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Full text of "Flora of Peru"

THE UNIVERSITY 

OF ILLINOIS 

LIBRARY 




OCT 7 1939 



FLORA OF PERU 



BY 

J. FRANCIS MACBRIDE 

ASSOCIATE CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 



V 

N NATURAL 

HISTORY 




THE LIBRARY OF THE 

NOV 191938 

UNIVERSITY OF ILLINOIS 

BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XIII, PART II, NUMBER 3 

OCTOBER 31, 1938 

PUBLICATION 428 



FEINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 



FLORA OF PERU 



J. FRANCIS MACBRIDE 



70. BERBERIDACEAE. Barberry Family 

The importance of this family in Peru centers about Berberis lutea 
and relatives, the "ccarhuascassa" or "palo amarillo," sometimes 
called "espino amarillo," whose yellow wood, with that ofBocconia, 
furnishes the beautiful, yellow dye seen in ponchos and other clothing 
to this day (Ruiz & Pavdri). 

1. BERBERIS L. Barberry 

Reference: Schneider, Bull. Herb. Boiss. II. 5: 33^*8, 133-148, 
391-403, 449-464, 655-670, and 800-831. 1904-1905. 

Shrubs with or without spines, these typically trifurcate. Leaves 

, simple in the Peruvian species, fasciculate, entire or more or less 

l aculeate-dentate. Flowers yellow or reddish yellow, variously dis- 

r- posed, with 3-4, most often 6, deciduous sepals and usually as many 

petals, these biglandular within. Filaments as many as the petals, 

. dentate or edentate. Ovules 1-12, laterally inserted at the base of 

< the 1-celled ovary, the fruit 2-3-seeded. Schneider has given a 
' careful and discriminating account of the Andean forms which, 
5 however, with more knowledge, probably will be found to constitute 

fewer species. For the purposes of this work it has seemed practical 

to follow earlier authors and group the forms primarily according to 

y> the nature of the inflorescence, which for well developed specimens 

^ of nearly all species is quite obvious. It may be stated that the work 

^ of P. E. Citerne, cited several times on the following pages, is a 

thesis of the Faculty des Sciences de Paris, Se"rie B, No. 183. 1892, 

very rare and sometimes filed in libraries under this citation. 

v? Omitted from the following synopsis is the name B. rotunda 

~ Macbr. Candollea 6: 3. 1934, which, as casual observation of the 

type has shown, has nothing in common with this family except 

; a certain similarity in foliage. Because of this, the specimen had 

w been laid in a cover with species of Berberis described the same 

< evening and perhaps flowers of Berberis became mixed with the 
* aberrant sheet. 

Flowers racemose (sometimes very shortly so), racemose-corymbose, 

or paniculate. 

^ Flowers borne in more or less peduncled panicles. 

665 



666 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves oblong-elliptic or obovate. 
Leaves distinctly paler beneath, often glaucous-pruinose. 
Leaves papillose beneath, the veins scarcely obvious. 

B. brumalis. 

Leaves epapillose, finely reticulate-veined on both sides. 
Leaves (beneath) and panicles puberulent. 

B. Hochreutinerana. 
Leaves and panicles glabrous B. Barbeyana. 

Leaves concolorous, not at all paler beneath, or at least 
lustrous on both sides. 

Leaves mostly 2-4 cm. wide. 
Leaves mostly spinose-denticulate; panicles glabrous. 

B. Beauverdiana. 
Leaves entire or sparsely spinulose; panicles typically 

granular-puberulent. 
Leaves obovate, often 4-5 cm. wide; panicles 4-5 cm. 

broad B. multiflora. 

Leaves oblong-elliptic, about 2 cm. wide; panicles 

strict, narrow B. paniculata. 

Leaves mostly 1-1.5 cm. wide B. loxensis. 

Leaves rotund or more or less 4-angled, dentate. 
Leaves epapillose, elevated-reticulate above, truncate at the 

base B. podophylla. 

Leaves papillose beneath, obscurely reticulate-veined above, 

cordate at the base B. buceronis. 

Flowers racemose or variously disposed but not obviously in 

peduncled panicles (cf. B. brumalis and B. loxensis), 
Leaves distinctly subrotund or subquadrate, epapillose or sub- 

epapillose beneath. 
Leaves spinulose-dentate, epapillose, lustrous above. 

B. Gayi. 
Leaves spinulose-dentate, subpapillose, dull on both sides. 

B. armata. 

Leaves entire, papillose above B. Weberbaueri. 

Leaves distinctly longer than broad. 

Leaves typically entire (exceptions seem to concern only an 
occasional leaf). 



FLORA OF PERU 667 

Venation densely and finely reticulate. 

Leaves ashy-pruinose beneath B. Jelskiana. 

Leaves concolorous. 

Leaves 3-4 cm. long, papillose B. monosperma. 

Leaves 1.5 cm. long or shorter, epapillose. 

B. boliviano,. 

Venation lax, if reticulate, as in B. flexuosa at maturity, 
neither continuously nor closely netted. 

Inflorescence glabrous, glaucous; pedicels 10-15 mm. long. 

B. commutata. 

Inflorescence more or less granular-glandular or puberu- 
lent; pedicels shorter. 

Leaves epapillose; spines none or reduced. 

B. Lobbiana. 

Leaves papillose, especially above; spines well devel- 
oped B. flexuosa. 

Leaves characteristically serrate or spinulose. 
Leaves longer than 1 cm. 

Leaves to 1.5 cm. wide, apically dentate, epapillose. 

B. Lobbiana. 
Leaves mostly wider, usually dentate below the middle, 

papillose. 
Venation densely reticulate; leaves sparsely spinulose- 

dentate B. peruviana. 

Venation loose, yellowish; leaves spinescent-serrate. 

B. dryandriphylla. 

Leaves small, rarely 1 cm. long, apically 3-angled-spinulose 
or entire B. Humbertiana. 

Flowers solitary or fasciculate (cf. B. brumalis). 
Leaves about as broad as long. 

Branchlets spinose, pubescent; leaves closely reticulate- veined 

beneath B. agapatensis. 

Branchlets glabrous, not spinose; venation lax, faint. 

B. Benoistiana. 
Leaves distinctly longer than broad. 

Leaves rather large, usually wider than 1 cm., longer than 2 cm. 
Leaves epapillose, to 3 cm. long, 13 mm. wide. . B. Lobbiana. 



668 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves papillose beneath, often larger. 

Leaf margin entire, the leaves chartaceous B. latifolia. 

Leaf margin spinose, the leaves coriaceous. 
Pedicels 6-10 mm. long, glabrate; leaves regularly spinose- 

dentate B. saxicola. 

Pedicels 4-5 mm. long, puberulent; leaves apically 

spinose-dentate B. carinata. 

Leaves mostly small, scarcely 10 mm. wide except sometimes in 

B. lutea var. conferta, rarely longer than 2 cm. 
Leaves evidently glaucous beneath. 

Leaves broadly obovate, about 2 cm. long, 1 cm. wide. 

B. carinata. 

Leaves oblanceolate, 5-8 mm. wide B. huanucensis. 

Leaves concolorous or nearly so, at least when dry. 
Leaves conspicuously callous-margined, papillose. 

B. cliffortioides. 

Leaves merely margined, epapillose unless obscurely so 
near the petiole, or waxy. 

Pedicels glabrous, 5-8 mm. long B. virgata. 

Pedicels typically granular-puberulent B. lutea. 

Berberis agapatensis Lechl. Berb. Amer. Austr. 44. 1857; 148. 

A shrub about 1.5 meters high, with divaricate, soon terete, 
slightly pubescent branchlets; nodal trifid spines 6-8 mm. long; 
leaves subcordate and entire at the base, dentate above the middle, 
2-2.5 cm. long, about as wide, glabrous, coriaceous, lustrous above, 
papillose, glaucous and conspicuously reticulate- veined beneath; 
petioles slightly pubescent, to 2 cm. long; pedicels 3-7 and 1-2 cm. 
long; filaments edentate; berry 8-10-seeded, the seeds oblong-obo- 
vate, angled. Fruit purple-violet, "twice as large as that of the 
common jumper." The type grew in an argillaceous schist. Neg. 
34505. 

Puno : Agapata to Sachapata, Lechler 2646, type. Bolivia. 

Berberis armata Citerne, Berber, et Erythrosp. 152. 1892; 813. 

Spines 3-5-furcate, to 5 cm. long; branchlets flexuous, smooth, 
whitish yellow; leaves in 2's or 3's, attenuate or subrounded at the 
base, 3-4 cm. long, 2.5-3.5 cm. wide, coriaceous, essentially epapil- 
lose, that is, subpapillose only beneath, dull, obscurely nerved on 
both sides, undulate and 5-6-spinulose-dentate or serrate, the 



FLORA OF PERU 669 

sounded tip mucronate; racemes simple or compound, few-flowered, 
shorter than the leaves, granular-puberulent; pedicels 4-6 mm. long; 
style distinct; flowers about 6 mm. broad, the crenulate sepals about 
as long as the petals. Negs. 14286, 34506. 

Cuzco(?):Ga2/, 1839-1840. Without locality: Dombey. 

Berberis Barbeyana Schneid. Bull. Herb. Boiss. II. 5: 816. 1905. 

Branchlets terete; spines none or short, rarely 1.5 cm. long; 
blades elliptic-oblong, 2-4.3 cm. long, 1.3-2 cm. wide, coriaceous, 
rarely with 1 or 2 teeth, distinctly whitish beneath, not papillose; 
petiole 3-8 mm. long; panicles 6-14 cm. long; pedicels 7-11 mm. 
long; bracts about 3 mm. long; flowers 9 mm. broad; stamen connec- 
tive scarcely obvious; style short; ovules 4. Neg. 27404. 

Amazonas: Chachapoyas, Mathews, type. 

Berberis Beauverdiana Schneid. Bull. Herb. Boiss. II. 5: 
817. 1905. 

Branchlets strongly sulcate, yellowish brown; spines weak, 5-7- 
parted, scarcely 8 mm. long; leaves elliptic or oblong, usually short- 
dentate, coriaceous, lustrous on both sides, little paler beneath, not 
distinctly papillose, 3-5 cm. long, 1.7-2.5 cm. wide; panicles to 9 cm. 
long, glabrous; pedicels 5-6 mm. long, the bracts about half as long; 
stamen connective obvious; style obsolete; ovules 2. B. Jamesonii 
Lindl. Journ. Hort. Soc. 5: 8. 1850, accredited to Peru by Schneider,' 
is from Ecuador; it is distinguishable by its roundish, brown-red 
branchlets, and pedicels 10-15 mm. long. Negs. 27405, 14287. 

Amazonas: Chachapoyas, Mathews, type. Cajamarca: Huama- 
lies, Weberbauer 1+199 ("a variety?; leaves entire and pedicels longer," 
Schneider in herb.). 

Berberis Benoistiana Macbr., sp. nov. 

B. agapatensis Lechl. similis, glabra; foliis ellipticis basi fere 
rotundatis infra medium usque ad apicem sparse spinuloso-dentatis, 
ad 4 cm. longis, 3 cm. latis, subtus perobscure nervatis haud reticu- 
latis perdense papillosis; sepalis fere 6 mm. longis; petalis ca. 4 mm. 
longis. At one time referred to Lechler's species of southern Peru, 
and obviously closely allied, but that shrub is somewhat puberulent 
and the strongly reticulate-veined leaves are relatively less papillose. 
The papillae on the under leaf surface of the new species are so 
abundant that they seem to form a tissue. Moreover, the habitat 
is strikingly different. B. Benoistiana was found on a fog-drenched 
summit in open, mossy places. A slender shrub, 1 meter high, the 



670 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

flowers bright yellowish red, the fruits black. Named for Dr. R. 
Benoist of the Natural History Museum, Paris, whose discriminating 
studies on the related flora of Ecuador may be commemorated here. 
Huanuco: Tambo de Vaca, 4,000 meters, Macbride 4455 (type in 
Herb. Field Mus.). 

Berberis boliviana Lechl. Berb. Amer. Austr. 21. 1857; 393. 
B. conferta HBK. var. boliviana Schneid. Bull. Herb. Boiss. II. 5: 
393. 1905. 

A branched shrub, the conspicuously spiny branchlets often arcu- 
ately spreading, little flexuous; spines slender, straight, spreading, 
usually trifid, about 15 mm. long; leaves 5-8-fasciculate, obovate- 
cuneate, sessile or subsessile, pale green, concolorous, entire, thinly 
callous-margined, subcoriaceous, mostly 12 mm. long and 5 mm. 
wide, finely and conspicuously reticulate-veined on both sides, epa- 
pillose; flowers in short racemes or subcorymbs, these 1-1.5 cm. long; 
pedicels obscurely but obviously puberulent, 2-3 mm. long; open 
flowers 4-5 mm. broad, the obovate sepals and petals subequal, 
about 3 mm. long; stigma large, sessile. Type collected by Pentland 
in Bolivia, near the Peruvian boundary. B. Keissleriana Schneid., 
808, of Bolivia has linear-spatulate leaves, definitely papillose 
beneath; B. Rechingeri Schneid., 808, similar, has broader, epapillose 
leaves, and simple or no spines. Neg. 34507. 

Cuzco: Pachar, 2,900 meters, bushy bank, Pennell 13690; fruit 
blue, glaucous. At 3,200 meters, Herrera 3028. Cuzco, Soukup 577. 
Bolivia. "Chehejche." 

Berberis brumalis Macbr. Candollea 6: 4. 1934. 

Glabrous, the long, yellowish green, nearly terete branches 
unarmed; leaves 5-7-fasciculate; petioles short-dilated at the base, 
strongly canaliculate, about 7 mm. long, articulate below the elliptic- 
oval blade, this about 4 cm. long and 2 cm. wide, rounded-acute 
at the base, spinulose-cuspidate at the apex, the revolute margin 
entire or with 2-3 remote, spinulose teeth, coriaceous, lustrous and 
obscurely reticulate- veined above, dull, paler, scarcely veiny but 
papillose beneath; pedicels 1 mm. long, glabrous; sepals and petals 
scarcely 3.5 mm. long, minutely ciliate; stamens apparently almost 
2 mm. long, the connective strongly produced. The type has no 
inflorescence, but has one dissected flower. The position of the 
species now appears to me to be doubtful. In leaves it simulates 
species with fasciculate flowers, but it matches no species seen by me. 

Peru: Without locality, Weberbauer 6799, type. 



FLORA OF PERU 671 

Berberis buceronis Macbr. Candollea 6: 3. 1934. 

A spiny shrub with flexuous, terete, glabrous branchlets and 2- 
3-fid spines, these 1.5 mm. thick at the base, 7 mm. long, strongly 
curved; petioles slender, 1-2.5 cm. long, minutely puberulent, about 
5-fasciculate; leaves subrotund, usually deeply cordate at the base, 
3-4 cm. long and broad, chartaceous, slightly lustrous above but 
scarcely veiny, dull, papillose, and prominently reticulate-veined 
beneath, undulate-spinulose-dentate; flowers in peduncled corymbs 
or subpaniculate, granular-pulverulent, 5 mm. long; pedicels to 
10 mm. long; stamens 3.5 mm. long; connective obtuse; stigma 
capitate. 

Peru : Without locality, Weberbauer 6994, type. 

Berberis carinata Lechl. Berb. Amer. Austr. 41. 1857; 394. 

A shrub 1 meter tall or taller, with terete, divaricate, unarmed, 
rusty- tomentose branchlets, glabrate in age; leaves cuneate-obovate, 
carinate-cartilaginous on the margins, the apex strongly 5-6-spinose- 
dentate, 2-2.5 cm. long, 8-12 mm. wide, lustrous above, conspicu- 
ously glaucescent and papillose beneath; pedicels somewhat shorter 
than the leaves; flowers nodding, the sepals 5, subrotund, dull 
yellow, the longer, yellow petals obovate, acuminate; fruit ovate, 
"lageniform." Var. echinata Diels, Notizbl. Bot. Gart. Berlin 11: 
782. 1933, has hard leaves with spines 2-2.5 mm. long. B. tomentosa 
R. & P. of Chile has larger leaves, pubescent beneath; Schneider and 
Lechler in error ascribe it to Peru. Neg. 34510. 

Puno: In schist, Agapata, Lechler 2644, type. Cuzco: Hills of 
Sacsahuaman, 3,500 meters, Herrera, type of the variety. Bolivia. 

Berberis cliffortioides Diels, Notizbl. Bot. Gart. Berlin 11: 
782. 1933. 

Young branches purple-glaucescent; spines 1-3-fid; leaves 
crowded, almost sessile, fleshy-coriaceous, narrowly oblanceolate, 
entire, acutely spinescent at the apex, with conspicuous, thick 
margins, 7-15 mm. long, 2-3.5 mm. wide, papillose and weakly 
nerved beneath, the midrib prominent; peduncle 6-10 mm. long, 
puberulent; sepals papillose-puberulent, the inner 4 mm. long, pale 
yellow. Noted by the collector as 30 cm. high. Contrasted by the 
author with B. rectinervia Rusby, Mem. Torrey Club 3, pt. 3: 5. 
1893, a glabrous plant of Bolivia, and apparently, with that, too 
near B. lutea, sens. lat. 

Junin : Puna in Cordillera de Punto in via a Huancayo ad Collor- 
bamba (Raimondi, type). 



672 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Berberis commutata Eichl. in Mart. Fl. Bras. 13, pt. 1: 231. 
1864; 804. B. divaricata Rusby, Mem. Torrey Club 4: 203. 1895. 

A glabrous shrub with light-colored, elongate branches, more or 
less well developed, trifid, yellowish spines, and yellowish green 
leaves, their nerves and veins rather prominent on both sides; leaves 
oblong-obovate, cuneate to the base, apiculate at the acutish to 
broadly rounded apex, entire or with 1-2 remote, weakly spinulose 
teeth, subcoriaceous, lustrous above, paler, dull, and papillose 
beneath, typically about 2 cm. wide, 6 cm. long, or half as long and 
as wide, or somewhat narrower; racemes more or less nodding, 
slightly shorter or longer than the leaves, the pedicels soon 10-15 mm. 
long; flowers 5 mm. long; style well developed, the stigma prominent. 
Weberbauer, 178, has recorded the species from the Department 
of Junin at 3,300 meters. Neg. 34513. 

Apurimac: Abancai (Weberbauer 5911; det. Schellenberg). 
Huanuco: Dunkafael, Sawada 78. Ayacucho: Hacienda Totora- 
bamba (Weberbauer 5485; det. Schellenberg). Cuzco: Gay 2191. 
Valle del Urubamba, Herrera 3395. Prov. Calca, Hammarlund 609. 
Pelechuco (Peru, fide Rusby), type of B. divaricata (Pearce). 
Bolivia. "Tancar." 

Berberis dryandriphylla Diels, Notizbl. Bot. Gart. Berlin 11: 
781. 1933. 

Glabrous, with trifid spines and black branchlets; blades coria- 
ceous, obovate-oblong, coarsely spinescent-serrate, 3-6.5 cm. long, 
1.5-2.5 cm. wide, lustrous above, papillose beneath and with light 
yellow nerves; petioles 2-8 mm. long; racemes sessile, lax, to 5 cm. 
long, the purple pedicels 1 cm. long; sepals elliptic, 5-6 mm. long, 
3-5 mm. wide, the petals nearly as large; stamens 3.5 mm. long, the 
connective obtuse; ovary 3 mm. long; style obsolete; ovules 2. 
Compared by the author withB. pichinchensis Turcz., with pubescent 
racemes. B. Trollii Diels, of Bolivia, has leaves 2-3 cm. wide, 
racemes 1.5-2 cm. long, and pedicels 5-6 mm. long. 

Cuzco: Valle del Forachtay, 2,150 meters (Herrera 1742). 

Berberis flexuosa R. & P. Fl. Peruv. 3: 52. pi. 281. 1802; 812. 

A densely branched shrub 2 meters high; branchlets slender, 
virgate, little angled, distinctly flexuous; spines slender, trifid, 1-3 
cm. long; leaves elliptic-obovate, acute at the short petiole, entire 
or with 2-3 teeth near the apex, becoming about 3 cm. long, 1.5 (-2) 
cm. wide, at maturity loosely reticulate-veined on both sides, papil- 
lose, especially above, paler and dull beneath; racemes (2-) 4-6 cm. 



FLORA OF PERU 673 

long, rather densely but obscurely puberulent; bractlets, even in age, 
puberulent, sessile, ovate-aristulate; pedicels finally 5-8 mm. long; 
flowers pale yellow, scarcely 5 mm. long; stamen connective pro- 
duced; style obvious; ovules 3-4; young berry lageniform. There 
may be two forms on the type sheet at Madrid, although all the 
material is referred by Diels to the type without question. The 
smaller branchlet has obovate leaves only 1.5 cm. long and 7 mm. 
wide, faintly veined, but apparently young; inflorescence 4-flowered, 
almost glabrous. The stem is very flexuous, as on the other branch, 
which doubtless is to be taken as the exact type. Neg. 12988. Ruiz 
and Pavon gave Tarma, Cheuchin, and Pillao as localities. The 
Paris Dombey material is much better. B. papillosa Benoist of 
Ecuador, which might be sought here, has larger leaves lustrous 
above, white-glaucescent beneath, and spinulose-dentate toward the 
apex. B. Rechingeri Schneid., 808, of Bolivia, has simple spines and 
leaves only half as large. My No. 1285, distributed as B. mono- 
sperma, had dark blue fruit used locally to make ink. 

Lima: Cheuchin, Ruiz & Pav6n. San Rafael, river canyon slopes, 
2,500 meters, 3133. Junin: Tarma, Dombey; Weberbauer 1726. 
Chinche, Yanahuanca, on grassy slope, 1285. Without locality, 
Haenke. "Haucampe," "huajanpe." 

Berberis Gayi Citerne, BerbeY. et Erythrosp. 152. 1892; 670. 

Spines trifid, scarcely 2 cm. long; branchlets straw-colored, 
glabrous; leaves elliptic to subrotund, about 3.5 cm. long and broad, 
or smaller, especially somewhat narrower, coriaceous, smooth and 
lustrous above, almost dull beneath, faintly reticulate-veined on both 
sides, epapillose, with about 16 spinulose teeth, short-petiolate, 
the base rounded-acute; racemes spreading, 2-4 cm. long, usually 
simple; bractlets subulate-acuminate, puberulent; fruiting pedicels 4 
mm. long; style obvious; stigma large; fruit lageniform. The fruit- 
ing racemes, except for the bractlets, are glabrous, but the flowers are 
unknown. Neg. 34521. 

Cuzco(?):Ga# 1164, type. 

Berberis Hochreutinerana Macbr. Candollea 6: 2. 1934. 

Spines trifid, to 7 mm. long, barely 1 mm. thick at the base; 
branchlets glabrous, subterete; leaves about 7-fasciculate, the blades 
oblong-oblanceolate, mostly 7-8 cm. long and 3 cm. wide, rounded 
at the apex, cuneate-attenuate to the petiole, entire, chartaceo- 
coriaceous, epapillose, lustrous and slenderly reticulate- veined above, 
paler and dull beneath, the rather prominent nerves pulverulent; 



674 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

petioles to 1 cm. long, articulate at the persistent sheath, this some- 
times nearly 1 cm. long; panicles many-flowered, solitary, distinctly 
puberulent, with spreading branchlets, to 15 cm. long; bracts 3-4 
mm. long; pedicels rarely nodding, to 7 mm. long; sepals to 5.5 mm. 
long, exceeding the obovate petals; stamens 3.5 mm. long, the con- 
nective little produced; style prominently capitate-dilated. 
Cuzco: Mount Acomayo, 3,100 meters, Weberbauer 6831. 

Berberis huanucensis (Schneid.) Macbr., comb. nov. B. glauca 
R. & P. in herb., the name untenable. B. virgata R. & P. var. 
huanucensis Schneid. Bull. Herb. Boiss. II. 5: 391. 1905. 

Similar to B. lutea, to which it could be referred as a variety, 
but at least the younger leaves conspicuously glaucous beneath; 
spines none, or trifid and rarely 9 mm. long; leaves to 2.5 cm. long, 
5-8 mm. wide, often 3-5-spinulose-dentate at the apex, densely 
granular-papillose beneath and above but the papillae often hidden 
by glaucous wax; bracts slightly ciliate; pedicels 1.5-3 cm. long, 
glabrous. Negs. 14313, 29227. 

Huanuco: Ruiz & Pavon, type. Chiclin, Sawada 85. "Chijchi." 

Berberis Humbertiana Macbr., sp. nov. 

Frutex glaberrimus; ramis vel ramulis virgatis haud flexuosis 
teretibus; spinis trifidis sat validis ca. 5 mm. longis; foliis fasciculatis 
sessilibus cuneato-obovatis apice 3-angulato-spinuloso-dentatis vel 
interdum lanceolatis, fere integerrimis et acuto-mucronatis, cartilagi- 
neo-marginatis, rigido-coriaceis, supra paullo nitidulis, subtus opacis 
prominente papillosis obscure laxeque reticulatis, 8-10 mm. longis 
4-5 mm. latis; racemis 6-15 mm. longis; pedicellis 2-2.5 mm. longis; 
floribus 4-6; sepalis subrotundatis cum petalis vix 3 mm. longis; sta- 
minibus obscure apiculatis; stylis nullis; fructibus globosis. With 
a superficial resemblance to B. lutea and its allies, this shrub is 
closely related to B. Keissleriana Schneid., 808, as remarked by 
Schneider on the type sheet, but that Bolivian plant is puberulent, 
has definitely petioled, narrower leaves, and more numerous flowers. 
B. Wettsteiniana Schneid., 809, similar, has much larger leaves, 
racemes to 4 cm. long, the flowers 6 mm. broad, on pedicels 3-5 mm. 
long. Citerne had given the Peruvian plant a herbarium name 
without, however, indicating the characters. It may commemorate 
the furtherance of botanical science by Professor H. Humbert, 
Director of the botanical museum, Museum of Natural History, Paris. 
Neg. 34532. 

