ex D. Don, Mem. Wern. Soc. 
4: 313. 1823, in syn. Cremanium coeleste D. Don, I.e. Miconia 
chrysopetala Naud. I.e. 233. 

Allied to M. clathrantha but the stouter, at first acutely angled, 
branches and the nerves beneath of the minutely serrulate-ciliate 



400 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves early faintly scurfy, becoming glabrous, the sparsely and 
openly branched inflorescence scurf y-puberulent, glabrate in age; 
leaves 3-nerved with an additional obscure marginal pair, often 
2 dm. long, 7-10 cm. wide, yellowish green (or drying brown), the 
transverse veins prominent; panicle to 2 dm. long, flowers sessile, 
yellowish, the often reddish suburceolate calyx 2 mm. long, the 
yellow petals less than 1 mm. long; style 4 mm. long, the stigma 
peltate. In type the leaves are nearly oblong, obovate, 4-6 cm. 
wide, sometimes only 2 cm. wide, 8 cm. long. Specimens Klug 635 
and 591 from Mishuyacu, very young, apparently might key here; 
they are especially marked by a minute, seemingly simple pubes- 
cence on the veinlets of the leaves beneath. Tree 8 meters high 
(Mexia). F.M. Neg. 17042. 

Huanuco: Chinchao and Chacahuasi, Ruiz & Pavdn, type. 
Pampayacu, region Churubamba in dense rain forest, 1,625 meters, 
Mexia 8247 (det. Gl.); Poeppig 1268.J\mm: Pichis Trail, Killip 
& Smith 25924 (det. Gl.). Bolivia. 

Miconia comosa Cogn. in Mart. Fl. Bras. 14, pt. 4: 408. 1888; 
867. 

Similar to M. ciliata but the petioles usually only 2-7 mm. long, 
the branchlets and leaves beneath on the nerves papillose-hirtellous, 
the marginal ciliation appressed and the floral bracts 3-5 mm. long. 
The leaves are entire except for the long but sparse ciliation. Section 
Amblyarrhena. F.M. Neg. 32334. 

San Martin: Tarapoto, Spruce 4438, type. 

Miconia compacta Gleason, Bull. Torrey Club 58: 230. 1931. 
M. semota Markgr. Notizbl. Bot. Gart. Berlin 12: 177. 1934. 

Branches terete, densely brown-stellate- tomentose; petioles stout, 
to 17 mm. long; leaves elliptic, 9-17 cm. long, about half as wide, 
acute or abruptly acuminate, rounded at base, entire, 3-nerved or 
3-5-plinerved, membranous, glabrous above except along the pri- 
maries, loosely stellate-pubescent below, especially on the veins; 
panicle racemiform, tomentose, the 6-merous sessile flowers in 
terminal glomerules on the short branches, these only 3-5 mm. 
long; bracts lanceolate, persistent, 2-3 mm. long; calyx about 3.5 
mm. long, the stellate trichomes sessile, the broadly ovate reflexed 
sepals 1.3 mm. long, not rounded but obtuse; filaments glandular, 
the connective of the larger tuberculate dorsally, shortly produced 
and glandular basally; ovary 3-celled, glandular; style glabrous, the 
stigma ovoid, 4 mm. long. Allied to M. glomerata Triana to which 



FLORA OF PERU 401 

the Tessmann collections were referred at Berlin. Suggests M . cin- 
chonaefolia DC., 745, which however has eglandular filaments; 
subulate anthers. Section Eumiconia. 

Loreto: Iquitos, Killip & Smith 26936, type; 27354; King 1050; 
Tessmann 3648; 3612. 

Miconia consi mills Pilger, Verh. Bot. Ver. Brandenb. 47: 
170. 1905. 

Compressed younger branches, petioles and inflorescence minutely 
brown-scurf y-puberulent; petioles 2-3.5 cm. long; leaves elliptical, 
narrowed at base, gradually acuminate, very membranous, bright 
green and glabrous except for a minute brown-punctate scurfiness on 
the 5 nerves beneath, to 2 dm. long, half as broad, loosely reticulate- 
veined; calyx shortly pedicellate, 2 mm. long; petals 4, yellowish- 
white, 3 mm. long; anthers lanceolate, slightly attenuate above, 
1-pored, 4 mm. long, the connective shortly spurred; ovary free; 
style to 1 cm. long, the stigma not at all enlarged. In spite of some- 
what pointed anthers, belongs in Glossocentrum. F.M. Neg. 17043. 

Loreto: Cerro de Escaler, 800 meters, Ule 6543, type. Rio 
Nanay, Williams 689. Florida, King 1997(1). "Isula-micuna." 

Miconia Cookii Gleason, Phytologia 1: 43. 1933. 

Similar to M. elongata; petioles slender, 2-3 cm. long; 'leaves 
abruptly acuminate to a short cusp, 3-plinerved; calyx subsessile, 
about 2.5 mm. long, very sparsely and minutely puberulent and 
brown-punctate, the triangular acute sepals nearly 1 mm. long; 
petals round-obovate, 2.2 mm. long; connective elevated and later- 
ally lobed; ovary 5-celled, the style 5.5 mm. long, the capitellate 
stigma barely expanded. Allied also to M. monzoniensis with 
crowded panicles, shorter petioles, broader leaves, 4-celled ovary, 
and to M. Bangii Cogn. Mem. Torrey Club 3, no. 3: 30. 1893, of 
Bolivia, the leaves broader, the stigma subpeltate. Apparently is 
M. Terera at least in part. Section Amblyarrhena. 

Cuzco: San Miguel, Urubamba Valley, 1,800 meters (Cook & 
Gilbert 1742, type). 

Miconia corymbiformis Cogn. Bull. Acad. Belg. se>. 3. 14: 
964. 1887; 929. 

A completely glabrous tree or the leaves with a few scattered and 
evanescent scales beneath; branches acutely tetragonous or nearly 
winged; petioles to 4 cm. long; leaves rigid-coriaceous, 1-2 dm. long, 
4-9 cm. broad, with 5 thick nerves, oblong or ovate-oblong, rounded 



402 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

at base, the entire margin lightly revolute; flowers densely corymbed 
in panicles to 1.5 dm. long, the pedicel 4-8 mm. long; calyx tube 
6 mm. long, the lobes broadly rounded, 1.5-2 mm. long; petals 
yellowish, broadly rounded, 4-5 mm. long, the style 5 mm. long. A 
Ruiz & Pavon collection labeled as from Peru was actually obtained 
at Guayaquil. Section Chaenopleura. 

Peru (probably). Ecuador; Colombia. 

Miconia crassifolia Triana, Trans. Linn. Soc. 28: 125. 1871; 881. 

Terete branches, leaves beneath and compact inflorescences even 
to the calyces (or these glabrate in age) scurfy stellate-tomentose 
with dark-brown short trichomes; petioles 3-6 mm. long; leaves sub- 
orbicular to ovate-oblong, entire or denticulate toward the acute or 
obtusish apex, fleshy-coriaceous, yellowish and glabrous but gran- 
ulose-papillate above, 5-plinerved with an additional faint marginal 
pair, mostly 6 cm. long, 4 cm. broad; pedicels 1-3 mm. long, the 
calyx 3-4 mm. long; petals suborbicular, 3 mm. long; connective 
bitubercled posteriorly; style sometimes stellate-puberulent, 4-5 mm. 
long, the stigma peltate. The leaf nerves are essentially or quite 
basal. Two leaf-forms have been named, var. parvifolia Cogn. and 
var. rotundifolia Cogn. Section Amblyarrhena. F.M. Neg. 25942. 

Amazonas: Chachapoyas, Mathews 1261; 1262; 3216, types; 
Williams 7555 (det. Gl.). 

Miconia crassipes Triana, Trans. Linn. Soc. 28: 116. 1871; 781. 

Branches obtusely 4-sided; petioles thick, 1-2.5 cm. long; leaves 
elliptic or elliptic-oblong, acute, auriculate-cordate at base, scarcely 
denticulate, coriaceous, glabrous above, pale or rusty scurfy-tomen- 
tose beneath, the indument compact, 12-18 cm. long, 6-10 cm. broad ; 
flowers glomerate on the subverticillate panicle branches, the ovoid- 
turbinate calyx 2.5 mm. long; petals retuse, 2-2.5 mm. long; ovary 
scurfy-puberulent above; style 4-5 mm. long. Section Eumiconia. 

Peru: (Mathews 1300). 

Miconia crassistigma Cogn. Bot. Jahrb. 42: 143. 1908. 

Subquadrate branchlets densely scurf y-stellate-puberulent; peti- 
oles 1-2 cm. long; leaves firm, oblong, acute, entire but minutely 
rigid-ciliate-denticulate, glabrous above, sparsely stellate-scurfy, 
mostly only on the nerves and slightly papillose-punctate beneath, 
3-nerved with an additional faint submarginal pair, 4-7 cm. long, 
2-2.5 cm. broad; panicle dense; flowers very minute, the calyx only 
1.5 mm. long, the white, slightly emarginate petals shorter than 1 



FLORA OF PERU 403 

mm.; pedicels 0.5-1 mm. long; style filiform, 2 mm. long. Allied to 
M. Mandonii, 911, of adjacent Bolivia, with petals twice as long, 
the calyx nearly 4 mm. broad on pedicels 1-6 mm. long. Section 
Cremanium. F.M. Neg. 17044. 

Huanuco: Prov. Huamalies, 2,500 meters, Weberbauer 3407. 

Miconia cremophylla Naud. Ann. Sci. Nat. se>. 3. 16: 228. 
1851; 919. 

Similar to M. coelestis; leaves glabrous above, sparsely brown- 
punctate-scurfy beneath, 1-1.5 dm. long, 4-5.5 cm. wide, only 
shortly narrowed at base; petioles 1-2 cm. long; flowers usually 
ternate; petals 1.5 mm. long; calyx obtusely lobed; style 3-4 mm. 
long. In the Peruvian specimens the leaves are subcoriaceous, yel- 
lowish green, remotely mucronulate on the margins, caudate-acu- 
minate, with 3 nerves and 2 faint additional ones; the indument is 
merely a pulverulence. Section Cremanium. F.M. Neg. 25944. 

Amazonas: Chachapoyas, Mathews 1297. Without locality, 
Pawn. Colombia; Bolivia. 

Miconia crocea (Desr.) Naud. Ann. Sci. Nat. se>. 3. 16: 245. 
1851; 897. Melastoma crocea Desr. in Lam. Encycl. 4: 55. 1797. 

Nearly glabrous but setose-barbate, usually conspicuously, at 
base of leaves beneath on the nerves; petioles 1-1.5 cm. long, the 
younger somewhat setulose; leaves ovate-oblong, obtusely or shortly 
acuminate, subrotund or acute at base, remotely denticulate, 3-5- 
nerved or nearly 3-plinerved, the outer nerves thin, 5-6 cm. long, 
2.5-3 cm. wide; panicle dense, the flowers 7-8-merous; pedicels 1-2 
mm. long; calyx 3-4 mm. long, the minute teeth acute; petals white, 
2.5-3 mm. long; style 6-7 mm. long, the stigma subpeltate. The 
Ruiz & Pavon collection cited in litt. as Peruvian was collected at 
Guayaquil. Section Cremanium. 

Peru: (Spruce 4971). Ecuador; Colombia. "Colca," "tin." 

Miconia cuneata [Berg] Triana (name), Trans. Linn. Soc. 28: 
113. 1871; 826. Staphydium cuneatum Berg ex Triana, I.e., in syn. 

Branches, petioles and panicles densely long-hirsute, the longer 
trichomes 2 mm. long, 3-5-stellately branched at tip; petioles excep- 
tionally stout, 0.5-1.5 cm. long; leaves oblong-lanceolate, long-ciliate 
and slightly undulate-denticulate, long-acuminate, sparsely setulose 
above, densely hirsute especially on the 3-5 nerves beneath, the 
trichomes many-parted at apex, the larger leaves 1.5-2 dm. long, 
5-9 cm. broad, the opposite leaf usually distinctly smaller; panicle 



404 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3-8 cm. long; calyx 3 mm. long, sessile; petals 2 mm. long; style 5-6 
mm. long. Section Eumiconia. F.M. Neg. 17045. 

San Martin: Rio Tocache, Poeppig 2046; 2047, types. 

Miconia cuspidata [Mart.] Naud. Ann. Sci. Nat. se"r. 3. 16: 
163. 1851; 809. Oxymeris cuspidata Mart, ex Triana, Trans. Linn. 
Soc. 28: 109. 1871. 

Branches nearly terete, the younger petioles, leaves beneath and 
calyces minutely scurfy-puberulent, the indument evanescent except 
in the panicles, these 5-7 cm. long; petioles 4-6 mm. long; leaves 
oblong-lanceolate, entire, submembranous, strikingly acuminate, the 
acumen slender or nearly filiform, drying black and lustrous, 6-10 
cm. long, 1.5-3 cm. broad; pedicels 2-3 mm. long; calyx tube 2 mm. 
long, the lobes 0.5 mm. long; petals about 3 mm. long; connective 
shortly spurred posteriorly, minutely biauricled anteriorly; style 
truncate at apex, 5-6 mm. long. Section Eumiconia. F.M. Neg. 
17047. 

Peru (probably). Amazonian Brazil. 

Miconia cyanocarpa Naud. Ann. Sci. Nat. se>. 3. 16: 221. 
1851; 899. 

Allied to M. caerulea; branches slender, shortly (typically) and 
densely hirsute; petioles to 3 cm. long; leaves ovate, more or less 
acuminate, rounded at base, thin and very green but hirtellous on 
both sides, more loosely so beneath, 5-7-nerved, 8-15 cm. long, 
usually half as wide; flowers densely glomerate, the clusters often 
continuous; calyx obviously dentate, 1.5 mm. long; petals scarcely 
1 mm. long; fruit blue, 3 mm. thick. The var. hirsuta Cogn. is 
densely long-hirsute on the branches. Section Cremanium. 

Junin: San Ramon, Killip & Smith 24690 (det. Gl.). Chancha- 
mayo Valley, Schunke 1739; 395. Puno: Sandia Valley, Weberbauer 
1067 (det. Cogn.); 278. Without locality (Haenke). Bolivia. 

Miconia decurrens Cogn. Bol. Mus. Goeldi 5: 253. 1909. 

Similar to M. aureoides; leaves membranous, broadly lanceolate, 
long-acuminate, 5-7-plinerved, glabrous both sides but pale beneath, 
18-25 cm. long, 5-8 cm. broad; lacerate calyx segments finally 
caducous, the calyx tube 2.5-3 mm. long, the membranous limb 
1.5-2 mm. long; petals white, only about 2 mm. long; stigma puncti- 
form. Petioles winged nearly to base by the decurrent leaf blade. 
Section Laceraria. 

Loreto: Iquitos, in woods (Ducke 7603, type). 



FLORA OF PERU 405 

Miconia densifolia Cogn. Bot. Jahrb. 42: 145. 1908. 

Allied to M. cremophylla but entirely glabrous and the branches 
acutely 4-sided or narrowly winged; petioles 4-8 mm. long; leaves 
narrowly oblong, narrowed to an obtuse base, 6-8 cm. long, 1.5- 
nearly 2.5 cm. broad, remotely callous-denticulate, 3-nerved; calyx 
about 1 mm. long, sessile, its lobes acute; flowers minute, the petals 
about 0.5 mm. long, aggregate; style extremely short, the stigma 
subpeltate. Section Cremanium. F.M. Neg. 17048. 

Huanuco: Prov. Huamalies, 2,500 meters, Weberbauer 3404, type. 
Playapampa, 4859 (?) (in fruit). 

Miconia dipsacea Naud. Ann. Sci. Nat. se>. 3. 16: 138. 1851; 775. 

Allied to M. amplexicaulis but the more rigid, nearly oblong 3- 
nerved leaves auriculate-cordate and densely stellate-tomentose 
beneath; calyx 3 mm. long, shortly 5-lobed, the pubescence persist- 
ing; petals rounded at apex, 3 mm. long; style 5-6 mm. long, the 
stigma subpeltate. Compact shrub or tree to 5 meters high. Sec- 
tion Eumiconia. F.M. Neg. 17007. 

Huanuco: Cueva Grande, 4790. Pozuzo, 4707; Valley of the 
Monzon, 900 meters, Weberbauer 3480 (det. Cogn.); 284. Pam- 
payacu, Kanehira 110; Cuchero, Poeppig 176. Chinchao, Ruiz & 
Pawn (det. Markgr.). Casapi, Mathews 1722.- Junin: La Merced, 
5541. Chanchamayo Valley, Weberbauer 2016 (det. Cogn.); 247. 
Ayacucho: Prov. Huanta, Weberbauer 5647 (det. Cogn.). Cuzco: 
Valle de Lares (Herrera 800). Rio Chaupimayo, Soukup 506, in 
part. "Ppuino-ppuino." 

Miconia dispar Benth. in Hook. Journ. Bot. & Kew Misc. 2: 
241. 1850; 791. 

Branches densely and shortly tomentose-puberulent with plumose 
trichomes; petioles 4-5 sided, 2-4 cm. long; leaves obscurely undulate- 
crenulate, ovate-oblong, smooth above, often drying black, fulvescent 
beneath with a dense pubescence of stellate trichomes, 5-nerved, 
2-3.5 dm. long, 9-18 cm. broad; panicle 1-2.5 dm. long, the short 
lateral branches opposite and mostly trifid, the flowers sessile and 
secund on these 3 forks; calyx shortly toothed, about 2.5 (-3.5) mm. 
long; petals pale rose-color, 3 mm. long; style 6-7 mm. long; anthers 
3-3.5 mm. long; connective produced, minutely gibbous posteriorly, 
obscurely bilobed anteriorly. Tree to 10 meters high. Section 
Eumiconia. F.M. Neg. 17051. 

Junin: Chanchamayo Valley, Schunke 303; A. 121. Cuzco: Rio 
Chaupimayo, Soukup 506, in part. Brazil. 



406 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia dodecandra (Desr.) Cogn. in Mart. Fl. Bras. 14, pt. 
4: 243. 1887; 740. Melastoma dodecandra Desr. in Lam. Encycl. 
4: 46. 1797. 

Candelabra-branched shrub or small tree, the younger branches, 
petioles, leaves beneath and panicles densely stellate-puberulent or 
tomentose; petioles elongate; leaves firm, ovate, shortly acuminate, 
somewhat rounded at base, usually 5-nerved, 12-18 cm. long, 6-7 
cm. broad; pedicels 4-10 mm. long, the geminate, obtuse, involucrate 
but caducous bracts 6-7 mm. long; flowers mostly 6-merous; calyx 
5-6 mm. long; petals thick, 7-8 mm. long; connective slightly gibbous 
dorsally ; style glabrous below. The calyx is distinctly lobed and per- 
sistently ashy-tomentose. Mexia 6454 has a pubescent style. 
Section Tamonea. 

Loreto: Mouth of Santiago, in dense forest, Mexia 6192 (det. Gl.). 
Rio Napo, Mexia 6454 (det. GL). Junin: La Merced, Killip & Smith 
23920 (det. Gl.); Schunke 1827. San Martin: Moyobamba, Woyt- 
kowski 29. Brazil to Mexico and the West Indies. "Calsoncillo." 

Miconia dolichorrhyncha Naud. Ann. Sci. Nat. se*r. 3. 16: 
166. 1851; 859. 

Branches scurfy- tomentose; petioles about 1 cm. long; leaves 
oblong, rounded at base, very long-caudate-acuminate, minutely 
reddish-stellate-hirtellous beneath, 5-nerved, 1.5-2 dm. long, 5-7 
cm. broad; panicles about 1 dm. long, the pedicels mostly 1-2 mm. 
long; calyx 1.5-2 mm. long; petals obtuse, 2 mm. long; anthers 2- 
cleft, the long-produced connective spurred posteriorly; style 3 mm. 
long; fruits yellow. Section Chaenanthera. F.M. Neg. 36292. 

Junin: Pichis Trail, Killip & Smith 25889. La Merced, 5589. 
San Martin: Moyobamba, Mathews 9291, type. Loreto: Rio Para- 
napura, Klug 3934. Bolivia. 

Miconia Donaeana Naud. Ann. Sci. Nat. se>. 3. 16: 244. 1851 
(name); 738. Chitonia caudata D. Don, Mem. Wern. Soc. 4: 317. 
1823, not M. caudata DC. 

Branches, as the 5 leaf nerves beneath, slightly minutely scurfy; 
petioles 4-7 mm. long; leaves ovate, or nearly rotund, more or less 
abruptly, narrowly and sharply caudate, glabrous above, 1.5-3 dm. 
long, 1-2 dm. wide; flowers sessile or subsessile, bibracteate, the 
bracts 2-4 mm. long, promptly caducous; calyx about 3 mm. long, 
densely scurf y-tomentose, obscurely lobed; petals roseate, 4 mm. 
long; filaments and connective densely glandular-puberulent; style 



FLORA OF PERU 407 

10-12 mm. long, the stigma peltate. Small shrub or tree (Schunke). 
Section Tamonea. F.M. Neg. 17052. 

