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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,