Cuzco (?): Gay, type in herb. Paris. Andes of Peru, Castelnau. 



FLORA OF PERU 675 

Berberis Jelskiana Schneid. Bull. Herb. Boiss. II. 5: 808. 1905. 

Branchlets glabrous; leaves subcoriaceous, oblong, entire, slen- 
derly elevated-reticulate-veined on both sides, attenuate to the petiole, 
about 2 cm. long, scarcely half as wide; racemes lax, 10-15-flowered, 
to 6 cm. long; bracts 2-3 mm. long; flowers 10-11 mm. broad; anthers 
without connective; ovules 2; style obsolete. Neg. 14296. 

Cajamarca: Cutervo, Jelski 232, type. 

Berberis latifolia R. & P. Fl. Peruv. 3: 52. pi. 282. 1802; 450. 

Glabrous; branchlets virgate, elongate; leaves entire but undu- 
late, obovate, the larger, with the petiolar base, 8 cm. long, 3 cm. 
wide, lustrous above, papillose beneath, prominently reticulate- 
veined on both sides, chartaceous; spines trifid, 3-6 mm. long, shorter 
than the slender petioles; flowers 3-6-fasciculate; peduncles very 
much shorter than the leaves; style developed; fruit oval, 3-4-seeded, 
the seeds obovate. Schneider describes the leaves as mucronate, 
but those of the Madrid material are rounded at the apex and not 
apiculate. Neg. 29228. 

Huanuco: In cold groves, Pillao, Ruiz & Pavon. "Huajanpe," 
"palo amarillo." 

Berberis Lobbiana Schneid. Bot. Jahrb. 42: 83. 1908. B. con- 
ferta HBK. var. Lobbiana Schneid. Bull. Herb. Boiss. II. 5: 394. 1905. 

Similar to B. Jelskiana; flowers 3-4-fasciculate or distinctly 
racemose and the inflorescence then to 4 cm. long; pedicels and 
bracts apically glandular; leaves obovate-lanceolate, entire or with 
2-6 short teeth above the middle, to nearly 3 cm. long and 13 mm. 
wide, presumably not papillose beneath because compared by the 
author with B. virgata.Negs. 14299, 30132-3. 

Huanuco: Monzon, 3,500 meters, Weberbauer 3805. Bolivia. 
"Quisca." 

Berberis loxensis Benth. PI. Hartw. 125. 1843; 819. 

Similar in general to B. paniculata; leaves narrowly obovate, 
rounded at the minutely apiculate apex, narrowed below to the 
distinct petiole, the larger about 4 cm. long, 15-17 mm. wide, 
typically entire, very finely reticulate- veined on both sides; peduncles 
leafless; bractlets puberulent; pedicels 2-4 mm. long; sepals and petals 
about 3 mm. long. Differs from B. multiflora in its smaller leaves 
and flowers, the latter in narrower panicles; also, according to 
Schneider, the leaves have an obvious hypoderm (a specific char- 
acter?). He referred Weberbauer 1+199 to B. Beauvardiana with the 



676 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

remark "leaves entire, pedicels longer." Some leaves show 1-3 short, 
appressed teeth and mostly they measure 3 cm. long, and 12 mm. 
wide, otherwise the specimen, in aspect at least, matches B. loxensis. 
Closely allied forms include: B. Warscewiczii (Klotszch) Hieron., 
819, leaves typically dentate but an entire-leaved form is doubtfully 
distinct from the present plant; B. Schwerini Schneid., 818, found 
near Peru, with larger flowers, is well marked, the leaves spinulose- 
dentate, drying brown beneath, obscurely reticulate-veined above; 
B. Hallii Hieron., 813, has shorter leaves and suggests B. flexuosa, 
the panicles being simple except near the base. Neg. 14301. 

Cajamarca: Huambos, 2,400 meters, Weberbauer 4199.- Huanuco: 
Yanano, 1,800 meters, along trail, 3728. Yata, 2,100 meters, 2289. 
Ecuador. 

Berberis lutea R. & P. Fl. Peruv. 3: 51. pi. 280. 1802; 392. 

Unarmed, the branchlets more or less densely puberulent; leaves 
oblanceolate, 1-2 cm. long, 5-9 mm. wide, nearly always entire, or 
2-4-spinose-dentate, sessile or subsessile, lustrous, perhaps slightly 
glaucescent beneath but drying concolorous; flowers mostly 3-6- 
fasciculate; pedicels finally 6-10 mm. long, at least minutely puberu- 
lent, mostly about half as long as the leaves; style distinct; fruit 
ovate-elliptic; seeds usually 2, rarely 3-4, oblong-obovate. The 
original description reads "shrub glabrous," but the Madrid material 
is all definitely and persistently puberulent. The yellow petals are 
2-3 times longer than the reddish yellow calyx. The longer branches 
are unbranched for several decimeters (Ruiz and Pavon). With its 
wood the Indians dyed "bayatas" and cottons a yellow color, 
beautiful and permanent, and because of its durability made tool 
handles from it. Their name means "yellow wood with spiny 
leaves." (Ruiz & Pavon.) Negs. 8605, 27416. 

Huanuco: On wooded rocks, Panao, Pillao, Chaclla, Mufia, 
Ruiz & Pavdn. Chaclla, 3,200 meters, Weberbauer 6689. Amazonas: 
Chachapoyas (Mathews). "Espino amarillo," "ccarhuascassa." 

Berberis lutea R. & P. var. conferta (HBK.) DC. Syst. 2: 14. 
1821; 393. B. conferta HBK. Nov. Gen. & Sp. 5: 69. pi J^O. 1821. 
B. phyllacantha Rusby, Mem. Torrey Club 6: 4. 1896. 

A shrub 1 meter high; young branchlets terete, yellowish, mi- 
nutely puberulent, the older ones short, stout, glabrous; spines 
shortly trifid, deciduous; leaves rarely entire, usually 3-5-spinulose- 
dentate near the apex, oblanceolate or obovate, short-petiolate, 
mostly 10 mm. long, 6-8 mm. wide, or 15-18 mm. long, very smooth 



FLORA OF PERU 677 

above, lustrous and faintly reticulate-veined on both sides, not at 
all paler beneath, waxy-celled but epapillose; flowers 1-4-fasciculate, 
5-6 mm. wide; pedicels puberulent-pilose, 3-8 mm. long. This is 
the typical form of the variety. Citerne, 116, found the hypoderm 
cellular, but that of B. lutea fibrous. As Baehni has suggested to me, 
this apparent difference may well be the result of environment. 

Among several described variations of B. confer ta the following 
have been found in Peru, but since they have not been studied by me 
in relationship to B. lutea, they are not transferred; their existence 
suggests that the last five species in the key may not all be distinct 
from them or from each other: var. Spruceana Schneid., 393, spines 
none or simple, leaves to 20 mm. long, 7 mm. wide, entire, reticulate- 
veined on both sides, pedicels to 15 mm. long; var. hypopyrrantha 
Schneid. Bot. Jahrb. 42: 82. 1908, leaves 12-30 mm. long, 6-20 mm. 
wide, narrowed at the base to a petiole 5 mm. long, pedicels hirsu- 
tulous, 10 mm. long, flowers orange, to 1 cm. wide, branchlets espi- 
nose; var. psiloclada Schneid. loc. cit., similar to var. hypopyrrantha, 
but short-spinescent, leaves somewhat smaller, flowers 1-3, yellow, 
reddish outside, pedicels 2-4 mm. long, glabrous. Neg. 14290 (var. 
hypopyrrantha). 

Lima: Rio Blanco, 4,500 meters, 3013. Cajamarca: Humboldt, 
type. Huambos, 3,000 meters, Weberbauer 4157; 260 (var. Spru- 
ceana). Ancash: Above Huaraz, 3,200 meters, Weberbauer 8225 
(type of var. psiloclada). Yungai, 3,700 meters, Weberbauer 3278. 
Puno: Cuyocuyo, 3,100 meters, Weberbauer 853; 183 (var. hypo- 
pyrrantha). On Bolivian boundary (Pentland; var. boliviano). 
Huanuco: Grassy hillside, 3,000 meters, 1846. Northeast of Hua- 
nuco, 3,660 meters, 2173. Hualgayoc, Martinet 1071. Ayacucho: 
Huanta, 3,400 meters, Weberbauer 5560. Cuzco: Paucartambo to 
Tres Cruces, 3,400 meters, Pennell 14162. Urubamba, Soukup 60. 
Ollantaitambo, 2,800 meters, Herrera 3449. 

Berberis monosperma R. & P. Fl. Peruv. 3: 52. 1802; 804. 

Branchlets pale, smooth, flexuous, glabrous; spines trifid, to 2.5 
cm. long; leaves fasciculate, lance-obovate to subelliptic, obtuse, 
mucronate, cuneate to the sessile or subsessile base, entire, narrowly 
callous-margined, dull, finely reticulate-veined, clearly papillose, 
especially above, 3-4 cm. long, 8-10 mm. wide; flowers racemose or 
corymbose, the nodding, puberulent pedicels about 10 mm. long; 
flowers about 10, and 4 mm. wide, apparently granular-puberulent, 
like the sessile bractlets. The Madrid material is somewhat dam- 
aged, so the character of the pubescence is not entirely clear; plant 



678 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

"glabrous" according to Ruiz and Pa von. The specimen referred 
here by Schneider may not accord with this description. Neg. 29229. 
Lima(?): Huamantanga, Nee. Junin: Tarma, Ruiz & Pavon. 
Ancash: Tallenga, 3,300 meters, Weberbauer 2851 (det. Schneider). 
Puno: Agapata, 3,000 meters (Lechler 2642). 

Berberis multiflora Benth. PI. Hartw. 124. 1843; 816. 

Very similar to B. Barbeyana, the leaves also without obvious 
hypoderm but not at all glaucescent beneath, to 3 cm. wide and 6 
cm. long, or larger; panicle branches typically puberulent but gla- 
brous in var. calvescens Schneid., 816. This is perhaps the earliest 
name for a group of too closely distinguished forms, mostly Ecua- 
dorean but to be expected in adjacent Peru. See note under B. 
loxensis. Neg. 14313. 

Peru: Probably. Ecuador. 

Berberis paniculata Juss. ex DC. Syst. 2: 12. 1821; 819. 

Spines (on flowering branch) trifid, straight, 5 mm. long, decidu- 
ous; branchlets glabrous, smooth, striate at first, angled; leaves 
fascicled, oblong-elliptic, short-acute, mucronate, attenuate to the 
base, 5 cm. long, 1.5-2 cm. wide (perhaps larger on the branches), 
lustrous and finely but manifestly reticulate-veined on both sides, 
especially beneath, epapillose, concolorous; petioles to 5 mm. long; 
peduncles bearing 1-3 leaves similar to those of the subtending 
fascicle but smaller; panicles strict, the flowering portion 5-8 cm. 
long, 2 cm. broad, distinctly granular-glandular or puberulent; pedi- 
cels 4-5 mm. long; sepals and petals subequal, about 4 mm. long. 
The type may be from Ecuador, the original label (Paris) reading 
simply "Peru." Neg. 34530. 

Peru: Without locality, Jussieu. 

Berberis peruviana Schellenb. Bot. Jahrb. 50: Beibl. Ill: 6. 
1913. 

Glabrous except the puberulent branchlet tips; spines trifid; 
petioles 7 mm. long; blades obovate, cuneate at the base, coriaceous, 
lustrous above, papillose, dull and densely reticulate- veined beneath, 
to 4 cm. long, 2.5 cm. wide, mostly smaller, all sparsely spinulose- 
dentate; racemes about twice as long as the leaves; pedicels 9 mm. 
long; sepals 5 mm. long; style fungiform, persistent on the blue-black 
berry. 

Cuzco: Prov. Convention, near mouth of Rio Pampas, 2,800 
meters, Weberbauer 5897, type. 



FLORA OF PERU 679 

Berberis podophylla Schneid. Bot. Jahrb. 42: 84. 1908. 

A glabrous, armed shrub, the large, trifid spines nearly 2 cm. 
long; leaves rotund or 4-angled, usually truncate at the base, with 
8-14 spreading teeth, epapillose but distinctly elevated-reticulate 
on both sides; petioles (5-) 8-14 mm. long; panicles many-flowered, 
to 5 cm. long; flowers yellow, 4-5 mm. broad; pedicels 4-5 mm. long, 
the very acute bracts 1-1.5 mm. long; style none. Neg. 14306. 

Ancash: Between Samanco and Huaraz, 3,000 meters, Weber- 
bauer 3120, 3137; 170. 

Berberis saxicola Lechl. Berb. Amer. Austr. 42. 1857; 392. 

A low, unarmed, glabrous shrub about 30 cm. high, with arcuate, 
terete, robust branchlets; leaves in fascicles of 5-12, obovate, attenu- 
ate to the short petiole, coarsely spinose-dentate, coriaceous, laxly 
reticulate-veined, lustrous above, pale waxy-papillose beneath, to 
5.5 cm. long and 2.5 cm. wide; pedicels 1-several, 2-5 mm. (10 in 
fruit) long; sepals 5-6, oblong-obovate, entire like the larger, obovate 
petals; fruit ovate, the style obvious, the stigma orbicular. Flowers 
orange, "larger than those of the common barberry"; apparently 
6-7 mm. wide. Suggested by Schneider to be a synonym of B. lutea 
but apparently the same as B. conferta HBK. var. hypopyrrantha 
Schneid. Negs. 34536, 34537. 

Puno: In rocks, Sachapata and Tabina, 4,700 meters, Lechler 2096, 
type. Cuzco: Paso de Tres Cruces, 3,700 meters, Pennell 13911; 
petals scarlet. Paucartambo, Soukup 389. Cuzco, 3,600 meters, 
Vargas 530. 

Berberis undulata Lindl. Journ. Hort. Soc. 5: 7. 1850; 809. 

Spines 3-5-parted; leaves fasciculate, oblong-lanceolate, coria- 
ceous, dull, scarcely reticulate- veined, undulate or sometimes spinose- 
dentate; racemes erect, subsessile, shorter than the leaves. Schneider 
suggests that this poorly described plant, apparently cultivated from 
Lobb material (from Peru?) may belong to his group Truxillenses, 
represented in Peru by B. Jelskiana. Unless the name can be placed 
more definitely, it is best to ignore it. 

Berberis virgata R. & P. Fl. Peruv. 3: 51. pi. 281. 1802; 391. 

Similar to B. lutea but glabrous and with many smaller leaves; 
spines none or simple, stipular, 4-7 mm. long; branches slender, 
whiplike, often several decimeters long without a branchlet; leaves 
typically entire, or rarely with 3-4 spinose teeth near the apex, 
narrowly oblanceolate, 7-13 mm. long, 3-5 mm. wide, apiculate, 



680 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

apparently concolorous, very lustrous above, slightly so beneath, 
finely reticulate- veined on both sides; pedicels mostly 3 and 5-8 mm. 
long; flowers small, the petals 3-4 mm. long, conspicuously exceeding 
the calyx; style sessile, mushroom-like; berry 2-seeded, the seeds 
obovate (Ruiz and Pavon). B. rariflora Lechl., of Bolivia, more 
stockily branched, has short, trifid spines, but is otherwise similar. 
Negs. 27422, 14314. 

Huanuco: Woods, Huasa-huasi, Pillao, Huarica, Ruiz & Pavon, 
type. Cuzco: Above Yanamanche, 4,000 meters, Weberbauer 4946; 
243, 244. 

Berberis Weberbaueri Schneid. Bot. Jahrb. 42: 83. 1908. 

A shrub 2 meters high, the young branchlets minutely puberulent, 
the older ones armed with trifid, rather stout spines to 1.5 cm. 
long; leaves obovate-elliptic to subrotund, truncate or narrowed at 
the base to the (4-8 mm. long) petiole, entire, 1.5-3(-4) cm. long, 
1.2-3 cm. wide, papillose and finely reticulate- veined above; racemes 
to 3.5 cm. long, minutely puberulent; flowers 4-10, yellow, 10-12 
mm. broad; pedicels 4-8 mm. long; style distinctly developed. In 
the obvious style, allied to B. monosperma, but in other respects to 
B. Jelskiana. Neg. 14318. 

Ancash: Ocros, 3,400 meters, Weberbauer 2680; 170. 

71. MENISPERMACEAE. Moonseed Family 
Reference: Diels, Pflanzenreich IV. 94. 1910. 

A family comprising low-climbing, herbaceous vines and more or 
less scandent shrubs whose often small pistillate and staminate 
flowers suggest the Euphorbiaceae. The leaves of some species 
are entire and variously lobed, most often palmately, on the same 
plant, the fruits being grape-like in appearance. Economic interest 
of the family lies in a species of Chondodendron (cf. polyanthum) 
and Anomospermum reticulatum, known as "pani," which, as shown 
recently by Smith and Krukoff, Bull. Torrey Club 64: 401-409. 
1937, are ingredients of the arrow poison "curare." The first named 
plant is known by the Indians as "atinupa," the latter as "iku." 
In spite of monographic treatments of the family, the genera remain 
identifiable only on technical characters, largely those of the fruit. 
A supplementary, artificial key has therefore been devised for the 
Peruvian plants, but with either key the genera are difficult to dis- 
tinguish. Most of the material cited here has been determined by 
Diels. 



FLORA OF PERU 681 

Artificial key (often applicable only to Peruvian species) 
Inflorescence spicate or racemose, or at least apparently so. 

Leaf nervation somewhat melastomaceous 3. Sciadotenia. 

Leaf nervation not at all melastomaceous (unless in D. remota with 

broadly ovate, truncate-based leaves). 

Stamens free, usually 3; petals basally thickened . 5. Disciphania. 
Stamens more or less united ; petals evenly fleshy. 

4. Odontocarya. 

Inflorescence cymose or compound. 

Petioles more or less contorted or geniculate at the base, or tumid 
below the blade, but at least not noticeably enlarged at the 
base (cf. Anomospermum) . 

Petioles tumid or triangularly broadened below the blade (little 
so in Somphoxylon Klugii with oblong and pinnate-nerved 
leaves) . 
Inflorescences 20 cm. long or shorter, or the leaves softly 

white-tomentose beneath 2. Chondodendron. 

Inflorescences mostly several decimeters long; leaves glabrate 

or puberulent beneath 6. Somphoxylon. 

Petioles not tumid below the blade, unless in Sciadotenia 

brachypoda. 
Leaves broadly ovate. 
Petals and sepals 6; leaves, if pubescent, deeply cordate at 

the base 4. Odontocarya. 

Petals 1 or 2 or none; leaves pubescent, or not deeply cordate 

at the base 1. Cissampelos. 

Leaves suboblong. 

Leaves entire 3. Sciadotenia. 

Leaves serrate 7. Synandropus. 

Petioles definitely enlarged at the base and straight (sometimes 
also tumid below the blade, or in Anomospermum to the base 
but straight). 
Leaves melastome-like; petals minute or none. 

Leaves glabrous; sepals 6, the outer 3 bracteolate. . 11. Abuta. 
Leaves white-pubescent beneath; sepals 6-18, imbricate. 

2. Chondodendron. 
Leaves not melastome-like; petals obvious. 

Inflorescence (staminate) crowded, much shorter than the 
petioles 10. Elissarrhena. 



682 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Inflorescence (at least the staminate) an elongate (5-15 cm.) 
panicle. 

Inflorescences 3 or 4 together or congested; petals not very 
thick 8. Hyperbaena. 

Inflorescences 1 or 2 together, open; petals very fleshy. 

9. Anomospermum. 

Technical key (based on Diels, op. cit.) 

Sepals of pistillate flowers 1 or 2, of the staminate 4; flowers minute, 
the staminate many; cotyledons not foliaceous; endosperm 
scarcely ruminate 1. Cissampelos. 

Sepals 6-36 in both kinds of flowers. 
Sepals valvate or spirally imbricate; endosperm none. 

Flowers pedicellate, paniculate 2. Chondodendron. 

Flowers in small cymes 3. Sciadotenia. 

Sepals more or less imbricate. 

Outer sepals often distinctly smaller but not minute; cotyledons 
foliaceous, except in Hyperbaena. 

Leaves membranous or subcoriaceous, often lobed; endosperm 
present. 

Inflorescence narrowly paniculate, pseudoracemose, or 
spicate. 

Stamens or staminodia usually 6, more or less connate. 

4. Odontocarya. 
Stamens 3-6, free; pistillate flowers without staminodia. 

5. Disciphania. 
Inflorescence amply paniculate. 

Leaves entire 6. Somphoxylon. 

Leaves serrate 7. Synandropus. 

Leaves coriaceous, entire; endosperm none. . .8. Hyperbaena. 
Outer sepals minute, bractlike; cotyledons not foliaceous. 
Petals conspicuous. 

Petals fleshy, strongly thickened; stems not fistulous. 

9. Anomospermum. 

Petals fleshy, but little thickened; stems fistulous. 

10. Elissarrhena. 

Petals minute or none. . .11. Abuta. 



FLORA OF PERU 683 

1. CISSAMPELOS L. 

Usually herbaceous vines with often peltate leaves and many- 
flowered, paniculate-corymbose, axillary inflorescences, or these 
borne on axillary branchlets. Staminate flowers with 4 sepals, these 
usually pilose dorsally, often spreading. Petals connate or very 
rarely free. Stamens connate. Pistillate flowers with 1 obovate 
sepal and usually only 1 shorter petal. Carpel 1, villous. Drupes 
fleshy. 

Leaves mostly and distinctly peltate; staminate inflorescences 

elongate, open. 

Leaves often more or less pilose, the hairs long; inflorescence 
pubescence loose, spreading; bracts all foliaceous. 

C. Iropaeolifolia. 

Leaves glabrate, the sparse pubescence minute; inflorescence 
cinereous-strigose-hispidulous; bracts all reduced. 

C. grandifolia. 

Leaves not peltate except sometimes in C. Pareira with the staminate 
inflorescence normally corymbose in the leaf axils. 

Leaves glabrous beneath or, if pubescent, not so densely that the 
leaf surface is hidden, if at all cordate, very shallowly and 
openly so; bracts all reduced. 
Leaf base, even if slightly cordate, tapering into the petiole; 

pistillate inflorescence simple C. andromorpha. 

Leaf base somewhat cordate, at the junction with the petiole 
rounded or truncate; pistillate inflorescence paniculate. 

C. fasciculata. 

Leaves densely pubescent beneath, the leaf surface often con- 
cealed ; staminate bracts reduced C. Pareira. 

Cissampelos andromorpha DC. Syst. Veg. 1: 539. 1818; 303. 

Petioles and blades sparsely pilose, the blades ovate-cordate, 
sub truncate or emarginate at the base, usually obtuse or retuse at the 
apex, mucronulate, 5-7 cm. long, 6-8 cm. wide, with 5-7 palmate 
nerves, these conspicuous beneath; staminate panicles often fascicu- 
late, cymosely branched, 5-25 cm. long; bracts obsolete; pistillate 
inflorescence simple; staminate sepals glabrous or sparsely pilose 
outside. Flowers yellow (Williams). 

Loreto: Florida, Rio Putumayo, 200 meters, King 2029. Mishu- 
yacu, in clearing, 100 meters, Klug 1008. Balsapuerto, in clearing, 
King 2893. Iquitos, Williams 8098; in clearing, Klug 967, 1008. 



684 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Caballo-cocha, Williams 2085. Pebas, Isern 2263. Rio Acre: Ule 
9384 (form with acuminate leaves). Widely distributed in South 
America. 

Cissampelos fasciculata Benth. Lond. Journ. Bot. 2: 361. 1843; 
304. 

Closely related to C. andromorpha, but the pistillate inflorescence 
paniculate, the staminate sepals usually densely pubescent outside, 
and the bracts mostly well developed.- Flowers brownish yellow 
(Klug). Juice from the leaves is employed as a remedy for diseases of 
the eye (King). Neg. 27515. 

Loreto: Florida, Rio Putumayo, 180 meters, King 2322. Lower 
Rio Nanay, Williams 405. Mishuyacu, Klug 1498. Central and 
South America. ' ' Aipoyo . ' ' 

Cissampelos grandifolia Triana & Planch. Ann. Sci. Nat. IV. 
17: 44. 1862; 305. 

Branches sulcate; petioles pilose, attached to the blade as much 
as 4 cm. from the margin; blades finally glabrous above, pubescent 
beneath, especially on the 5 prominent nerves, very broadly ovate or 
suborbicular, sometimes shallowly crenate, 7-15 cm. long and broad; 
staminate panicle ample, to 20 cm. long, the primary branchlets soli- 
tary or few, densely hispidulous-tomentulose; sepals sericeous- 
pilose; drupes tuberculate, pilose, 5-6 mm. long. 

Loreto: Leticia, Williams 3049, 3051. Pumayacu, between 
Balsapuerto and Moyobamba, 600-1,200 meters, in forest, Klug 
3230. Mishuyacu, 100 meters, in clearing, Klug 773. Rio Napo, 
near Mazan, 110 meters, overflowed bank, Mexia 6 463 a. Central 
and South America. 

Cissampelos Pareira L. Sp. PI. 1031. 1753; 286. 