Junin: Chanchamayo Valley, Schunke 282; 338. Huanuco: 
Cuchero, Ruiz & Pavon; Dombey; Poeppig 1213. Chinchao, Ruiz 
& Pavon. San Martin: San Roque, Williams 7002; 7055 (det. Gl.). 
"Muena-mullaca," "nucnu mullaca." 

Miconia Duckei Cogn. Bol. Mus. Goeldi 5: 254. 1909. 

Younger parts, as the leaves beneath, at least on the 5 nerves 
shortly and densely stellate-tomentose, the nerves basal or nearly, 
or 3 distinctly above the leaf base; petioles slender to 3.5 cm. long; 
leaves elliptic-obovate, obtuse at base, apically obtusish to shortly 
acuminate, entire or crenulate, membranous, pale green above and 
finally glabrous, minutely and rather sparsely stellate-pilose beneath, 
15-17 cm. long, 8-10 cm. broad; panicle 2-6 cm. long, the flowers 
sessile, densely congested, or in nearly continuous glomerules or 
these rarely at the ends of branchlets; calyx ashy-hirtellous, the 
lacerate limb puberulent, about 2 mm. broad, about as long; petals 
white, narrowly obovate, to 2 mm. long; anthers eglandular; style 
filiform, glabrous, 5 mm. long, the stigma punctiform. The follow- 
ing material was distributed as M. capitata, a species with different 
calyx. The type has 5-nerved, nearly entire, obtusish leaves, but 
apparently the leaves vary as described here. Section Laceraria. 
F.M. Neg. 25952. 

Loreto: Iquitos, in woods (Ducke 7586, type); Killip & Smith 
27284; Williams 1327; 1338. Pebas, Williams 1634. La Victoria, 
Williams 2993. 

Miconia dumetosa Cogn. Bot. Jahrb. 42: 144. 1908. 

Densely branched shrub 2 meters high, the slender branches, as 
the 3 leaf nerves beneath, stellate-puberulent; petioles 1-3 mm. long; 
leaves coriaceous, obovate-oblong, basally acute, obtuse, in age gla- 
brous above, 1.5-2.5 cm. long, 7-13 mm. broad; panicle dense, to 
2.5 cm. long; flowers sessile or subsessile, 5-merous; calyx campanu- 
late-ovoid, slightly scurfy, 1.5 mm. long, the minute teeth acute, 
petals scarcely 0.5 mm. long; style glabrous, 1.5 mm. long, the 
stigma peltate. Near M. loxensis (Bonpl.) DC. Section Cremanium. 
F.M. Neg. 17053. 

Amazonas: Tambo Ventillas, 2,400 meters, east of Chachapoyas, 
Weberbauer 4389, type. 

Miconia egensis Cogn. in Mart. Fl. Bras. 14, pt. 4: 374. 
1887; 849. 



408 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branches obtusely quadrate, often somewhat 4-sulcate, the 
younger minutely stellate-scurfy; petioles about 1 cm. long; leaves 
ovate or ovate-oblong, acutely acuminate, acute or obtuse at base, 
glabrous except for a minute scurfy puberulence on the leaf-nerves 
beneath, this evanescent, more prominent in the inflorescence, to 
2.5 dm. long, 1-1.5 dm. broad, 3 (-S)-nerved or shortly plinerved, 
the outer pair faint; panicles 1 dm. long, the crowded flowers sessile 
or subsessile; calyx 2 mm. long, scarcely lobed, the limb caducous; 
petals 2 mm. long; stamens subequal, filaments glabrous; connective 
shortly produced but simple; style 5 mm. long, clavate or subpeltate 
above. Three-meter tree with white flowers (Klug). Section Glos- 
socentrum. F.M. Neg. 17054. 

Loreto: Williams 1637; 2703 (det. Gl.). Florida, 2258 (det. GL). 
Brazil. "Acano-fue-paroe," "huirima-ey" (Huitoto). 

Miconia elaeagnoides Cogn. in Mart. Fl. Bras. 14, pt. 4: 390. 
1887; 858. Miconia dichrophylla Macbr. Field Mus. Bot. 4: 189. 
1929(?). 

Branches acutely tetragonous, often strongly flattened toward 
the tip, the younger with the petioles, peduncles and panicles densely 
squamulose-pubescent; leaves membranous or rather firm, opposite, 
oblong-ovate or oblong-lanceolate, mostly 1.5 dm. long, 5 cm. broad, 
sometimes apparently twice as large, acute at base, acuminate, green 
and glabrous above, densely silvery or rusty-lepidote beneath, 5- 
plinerved; flowers 5-merous, sessile; calyx campanulate, about 2 mm. 
long, minutely triangular-dentate; petals suborbicular, about 1.5 
mm. long, glabrous; anthers oblong-linear, the terminal pore mod- 
erately wide, the connective obscurely prolonged at base, minutely 
bicalcarate on both sides; style truncate, short. Description largely 
from the type of M. dichrophylla; Gleason, Bull. Torrey Club 58: 242, 
is probably correct in suggesting that only one species is concerned, 
the Cogniaux specimens being young, and the sepals probably are 
not truly "linear-subulate." Section Eumiconia, fide Gleason. F.M. 
Neg. 25953. 

Junin: Slender tree, La Merced, 5495 (type, M. dichrophylla); 
Killip & Smith 26379 (det. Gl.). San Martin: Williams 5891; 6421; 
6472; 6586 (det. Gl.). Loreto: Williams 4886 (det. GL). Bolivia; 
Brazil. "Ubiamba." 

Miconia elongata Cogn. Bull. Torrey Club 17: 91. 1890; 874. 

Glabrous shrub with rather acutely tetragonous branches, and 
thin lanceolate leaves, narrowed to base and decurrent on the peti- 



FLORA OF PERU 409 

oles, these to 1 cm. long or longer; leaves 3-nerved with an additional 
marginal pair, 1-2 dm. long, 3.5-5 cm. broad; panicle 7-8 cm. long; 
pedicels 1-2 mm. long, medially articulate; calyx ovoid, 1.5 mm. 
long; petals white, obovate, less than 1 mm. long; style 2-2.5 mm. 
long, the stigma peltate. Section Amblyarrhena. Shrub 3 meters 
high (Klug). 

Loreto: Pumayacu, Klug 3223 (det. Gl.). Bolivia. 

Miconia eriocalyx Cogn. in Mart. Fl. Bras. 14, pt. 4: 342. pL 69. 
1887; 823. 

Branches sub terete, evanescently tomentose; petioles 2-5 cm. 
long; leaves minutely crenulate, ovate, cordate at base, acuminate, 
glabrous above, rusty-stellate beneath, 2-3.5 dm. long, 1-2 dm. 
broad, 5-nerved, thick and rigid; flowers crowded on the white stel- 
late- tomentose panicle branches; calyx densely tomentose, about 3 
mm. long, the glabrous petals as long, the style at least twice as 
long; stamens subequal, the scarcely produced connective obscurely 
biauricled anteriorly, scarcely little gibbous posteriorly. The inflo- 
rescence branches are often clothed with a deep felt-like indument. 
A tree 8-10 meters high (Killip & Smith). Section Eumiconia. F.M. 
Neg. 32335. 

San Martin: Tocache, Poeppig. Loreto: Yurimaguas, Killip 
& Smith 27593 (det. Gl.). Brazil. 

Miconia erioclada Triana, Trans. Linn. Soc. 28: 111. 1871; 814. 

Branches subterete, rather densely to conspicuously long-hir- 
sute- villous, the trichomes simple, some a little barbellate basally; 
petioles 0.5-2 cm. long; leaves unequally dentate, sometimes ob- 
scurely, elliptic-oblong, subrotund at base, acuminate, more or less, 
often sparsely, appressed-pilose both sides, 5-nerved, 1-2.5 dm. 
long, 5-10 cm. broad; panicle narrow, few-flowered; bractlets 2-3 
mm. long; calyx to 3 mm. long, distinctly 5-lobed; petals white, 
2 mm. long, connective obscurely bituberculate; style 5 mm. long. 
Common at altitudes of less than 700 meters (Gleason). Section 
Eumiconia. F.M. Neg. 32336. 

Junin: La Merced, sandy brushy river flat, 1-meter bush, 5546; 
Killip & Smith 23469. Huanuco: Cuchero, Ruiz & Pavon; Poeppig 
1049; 1451; 1673. San Martin: Tarapoto, Spruce; Williams 6758 
(det. Gl.). Loreto: Yurimaguas, Poeppig; Williams 294; 3210; 
4940; 4999 (all det. Gl.). Rio Itaya, Killip & Smith 29504. 
Soledad, Killip & Smith 29554. Cuzco: Paucartambo, Soukup 385 
(det. Standl.). Brazil; Bolivia. 



410 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia eugenioides Triana, Trans. Linn. Soc. 28: 109. 
1871; 811. 

Branches terete, the younger densely yellowish stellate-scurfy- 
pulverulent; petioles 1 cm. long; leaves firm, lustrous, ovate or oval- 
elliptic, caudate-acuminate, entire, glabrous or minutely stellate- 
puberulent on the 5 nerves beneath, 2 of these fainter, to 12 cm. long, 
7 cm. broad, often smaller; panicles broad, to 1 dm. long; pedicels 
2-3 mm. long; calyx 1.5-2 mm. long, glabrous above; petals acute, 
3 mm. long; connective anteriorly bituberculate, posteriorly produced 
into a short, truncate appendage; style 5-6 mm. long. Six meters 
high, with white flowers. Suggests Leandra in acute petals. Sec- 
tion Eumiconia. Illustrated, Mart. Fl. Bras. 14, pt. 4, pi. 65. 

Loreto: La Victoria, Williams 2966 (leaves narrower than type, 
GL). Florida, King 2142 (det. Gl.). Brazil. "Uifiguiray." 

Miconia expansa Gleason, Bull. Torrey Club 63: 536. 1936. 

Nearly glabrous shrub 3 meters high, the upper branches exactly 
4-angled; petioles stout, the upper 2-3 cm. long; leaf blades ample, 
membranous, obovate-oblong or broadly elliptic, shortly and 
abruptly apiculate, entire but sparsely glandular-ciliate, broadly 
cuneate at base, 5-plinerved; panicle ample, glabrous unless obscurely 
scurfy at nodes, the branches all evidently dilated above and strongly 
4- winged; calyx campanulate, glabrous, 4 mm. long, the broadly 
ovate sepals 2 mm. long and nearly as broad, minutely glandular- 
setose at tip; petals fleshy, white, nearly 5 mm. long, 4 mm. wide; 
filaments sparsely glandular as connective near the base; style very 
stout, about 5.5 mm. long, the capitate stigma nearly 2 mm. broad; 
ovary inferior, 5-celled, costate apically. Section Amblyarrhena. 

Loreto: In forest at Pumayacu, 600-1,200 meters, Klug 3201, 
type. 

Miconia falcata Cogn. Bot. Jahrb. 42: 139. 1908. 

Apparently scarcely distinct from M. dipsacea; leaves arcuate- 
falcate, 7-nerved; panicle branches branched; stigma scarcely dilated, 
the filiform style 5-6 mm. long. Calyx 2 mm. long; 5-meter shrub, 
the branches very stout. Section Eumiconia. F.M. Neg. 17058. 

Cuzco: Santa Ana, Prov. Convention, 1,500 meters, Weberbauer 
4998. 

Miconia fallax DC. Prodr. 3: 181. 1828; 788. 

Very similar to M. argyrophylla ; leaves obtuse or shortly and 
obtusely acuminate; petals rounded or subretuse, glabrous, 3.5-4 



FLORA OF PERU 411 

mm. long; connective below the cells not or scarcely produced, 
minutely tuberculate posteriorly; style 7-9 mm. long; fruit broadly 
subglobose, 10-sulcate, 5-6 mm. thick. Section Eumiconia. 

Junin: San Ramon (Killip & Smith 24.775, "an abnormal form 
with longer leaves" Gleason). Paraguay to the Guianas. 

Miconia filamentosa Gleason, Bull. Torrey Club 65: 579. 1938. 

Upper branches terete, slender, finely stellate-puberulent, as 
the petioles, leaves beneath on the veins and the panicles, including 
the calyces; leaves oblong-lanceolate, narrowed to the minutely 
cordate base, often conspicuously crenate, narrowly acuminate, 
3-nerved or nearly 3-plinerved, glabrous above, to 12 cm. long, 3 cm. 
broad; panicle 6-8 cm. long and broad, the 5-merous flowers sessile 
in 3-flowered cymes but the lateral flowers apparently pedicellate; 
calyx about 2 mm. long, the rounded or acute sepals 0.5 mm. long; 
petals scarcely retuse, glabrous, 3 mm. long; stamens dimorphic, 
the slender filaments 4.6 or 2.3 mm. long; connective of larger 
stamens nearly enclosing the filaments by its extended oblique 
rounded lobes, of the smaller, produced with one dorsal and two 
lateral nodes; ovary 4-celled; style slender, glabrous, 5.5 mm. long, 
the stigma minutely capitate. Probably nearest M. Sprucei Triana. 
Section Eumiconia. 

Loreto: Mishuyacu, near Iquitos, Klug 938, type. Rio Itaya, 
Williams 3306, in fruit, the leaves less crenate, the panicle to 2 dm. 
long. Rio Mazan, Schunke 160; Rio Itaya, Williams 3298. Brazil. 

Miconia firma Macbr., spec. nov. 

Ramis robustis superne acute 4-angulatis et sulcatis demum 
solum quadratis glabris; petiolis supra sulcatis minute hirsutis 
apice tuberculo-dilatatis ad 4 cm. longis; foliis ovato-ellipticis, basi 
rotundatis, breviter acuminatis, integris, 5-nerviis, supra planis, 
sparse scabro-setulosis ut videtur glabratis, subtus dense fulvo- 
hirsutis, ad 23 cm. longis fere 10 cm. latis rigide coriaceis; paniculis 
glabratis vel sparse minuteque hispidulis, dense multifloris, 14 cm. 
longis, ad 5.5 cm. latis; floribus 4-meris; pedicellis circa 1 mm. longis; 
calyce campanulato, glabro, obscure lobato, circa 1.5 cm. longo; 
petalis late oblongo-ellipticis, glabris, obtusis fere 2.5 mm. longis, 
staminibus glabris; filamentis 1 mm. longis, antheris fere oblongis 
vel vix obovoideis, 1.5 mm. longis, poris 2 dehiscentibus, connectivo 
simplice; stylo 2.5 mm. longo, stigmate subpeltato; ovario libero valde 
costato. Ovary completely free and strongly ribbed as in few species. 

Cajamarca: Weberbauer 6389, type. 



412 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia flaccida Gleason, Bull. Torrey Club 58: 226. 1931. 

Branches and panicle-branches acutely 4-angled and narrowly 
winged, glabrous as the entire plant unless finely scurfy at the nodes 
and on the calyx and petals without; petioles slender, 2-3 cm. long; 
leaves thin, pale-green, narrowly elliptic, 2.5-3.5 dm. long, 6-7 cm. 
wide, long-cuneate at base, long-acuminate, entire, 3-plinerved; 
panicle ample, the flowers sessile or subsessile, 5-merous, calyx 
tubular-campanulate, about 3.5 mm. long, the broadly triangular 
sepals acute; petals 4 mm. long, strongly inequilateral, deeply retuse; 
stamens about 15, isomorphic; filaments glabrous, 4 mm. long; 
anthers stoutly subulate, nearly straight, the connective not produced 
or appendaged; ovary 5-celled, glabrous as the style, this slender, 
7 mm. long, the stigma capitate. Shrub 2-3 meters high, the winged 
stems hollow; the stamens suggest those of Tococa (Gleason). Sec- 
tion Tamonea. 

Junin: Pichis Trail, 1,600 meters, Kittip & Smith 25625. 

Miconia flaviflora Gleason, Bull. Torrey Club 63: 534. 1936. 

Allied to M. acuminifera, but the leaves oblong-ovate, 10-15 (30) 
cm. long, 3.5-5.5 (9) cm. broad; irregular calyx lobes about 0.5 mm. 
long; petals narrowly oblong-obovate, obtuse, 3.5 mm. long, 1.2 mm. 
broad, reflexed at anthesis, minutely puberulent without; filaments 
glabrous, the anthers nearly isomorphic, the connective only of the 
smaller minutely 3-lobed basally. Early collections in bud were 
referred to M. acuminifera, but flowering material shows that the 
petals are only half as long as in that species, the calyx much more 
deeply lobed (Gleason). Suggests M. cremophylla which may not 
be correctly placed. Section Tamonea. 

San Martin: Near Moyobamba, Klug 3697, type; (Williams 
74.21, det. Gl.). Junin: Near La Merced, Kittip & Smith 2^000; 
25575; 25997 (det. Gl.). Costa Rica. 

Miconia floccosa Cogn. Bot. Jahrb. 42: 140. 1908. 

Branches obtusely quadrate, densely clothed, as the leaves 
beneath, with a felt-like stellate tomentum; petioles 0.5-1.5 cm. 
long; leaves rigid, thick coriaceous, oblong-lanceolate, shortly 
narrowed at base and apex, soon glabrate above, 3-nerved, but the 
marginal pair nearly concealed in the tomentum, 7-11 cm. long, 2-3 
cm. wide; panicle 6-10 cm. long, the sessile aggregate calyces 4-5 mm. 
long; petals rigid, obtuse, 5 mm. long, glabrous; style glabrate, 4-5 
mm. long, the stigma little enlarged. Allied to M. lanata Triana. 
Section Eumiconia. F.M. Neg. 17059. 



FLORA OF PERU 413 

Junin: West of Huacapistana, 3,200 meters, Weberbauer 2200, 
type. 

Miconia fruticulosa Cogn. Bot. Jahrb. 42: 146. 1908. 

Completely glabrous shrub 0.5 meters high, the slender upper 
branches acutely tetragonous; petioles 4-10 mm. long; leaves coria- 
ceous, spinulose-serrate, ovate-cordate, shortly acuminate, 3-nerved, 
2.5-4.5 cm. long, 13-26 mm. broad; panicles 2-5 cm. long, the 
4-merous flowers on pedicels 3-5 mm. long; calyx tube 4 mm. long, 
the obtuse lobes 1.5 mm. long, denticulate below the tip; petals 
yellowish white, thick, rounded and shortly apiculate, 5 mm. long; 
filaments strongly compressed, 2.5 mm. long; style glabrous, filiform, 
5-6 mm. long, the stigma capitate. Allied to M. epiphytica Cogn. 
Section Chaenopleura. F.M. Neg. 17064. 

Huanuco: Above Monzon, 3,400 meters, Weberbauer 3383, type. 

Miconia galactantha Naud. Ann. Sci. Nat. ser. 3. 16: 245. 1851; 
918. M. serrulata [Pavon] Triana, Trans. Linn. Soc. 28: 129. 1871, 
not Naud. Melastoma serrulata Pav. ex D. Don, Mem. Wern. Soc. 4: 
314. 1823, in syn. Cremanium serrulatum D. Don, I.e. 

Near to M. clathrantha but early scurfy, especially on the nerves 
beneath, and the closely net-veined, minutely serrulate, oblong- 
elliptic leaves much firmer, acute, 6-11 (13) cm. long, 1.5-3 (3.5) 
cm. broad, on petioles only 1 cm. long; calyx shortly and obtusely 
lobed, 1.5 mm. long; petals nearly 1 mm. long, retuse. Small flower- 
fascicles almost contiguous. Weberbauer 5585, referred by Cogn. to 
M. polyneura, is rather this species according to Gleason in herb., 
but the flowers, even though young, seem to be larger, and mostly 
pedicellate. Section Cremanium. F.M. Neg. 17165. 

Huanuco: Chinchao & Cuchero, Ruiz & Pavdn, type. Yanano, 
3744; 4924(1) (in fruit). Junin: Chanchamayo Valley, Schunke 169. 

Miconia glaberrima (Schlecht.) Naud. Ann. Sci. Nat. se'r. 3. 
16:243. 1851; 916. 

Completely or essentially glabrous slenderly branched shrub, the 
upper branches obtusely tetragonous; petioles to 3.5 cm. long; leaves 
rather firm, ovate-oblong, shortly acuminate, obtuse or acutish at 
base, entire or nearly, about 1 dm. long, 3-5 cm. broad, 3-nerved; 
panicle often much less than 1 dm. long; calyx subhemispheric, 1.5 
mm. long, minutely toothed; petals 1 mm. long; style 1-3 mm. long; 
fruit white. The Peruvian plant is var. australis Macbr. Field Mus. 
Bot. 4: 184. 1929, the leaves abruptly obtuse at base or even sub- 
emarginate, panicles 3-4 cm. long, sparsely and extremely minutely 



414 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

scurf y-puberulent; calyx teeth little obvious; is perhaps M. theaezans, 
but the anthers are 2-pored. Section Cremanium. F.M. Neg. 17067. 
Junin: Hacienda Schunke, La Merced, open shrub 1-2 meters 
high, 5769, type (var.); 5771. Mexico and Central America. 