More or less pubescent, the staminate inflorescence subcorym- 
bose, in the axils of normal leaves; the only Peruvian species with 
this character. Many forms have been described as varieties, the 
Peruvian plants being mostly var. Haenkeana Presl (ex Diels, 
Pflanzenreich IV. 94: 292. 1910; C. Haenkeana Presl, Rel. Haenk. 2: 
80. 1835; type collected in Peru by Pavon; C. hirsutissima Presl, loc. 
cit.; type from Peru, Poeppig 1293; another synonym is C. cor data 
Ruiz, mss.), differing from the type form in having only the upper 
leaves peltate, often larger and heavier. Illustrated, Pflanzenreich, 
loc. cit. 

San Martin: San Roque, 1,400 meters, in forest, Williams 7256, 
7375. Tarapoto, 750 meters, Williams 5564; Spruce 4409; Ule 523. 






FLOI& OF PERU 685 

Huanuco: Posuso, Cochero, Chacahuasi, Ruiz & Pavon. Pam- 
payacu, Kanehira 83; at 1,050 meters, climbing over shrubs, 5022. 
Yanano, 1,800 meters, 3758. Cochero, Poeppig 1293. Between 
Chaclla and Mufia, 2,100 meters, Weberbauer 6710, 6702Cuzco: 
Quebrada Versalles, Diehl 2448. Without locality, Gay 1031, 1019. 
Junin: Tarma, Weberbauer 6523, 1993; 247. Tropics of both hemi- 
spheres. "Vaca-nahui-huasca" (Williams). 

Cissampelos tropaeolifolia DC. Syst. Veg. 1: 532. 1818; 299. 

Petioles 6-8 cm. long, more or less hispid; blades peltate, ovate 
or subtriangular-orbicular, truncate or rotund at the base, long- 
mucronate, 4-7 cm. long, 3.5-7 cm. wide; staminate inflorescence 
with slender peduncles and branchlets; carpels glabrous. 

Junin: La Merced, 700 meters, Killip & Smith 23443. Chan- 
chamayo, Isern 2416. Chacahuasi, Ruiz & Pavon. San Martin: 
San Roque, 1,400 meters, Williams 7456. Huanuco: Posuso, 4594. 
Cochero, Dombey, type. Pampayacu, 5059. Loreto: Moyobamba, 
Weberbauer 4517, 4521. El Recreo, Williams 3936. Rio Itaya, 110 
meters, Killip & Smith 29384; Williams 3400, 3374. San Antonio, 
Rio Itaya, 110 meters, in forest, Killip & Smith 29497. Rio Nanay, 
Williams 518. Caballo-cocha, Williams 2405. Balsapuerto, Klug 
2911. Sandia: Chunchosmayo, Weberbauer 1188. Rio Acre: Ule 
9385. Extending to Brazil and Central America. 

2. CHONDODENDRON R. & P. 

Climbing shrubs with tuberculate bark and entire leaves. Pani- 
cles axillary, solitary or fasciculate from older branches. Staminate 
flowers with 6-18 sepals, the outer minute, pubescent within, the 
inner glabrate but ciliate, the tips finally reflexed. Petals 6, gla- 
brous. Stamens 6 and free or 3 and connate, the anthers longitu- 
dinally dehiscent. Pistillate flowers similar, with minute or no 
staminodia. Carpels 6; style obsolete, the stigma linguiform. 
Drupes stipitate, spreading; endosperm none. The name was spelled 
Chondrodendron by Miers. 

Besides the following there is a sterile specimen apparently 
referable to the genus from Pebas, Loreto, collected by Castelnau, 
who noted it as used by the Oregones, less frequently by the Yaguas, 
as an arrow and fish poison under the name "pani." This was named 
Cocculus toxicoferus Wedd. in Castelnau, Exped. Ame"r. Sud 5: 22. 
1851. It may be described as follows: Scandent shrub with thin, 
smooth, gray bark, sparsely brownish-rugulose; branches terete, 
striate, glabrous; leaves (young) palmate-ovate, acutish at the sub- 



686 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

peltate base, abruptly and narrowly acuminate, 3-5-nerved; marginal 
nerves scarcely half the leaf length, the 3-5 secondary veins anasto- 
mosing toward the tip of the leaf arcuately with the primaries and 
the midrib, glabrous throughout, the upper surface lustrous, pale 
green, the lower glaucous; petiole greatly elongate, subequaling the 
leaf blade; flowers unknown (but said to flower in September, fruit 
in December). 

Sepals 6, all tomentulose outside. 

Outer sepals less than 1 mm. long C. candicans. 

Outer sepals more than 1 mm. long. 

Principal basal leaf nerves 3; petals pilose C. polyanthum. 

Principal basal leaf nerves 5; petals glabrous C. iquitanum. 

Sepals more numerous, the inner ones glabrous C. tomentosum. 

Chondodendron candicans (Rich.) Sandw. Kew Bull. 342. 
1930. Abuta candicans Rich, ex DC. Syst. Veg. 1: 543. 1818. Sciado- 
tenia candicans Diels, Pflanzenreich IV. 94: 87. 1910. 

Leaves rather chartaceous, glabrous and lustrous above, cinereous- 
crustaceous-tomentulose beneath, narrowly ovate, gradually acute or 
acuminate, about 20 cm. long and half as wide or, according to 
Sandwith, 6-17 cm. long, 3-10 cm. wide, 3 of the 5 basal nerves 
stout; staminate inflorescence paniculate, cinereous-tomentulose, to 
15 cm. long, 4-10 cm. wide at the base; pedicels 2-10 mm. long; 
outer sepals 0.8 mm. long, 0.6 mm. wide, the inner to 1.5 mm. long, 
12 mm. wide; petals 6, glabrous, to 0.6 mm. long; fruiting inflores- 
cence 2-5 cm. long, the densely pilose pedicels spreading, 8-12 mm. 
long; drupes often 3, densely brownish-tomentose, the free stipes 
3-5 mm. long. Description of flowers from Sandwith, of fruit from 
Diels, based on the Huber material. 

Rio Acre: Huber 4286. Brazil; Guianas. 

Chondodendron iquitanum Diels, Notizbl. Bot. Gart. Berlin 
9: 997. 1926. 

Adult branches glabrous; petioles minutely pubescent, 8-18 cm. 
long; blades coriaceous, glabrous above, pale and minutely tomentose 
beneath, broadly ovate or ovate-elliptic, acuminate, 18-26 cm. long, 
12-17 cm. wide, the 5 primary as well as the secondary nerves promi- 
nent beneath; staminate inflorescence 8-10 cm. long, sericeous- 
tomentose, the pedicels 2.5-3 mm. long; outer sepals 1.3 mm. long, 
the inner 2.3 mm. long; petals obovate, 1.2 mm. long, equaled by the 



FLORA OF PERU 687 

glabrous stamens. Allied to C. polyanthum, but with shorter, 
stouter pedicels and 5-7 basal leaf nerves. Neg. 27514. 

Loreto: Mouth of Rio Santiago, in upland forest, Tessmann 4196. 

Chondodendron polyanthum Diels, Pflanzenreich IV. 94: 78. 
1910. Hyperbaena polyantha Diels, Verh. Bot. Ver. Brandenb. 50: 
73. 1908. 

Similar to C. tomentosum, but much less pubescent, the leaves 
glabrous above, and with 2 basal nerves; inner sepals 2-2.5 mm. 
long; petals slightly pilose, to 2 mm. long. Type from Brazil, 
with leaves 8-15 cm. long, the pedicels slender, 3-6 mm. long. Neg. 
4985. 

Loreto: San Antonio, Rio Itaya, Kittip & Smith 29337. Balsa- 
puerto, Killip & Smith 28665. Fortaleza, near Yurimaguas, Klug 
2782. Pebas, Castelnau. Florida, Rio Putumayo, 200 meters, in 
forest, Klug 2042. Brazil. "Atinupa." 

Chondodendron tomentosum R. & P. Syst. 261. 1798; 79. 
Epibaterium tomentosum Pers. Syn. PI. 2: 561. 1807. Cocculus 
Chondodendron DC. Syst. 1: 522. 1818. Botryopsis Spruceana Eichl. 
Flora 47: 394. 1864. Chondodendron cretosum Miers, Contr. Bot. 
3: 312. 1871. 

Branchlets, petioles, and especially the blades beneath velutinous- 
pubescent; petioles 8-12 cm. long, apically enlarged; blades sparsely 
pubescent or glabrous above, subcordate-ovate or suborbicular, obtuse 
or somewhat emarginate, obscurely crenate, sometimes 10-15 cm. 
long and about as broad, but often smaller; the basal nerves 5, the 
lateral 3-4; panicles 10-25 cm. long; 9 outer sepals about 1 mm. long, 
the 6 inner ones 3.5 mm. long; fruiting pedicels enlarged; drupes 
ellipsoid, 12 mm. long, the stipes 3-4 mm. long. Illustrated, Mart. 
FI. Bras. 13, pt. 1: pi 48 (fls.). Neg. 34500. 

Junin: Chacahuasi, Ruiz & Pavon, type. Rio Perene*, 600 
meters, Kittip & Smith 25175. San Martin: Rio Mayo, Williams 
6276. Tarapoto, Spruce 4474; Ule. Chazuta, 260 meters, in forest, 
Klug 4044- Loreto: Rio Ucayali, Tessmann 3342, 3157. Mouth 
of Rio Santiago, Tessmann 4333. Colombia and Panama. 

3. SCIADOTENIA Miers 

Similar to Chondodendron, but the staminate sepals 18-36. 
Petals pubescent without. Anthers obliquely dehiscent. Pistillate 
flowers without petals. Carpels 6-16, connate at the base. 



688 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves cordate or truncate at the base S. ramiflora. 

Leaves narrowed to the base. 

Leaves 3-nerved S. brachypoda. 

Leaves 5-nerved S. similis. 

Sciadotenia brachypoda Diels, Pflanzenreich IV. 94: 84. 1910. 

Similar to S. ramiflora but less pubescent, the petioles 4-6 cm. 
long, the blades acute at the base, 10-20 cm. long, half as wide, long- 
acuminate, 3-nerved at the base; fruiting peduncles 1.5-2.5 cm. 
long, densely hispidulous-pilose. Probably in Amazonian Peru, since 
it has been found on the Rio Futumayo. S. amazonica Eichl. and 

5. Sprucei Diels have axillary pistillate peduncles; leaves in the former 
only 3.5 cm. wide, in the latter 5-7 cm. Neg. 4986. 

Peru: Probably. Amazonian Brazil. 

Sciadotenia ramiflora Eichl. in Mart. Fl. Bras. 13, pt. 1: 202. 
pi. 47. 1864; 84. 

Young branches shortly rusty-pilose; petioles 2.5 to nearly 4 cm. 
long; blades thin, glabrous and shining above, sparsely pilose beneath, 
cordate-ovate or lance-cordate, rarely truncate at the base, acuminate 
at the apex, 7.5-13 cm. long, 2.5-6.5 cm. wide, with 5 basal nerves; 
pistillate inflorescences supra-axillary, mostly solitary; drupes 6, 
obovoid, pilose, on short-connate stipes, 8-12 mm. long. Negs. 
30149, 4986. 

Loreto: Yurimaguas, Poeppig 2271, type. Mishuyacu, 100 
meters, Klug 1299. 

Sciadotenia similis Moldenke, ined. 

Leaves slender-petiolate, glabrous, subcoriaceous, the blades 
broadly elliptic to elliptic-ovate, 10-15 cm. long, abruptly long- 
acuminate, rounded and abruptly contracted at the base, somewhat 
lustrous, very pale beneath; drupes narrowly obovoid, about 1.5 cm. 
long and 1 cm. wide, rounded at the apex. 

Loreto: La Victoria, on the Amazon, in forest, Williams 3075. 

4. ODONTOCARYA Miers 

Scandent shrubs with verrucose and fistulous branchlets and 
long-petioled, more or less cordate-ovate or elliptic, rather thin leaves 
with 5-7 basal nerves. Staminate inflorescence pseudoracemose, the 
flowers fasciculate; pistillate inflorescence racemose, the flowers 
solitary; sepals membranous, concave, 6, the outer 3 smaller. Petals 

6, the margins inflexed. Stamens usually 6, more or less connate, 
the anthers dehiscent by a vertical slit. Pistillate flowers with 6 



FLORA OF PERU 689 

filamentose staminodia. Carpels 3, glabrous, the sessile stigma 
radiately incised. Drupes fleshy, with a scabrous-tuberculate and 
coarsely pilose endocarp. Similar to Somphoxylon except for the 
inflorescence; see Diels, Notizbl. Bot. Gart. Berlin 13: 27. 1936. 

Leaves truncate or very shallowly cordate at the base, glabrous 
beneath or nearly so. 

Leaves 5-6 cm. wide and distinctly longer. 

Acumen of the leaf 5 mm. long; inner sepals about 1.5 mm. long. 

0. diplobotrya. 
Acumen 2-3 mm. long; inner sepals about 3 mm. long. .0. Ulei. 

Leaves 9-11 cm. wide and nearly as long 0. floribunda. 

Leaves usually cordate at the base, pubescent beneath . .0. paupera. 

Odontocarya diplobotrya Diels, Pflanzenreich IV. 94: 172. 
1910. 

Glabrous or nearly so; branchlets slender, sulcate-striate; petioles 
flexuous, 4-7 cm. long; blades thin, paler beneath, emarginate or 
truncate at the base, the conspicuous tip subobtuse, 8-9 cm. long, 
about 6 cm. wide, the 3 basal nerves prominent beneath; staminate 
inflorescence compound, 10 cm. long, the lower subracemose, spread- 
ing branchlets about 1 cm. long; flowers solitary or in small fascicles; 
outer sepals ovate, the inner obovate, 1.5 mm. long; petals 1.2 mm. 
long; stamens more or less connate. Neg. 4988. The Ayacucho 
specimen has slightly larger flowers (Smith). The related 0. Schim- 
pffi Diels, Notizbl. Bot. Gart. Berlin 13: 27. 1936, of Ecuador has 
only 3 stamens. 

Loreto: Yurimaguas, Poeppig, type; Williams 5241, 5126. La 
Victoria, Williams 2657, 2593. Rio Ucayali, Tessmann 3265, 3356. 
Junin: Chanchamayo, Raimondi; Martinet. Ayacucho: Rio Apuri- 
mac valley (Killip & Smith 22855; det. Smith). Brazil. 

Odontocarya floribunda Diels, Notizbl. Bot. Gart. Berlin 6: 
133. 1914. 

Scandent, with lenticellate, striate branches; petioles 8-11 cm. 
long; blades glabrous except for the minutely puberulent nerves 
beneath, cordate at the base, slightly decurrent on the petiole, sub- 
orbicular, acuminate, 9-11 cm. wide, nearly as long, the 3-5 primary 
nerves prominent beneath; pedicels slender; outer sepals broadly 
ovate, 1.3 mm. long, the narrower inner ones 3.5-4 mm. long; petals 
2-2.5 mm. long; stamens connate nearly to the middle, about 2 mm. 



690 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long. Similar in staminate inflorescence to 0. diplobotrya. Neg. 
27513. 

Rio Acre: Seringal San Francisco, Ule 9381, type. 

Odontocarya paupera (Griseb.) Diels, Pflanzenreich IV. 94: 
172. 1910. Cocculus pauper Griseb. Gott. Abh. 7: 162. 1857. 

Petioles slender, slightly pilose, 5-10 cm. long; blades thin, 
minutely scabrous above, usually somewhat pubescent, especially on 
the nerves beneath, more or less distinctly cordate or sometimes 
truncate and subhastately trilobed, obtuse or acute, to 15 cm. long 
and nearly as broad, often much smaller, with 3-7 basal nerves; 
staminate inflorescence slender, to 15 cm. long, the flowers in fascicles 
of 5-6, on pedicels 2.5-4 mm. long; outer sepals ovate-elliptic, the 
inner obovate-elliptic, 2.5-3 mm. long; petals 2-2.5 mm. long; 
filaments more or less connate; pistillate inflorescence with pedicels 
5-8 mm. long; carpels 3, with a 3-lobed stigma; drupes 1 cm. long. 
Illustrated, Mart. Fl. Bras. 13, pt. 1: pi. 36. Neg. 30151. 

Loreto: Yurimaguas, Williams 4235, 5168. Caballo-cocha, 
Williams 2330. San Martin: Zepelacio, 1,200 meters, in forest, 
Klug 3317. West Indies; Central and South America. 

Odontocarya Ulei Diels, Notizbl. Bot. Gart. Berlin 6: 133. 1914. 

Similar to 0. diplobotrya, but the ovate leaves subobtuse at the 
base and the outer sepals only 0.6-0.8 mm. long, the inner ones 3 mm. 
long, the petals 1.5 mm. long. Neg. 4989. 

Rio Acre: Seringal San Francisco, Ule 9380, type. 

5. DISCIPHANIA Eichl. 

Perennial vines from bulbous roots. Leaves usually cordate, 
sometimes 3-7-lobed, the palmate nerves 5-7. Flowers in simple, 
axillary spikes or pseudoracemes. Staminate sepals usually 6 and 
subequal, thin or fleshy. Petals much smaller, fleshy, more or less 
excavate at the base. Stamens 3 or rarely 6, free. Pistillate sepals 
and petals subequal. Staminodia none. Carpels 3, free, the stigma 
discoid, subsessile. Drupes often solitary, fleshy, the endocarp 
woody. 

Leaves, at least in part, distinctly cordate at the base. 
Plants not densely pilose. 

Leaf sinus closed D. clausa. 

Leaf sinus open. 

Leaves all cordate at the base, strigose D. convolvulacea. 



FLORA OF PERU 691 

Leaves all cordate at the base, glabrous D. Tessmannii. 

Leaves slightly cordate at the base, some of them truncate, 

glabrate D. Ernstii. 

Plants densely long-pilose D. lobata. 

Leaves truncate or rounded at the base. 

Leaves 5-nerved from the base D. remota. 

Leaves pinnate-nerved D. Killipii. 

Disciphania clausa Diels, Pflanzenreich IV. 94: 176. 1910. 

Glabrous (except sepals), the branchlets drying purple; petioles 
6-8 cm. long; blades thin, paler beneath, cordate-ovate or broadly 
elliptic, 8-11 cm. long, 6-8 cm. wide, 5-nerved; pistillate inflorescence 
7-8 cm. long; sepals free, papillose- velutinous within at the base, the 
subovate outer ones 7 mm. long, the inner narrower and shorter; 
petals narrowly obovate, 2.5-3 mm. long; drupes subglobose, 18 mm. 
long. Type from northern Amazonian Brazil. Neg. 4999. 

Rio Acre: Ule 9383 (det. Diels). Brazil. 

Disciphania convolvulacea (Poepp.) Diels, Pflanzenreich IV. 
94: 178. 1910. Chondodendron convolvulaceum Poepp. Nov. Gen. & 
Sp. 2: 65. pi. 190. 1838. Odontocarya convolvulacea Miers, Contr. 
Bot. 3: 62. 1871. 

Distinctly hispidulous-pilose on all parts; petioles 9-18 cm. long; 
blades cordate-acuminate, about 10 cm. long, 7-10 cm. wide, 5-7- 
nerved; pistillate spikes finally 15 cm. long, on peduncles 3-4 cm. 
long; bracts 3-4 mm. long; outer sepals elongate-ovate, much exceed- 
ing the inner ones; carpels 3; stigma sessile; sepals finally 7 mm. long, 
tardily deciduous; fruits berry-like. D. micrantha Diels, Amazonian, 
is glabrous, and aberrant in the genus by its 6 stamens and unequal 
sepals, the outer ones smaller. The Ecuadorean D. appendiculata 
Diels has petioles 4-5 cm. long and.appendaged petals. Neg. 5000. 

Huanuco: Cochero, Poeppig 1394- "Uva del monte." 

Disciphania Ernstii Eichl. Jahrb. Bot. Gart. Berlin 2: 329. 
pi 12. 1883; 175. 

Glabrous or nearly so; petioles 5-7 cm. long; blades thin, paler 
beneath, 7-15 cm. long and broad, cordate, entire or irregularly 3-7- 
angulate-lobed, 5-7-nerved; staminate spikes about 6 cm. long, the 
sepals 4 times as long as the fleshy petals; pistillate flowers similar, 
but the petals and sepals connate at the base, the sepals 4-5 mm. 
long, the petals 1.5 mm. long; carpels 3, the stigma subsessile; drupes 



692 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

mostly solitary, scarlet, the endocarp winged. Compare D. micran- 
tha, under D. convolvulacea. Neg. 5001. 

Junin: La Merced, 600 meters, cultivated, but said to be native, 
5466. Loreto: Huallaga, Yurimaguas, Williams 4654- Venezuela. 

Disciphania Killipii Diels, Notizbl. Bot. Gart. Berlin 11: 782. 
1933. 

Herbaceous; leaves ovate-elliptic, acuminate, rounded or sub- 
truncate at the base but not cordate nor at all peltate, glabrous, 
8-10 cm. long, 4.5-5.5 cm. wide, the petiole 5-6 cm. long, the primary 
nerves 3-5; staminate spikes solitary, axillary; sepals joined and cup- 
like at the base, then stellate-spreading, salmon pink, fleshy, 5-6.5 
mm. long, 3-3.5 mm. wide, the outer ones slightly longer and nar- 
rower; petals broadly obtriangular; stamens 3. Apparently the 
spikes may sometimes be paniculate-racemose, but perhaps, as the 
author suggests, an anomaly. The species is unique in its epeltate 
leaves. 

Loreto: Iquitos, 100 meters, in forest, Killip & Smith 27041, type. 

Disciphania lobata Eichl. in Mart. Fl. Bras. 13, pt. 1: 169. 
pi. 36. 1864; 178. 

Like D. convolvulacea, but more reddish-villous, the leaves some- 
times 20 cm. long and broad, the basal sinus very deep; bracts about 
2 mm. long; drupes angulate-oblong, hispid. Illustrated, Pflanzen- 
reich, loc. cit. /. 63. Flowers red-brown. 

Loreto: Mishuyacu, 100 meters, in forest, King 780, 423. Brazil. 

Disciphania remota Diels, Notizbl. Bot. Gart. Berlin 11: 782. 
1933. 

Herbaceous; longer petioles about 8 cm. long; blades glabrous, 
broadly or narrowly ovate, truncate or very slightly emarginate at 
the base, acuminate, 3-5-nerved, 12-15 cm. long, 6-10 cm. wide; 
staminate spikes slender, with the peduncle to 25 cm. long, the 
remote flowers glabrous; sepals elliptic-ovate, concave, incurved at 
the tip, 4.5 mm. long, 2.5 mm. wide; petals 6, slightly fleshy, 1.5 mm. 
broad, the back very shortly produced; stamens 0.8-1 mm. long, the 
connective dilated . Allied to D. Ernstii Eichl., but the leaves not 
lobed and the flowers remote. 

Loreto: Puerto Arturo, Yurimaguas, 135 meters, in forest, Killip 
& Smith 27892, type; Williams 4649. Yurimaguas, Williams 5353. 



FLORA OF PERU 693 

Disciphania Tessmannii Diels, Notizbl. Bot. Gart. Berlin 9: 
137. 1924. 

A glabrous, herbaceous vine with striate stems; petioles genicu- 
late, 6-8.5 cm. long; blades membranaceous, cordate-hastate-ovate, 
acutely acuminate, 6-7 cm. long, 4 cm. wide, palmately 5-nerved, 
the nerves scarcely prominent; staminate spikes pedunculate, the 
bracts obsolete(?); flowers sessile, yellowish green, the 6 sepals 
rotately spreading, the outer ones 5 mm. long, half as broad, the 
inner 5-6 mm. long, 3.5 mm. broad; petals 6, with a narrow, tongue- 
shaped appendage. D. appendiculata Diels, of Ecuador, has broader, 
7-nerved, pubescent leaves. Neg. 27512. 

Loreto : Yarina-cocha, middle Rio Ucayali, Tessmann 8385, type. 

6. SOMPHOXYLON Eichl. 

Vines with entire, pinnately nerved (except for the basal nerves) 
leaves. Staminate flowers with 6 sepals, the 3 larger, inner ones 
broadly elliptic and concave. Petals 6. Stamens 3, connate to the 
anthers; anthers vertically dehiscent. Panicles very large, with 
widely spreading branches, the flowers subspicate, in glomerules of 
2-5. Illustrated, Pflanzenreich IV. 94:/. 62. 

Petioles rarely more than 1.5 cm. long S. Klugii. 

Petioles 3-8 cm. long. 

Petals 6; stamens 3 S. Wullschlaegelii. 

Petals 3; stamens reduced to 1 S. deminutum. 

Somphoxylon deminutum Diels, Notizbl. Bot. Gart. Berlin 9: 
997. 1926. 

Similar to S. Wullschlaegelii, but remarkable in the reduction of 
the petals to 3 and the stamens to 1; leaves broadly ovate, excisely 
cordate at the base, about 15 cm. long, 10-11 cm. wide, the primary 
lateral nerves about 5; staminate panicle glabrous, abundantly 
floriferous; outer sepals 0.5 mm. long, the inner 1.5 mm. long, obo- 
vate; petals with inflexed margins, 1 mm. long. 

Loreto: San Isidro, mouth of Rio Pastaza, in low, flooded woods, 
Tessmann 4976, type. 

Somphoxylon Klugii A. C. Smith, Bull. Torrey Club 58: 92. 
1931. 

Branchlets striate; petioles slightly swollen at the base, 12-16 mm. 
long; blades oblong or ovate-oblong, rounded at the base, acuminate 
or caudate, 13-16 cm. long, 5-6 cm. wide, the lateral nerves 7-8; 



694 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

staminate inflorescence to 30 cm. long and broad, the primary 
branches 12-15 cm. long, densely but minutely puberulent; outer 
sepals ovate-deltoid, the ovate inner ones 1.8 mm. long; petals 1.4 
mm. long; anther column 1.5 mm. long. 

Loreto: Mishuyacu, near Iquitos, Klug 579, type; 276. La 
Victoria, Williams 2663. 

Somphoxylon Wullschlaegelii Eichl. in Mart. Fl. Bras. 13, 
pt. 1: 206. pi. 37. 1864. S. Ulei Diels, Verh. Bot. Ver. Brandenb. 50: 
74. 1908. 

Branches terete, not at all striate; petioles 3-8 cm. long, some- 
what enlarged below the subcoriaceous blades, these lustrous above, 
dull and glabrous or papillose-puberulent beneath, ovate-elliptic, 
the base rounded or subcordate, the apex acuminate or cuspidate, 
sometimes 25 cm. long and 19 cm. wide, the lateral nerves 5-8; 
staminate panicles very minutely puberulent, to 60 cm. long and 
45 cm. wide, the primary branches 20-30 cm. long, the secondary 
branches 3-5 cm. long; flowers yellowish white; outer sepals ovate, 
the broader inner ones 2-3.5 mm. long; petals 1.5-3 mm. long; 
anther column 1.5-2.5 mm. long. 

Loreto: San Antonio, Ule 6696; type of S. Ulei. Brazil; Guianas. 

7. SYNANDROPUS A. C. Smith 

A woody vine, the branchlets with loose, thin bark, the serrate, 
petiolate leaves palmately nerved at the base, pinnately nerved 
above. Pistillate inflorescence (staminate unknown) ample, panicled, 
with spreading branches, arising below the leaves, the flowers fasci- 
cled. Sepals 6, the outer small, the inner ovate, thin. Petals 3, 
opposite the 3 stamens whose anthers they enclose. Allied to Odon- 
tocarya and Somphoxylon, from both of which it differs in its less 
completely fused stamens, in having 3 instead of 6 petals, and 
usually, further, from the first in having only 3 stamens. The 
author suggests that each petal represents a complete fusion of 2. 
The validity of the genus evidently must depend on characters of 
pistillate flowers or fruit, because elsewhere, as in Disciphania, the 
number of petals and stamens may vary, and the number of stamens 
and degree of stamen connation varies in Odontocarya. Under these 
circumstances the validity of the present genus is extremely doubtful. 

Synandropus membranaceus A. C. Smith, Bull. Torrey Club 
58: 93. 1931. 

Petioles slender, at first pilose, 1-5 cm. long; blades oblong, cor- 
date at the base, obtusely acuminate, 8-10 cm. long, 3-4.5 cm. wide, 



FLORA OF PERU 695 

glabrous or sparsely pilose beneath, 5-9-nerved; inflorescence gla- 
brous, to 25 cm. long, minutely bracteolate; pedicels to 1.5 mm. long; 
inner sepals 1.5 mm. long; petals 1 mm. long and broad, the margins 
inflexed; filaments fused half their length; anthers 2-celled, opening 
by lateral, vertical clefts. Probably to be found in Amazonian Peru. 

Peru: Probably. Amazonian Brazil. 

8. HYPERBAENA Miers 

Leaves coriaceous, entire, and, except for the 3-5 basal nerves, 
pinnately nerved. Staminate inflorescence paniculate, with cymose 
branchlets, the pistillate less branched. Sepals 6, the 3 inner ones 
larger, all imbricate. Petals 6, somewhat fleshy. Stamens 6, free, 
vertically and laterally dehiscent. Carpels 3, free, gibbous. Endo- 
sperm none. 

Hyperbaena domingensis (DC.) Benth. Journ. Linn. Soc. 5: 
Suppl. 2: 50. 1861; 200. Cocculus domingensis DC. Syst. 1:528. 1818. 

Scandent, the branchlets and bractlets of the finely branched, 
lax inflorescences rusty-pilose; petioles 1.5-5 cm. long, geniculate 
below the blade, this mostly 5-8 cm. wide, 8-10 cm. long, glabrous 
in age, lustrous above, the basal nerves usually not more conspicuous 
than the lateral ones; inflorescences 5-20 cm. long or longer, the 
staminate decompound, the pistillate less branched; outer sepals 1.5 
mm. long, 1 mm. broad or narrower, the inner scarcely longer, strongly 
concave, nearly as broad as long; petals subrotund, about as long as 
the sepals; stamens 1 mm. long; pistillate flowers similar, the stami- 
nodia 0.4 mm. long; drupes 1.2-1.6 cm. long, scarlet, ripening black- 
violet, lustrous. Illustrated, Deless. Icon. 1: pi. 96; Pflanzenreich 
IV. 94: 198 (flowers and fruit). 

Libertad: Ongon, 1,200 meters, Weberbauer 7061. Bolivia to the 
Guianas and West Indies. 

9. ANOMOSPERMUM Miers 

Leaves entire, reticulate-veined. Flowers axillary or supra- 
axillary. Sepals 6, the outer minute, the much larger inner ones 
elliptic or suborbicular. Petals 6, smaller than the inner sepals. 
Anther cells longitudinally dehiscent. Pistillate flowers with 6 
staminodia. Carpels 3, free; stigma ligulately dilated. Drupes 
lustrous, coriaceous. Endosperm abundant. 

Leaves not triplinerved. 

Leaves glabrous; inner sepals 6-8 mm. long A. Schomburgkii. 



696 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves minutely puberulent beneath; inner sepals about 1 mm. long. 

A. minuliflorum. 
Leaves triplinerved. 

Leaves coriaceous, very densely and minutely reticulate-veined 
beneath, the nerves scarcely elevated on the upper surface. 

A. chloranthum. 

Leaves rather thin, laxly reticulate-veined, the nerves strongly 
elevated on the upper surface A. japurense. 

Anomospermum chloranthum Diels, Notizbl. Bot. Gart. 
Berlin 6: 132. 1914. 

Glabrous except the sparsely pilose younger branches; petioles 
5-6 cm. long; blades coriaceous, broadly obovate, triangularly 
narrowed to the base, acuminate, 10-12 cm. long, about half as wide, 
nearly triplinerved; staminate flowers paniculate, on leafless branch- 
lets 10-18 cm. long and 2-3 cm. broad; peduncle 2 cm. long; flowers 
greenish, the outer sepals 1.2 mm. long, 1 mm. wide, the inner 5 mm. 
long, nearly as broad; petals 2-2.5 mm. long and wide, connivent 
over the stamens, these 1.7 mm. long, with small, erect or slightly 
oblique anthers. Distinct by its broadly obovate, nearly 3-nerved 
leaves, but in the absence of fruit the relationship is unknown. Tess- 
mann 4579 has nearly elliptic but triplinerved leaves. Neg. 4996. 

Loreto: Rio Itaya, Williams 3414- Pongo de Manseriche, Tess- 
mann 4689, 4579. Rio Acre: Seringal San Francisco, Ule 9388, type. 

Anomospermum japurense (Mart.) Eichl. Flora 47: 388. 1864. 
Cocculus japurensis Mart. Flora 24: Beibl. 2: 44. 1841. 

Leaves slender-petiolate, the blades thin, glabrous, concolorous, 
lustrous, ovate or lance-ovate, acutely short-cuspidate, usually 
rounded at the base, 7.5-10 cm. long, 3-5 cm. wide; flowering pedicels 
2-6 mm. long; drupes obovoid, almost 3 cm. long and 1.5 cm. in 
diameter. 

Loreto: Yurimaguas, in forest, 200 meters, Williams 3860 (det. 
Moldenke). Amazonian Brazil. 

Anomospermum minutiflorum Diels, Notizbl. Bot. Gart. 
Berlin 9: 996. 1926. 

A liana as much as 25 meters long, the stems to 17 cm. thick; 
petioles straight, thickened throughout, glabrous; blades coriaceous, 
minutely reticulate-veined on both sides, lustrous, minutely pilose 
beneath, broadly elliptic or oblong-elliptic, acuminate, 10-15 cm. 
long, 5-6.5 cm. wide, the primary nerves 6-8, obscure above; panicles 



FLORA OF PERU 697 

solitary or binate, to 15 cm. long, the subspicate cymules only 2.5 mm. 
long, the rachis and peduncle cinereous-pilose; flowers greenish, the 
outer sepals 0.5 mm. long, the inner broadly elliptic-obovate, 1.2 
mm. long; petals subspatulate, 0.5 mm. long, the stamens 1 mm. 
longer. Neg. 27510. A. Ulei Diels, Amazonian, has larger, ovate, 
merely acutish leaves and smaller flowers, the inner sepals about 
4 mm. long. 

Loreto: Mouth of Rio Santiago, in upland woods, Tessmann 
4565, type; 4626. 

Anomospermum Schomburgkii Miers, Ann. Nat. Hist. II. 
7:39. 1851; 186. 

Glabrous except for the puberulent branchlet tips and the cilio- 
late, bracteolate outer sepals; petioles 1-1.5 cm. long; blades coria- 
ceous, lustrous above, oblong-elliptic, acuminate, mucronulate, 
about 10 cm. long, 3.5-4 cm. wide, with 2 marginal basal nerves free 
to the third or fifth lateral nerve, finely reticulate; flowers borne 
loosely on leafy or leafless branchlets, solitary or few-fascicled; 
pedicels 1-1.5 cm. long; bractlets broadly triangular; outer sepals 
broader than long, scarcely 1 mm. long; inner sepals roundish, 6-8 
mm. long, glabrous, yellow; petals very fleshy, strongly arched 
apically to enclose the stamens or staminodia, 2.5-3 mm. long and 
about as wide; drupes straight, 2-2.5 cm. long, fleshy-coriaceous, 
with ligneous endosperm. The Peruvian plant is var. lucidum 
(Miers) Diels, with the leaves exceedingly lucid above and obsoletely 
veined. Illustrated, Pflanzenreich IV. 94: 187. Neg. 30148. 

Loreto: Iquitos, Klug 1370. Maquisapa, upper Rio Nanay, 
climbing over trees and shrubs, Williams 1191. San Martin: 
Zepelacio, 1,100 meters, in forest, Klug 3739, 3756; flowers yellow. 
Brazil; Guianas. 

10. ELISSARRHENA Miers 

Well marked by the fistulous, tomentose branches, long-petioled, 
glabrous leaves, and tomentulose, short-peduncled, congested in- 
florescence. Flowers (pistillate unknown) with 3 minute, broadly 
ovate and 3 much larger, suborbicular sepals. Petals 6, glabrous, 
broadly reniform. Stamens 6, free, the filaments exteriorly strongly 
convex. Anther cells horizontally valvate-dehiscent. 

Elissarrhena grandifolia (Eichl.) Diels, Pflanzenreich IV. 94: 
189. 1910. Anomospermum grandifolium Eichl. in Mart. Fl. Bras. 
13, pt. 1: 169. pi. 37. 1864. 



698 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles 5-14 cm. long; blades thin, paler beneath, narrowed or 
truncate at the base, obtuse or acute at the apex, sometimes 30 cm. 
long and 15 cm. wide, with 3-5 basal nerves; inflorescence 2.5-5 cm. 
long; bracts very minute; larger sepals 4 mm. long, 3.5 mm. broad. 
Illustrated, Pflanzenreich IV. 94: 189. Neg. 19130. 

Loreto: Pongo de Manseriche, Tessmann 3834- Balsapuerto 
(Killip & Smith 28665; det. Smith). San Antonio, on Rio Itaya 
(Killip & Smith 29337; det. Smith). Rio Acre: Ule 9389. Brazil. 

11. ABUTA Aubl. 

Leaves coriaceous. Staminate inflorescence usually panicled and 
compound, the pistillate laxly racemose. Sepals 6, the outer bract- 
like. Petals minute or none. Stamens 6, connate or free, the cells 
dehiscent by vertical or transverse slits. Pistillate flowers with 6 
or fewer staminodia. Carpels 3, free, with sessile, subulate, recurved 
stigmas. Drupes short-stipitate or attenuate at the base. Endo- 
sperm present. 

Leaves large, mostly 16-30 cm. long, the nerves very prominent on 
the upper surface A. grandifolia. 

Leaves small, chiefly 10 cm. long or less, the nerves not prominent 
on the upper surface A. Klugii. 

Abuta grandifolia (Mart.) Sandwith, Kew Bull. Misc. Inf. 
1937: 397. 1937. A. concolor Poepp. & Endl. Nov. Gen. & Sp. 2: 
64. pi. 188. 1838; 197. 

A shrub or small tree with strict, finally glabrous branches; 
petioles tumid at the base, gradually enlarged toward the apex, 
2.5-8 cm. long; blades pale green, glabrous, ovate-oblong or nearly 
oblanceolate, acuminate or cuspidate, 10-20 cm. long, 6-12 cm. wide, 
with 3 very prominent, palmate nerves; staminate inflorescence 
2-8 cm. long; anthers transversely dehiscent, the cells finally often 
confluent, the filaments glabrous; drupes ellipsoid, glabrous, yellow- 
ish, 2-2.5 cm. long. A. panurensis Eichl., of northern Brazil, 
similar, has hispidulous filaments. A. Grisebachii Triana & Planch, 
and A. Imene (Mart.) Eichl., both Amazonian, have longitudinally 
dehiscent anthers; the leaves of the former are tomentulose beneath, 
of the latter, glabrous. The roots of A. concolor, boiled in water, 
serve as a remedy for anemia (Williams). 

San Martin: Pongo de Cainarachi, Klug 2757. Loreto: Caballo- 
cocha, Williams 2435. Tira Doble, Rio Nanay, Williams 1051. 
Iquitos, Williams 8013; Tessmann 5334. Rio Itaya, Williams 3500. 



FLORA OF PERU 699 

Yurimaguas, Williams 4713; Killip & Smith 29101, 28708. Mishu- 
yacu, 100 meters, in forest, Klug 2526; a tree of 7 meters. Balsa- 
puerto, 220 meters, in forest, Klug 3006; a tree of 4 meters. 
Between Yurimaguas and Balsapuerto, 150 meters, Killip & Smith 
28295. Rio Acre: Ule 9387, 9386. "Sanango," "caimitillo," 
"trompetero-sanango." French Guiana; Brazil; Bolivia. 

Abuta Klugii Moldenke, ined. 

A woody vine, with the aspect of Anomospermum, the slender 
branches glabrous; leaves slender-petiolate, the blades ovate to 
oblong-ovate or elliptic-ovate, abruptly acuminate, obtuse or 
narrowly rounded at the base, 3-nerved, coriaceous, somewhat 
lustrous, the finer venation very inconspicuous; inflorescence sparsely 
griseous-sericeous. 

Loreto: Mishuyacu, near Iquitos, 100 meters, in forest, Klug 
1415, type. Also in Amazonian Brazil. 

72. WINTERACEAE 

Reference: Robert P. McLaughlin, Trop. Woods 34: 21-22. 1933. 
This family, which often has been included in Magnoliaceae, has 
a different wood structure (cf. McLaughlin, op. cit. 35), and is 
distinguished readily by its exstipulate, pellucid-punctate leaves and 
never conelike fruits, the carpels being arranged circularly more or 
less in a single whorl. 

1. DRIMYS Forst. 

An evergreen shrub or tree with light-colored, smooth, aromatic 
bark. Sepals and petals in whorls. Flowers in axillary or terminal 
cymes or umbels. Stamens several, the anther cells distinct. 

Drimys Winter! Forst. Char. Gen. 84. pi. 42. 1776. D. grana- 
tensis Mutis ex L. f. Suppl. 269. 1781. D. Winteri Forst. f. granatensis 
Eichl. in Mart. Fl. Bras. 13, pt. 1: 135. 1864. 

Leaves oblong-obovate, 7-13 cm. long, more or less glaucous 
beneath; flowers white, with 6 or more petals; fruit purplish black. 
Sometimes a tree to 18 meters. Botanists are not in agreement as 
to the specific limitation of this widely distributed plant, which 
exhibits great variation in the development of the inflorescence and 
in the leaves; cf. Miers, Contr. Bot. 1: 123-137. pi. 25-27. 1851-61, 
and Hauman, Comm. Mus. Nac. Buenos Aires 2: 50. 1923, who 
recognize a number of species. Eichler, on the other hand, in Mart. 
Fl. Bras. 13, pt. 1: pi. 30, 31, considers that there is one species 
with a number of forms. 



700 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Eichler and many others have related the story of the plant's 
discovery by Captain Winter who, accompanying Sir Francis Drake 
on the circumnavigation of 1577-1580, remained in the Magellan 
region after a storm to let his men recuperate, and used the bark of 
Drimys against scurvy, after which time it enjoyed great popularity 
in medicine for many years. It is now used only locally or as a 
condiment. 

Amazonas: Chachapoyas (Weberbauer, 262). Central and South 
America; Australia; New Caledonia. "Winter's bark," "canela." 

73. ANNONACEAE. Anona Family 
By Rob. E. Fries 

Ligneous plants with the leaves alternate, entire, and exstipulate. 
Flowers generally trimerous. Sepals 3, rarely 2, valvate or imbricate. 
Petals generally 6, in 2 series, valvate or imbricate, the inner some- 
times rudimentary or absent. Stamens generally numerous; anther 
cells adnate, the connective usually expanded above the anther and 
truncate. Carpels numerous, rarely few or solitary, generally 
free; ovules 1 or more. Fruiting carpels sessile or stipitate, free 
(monocarps) or united in a dry or fleshy mass. Seeds with or without 
an aril, with copious, ruminate endosperm and minute embryo. 
The large family Annonaceae, which is of pan tropic distribution, 
is represented in America by 33 genera, of which 17 are represented in 
the flora of Peru. For some years the author has been engaged on 
a monographic revision of the American genera and has published 
his results in Acta Horti Bergiani, volumes 10 and 12, to which the 
reader is referred. This revision is now nearly completed; of the 
South American genera only the largest and also the most difficult 
genus, Guatteria, remains. A preliminary revision has been under- 
taken but it can not be helped that the synopsis of the Peruvian 
Guatterias given below should be regarded as provisional. Further 
researches certainly will show that many additions and corrections 
will be necessary. 

The difficulty in the taxonomic treatment of Annonaceous plants 
is above all due to the fact that the material accessible at present 
is frequently incomplete. Numerous species are known with flowers 
only, others only in the fruiting stage. For this reason a comparison 
of the species is often impossible, which is particularly evident when 
it comes to the construction of the analytic keys. The keys given 
here are thus in many cases quite artificial and do not pretend to 



FLORA OF PERU 701 

show the true relations of the species. The same applies also to the 
arrangement of the genera. 
Petals imbricate. 

Pedicels with bracts; anthers not transversely septate. 
Pubescence of simple hairs. 
Ripe carpels free. 
Flowers axillary. 
Ovules basal. 

Pedicels with a bract above the articulation; connec- 
tive of the stamens not expanded above the 

anther 1. Oxandra. 

Pedicels without a bract above the articulation; con- 
nective expanded and truncate above the anther. 

2. Guatteria. 

Ovules lateral, sometimes near the upper end of the ovary. 
Petals rotund to oblong, obtuse. 
Pedicels without bracts above the articulation. 

3. Pseudoxandra. 
Pedicels with bracts above the articulation. 

4. Cremastosperma. 
Petals linear-lanceolate, gradually attenuate to the 

acute apex 5. Ruizodendron. 

Flowers opposite the leaves 6. Malmea. 

Carpels united into a fleshy mass in fruit 7. Fusaea. 

Pubescence of stellate hairs or stellate scales 8. Duguetia. 

Pedicels entirely without bracts; anthers transversely septate. 

9. Porcelia. 
Petals, at least the outer ones, valvate. 

Pedicels entirely without bracts; flowers solitary; inner petals 
large, saccate, with involute margins; anthers transversely 

septate 10. Cymbopetalum. 

Pedicels with bracts; inner petals not saccate. 
Ovules more than one, lateral. 
Anthers not transversely septate. 

Flowers spherical in bud, all bisexual; petals rotund-ovate. 

11. Unonopsis. 

Flowers elongate in bud, polygamous or dioecious; petals 
linear-oblong 12. Diclinanona. 



702 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Anthers transversely septate 13. Xylopia. 

Ovules 2, basal, erect 14. Anaxagorea. 

Ovule 1, basal or nearly so. 

Flowers axillary; monocarps free, stipitate. 

Pedicels long, with a bract above the articulation. 

11. Unonopsis. 

Pedicels very short (flowers nearly sessile), not bracteate 
above the articulation 15. Guatteriopsis. 

Flowers not axillary; carpels united into a more or less fleshy 
mass in fruit. 

Outer petals without wings or spurs 16. Annona. 

Outer petals produced into wings or spurs .... 17. Rollinia. 

1. OXANDRA A. Rich. 

Trees or shrubs. Flowers small, solitary in the axils or in few- 
flowered, axillary inflorescences; pedicels articulate a little above the 
base, bearing a few small, distichous bracts below and one above 
the articulation. Sepals imbricate in bud, persistent. Petals all 
rather alike, thin, orbicular, ovate, or oblong, imbricate, black when 
dried. Stamens 6-20, the connective elongate above the anther into 
a triangular to lanceolate appendage. Carpels rather few (to 13); 
ovule 1, basal, erect. Monocarps shortly stipitate or nearly sessile, 
one-seeded. A medium-sized genus, centering about the Amazon 
and its tributaries and in Guiana (15 species); one of them and 3 
others occur in Colombia and Panama, 2 in the West Indies, and 
2 in southern Brazil. 

Leaves acute or rounded at the base, without a tooth on each side 
of the petiole. 

Leaves cuneate at the base, not verrucose. 

Flowers several, in branched inflorescences; pedicels minutely 
hairy 1.0. Riedeliana. 

Flowers 1-2; pedicels glabrous 2. 0. Espintana. 

Leaves rounded or very shortly acute at the base, more or less 
verrucose. 

Leaves small, up to 2 or rarely 3 cm. broad; midrib deeply sunken 
on the upper surface 4. 0. acuminata. 

Leaves larger, more than 3 cm. broad; midrib flat or prominent 
on the upper surface. 



FLORA OF PERU 703 

Monocarps spherical, on stipes 1 cm. long; leaves rounded, 
but not emarginate, at the base 3. 0. sphaerocarpa. 

Monocarps ellipsoidal, on short stipes 2-3 mm. long. 

Leaves glabrous, rounded but not emarginate at the base. 

5. 0. euneura. 

Leaves hairy beneath and on the midrib above, rounded 
and slightly emarginate at the base. 6. 0. macrophylla. 

Leaves truncate at the base, with a small tooth on each side of the 
petiole 1.0. xylopioides. 

1. Oxandra Riedeliana R. E. Fries, Arkiv Bot. 5, pt. 4: 2. 

pi. 2, f. 7-9. 1905. 

A small tree, 15 meters high; youngest branchlets minutely 
sericeous and soon glabrous; leaves membranaceous, usually yellowish 
green when dried, glabrous and lustrous, lanceolate or rhombic- 
lanceolate, broadest at the middle and equally narrowed at both ends, 
cuneate and very acute at the base, long-acuminate to the obtuse 
apex, 8-12 cm. long, rarely longer; midrib prominent on both sides; 
flowers several, in small, sessile inflorescences; pedicels, bracts, and 
sepals minutely appressed-hairy; flower buds ovoid; sepals about 
1 mm. long, ciliate; petals glabrous, ciliate, elliptic to oblong- 
elliptic, about 6 mm. long. Neg. 19322. 

Loreto: Iquitos, Tessmann 5383. Known also from several 
localities in Brazil, on the Amazon and its southern tributaries. 

2. Oxandra Espintana (Spruce) Baill. Hist. PL 1: 207. 1868. 
Bocagea Espintana Spruce ex Benth. Journ. Linn. Soc. 5: 71. 1861. 
0. aromatica Tr. & PL Ann. Sci. Nat. IV. 17: 36. 1862. B. aromatica 
Britton, Bull. Torrey Club 16: 14. 1889. 0. ovata Rusby, Descr. 
New Sp. S. Amer. PL 19. 1920. 

A small tree, 10 meters high; leaves ovate-elliptic, lanceolate or 
obovate-lanceolate, usually gradually narrowed at the apex, cuneate 
at the acute or rarely obtuse base, glabrous, lustrous and, finally, 
reticulate on both sides, 6-12 cm. long; midrib a little impressed on 
the upper side; flowers 1-2, on very short, glabrous pedicels 1-2 cm. 
long; bracts, sepals, and petals glabrous; flower buds globular to 
ovoid. Negs. 21296, 27550. 

San Martin: Tarapoto, Spruce 4920 (type). Juanjui, upper Rio 
Huallaga, Klug 3814. Also in Colombia and western Amazonian 
Brazil. 



704 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3. Oxandra sphaerocarpa R. E. Fries, Acta Hort. Berg. 12: 
198. 1934. 

Leaves membranaceous, oblong to elliptic-oblong, 1-1.5 mm. long, 
cuspidate, rounded at the base, glabrous above, with a few hairs on 
the midrib when young, covered beneath with long, appressed hairs, 
finally more or less glabrescent, rather thinly verrucose on both 
sides; midrib flat above; inflorescences 1-2-flowered; fruits on rigid, 
glabrous pedicels 5-7 mm. long; monocarps spherical, glabrous, on 
stipes 1 cm. long. 

Loreto: Lower Rio Nanay, Williams 475 (type), 478. 

4. Oxandra acuminata Diels, Bot. Jahrb. 37: 410. 1906. 

A tree up to 20 meters high; young branchlets densely covered 
with erect, golden-ferruginous hairs, soon glabrescent; leaves small, 
lanceolate-oblong, more or less abruptly contracted at the apex into 
a cusp 2-3 cm. long, rounded or shortly acute at the base, glabrous, 
densely verruculose beneath, up to 10 cm. long and 3 cm. broad; 
midrib deeply impressed above; inflorescences sessile in the axils 
of detached leaves, 1-3-flowered; pedicels glabrous, 2-3 mm. long; 
bracts and calyx sericeous and ciliate; flower buds short-cylindric- 
trigonous; petals 6-7 mm. long. 