Miconia glaucescens Triana, Trans. Linn. Soc. 28: 115. 
1851; 830. 

Upper branches slightly compressed, puberulent; leaves obovate- 
lanceolate, decurrent into the 3-6 mm. long petiole, caudate-acumi- 
nate, green above, pale and puberulent beneath with slender appressed 
trichomes on the nerves, these 3, with a very faint marginal additional 
pair, 10-17 cm. long, 3-6 cm. broad; panicle narrow, lax, to 1.5 dm. 
long, the sessile flowers ternate; calyx oblong or subcampanulate, 
acutely dentate, 2.5-3 mm. long, glabrate; petals 1.5 mm. long, 
obtuse; anthers subulate, unappendaged; style 4-6 mm. long. The 
mature leaves are only and minutely brown-lepidote-pubescent 
beneath, a little plinerved by reason of the decurrent base. Section 
Eumiconia. F.M. Neg. 17068. 

Puno: San Gaban, Lechler 2402, type. 

Miconia glandulifera Cogn. Melast. 737. 1891. 

Resembles M. ciliaris Triana but petioles 2-3 cm. long, leaves 
5-nerved and glabrous above, calyx rather more densely scurfy- 
tomentose, about 3 mm. long, the lobes somewhat irregular; petals 
scurfy-puberulent without, anthers glandular below. Cf. M. muri- 
cata, which is probably the earlier name. Section Tamonea. F.M. 
Neg. 25962. 

Huanuco: Casapi, Mathews 1728, type. Junin: Pichis Trail, 
Killip & Smith 25451. Perene" Bridge, 25373 (det. Gl.). Loreto: 
Yurimaguas, Williams 4100 (det. Gl.). 

Miconia glomerata Triana, Trans. Linn. Soc. 28: 110. 1871; 776. 

Upper branches densely stellate- tomentose; petioles stout, 1-3 
cm. long; leaves oblong-lanceolate, cordate, gradually acute, gla- 
brous and light green above, as also beneath in age, or there scurfy 
on the nerves and drying brown, 3-nerved with an additional very 
faint marginal pair, 1-3 dm. long, 7 cm. wide; flowers densely capi- 
tate-glomerate at the ends of the short simple panicle branches, the 
glomerule often occupying the entire branch; fruiting calyx 3 mm. 
long, the lobes broadly rounded, two-thirds mm. long. Cf. M. 
Martiniana and note. F.M. Neg. 29501. 

Huanuco: Chinchao, Ruiz & Pavon, type. San Anton, Nee. 



FLORA OF PERU 415 

Miconia glutinosa Cogn. Bot. Jahrb. 42: 141. 1908. 

Similar to M. alypifolia but the branches broadly 4-winged, the 
younger with the petioles, peduncles and calyces glutinous; leaves 
obtuse at base, densely appressed-tomentose beneath, 6-10 cm. long, 
3-4.5 cm. broad; calyx limb subtruncate. Section Amblyarrhena. 
F.M. Neg. 17070. 

Puno: Between Sandia and Chunchusmayo, 2,400 meters, Weber- 
bauer 1325, type. 

Miconia grandifolia Ule, Notizbl. Bot. Gart. Berlin 6: 359. 1915. 

Glabrate shrub or tree, only the younger parts with the leaf 
nerves beneath and the ample panicles obscurely and most minutely 
stellate-pulverulent; leaves subsessile, firm or subcoriaceous, even to 
4 dm. long and half as wide, elliptic or ovate-elliptic, shortly cuspi- 
date-acuminate, amplexicaul at base, entire or repandly margined, 
shortly plinerved, 3 nerves prominent with 2 fainter submarginal 
ones; flowers sessile, 5-merous, calyx tube oblong-urceolate, the tri- 
angular teeth acute, the former 3 mm., the latter 0.5 mm. long; 
petals white, 3 mm. long, 2 mm. broad; anthers nearly equal, linear, 
the connective anteriorly tuberculate-lobed; style thick, 3.5 mm. 
long, the stigma capitate. Apparently nearly M. pandurata. Sec- 
tion Eumiconia. 

Loreto: Leticia, Ule 6207, type. Brazil. 

Miconia granulosa (Bonpl.) Naud. Ann. Sci. Nat. se"r. 3. 16: 
218. 1851; 910. Melastoma granulosa Bonpl. Melast. 25. pi. 12. 1816. 

Branches stout, obtusely tetragonous, young panicles and petioles 
granular puberulent; petioles 1-3 cm. long; leaves oblong-ovate, 3- 
nerved, rounded at base, acuminate, glabrous above, more or less 
densely granular-tomentose beneath with bright brown trichomes, 
10-15 cm. long, 3-5 cm. wide, thick-coriaceous; flowers sessile, con- 
gested, calyx scurfy, shortly toothed, 2.5 mm. long; petals white or 
roseate, 1 mm. long; style 4 mm. long, the stigma obtuse. The leaf- 
pubescence is soon partly deciduous. Section Cremanium. F.M. 
Neg. 36260 (in part). 

Peru (probably). Bolivia; Colombia. 

Miconia Grayana Cogn. Melast. 896. 1891. 

Glabrous, with slender, elongate, subterete branches that are 
supplied at the nodes with membranous sleeve-like stipules; petioles 
1-2 cm. long; leaves rather firm, remotely subulate-denticulate, 
lanceolate, shortly acuminate, acute at base, 3-plinerved, the nerves 



416 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

deeply impressed above, 5-9 cm. long, 1.5-3 cm. wide; panicles 4-6 
cm. long, the flowers 6-merous on pedicels 2-4 mm. long; calyx cam- 
panulate, slightly constricted below the middle, narrowly and 
acutely but shortly 6-dentate, 4 mm. long; petals 2 mm. long; stamens 
12; style 6 mm. long, the stigma capitate. Section Amblyarrhena. 
Peru(?) : Grisar, type. 

Miconia Griffisii Macbr. Field Mus. Bot. 4: 188. 1929. 

Branches soon terete, densely setulose-hirtellous, with only 
simple trichomes; petioles 7-12 mm. long; leaves subcoriaceous, 
ovate-lanceolate, acute or shortly acuminate, rounded or subacute 
at base, glabrous above, sparsely hirtellous on the 3 nerves and 
veins beneath, mostly 8 cm. long, 3-3.5 cm. broad; panicles densely 
flowered, about 1 dm. long; flowers 4-merous, on pedicels 5 mm. 
long; calyx turbinate-campanulate, glabrate, 4 mm. long, distinctly 
lobed, the lobes subrotund, the outer teeth reduced to tubercles; 
petals white, suborbicular, about 3 mm. long; style 5-6 mm. long, 
the stigma peltate. There is an outer pair of marginal nerves. 
Shrub 1-1.5 meters high, named for C. N. Griffis, of Lima. M. 
pichinchensis Benth., collected by Ruiz & Pavon from Guayaquil, 
somewhat similar, has thin, subsessile leaves. Section Chaenopleura. 

Huanuco: Tambo de Vaca, 4382, type. 

Miconia grisea Cogn. Bot. Jahrb. 42: 147. 1908. 

Much-branched shrub related to M. chionophila Naud., the 
branches densely scurfy with minutely barbellate, rather long 
trichomes, the leaves slightly stellate-scurfy especially on the 3 
nerves beneath, 1.5-2 cm. long, 8-16 mm. broad; petioles 3-6 mm. 
long; panicle little if at all branched, the few to many flowers nodding 
on arcuate pedicels 2-4 mm. long; petals white, 3 mm. long; style 
glabrous, 4 mm. long, the stigma punctiform. There is an additional 
pair of submarginal nerves barely noticeable in the larger leaves. 
Section Chaenopleura. F.M. Neg. 17071. 

Huanuco: Near Monzon, 3,400 meters, Weberbauer 3346, type. 

Miconia guianensis (Aubl.) Cogn. Jahrb. Bot. Gart. Berlin 4: 
280. 1886; 741. Tamonea guianensis Aubl. PI. Guian. 2: 658. 1775. 

Branches densely scurfy-pulverulent, soon glabrate; petioles 
elongate; leaves subcoriaceous, oblong-ovate, acuminate, glabrous 
above, usually rather thinly stellate-tomentulose beneath, the tri- 
chomes minute, 3-5 (-7)-nerved; inflorescence ample, the fascicled 
short branches that simulate pedicels flattened or quadrate, the 



FLORA OF PERU 417 

sessile flowers subtended by 2 obovate bracts 6-11 mm. long, con- 
spicuous only in bud; petals 6-8 mm. long; calyx finally glabrous; 
filaments glabrous; connective distinctly elevated below and minutely 
bicalcarate at the prolonged base. The Poeppig specimen not 
studied by me; the other Peruvian specimens, broader leaved in 
bud, are doubtful. Section Tamonea. 

Loreto: Prov. Mainas, Poeppig. Yurimaguas, Williams 4036; 
4458; 4693. Bolivia to the West Indies. 

Miconia hamata Cogn. Bot. Jahrb. 42: 141. 1908. 

Branches obscurely quadrate, the younger densely clothed with 
subappressed, mostly plumose trichomes; petioles 5-15 mm. long; 
leaves subcoriaceous, narrowly ovate, rounded at base, shortly 
acuminate, 7-plinerved, strigose above with spreading hook-setae, 
these definitely thickened at base, densely long-villous beneath, 
6-8 cm. long, 2.5-4 cm. broad; flowers 5-merous, sessile, in very 
small heads; calyx densely villous, minutely and broadly lobed, 2 mm. 
long; petals white, 1.5 mm. long; filaments capillary, glabrous as 
the style, both about 3 mm. long. Allied to M. capitellata Cogn., 
892, of Ecuador, with broader 5-plinerved leaves and deep yellow 
petals 2 mm. long. Section Amblyarrhena. F.M. Neg. 17074. 

Amazonas: Near Tambo Bagazan, 2,200 meters, Weberbauer 
4447, type. 

Miconia Herrerae Gleason, Amer. Jour. Bot. 19: 745. 1932. 

Flowering branches stout, simple, slightly compressed, thinly 
stellate-tomentulose toward the tips; petioles stout, 15-20 mm. 
long; leaves subcoriaceous, elliptic, acute at both ends, entire, 3- 
nerved, with an additional pair of submarginal veins, to 18 cm. long 
and half as wide, soon glabrous above, loosely but permanently 
stellate-pubescent beneath, the veins reticulately raised; panicle 
ample, its branches sparsely stellate-tomentulose; flowers 5-8- 
usually 6-merous, sessile in small glomerules; calyx lightly stellate- 
puberulent, about 3 mm. long, the membranous sepals rotund, 
0.3 mm. long; petals nearly 2.5 mm. long, glabrous, retuse; stamens 
about 20, the filaments straight or geniculate below the apex; 
connective produced into 4 minute lobes; style glabrous, 5 mm. 
long, the stigma capitate. Cf . M. Pavoniana. Section Glossocentrum. 

Cuzco: Cedrobamba, Urubamba Valley, 2,200 meters, Herrera 
1575, type; 3194; 3221. 

Miconia heteromera Naud. Ann. Sci. Nat. se"r. 3. 16: 124. 
1851; 812. 



418 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Typically upper branches and younger parts granulose-sciirfy 
and densely brown-hirsutulous with spreading trichomes, rarely 
glabrate; petioles 0.5-1 cm. long; leaves rather strongly unequal, 
5-6 (-10) cm. long, 2-2.5 (-3) cm. broad, oblong-lanceolate, long- 
acuminate, sparsely setulose on both sides, 3-nerved; panicles 
racemiform, few-flowered, only 2-3 cm. long, the shortly pedicellate 
flowers usually 6-merous; calyx 2.5-3 mm. long; petals obtuse, 4 mm. 
long; style little enlarged apically, 6 mm. long. The setose trichomes 
typically are more persistent than the stellate. But Gleason refers 
here Williams 2688; 2985; 3011 from Loreto, glabrous except 
"thinly furfuraceous on petioles and stems." Section Eumiconia. 
F.M. Neg. 17076. 

San Martin: Tocache, Poeppig 1911, type. Loreto: La Victoria, 
Williams 2985; 3011; 2688. 

Miconia hygrophila Naud. Ann. Sci. Nat. se>. 3. 16: 220. 
1851; 905. 

Branches obscurely tetragonous, the upper, as the petioles and 
panicles, scurfy tomentose; petioles 1-3 cm. long, stout; leaves oblong- 
elliptic, acute, the entire margin revolute, coriaceous, reddish and 
nearly glabrous to densely hirtellous with plumose trichomes beneath, 
especially on the 3 principal nerves; panicles ample, much branched, 
the ebracteate flowers sessile; calyx shortly 5-dentate; petals retuse, 
white, 1 mm. long; stigma minutely capitate. Section Cremanium. 

Ayacucho: Ccarrapa, Killip & Smith 22457.- Junin: Carpapata, 
Killip & Smith 244-71 (det. Gl.). Cuzco: Rio Chaupimayo, Soukup 
590. 

Miconia ibaguensis (Bonpl.) Triana, Trans. Linn. Soc. 28: 110. 
1871; 815. Melastoma ibaguensis Bonpl. Melast. 105. pi. 45. 1816. 

Upper branches often minutely stellate as well as densely hirsute- 
setulose with brown, somewhat spreading trichomes; petioles 0.5- 
1 cm. long; leaves oblong-ovate-lanceolate, often minutely serrulate 
and ciliate, setulose-hirsute beneath, sometimes also more or less 
above, shortly 5-plinerved (sometimes 5-nerved, 3 principal nerves), 
6-12 cm. long, (2) 2.5-5 cm. broad; calyx minutely stellate-villous 
and somewhat setulose, 5-lobed, 3 mm. long; petals 2.5-3 mm. long, 
white, yellowish or rarely roseate; connective simple or minutely 
tuberculate, anteriorly biauriculate; style 5-7 mm. long, the stigma 
somewhat infundibuliform. The pubescence even on the leaves is 
loose or spreading. Var. glabrata (Steud.) Cogn. is the form with 



hirsute branches, the leaves usually entire, lustrous and glabrous or 
glabrate above. Section Eumiconia. 

Junin: La Merced, 5488; 5534. Chanchamayo Valley, Schunke 
347; 357; 1543. San Ramon, Kittip & Smith 24781 . Huanuco : 
Monzon, Weberbauer 3474 (det. Cogn.). Pozuzo, 4568. Pampayacu, 
5020. Cuchero and Casapi, Poeppig 4; 98; 1403. San Martin: 
Moyobamba, (Fielding, the var.); Mathews 1294; 1717; 3215; 
(Fielding 1294)- Tarapoto, Ule 6457 (det. Pilger); Poeppig; Spruce 
4233; Williams 5480; 5630; 6151; 6428; 6433. San Roque, Williams 
7530. Brazil to Mexico and the West Indies. "Mullaca." 

Miconia icosandra Gleason, Bull. Torrey Club 58: 239. 1931. 

Small, completely glabrous (essentially) tree with stout, obtusely 
4-angled branches and membranous, narrowly oblong, acuminate, 
entire, shortly 5-plinerved leaves; petioles slender, 1.5-2.5 cm. long; 
leaves to 16 cm. long, a third as broad, acute at base, obscurely pul- 
verulent beneath; panicles to 1.5 dm. long, the 5-merous flowers 
crowded in small cymules, the lateral subsessile; calyx glabrous, 
about 2.5 mm. long, the broadly triangular acute sepals 0.4 mm. 
long; petals 3 mm. long, obliquely retuse, minutely pulverulent on 
both sides; stamens about 20; filaments filiform; connective shortly 
produced into an oblong dorsal lobe and 2 broadly rounded lateral 
lobes; ovary 4-celled, glabrous, the placentae nearly basal, extending 
radially into the cells and adnate to the bottom of the ovary, with 
10-13 ovules in each cell; style 6 mm. long, the stigma capitate. 
There are usually some sterile filaments; the arrangement of the 
placentae is most unusual, although there are species in which the 
ovules are reduced to one in each cell at base (Gleason). Section 
Glossocentrum. 

Junin: Enenas, Pichis Trail, 1,600 meters, Killip & Smith 25781, 
type. 

Miconia impetiolaris (Sw.) D. Don, Mem. Wern. Soc. 4: 316. 
1823; 775. Melastoma impetiolaris Sw. Prodr. 70. 1788. 

Allied to M. amplexicaulis but the leaves elliptic, somewhat pli- 
nerved, usually undulate-toothed, sometimes entire, subamplexicaul 
and auriculate at base; panicle branches long, spreading (when well- 
developed), interruptedly spike-like; calyx campanulate, shortly 
lobed, 2.5-3 mm. long, typically stellate, in Peru often glabrate; 
petals white, retuse, 2-3 mm. long; connective prolonged, recurved 
around filament, at the end obscurely 3-lobed or truncate (Gleason) ; 
style 4 mm. long. M. grandifolia is nearly this but that, at least as 



420 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to type, is merely pulverulent even to the narrow inflorescence. A 
shrub or small tree. Section Eumiconia. 

Huanuco: Near confluence of the Rio Huallaga and the Rio 
Cayumba, 794 meters, Mexia 8268 (det. Gl.). Bolivia to Mexico 
and the West Indies. 

Miconia inamoena Pilger, Verh. Bot. Ver. Brandenb. 47: 

171. 1905. 

Branches and petioles densely hirsute-setulose with reddish tri- 
chomes, these softer and more crisped on the leaves beneath, shorter 
and fewer on the leaves above; petioles stout, elongate; leaves to 2 
dm. long, half as broad, 7-nerved, membranous, elliptic, sometimes 
slightly cordate at base, shortly narrowed and abruptly short acu- 
minate at the tips; flowers crowded, sessile, few; calyx tube broadly 
campanulate, about 2 mm. long, the 5 subulate lobes acute; petals 
roseate, broadly elliptic, irregularly retuse; stamens 10, the thick, 
broad filaments slightly pilose, 2 mm. long, the longer style with 
capitate stigma. Allied by author to M. obscura (Bonpl.) Naud. 
Section Amblyarrhena. F.M. Neg. 17080. 

Loreto: Cerro de Escaler, 1,200 meters, Ule 6735, type. 

Miconia integrifolia Cogn. Melast. 936. 1891. 

Branches obscurely quadrate, somewhat scurfy (and with a few 
scattered setae) puberulent as the petioles and thyrsoid panicles; 
petioles 0.5-1 cm. long; leaves coriaceous, narrowly ovate, minutely 
cordate at base, acute, entire, obscurely 3 (2 obscure outer nerves) 
-5-plinerved, glabrous and lustrous above, faintly hirtellous in 
nerve axils beneath, 5-8 cm. long, 2.5-3.5 cm. broad; panicles as 
long, the flowers congested, 4-merous, on pedicels 1-3 mm. long; 
calyx subhemispheric, 2.5 mm. long; petals broadly subrotund, 1.5-2 
mm. long; style sparsely hirtellous, 3 mm. long, the stigma capitate. 
Cf . M. laurina, to which, as to Ruiz & Pavon specimen from Vitoc, 
this description applies except that the anthers are seemingly those 
of Section Cremanium; however, the pores are soon open. Section 
Chaenopleura. 

San Martin: Valle de Vitoc, Isern 2343. Without locality, Ruiz 
& Pavon (Herb. Boissier). 

Miconia juruensis Pilger, Verh. Bot. Ver. Brandenb. 47: 

172. 1905. 

Tree, essentially glabrous, the pubescence where present only a 
very sparse pulverulence; petioles 1-2 cm. long; leaves membranous 



FLORA OF PERU 421 

or firm, elliptic, more or less rounded at base, shortly acuminate, 
mostly 10-12 cm. long, 3-5 cm. broad, 3-plinerved, the outer addi- 
tional nerves faint, typically opaque on both sides; flowers subsessile, 
the panicles about 1 dm. long; calyx tube 3.5-4 mm. long, campanu- 
late-cylindrical, the dilated limb scarcely denticulate; petals white, 
5, irregularly elliptic and emarginate apically, 3.5-4 mm. long; sta- 
mens 10, connective somewhat stipitate-glandular; style finally 7 
mm. long, the stigma minutely capitate. Section Eumiconia. F.M. 
Neg. 17084. 

Loreto: Mouth of Santiago, Tessmann 4218 (det. Markgr.). 
Lower Rio Nanay, Williams 481 (det. Gl.). Yurimaguas, Williams 
4230; 5186. Brazil; Bolivia. "Caracha caspi." 

Miconia Klugii Gleason, Bull. Torrey Club 58: 240. 1931. 