Junin: Tarma, near La Merced, Weberbauer 1897 (type). Also 
in Brazil on the rivers Jurua and Madeira. 

5. Oxandra euneura Diels, Notizbl. Bot. Gart. Berlin 10: 173. 
1927. 

A small tree, 8 meters high, the young branchlets covered with a 
short and dense fulvous pubescence; leaves very shortly petiolate 
(1-2 mm. long), oblong to elliptic, very abruptly contracted at the 
apex into a cusp 1-1.5 cm. long, rounded but not emarginate at the 
base, glabrous and densely verruculose-punctate on both sides, 10-20 
cm. long and 4-6.5 cm. broad; midrib prominent on both sides; 
pedicels 3-4 mm. long, pubescent; monocarps ovoid, nearly sessile. 

Loreto: Yurimaguas, Killip & Smith 28029, 28036. Mishuyacu, 
King 135. Lower Rio Itaya, Soledad, Tessmann 5320 (type). "Es- 
pintana." 

6. Oxandra macrophylla R. E. Fries, Acta Hort. Berg. 10: 
173. 1931. 

A shrub or small tree, the young branchlets densely covered with 
short, patent, fuscous hairs; petioles only 1-2 mm. long; lamina 
large (17-25 cm. long and 6-8 cm. broad), ovate to oblong-ovate, 



FLORA OF PERU 705 

abruptly narrowed into a cusp 1.5-2 cm. long, rounded and a little 
cordate at the base, very densely verruculose-punctate on both sides; 
pedicels of the fruits glabrous, 3-4 mm. long; monocarps ellipsoidal, 
on stipes about 3 mm. long. 

Loreto: Yurimaguas, Killip & Smith 28752 (type) ; Williams 4129. 

7. Oxandra xylopioides Diels, Notizbl. Bot. Gart. Berlin 10: 
172. 1927. 

A tall tree, reaching 20-25 meters; leaves rigid, linear-oblong, 
gradually narrowed to the apex, more or less truncate at the base 
with a small tooth on each side of the short petiole, 8-12 cm. long and 
about 1.5 cm. broad, both sides smooth, the upper glabrous, the 
lower covered when young with long, appressed hairs but very soon 
quite glabrous; midrib impressed above, prominent below, the nerves 
and veins not conspicuous; flowers solitary on short, densely brac- 
teate pedicels; petals narrowly ovate, 7-8 mm. long. Neg. 27553. 

Loreto: Upper Maranon, at the mouth of Rio Santiago, Tess- 
mann 4572 (type). Yurimaguas, Killip & Smith 28317. Amazonian 
Brazil. "Espintano." 

2. GUATTERIA R. & P. 

Shrubs or trees, the pubescence of simple hairs. Sepals valvate. 
Petals imbricate, usually hairy, especially so outside at the base. 
Stamens numerous; connective expanded above the anther into a 
truncate disk. Carpels numerous; ovule 1, basal, erect. Monocarps 
ovoid or ellipsoid, or shortly cylindrical-ellipsoid, one-seeded, sti- 
pitate. The genus is the largest of the family Annonaceae, the 
species numbering nearly 200. All are American, with a distribution 
from southern Mexico and the West Indies to Bolivia and southern 
Brazil. 

Flowers terminal 1. G. terminates. 

Flowers axillary. 

Leaves not verruculose-punctulate. 

Leaves small or larger, up to 20-25 cm. long; pedicels generally 
more than 1 cm. long; monocarps (when known) ovoid, 
ellipsoid, or pyriform. 
Lower side of the leaves more or less hairy but never entirely 

covered with appressed, persistent hairs. 
Leaves small to medium-sized (less than 7 cm. broad). 
Leaves lanceolate to narrowly oblong. 
Pedicels hairy. 



706 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Young branchlets very densely covered with short, 
patent hairs; connective disk short-hairy. 

2. G. Schomburgkiana var. holosericea. 
Young branchlets with sparse, more or less patent 
hairs or appressed-hairy; connective disk gla- 
brous. 

Young branchlets and the lower side of the leaves 

with long and more or less patent hairs; 

midrib hairy above, at least when young. 

Leaves rounded or acute at the base, but not 

emarginate. 

Pedicels 1-1.5 cm. long. 
Flowers small; petals up to 12 mm. long. 

3. G. Augusti. 
Flowers larger; petals 20-25 mm. long. 

4. G. alutacea. 
Pedicels 2-4 cm. long. 

Leaves lanceolate, long-acute at the base, 

1.5-2.5 cm. broad 5. G. Klugii. 

Leaves oblong to lanceolate, rounded or 
very shortly acute at the base, 3-4 cm. 
broad. 

Leaves glabrous above when young 
(except on the midrib); branchlets 

thinly hairy 6. G. juruensis. 

Leaves hairy above when young; branch- 
lets with dense, patent hairs. 

Leaves usually rounded-acute at the 
base; network of the veins dense 
and nicely prominent beneath. 

7. G. hirsuta. 
Leaves oblong, rounded at the base; 

network with larger meshes. 

8. G. excelsa. 
Leaves rounded and emarginate at the base. 

9. G. trichoclonia. 

Young branchlets and the lower side of the leaves 
with short and appressed hairs; midrib quite 
glabrous above. 



FLORA OF PERU 707 

Leaves 15-20 cm. long, with a long and very 
sharp acumen 10. G. acutissima. 

Leaves 9-12 cm. long, contracted into a short 
and blunt cusp 11. G. elata. 

Pedicels quite glabrous, black and lustrous. 

12. G. modesta. 

Leaves oblanceolate (broadest above the middle). 
Leaves with a cusp 1-1.5 cm. long. . 13. G. Tessmannii. 

Leaves obtuse at the apex, not cuspidate. 

14. G. chlorantha. 

Leaves elliptic, rarely elliptic-obovate. 
Leaves long-cuneate-acute and decurrent at the base. 
Leaves 8-12 cm. long and 3-5 cm. broad; pedicels 
and flowers ferruginous-tomentose. 

15. G. eugeniaefolia. 

Leaves 15-20 cm. long and 5-7 cm. broad; pedicels 
and flowers gray-sericeous. 

16. G. phanerocampta. 

Leaves rounded or very shortly acute at the base. 
Petals not glaucous, from a narrow base obovate, 

the inner ones retuse. 

Flowers and pedicels rufous-pilose; sepals decidu- 
ous 17. G. calliantha. 

Flowers and pedicels ferruginous-hairy; sepals 

not deciduous 18. G. coeloneura. 

Petals glaucous, from a broad base oblong-ovate, all 

rounded at the apex. 
Leaves shortly acute at the base. 

19. G. pleiocarpa. 
Normal leaves rounded at the base. 

Monocarps rounded at the apex, about 12 mm. 
long; pericarp fleshy; stipes about 15 mm. 

long 20. G. glauca. 

Monocarps apiculate, with a dry pericarp, 
8-9 mm. long; stipes 3-10 mm. long. 

21. G. juninensis. 

Leaves about 20 cm. long or more, 7-10 cm. broad, elliptic 
to elliptic-obovate. 



708 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves with patent hairs beneath 22. G. ucayalina. 

Leaves with appressed hairs beneath. 

Leaves dark red -brown when dried, smooth above; 
upper side of the midrib hairy toward the base. 

23. G. Guentheri. 

Leaves yellowish green when dried, with prominent 
veins above; midrib quite glabrous above. 

24. G. spectabilis. 
Lower side of the leaves entirely covered with appressed, 

persistent hairs 25. G. hyposericea. 

Leaves very large (when normally developed 30-50 cm. long), 
oblong and long-petiolate; pedicels 5-10 mm. long; mono- 
carps large, short-cylindric, on stipes 3-6 mm. long. 
Leaves acute at the base; sepals 6-8 mm. long; petals lanceo- 
late, narrowed to the obtuse apex ... 26. G. megalophylla. 
Leaves rounded at the base; sepals more than 1 cm. long; 
petals broadly obovate, broadly rounded at the apex. 

27. G. melosma. 
Leaves verruculose-punctulate. 

Leaves medium-sized (rarely more than 20 cm. long) and 
generally broadest at the middle (if cuneate, not more than 
6 cm. broad). 
Leaves rounded or rounded-acute at the base. 

29. G. macrocarpa. 
Leaves long-acute at the base. 
Verrucae on the leaves rather sparse, not touching each other. 

Petioles 2-3 mm. long 28. G. Dielsiana. 

Petioles 8-10 mm. long 30. G. peruviana. 

Verrucae very small and dense, more or less touching each 

other 31. G. decurrens. 

Leaves large (20-35 cm. long), obovate or oblanceolate. 

Young branchlets appressed-hairy 32. G. amazonica. 

Young branchlets covered with patent or reflexed hairs. 
Leaves obovate, gradually attenuate into a narrow, obtuse 
base; petals linear-oblong, retuse at the truncate apex. 

33. G. multivenia. 

Leaves oblanceolate, narrowed to the broad, rounded base; 
petals ovate-elliptic, obtuse to rounded at the apex. 

34. G. excellens. 
Insufficiently known species 35. G. eriopoda. 



FLORA OF PERU 709 

1. Guatteria terminalis R. E. Fries, sp. nov. Guatteria glauca 
Hohenacker in sched., non R. & P. 

Ramuli novelli primo ferrugineo-, dein cinerascenti-sericei; 
foliorum petiolus 2-4 mm. longus; lamina rigida, lucida, supra ab 
initio glabra, subtus primo adpresse hirsuta demumque glabra, late 
rhombeo-lanceolata, basin versus cuneato-acutata et lato-decurrens, 
apice brevius et obtusiuscule acutata, 5-9 cm. longa et 2.5-3.5 cm. 
lata; reticulum venularum densissimum, utroque latere elevatum; 
flores solitarii in ramulis foliosis terminales; pedicellus articulatus, 
supra articulationem 1.5-2 cm. longus, rigidus, sericeus; sepala 
triangulari-ovata, ad 5 mm. longa; petala extus densissime longeque 
sericea; carpella 40-50. 

Peru: Tatanara, Lechler 2648 (type in Herb. Upsal.). 

2. Guatteria Schomburgkiana Mart. Fl. Bras. 13, pt. 1: 38. 

1841. Guatteria vestita Kl. in R. Schomb. Fauna und Flora Brit. 
Guiana 979. 1848 (nomen nudum). Annona Hostmannii Steud. Flora 
26, pt. 2: 754. 1843. 

2a. Guatteria Schomburgkiana var. holosericea R. E. Fries, 
var. nov. 

Ramuli novelli, petioli et nervus medius subtus pilis densissimis 
brevibus patentissimis rubiginosis vestiti, holosericei; folia lanceo- 
lata, sat abrupte in cuspidem 1-1.5 cm. longam contracta, basi 
rotundato-acuta et breviter decurrens, supra demum glabra et lucida, 
8-9 cm. longa et ca. 2.5 cm. lata; flores parvi, 5-10 mm. longe 
pedicellati; petala exteriora oblongo-oblanceolata, ca. 8 mm. longa 
et 3.5 mm. lata, interiora obovato-elliptica, 4-5 mm. lata. Con- 
nectivi appendix breviter pilosa. Neg. 14210. 

Loreto: Florida, Rio Putumayo, at mouth of Rio Zubineta, in 
forest, Klug 2259 (type in Herb. Mus. Stockholm). Huitoto Indian 
name, "Ag-guio." 

The variety differs from the type (of Guiana) especially in 
the very dense pubescence of short, patent hairs. 

3. Guatteria Augusti Diels, Notizbl. Bot. Gart. Berlin 9: 51. 
1924. 

A tree about 10 meters high, young branchlets covered with 
loosely appressed hairs; leaves on petioles 2-3 mm. long, rigid, gla- 
brous above except on the midrib, sparsely clothed beneath with 
long, decumbent hairs, lanceolate-oblong, short-acute at the base, 
attenuate at the apex into a long, narrow acumen, 7-11 cm. long 



710 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and 2-3 cm. broad; flowers 1-2, pedicels 1-1.5 cm. long, pilose; 
sepals about 7 mm. long, broadly ovate, acuminate, long-hairy out- 
side, tomentose inside; corolla finally stellate-expanded; petals long- 
sericeous outside, reaching 12 mm. in length, the outer lanceolate, 
rather acute, about 4 mm. broad, the inner ovate, obtuse, 5-6 mm. 
broad. 

Peru: On the Rio Mixiollo, a left tributary of Rio Huallaga, 
below Ongon, altitude 1,200-1,300 meters, Weberbauer 7062 (type). 

4. Guatteria alutacea Diels, Verh. Bot. Ver. Brandenb. 47: 
126. 1905. 

A shrub or small tree, 5-10 meters high, the young branches 
with patent hairs; leaves on petioles about 4 mm. long, sericeous 
above when very young, soon glabrescent, appressed-hairy beneath, 
oblong-lanceolate and long-acuminate, acute at the base, 10-18 
cm. long and 2.5-4 cm. broad; pedicels generally solitary, rigid, 
pilose, 10-15 mm. long; sepals finally recurved, broadly ovate, long- 
sericeous outside, 5-7 mm. long; petals puberulous, yellow-sericeous 
outside at the base, oblong-ovate, the inner 20-25 mm. long, the 
outer a little shorter. Neg. 27536. 

San Martin: Tarapoto, Ule 6427 (type); Spruce 4270; Mathews 
465, 1423; Williams 5503, 6510. 

4a. Guatteria alutacea var. angustifolia R. E. Fries, var. nov. 

A typo differt folis anguste lanceolatis, apicem versus sensissime 
longeque angustatis, basi longe acutatis, 12-16 cm. longis et 2-2.8 cm. 
latis. 

San Martin: Tarapoto, Spruce 4270a (type, in Herb. Berlin). 

5. Guatteria Klugii R. E. Fries, sp. nov. 

Frutex ca. 2 m. altus, ramulis novellis pilis patentibus flaves- 
centibus vestitis; folia 2-3 mm. longe petiolata, rigida, supra ab 
initio glabra costa impressa diu hirsuta excepta, subtus pilis adpressis 
sparse vestita, anguste lanceolata, utrinque longe acutata, 7-11 cm. 
longa et 1.5-2.5 cm. lata; pedicelli solitarii, graciles, pilosi, 3-4 cm. 
longi; sepala ovata, parce sericea, 7 mm. longa; petala lanceolata, 
extus praesertim basin versus flavescenti-sericea, ceterum tomentella, 
interiora demum 18-20 mm. longa et 8-9 mm. lata, exteriora paulo 
minora et angustiora. 

San Martin: San Roque, in forest, Williams 6976. Zepelacio, 
near Moyobamba, Klug 3299 (type, in Herb. Mus. Stockholm). 



FLORA OF PERU 711 

6. Guatteria juruensis Diels, Verh. Bot. Ver. Brandenb. 47: 
126. 1905. 

A large tree, up to 30 meters high, the young branchlets sparsely 
pilose; leaves membranaceous, on petioles 3-4 mm. long, glabrous 
above and very thinly hairy or glabrous beneath, narrowly oblong, 
rounded or very shortly acute at the base, acuminate, 10-14 cm. 
long and 3-4 cm. broad; pedicels 2-4 cm. long, solitary, slender, 
provided with a few patent hairs; sepals broadly ovate, sparsely 
hairy outside, tomentellous inside, reflexed and pendulous, 8-10 mm. 
long; petals minutely sericeous, longer-ferruginous-sericeous at the 
base outside, the inner oblong-obovate, often crenulate at the apex, 
2-2.5 cm. long and 1 cm. broad, the outer slightly smaller; stamens 
1 mm. long. Neg. 14194. 

Loreto: La Victoria on the Amazon, Williams 2864- Leticia, 
Williams 3157. Also in western Amazonian Brazil (Rio Jurua, 
Ule 5010, type). 

7. Guatteria hirsuta R. & P. Syst. Veg. Fl. Peruv. 1: 146. 
1798. 

A shrub, with young branchlets covered with long, patent, ferrugi- 
nous hairs, soon glabrescent and blackening; leaves on petioles 3-5 
mm. long, clothed at first on both sides with decumbent hairs, glabres- 
cent and lustrous above, lanceolate or lanceolate-oblong, shortly acute 
at the base and shortly decurrent, long and gradually narrowed at 
the apex into an obtuse acumen, 10-15 cm. long and 3-4 cm. broad, 
veins forming a dense network, prominent, especially on the lower 
surface; flowers 1-2; pedicels 2-2.5 cm. long, densely rufous-hirsute; 
sepals long and densely rufous-sericeous; petals not seen; mono- 
carps (immature) clavate, glabrous, obtuse, 9-12 mm. long and 5-6 
mm. thick, narrowed into a stipe 15 mm. long. Negs. 14193, 27543. 

Huanuco: "In nemoribus versus Chinchao vicum," Ruiz (type). 

8. Guatteria excelsa Poepp. ex Mart. Fl. Bras. 13, pt. 1: 38. 
1841. 

Young branchlets hirsute, the hairs long, patent, ferruginous; 
petioles about 4 mm. long; lamina membranaceous, 12-16 cm. long 
and 2.5-4 cm. broad, blackening when dried, at first appressed-hairy 
above, later glabrescent except on the midrib, the lower surface thinly, 
on the midrib densely, clothed with long, straight, patent hairs, nar- 
rowly oblong with rounded and shortly decurrent base, terminating at 
the apex in an obtuse 2-3 cm. long acumen, lateral nerves 18-20 pairs, 
the network of the slightly prominent veins with large meshes; 



712 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

flowers solitary; pedicels hirsute, 15-25 mm. long, articulate near 
the middle; sepals ovate, rufous-hirsute outside, tomentellous inside, 
about 8 mm. long; petals stellate-expanded, all rhombic-obovate 
and crenulate at the apex, rufous- tomentellous; stamens 1 mm. long. 
Neg. 14185. 

Peru: Pampayaco, Poeppig 139 (1467), type. 

As the material of G. excelsa and G. hirsuta accessible at present 
is very incomplete, it has been difficult to state exact differences 
between them (see the key). Perhaps they are not specifically 
separate. 

9. Guatteria trichoclonia Diels, Notizbl. Bot. Gart. Berlin 
11:77. 1931. 

A tree, 5-10 meters high, young branchlets densely covered with 
long, straight, patent, yellow hairs; leaves on petioles 2-3 mm. 
long, long-hirsute on both sides but especially at the margins and 
on the midrib beneath, lanceolate-oblong, acuminate, rounded and 
a little cordate at the base, 7-18 cm. long and 2.5-6 cm. broad; 
pedicels 1.5-3 cm. long, patently hairy; sepals ovate, up to 10 mm. 
long, hairy; petals broadly lanceolate, the inner a little larger, 
sericeous, nearly 2 cm. long; monocarps obovate or ellipsoid, acute 
at both ends or obtuse at the apex, 7-8 mm. long and 4-5 mm. 
thick, their stipes slender, 6-10 mm. long. Neg. 14217. 

Loreto: Mainas, Poeppig 1889. Also in Bolivia. 

10. Guatteria acutissima R. E. Fries, sp. nov. 

Arbor 9 m. alta, ramulis novellis sericeis mox glabrescentibus 
et nigricantibus; foliorum petiolus ca. 5 mm. longus; lamina 
rigida, exsiccata nigrescens, adulta supra glaberrima nitidaque, 
subtus sparse adpresse hirsuta, oblongo-lanceolata, ad medium latis- 
sima, utrinque aequaliter longeque angustata, ad basin acutam con- 
spicue decurrens, apice acumine angusto acutissimo 1-1.5 cm. longo 
ornata, 14-20 cm. longa et 4-5.5 cm. lata; flores in axillis solitarii; 
pedicelli rigidi, recti, ferrugineo-sericei, ca. 1 cm. longi; sepala 
reflexa, triangulari-ovata, extus sericea, 4-5 mm. longa; petala 
oblongo-oblanceolata, apice rotundata, 15-18 mm. longa; exteriora 
5-6, interiora 7-8 mm. lata, puberula et basi extus ferrugineo-sericea. 

Loreto: Mishuyacu, near Iquitos, in forest, Klug 1268 (type in 
Herb. Field Mus.). 

11. Guatteria elata R. E. Fries, sp. nov. 

Arbor 20-30 m. alta, ramulis novellis adpresse pilosis, cito 
glabrescentibus et nigricantibus; folia 5-7 mm. longe petiolata, 



FLORA OF PERU 713 

coriacea, exsiccata nigricantia, supra glabra laevia et lucida, subtus 
pilis laxis persistentibus adpressis obsita, oblongo-elliptica, rarius 
oblongo-oblanceolata, sat abrupte in cuspidem ca. 1 cm. longam 
obtusam contracta, basi acuta et decurrens, 9-12 cm. longa et 3-4 
cm. lata; flores 1-2, pedicellis 1-1.2 cm. longis rufo-sericeis sustenti; 
sepala ovata, extus sericea, ca. 5 mm. longa; petala exsiccata 
subglauca, extus minute sericea, primo oblongo-obovata, demum in 
laminam rotundatam dilatata, interiora demum 18 mm. longa et 
15 mm. lata, exteriora paulo minora; stamina 1 mm. longa. 

Loreto: Mishuyacu, near Iquitos, Klug 826. Also collected in 
Brazil at Sao Paulo de Olivenca, near the Peruvian frontier ( Krukoff 
8356; type in Herb. Mus. Stockholm). 

Klug 649 probably belongs to this species but differs in its more 
oblanceolate leaves, less hairy pedicels, and other details. 

12. Guatteria modesta Diels, Notizbl. Bot. Gart. Berlin 9: 
139. 1924. 

A tree with quite glabrous, black branchlets; leaves rigid, 
glabrous on both sides, lustrous above, lanceolate and broadest at 
the middle, long-attenuate and acute at both ends, decurrent at 
the base, 10-16 cm. long and 3.5-5 cm. broad; flowers solitary; 
pedicels slender, quite glabrous and blackening, about 2 cm. long; 
sepals broadly ovate, nearly glabrous, about 4 mm. long, deciduous; 
petals small, sericeous outside, elliptic-ovate, obtuse, the outer 
9-10 mm. long and 4-5 mm. broad, the inner a little shorter and 
broader; stamens 1.2 mm. long; monocarps ellipsoid, obtuse, about 
15 mm. long and 8 mm. thick, abruptly narrowed into a stipe 12-16 
mm. long. Neg. 14202. 

Loreto: Middle Rio Ucayali, Yarina-cocha, Tessmann 31+76 (type). 
Lower Rio Huallaga, Williams 4896. "Cara huasca," "espintanal." 

13. Guatteria Tessmannii R. E. Fries, sp. nov. 

Arbor ca. 15 m. alta, ramulis novellis minute ferrugineo-sericeis; 
foliorum petiolus 7-10 mm. longus; lamina supra mox glabrata et 
lucida, subtus pilis sparsis brevibus adpressis instructa vel subglabra, 
oblanceolata, basi longe cuneata et decurrens, apice in acumen 1.5-2 
cm. longum obtusum sat abrupte contracta, 12-18 cm. longa et 3.5- 
5 cm. lata; flores 2-3 axillares; pedicelli stricti, ferrugineo-sericei, 
12-15 mm. longi; sepala mox reflexa, 5-6 mm. longa; petala oblonga, 
apice rotundato-truncata et saepe crenulata, extus ferrugineo-sericea, 
12-14 mm. longa et 4.5-7 mm. lata; stamina 1 mm. longa; carpella 
numerosissima, germine rufo-hirsuto. 



714 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Upper Rio Maranon, mouth of Rio Santiago, Tessmann 
4651 (type in Herb. Berlin). 

14. Guatteria chlorantha Diels, Notizbl. Bot. Gart. Berlin 
9: 139. 1924. 

A tree with sericeous branchlets; leaves rigid, when adult glabrous 
and lustrous above, sparsely appressed-hairy beneath, 8-12 cm. long 
and 2.5-4 cm. broad, oblanceolate, gradually narrowed to the very 
acute base, and long-decurrent on a petiole 5-10 mm. long; flowers 
issuing from the lower part of the branchlets, 1-2 in the axils of 
fallen leaves, the pedicels about 1 cm. long; sepals broadly ovate, 
5-6 mm. long, deciduous; petals obovate-elliptic, narrowed at the 
base, rounded at the apex, puberulous, 1.5-2 cm. long and 8-10 mm. 
broad; stamens 1 mm. long. Neg. 14179. 

Loreto: Middle Rio Ucayali, Yarina-cocha, Tessmann 3423 (type). 
"Yana waska," "yuno" (Chama). 

15. Guatteria eugeniaefolia A. DC. in sched. (in herb. DC.). 
G. ovalis Mart. Fl. Bras. 13, pt. 1: 38. 1841 (non R. & P.). 

Young branchlets thinly tomentellous; leaves on petioles 3-6 
mm. long, hairy above when young, finally quite glabrous and 
lustrous, sparsely appressed-hairy beneath, elliptic-obovate, cuneate 
and decurrent at the base, abruptly contracted at the apex into a 
cusp 1 cm. long, 8-12 cm. long and 3-5 cm. broad, veins equally 
prominent on both sides, forming a dense network; flowers solitary; 
pedicels 12-15 mm. long, tomentose; sepals 4-5 mm. long, tri- 
angular-ovate, ferruginous- tomentose; petals 12-15 mm. long, 
rounded at the apex, ferruginous-tomentose on the outside, yellow- 
ish-gray-tomentellous inside, the outer oblong, about 5 mm. broad, 
the inner narrowly obovate, 7 mm. broad; connective disk um- 
bonate; monocarps (only unripe ones seen) numerous, obovoid, on 
pedicels 5-7 mm. long. Neg. 27539. 

Loreto: Tocache, Mainas, Poeppig 1859 (type in Herb. DC.). 

16. Guatteria phanerocampta Diels, Notizbl. Bot. Gart. 
Berlin 11: 76. 1931. 

Young branchlets sericeous, soon glabrescent; leaves niembra- 
naceous, 15-20 cm. long and 5-7.5 cm. broad, pale green when dried, 
glabrous above and very sparsely appressed-hairy beneath, lanceolate- 
elliptic or rarely obovate, rounded-acute and long-decurrent at the 
base, ending at the apex in an acumen 1-3 cm. long; flowers solitary, 
on sericeous pedicels 2-3 cm. long; corolla stellate-expanded, 3-4 cm. 



FLORA OF PERU 715 

across; petals gray-sericeous outside, the outer rhombic-elliptic and 
narrowed to the obtuse apex, 12-18 mm. long and 7-10 mm. broad, 
the inner oblong-obovate, rounded at the apex, slightly longer and 
broader than the outer; stamens 1.8-2 mm. long. 

Loreto: Upper -Rio Itaya, San Antonio, Williams 3436 (type). 
"Charahusca" (Quechua). 

17. Guatteria calliantha R. E. Fries, sp. nov. 

Arbor ca. 15 m. alta; folia 5-7 mm. longe petiolata, mem- 
branacea, supra laevia et adulta glabra, subtus primo ferrugineo- 
sericea, glabrescentia sed pilis nonnullis longis instructa, elliptico- 
obovata, obtuse cuspidata, ad basin contracta et acute decurrens, 
10-15 cm. longa et 5-6 cm. lata; nervi validiores supra impressi; 
flores in axillis inferioribus 2-4 (rarius 1), pedunculo ca. 1 cm. longo 
rubiginoso-crinito sustenti; sepala demum reflexa, rotundato-ovata, 
extus rubiginoso-hirsuta, ca. 6 mm. longa; petala extus ad basin 
rubiginoso-sericea, ceterum minutissime sericea et demum fere 
glabrescentia, e basi angusta obovato-spathulata; interiora apice 
retusa 18-22 mm. longa et 10-13 mm. lata, exteriora paulo minora. 

Loreto: Lower Rio Itaya, Soledad, Tessmann 5164 (type in Herb. 
Mus. Berlin). 

18. Guatteria coeloneura Diels, Bot. Jahrb. 37: 408. 1906. 

A tree about 10 meters high, the young branchlets ferruginous- 
pilose; leaves coriaceous, on petioles 3-5 mm. long, glabrescent 
above and sparsely hairy below even when mature, elliptic and 
shortly acuminate, shortly acute at the base, 8-14 cm. long and 3-6 
cm. broad, lateral nerves impressed on the upper side; pedicels 1-2, 
ferruginous- tomentose, 1.5-2 cm. long; sepals rounded-ovate, decid- 
uous, 8-9 mm. long; petals ferruginous-sericeous, especially at the 
base outside, broadly obovate, rounded or truncate at the apex, 
the inner 12-13 mm. long and 11-12 mm. broad, the outer a 
little shorter and narrower; monocarps ellipsoid, 8-9 mm. long 
and 5-6 mm. broad, their stipes thick, straight, 5-6 mm. long. 
Neg. 14182. 

Huanuco: In the mountains southwest of Monson, Weberbauer 

3548 (type). 

19. Guatteria pleiocarpa Diels, Bot. Jahrb. 37: 409. 1906. 

A shrub 5 meters high, the young branchlets clothed with soft, 
patent, rufous hairs; leaves coriaceous, on petioles 5-8 mm. long, 



716 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glabrous above when adult, loosely appressed-hairy beneath, oblong- 
elliptic, shortly acute at the base, acuminate, 14-16 cm. long and 
5-6 cm. broad; pedicel 1 (rarely 2), straight, rufous-pilose, 1.5-2.5 
cm. long; sepals recurved, 7-8 mm. long; petals glaucous when 
dried and minutely ferruginous-tomentellous, oblopg-ovate, the outer 
about 10 mm. long and 5 mm. broad, the inner 13-14 mm. long, 
7 mm. broad; monocarps numerous, ellipsoid or obovoid, apiculate, 
7-8 mm. long and 4 mm. broad, stipes very slender, 1.5-2 cm. 
long. Neg. 14206. 

Loreto: Moyobamba, Weberbauer 4530 (type). 

20. Guatteria glauca R. & P. Syst. Veg. Fl. Peruv. 1: 145. 1798. 

Young branchlets sparsely clothed with long, rather appressed 
hairs; leaves on petioles 4-7 mm. long, coriaceous, thinly appressed- 
hairy on both sides when young, soon glabrescent, elliptic to oblong- 
elliptic, rounded or sometimes shortly acute at the base, abruptly 
contracted at the apex into a short, obtuse cusp, 12-20 cm. long and 
5-7 cm. broad; flowers solitary; pedicels 1-2 cm. long, softly hairy; 
sepals rounded-ovate, about 7 mm. long, recurved; petals elliptic, 
ferruginous-sericeous at the base outside, tomentellous on the upper 
part, inner petals up to 17 mm. long and 10 mm. broad, the outer 
a little smaller; monocarps ellipsoid-pyriform, rounded at the apex, 
about 12 mm. long and 8 mm. broad, stipes about 1.5 cm. long. 
Neg. 14187. 

Huanuco: "In nemoribus versus Cochero et Chinchao," Ruiz 
(type). 

21. Guatteria juninensis R. E. Fries, sp. nov. 

Frutex vel arbor parva, ramulis novellis minute pilosis, cito 
glabrescentibus; foliorum petiolus 3-5 mm. longus; lamina coriacea, 
adulta supra glaberrima nitidaque, subtus pilis laxissimis brevibus 
adpressis obsita, oblonga vel oblongo-elliptica, basi rotundata, apicem 
versus sensim vel modice abrupte contracta obtuseque acuminata, 
10-20 cm. longa et 4-6.5 cm. lata; nervi laterales paulo impressi; 
flores solitarii; pedicelli rigidi, 1.5-2.5 cm. longi, breviter pilosi; 
sepala rotundato-ovata, obtusa, extus sericea, 3-4 mm. longa, mox 
reflexa; petala ovato-elliptica, apice late rotundata, extus basi 
sericea ceterumque tomentella, interiora 10-14 mm. longa et 6-10 
mm. lata, exteriora paulo minora; monocarpia pyriformia, breviter 
acuta, 8-9 mm. longa et 5 mm. lata, stipitibus 3-5 mm. longis (rarius 
longioribus) sustenta. 



FLORA OF PERU 717 

Junin: La Merced, Hacienda Schunke, above San Ramon, 
Macbride 5753, 5786; Schunke A91; Killip & Smith 24877. Chan- 
chamayo Valley, Schunke 267 (type in Herb. Field Mus.). East of 
Quirimi Bridge, near La Merced, Killip & Smith 23975. Colonia 
Perene", Killip & Smith 24949. 

22. Guatteria ucayalina Huber, Bol. Mus. Goeldi 4: 560. 1906. 
Young branchlets densely ferruginous-hirsute; leaves rigidly 

membranaceous, elliptic-obovate, abruptly acuminate at the apex, 
acute at the base, glabrous above except on the midrib and thicker 
nerves, sparsely hairy beneath, with patent hairs more dense on the 
nerves, 15-25 cm. long and 7-10 cm. broad; pedicels solitary, 
glabrescent, about 3.5 cm. long in fruiting stage, articulate a little 
above the base; monocarps numerous, pyriform, apiculate, 1-1.2 
cm. long and 7 mm. broad, glabrous, narrowed at the base into a 
very slender stipe 3-3.5 cm. long. Neg. 14218. 

Peru: Quebrada Grande del Cerro de Canchahuaya, Huber 1431 
(type). 

23. Guatteria Guentheri Diels, Notizbl. Bot. Gart. Berlin 10: 
169. 1927. 

A large tree, up to 25 meters high, the young branchlets sericeous; 
leaves on petioles 7-10 mm. long, large, rigid, dark brown when 
dried, glabrous and lustrous above except on the lower part of the 
sunken midrib, densely sericeous beneath at first, finally almost 
glabrescent, elliptic, shortly cuspidate at the apex, narrowed to the 
acute or sometimes rather obtuse base, 18-22 cm. long and 7-9.5 cm. 
broad; flowers 1-2, often produced from old branches; pedicels 
straight, rigid, appressed-hairy, 2-3 cm. long; sepals recurved, 
7-8 mm. long; petals obovate, puberulous and ferruginous-sericeous 
outside at the base, 3-3. 5 cm. long, the margins retrorse. Neg. 14189. 

Loreto: Upper Rio Maranon, at the mouth of Rio Santiago, 
Tessmann 4387 (type). Also in Amazonian Brazil. 

24. Guatteria spectabilis Diels, Notizbl. Bot. Gart. Berlin 9: 
138. 1924. 

A large tree, about 25 meters high, with a trunk half a meter 
thick, the young branchlets appressed-hairy, soon glabrescent; leaves 
coriaceous, 18-25 cm. long and 6-10 cm. broad, glabrous and lus- 
trous above, appressed-hairy beneath, elliptic, abruptly contracted 
to a nearly round base but rather long-decurrent on the petiole, 
which is 8-10 mm. long, cuspidate at the apex; flowers 1-2 in the 



718 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaf axils; pedicels 2.5-3.5 cm. long, sericeous; sepals finally pendu- 
lous, 0.8-1.2 cm. long; petals oblong-obovate, rounded at the apex, 
puberulous and ferruginous-tomentose outside at the base, attaining 
3.5 cm. in length when fully developed, the interior a little larger 
than the outer. Neg. 14215. 

Loreto: Middle Rio Ucayali, Yarina-cocha, in forest, Tessmann 
3235 (type). Upper Rio Maranon, at the mouth of Rio Santiago, 
Tessmann 1+638. 

25. Guatteria hyposericea Diels, Notizbl. Bot. Gart. Berlin 
11: 76. 1931. 

A medium-sized tree, the young branchlets densely sericeous; 
petioles 7-8 mm. long; lamina coriaceous, soon glabrous above, very 
densely covered beneath with appressed hairs, lanceolate-oblong, 
gradually narrowing at both ends, long-acuminate at the apex, 
decurrent at the base, 12-20 cm. long and 3-7 cm. broad; pedicels 
1-2, sericeous, 1.3-1.8 cm. long; sepals and petals densely ferrugi- 
nous-sericeous or tomentellous; petals broadly elliptic, 11-13 mm. 
long, 6-8 mm. broad; stamens 1.3 mm. long, the connective disk 
umbonate; monocarps ellipsoid-obovoid, shortly acute, glabrous, 
10 mm. long and 6 mm. thick, abruptly contracted into a slender 
stipe 10-15 mm. long. 

San Martin: Upper Rio Huallaga, Tarapoto, Williams 6323, 
6671. Lower Rio Huallaga, Williams 4129 (type). "Jana huasca." 
Also in Amazonian Brazil (State of Para). 

26. Guatteria megalophylla Diels, Verh. Bot. Ver. Brandenb. 
47: 127. 1905. 

A small tree, up to 15 meters high, the young branchlets sericeous; 
petiole 1.5-3 cm. long, its lower part (the real petiole) thickened and 
black, its upper half narrow-winged; lamina very large (30-50 cm. 
long and 7-15 cm. broad), rigid, glabrous and smooth above, very 
thinly appressed-hairy or glabrous beneath, oblong to oblanceolate, 
acute at the base, long-acuminate; pedicels 1-2, up to 8 mm. long, 
thick; sepals broadly ovate, golden-sericeous outside, 6-8 mm. long 
and broad; petals coriaceous, golden-sericeous outside, 12-14 mm. 
broad, the outer petals elongate-ovate, narrowed to the obtuse apex, 
18-25 mm. long, the inner ovate-elliptic, less narrowed to the rounded 
apex, 22-27 mm. long; monocarps oblong-ellipsoid, 18-22 mm. 
long, abruptly narrowed into a thick stipe 3-6 mm. long. Neg. 14199. 

Loreto: Upper Rio Maranon, mouth of Rio Santiago, Tessmann 
4325. Between Yurimaguas and Balsapuerto, on the lower Rio 






FLORA OF PERU 719 

Huallaga, Killip & Smith 28260. Santa Rosa, below Yurimaguas, 
Kittip & Smith 28780. Iquitos, Killip & Smith 27326. "Envira." 
Also in Amazonian Brazil ( Ule 5630, type). 

27. Guatteria melosma Diels, Notizbl. Bot. Gart. Berlin 
10: 170. 1927. 

A tree, up to 15 meters high; leaves very large, the petiole 
1.5-3 cm. long, thickened below the middle, narrow-winged above; 
lamina coriaceous, glabrous above, thinly appressed-hairy beneath, 
oblong, abruptly contracted at the base and decurrent, cuspidate 
(1-2 cm. long) at the apex, 30-40 cm. long and 6-12 cm. broad; 
flowers axillary, 1-2; pedicels 5-10 mm. long; sepals rigid, rounded- 
ovate, ferruginous-sericeous outside, about 1.2 cm. long and broad; 
petals obovate, broadly rounded at the apex, golden-sericeous outside 
near the base, silvery-sericeous toward the apex, 1.8-2.5 cm. long; 
monocarps oblong-ellipsoid, 2-2.2 cm. long and about 1 cm. broad, 
on stipes 4-5 mm. long. Neg. 14200. 

Loreto: Upper Rio Maranon, Iquitos, Tessmann 5039 (type), 
5352. Mishuyacu, near Iquitos, King 719, 1112. 

28. Guatteria Dielsiana R. E. Fries, nom. nov. G. ucayaliana 
Diels, Notizbl. Bot. Gart. Berlin 9: 138. 1924. 

A small tree with appressed-hairy branchlets; leaves medium- 
sized, rather thinly verruculose-punctulate, glabrous above from the 
beginning, densely sericeous beneath when young, later glabrescent 
and finally sparsely appressed-hairy or nearly glabrous, narrowly 
oblong, acuminate, gradually narrowed to the base and decurrent on 
the (2-4: mm. long) petiole, 10-18 cm. long and 3.5-5 cm. broad; 
pedicels 1-3, sericeous, short; sepals appressed, about 7 mm. long; 
petals silvery-sericeous outside, 18-20 mm. long, the outer nar- 
rowly lanceolate, acute, the inner ovate-lanceolate, obtuse; fruits 
unknown. 

Loreto: Rio Ucayali, Yarina-cocha, forest near the lake, Tess- 
mann 3212 (type). 

29. Guatteria macrocarpa R. E. Fries, sp. nov. 

Arbor vel frutex; ramuli novelli pilis patulis longiusculis flaves- 
centibus dense velutino-sericei ; foliorum petiolus ca. 5 mm. longus; 
lamina (exsiccata) flavovirens, utroque latere verruculoso-punctata, 
supra glaberrima, subtus primo dense sericea, adulta setulis sat 
decumbentibus sparse vestita, oblongo-elliptica, basi rotundata et 
ima basi tantum brevissime decurrens, apice abrupte 5-10 mm. longe 



720 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cuspidata, 14-20 mm. longa et 4.5-6.5 mm. lata; flores ignoti; 
monocarpia cylindrico-oblonga, apice rotundata et minute apiculata, 
basi obtusiuscula, demum glabra, ca. 24 mm. longa et 9 mm. lata, 
stipitibus ca. 8 mm. longis crassis sustenta. 

Loreto: Santa Rosa, lower Rio Huallaga, below Yurimaguas, in 
dense forest, Kittip & Smith 28965 (type in Herb. Field Mus.). 

30. Guatteria peruviana R. E. Fries, sp. nov. 

Arbor 6-7 m. alta, ramuli novellilonge strigosi; foliorum petiolus 
ca. 10 mm. longus; lamina papyracea, utroque latere creberrime 
verruculoso-punctata, adulta supra glaberrima, subtus pilis longius- 
culis adpressis plus minusve sparse obsita, oblongo-elliptica, basi acuta 
et decurrens, apice abrupte in cuspidem 1-2 cm. longam contracta, 
15-20 cm. longa et 5.5-7.5 cm. lata; pedicelli erecti, stricti, strigosi, 
ca. 8 mm. longi; sepala baud recurvata; petala erecto-patula, crassa, 
oblonga, apice rotundata, extus ferrugineo-sericea, 15 mm. longa (vel 
longiora?), 6-7 mm. lata; carpella 20-25; monocarpia cylindrico- 
elongata, in triente superiore latissima, 6-8 mm. longe stipitata. 

Loreto: Along Rio Maranon, near mouth of Rio Tigre, on wooded 
banks, Kittip & Smith 27522 (type in U. S. Nat. Herb.). 

31. Guatteria decurrens R. E. Fries, sp. nov. 

Arbor 6-8 m. alta, ramulis novellis pilis oblique patentibus 
vestitis; foliorum petiolus 3-5 mm. longus; lamina membranacea, 
flavovirens, creberrime verruculoso-punctata. supra glabra, subtus 
pilis pallidis laxis, in nervis densioribus, decumbentibus obsita, 
oblanceolata, 1-2 cm. longe cuspidata, basin versus longissime sensim- 
que angustata, ima basi obtusa, 16-23 cm. longa et 5-6 cm. lata; 
flores ignoti; fructus pedicellus 2.5-3 cm. longus, rigidus, strigosus; 
monocarpia pauca, elongato-ellipsoidea, obtusa, basi acuta, 17-19 
mm. longa, 7-8 mm. crassa, 10-13 mm. longe stipitata. 

Loreto: Soledad, on Rio Itaya, in dense forest, Killip & Smith 
29585 (type in U. S. Nat. Herb.). 

32. Guatteria amazonica R. E. Fries, sp. nov. 

Frutex vel arbor parva, ad 6 m. alta; ramuli novelli primo pilis 
rigidis adpressis dense sericei, mox glabrescentes; foliorum petiolus 
3-5 mm. longus; lamina papyracea, verrucis minutissimis densissime 
punctata, supra ab initio glabra, subtus primo dense flavescenti- 
sericea, glabrescens et rubescens, demum pilis disseminatis longis 
adpressis instructa, oblanceolata vel obovata, basin obtusam versus 



FLORA OF PERU 721 

rotundato-cuneata, apice in cuspidem ca. 2 mm. longam sat abrupte 
contracta, 25-35 cm. longa et 8-12 cm. lata; flores ignoti; fructus 
in trunco vel ramis crassis glomerati, pedicellis rigidis adpresse hir- 
sutis 15-22 mm. longis sustenti; monocarpia elongato-ellipsoidea, 
16-18 mm. longa et 7-8 mm. crassa, in stipites 8-9 mm. longos 
abrupte contracta; pericarpium siccum tenue. 

Loreto: Chimbote, on Rio Amazonas, Kuhlmann (Herb. Rio de 
Janeiro 24260; type in Herb. Stockholm). Also in western Amazo- 
nian Brazil (Sao Paulo de Olivenca, Krukoff 8500) . 

33. Guatteria multivenia Diels, Notizbl. Bot. Gart. Berlin 
10: 171. 1927. 

A tree about 15 meters high, the young branchlets densely covered 
with short, patent, soft hairs; petioles 3-6 mm. long; leaves thin, 
very densely verruculose-punctulate, glabrous above when adult, at 
first densely golden-sericeous beneath, later thinly clothed with 
patent, long hairs, obovate, gradually narrowed to the obtuse base, 
rounded at the apex and abruptly cuspidate, 20-35 cm. long and 6-12 
cm. broad; flowers produced from thick branches; pedicels thick 
rigid, 1-1.5 cm. long; sepals recurved, 7-9 mm. long; petals seri- 
ceous on the outside, oblong, the outer 18-20 mm. long and 7-8 mm. 
broad, the inner up to 25 mm. long, emarginate at the apex; fruits 
not known. Neg. 14203. 

Loreto: Lower Rio Itaya, Soledad, Tessmann 5192 (type). 

34. Guatteria excellens R. E. Fries, sp. nov. 

Arbor 3 m. alta, ramulis novellis crassis fusco-velutinis; foliorum 
petiolus 2-4 mm. longus; lamina utroque latere creberrime verrucu- 
loso-punctata, adulta supra glabra, subtus (praesertim in nervis) 
pilis decumbentibus obsita, oblanceolata, basi inaequilatera et 
rotundata, apice in cuspidem 2-2.5 cm. longam acutissimam abrupte 
contracta, 35-50 cm. longa, 10-15 cm. lata; nervi laterales 20-30 
utrinque; flores solitarii, pedicellis crassis rufo-velutinis ca. 1 cm. 
longis sustenti; sepala 10 mm. longa; petala ovato-elliptica, 
omnia ca. 15 mm. longa et extus argenteo-sericea; monocarpia 
cylindrico-ellipsoidea, apice rotundata et minutissime apiculata, 
ca. 20 mm. longa et 10-11 mm. crassa, 5 mm. longe stipitata, peri- 
carpio carnoso. 

Loreto: Mishuyacu, near Iquitos, in forest, Klug 1273 (type in 
Herb. Field Mus.). 

35. Guatteria eriopoda DC. (ex Dunal, Monogr. Anon. 130. 
1817) is an insufficiently known species, described from material 



722 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

collected by Dombey near Cochero in Peru. It is probably related 
to G. hirsuta and G. excelsa. Perhaps Poeppig 1786 (from Mainas) 
belongs to this species, but the material is incomplete, since developed 
flowers and fruits are missing. Neg. 34471. 

3. PSEUDOXANDRA R. E. Fries 

Shrubs or trees with medium-sized, oblong or lanceolate leaves. 
Flowers axillary, solitary or in few-flowered inflorescences; pedicels 
very short, articulate, with distichous bracts below the articulation 
but without any bract above this. Flower buds globose. Sepals 
imbricate, small. Petals concave, with thin margins, imbricate, 
ovate-rotund, all similar but the inner thicker. Stamens numer- 
ous, the connective expanded above the pollen sacs into a disk. 
Carpels several; ovule 1, lateral and usually attached to the top of 
the ovary, pendulous. Monocarps globose, shortly stipitate, one- 
seeded. A small genus, containing six species in the Amazonian 
region and British Guiana, only two recorded from Peru. 

Leaves lustrous above, rounded and only shortly decurrent at the 
base; lateral nerves conspicuously thicker than the veins; flowers 
larger; petals 11-12 mm. long, the outer densely sericeous 
outside 1. P. Williamsii. 

Leaves dull, conspicuously acute at the base; lateral nerves slender, 
of about the same thickness as the veins; flowers small; petals 
6-8 mm. long, nearly glabrous 2. P. polyphleba. 

1. Pseudoxandra Williamsii R. E. Fries, Acta Hort. Berg. 12: 
227. /. 2b-c. 1937. Cremastosperma Williamsii R. E. Fries, op. cit. 
206. 1934. 

A shrub, the youngest branchlets sericeous; leaves membrana- 
ceous, oblong, rather abruptly narrowed into a short cusp (1-1.5 
cm. long), rounded-acute at the base and shortly decurrent to a 
petiole 4-5 mm. long, lustrous and clothed when young with long, 
appressed hairs on both surfaces, finally almost glabrous, 12-20 
cm. long and 3-5 cm. broad; network of veins dense, nicely prominent 
on both sides; flowers sericeous, their pedicels about 5 mm. long; 
sepals and outer petals gray-sericeous outside, the inner petals seri- 
ceous only along the midrib; all petals of the same size, broadly 
ovate, 11-12 mm. long. 

Loreto: Yurimaguas, Williams 3960 (type). 

2. Pseudoxandra polyphleba (Diels) R. E. Fries, Acta Hort. 
Berg. 12: 230. 1937. Unonopsis polyphleba Diels, Verh. Bot. Ver. 



FLORA OF PERU 723 

Brandenb. 47: 131. 1905. Cremastosperma polyphlebum R. E. Fries, 
Acta Hort. Berg. 10: 331. 1931. 

A shrub or tree 20 meters high; leaves membranaceous, opaque, 
elliptic or elliptic-oblong, very abruptly cuspidate, acute and decur- 
rent at the base, glabrous on both sides or with a few hairs beneath, 
12-18 cm. long and 4-6 cm. broad; sepals 1-2 mm. long; petals 
nearly glabrous, about 8 mm. long, their margins very thin, mem- 
branaceous and crisp; stamens 2 mm. long. 

Loreto: Iquitos, Mexia 6416. Also in Amazonian Brazil. 

4. CREMASTOSPERMA R. E. Fries 

Shrubs or trees with 1, rarely 2, axillary flowers, or sometimes 
cauliflorous. Flowers medium-sized, generally glabrous; pedicels 
articulate above the base, nearly always with one small bract above 
the articulation. Sepals free, imbricate in bud, with thin, ciliolate 
margins. Petals much longer than the calyx, rigid, imbricate, with 
thin edges. Stamens numerous, the connective expanded above the 
anther into a disk. Carpels several; ovule one, attached close to the 
upper end of the ovary, pendulous, sometimes located lower down. 
Monocarps free, stipitate, one-seeded. Twelve species are known 
hitherto, inhabiting the most tropical parts of South America, the 
eastern slopes of the Peruvian and Bolivian Andes, Amazonian 
Brazil, and French Guiana. 

Flowers small to medium-sized; petals 10-15 mm. long. 

Pedicels glabrous, produced from the axils of persistent or some- 
times fallen leaves. 

Petals oblong; pedicels very slender, up to 3 cm. long. 

1. C. gracilipes. 
Petals round or elliptic; pedicels thicker. 

Pedicels 5 cm. or longer. 

Leaves lanceolate, gradually long-attenuate at the apex, 
broadest at the middle 2. C. pendulum. 