Branches stout, roundly 4-angled, very closely and minutely 
lepidote as the stout, angled petioles, these 2-6 cm. long; leaves firm, 
elliptic, acute at base, abruptly acuminate, 3-plinerved, glabrous 
and lustrous above, densely covered beneath with a thick brown 
lepidote indument, sometimes 2 dm. long or longer by half as wide; 
veins strongly impressed above; inflorescence freely branched, to 2 
dm. long, closely but thinly lepidote; flowers sessile, 5-merous; 
calyx campanulate, nearly 2 mm. long, its walls fleshy; sepals minute 
points; petals cuneate-obovate, 1.8 mm. long, 1 mm. wide, obliquely 
truncate or slightly retuse; connective lobed basally; ovary 3-celled, 
glabrous, the style 2.5 mm. long, gradually thickened to the trun- 
cate stigma. Section Glossocentrum. 

Loreto: Iquitos, 10-meter tree, Klug 1141, type. 

Miconia lacera (Bonpl.) Naud. Ann. Sci. Nat. se>. 3. 16: 152. 
1851 ; 866. Melastoma lacera Bonpl. Melast. 9. pi 5. 1816. 

Slender, branched shrub well marked by the long, often bright 
red pilosity of the upper stems and the leaf -margins, the trichomes 
even to 5 mm. long; petioles 1-2 cm. long; leaves thin, 7-15 cm. 
long, half as wide, 3-nerved, sparsely setose on both sides or some- 
what villous beneath, the trichomes, especially above, usually scat- 
tered, in general oblong-lanceolate, acuminate, minutely denticulate; 
panicles 5-10 cm. long, the long ciliate bractlets 1-3 mm. long; 
calyx tube 2 mm. long, the interior very long laciniate-setose lobes 
little shorter; petals rose, 3-4 mm. long; style 3 mm. long. Section 
Amblyarrhena. 

Loreto: Florida, 1 meter high, the flowers white, Klug 2230 (det. 
Gl.). Brazil to Mexico and the West Indies. "Chosero-ey" (Huitoto) . 



422 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia lasiocalyx Cogn. Bull. Torrey Club 23: 278. 1896. 

Obtusely tetragonous branches and petioles shortly and densely 
hirtellous-puberulent or hirtellous-scurfy; petioles 2-3.5 cm. long; 
leaves membranous, obscurely denticulate, ovate-oblong-elliptic, 
acuminate, rounded at base, very shortly and appressed-setulose 
above, minutely hirtellous beneath, shortly 5-7-plinerved, 1.5-2 dm. 
long, 6-8 cm. wide; flowers 5-merous, sessile, rather crowded, the 
pyramidal panicle with divaricate branches; calyx almost shaggy- 
hirsute, the tube 3 mm. long; petals broadly obovate, subtruncate, 
1 mm. long; anthers 2-pored apically; style 3 mm. long, the stigma 
peltate. Aff. M. costaricensis and the anthers 2-pored according to 
the author; nevertheless referred by him to Section Amblyarrhena, 
with "anthers minutely 1-pored." 

Cuzco: Near Rio Yanamayo, Pennell 14053. Bolivia. 

Miconia lasiostyla Gleason, Bull. Torrey Club 58: 243. 1931. 

Young branches obtusely tetragonous and lightly sulcate, densely 
clothed with branched brown trichomes; petioles stout, 1-2 cm. long; 
leaves membranous, obovate-oblong, entire, acute, narrowed from 
above the middle to an obtuse base, 5-7-plinerved, glabrous above 
except shortly stellate-pubescent on the primaries, densely brown- 
stellate-pubescent beneath, to 2 dm. long, 7.5 cm. wide; panicles 
ample, the trichomes densely stellate-tomentose on the glomerate, 
subsessile, globose-campanulate calyces, these nearly 5 mm. long, 
sessile and much-branched; sepals broadly rotund, glabrate, the outer 
teeth stout, conic; petals subrotund, 2.5 mm. long, somewhat retuse; 
filaments glandular, below flattened, above terete; anthers oblong, 
obtuse, the connective not produced, simple; ovary 4-celled, many- 
ovuled, apically glandular; style densely glandular, about 5 mm. 
long, the stigma peltate. Differs in pubescence, closely glomerulate 
flowers and shape of leaves from M. majalis Cogn. and M . floribunda 
(Bonpl.) DC. (Gleason). Section Amblyarrhena. 

Huanuco: Tree 8 meters high, Pan de Azucar, Sawada 77. 

Miconia latifolia (D. Don) Naud. Ann. Sci. Nat. se>. 3. 16: 
244. 1851; 921. Cremanium latifolium D. Don, Mem. Wern. Soc. 
4: 312. 1823. 

Upper branches obtusely quadrate, glabrate, obscurely pilose; 
leaves serrulate, cordate-ovate, acute, 7-9 (-10) cm. long, 3-5.5 cm. 
wide, smooth and lustrous above, pilose-setulose on the nerves and 
the veins beneath; inflorescence glabrous, the many flowers 4-merous, 



FLORA OF PERU 423 

white, small, on pedicels 1.5 mm. long. Inadequately known; cf. 
M. andina. Section Cremanium. F.M. Negs. 17091; 29502. 

Junin: Vitoc, Ruiz & Pavon, type. 

Miconia laurina (D. Don) Naud. Ann. Sci. Nat. se>. 3. 16: 245. 
1851 (name) ; 899. Cremanium laurinum D. Don, Mem. Wern. Soc. 
4: 313. 1823. 

Branches pilose; leaves lanceolate, obtuse, glabrous above, pilose 
beneath, 3-nerved; panicle racemose; flowers 5- or 6-merous, pedicel- 
late, nodding, small, white. No specimen so named has been found 
in the Ruiz and Pavon collections, but a plant from Vitoc (Ruiz & 
Pavon) may belong here; however, it could as well be referred, as 
to foliage at least, to M. integrifolia. Section Cremanium. 

Huanuco: Pampayacu, Sawada P. 16(1). Without locality 
(Pavdn). 

Miconia Lechleri Triana, Trans. Linn. Soc. 28: 124. 1871; 891. 

Branchlets and leaves beneath densely stellate-tomentose, the 
latter ovate-cordate, long-subcaudate-acuminate, shortly strigose 
above, 9-nerved, lightly foveolate beneath, 1-1.5 dm. long, 7-10 cm. 
wide; petioles setose-barbate above, 2-3 cm. long; panicle lax, 
1-1.5 dm. long, glabrous as the sessile calyces, these acutely dentate, 
2.5-3 mm. long; petals triangular-oblong, 2 mm. long; connective 
simple; style 4 mm. long; fruit 3-4 mm. thick. Allied by Cogniaux 
to M. Ruizii, now fide Gleason in Section Eumiconia, a similar 
Clidemia-like shrub. 

Puno: Tatanara, Lechler 2562, type. 

Miconia lepidota [Schrank & Mart.] DC. Prodr. 3: 180. 1828; 
788. Melastoma lepidota Schrank & Mart, ex DC. I.e., in syn. 

A shrub with conspicuous foliage, the leaves dark green above, 
rusty-cinereous beneath with a dense, compact, stellate-lepidote 
indument, the brownish silvery scales persistently concealing the 
leaf surface; upper branches compressed, becoming glabrate; petioles 
1-2.5 cm. long; leaves ovate-oblong, shortly acuminate, firm, 3- 
nerved with an additional fainter pair, 8-20 cm. long, half as wide; 
flowers borne secundly, the panicle much branched; calyx 10-costate, 
2.5 mm. long; petals 2 mm. long, retuse; connective anteriorly bicalca- 
rate; style 5 mm. long, the stigma somewhat peltate; fruit black, 
2 mm. thick. The scarcely significant var. grandifolia Cogn. has 
leaves 10-25 cm. long, 8-13 cm. wide, the petioles 2-4.5 cm. long. 
Section Eumiconia. F.M. Neg. 17094. 



424 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Yurimaguas, Poeppig 2458; Klug (the var.); Killip & 
Smith 29954 (det. Gl.). Bolivia to Guiana. 

Miconia lilacina Triana, Trans. Linn. Soc. 28: 129. 1871; 914. 

Glabrous, the slender branches obtusely 4-sided ; leaves coriaceous, 
sessile, pale and opaque above, drying darker beneath, ovate- 
lanceolate, cordate and somewhat amplexicaul at base, long-acumi- 
nate, 5-nerved, 7-10 cm. long, 3-4 cm. broad; panicle pyramidal, 
5-10 cm. long; pedicels 1-2 mm. long, articulate above the middle; 
calyx 2 mm. long, the coriaceous limb shortly and acutely lobed; 
petals lilac, obtuse, less than 1 mm. long; connective perfectly 
smooth; ovary glabrous; style 2-3 mm. long, the blackish stigma 
capitate; fruit blue, 6 mm. thick. Section Cremanium. 

Cuzco: Pearce. Without locality (Haenke). 

Miconia livida Triana, Trans. Linn. Soc. 28: 121. 1871; 876. 

Completely glabrous with terete, very slender branches and 
narrowly lanceolate, long-caudate, acuminate, 5-plinerved, mem- 
branous leaves, their margins lightly undulate and minutely setulose- 
denticulate, usually about 10 cm. long, 1.5-2.5 cm. broad; petioles 
1 cm. long; panicle many-flowered, 6-10 cm. long; pedicels 2-3 mm. 
long; calyx ovoid, distinctly and acutely lobed, 2 mm. long; petals 5, 
broadly obovate, 1.5 mm. long; connective simple; style filiform, 
glabrous, scarcely enlarged apically, 4-5 mm. long. Leaves with 
3 nerves above the base and 2 faint marginal basal ones. Section 
Amblyarrhena. F.M. Neg. 29503. 

Huanuco: Chinchao, Ruiz & Pavdn. Without locality (Mathews 
1180}. Distrito Churubamba, 1,725 meters, Mexia 8238. Bolivia. 

Miconia longifolia (Aubl.) DC. Prodr. 3: 184. 1828; 848. 
Melastoma longifolia Aubl. PI. Guian. 1: 432. pi. 170. 1775. 

Shrub or small tree with acutely 4-angled young branches, gla- 
brous or glabrate throughout, the sparse or evanescent indument 
merely a stellate pulverulence; petioles slender, 5-15 mm. long; 
leaves usually in whorls of 3, shortly 3-plinerved, oblong-lanceolate, 
acuminate, acute at base, 8-18 cm. long, 2.5-6 cm. broad; panicles 
widely branched, the principal branches usually several at each node; 
flowers 5-merous, subsessile, fragrant; calyx broadly campanulate, 
about 2 mm. long, the sepals minute; petals about 1 mm. long, 
puberulent; connective prolonged into a broad, obtuse or truncate 
dorsal lobe; style terete, 3-4 mm. long, the stigma truncate or 
punctif orm . Section Glossocentrum. 



FLORA OF PERU 425 

Loreto: Pongo de Manseriche, Tessmann 4100 (det. Markgr.). 
Iquitos, King 2236 (det. Standl.). Rio Itaya, Williams 3402; 3280; 
3506. Florida, King 2236; 2142. Rio Huallaga, Williams 4912; 
4769; 5350. Fortaleza, near Yurimaguas, King 2825. Southern 
Brazil, the Andes and the West Indies. "Rifari," "echapa-ey" 
(Huitoto). 

Miconia longiracemosa Gleason, Bull. Torrey Club 58: 
232. 1931. 

Simulates closely in vegetative characters M. pteropoda but the 
vein areoles on the lower side of the leaves are nearly always 5-6- 
sided and only 0.3 mm. across and, more especially, the anthers are 
dimorphic, the connective of the larger prolonged 0.4 mm. at base 
into a single semi-circular flat appendage, that of the smaller pro- 
longed 0.2 mm. into a triangular dorsal and 2 broadly triangular 
deflexed lateral lobes. As Gleason suggests, the value of the anther 
connective as a character in classification here is not fully known. 
Cf . also M. prasina. Section Eumiconia. 

Loreto: Lower Rio Huallaga, Williams 4249, type; same region 
or Rio Itaya and Rio Nanay, Williams 115; 604; 3878; 4038; 4167; 
4349 (all det. Gl.). Mishuyacu, near Iquitos, King 339(1}. Brazil; 
Colombia. "Bucacuru caspi." 

Miconia longisepala Gleason, Bull. Torrey Club 63: 536. 1936. 

A small, glabrous tree, the upper branches obscurely 4-angled; 
petioles slender, 10-15 mm. long; leaves elliptic, abruptly and nar- 
rowly acuminate into a linear acumen 10-15 mm. long, clearly 
cuneate into the petiole, entire, 3-nerved, not counting the faint 
outer pair, firm, 6-8.5 cm. long, 2.5-3.5 cm. broad; pedicels slender, 
to 0.5 mm. long, but seemingly about 8 mm. long because nodose 
below the flowers; calyx subglobose, about 3.5 mm. long, the spread- 
ing or reflexed linear-oblong lobes acute, 3.2 mm. long, the outer 
linear terete teeth about 1 mm. long; petals reflexed, broadly oblong, 
inequilaterally retuse, 5.6 mm. long; filaments thick, flattened, 
densely glandular, the glands subsessile; connective simple; ovary 
5-celled, glabrous; style stout, 8 mm. long, glandular, the glands 
below the middle minutely stipitate; stigma peltate, 2-3 mm. broad. 
Sepals remarkably long, nearly equaling the tube. Section 
Amblyarrhena. 

Loreto: 5-meter tree in forest, 600-1,200 meters, Pumayacu, 
King 3233, type. 



426 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia loretensis Pilger, Verb. Bot. Ver. Brandenb. 47: 
172. 1905. 

Younger branches strongly flattened; leaves lanceolate or lanceo- 
late-elliptic, gradually narrowed to the obtuse point and slightly 
decurrent into the 5-10 mm. long petiole, glabrous or faintly scurfy 
on the veins, drying black above, rather pale yellow-green beneath, 
7-9 cm. long, about 2 cm. broad, shortly 3-plinerved with 2 partial 
marginal nerves; panicle dense, 7-8 cm. long, narrow, the branches 
subsimple; calyx subsessile, tomentulose, obconic-campanulate, 
denticulate apically, about 3 mm. long; petals obovate-elliptic, 
white, 3 mm. long; connective shortly subauriculately produced and 
denticulate; style 4 mm. long, the stigma minutely capitate; ovary 
3-celled. Allied to M. prasina. Section Eumiconia. F.M. Neg. 17100. 

Loreto: Cerro de Escaler, 1,400 meters, Ule 6734, type. 

Miconia lugubris Cogn. Bot. Jahrb. 42: 142. 1908. 

Branches slender, subterete, with petioles and peduncles shortly 
and densely sordidly stellate-pilose; petioles 6-12 mm. long; leaves 
oblong-elliptic, acutish base and apex, entire, 3-nerved, 4-6 cm. 
long, 1.5-2.5 cm. broad, minutely setulose above, the trichomes 
enlarged at base, minutely and densely foveolate beneath and shortly 
and sparsely setulose, the nerves and veins rather densely stellate- 
puberulent; panicle broad, many-flowered, 4-7 cm. long, the flowers 
crowded on the simple branches; calyces sessile, urceolate, 1.5 mm. 
long, scarcely scurfy, the acute minute teeth shortly pilose; petals 
white, scarcely 1 mm. long; filaments capillary as the style, this 
glabrous, 2 mm. long, the stigma minutely capitate. Allied to 
M. atrofusca and M. caerulea. M. caelata (Bonpl). DC., 906, as to 
type from Ecuador, is glabrous and bullate above, the calyx 2.5-3 
mm. long; cf. M. atrofusca. Section Cremanium. F.M. Neg. 17101. 

Huanuco: Southwest from Monzon, 2,500 meters, Weberbauer 
3394, type. 

Miconia Malatestae Macbr. Field Mus. Bot. 4: 181. 1929. 

A stout, nearly glabrous, little-branched shrub 3 meters high, 
the upper branches obtusely tetragonous and sulcate; petioles thick, 
1.5-2 cm. long; leaves fleshy-coriaceous, deep green and sublustrous 
above, drying yellowish, entire or obscurely and remotely ciliate- 
denticulate, oblong or ovate-elliptic, obtuse or shortly acute, some- 
what decurrent at base into the petiole, mostly about 2 dm. long and 
half as broad, 3-nerved, with a pair of faint, additional, marginal 
nerves; panicle broadly pyramidal, 1-2 dm. long; calyces glabrous, 



FLORA OF PERU 427 

subsessile, shortly and acutely lobed, about 4 mm. long; petals 
white, broadly obovate, 3 mm. long; stamens 4.5 mm. long; anthers 
linear, apparently 1-pored, nearly 3 mm. long, shortly bituberculate 
at base; connective produced about 1 mm.; style 5 mm. long, the 
stigma peltate; fruit purple, 6 mm. thick. Named for Senor Enrique 
Malatesta of Huanuco. Near M. obovalis Naud., 803, with obovate 
plinerved leaves with prominent transverse veins and smaller calyces. 
Section Eumiconia(l). 

Huanuco: Hacienda Villcabamba on Rio Chinchao, coarse 
stalked shrub, 5176, type. 

Miconia Martiniana Gleason, Bull. Torrey Club 58: 233. 1931. 

Upper branches bisulcate, very densely brown-scurf y-tomentose; 
petioles stout, 3-5 cm. long, similarly pubescent; leaves narrowly 
obovate-oblong, acuminate, narrowed from the middle to the sub- 
cordate base, more or less obscurely undulate-serrate, 5-plinerved, 
soon glabrous and lustrous above except for the tomentose mid- 
nerve, softly stellate-pubescent, especially on the veins, beneath, 
to 3 dm. long, 14 cm. wide; flowers 5-merous, sessile, in dense glom- 
erules on the few elongate, 4-sulcate, densely stellate-tomentose 
inflorescence branches; calyx campanulate, tomentose, about 3.5 
mm. long, the acute lobes minute; petals cuneate, inequilaterally 
retuse, stellate without, nearly 3 mm. long; stamens dimorphic, the 
filaments 3.5 or 3 mm. long; connective of larger anthers expanded 
at base into 2 conspicuous lateral lobes and 2 minute dorsal ones or 
of the smaller into very slender lateral lobes and 1 minute dorsal 
one; ovary half inferior, 3-celled, apically pubescent and prolonged 
into 5 erect flat lobes; style glabrous, 7 mm. long, the stigma capi- 
tate. Cf. M. glomerata; also M. Pentlandii Naud., 771, Bolivian, 
the oblong leaves about 16 by 4 cm., the petals 1.5-2 mm. long. 
Section Eumiconia. 

San Martin: San Roque, 1,350-1,500 meters, Williams 7212, 
type; 7529. 

Miconia Martiusiana DC. Prodr. 3: 186. 1828; 850. 

Nearly glabrous; branches obtusely quadrate; petioles 1-1.5 cm. 
long; leaves nearly oval or broadly elliptic, 8-12 cm. long, 4-8 cm. 
wide, acute, rounded or barely acute at base, chartaceous-coriaceous, 
lustrous above, very shortly 5-plinerved, only the 3 middle nerves 
prominent; panicle about 1 dm. long; calyces subsessile, 2 mm. long, 
the limb caducous, distinctly 5-lobed; petals 1.5-2 mm. long; style 



428 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

filiform, the stigma scarcely enlarged. Section Glossocentrum. F.M. 
Neg. 17016. 

Loreto: Mishuyacu, King 22; 1082; 1090 (all det. Gl.). Brazil. 

Miconia Matthaei Naud. Ann. Sci. Nat. se>. 3. 16: 176. 1851; 
812. 

Branches densely long-reddish-hirsute, the stiff trichomes in- 
curved; petioles 1-2 cm. long; leaves lanceolate or oblong-lanceolate, 
often shortly caudate-acuminate, similar, 1-2 dm. long, 4-6 cm. broad, 
glabrous above, rather densely and crisply reddish-hirsutulous 
beneath, especially on the 3 nerves, these sometimes arising shortly 
above the leaf base; panicle softly red-villous, 5-8 cm. long, the 
bractlets 3-5 mm. long, the sessile 5-merous flowers crowded; calyx 
distinctly 5-lobed, the tube turbinate; petals 8 mm. long, style 
elongate, the stigma subpeltate; connective scarcely produced, 
anteriorly bilobed-glandular; fruit black, 3 mm. thick. Section 
Eumiconia. F.M. Neg. 36274. 

San Martin: Tocache, Poeppig 1913. Moyobamba, Mathews 
1299, type; Weberbauer 4706 (distr. as M. stelligera) . Loreto: 
Masisea, Killip & Smith 26854 (det. Gl.). Amazonian Brazil; 
Bolivia; Trinidad; Central America. 

Miconia mazanana Macbr., spec. nov. 

Ramulis ad apicem, petiolis superioribus et pedunculis plus 
minusve dense setulosis praecipue ad nodos; petiolis solum 5-10 mm. 
longis; foliis ellipticis, basi subacutis, apice caudatis, breviter vel vix 
5-plinerviis; calyce leviter furfuraceo; petalis fere glabris; filamentis 
sparse et minutissime hirsutulis ut videtur haud glandulosis; con- 
nectivo basi glanduloso. Apparently much like M. Donaeana 
except for the setose pubescence, glabrate petals, eglandular fila- 
ments, in the last character similar to the otherwise different M. 
amazonica et al. A shrub 3 meters high, the stem 10 cm. in circum- 
ference; flower lilac (Schunke). 