Leaves obovate-elliptic, shortly narrowed at the apex and 
usually broadest above the middle. 

3. C. pedunculatum. 
Pedicels short, less than 3 cm. 

Leaves narrowing at the apex into a filiform cusp, this 
2-3 cm. long 4. C. longicuspe. 

Leaves with a short and broad cusp. .5. C. megalophyllum. 



724 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels covered with a short, dense pubescence; flowers produced 

from old, thick branches 6. C. cauliflorum. 

Flowers larger; petals about 25 mm. long 7. C. peruvianum. 

1. Cremastosperma gracilipes R. E. Fries, Acta Hort. Berg, 
pt. 26, 10: 325. 1931. 

A shrub or small forest tree with slender, glabrous branchlets, 
provided on the youngest parts with a more or less conspicuous, 
narrow ridge; leaves 10-15 cm. long and 3-6 cm. broad, membra- 
naceous, glabrous, oblong or oblanceolate, rounded and obtuse or 
very shortly acute at the base, abruptly contracted at the apex into 
a blunt cusp 1-2 cm. long; flowers small; pedicels very thin and 
glabrous, about 3 cm. long; petals blackening when dried, glabrous, 
all of about the same size, oblong-elliptic and 10-13 mm. long when 
fully developed, their margins thin and crisp. 

Loreto: Upper Rio Maranon, Puerto Mele"ndez, Tessmann 4748 
(type). Puerto Arturo, near Yurimaguas, Williams 5085, 5287, 5296. 
Yurimaguas, Santa Rosa, Williams 4919; Killip & Smith 28961. 

2. Cremastosperma pendulum (R. & P.) R. E. Fries, Acta 
Hort. Berg. 10: 325. 1931. Guatteria pendula R. & P. Syst. Veg. 
Fl. Peruv. 1: 146. 1798. 

Leaves on petioles 6-7 mm. long, rigid, lustrous, minutely seri- 
ceous when young, soon quite glabrous, lanceolate, broadest at the 
middle and narrowing equally at the acute base and at the acuminate, 
obtuse apex, 14-25 cm. long and 3-7 cm. broad; midrib prominent 
on both sides, the lateral nerves divergent at a very acute angle; net- 
work of veins dense and nicely prominent on both sides; flowers 
solitary; pedicels up to 10 cm. long, slender, black; sepals and 
petals glabrous but ciliate at the margins, blackening when dried; 
petals all of equal length (10-12 mm. long), the outer orbicular, the 
inner a little narrower; monocarps ellipsoid, black, on stipes 13-14 
mm. long. Negs. 29211, 27527. 

Huanuco: Vitoc, Ruiz & Pavon. 

3. Cremastosperma pedunculatum (Diels) R. E. Fries, Acta 
Hort. Berg. 10: 48. /. 6a-c. 1930. Aberemoa pedunculata Diels, Bot. 
Jahrb. 37: 409. 1906. Duguetia pedunculata Macbr. Field Mus. Bot. 
4: 172. 1929. Guatteria socialis Macbr. op. cit. 171. 

A tree, 12 meters high, the young branchlets appressed-ferrugi- 
nous-hirsute; leaves 15-20 cm. long and 5-8 cm. broad, rigid, finally 
quite glabrous and lustrous on both sides, oblanceolate-elliptic, 



FLORA OF PERU 725 

tapering into a blunt cusp, long-cuneate at the base and shortly 
decurrent on the (7-10 mm. long) petiole; flowers 1 or rarely 2, 
axillary on young branches; pedicels very long, up to 10 cm., straight, 
black and rugose; sepals rounded, 2-3 mm. long, ciliate; petals 
ovate-elliptic, all of the same length (about 15 mm.), the inner a 
little narrower than the outer; monocarps pyriform, their stipes 
thick, straight, rigid, about 16 mm. long. 

Loreto: Near Moyobamba, Weberbauer 4558 (type) ; Klug 3726. 
Junin: Chanchamayo Valley, Schunke 395. 

Schunke states that his plant is a liana, but as it agrees with the 
typical C. pedunculatum in the leaves, flowers, and other characters, 
his specimen (the type of Guatteria socialis Macbr.) may be regarded 
as C. pedunculatum. 

4. Cremastosperma longicuspe R. E. Fries, Acta Hort. Berg. 
12: 203. 1934. 

A small shrub, usually less than one meter high; young branchlets 
glabrous; leaves membranaceous, glabrous on both sides or some- 
times with a few hairs on the midrib, obovate, abruptly contracted 
into a (2-3 cm. long) very narrow and filiform cusp, cuneate and 
acute at the base, 14-23 cm. long and 6-9 cm. broad; network of the 
veins large-meshed; flowers 1-2, axillary on young branches; pedicels 
up to 2 cm. long, glabrous, obviously thickened toward the top; petals 
not blackening when dried, thick, glabrous, all rather alike, ovate to 
orbicular, 12-18 mm. long; monocarps clavate-ellipsoid, their stipes 
1 cm. long or slightly more; pericarp glabrous, fragile, the single seed 
attached to the upper end of the ovary, pendulous. 

Loreto: Mainas, Poeppig 2091 (type). Yurimaguas, Williams 
4092; Kuhlmann (Herb. Rio de Janeiro 24265). 

5. Cremastosperma megalophyllum R. E. Fries, Acta Hort. 
Berg. 10: 329. 1931. 

A small tree with thick, glabrous branchlets; leaves unusually 
long-petiolate (up to 15 mm. long), very large, 20 cm. long or more 
(sometimes attaining 45 cm.), 10-17 cm. broad, elliptic to obovate, 
more or less acute at the base and abruptly narrowed into a broad, 
obtuse cusp 1-1.5 cm. long, quite glabrous and lustrous on both 
sides; flowers solitary in the axils of fallen leaves, sometimes on 
rather thick branches; pedicels thin, glabrous, 2 cm. long; perianth 
glabrous, white to light yellow, but blackening when dried; sepals 



726 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rounded, 5-6 mm. long, ciliolate; petals all alike, orbicular, with thin, 
ciliolate margins, 13-15. mm. long and broad. 

Loreto: Boca de Pebas, Ducke (Herb. Rio de Janeiro 19620, type). 
Balsapuerto, King 3069. 

6. Cremastosperma cauliflorum R. E. Fries, Acta Hort. 
Berg. 10: 330. 1931. 

A small tree, 9 meters high, the youngest branchlets tomentellous; 
leaves large, up to 30 cm. long and 12 cm. broad, blackening when 
dried, glabrous and lustrous on both sides, with a few appressed, 
small hairs on the midrib and larger veins beneath, where the smaller 
veins form a very dense and nicely prominent network; base of the 
leaves acute and shortly decurrent; flowers clustered on thick, 
old branches; pedicels about 2.5 cm. long, covered with dense, short 
pubescence; petals all of the same length (about 15 mm. long), the 
outer ovate, the inner a little narrower, oblong-ovate, all covered on 
the outside and on the upper part of the inside with a yellowish gray, 
soft pubescence. 

Loreto: Mishuyacu, near Iquitos, Klug 902 (type). 

7. Cremastosperma peruvianum R. E. Fries, Acta Hort. 
Berg. 12: 204. 1934. 

A small tree with thick, glabrous branchlets; leaves very large 
(50-55 cm. long and 11-16 cm. broad), elliptic or oblong, abruptly 
cuspidate at the apex, broadly rounded at the base, glabrous and 
lustrous on both sides; flowers solitary in the axils of fallen leaves; 
pedicels about 17 mm. long, after an thesis elongated to 30 mm., 
glabrous, with a few bracts 1 mm. long near the base, articulate but 
without any bract above the articulation ; sepals orbicular, glabrous 
but ciliolate, 3-4 mm. long; petals pale yellow, green at the base, 
not blackening when dried, glabrous, elliptic to narrowly spatulate, 
of almost equal length (22-27 mm.). 

Loreto: Upper Rio Maranon, mouth of Rio Santiago, Tessmann 
4*7* (type). 

5. RUIZODENDRON R. E. Fries 

A tree with medium-sized leaves; network of the veins dense, 
prominent. Flowers axillary, bisexual. Sepals free, imbricate in 
bud. Petals free, all rather alike, membranaceous, linear-lanceolate, 
imbricate. Stamens numerous, the connective expanded above the 
pollen sacs into a truncate disk. Carpels numerous; ovary glabrous, 
oblong-ovoid, with a sessile, hairy stigma; ovule solitary, lateral, 



FLORA OF PERU 727 

attached below the middle of the cavity, obliquely ascending. 
Monocarps free, stipitate, fleshy, depressed and transversely ovoid. 
Seed solitary, horizontal. A monotypic genus. 

1. Ruizodendron ovale (R. & P.) R. E. Fries, Arkiv Bot. 28B, 
No. 4: 3. 1936. Guatteria ovalis R. & P. Syst. Veg. Fl. Peruv. 1: 
146. 1798. 

A tree about 5 meters high, the young branchlets appressed- 
hirsute; petioles 5-8 mm. long; lamina elliptic, ovate, or lanceolate- 
ovate, rounded at the base but very shortly decurrent on the petiole, 
obtuse at the apex, 10-20 cm. long and 5-9 cm. broad, glabrous 
above, glaucous and nearly glabrous beneath; flowers solitary on 
pedicels 4-6 mm. long; sepals ovate-lanceolate, sericeous on the 
outside, 5 mm. long; petals white but blackening when dried, linear- 
lanceolate and long-attenuate to the acute apex, finally glabrous, 
4-4.5 cm. long and 3-7 mm. broad; monocarps about 10 mm. high 
and 12-15 mm. broad, glabrous. Neg. 29210. 

Huanuco: Posuso, Ruiz & Pavdn (type). San Martin: Juanjui, 
upper Rio Huallaga, Klug 3798. Also in Bolivia. 

6. MALMEA R. E. Fries 

Trees or shrubs. Flowers medium-sized or large; inflorescences 
few- to several-flowered, opposite the leaves; pedicels articulate 
immediately above a nearly basal bract and bearing another bract 
above the articulation. Sepals small, with thin margins, imbricate 
in bud. Petals much longer than the sepals, blackening when dried, 
orbicular-elliptic, imbricate in bud, their margins thin and more or 
less crispate. Stamens very numerous, the connective expanded 
above the anther into a truncate disk. Carpels numerous, containing 
one basal, erect ovule. Monocarps free, stipitate, one-seeded. 
Of this genus eleven species are known, six inhabiting the Amazonian 
region and Guiana, one native of Bahia, one of Ecuador, and three of 
Central America (Mexico to Panama). 

Leaves acute at the base. 

Leaves not cuspidate; pedicels more than 1 cm. long. 

Leaves smaller (up to 10-11 cm. long); outer petals shorter than 

the inner 1. M. Dielsiana. 

Leaves larger; all petals of equal length 2. M. Raimondii. 

Leaves cuspidate; pedicels 0.5 cm. long or shorter. 

4. M. cuspidata. 
Leaves rounded at the base. . . .3. M. xanthochlora. 



728 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. Malmea Dielsiana (Saff.) R. E. Fries, Acta Hort. Berg. 
10: 42. /. 56. 1930, and 10: 319. 1931. Cymbopetalum Dielsianum 
Saff. ex Diels, Notizbl. Bot. Gart. Berlin 9: 1156. 1927. 

A tree or shrub, the youngest branchlets ferruginous-hirsute; 
leaves small (up to 12 cm.), elliptic or lanceolate, cuneate at the 
base, gradually narrowed to the blunt apex, at first appressed-hairy 
beneath, soon glabrate except on the midrib; inflorescences on 
peduncles 0.5 cm. long, 1 (-2) -flowered; pedicels 2-2.5 cm. long; 
petals orbicular or elliptic, all glabrous, or the outer bearing a few 
small, appressed hairs, the inner petals about 18 mm. long, the outer 
9-12 mm.; monocarps oblong-ellipsoid, about 15 mm. long, shortly 
and densely hairy, their stipes up to 2.5 cm. long. 

San Martin: Tarapoto, Ule 10 (type); Williams 6710. 

2. Malmea Raimondii (Diels) R. E. Fries, Acta Hort. Berg. 
10: 320. 1931. Guatteria Raimondii Diels, Notizbl. Bot. Gart. Berlin 
11:75.1931. 

Leaves large, coriaceous, glabrous on both sides and lustrous, 
elliptic or oblong-elliptic, broadest at the middle and narrowed equally 
to the acute base and the obtuse apex, 10-22 cm. long and 4-7 cm. 
broad; midrib prominent on both sides; inflorescences very shortly 
pedunculate (1-) 2-flowered; pedicels thick, usually curved, up to 3.5 
cm. long; sepals and petals densely and shortly pubescent; petals 
all alike, nearly orbicular, undulate at their margins, 2-2.5 cm. long; 
connective disks conspicuously hairy. Neg. 14209. 

San Martin: Tarapoto, Spruce; Raimondi 238, 1974 (types); 
Williams 6226. "Sabina." 

3. Malmea xanthochlora (Diels) R. E. Fries, Acta Hort. 
Berg. 10: 45. /. 5a. 1930. Guatteria xanthochlora Diels, Notizbl. Bot. 
Gart. Berlin 10: 169. 1927. 

A small tree, about 6 meters high; leaves rigid, appressed- 
hirsute when young, soon quite glabrous and lustrous on both sides, 
lanceolate-oblong, rounded and obtuse at the base, gradually 
narrowed to the obtuse apex, up to 22 cm. in length and 7 cm. 
in breadth; inflorescences very shortly pedunculate, few- to 4- 
flowered, but developing only one flower at a time; pedicels 1-1.5 
cm. long; sepals ferruginous-pilose on the outside; petals all alike, 
elliptic, rounded at the apex, undulate on the margins, 2.5-3 cm. 
long, glabrous. 

Loreto: Upper Rio Maranon, at the mouth of Rio Santiago, 
Tessmann 4426, 4553 (type). 



FLORA OF PERU 729 

4. Malmea cuspidata Diels, Notizbl. Bot. Gart. Berlin 11: 
78. 1931. 

Leaves sericeous-pilose when young, soon quite glabrous, coria- 
ceous and lustrous above, obovate-elliptic, contracted at the apex 
into a cusp 1-1.5 cm. long, acute at the base, 10-12 cm. long and 
3.5-5 cm. broad; flowers solitary, on short (about 5 mm. long), 
pilose pedicels; sepals small, densely pilose; outer petals pilose 
outside and ciliate on the margins, up to 16 mm. long, the inner 
glabrous and of about the same size. 

Loreto: Rio Itaya, Williams 113 (type). 

7. FUSAEA Saff. 

Shrubs or trees with large, oblong leaves. Flowers large, solitary 
or in few-flowered inflorescences, terminal on very reduced, axillary 
or supra-axillary branchlets. Perianth segments rigid; sepals large, 
valvate, finally detached at the base; petals free or connate at the 
base, much longer than the sepals, oblong-lanceolate to rhombic- 
obovate, imbricate in bud, densely covered with long, appressed 
hairs. Stamens numerous, on a low, convex torus; connective 
terminating in a truncate disk; outer stamens sterile, longer than 
the fertile. Carpels very numerous, densely packed in a central 
cavity on the torus; ovule one, basal, erect. Fruiting carpels fused 
into a spherical, solid mass. Only three species known, all recorded 
from Peru, two of them, so far as known, endemic in that country. 

Petals free (Sect. Eu-Fusaea R. E. Fries). 

Leaves rounded or shortly acute at the base; nerves and veins 
impressed on the upper surface; staminodes 2-2.5 mm. broad, 

rounded at the apex 1. F. longifolia. 

Leaves long-acute at the base and long-decurrent; veins not 

impressed on the upper surface; staminodes about 1.5 mm. 

broad, gradually narrowed to the obtuse apex . 2. F. decurrens. 

Petals connate at the base, forming a short perianth tube (Sect. 

Synpetalantha R. E. Fries) 3. F. peruviana. 

1. Fusaea longifolia (Aubl.) Saff. Contr. U. S. Nat. Herb. 18: 
65. /. 73-74. 1914. Annona longifolia Aubl. Hist. PI. Guian. 1: 615. 
pi. 248. 1775. Duguetia longifolia Baill. Adansonia 8: 327. 1868. 
Aberemoa longifolia Baill. Hist. PI. 1: 205. /. 233-235. 1868. Annona 
rhombipetala R. & P. Fl. Peruv. 5: pi. 489 (ined.) ex G. Don, Gen. 
Syst. 1: 87. 1831. F. rhombipetala Macbr. Field Mus. Bot. 4: 172. 
1929. Uvaria spectabilis DC. ex Dun. Monogr. Anon. 92. 1817. 



730 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A shrub or small tree, the young branchlets ferruginous-hirsute; 
leaves rigid, oblong or rarely oblanceolate, long-cuspidate, shortly 
acute or more or less rounded at the base, quite glabrous above and 
finally nearly glabrous or sparingly hairy beneath, very densely 
verrucose-dotted, 12-30 cm. long and 4-10 cm. broad; nerves and 
veins impressed on the upper surface; sepals broadly ovate, 1-1.6 cm. 
long, connate at the base; petals free, the outer 3-4 cm. long, the 
inner 4-4.5 cm.; staminodes oblong-obovate, puberulous, 4-6 mm. 
long and 2-2.5 mm. broad; fruit fleshy and edible, of the size of a 
small apple (about 6 cm. in diameter), glabrous and smooth or 
areolate. 

Peru: Rivero. Huanuco: Pueblo Nuevo and Chicoplaya, Ruiz 
(type of Annona rhombipetala) . Loreto: Mishuyacu, near Iquitos, 
King 865. 

2. Fusaea decurrens R. E. Fries, Acta Hort. Berg. 12: 207. 1934. 
A small tree, about 7 meters high, with slender, rufous-strigose 

branchlets; leaves finally glabrous and densely verruculose, very 
lustrous above, oblanceolate or rarely oblong-lanceolate, cuspidate, 
narrowed to the base and long-decurrent, forming narrow wings on 
the upper part of the petiole, 18-22 cm. long and 4-6.5 cm. broad; 
midrib and principal nerves sunken on the upper surface, the smaller 
veins not impressed and hardly visible; petals densely silvery- or 
ferruginous-sericeous on both sides, rather abruptly contracted into 
an obtuse cusp, the outer about 2.5 cm. long and 1 cm. broad, the 
inner about 3 cm. long, 1.5 cm. broad; staminodes linear-oblong, 
gradually narrowing from the middle to the obtuse apex, 5 mm. long 
and 1.2-1.7 mm. broad; fruit not known. 

Loreto: Mishuyacu, near Iquitos, Klug 952, type. 

3. Fusaea peruviana R. E. Fries, Acta Hort. Berg. 12: 273. 
1937. 

A small tree; youngest branchlets with patent, ferruginous hairs; 
leaves 1&-30 cm. long, 4-8 cm. broad, membranaceous, opaque, 
glabrous above, finally so beneath, usually conspicuously verruculose, 
oblanceolate-oblong, long-narrowed to the obtuse base, gradually 
or rather abruptly contracted at the apex into a narrow cusp about 
3 cm. long; inflorescences 1-2-flowered, sessile, supra-axillary from 
about the middle of the internode; petals densely golden-sericeous 
outside, especially near the base, connate at the base into a cylindric 
tube, 1 cm. long and 1.5 cm. wide; corolla lobes oblong-obovate, 
the outer obtuse and about 2.5 cm. long, 1-1.2 cm. broad, the inner 



FLORA OF PERU 731 

broadly rounded at the apex, 3-3.5 cm. long and 1.7-2 cm. broad; 
fruits unknown. 

Loreto: Yurimaguas, Kuhlmann (Herb. Rio de Janeiro 24.362; 
type). 

8. DUGUETIA St. Hil. 

Trees or shrubs with stellate hairs or stellate scales. Inflores- 
cences few- or many-flowered, more or less strictly opposite the leaves 
or sometimes produced from older branches or from the trunk. 
Flowers medium-sized or large, bisexual. Sepals valvate. Petals 
free, imbricate in bud, generally coriaceous. Stamens numerous; 
connective generally (in all Peruvian species) expanded above the 
anther into a truncate disk. Carpels numerous; ovule solitary, basal, 
erect. Monocarps sessile, closely crowded, ligneous or fleshy, easily 
separable or more or less coherent. A large genus of about 65 
species, inhabiting a vast area from Panama and the southernmost 
islands of the West Indies south to Paraguay. 

Petals densely covered on the outside with stellate scales. 
Sepals quite or nearly free. 
All petals of equal breadth, up to 2.5 cm. long. 

Leaves completely covered beneath with very dense scales. 

1. D. odorata. 
Adult leaves with scattered scales beneath. 

Leaves lanceolate-elliptic, at most 3-4 times longer than 

broad. 
Leaves thin, smaller and narrower (up to about 5 cm. 

broad) ; petals oblong 2. D. quitarensis. 

Leaves rigid, larger and broader (5-9 cm. broad); petals 

obovate-spatulate 3. D. Tessmannii. 

Leaves oblong, 5 times longer than broad. 4. D. peruviana. 
Inner petals obviously narrower than the outer, all about 4.5 cm. 

long 5. D. macrophylla. 

Sepals large, conspicuously connate at the base . . 6. D. hadrantha. 
Petals softly tomentellous with short, stellate hairs. 

Young branchlets covered with appressed, stellate scales; mono- 
carps about 2 cm. long, with conic and acute apex. 

7. D. Spixiana. 

Young branchlets with small, rigid, stellate hairs; monocarps 
truncate and plane at the apex, about 1 cm. long. 

8. D. latifolia. 



732 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. Duguetia odorata (Diels) Macbr. Field Mus. Bot. 4: 172. 
1929. Aberemoa odorata Diels, Notizbl. Bot. Gart. Berlin 10: 171. 
1927. 

A shrub or small tree, up to 12 meters high; leaves narrowly 
elliptic to oblanceolate, long and gradually attenuate at both ends, 
15-30 cm. long and 4-9 cm. broad, quite glabrous above, even when 
young, completely covered beneath with persistent, silvery scales; 
flowers produced from older branches; sepals with gold-colored scales 
outside; petals spatulate-elliptic, rounded at the apex, silvery-scaly 
outside, stellate-hairy inside, 17-20 mm. long. Neg. 14229. 

Loreto: Miraflores, at the mouth of Rio Tigre, Tessmann 3745, 
type. Mishuyacu, near Iquitos, Klug 1587. Also in western 
Amazonian Brazil. 

2. Duguetia quitarensis Benth. in Hook. Lond. Journ. Bot. 2: 
361. 1843. Aberemoa quitarensis R. E. Fries, Sv. Vet. Akad. Handl. 
34, pt. 5: 20. 1900. Duguetia ibonensis Rusby, Mem. N. Y. Bot. 
Card. 7: 246. 1927. 

A small tree, the young branchlets wholly covered with round, 
yellow scales; leaves on petioles 3-7 mm. long, lanceolate to elliptic, 
acute at both ends, 10-20 cm. long and 3-5.5 cm. broad, glabrous 
above, even when young, at first densely scaly beneath but soon 
nearly glabrate and provided only with scattered, stellate scales; 
inflorescences 1-2-flowered, the pedicels 3-7 mm. long; sepals elliptic- 
oblong, with dense golden scales outside, 12-13 mm. long; petals all 
alike, ligulate-oblong, rounded at the apex, 15-20 mm. long and 
6-7 mm. broad, with very dense silvery to yellow scales outside; fruit 
3-4 cm. in diameter; monocarps glabrous, shortly acute. Neg. 14231. 

Loreto: Lower Rio Huallaga, Williams 4907. Also in Bolivia, 
Amazonian Brazil, and British Guiana. 

3. Duguetia Tessmannii R. E. Fries, Acta Hort. Berg. 12: 52. 
1934. 

A tree 10-13 meters high, the young branchlets with dense golden 
scales; petioles 7-13 mm. long; lamina rigid, glabrous and lustrous 
above, thinly scaly beneath, elliptic to elliptic-oblong, acute at both 
ends, 15-25 cm. long and 5-9 cm. broad; pedicels 7-15 mm. long; 
sepals ovate or rounded-ovate, 11-15 mm. long and 9-11 mm. 
broad; petals obovate-spatulate, rounded at the apex, up to 24 mm. 
long and 8-12 mm. broad, the inner a little broader than the outer, 
all with densely golden scales outside. Neg. 28602. 



FLORA OF PERU 733 

Loreto: Ucayali, Yarina-cocha, Tessmann 3201. Upper Maranon, 
at the mouth of Rio Santiago, Tessmann 4373, type. Lower Rio 
Huallaga, below Yurimaguas, Killip & Smith 28877. Also in western 
Amazonian Brazil (Rio Purus). 

Closely related to Duguetia quitarensis and possibly only a variety 
of that species. 

4. Duguetia peruviana (R. E. Fries) Macbr. Field Mus. Bot. 
4: 172. 1929. Aberemoa peruviana R. E. Fries, Sv. Vet. Akad. Handl. 
34, pt. 5: 22. pi. 3, f. 1-3. 1900. Annona conifera Ruiz in sched. 

Young branchlets densely stellate-scaly, yellow; leaves on petioles 
2-4 mm. long, oblong or oblong-lanceolate, with an acumination 
1-1.5 cm. long, shortly rounded-acute at the base, glabrous and 
lustrous above, olivaceous beneath and provided with numerous 
yellow, stellate scales, not covering the whole surface; inflorescences 
opposite the leaves, sessile, 1-3-flowered ; pedicels about 7 mm. long; 
sepals oblong-ovate, 12-13 mm. long, with golden scales outside; 
petals rounded at the apex, about 15 mm. long, the outer ovate, 
8 mm. broad, the inner obovate, 10 mm. broad. Neg. 14230. 