Loreto: Rio Mazan, Jose M. Schunke 374, type. 

Miconia media (D. Don) Naud. Ann. Sci. Nat. se>. 3. 16: 244. 
1851; 915. Cremanium medium D. Don, Mem. Wern. Soc. 4: 313. 
1823. C. tinctorium DC. Prodr. 3: 193. 1828. M. tinctoria Naud. I.e. 
229. 

Young branches and petioles somewhat ciliate-pilose, the latter 
1-2 cm. long; leaves ovate-oblong, subrotund to acutish at base, 
obtusely acute, subentire or callous-serrulate, smooth, 3-nerved 
(with a fainter outer pair), 5-8 cm. long, 2-3 cm. wide; panicles 6-8 



FLORA OF PERU 429 

cm. long; pedicels 1-2 mm. long; calyx distinctly toothed, suburce- 
olate, 2.5 mm. long; petals subrotund, 1.5 mm. long; style 3 mm. 
long, the stigma peltate. Near M. crocea but the leaves are slender 
and longer and less definitely pubescent; the white flowers have an 
unpleasant odor. Section Cremanium. F.M. Neg. 17017. 

Huanuco: Mito, 1462. Pampayacu, Sawada P25; P29; P93. 
Southeast of Huanuco, 2099. Huassa-huassi and Pillao, Ruiz & 
Pavon, type; Dombey. Cuzco: Paucartambo, Vargas 321 (det. 
Standl.). "Tire," "tire-morado," "tiri," "tiri-cimaron." 

Miconia megaphylla Gleason, Bull. Torrey Club 59: 363. 1932. 

Stems stout, obscurely 4-angled, very thinly tomentulose, 
appearing glabrous; petioles stout, 2-3 mm. long; leaves broadly 
elliptic, firm, glabrous above, compactly and minutely ashy-tomen- 
tulose beneath, 9-13-plinerved, broadly rounded at base, shortly 
acuminate, obscurely crenate; bracts oblong, 5-8 mm. long, decid- 
uous; panicle appressed-stellate-pubescent, the 6-merous sessile 
flowers glomerate; calyx tubular, the coriaceous ovate sepals 3.3 
mm. long, the outer teeth salient at nearly a right angle, 3-sided, 
keeled on the inner side; stamens equal; connective simple; ovary 
4-celled with glabrous conic summit, 12-ribbed; style 10 mm. long, 
the stigma truncate. The leaf-pubescence is so fine and closely 
appressed that the leaves appear merely ashy beneath, the veins 
glabrous; in the Peruvian material, the panicle is extremely narrow, 
in the type pyramidal. Section Jucunda. 

San Martin: Near Moyobamba, 1,200 meters, 3 meters high, the 
flowers lilac-rose, Klug 3518 (det. Gl.). Bolivia. 

Miconia membranacea Triana, Trans. Linn. Soc. 28: 115. 
1871; 830. 

Related to M. glaucescens and similar but the leaves broader, 6-9 
cm. wide, extremely thin, sessile and distinctly 3-plinerved, the 
calyx more or less furfuraceous-pubescent, the petals subacute and 
the stigma minutely capitate; anther connective a little enlarged 
posteriorly. The brown lepidote-punctae do not conceal the under 
leaf surface. Section Eumiconia. 

Huanuco: Valley of the Monzon, 700 meters, Weberbauer 3682 
(det. Cogn.); 285. San Martin: Uchiza, Ruiz & Pavon, type. F.M. 
Neg. 17109. 

Miconia Miles-Morgani Macbr. Field Mus. Bot. 4: 184. 1929. 

Branches slender, nearly terete, early more or less scurfy-stellate 

as the petioles and peduncles, the former slender, 1-1.5 cm. long; 



430 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves broadly ovate, rounded or slightly cordate at base, scarcely or 
not at all acute, obscurely undulate-crenulate, membranous, minutely 
and sparsely stellate-pubescent above, becoming glabrous and some- 
what reticulate bullate, rather sparsely stellate on the 5 nerves and 
reticulate veins beneath, mostly 5 cm. long, 3 cm. broad; panicles 
narrowly pyramidal, about 8 cm. long; pedicels 1 mm. long, the 
flowers sometimes nodding, 5-merous; calyx nearly glabrous, ob- 
scurely denticulate, 2 mm. long, as also the white petals; anthers 
subcuneiform, broadly 2-pored, 1.5 mm. long; style 2 mm. long, the 
stigma peltate. The marginal nerves are faint; a bush to 1 meter 
high, with fragrant white flowers, apparently related to M . peru- 
viana. Named for Miles Morgan, of the Cerro de Pasca Copper 
Corporation. Section Cremanium. 

Huanuco: Between Muna and Tambo de Vaca, 1+318, type. 

Miconia minutiflora (Bonpl.) DC. Prodr. 3: 189. 1828; 855. 
Melastoma minutiflora Bonpl. Melast. 50. pi. 22. 1816. 

Divaricately branched even to the inflorescence, the slender 
branches slightly compressed, glabrous or nearly so or only with some 
scattered, minute, stellate trichomes; petioles 3-6 mm. long; leaves 
oblong-lanceolate, often shortly narrowed at base, acute to caudate- 
acuminate, 8-12 cm. long, 2-5 cm. broad, 3-nerved; panicles nearly 
as broad as long; flowers subsessile, the calyx limb deciduous, indis- 
tinctly 5-lobed, the petals white, 2-2.5 mm. long; style 4 mm. long, 
enlarged above, truncate. The calyx is scarcely 1.5 mm. long. 
Here probably belongs M. Candolleana Triana, 854, and M. pusilli- 
flora (DC.) Triana, 864, as to Peruvian material so named. Calyx 
lobes at most 0.5 mm. long, acute or subacute, with broad rounded 
sinuses; stamens barely dimorphic, the lateral lobes of the connective 
of the smaller not salient or none, those of the larger, small or none 
(Gleason, Bull. Torrey Club 55: 117. 1928). Section Glossocentrum. 

Junin: Chanchamayo Valley, Schunke 422; 268; 1506. La 
Merced, 5476 (T) (compare M. Regelii). Huanuco: Cuchero, Poeppig 
116; 1741- Chinchao, Ruiz & Pavon. San Martin: Tarapoto 
(Fielding 1268). Moyobamba, Mathews 1268. Loreto: Williams 
1616 (det. GL). Pongo de Manseriche, Tessmann 3643 (det. Markgr. 
as M. Candolleana). Florida, Klug 2177 (det. Standl.). Iquitos, 
King 736; Killip & Smith 27346. Northern South America and 
Trinidad. "Yanabarigu-ey" (Huitoto), "carachupa sacha." 

Miconia modica Macbr. Field Mus. Bot. 4: 182. 1929. 
Upper branches obtusely tetragonous, sulcate, with the petioles, 
panicles, calyces and leaves beneath, especially on the nerves, shortly 



FLORA OF PERU 431 

and densely scurfy-stellate with brown trichomes; petioles 4-7 cm. 
long; leaves shortly 5-plinerved, rather firm, entire or obscurely and 
remotely callous-denticulate, ovate-elliptic, obtuse or subrounded 
at base, gradually obtusely acuminate, mostly 2 dm. long and half 
as broad, glabrous above; transverse veins prominent beneath; pani- 
cles ample; pedicels 1-2 mm. long; flowers 5-merous; calyx about 3 
mm. long; petals white, 4 mm. long, puberulent; filaments and style 
glandular; anthers oblong, not at all ventricose, scarcely 2 mm. 
long; style to 5 mm. long, the stigma minutely capitate. A speci- 
men only in bud, Sawada 57, from Pampayacu, with leaves pilose 
beneath, probably is related. Sawada 85, from Chinchao, also only 
in bud, is glabrate; both approach M. majalis Cogn., 886, et al. 
Section Amblyarrhena. 

Ayacucho: Choimacota Valley, in evergreen forest, 8 meters high, 
Weberbauer 7552, type. 

Miconia mollis Triana, Trans. Linn. Soc. 28: 113. 1871; 822. 

Branches and leaves beneath cinereous with a dense pilose, 
stellate tomentum; petioles 3-7 mm. long; leaves elliptic-oblong, 
3-nerved with 2 additional partial submarginal nerves, minutely cor- 
date at base, shortly acuminate, in age only scabrous above, 1-2 dm. 
long, 3-7 cm. broad, acutely and rather unevenly dentate; panicles 
5-10 mm. long; bractlets 2-5 mm. long; flowers sessile or nearly so, 
moderately crowded at the ends of the panicle branches; calyx 5 mm. 
long, suburceolate; petals stellate-tomentose, 4 mm. long; connective 
smooth; style to 9 mm. long. Well marked by the leaf dentation, 
the teeth 3-5 mm. long. Section Eumiconia. F.M. Neg. 21190. 

San Martin: Lamas, Mathews 1292. Tarapoto, Spruce 4177; 
Williams 6718 (det. Gl.). 

Miconia monzoniensis Cogn. Bot. Jahrb. 42: 140. 1908. 
Miconia pterogona Gleason, Amer. Jour. Bot. 19: 746. 1932. 

Nearly glabrous shrub with slender, acutely more or less wing- 
angled branches, firm or coriaceous, oblong or elliptic-obovate, 
sharply acute, entire, shortly 3-plinerved leaves and 5-merous 
flowers borne in dense panicles 5-7 cm. long; petioles stout, 2-5 mm. 
long; leaves rather lustrous above, obtuse at base, 5-8 (15) cm. long, 
half as broad, the somewhat arachnoid-scurfy nerves very prominent 
beneath, with numerous widely spreading transverse veins; calyx 
tube fleshy, 3 mm. long, the teeth 0.5-0.7 mm. long; petals white, 
obliquely and broadly ovate, glabrous, 2 mm. long; stamens gla- 
brous, connective simple, the style 3-4 (4.5) mm. long, slightly 



432 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

enlarged apically; ovary 3-celled (Gleason). The sessile or sub- 
sessile flowers are compactly borne; allied to M. elongata. Erect 
tree 6 meters high, the fruit pink (Mexia). The Mexia specimen in 
Field Museum Herbarium seems to be M. livida. The type of 
Gleason's species has more stoutly winged branches, somewhat 
larger leaves, and is obscurely appressed-arachnoid or glutinous- 
scurfy, characters in degree found in type of Cogn. from same 
region. Section Amblyarrhena. F.M. Neg. 17117. 

Huanuco: Southwest of Monzon, 2,400 meters, Weberbauer 3420, 
type. Distrito Churubambo, in forest, 1,750 meters, Mexia 8238 
(det. Gl.). Pan de Azucar, Sawada 79, type, M. pterogona. 

Miconia mucronata (Desr.) Naud. Ann. Sci. Nat. se"r. 3. 16: 
120. 1851; 732. Melastoma mucronata Desr. in Lam. Encycl. 4: 
46. 1797. Melastoma holosericea L. Sp. PI. 390. 1753. M. holosericea 
(L.) Triana, Trans. Linn. Soc. 28: 101. 1871 (not DC.). 

Branches glabrate or with the leaves beneath more or less velvety 
stellate, the indument often so compact that the leaf-surface appears 
painted; petioles 1-3 cm. long; leaves coriaceous, ovate-elliptic, 
abruptly acuminate or long-caudate, acute or cuneate at base, gla- 
brous above, 3-5-nerved or 3-5-subplinerved, 1-4 dm. long, 6-12 cm. 
broad; flowers 6-9 mm. long, sessile, usually 6-merous, umbellately 
disposed in many-flowered panicles; bracts 3-4 mm. long, caducous; 
calyx tube 6 mm. long, the rounded sepals 2.5-3.5 mm. long, cal- 
loused by the outer tooth, deciduous; petals 9 mm. long, slightly 
retuse; filaments 6-8 mm. long, equaled by the anthers, these sub- 
ulate, the extended connective with 2 obtuse basal lobes and a short 
erect dorsal spur; ovary usually 4-celled; style to 18 mm. long and 
stigma truncate. Section Jucunda. 

The pubescence of this and other species with similar indument 
has been used for tinder. 

San Martin : Lamas, Williams 6435 (det. Gl.) . Near Moyobamba, 
1,100 meters, King 3342. Junin: San Ramon (Killip & Smith 
24766; 24800, det. Gl.). Bolivia and Brazil to Trinidad. 

Miconia muricata (D. Don) Triana, Trans. Linn. Soc. 28: 102. 
1871; 756. Axinaea muricata D. Don, Mem. Wern. Soc. 4: 321. 1823. 

Flowering branches very shortly scurfy or muricate, subquadrate; 
petioles 2-4 cm. long; leaves elliptic-obovate, subcordate at base, 
caudate-acuminate, densely crenulate-denticulate, glabrous above, 
sparsely scurfy beneath, firm, 5-nerved, 1.5-3 dm. long, 10-17 cm. 
wide; panicles to 2 dm. long; pedicels 1-7 mm. long; flowers 5-merous; 



FLORA OF PERU 433 

calyx densely muricate, 6 mm. long, the rounded lobes with a 
minute dorsal tooth. The conical tubercled calyx is usually muricate- 
scurfy or early merely puberulent, but often the conical tubercles 
on the fruiting calyx are conspicuous, sometimes even to 0.5 mm. 
long. In spite of the many collections made, M. muricata has never 
been found in flower, simply, perhaps, because in flower it has been 
called M. glandulifera. Section Octomeris. F.M. Negs. 34153; 17120. 
Huanuco: Pozuzo, 4573(1) (in fruit). Junin: Vitoc, Ruiz & Pavdn, 
type. La Merced, Kittip & Smith 23526; 24085 (det. GL). 

Miconia myrianthaBenth.inHook. Journ.Bot.2: 314.1840; 855. 

Similar to M. minutiflora; sepals triangular-ovate, scarcely 1 mm. 
long, acute, reflexed at anthesis, with a minute outer tooth, the 
intervening sinuses narrow and acute; stamens distinctly dimorphic, 
the larger with connective prolonged into a dorsal appendage, with 
2 narrow lateral lobes directed forward, the smaller with narrow spur 
and 2 short lobes; the calyx is often stellate-puberulent. Lowlands. 

Peru (probably). Trinidad to Bolivia. 

Miconia neriifolia Triana, Trans. Linn. Soc. 28: 127. 1871; 
907. M. nervifolia Triana, I.e. 181 (index). 

Stoutly branched, the branches obtusely 4-sided, the petioles 
thick, 3-5 mm. long, the leaves fleshy coriaceous, narrowly lanceo- 
late, entire, obtuse, 7-10 cm. long, 13-18 mm. broad, smooth, pale 
and opaque above, densely appressed stellate-tomentose with rusty 
trichomes, as also the compactly branched panicle; petioles 3-5 mm. 
long; calyx 2.5 mm. long, the limb 5-lobed; petals suborbicular, 2 
mm. long; connective dorsally bituberculate at base; ovary glabrous, 
the style short, the stigma truncate. The strict panicle branches 
are appressed-ascending with thick pedicels; the transverse leaf- 
veins are manifestly numerous beneath. The var. brevifolia Cogn. 
Bot. Jahrb. 42: 143. 1908 has leaves 3.5-5 cm. long, 12-19 mm. 
broad, the flowers a little larger. Section Cremanium. F.M. Neg. 
17122 (var.). 

Amazonas: Chachapoyas (Mathews 24, type). Huanuco: South- 
west of Monzon, 3,200 meters, Weberbauer 3356, the var. Chinchao, 
Sawada 58. Wet, rocky montana, Playapampa, 4866. 

Miconia nervosa (Smith) Triana, Trans. Linn. Soc. 28: 111. 
1871; 818. Melastoma nervosa Smith in Rees, Cycl. 23: n. 31. 1822. 

A shrub usually about 2 meters high with strikingly 5-7-plinerved 
leaves typically ovate, acuminate and well-attenuate into a short 
petiole (this 1-2 cm. long) or subsessile, somewhat scabrous-setulose 



434 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

above, subappressed-pilose beneath, 1.5-3 dm. long, 5-11 cm. 
broad, usually membranous; branches and panicles setulose or some- 
what villous (the trichomes often light brown), the panicle extremely 
narrow, the flowers congested on the branches that are rarely longer 
than 1 cm.; calyces sessile to 4 mm. long; petals red, about 4 mm. 
long; connective simple; ovary nearly free; style 5 mm. long. The 
var. mediana Macbr. Field Mus. Bot. 4: 181. 1929, has the leaves 
tapering to petioles 2-4 cm. long, the pubescence shortly hirsutulous 
beneath, the petals white. Very common in Loreto. M. ceramicarpa 
(DC.) Cogn., 819, of the Amazon and the Guianas, mentioned by 
Williams under the name "yutobanco," is probably, as to Peru, 
M. nervosa. In any case there may be only one species; the leaves 
of M. ceramicarpa are more shortly plinerved, more broadly cuneate 
to rounded at base, less pubescent. Section Eumiconia. 

Huanuco: Tingo Maria, Stork & Horton 9554. Junin: Hacienda 
Schunke, La Merced, 5811, type, the var.; 5507, the var. San 
Martin: Williams 6074. (det. Gl.). Chicoplaya, Ruiz & Pawn. 
Without locality, Mathews 1288. Loreto: On the Rio Ucayali, 
Castelnau. Yarina-Cocha, Tessmann 3203 (det. Markgr.). Prov. 
Minas, Poeppig 2557, part. Rio Itaya, Williams 3261 ; Killip & Smith 
29665. Iquitos, Williams 3728; 3729. Pebas, Williams 1869; 1601a; 
1682; 1598. Caballo-Cocha, Williams 2043; 2321. Rio Nanay, 
Williams 784; 685. La Victoria, Williams 3083; 2581. Masisea, 
Killip & Smith 26855. Yurimaguas, Killip & Smith 27967; Williams 
4700; 3879; 4344; 4801. Rio Acre: Krukoff 5358. Bolivia and 
Brazil to Trinidad and Mexico. "Sardina mullaca," "millua mul- 
laca," "atun mullaca." 

Miconia nigricans Cogn. Bot. Jahrb. 42: 146. 1908. 

Branches obscurely angled, the younger, as the petioles and 
broadly pyramidal panicles including the calyces, densely stellate- 
scurfy; petioles slender, 4-7 mm. long; leaves coriaceous, soon gla- 
brous above, early moderately stellate-scurfy beneath, becoming 
glabrate, narrowly oblong-lanceolate, rounded at base, acute or 
shortly acuminate, entire, 3-nerved, 6-9 cm. long, 1.5-2.5 cm. 
broad; panicles dense, 3-5 cm. long, the 5-merous flowers sessile; 
calyx broadly campanulate, 3 mm. long, minutely and subacutely 
toothed; petals white, 2 mm. long; stamens and style glabrous, the 
latter to 2.5 mm. long, the stigma peltate. Allied to M . rufa Triana, 
of Cuba. Section Chaenopleura. F.M. Neg. 17124. 

Amazonas: Tambo Ventillas, east of Chachapoyas, 2,400 meters, 
Weberbauer 4392, type. 



FLORA OF PERU 435 

Miconia nitida (D. Don) Naud. Ann. Sci. Nat. se"r. 3. 16: 244. 
1851; 934. Cremanium nitidum D. Don, Mem. Wern. Soc. 4: 312. 
1823. M. macrostoma Naud. I.e. 235. 

Glabrous or the young branches near the tip with a line of 
evanescent scurfy pubescence between the nodes; leaves very 
minutely and remotely spinulose denticulate, rigid, slightly lustrous 
on both sides, ovate, mostly obtuse or acutish, subcordate at base, 
3-nerved, 3-5 cm. long, about 3 cm. broad; petioles 2-4 (-12) mm. 
long; panicles nodding, few-flowered; pedicels 1-5 mm. long; calyx 
6 mm. long, the lobes broadly rotund; petals 4, white, thick, exserted 
about 5 mm. ; filaments glandular pilose. Stems said to root below, 
the branches slender. The Ruiz & Pavon specimen was at one time 
referred by Triana to M. thyrsiflora, that is, M. thyrsoidea according 
to Cogniaux. The leaves are finely impressed reticulate above. 
Section Chaenopleura. F.M. Negs. 17179; 29507; 25991. 

Peru: Sariapata, Ruiz & Pavdn, type. 

Miconia nobilis Gleason, Bull. Torrey Club 58: 235. 1931. 

Upper branches stout, roundly 4-angled and densely brown- 
tomentose; petioles 1 cm. long, 3-angled; leaves thin, obovate, acute 
at base, abruptly acuminate, entire, to 4.5 dm. long and half as wide, 
5-plinerved (the outer pair submarginal), glabrous above, sparsely 
and thinly lanate beneath, the trichomes short, appressed; panicle 
ample, much branched, its strongly sulcate branches densely brown 
lanate; flowers 5-merous, sessile in small lateral glomerules; calyx 
campanulate, about 3.5 mm. long, soon glabrous, the sepals de- 
pressed, only 0.2 mm. long; petals nearly 3 mm. long; stamens 
somewhat dimorphic, linear, the connective produced and marginally 
glandular; ovary 3-celled; style stout, glabrous, nearly 3 mm. long, 
the stigma capitate. Allied to M. dipsacea and related species. 
Collectors noted calyx as rich pink, corolla white. Section 
Eumiconia. 

Junin: San Nicolas, Pichis Trail, tree 4-5 meters high, in dense 
forest, 1,100 meters, Kittip & Smith 26006, type. 

Miconia obscura (Bonpl.) Naud. Ann. Sci. Nat. se>. 3. 16: 
243. 1851; 889. Melastoma obscura Bonpl. Melast. 121. pi. 52. 
1816. Clidemia obscura DC. Prodr. 3: 162. 1828. 

Upper branches and inflorescences shortly red-brown hirtellous; 
petioles 1-2.5 cm. long; leaves elliptic, acute, entire, subcoriaceous, 
green, but the 3 nerves and the 2 fainter marginal ones beneath 
densely reddish-brown-villous, the veins and surface lightly so, 



436 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

more or less tuberculate or bullate or sparsely asperous above, 1-1.5 
dm. long, 4-6 cm. broad; panicle pyramidal, to 1.5 dm. long, the 
sessile flowers crowded; calyx 3 mm. long; petals ovate, 2 mm. long; 
style 4 mm. long, the stigma lightly dilated. An attractive compact 
shrub. The Ecuadorian type is plinerved, bullate-asperous above. 
The Ruiz & Pavon specimen is merely sparsely asperous above, the 
calyx densely scurfy and setulose. Section Amblyarrhena. F.M. 
Neg. 37460. 

Huanuco: Chinchao, Ruiz & Pav6n(t). Cuchero, Poeppig 120; 
(1 502; 1 726) (?) . Ecuador. 

Miconia opacifolia Macbr. Field Mus. Bot. 4: 184. 1929. 

A small tree about 3 meters high, completely glabrous except 
for a minute scabrosity on the upper side of the leaves, these sessile, 
membranous, drying yellowish above, rather purplish beneath, 
obscurely and remotely undulate denticulate, ovate-oblong or -lanceo- 
late, basally cordate, gradually acuminate, 1.5-2 dm. long, mostly 
7 or 8 cm. broad, 5-7-nerved; panicles pyramidal, about 1 dm. long; 
flowers 5-merous, minute, subsessile; calyx shortly lobed, scarcely 
1 mm. long; petals apparently white; anthers obovoid, 2-pored, 
truncate; style included. Flowering branches somewhat sulcate. 
Apparently allied to M. lilacina Triana. The leaves, living, were 
velvety in appearance. Section Cremanium. 

Huanuco: Hacienda Villcabamba, Rio Chinchao, 5197, type. 

Miconia Ottikeri Macbr. Field Mus. Bot. 4: 185. 1929. 

A shrub about 1 meter high with glabrous, acutely 4-sided and 
somewhat winged branches; petioles 1.5-2 cm. long, setose above 
with long black trichomes; leaves coriaceous, minutely callous- 
ciliate-denticulate, ovate-acuminate, subrotund at base, glabrous 
above, yellowish beneath and glabrous except for the black pilose 
nerves, especially the middle one, mostly 7 cm. long, 3-3.5 cm. 
broad, definitely 3-nerved; marginal nerves nearly obsolete; panicles 
narrow, lax, about 1 dm. long; pedicels 2 mm. long; calyx glabrous, 
subhemispheric, shortly 5-lobed, 3-4 mm. long; fruit red, 5-6 mm. 
thick. Apparently to be compared with M. media but the flowers 
unknown; it also suggests M. flavescens Cogn., 930. Named for Mr. 
Ottiker of the Ferrocarril Central del Peru. Section Chaenopleura. 

Huanuco: Tambo de Vaca, 4430, type. 

Miconia pachydonta Gleason, Phytologia 1: 41. 1933. 
Allied and in general character rather similar to M. megaphylla; 
petioles 5-10 mm. long; leaves obovate-oblong, narrowed or sub- 



FLORA OF PERU 437 

cuneate at base, 12-20 cm. long, 5-8 cm. wide; outer teeth of the 
deciduous sepals large, pyramidal, spreading at nearly right angles, 
the calyx densely and closely ashy-tomentose; style glandular- 
puberulent at base. According to the author, near M. gratissima 
Benth., known from the Amazon Valley, but the leaves much broader, 
the calyx more densely tomentose and the ovary not spinulose tipped. 
Section Jucunda. 

Loreto: In dense forest at Florida, Klug 2143, type; 2223. 
"Jucaguino-ey," "chaita-nargu-ey" (Huitoto). 

Miconia pandurata Triana, Trans. Linn. Soc. 28: 108. 1871; 
807. 

Small, little-branched shrub 12 dm. high, the strong, glabrous 
branches obtusely tetragonous; leaves membranous, brown above, 
purplish beneath, glabrous or scarcely scurfy, 5-plinerved (lateral 
nerves arising about the middle), sessile, rounded-cordate from 
the base, pandurate, acuminate, entire, 3-4 dm. long, 12-16 cm. 
broad; panicles broadly pyramidal, 2 dm. long, the branches divari- 
cate; calyx densely scurfy-pubescent, 2.5 mm. long, the somewhat 
dilated limb obscurely 5-dentate. Section Eumiconia. 

San Martin: Campana Mountains, 1,500 meters, (Spruce, type). 
Colombia. 

Miconia papillosa (Desr.) Naud. Ann. Sci. Nat. se>. 3. 16: 
216. 1851; 892. Melastoma papillosa Desr. in Lam. Encycl. 4: 48. 
1797. 

Terete younger branches and calyces densely scurfy-tomentose 
as also the petioles and panicle-branches, but these usually with a 
few stiff simple trichomes intermixed; petioles 2-4 cm. long; leaves 
mostly ovate-oblong, obtusish, subcordate at base, crenulate, sub- 
membranous, 7-nerved, 6-10 cm. long, 3.5-5.5 cm. broad, conspicu- 
ously and coarsely bullate above and with a few short setae or 
glabrous, correspondingly deeply foveolate beneath, densely to rather 
sparsely crinkly hirtellous on the reticulate veins, the nerves scurfy 
with minute barbellate trichomes; flowers mostly sessile, crowded 
in panicles 5-10 cm. long; calyx 3 mm. long; petals subrotund, 
1.5 mm. long; style 4-5 mm. long, the stigma subcapitate. Suggests 
a little M. trichrona but the leaves narrower and smaller, branches 
scurfy and flowers crowded but not glomerate. Section Amblyar- 
rhena. F.M. Neg. 25998. 

Peru (probably). Bolivia; Ecuador. 



438 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia pauciglandulosa Naud. Ann. Sci. Nat. se>. 3. 16: 
183. 1851; 801. 

Glabrous or scarcely puberulent at the tips of the slightly flat- 
tened branches and on the leaf nerves beneath; petioles 1-2 cm. 
long; leaves broadly ovate, rounded at base, acuminate, 1-1.5 dm. 
long, 5-9 cm. broad, membranous, shortly 5-plinerved; panicles 
scarcely 1 dm. long, the 5-merous flowers solitary at the tips of the 
panicle-branches or lateral; calyx tube 2 mm. long, the triangular 
lobes 0.7 mm. long; petals retuse, 2.5-3 mm. long; connective 
minutely produced at base and with a few pedicellate glands; 
style 5-6 mm. long. The dried panicles, and to some extent the 
leaves, have a metallic tinge. Section Eumiconia. F.M. Neg. 26000. 

Huanuco: Cuchero and Pampayacu, Poeppig 2; 108; 1780, type 
colls. 

Miconia Pavoniana Naud. Ann. Sci. Nat. se>. 3. 16: 180. 1851; 
903. 

Branches stout, obtusely angled, reddish; petioles 1-1.5 cm. long; 
leaves oblong-elliptic, acutish at base, acuminate, subcoriaceous, 
rusty-red tomentose-stellate beneath, dark green above, entire or 
crenulate toward the tip, 5-nerved (or shortly plinerved), 1-2 dm. 
long, 6-7 cm. wide; panicle 1-2 dm. long, the usually 5-merous sessile 
flowers glomerate; calyx truncate, 3 mm. long; petals 2 mm. long; 
style 4 mm. long, the stigma subclavate. Sawada 65 has smaller 
plinerved leaves. Cf. M. Herrerae. Section Cremanium. F.M. 
Neg. 36272. 

Huanuco: Tambillo, southwest of Panao, 3568. Pampayacu, 
Kanehira 76. Pillao, in 1787, Ruiz & Pavon, type. Quebrada Pana- 
huanca, (Mathews 872). Pan de Azucar, Sawada 6'5(?). 

Miconia pedicellata Cogn. Melast. 875. 1891. 

Glabrous; petioles 3-7 mm. long; leaves rigid but fragile in the 
herbarium, pale green above, brown beneath, narrowly lanceolate, 
acutish at base, long-acuminate, 8-12 cm. long, merely 2-2.5 cm. 
broad, 5-plinerved or rather 3-plinerved, the outer nerves very faint 
and not obvious above; panicle subcorymbose, 2-3 cm. long; pedicels 
2-5 mm. long; calyx tube 2 mm. long, the lobes less than 1 mm. 
long; petals 5, ovate, 3 mm. long; style somewhat enlarged apically, 
4-5 mm. long. Section Amblyarrhena. F.M. Neg. 32339. 

San Martin: Tarapoto, Campana Mountain, Spruce 434-2. Costa 
Rica; Colombia. 



FLORA OF PERU 439 

Miconia peruviana Cogn. Melast. 909. 1891. 

Branches obscurely 4-sided, the younger densely scurfy-hirtel- 
lous; petioles 5-10 mm. long; leaves narrowly ovate, basally obtuse, 
acute, 5-nerved, denticulate-ciliate, stellate-scurfy beneath, especially 
on the nerves, shortly and densely setulose-hirtellous above, 4-7 cm. 
long, 2-4 cm. wide, the numerous veins prominent beneath; panicles 
densely flowered, 5-6 cm. long, the pedicels scarcely 1 mm. long; 
calyx glabrous, obscurely 5-denticulate; style subclavate. Drying 
yellowish brown and suggesting Clidemia in general aspect. Section 
Cremanium. F.M. Neg. 17129. 

Puno: Tabina, Lechler 2066, type. 

Miconia phaeophylla Triana, Trans. Linn. Soc. 28: 113. 
1871; 825. 

Tree 6 meters high, the upper flattened branches and the leaves 
beneath densely and minutely rusty-stellate, the tiny trichomes sub- 
lepidote; petioles 2-2.5 cm. long; leaves ovate or oblong-elliptic, 
decurrent to the petiole, obtusely caudate-acuminate, at first some- 
what scurfy, in age glabrous and lustrous above, fleshy coriaceous, 
5-nerved or shortly 5-plinerved, 1-1.5 dm. long, 4-6 cm. broad; 
panicle broadly pyramidal or subcorymbose; flowers subsessile, 
subumbellate at the tips of the panicle-branches; calyx turbinate- 
campanulate, 2.5-3 mm. long; petals white, 2.5 mm. long; anther- 
connective cordately auricled; style 4 mm. long. Section Eumiconia. 
F.M. Neg. 26005. 

San Martin: Tarapoto, Spruce 4861, type. 

Miconia pileata [Schrank & Mart.] DC. Prodr. 3: 180. 1828; 
867. Melastoma pileatum Schrank & Mart, ex DC. I.e. in syn. 

Marked, as the closely related M. lacera, by the conspicuous 
ciliation on the leaf margins, but the trichomes shorter and stiffer, 
often sparser, as also on the branches; leaves firm, 3-nerved but 
with an additional fainter marginal pair; calyx 2-2.5 mm. long, the 
inner lobes much shorter than the tube, entire; petals 2 mm. long; 
style 2 mm. long; connective simple. Section Amblyarrhena. F.M. 
Neg. 26007. 

San Martin: Vicinity of Tarapoto, Williams 6369; 6390; 7306; 
7412 (all det. Gl.). Brazil; Colombia. "Casha mullaca," "millua 
mullaca." 

Miconia Pilgeriana Ule, Notizbl. Bot. Gart. Berlin 6: 363. 1915. 
Younger branches, petioles and panicles most minutely scurfy- 
stellate; petioles 8-13 mm. long; leaves oblong-elliptic or ovate-ob- 



440 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, long and mucronulately acuminate, acute but shortly so at the 
rounded base (typically; sometimes even decurrent), the margins 
slightly undulate, 3-nerved with an outer fainter pair, firm, glabrous 
above unless for the nerves, more or less minutely stellate, scurfy- 
puberulent beneath, about 5 cm. wide, 12 cm. long; panicles nearly 
as broad as long, the slender branches spreading-ascending; flowers 
subsessile, 5-merous; calyx tube campanulate, truncate, indistinctly 
denticulate, 2 mm. long; petals elliptic, 2 mm. long; style thick, 1.5 
mm. long, glabrous as the linear stamens, but these appearing short 
and broad. Stamens distinctive, being sterile and nearly terete 
below while the distal portion broadens out into obovate fertile 
pollen sacs opening by a wide terminal pore similar to those of 
M. stellipilis Cogn. of Bolivia, with differently shaped petals (Glea- 
son). Section Glossocentrum. F.M. Neg. 17134. 

Loreto: Iquitos, Tessmann 5062; 3618 (det. Markgr.); 3034; 
King 1350. Pebas, Williams 1714; 1779; 1718. Caballo-Cocha, 
Williams 2155; 2319 (distr. as M. calvescens). Rio Itaya, Wittiams 
160; 3339. Florida, King 2258. Brazil. "Rifai," "huirima-ey" 
(Huitoto), "palo bianco," "mullaca." 

Miconia Poeppigii Triana, Trans. Linn. Soc. 28: 107. 1871; 808. 
Miconia congesta Cogn. Repert. Sp. Nov. 8: 2. 1910, fide Gleason. 

A white-barked tree sometimes 25 meters high, very strict in 
habit, with green flowers and acutely angled upper branches that are 
evanescently scurf y-puberulent as the petioles above, these 1-2 cm. 
long; leaves elliptic, acute at base to rather long-attenuate, acumi- 
nate, firm or rigid, glabrous or scarcely pulverulent on the nerves 
beneath, shortly 5-plinerved (3 prominent ones, the outer 2 faint), 
1-2 dm. long, 2.5-5 cm. broad; panicles many-flowered; petals 
oblong-obovate, 2 mm. long; anthers unequal, the connective of the 
larger curved at base and with 2 minute lateral teeth or spurs; 
calyx scurf y-puberulent, distinctly 4-5-lobed, 1.5 mm. long, obvi- 
ously pediceled; style soon 3 mm. long, the stigma peltate; fruit 
black, 3 mm. thick. Description of flowers from Williams 3698. 
Section Eumiconia. F.M. Neg. 17135. 

San Martin: Tocache, Poeppig 1912, type. Loreto: Yurimaguas, 
Tessmann 3766 (det. Markgr.); Wittiams 4147; 4373, etc. (det. Gl.). 
Iquitos, Williams 3698 (det. GL). La Victoria, Williams 2900. Rio 
Itaya, Wittiams 3340. Bolivia; Brazil. "Rifari," "rupina." 

Miconia polyneura Triana, Trans. Linn. Soc. 28: 128. 1871; 909. 

Closely allied to M. peruviana but nearly glabrous except the 

scurfy-puberulent leaves and calyx; branches annulately enlarged 



FLORA OF PERU 441 

at the nodes; petioles 1-2.5 cm. long; leaves shortly and obtusely 
acuminate, callous-denticulate, subcoriaceous, pale and scurfy, 
especially on midrib beneath, 7-9 cm. long, 3-5 cm. broad; panicles 
broadly pyramidal, 1 dm. long; pedicels 1-3 mm. long; calyx 2 mm. 
long; petals subrotund, 1.5 mm. long; connective obscurely bitu- 
bercled; style 3^4 mm. long, the stigma minutely capitate. Gleason 
in herb, has questioned the determination of the Weberbauer col- 
lection, which seems to merge with M. galactantha. M. loxensis 
(Bonpl.) DC., 911, to be expected in adjacent Peru, has setose 
denticulate leaves, subclavate stigma. Section Cremanium. F.M. 
Neg. 26013. 

Ayacucho: Prov. Huanta, 3,100 meters, Weberbauer 5585 (det. 
Cogn.). Without locality, Bonpland. Colombia. 

Miconia prasina (Sw.) DC. Prodr. 3: 188. 1828; 805. Mela- 
stoma prasina Sw. Prodr. 69. 1788. 

Essentially glabrous or the branches obscurely puberulent; peti- 
oles 5-20 mm. long, the oblong-lanceolate leaves often decurrent on 
them, drying yellowish green, acute, glabrous or at first a little 
puberulent beneath, 3-5-plinerved or scarcely so, 1-2 dm. long, 3-8 
cm. wide; panicle usually ample, the branches opposite, the sessile 
or subsessile 5-merous flowers crowded toward their tips; calyx nar- 
rowly campanulate, about 3 mm. long, thinly scurf y-puberulent, 
acutely or obtusely toothed, the outer and inner teeth subequal; 
petals 2-3 mm. long; connective bilobed laterally; ovary 3-celled; 
style to 6 mm. long, slightly dilated to the truncate stigma. 
Resembles in fruit M. glaberrima. Sometimes a tree 7 meters high. 
Gives a black dye. Section Eumiconia. 

Puno: Valley of the Sandia, Weberbauer 1113 (det. Cogn.); 278. 
Loreto: Rio Nanay, Williams 572; 974- Yurimaguas, Williams 
4807; Killip & Smith 28929. Paraguay to Mexico and the West 
Indies. "Mullaca Colorado," "mullu caspi," "isula micuna," 
"tintureira." 

Miconia pseudo-centrophora Cogn. Melast. 893. 1891. 

Branches subterete, scurfy-puberulent toward the tips as also 
the petioles and inflorescence, the former 1-2 cm. long, the latter 
pyramidal, 1 dm. long; leaves drying rigid and dark-colored above, 
where glabrous but minutely bullate, slightly foveolate and ashy- 
stellate-punctate beneath, rather closely mucronulate-dentate, 5- 
nerved, 6-10 cm. long, 2.5-4 cm. wide; pedicels 1-2 mm. long; 
calyx 2.5 mm. long; petals subrotund, 1.5 mm. long; style 4 mm. 



442 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, the stigma subcapitate. In the Peruvian specimen the leaves 
are sparsely scurfy only on the veins beneath or glabrate, the pani- 
cle 5-10 cm. long, the short appressed branches with glomerate 
flowers. Doubtfully distinct from M. centrophora, at least not 
separable on the character of plane or bullate leaves. Section 
Amblyarrhena. F.M. Neg. 17138. 

Ancash: Huaraz, 3,000 meters, Weberbauer 3259 (det. Cogn.). 
Ecuador. 

Miconia pterocaulon Triana, Trans. Linn. Soc. 28: 114. 
1871; 794. 

Branches stout, broadly 4-winged; petioles strongly flattened, 
1-2.5 cm. long; leaves entire or the margins scarcely undulate, 
broadly oblong-lanceolate, long-attenuate to the subacute base, 
shortly acuminate, submembranous, glabrous above, silvery beneath 
with an extremely fine appressed tomentum, 2-3 dm. long, 7-11 cm. 
wide; panicle narrow, the branches simple or bifid, 1-1.5 dm. long; 
calyx 2 mm. long, shortly 5-lobed; fruit globose, 3-4 mm. thick. 
The stellate gray-brown indument on the leaves beneath is so dense 
and so fine that it appears to be a mere discoloration. Section 
Eumiconia. F.M. Neg. 26014. 

Peru(?) : Without locality, Ruiz & Pavon (det. Markgr.). Colombia. 

Miconia pteropoda Benth. in Hook. Journ. Bot. 2: 314. 1840; 
803. 

Closely allied to M. prasina but the petioles apically wing-mar- 
gined by the decurrence of the leaves, these larger, even to 2.5 dm. 
long, 7-9 cm. broad; panicle branches divaricate; calyces distinctly 
but extremely minutely puberulent, the outer teeth obsolete or 
tubercle-like, the inner depressed; petals obscurely puberulent with- 
out; stamens isomorphic, the connective with 2 lateral lobes that 
are deflexed anteriorly (Gleason). The leaves are more distinctly 
plinerved than usual in the similar species; cf. also M. longiracemosa. 
Section Eumiconia. F.M. Neg. 26015. 

San Martin: Near Moyobamba, 7-meter tree with cream-colored 
flowers, 1,100 meters, King 3312; 3643 (det. Gl.). Loreto: Itaya, 
inundated bank, shrub or tree to 8 meters, with white flowers, 
Mexia 6484 (det. Gl.). La Victoria, Williams 2707; 2987. Pebas, 
Williams 1636. Rio Itaya, Williams 115. Florida, King 2127. 
Brazil to the Guianas and Panama. "Acano-fue-peroi" (Huitoto), 
"y ana 'P an g a >" "bucacuru caspi." 



FLORA OF PERU 443 

Miconia puberula Cogn. in Mart. Fl. Bras. 14, pt. 4: 386. 
pi. 78. 1887; 856. 

Similar to M. minutiflora but the narrowly ovate leaves somewhat 
cordate at base and evanescently more or less rusty-floccose-puberu- 
lent, at least on the 5 nerves beneath; calyx limb truncate or obscurely 
lobed, only 1 mm. long, petals 1.4 mm. long; connective very shortly 
spurred; style 2 mm. long. The sometimes subsessile leaves are 
glabrous and chartaceous in age. A shrub 1-5 meters high with 
blue fruit (Ule). Section Glossocentrum. F.M. Neg. 26016. 

San Martin: Moyobamba, 800 meters, Weberbauer 44-92 (det. 
Cogn.); 290. Near Moyobamba, Klug 3362. Tarapoto, (Spruce; 
Mathews, type colls.); Ule 6745; 6746; Williams 5376; 5628; 5702; 
6150; 6425; 7103 (mostly distr. as M. minutiflora}. Junin: San 
Ramon, Killip & Smith 24790. "Uchu mullaca." 

Miconia pubipetala Miq. Stirp. Surinam. 50. 1850; 743. 
Diplochita parviflora Benth. in Hook. Journ. Bot. 2: 302. 1840. 
Miconia parviflora (Benth.) Cogn. in Mart. Fl. Bras. 14, pt. 4. 249. 
1887, not Naud. 

Younger branches, leaves beneath and calyces minutely or ob- 
scurely brown-stellate- tomentose; petioles 1.5-4 cm. long; leaves 
rather thin, usually drying black and lustrous above, obovate to 
elliptic-oblong, to 3 dm. long, about half as broad, often falcately 
acuminate, obtuse or rounded at base, 3-5-nerved, the outer nerves 
submarginal ; panicles to 2 dm. long, with 2-several branches at each 
node; flowers 5-merous, in terminal cymules, the pedicels 2-4. mm. 
long; calyx ashy-tomentulose, 4 mm. long; petals narrowly oblong, 
to 7 mm. long, minutely scurfy; connective appendaged, that of 
the episepalous stamens with a few stalked glands; ovary minutely 
pubescent, the puberulent style to 13 mm. long, the stigma truncate. 
Inflorescence bracts lacking or minute. Section Tamonea. 

Loreto: Williams 901 (det. Gl.). Trinidad; Guianas. "Sachi 
mullaca." 

Miconia Pulgari Macbr. Field Mus. Bot. 4: 182. 1929. 

Nearly glabrous straggling shrub with slender terete branches, 
these, as the petioles and peduncles, sometimes with a few crisped 
trichomes; petioles 8-10 mm. long; leaves entire or minutely and 
appressed-ciliate, oblong-lanceolate, acute at base, subabruptly and 
obtusely acuminate, membranous, glabrous above, sparsely pilose- 
hirtellous beneath only on the nerves and veins, 3-plinerved, not 
counting the marginal nerves, about 1 dm. long and 3 cm. broad, 



444 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the transverse veins and veinlets prominent beneath; panicles 
4-6 cm. long; flowers pinkish-white, 5-merous, on pedicels 1 mm. 
long; calyx glabrous or slightly granulose-stellate, denticulate, 
nearly 4 mm. long; petals broadly obovate, 2.5 mm. long; anthers 
narrowly obovoid, minutely 1-pored; style 5 mm. long, the stigma 
somewhat capitate. Named for Senor Francisco Pulgar of Panao. 
Nearly M. pedicellata, but that species completely glabrous. Section 
Amblyarrhena. 

Huanuco: Huacachi, near Muna, 4145, type. 

Miconia pulverulenta R. & P. Syst. Veg. 1: 104. 1798; 902. 
Cremanium pulverulenta (R. & P.) D. Don ex Triana, Trans. Linn. 
Soc. 28: 126. 1871. M. ramosipila Macbr. Field Mus. Bot. 4: 185. 
1929. 

Branches stout, toward the terminal inflorescence more or less 
shortly pubescent with a sordid scurfy tomentum of plumose tri- 
chomes; petioles 1-2.5 cm. long or longer; leaves subcoriaceous, 
entire or obscurely undulate-denticulate, elliptic-oblong, rounded or 
emarginate at base, acute or acuminate, glabrous and obscurely 
bullate above, yellowish stellate and plumose-hirtellous (at least on 
nerves) beneath, the trichomes scarcely concealing the surface, 
3-5-nerved with a submarginal additional fainter pair, 1-1.5 (2) dm. 
long, 5-7 (10) cm. wide; panicle pyramidal or sometimes nearly 
as broad as long, the glomerules of sessile flowers more or less approxi- 
mate in spikes; calyx glabrate, 1.5 mm. long, shortly and obtusely 
lobed; petals obovate, 1 mm. long; style 1.5 mm. long, subclavate 
or apparently (my coll.) peltate. The leaves dry a yellow-brown, 
a lighter greenish yellow beneath. My species has been retained by 
Gleason in herbarium, but I fail to find a sound basis for its separa- 
tion, although the anthers are seemingly sublinear, minutely pored. 
Section Cremanium. F.M. Neg. 17144. 

Huanuco: Chinchao and Huanuco, Ruiz & Pav6n, type. Muna 
and Tambo de Vaca, 4321 (type, M. ramosipila); 4341. Mito, 3411. 

Miconia punctata (Desr.) D. Don, Mem. Wern. Soc. 4: 316. 
1823; 793. Melastoma punctata Desr. in Lam. Encycl. 4: 50. 1797. 

A stoutly branched shrub, the upper branches, as those of the 
inflorescence, acutely angled and densely covered with a red-brown 
punctate-lepidote indument, this clothing also the leaves beneath; 
petioles deeply sulcate, 1-2 cm. long; leaves glabrous above, 3-nerved 
with also an obscure nearly marginal pair, lanceolate, entire, narrowed 
at each end, 10-18 cm. long, 3-6 cm. broad; panicles usually ample; 



FLORA OF PERU 445 

calyx 2 mm. long; petals 2.5 mm. long, style clavate, 4-4.5 mm. long, 
apically truncate; connective prolonged straight back and irregu- 
larly 1-3-lobed (Gleason). Section Eumiconia. F.M. Neg. 38248. 
Illustrated, Bonpland, Melast. pi. 40. 

Loreto: 5-meter tree with white flowers, Klug 2126 (det. Gl.). 
Venezuela and the West Indies. "Ucaguino-ey" (Huitoto). 

Miconia pyrifolia Naud. Ann. Sci. Nat. ser. 3. 16: 164. 1851; 808. 

Related to M. Poeppigii; essentially glabrous, the branches 
nearly terete; leaves shortly acuminate, lustrous above, 7-12 cm. 
long, 4-7 cm. broad; panicles 4-10 cm. long; calyx glabrous or nearly 
so, very shortly 5-lobed, 2.5 mm. long; petals nearly glabrous, about 
2 mm, long; connective well-produced, trifidly dilated at base; 
style 4 mm. long, the stigma subclavate. Very smooth or merely 
evanescently pulverulent on the young parts and inflorescence. 
Section Eumiconia. Illustrated, Mart. Fl. Bras. 14, pt. 1: pi. 62. 

Loreto: Prov. Mainas, Poeppig 2768, part. Brazil. 

Miconia quadrifolia Naud. Ann. Sci. Nat. se"r. 3. 16: 223. 
1851; 910. 

Branches toward the tips, as the inflorescence and the leaves on 
the 3 nerves beneath, scurfy-tomentose, the latter 4-verticillate, 
sessile or subsessile, entire, oblong, long-acuminate, rounded at base, 
9-12 cm. long, scarcely 3 cm. broad; panicles pyramidal, 6-8 cm. 
long; flowers minute, the calyx less than 1 mm. long, subsessile, 
subtruncate; petals subrotund, 1 mm. long; stigma punctiform. 
The Weberbauer specimen is petiolate. Section Cremanium. F.M. 
Neg. 36294. 

Huanuco: Casapi, Mathews 1724, type. Junin: Between Huaca- 
pistana and Palca, 2,000 meters, Weberbauer 2318 (det. Cogn.); 247. 

Miconia radula Cogn. Bull. Acad. Belg. ser. 3. 14: 957. 1887; 
889. M. sphagnophila Macbr. Field Mus. Bot. 4: 183. 1929, fide 
Gleason. 

Branches, petioles and leaves beneath densely hirsute-setulose, 
the trichomes mostly enlarged at base, reddish, those of the leaf- 
nerves beneath often stouter; petioles 5-10 mm. long; leaves ovate- 
elliptic, rounded at base, obtuse or scarcely acute, rigid, densely 
conic-bullate-setulose above, 5-nerved (Cogniaux) or plinerved, 3.5-6 
cm. long, 1.5-4 cm. broad; flowers crowded, subsessile, 5-merous, the 
asperous panicle a few cm. long; calyx 3-4 mm. long, the setae slender 
or conic-based, the lobes short, obtuse; petals suborbicular, 2.5 mm. 



446 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long; filaments and style typically glandular-pilose, the stigma 
peltate. The type by Andr6 from Ecuador has 5-nerved leaves 
according to Cogniaux, but they are actually plinerved; Gleason, 
Bull. Torrey Club 58: 244 in referring here the following collections, 
has accepted, probably rightly, considerable variation, particularly 
as regards the character of the calyx pubescence and the lack of 
glands on filaments in the collection by Killip and Smith, elsewhere 
used as a specific character. M. capitellata Cogn., 892, of Ecuador 
has somewhat larger acutish leaves on petioles 1-2 cm. long, the 
panicle twice as long with densely capitate sessile flowers. Section 
Amblyarrhena. 

Huanuco: 1-meter shrub in sphagnum, 4503 (type, M. sphagno- 
phila). Junin: Pichis Trail, (Killip & Smith 25908}. Amazonas: 
Chachapoyas, 2,400 meters, Weberbauer 4403 (det. Cogn.); 263. 
Ecuador. 

Miconia radulaefolia (Benth.) Naud. Ann. Sci. Nat. se*r. 3. 
16: 243. 1851; 818. Clidemia radulaefolia Benth. in Hook. Journ. 
Bot. 2: 309. 1840. 

Branches subterete, these, as the petioles and narrow panicles, 
appressed to spreading pilose-setulose; petioles 1-2 cm. long; leaves 
oblong-lanceolate, attenuate to the acute base, membranous, shortly 
bullate-scabrous above, minutely subappressed-setulose beneath, the 
trichomes nearly concealing the minutely foveolate surface, crenu- 
late-ciliate, shortly 3-5-plinerved, 1-2 dm. long, 3-6 cm. broad; 
flowers sessile, congested, the panicle 5-15 cm. long; calyx 3 mm. 
long, densely hirsute-pilose, the lobes short, the outer teeth just 
equaling the inner; petals 5 mm. long, style 7-8 mm. long. M. 
scrobiculata Cogn., Amazonian, 817, with leaves obtuse or rounded 
at base, branches and calyces long-pilose, is nearly Williams 8231 
and perhaps not specifically separable, but Gleason distinguishes it 
by the outer calyx tooth projection 0.6 mm. beyond the sepals. 
Section Eumiconia. F.M. Neg. 17146. 

Loreto: Iquitos, Killip & Smith 27307; 29662 (det. Gl.); Williams 
8231. British Guiana. "Mullaca mullaca." 

Miconia Regelii Cogn. in Mart. Fl. Bras. 14, pt. 4: 405. pi. 83. 
1888; 865. 

Branches little compressed, with the petioles and panicles 
densely and minutely scurf y-puberulent; petioles 5-10 mm. long; 
leaves ovate-elliptic, acute at base, shortly acuminate, merely 
granular-scurfy, principally on the nerves, or glabrate beneath, 



FLORA OF PERU 447 

glabrous above, firm, 3-nerved, with also a submarginal very faint 
pair, 6-11 cm. long, 3-5 cm. broad; panicles ample, the branches 
arcuate, the flowers sessile, minutely bibracteolate; calyx 1.5 mm. 
long, the white petals as long, the style twice as long, slightly en- 
larged apically; connective simple. Section Chaenanthera. 
San Martin: Tarapoto, Spruce. Brazil. 

Miconia retusa Pilger, Verh. Bot. Ver. Brandenb. 47: 174. 
1906. M. Macbridei Gleason, Bull. Torrey Club 58: 246. 1931. 

Glabrous shrub with slender virgate terete branches and terminal 
thyrsoid panicles of white flowers borne well above the leaves, these 
crowded, subcoriaceous, cordate-subsessile or almost amplexicaul, 
ovate, entire or callous-denticulate toward the acuminate tip, 3- 
nerved, with an additional outer obscure pair, 5.5-7 cm. long, 2.5- 
3.5 (5) cm. broad; panicles 7-9 cm. long, lax in age, the branches 
somewhat spreading; flowers 4-merous; pedicels 2 mm. long; calyx 
tube campanulate, red-puncticulate, 2 mm. long, the acute teeth 
1 mm. long, equaled by the appressed acute outer teeth (Gleason); 
petals irregularly retuse, thick, white, 2-2.5 mm. long; filaments 
and anthers 2 mm. long, the connective simple; style stout, finally 
exserted 3 mm., the stigma not at all enlarged; ovary inferior, 
4-celled. Gleason remarked regarding his species, "closely related 
to M. retusa, which has a small lax inflorescence and narrower 
leaves"; he did not know the flowers, and these, present in the 
Field Museum specimens, seem to match the description of M. retusa; 
the photograph shows a great similarity, the slight differences in 
leaves and inflorescence being in all probability within the natural 
range of variation to be expected in the species. Section Chaenopleura, 
fide Gleason. F.M. Neg. 17147. 

San Martin: Tarapoto, 750 meters, Williams 5596 (type, M. Mac- 
bridei). Loreto: Cerro de Escaler, 1,100 meters, Ule 6366, type. 

Miconia riparia Triana, Trans. Linn. Soc. 28: 117. 1871; 766. 

Densely leafy, the leaves nearly linear, long-acuminate, the 
slender branches sordidly tomentulose with gray stellate trichomes, 
some longer barbellate ones intermixed; petioles 3-6 mm. long; 
leaves glabrate, 3-nerved, obscurely if at all denticulate, firm or 
thin, to 12 cm. long, 3-15 mm. broad; flowers bracteolate, verticillate 
in a spike 3-5 cm. long, the linear bractlets 4-5 mm. long; fruiting 
calyx sparsely stellate-pilose, obscurely 5-lobed; fruit black, succu- 
lent, subglobose, 4-5 mm. thick. Section Eumiconia. F.M. Neg. 
26025. 



448 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Rio Huallaga, Klug 4009. Loreto: Yurimaguas, 
Spruce 4589, type. Pongo de Manseriche, shrub 1 meter high, 
fruit red, Mexia 6220 (det. GL). Rio Maranon Valley, Kittip & 
Smith 29136 (det. Gl.). 

Miconia rotundifolia (D. Don) Naud. Ann. Sci. Nat. se>. 3. 
16: 235. 1851; 935. Cremanium rotundifolium D. Don, Mem. Wern. 
Soc. 4: 311. 1823. 

Low, the stems creeping, branching, terete, shortly pilose with 
obscurely plumose trichomes; petioles 3-7 mm. long; leaves sub- 
crenulate and ciliate, suborbicular, sparsely setulose above and hirtel- 
lous beneath, 5-nerved including the partial submarginal ones, nearly 
as broad as long, mostly 1.5-2 cm.; flowers often ternate, subsessile 
or some pediceled, to 3 mm. long; petals 2-2.5 mm. long, sub truncate; 
calyx 3 mm. long, the lobes short; fruit 5 mm. thick. A yellow dye 
called "Olla-olla," made from this plant, Bocconia frutescens, and 
several others mixed together, is used by the Peruvian Indians to 
color their bodies (Ruiz & Pavon). Section Chaenopleura. F.M. 
Neg. 17152. 

Huanuco: Playapampa, 3,000 meters, 4490 (det. GL). Huassa- 
huassi, Ruiz & Pavon; Dombey, type. 

Miconia rubens (Swartz) Naud. Ann. Sci. Nat. se>. 3. 16: 169. 
1851; 921. Melastoma rubens Swartz, Prodr. 71. 1788. 

A completely glabrous shrub 3-4 meters high with slender, 
obtusely tetragonous branches and oblong-elliptic, minutely dentic- 
ulate, thin and pellucid leaves, these 3-nerved, with also a faint outer 
pair, or shortly 3-plinerved, acuminate, acute or attenuate at base, 
6-9 cm. long, 2-3 cm. broad; petioles 1-1.5 cm. long; panicles 4-7 
cm. long; calyces subsessile, 2 mm. broad; petals white, 0.7 mm. 
long; style 1.5 mm. long, the stigma peltate. In spite of the author- 
ity for it one wonders if this identification is correct in view of the 
range. Section Cremanium. 

Cajamarca: Near Hualgayoc, 2,700 meters, Weberbauer 4105 
(det. Cogn.); 259. Venezuela to Jamaica. 

Miconia rubiginosa (Bonpl.) DC. Prodr. 3: 183. 1828; 823. 
Melastoma rubiginosa Bonpl. Melast. 109. pi. 1+7. 1816. 

Twigs, petioles, leaves beneath and compact panicles, including 
the calyces, densely red-brown-tomentose with stellately branched 
trichomes, some of them minutely stiped; petioles thick, 2-6 mm. 
long; leaves ovate, shortly acuminate, entire, drying black and 



FLORA OF PERU 449 

lustrous above, subcoriaceous, 6-10 cm. long, 3-6 cm. broad; flowers 
5-merous, crowded, ebracteolate; calyx obscurely lobed, about 3 
mm. long; petals white, rounded, 2 mm. long; connective spurred 
posteriorly, obscurely biauricled anteriorly; style 4 mm. long; fruit 
black, 3-4 mm. thick. The reddish-brown stellate tomentum is 
often more or less deciduous from the leaves beneath in age. Section 
Eumiconia. F.M. Negs. 17153; 36295. 

San Martin : Tarapoto, Mathews 1290; Spruce 4265. Moyobamba, 
Mathews 1289. Loreto: Canela Ucsha, Vie 6738; 67S9 (det. Pilger). 
Brazil ; Bolivia to Colombia and Puerto Rico. 

Miconia rufescens (Aubl.) DC. Prodr. 3: 180. 1828; 777. 
Melastoma rufescens Aubl. PL Guian. 1: 408. pi. 157. 1775. 

Shrub, striking by virtue of the markedly reticulate-bullate, 
subsessile, oval-elliptic leaves and the almost shaggy pubescent 
branchlet tips, the long pubescence of barbellate trichomes extend- 
ing into the narrow crowded panicle of globosely glomerate flowers; 
petioles rarely 5 mm. long; leaves ovate, cordate, often crenulate- 
denticulate, densely reddish stellate-hirsute beneath, 5-7-nerved, 
8-15 cm. long, 5-10 cm. broad; panicles 1-2 dm. long; calyx densely 
long-pilose, 4 mm. long, minutely 5-lobed; petals white(?), glabrous, 
retuse, 4-5 mm. long; style to 9 mm. long; fruit subglobose, 4 mm. 
thick, edible according to Weberbauer. Shrub 1-3 meters high, 
with lilac rose petals (Klug). Section Eumiconia. 

Cajamarca: Prov. Jan, 1,200 meters, Weberbauer 6259.- San 
Martin: Tarapoto, Ule 6450 (det. Pilger); Spruce 4266; Williams 
5422; 7438. Moyobamba (Mathews 1288)', Weberbauer 4495(1} (det. 
Cogn.); 290; Klug 3363 (det. Gl.). Cuzco(?): Gay. Brazil to 
Colombia and British Guiana. "Mullaca," "sira sira." 

Miconia rugifolia Triana, Trans. Linn. Soc. 28: 125. 1871; 894. 

Upper branches tetragonous, scurfy-pubescent; petioles 6-8 mm. 
long; leaves firm, drying fragile, glabrous and bullate above, corre- 
spondingly closely foveolate beneath, where also sparsely hirtellous, 
5-nerved, crenulate, lanceolate, obtuse at base, to 1.5 dm. long, 
2.5-4 cm. broad; panicles pyramidal, short; calyces sessile, 1.5 mm. 
long, the oblong petals 1 mm. long, the style 3 mm. long; connective 
bituberculate anteriorly. Section Amblyarrhena. 

Puno: San Gaban, Lechler 2196, type. 

Miconia Ruizii Naud. Ann. Sci. Nat. seY. 3. 16: 192. 1851; 890. 

Densely but finely hirsutulous except the leaves beneath, these 

shortly villous-tomentose with simple, crisped trichomes, minutely 



450 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

bullate above but the surface nearly concealed by the pubescence, 
the trichomes not at all or little enlarged at base; petioles 1-2.5 cm. 
long; leaves ovate, acuminate, essentially entire, 7-nerved or shortly 
plinerved, 10-16 cm. long, 5-9 cm. broad; flowers sessile, more or 
less crowded, the panicles 5-7 cm. long, about as broad, the trichomes 
on the branchlets and calyx harsher than elsewhere but slender; 
calyx lobes membranous, short, obtuse; petals broadly cordiform, 
1.5 mm. long; filaments glabrous; connective simple; style 2-3 mm. 
long, the stigma peltate. Aspect of a Clidemia. Section Eumiconia, 
fide Gleason, Amblyarrhena, fide Cogn. F.M. Neg. 26027. 

Lima: Near Lima, Ruiz & Pavdn, type; Mathews 3217. Junin: 
Pichis Trail, Killip & Smith 25628; 25748 (det. Gl.). Ecuador; 
Bolivia. 

Miconia salicifolia (Bonpl.) Naud. Ann. Sci. Nat. seY. 3. 16: 
234. 1871; 922. Melastoma salicifolia Bonpl. ex Naud. I.e. M. 
rosmarinifolia Pav. ex Triana, Trans. Linn. Soc. 28: 127. 1871. 

Densely leafy, shortly branched shrub well characterized by 
oblong-linear leaves, these glabrous above, densely rusty-scurfy- 
tomentose beneath, obtuse, the margins becoming revolute, 2-3.5 
cm. long, only 2-7 mm. wide; petioles 2-4 mm. long; panicle racemi- 
form, few-flowered, nodding, the flowers 4-merous, the calyx 3 mm. 
long, subtruncate, the petals 1 mm. long, the style 4 mm. long with 
thick, ovoid stigma. The pubescence is felt-like. Section Cre- 
manium. F.M. Neg. 17157. 

Peru(?): Ruiz & Pavon. Junin: Altos de Vitoc, Isern 584; 574- 
Ecuador; Colombia. 

Miconia sanguinea (D. Don) Triana, Trans. Linn. Soc. 28: 125. 
1871; 896. Tococa sanguinea D. Don, Mem. Wern. Soc. 4: 305. 1823. 
Cremanium hispidissimum DC. Prodr. 3: 192. 1828, et Me"m. Melast. 
80. 1828. M. hispidissima (DC.) Naud. Ann. Sci. Nat. se>. 3. 16: 
191. 1851. 

Upper branches, as the petioles and panicles, spreading pilose- 
hispid; petioles 3-6 cm. long; leaves ovate-oblong, rounded at base, 
hirtellous both sides, membranous, entire or minutely denticulate, 
5-7-nerved, 1.5-2 dm. long, 6-10 cm. broad; panicle broadly pyra- 
midal, to 1.5 dm. long; pedicels 3-8 mm. long; calyx hirsute, 3-4 mm. 
long; petals 4-5 mm. long; filaments and style glandular-pilose, the 
latter stout, 6-9 mm. long. A 3-8-meter shrub or a tree; flowers 
white, fragrant. Section Amblyarrhena. F.M. Neg. 38253. 



FLORA OF PERU 451 

Huanuco: Cushi, 4824; Cuchero and Chinchao, Dombey; Ruiz 
& Pavdn, type; Poeppig 1003; Rio Isabel, Mexia 8114; Stork & 
Horton 9444- Junin: Chanchamayo Valley, Weberbauer 1985 (det. 
Cogn.); 247; Isern 2125. San Martin: Valley of the Monzon, 1,200 
meters, Weberbauer 3499 (det. Cogn.); 284. "Terye." 

Miconia saxatilis Macbr. Field Mus. Bot. 4: 186. 1929. 

Shrub about 3 meters high, the upper branches acutely tetrago- 
nous, enlarged at the nodes and, like the petioles, peduncles and 
inflorescence branches, sordidly and densely stellate-pubescent; peti- 
oles 2-4.5 cm. long; leaves broadly ovate-elliptic, rather abruptly 
narrowed at both ends, shortly acuminate, mostly 15-18 cm. long, 
7.5-9 cm. broad, entire or somewhat undulate-denticulate, mem- 
branous, glabrous and green above, paler and finely stellate-puber- 
ulent on the nerves and veins beneath, shortly 3-plinerved with an 
additional marginal pair; panicles densely flowered, about 8 cm. 
long; flowers 5-merous, subsessile; calyx glabrate, obtusely denticu- 
late, scarcely 1 mm. long, the petals hardly half as long; anthers 
subcuneiform, nearly 0.5 mm. long, 2-pored; style about 1 mm. long, 
the stigma peltate. Slender shrub of wet rocky upland. Section 
Cremanium. 

Huanuco: Playapampa, 4881, type. 

Miconia sclerophylla Triana, Trans. Linn. Soc. 28: 119. 
1871; 846. 

Younger branches (these compressed) and leaves beneath densely 
scurf y-puberulent; petioles 1-2 cm. long; leaves fleshy-coriaceous, 
fragile in drying, oblong-lanceolate, acuminate, subcordate or acute 
at base, glabrate above, rufescent beneath, entire, 5-nerved, 6-10 
cm. long, 2-3.5 cm. broad; panicle pyramidal, the sessile flowers 
densely glomerate on the spreading branches, 5-7 cm. long; calyx 
campanulate, scurf y-tomentose, 2 mm. long, the persisting limb 
acutely lobed; petals obovate, longer than the calyx; anthers linear, 
obliquely 1-pored, the simple connective gradually attenuate with 
the articulate filament. 

Peru(?): (Herb. Fielding), perhaps by Pawn. 

Miconia scorpioides (Schlecht. & Cham.) Naud. Ann. Sci. Nat. 
se'r. 3. 16: 243. 1851; 782. Melastoma scorpioides Schlecht. & Cham. 
Linnaea 5: 564. 1830. 

Upper branches pulverulent-stellate, strongly compressed and 
narrowly wing-margined alternately in opposite directions between 



452 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the nodes; leaves sessile or subsessile, narrowed to base or sometimes 
auriculate, obovate-oblong, apiculate or shortly acuminate, becom- 
ing glabrous above, sparsely stellate-scurfy beneath, the minute 
trichomes dot-like, 3-plinerved with a pair of submarginal ones, 
1.5-2.5 dm. long, 5-9 cm. broad; panicles narrow, the branches 
usually bifid, scorpioid and secundly flowered, pulverulent; calyx 
2 mm. long; petals rounded, 2.5 mm. long; style 6 mm. long; fruit 
broadly subglobose, lightly 10-costate. A tree 4 meters high (Klug). 
Section Eumiconia. F.M. Neg. 17161. 

Huanuco: Near Cuchero, Poeppig i7&?. Loreto: Lower Rio 
Huallaga, Williams 4266; 4617. Florida, Klug 2365; 2801 (det. 
Standl.). Brazil to Mexico. 

Miconia secundifolia Cogn. Bot. Jahrb. 42: 145. 1908. 

A 2-meter shrub with scarcely quadrate, stout, simple branches, 
the younger, as the petioles and peduncles, densely granular or 
papillose-stellate-puberulent; petioles 1-2 cm. long; leaves borne 
secundly, coriaceous, glabrous above except the puberulent mid- 
nerve, stellate-puberulent mostly only on the 5 nerves beneath 
where markedly reticulate veined, entire, ovate, rounded and lightly 
emarginate at base, shortly and obtusely acuminate, 7-10 cm. long, 
4.5-5 cm. broad; panicles broadly pyramidal, 6-8 cm. long, the 
branches and branchlets divaricate, the 5-6-merous flowers on pedi- 
cels 1-3 mm. long; calyx broadly campanulate, the tube 4 mm. 
across, the small teeth remote at base, acute, 1 mm. long; petals 
white, obliquely obovate, fleshy, 2.5 mm. long; anthers arcuate, 1.5 
mm. long; style glabrous, clavate, 3 mm. long, the stigma peltate. 
Affine M. rigida Triana. Section Chaenopleura. F.M. Neg. 17159. 

Amazonas: East of Chachapoyas, 2,200 meters, Weberbauer 4427, 
type. 

Miconia semisterilis Gleason, Bull. Torrey Club 63: 535. 1936. 

Upper branches acutely 4-angled and 2-sulcate as also the thick 
angled petioles these 1 cm. long and densely ashy-lepidote like 
the leaves beneath and the angled panicle branches even to the 
calyces, these sessile, 5-merous, scarcely lobed, about 1.5 mm. long; 
leaves narrowly elliptic-acuminate at both ends, often 17 cm. long, 
6 cm. wide, glabrous and opaque above, papyraceous, 5-plinerved, 
the outer pair submarginal; panicle 8-10 cm. long; petals white, 
glabrous, unequally truncate, 1.8 mm. long, 1.6 mm. wide; stamens 
nearly isomorphic, the slender filaments 1.6 mm. long, the anthers 
about as long, sterile below the middle, somewhat obovate in the 



FLORA OF PERU 453 

terminal portion, the connective produced into 2 minute lateral 
lobes and 1 (rarely 2) narrowly oblong dorsal appendage; ovary 
free and glabrous above, apparently 3-celled; style stout, about 2.5 
mm. long, glabrous, the stigma truncate. The scales are brownish 
on the stems, ashy on the leaves. Suggests M. Klugii, in which the 
anthers are fertile nearly to base; the anthers are similar in form and 
sterile for more than half their length in M. Pilgeriana and M. 
stellipilis Cogn., Bull. N. Y. Bot. Gard. 4: 360. 1907, also with 
similar appendages to the connective but these lack the sharply 
angled stems and the lepidote indument; all three, as M. semi- 
sterilis, belong to section Glossocentrum (Gleason). 

Loreto: Pumayacu, in forest, small tree 8 meters high, Klug 
3224, type. 

Miconia serialis DC. Prodr. 3: 182. 1828; 790. 

Subterete branches somewhat, the leaves beneath thickly, rusty- 
floccose-tomentose; petioles 1-2 cm. long; leaves elliptic-oblong to 
broadly elliptic-ovate, shortly attenuate at base, acutely short- 
acuminate, in age glabrous and lustrous above, 5-nerved or some- 
times shortly 3-plinerved, 1-2 dm. long, 4-8 cm. wide; panicles broad, 
to 1.5 dm. long, the branches bifid or trifid, the flowers secund, their 
petals obliquely retuse, 1.5-2 mm. long; calyx 2 mm. long; connective 
strongly dilated, basally bilobed anteriorly, tuberculate posteriorly; 
connective produced; style about 3 mm. long; fruit 10-sulcate, 3-3.5 
mm. thick. Common in Loreto. Like several other species, rich in 
tannin. Section Eumiconia. F.M. Neg. 17164. 

San Martin: Tocache, Poeppig 1863; Moyobamba, Weberbauer 
4531 (det. Cogn.); 290. Puno: San Gaban, Lechler 2467. Loreto: 
Iquitos, Tessmann 5391; Williams 1454; 1508; 3631; Killip & Smith 
29857; Klug 36; 465. Rio Mazan, Schunke 158 (det. Standl.). 
Yurimaguas, Williams 3836; 4718; Killip & Smith 27948; Poeppig 
2407b. Bolivia to Venezuela. "Carachi-caspi," "canela de velha." 

Miconia serrulata (DC.) Naud. Ann. Sci. Nat. se>. 3. 16: 118. 
1851; 734. Diplochita serrulata DC. Prodr. 3: 177. 1828. Chitonia 
macrophylla D. Don, Mem. Wern. Soc. 4: 319. 1823. M. macrophylla 
Triana (DC.) Cogn. in Mart. Fl. Bras. 14, pt. 4: 239. 1887. 

Nearly the same as M. bubalina, probably not distinct, but the 
rounded or lightly emarginate leaves 5-7-nerved and sometimes 
glabrate beneath; style 11-13 mm. long. The var. Hostmanni 
(Steud.) Cogn. has oblongish 5-nerved leaves, subabruptly acuminate. 
Section Tamonea. 



454 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Junin: Near Peren Bridge, Kittip & Smith 25297 (det. Gl.). 
San Martin: San Roque, Williams 7793 (det. GL). Loreto: Iquitos, 
Tessmann 3596; Williams 1481; 3647, the var. (det. Markgr.). 
Mouth of Santiago, Tessmann 4510 (glabrate form, det. Markgr.). 
Florida, shrub with lilac and white or white flowers, Klug 2336; 
2261 (det. GL). Pro, Williams 1996. La Victoria, Williams 2770. 
Yurimaguas, Williams 4569. Pebas, Williams 1629; 1631. Rio 
Mazan, Schunke 4 (det. Stand!.). Puno: Lechler 2115. Bolivia to 
the West Indies. "Difari de hoja ancha," "sesepe-o." 

Miconia setinervia Cogn. Bot. Jahrb. 42: 143. 1908. 

Branches obscurely quadrate, very shortly and rather sparsely 
setulose as the leaves beneath, principally on the nerves and veins, 
otherwise glabrous or the calyx obscurely puberulent; petioles 4-6 
mm. long; leaves lanceolate, acute at both ends, entire, 5-nerved, 
rigid but extremely fragile in the herbarium, drying black above, 
paler beneath, with numerous transverse veins, 5-7 cm. long, 1.5- 
2.5 cm. broad; panicle pyramidal, 6-10 cm. long; calyx campanulate- 
urceolate, 1 mm. long, the short, broadly ovate teeth acute; petals 
whitish, obovate, scarcely more than 0.5 mm. long; filaments capil- 
lary, glabrous, as the filiform style, this 1.5 mm. long, the stigma 
peltate. Allied to M. cyanocarpa Naud. Section Cremanium. 
F.M. Neg. 17160. 

Puno: Vicinity of Tambo Yuncacoya, between Sandia and 
Chunchusmayo, 1,800 meters, Weberbauer 1145, type. 

Miconia setulosa Cogn. Melast. 898. 1891. 

Stoutly branched, the branches obtusely tetragonous, the younger, 
as the petioles and panicles, long-setose-hirsute, the petioles 3-6 mm. 
long; leaves lanceolate, basally subacute, obtusish, spinulose-denticu- 
late, rigid, glabrous above, sparsely setose only on the 5 nerves 
beneath, 6-8 cm. long, 18-22 mm. broad; panicle thyrsoid, 5-10 cm. 
long, the usually 5-merous flowers crowded; pedicels 1-2 mm. long; 
calyx glabrous, minutely dentate, 3 mm. long; petals 2 mm. long; 
anthers bipored; style 3 mm. long, apically truncate; fruit black, 
3.5-4 mm. thick. Section Cremanium. 

Puno: Tatanara (Lechler 2637, type). 

Miconia spennerostachya Naud. Ann. Sci. Nat. se>. 3. 16: 
187. 1851; 801. M. nectaria Macbr. Field Mus. Bot. 4: 190. 1929. 
M. Aspiazui Macbr. I.e. 

Similar to M. pauciglandulosa but usually at once separable from 
that and other species by the presence of one to several cup-like 



FLORA OF PERU 455 

appendages, usually somewhat cone-shaped at base on the petioles 
and leaf bases above; petioles sometimes several cm. long, the leaves 
often shortly decurrent on them; flowers subsecund; calyx limb 
indistinctly lobed; connective not or scarcely produced, eglandular; 
petals obliquely subtruncate, 2 mm. long. The nectaries were not 
previously described. Section Eumiconia. F.M. Neg. 26030. 

Junin: La Merced, Hacienda Schunke, slender tree, 5738 (type, 
M. nectaria). La Merced, 5586 (type, M. Aspiazui). San Martin: 
Moyobamba, Mathews 1305, type. Pongo de Cainarache, 5-meter 
tree, the flowers white, King 2620 (det. Standl.). Loreto: Pebas, 
Williams 1686 (det. Gl.). Yurimaguas, Kittip & Smith 28131 (det. 
Gl.). Bolivia. 

Miconia stelligera Cogn. in Mart. Fl. Bras. 14, pt. 4: 275, 
pi 56. 1887; 778. 

Allied to M. barbinervis but often less pubescent, the thin leaves 
glabrate above, to 2.5 dm. long, 1 dm. wide; calyx shortly stellate- 
hirtellous and petals stellate-scurfy without, subrounded or apicu- 
late; style 7-8 mm. long, the stigma truncate. The stalked stellate 
pubescence is present in greater or less degree but mostly on the 
branches and petioles; leaves, except for the 3 principal nerves 
beneath, often glabrate. Section Eumiconia. F.M. Neg. 17171. 

San Martin: Pongo de Cainarache, 2.5 meters high, Klug 2629 
(det. Gl.). Junin: Chanchamayo Valley, Schunke 398. Loreto: 
Yurimaguas, Poeppig 2377 pt.; Ule 6265 (det. Pilger). Brazil. 
"Sira sira," "mullaca." 

Miconia stellipilis Cogn. Bull. N. Y. Bot. Gard. 4: 360. 1907. 

Younger branches, petioles and peduncles more or less densely 
stellate-puberulent; petioles 5-12 mm. long; leaves ovate-oblong, 
firm, rounded at base, caudately acuminate, entire, glabrous above 
at maturity, minutely stellulate on the 5 nerves and on the veins 
beneath or apparently glabrate, 8-16 cm. long, 3.5-6.5 cm. wide; 
panicles mostly branched; calyx campanulate, stellate-puberulent 
typically, in the Peruvian plant nearly glabrous, 1 mm. long and 
broad, sessile; petals ovate, obtuse, 1 mm. long; anthers as in M. Pil- 
geriana. The following collection may be distinct from the Bolivian 
type, not seen, which is, according to description, more pubescent, 
the trichomes papillose-stellate as in the apparently closely related 
M. dolichorrhyncha Naud. 

Cuzco: Marcapata Valley near Tio, 1,900 meters, Weberbauer 
7852. Bolivia. 



456 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Miconia stenostachya [Schrank & Mart.] DC. Prodr. 3: 181. 
1828; 789. Melastoma stenostachyum Schrank & Mart, ex DC. I.e. 
in syn. 

Near M. argyrophylla, the similarly bicolored leaves (green above, 
white appressed-lanate beneath) merely acute or shortly and obtusely 
acuminate, obscurely undulate-denticulate, and the connective not 
obviously produced if at all, minutely tubercled posteriorly, slightly 
bilobed anteriorly. The calyx becomes 10-costate, similar to 
M. albicans, differing in the connective. Section Eumiconia. 

Huanuco: Chinchao, Ruiz & Pavdn. Casapi and Cuchero, 
Poeppig 1437. San Martin: Near Tarapoto, Ule 6458 (det. Pilger); 
Williams 5423; 5629; 5678; 6330; 6419; 6427 (det. Gl.). Valley of 
Monzon, Weberbauer 3475 (det. Cogn.); 284. Near Moyobamba, 
1-meter shrub, the flowers yellow, Klug 3265 (det. Gl.). Ayacucho: 
Killip & Smith 22705 (det. Gl.). Bolivia to Mexico and the West 
Indies. "Caracha caspi." 

Miconia subglabra Cogn. in Mart. Fl. Bras. 14, pt. 4: 412. 
1888; 885. 

Branches slender, subterete, as the petioles and peduncles, 
minutely or granularly stellate-puberulent; petioles 1-1.5 cm. long; 
leaves oblong-elliptic, lanceolate, subrotund at base, shortly acumi- 
nate, rather firm, glabrous (or the midnerve scurfy above) except 
for a few scattered setae on the nerves beneath, 5-plinerved, the outer 
2 nerves fainter or the nerves sometimes nearly basal, 1-1.5 dm. 
long, 3-4.5 cm. broad; flowers glomerate, sessile, the rusty puberulent 
panicles 7-10 cm. long; calyx distinctly lobed, 3 mm. long; petals 
suborbicular, 2 mm. long; filaments and style eglandular, the latter 
4-5 mm. long, the stigma subpeltate; connective not produced 
anteriorly, subbicalcarate. The leaves dry pale; the calyx is thickly 
granular-scurfy. Section Amblyarrhena. F.M. Neg. 17174. 

San Martin: Tarapoto, Spruce 4842, type. 

Miconia Terera Naud. Ann. Sci. Nat. s<r. 3. 16: 196. 1851; 875. 

Completely glabrous and similar to M. pedicellata; leaves elliptic, 
shortly acuminate, thin but firm, 1.5-2 dm. long, 6-7 cm. broad; 
petioles 1-3 cm. long; calyx (2) 3-4 mm. long, shortly and obtusely 
5-lobed; petals 2-2.5 mm. long, oblong-elliptic or slightly obovate, 
retuse; style 3 mm. long, scarcely clavate; anthers 1.5 mm. long. 
The Junin specimens, with only blue fruits, are not surely this 
species. Tree 3-5 meters high. Section Amblyarrhena. F.M. Neg. 
29506. 



FLORA OF PERU 457 

Huanuco: Chicoplaya, Ruiz & Pavon, type; Dombey; (Lechler 
2526). Junin: Vitoc, Isern 234.2. La Merced, 700 meters, Killip 
& S