Peru: Without locality, Ruiz, type. 

5. Duguetia macrophylla R. E. Fries, Acta Hort. Berg. 12: 
63. 1934. 

A tree 7-9 meters high; leaves rigid, large (35-45 cm. long and 
10-13 cm. broad), oblanceolate, abruptly contracted into an acute 
cusp, long and gradually narrowed to the rounded base, glabrous 
above and finally very thinly stellate-scaly or nearly glabrous 
beneath; pedicels about 1 cm. long, thick; sepals broadly ovate, 
acuminate, with dense rufous scales outside, 15-18 mm. long; petals 
white, blackening when dried, grayish-stellate-scaly outside, about 
4.5 cm. long, obtuse, the outer oblong and 1.5 cm. broad, the inner 
linear-oblong, 0.9 cm. broad; stamens 3 mm. long; connective disk 
puberulous; monocarps 2-2.5 cm. long, glabrous, in the center of 
the truncate top provided with a cusp 2 mm. long. 

Loreto: Iquitos, Killip & Smith 27291, type. Alto Rio Itaya, 
Williams 3430. 

6. Duguetia hadrantha (Diels) R. E. Fries, Acta Hort. Berg. 
12: 65. 1934. Aberemoa hadrantha Diels, Verh. Bot. Ver. Brandenb. 
47: 128. 1905. 

A tree 6-12 meters high, the young branchlets densely scaly; 
leaves on petioles 3-5 mm. long, rigid, obovate-oblanceolate or more 



734 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rarely oblong, abruptly cuspidate, long-narrowed to the acute base, 
8-20 cm. long and 4-7 cm. broad, glabrous and lustrous above, 
very thinly scaly beneath; pedicels 5-6 mm. long, with dense golden 
scales; sepals ovate, obtuse, striate, scaly outside, 15-20 mm. long, 
connate below for 3-5 mm.; petals oblong-ovate, obtuse, densely 
yellowish-gray-scaly outside; stamens about 2 mm. long, the disk 
of the connective setulose; fruit unknown. Neg. 14225. 

Junin: Between Pampa Hermosa and Pangoa, Raimondi 2985. 
Also in western Amazonian Brazil ( Ule 5794, type) . 

7. Duguetia Spixiana Mart. Fl. Bras. 13, pt. 1: 23. pi. 5,111, 
1841. Aberemoa Spixiana R. E. Fries, Sv. Vet. Akad. Handl. 34, 
pt. 5: 22. 1900. 

A small or medium-sized tree or a tall shrub; young branchlets 
scaly; leaves membranaceous, 15-25 cm. long and 5-8 cm. broad, 
oblong-elliptic or narrowly obovate, abruptly cuspidate, rounded- 
acute or sometimes cuneate at the base, smooth and glabrous above, 
even when young, at first densely covered beneath with round, 
appressed scales, soon glabrescent and finally thinly scaly; inflores- 
cences opposite the leaves, 1-4-flowered; buds globular, apiculate; 
sepals densely scaly outside; petals elliptic-orbicular, rounded at the 
apex, 2-2.5 cm. long, clothed with a short, dense pubescence of soft, 
stellate hairs yellowish gray in color; fruit ellipsoid, 6-7 cm. long and 
4-5 cm. broad; monocarps about 2 cm. long, the apex conic, acute. 
-Neg. 6491. 

Loreto: Lower Rio Itaya, near Soledad, Tessmann 5301. Alto 
Rio Itaya, Williams 3292, 3455. Lower Rio Huallaga, Yurimaguas, 
Williams 4348. Amazonian Brazil (Martins 3010, type). 

8. Duguetia latifolia R. E. Fries, Acta Hort. Berg. 12: 103. 
1934. 

A small tree, up to 10 meters high, the young branchlets clothed 
with small, deeply divided, stellate scales or stellate hairs; leaves 
very large, up to 30-40 cm. long and 8-12 cm. broad, cuspidate, 
rounded-acute or sometimes rather obtuse at the base, glabrous 
above and finally nearly glabrous beneath, with a few very small, 
rigid, stellate hairs (or deeply divided, stellate scales) ; inflorescences 
often produced from older branches, many-flowered; pedicels thick, 
rigid, 1-1.5 cm. long, with dense yellow scales, tomentellous; flower 
buds ovoid, acuminate; sepals ovate, acute, about 10 mm. long; 
petals all alike, spatulate, broadly rounded at the apex, 15-18 mm. 
long, rather thinly clothed with short, gray, stellate hairs; fruit 



FLORA OF PERU 735 

spherical, 2-3 cm. across; monocarps about 1 cm. long, the apex 
truncate, gray-tomentellous. 

Loreto: Between Yurimaguas and Balsapuerto, Killip & Smith 
28240, type. Also in the western part of Amazonian Brazil. 

9. PORCELIA R. & P. 

Trees. Leaves ovate to lanceolate-oblong, nearly glabrous, 
with translucent glands. Flowers medium-sized, bisexual or polyga- 
mous, terminal or opposite the leaves, often produced on axillary, 
very reduced branchlets (pseudo-axillary flowers) ; pedicels articulate 
at the base, without bracts. Perianth gray-tomentellous; sepals 
small; petals imbricate in bud and finally widely spreading, all of 
about the same size, oblong-ovate or obovate. Stamens numerous; 
pollen sacs divided by transverse membranes into a row of small 
chambers; connective expanded above the anther into a truncate 
disk. Ovules numerous, lateral. Monocarps large, free, stipitate, 
ellipsoid-cylindric, with large, reniform seeds. A small genus of four 
species, one of them in Peru and adjacent Bolivia, one in Bolivia and 
western Brazil, and two in South Brazil (Goyaz, Minas Geraes, and 
Sao Paulo). 

1. Porcelia nitidifolia R. & P. Syst. Veg. 1: 144. 1798. P. Saf- 
fordiana Rusby, Mem. N. Y. Bot. Gard. 7: 242. /. 2. 1927. P. pon- 
derosa Rusby, op. cit. 245. Gftta#ma(?) ponderosa Rusby, Bull. 
N. Y. Bot. Gard. 6: 504. 1910. 

A large tree with ochraceous, puberulous young branchlets, soon 
glabrescent; leaves soon quite glabrous, oblong, gradually narrowed 
at the apex, more or less rounded at the base, but acute near the 
petiole and obviously decurrent, 15-18 cm. long and 4-5 cm. broad; 
flowers on pedicels 2-3 cm. long; petals up to 16 mm. long; stamens 
2 mm. long, the connective appendix velutinous. 

Huanuco: Chacahuasi, Ruiz & Pawn, type. San Martin: 
Zepelacio, near Moyobamba, Klug 3750. Also in Bolivia: Rurre- 
nabaque, Rusby 1751, type of Porcelia Saffordiana; and Williams 
1479, type of Porcelia (Guatteria) ponderosa. 

In eastern Bolivia and adjacent Brazil another species occurs: Por- 
celia Steinbachii (Diels) R. E. Fries, which may also be found in Peru. 
It is closely related to Porcelia nitidifolia but is characterized by 
shorter leaves of more ovate form, more rounded and less decurrent 
at the base. 



736 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

10. CYMBOPETALUM Benth. 

Trees and shrubs with large, rather thin leaves. Flowers solitary, 
terminal or, usually, produced from the internodes above the axils, 
only rarely from an axil (pseudo-axillary flower) ; pedicels articulate 
at the base, without bracts. Perianth segments valvate in bud; 
sepals small; outer petals flat and thin, the inner thick and fleshy, 
cymbiform, with involute margins. Stamens numerous; pollen sacs 
divided into small chambers by transverse membranes; connective 
expanded above the anther into a truncate disk. Fruit of several 
free, oblong-cylindric monocarps, containing several (4-14) lateral 
seeds. A small genus of 9 species, 5 of them native of Central 
America and 4 of the Amazonian region. One of them, the eastern 
species, C. brasiliense (Veil.) Benth., extends north to Guiana and 
Trinidad, and south to Rio de Janeiro. Only two species are 
recorded from Peru. 

Young branchlets and lower surface of the leaves, especially the 
thicker nerves, clothed with persistent hairs; flowers large 
(inner petals 3-4 cm. long), on very long pedicels (up to 20-30 
cm., rarely shorter) 1. C. longipes. 

Young branchlets and leaves soon glabrous; flowers smaller (inner 
petals about 1.5 cm. long), on pedicels 3-6 cm. long. 

2. C. Tessmannii. 

1. Cymbopetalum longipes Diels, Verh. Bot. Ver. Brandenb. 
47: 132. 1905. 

A low tree with pubescent branchlets; leaves rather rigid, glabrous 
above, hirsute beneath, especially on the nerves, ovate-elliptic or 
ovate-lanceolate, narrowed from the middle to both ends, usually 
acute and oblique at the base, gradually attenuated toward the 
apex, 12-24 cm. long and 4-11 cm. broad; flowers large, the very long 
pedicels reaching 10-20 cm., rarely a little shorter; outer petals 
about 2 cm. long, the inner 3-4 cm., very thick and broadly involute 
at the margins, the midrib and lateral nerves impressed on the out- 
side of the petals. Neg. 27529. 

Huanuco: Vitoc, Ruiz & Pavon. Loreto: Yurimaguas, Williams 
2831. Moyobamba, Weberbauer 4512. San Martin: Tarapoto, 
Spruce 4097, type. Juan Guerra, near Tarapoto, Ule 6912; Williams 
6875, 6876, 6877, 6920. Alto Rio Huallaga, Williams 6161, 6305, 
6512. Tocache, Poeppig. Also known from Brazil (western Ama- 
zonian region) and Bolivia. 



FLORA OF PERU 737 

In Amazonian Brazil, extending west to near the Peruvian 
border, another species of this genus occurs, Cymbopetalum odoratis- 
simum Barb. Rodr., which is very closely related to C. longipes. It 
has the same large flowers and long pedicels, but is distinguished by 
larger leaves with longer tips, quite glabrous and densely verrucu- 
lose-punctulate. It will probably also be found in Peru. 

la. Cymbopetalum longipes var. minor R. E. Fries, var. nov. 

Typo omnino minor; frutex humilis; folia 5-7 cm. longa, 
1.5-2.2 cm. lata; pedicelli breviores, ca. 4 cm. longi; flores minores, 
petala interiora (nondum rite evoluta?) 1 cm. longa. 

San Martin: Juan Guerra, near Tarapoto, Williams 6922 (type in 
Herb. Field Mus.). 

A noteworthy form, agreeing with the type in all characters (leaf 
shape, pubescence, etc.), differing only in the much smaller dimen- 
sions of all parts. 

2. Cymbopetalum Tessmannii R. E. Fries, Acta Hort. Berg. 
10: 188. pi. 7. 1931. 

A low tree, the youngest branchlets very soon quite glabrous, 
verruculose; leaves membranaceous, glabrous, oblong-elliptic or 
oblanceolate, cuspidate, rounded-acute at the base, 10-18 cm. long; 
pedicels glabrous, up to 6 cm. long but usually shorter, produced 
above the axil on one-fourth to one-half the height of the internode; 
flowers rather small, the outer petals about 1 cm. long, the inner 1.5 
cm. long, their margins narrowly involute. 

Loreto: Lower Rio Itaya, near Soledad, Tessmann 524.2, type. 
Upper Rio Itaya, Williams 3513. 

11. UNONOPSIS R. E. Fries 

Shrubs or small to medium-sized trees. Inflorescences axillary 
or sometimes produced from older branches or from the trunk; 
pedicels articulate above the basal bract, and provided with a second 
bract above the articulation. Flowers small; flower buds spherical. 
Sepals minute, valvate. Petals rigid, valvate, ovate or rotund, 
concave, especially the inner thick. Stamens numerous; connective 
expanded above the anther into a truncate disk. Carpels several to 
numerous; ovules few (to 6-8), lateral and horizontal, or else solitary 
and attached near the lower or upper end of the ovary. Monocarps 
free, stipitate. About 24 species, distributed from Honduras and 
West Indies to South Brazil; 9 species are recorded from Peru. 



738 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ovules 2 or more, lateral, horizontal. 
Young branchlets densely covered with soft, patent hairs. 
Leaves elliptic to elliptic-obovate, rounded at the base. 

1. U. Williamsii. 

Leaves oblanceolate, acute at the base 2. U. peruviana. 

Young branchlets quite or nearly glabrous, or with short- 

appressed hairs. 

Monocarps smaller; seeds 7-9 mm. in diameter .4. U. Matthewsii. 
Monocarps larger; seeds 10-14 mm. in diameter. 

Leaves broadest at the middle or below 3. U, grandis. 

Leaves broadest above the middle. 

Branchlets slender; leaves oblanceolate, 3-5 cm. broad. 

5. U. gracilis. 
Branchlets thicker and rigid; leaves obovate, 5-7 cm. broad. 

6. U. obovata. 
Ovule 1. 

Outer petals thickened toward the apex, their margins thick and 

truncate; ovule pendulous from near the top of the ovary. 

Young branchlets glabrous; leaves broadest at the middle, acute 

at the base; inflorescences many-flowered . 7. U. floribunda. 

Young branchlets shortly tomentellous; leaves broadest above 

the middle, obtuse at the base; inflorescence few-flowered. 

8. U. venificiorum. 

Outer petals not thickened, their margins thin, even at the apex; 
ovule erect from near the base of the ovary . . 9. U. stipitata. 

1. Unonopsis Williamsii R. E. Fries, Acta Hort. Berg. 12: 238. 
/. 6f, pi 1. 1937. 

A shrub or small tree; young branchlets, petioles, and pedicels 
densely tomentose; leaves elliptic to elliptic-obovate, rounded and 
obtuse at the base, abruptly cuspidate, finally quite or nearly gla- 
brous, but the midrib clothed with long, persistent, soft hairs above, 
very densely and minutely verruculose-punctate, 15-20 cm. long, 5-9 
cm. broad; flowers solitary or in few-flowered inflorescences; sepals 
about 3 mm. long; outer petals ovate, ferruginous-sericeous outside, 
about 12 mm. long; inner petals thicker, about 10 mm. long; ovules 
5-6, rarely fewer. 

Loreto: Alto Rio Itaya, San Antonio, Williams 3449, type. 
Also in Amazonian Brazil. 



FLORA OF PERU 739 

2. Unonopsis peruviana R. E. Fries, sp. nov. 

Arbor parva, ramulis novellis ferrugineo-tomentosis; foliorum 
petiolus 3-5 mm. longus, crassus, tomentosus; lamina primo utroque 
latere sericea, demum fere omnino glabrescens, nervo medio tamen 
supra persistenter tomentoso, oblanceolata, cuspidata, basin acutam 
versus longe cuneata, 12-16 cm. longa et 4-5 cm. lata; florum 
pedicelli crassi, ferrugineo-tomentelli, fructiferi ca. 15 mm. longi; 
sepala in discum ca. 8 mm. latum cqnnata; petala exteriora rotun- 
data, extus sericea, ca. 12 mm. longa; stamina 3 mm. longa; 
ovaria cinereo-tomentosa, ovulis 2-3 parata; monocarpia globosa 
vel breviter globoso-ellipsoidea, 12-15 mm. diam., stipitibus crassis 
7-8 mm. longis sustenta; semina (1-2 in monocarpiis examinatis) 
ca. 10 mm. diam., castanea. 

Loreto: Rio Nanay, near Altura, in forest, Williams 841- Maqui- 
sapa, on the upper Rio Nanay, Williams 1184 (type; Herb. Field 
Mus. No. 608,102), "Cura-caspi" (Inca), "espintana." 

3. Unonopsis grandis (Benth.) R. E. Fries, Sv. Vet. Akad. 
Handl. 34, pt. 5: 28. 1900. Trigyneia grandis Benth. Journ. Linn. 
Soc. Bot. 5: 70. 1861. 

A shrub or small tree, about 6 meters high, the young branchlets 
very shortly puberulous; leaves on petioles 4-6 mm. long, coriaceous, 
glabrous above (even on the midrib), minutely sericeous beneath 
and finally quite glabrous, elliptic-ovate and more or less oblique, 
rounded or shortly rounded-acute and decurrent at the base, long- 
acuminate at the apex, 12-24 cm. long and 5-8.5 cm. broad, the 
veins impressed above; inflorescences with several flowers; pedicels 
slender, sericeous, 10-12 mm. long; outer petals thinly gray-sericeous, 
attaining 7 mm., the inner 5-6 mm. long; carpels few (4-7), the ovules 
5; monocarps about 20 mm. long and 13-15 mm. broad. Neg. 14241. 

Loreto: Maquisapa, on the upper Rio Nanay, Williams 1204. 
Iquitos, Williams 3645. Also in Amazonian Brazil and Venezuela 
(Spruce 3163, type). 

4. Unonopsis Matthewsii (Benth.) R. E. Fries, Sv. Vet. Akad. 
Handl. 34, pt. 5: 28. 1900. Trigyneia Matthewsii Benth. Journ. 
Linn. Soc. Bot. 5: 69. 1861. 

A shrub or tree, up to 15 meters high, the branchlets rather 
appressed-hairy, soon glabrate; leaves on petioles 3-4 mm. long, 
sericeous on both sides when young, but soon glabrescent or with a 
few appressed hairs on the midrib beneath, oblong-elliptic or obovate- 
oblanceolate, usually acute at the base and shortly decurrent, rather 



740 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

abruptly cuspidate, 10-20 cm. long and 3-8 cm. broad; inflorescences 
in the axils of leaves, usually many-flowered; pedicels slender, 
appressed-hirsute, 1-2 cm. long; outer petals sericeous outside, 
5-6 mm. long; carpels 4-10, with 3-5 ovules; monocarps rufous, 
oblong-cylindric, up to 25 mm. long and 8-10 mm. broad, often 
shorter or even globose, if only one seed is developed; seeds 7-9 mm. 
across. Negs. 27564, 34457. 

San Martin: Tarapoto, Mathews 1421, type; Spruce 4402; Ule 
6341 ; Williams 5386, 5455, 5462, 5467, 5562. Lamas, near Tarapoto, 
Williams 6490. Juanjui, Alto Rio Huallaga, King 3839. Chazuta, 
Rio Huallaga, King 4062. Rio Acre: San Francisco, Ule 9369. 

5. Unonopsis gracilis R. E. Fries, Acta Hort. Berg. 12: 247. 
/. 5, f-h. 1937. 

A small tree with slender branches, the young branchlets thinly 
sericeous and soon glabrescent; leaves on petioles 3-5 mm. long, 
glabrous on both surfaces, oblanceolate or narrowly oblanceolate, 
long-acuminate, still longer narrowed to the acute and shortly 
decurrent base, 12-20 cm. long and 2.5-5 cm. broad, rarely broader; 
inflorescences few-flowered, on peduncles 2-3 mm. long; pedicels 
very slender, sericeous, 18-25 mm. long; outer petals thinly sericeous 
outside, 7-8 mm. long; carpels very few (2-5); ovules 4; monocarps 
fleshy and orange-red when fresh, oblong-ellipsoid, up to 3.5 cm. 
long and 1.5-2 cm. in diameter, their stipes thick, 3-5 mm. long; 
seeds disciform, about 12 mm. across. 

Loreto: Yurimaguas, Killip & Smith 27985. Yurimaguas, For- 
taleza, Williams 4188, type. Yurimaguas, Paranapura, Williams 
4608. Yurimaguas, Puerto Arturo, Williams 5346. Santa Rosa, 
Killip & Smith 28824. Balsapuerto, Killip & Smith 28583. 

6. Unonopsis obovata R. E. Fries, Acta Hort. Berg. 12: 249. 
/. 5, a-e. 1937. 

A shrub or small tree, the young branchlets sericeous, soon 
glabrescent; leaves on petioles 3-5 mm. long, finally glabrous on 
both sides, but with a few minute hairs on the thicker nerves beneath, 
obovate, generally abruptly contracted into a cusp 1-2 cm. long, 
acute or obtuse at the base, 12-17 cm. long and 5-7 cm. broad; 
inflorescences in the axils of persistent or recently fallen leaves, 
bearing 1-2 fruits (flowers not seen); pedicels rigid, 1-2 cm. long; 
monocarps few (up to 6), ellipsoid-oblong, fleshy and orange when 
fresh, glabrous, up to 2.5 cm. long and 1.3-1.7 cm. broad; stipes 
short and thick, 3-5 mm. long; seeds up to 4, disciform. 



FLORA OF PERU 741 

Loreto: Yurimaguas, Killip & Smith 27985, type; Williams 4112. 
San Martin: Tarapoto, Williams 6303. 

7. Unonopsis floribunda Diels, Notizbl. Bot. Gart. Berlin 9: 
137. 1924. 

A tree up to 20 meters high, with slender, glabrous branchlets; 
leaves on petioles 3-6 mm. long, glabrous on both sides, elliptic or 
lanceolate, rather equally narrowed to both ends, acute at the base, 
12-25 cm. long and 4-8 cm. broad; inflorescences freely branched and 
many-flowered, issuing from leafless parts of the branches; pedicels 
10-20 mm. long, slender; flowers small, the outer petals 7-10 mm. 
long, thinly gray-sericeous outside; carpels numerous (10-22 counted) ; 
ovule one, pendulous, attached near the upper end of the ovary; 
monocarps globose-clavate, on stipes 7-10 mm. long. Neg. 14240. 

Loreto: Ucayali, Cashibo Playa, Tessmann 3262, type. Mishu- 
yacu, near Iquitos, Klug 974- An apparently common species in the 
western regions of Amazonian Brazil, entering the eastern part 
of Peru. 

8. Unonopsis veneficiorum (Mart.) R. E. Fries, Acta Hort. 
Berg. 12: 259. 1937. Guatteria veneficiorum Mart, in Buchner, 
Repert. Pharmacie 36: 344. 1830; Fl. Bras. 13, pt. 1: 34. 1841. 

A shrub or tree, up to 10 meters high, the young branchlets very 
shortly but densely rufous- tomentellous; leaves on petioles 3-7 mm. 
long, oblong-oblanceolate or obovate, long-acuminate, usually 
long-cuneate toward the rounded base, 20-30 cm. long and 5-10 cm. 
broad, strigose on the thicker nerves when young, but soon quite 
glabrous; inflorescences in the axils of persistent or fallen leaves, 
sessile, few-flowered, their internodes very short; pedicels sericeous, up 
to 2.5-3 cm. long; outer petals 8-9 mm. long and broad, rufous- 
sericeous; carpels about 12; ovule solitary, pendulous from near the 
upper end of the ovary; monocarps on stipes 1-2 cm. long, clavate- 
ellipsoid, 1.5-2 cm. long; seed ellipsoid and slightly compressed from 
the sides, 11-15 mm. long. 

Loreto: San Antonio, on Rio Itaya, Killip & Smith 29340, 
29370, 29530; Williams 3408. Soledad, Rio Itaya, Killip & Smith 
29723, 29733. Iquitos, Tessmann 5131; Killip & Smith 27086; 
Williams 3772. Mishuyacu, near Iquitos, Killip & Smith 29935; 
Klug 1556. Caballo-cocha, on the Amazon River, Williams 2320. 
Pebas, Williams 1941. Also in western Amazonian Brazil (Martins, 
type; Krukoff 8149). 



742 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

9. Unonopsis stipitata Diels, Verb. Bot. Ver. Brandenb. 47: 
130. 1905. 

A small tree, 5 meters high, the young branchlets fulvous- tomentel- 
lous, soon glabrous; leaves on thick petioles 4-8 mm. long, rigid, 
glabrous and lustrous above, glabrous and sometimes with sparse 
hairs on the midrib beneath, obovate or obovate-oblong, rounded 
at the apex and very abruptly cuspidate, long-attenuate toward the 
rounded-acute or cuneate base, 20-28 cm. long and 5-9 cm. broad; 
inflorescences issuing from rather thick branches; outer petals ful- 
vous-sericeous outside, often striate, about 12 mm. long, their 
margins thin up to the apex but the petals nevertheless valvate in 
bud; connective appendix papillose; carpels very numerous, 50-60 
or even more; ovule solitary, erect, attached very near the base of 
the ovary; monocarps obliquely clavate-ellipsoid, 11-15 mm. long 
and on stipes 1-2 cm. long. 

Loreto: Leticia, Ule 6191, type. Lower Rio Nanay, Williams 
650. Distributed eastward to Amazonian Brazil, French Guiana, 
and Surinam. 

12. DICLINANONA Diels 

Small trees with rather large, oblong to oblanceolate-obovate 
leaves. Inflorescences axillary. Flowers medium-sized, polygamous 
(or dioecious?). Sepals small, free, valvate. Petals valvate, linear 
to oblong, concave at the base on the inner side, with 2 oblong 
glands at the margins. Stamens numerous in the staminate flowers, 
few in the bisexual, their connectives expanded and truncate above 
the pollen sacs. Carpels few, globose-ovoid, pilose; stigma sessile, 
glabrous; ovules 6-9, lateral, horizontal. Monocarps globose, with 
thick and ligneous pericarp, indehiscent; seeds few, flat, horizontal. 
Only two species known, both occurring in Peru. 

Leaves acute and decurrent at the base, gradually continuing in the 
edges of the 10-15 mm. long petiole; pedicels very slender, 
1-2 cm. long; petals 1-2 mm. broad 1. D. Tessmannii. 

Leaves decurrent as narrow wings on the petiole, the lower end of 
the wings suddenly contracted; petioles 4-6 mm. long; pedicels 
thicker, 5-8 mm. long; petals 2.5-3.5 mm. broad . . 2. D. calycina. 

1. Diclinanona Tessmannii Diels, Notizbl. Bot. Gart. Berlin 
10: 174. 1927. 

A small tree, up to 6 meters high; leaves large (15-25 cm. long 
and 5-8 cm. broad), obovate-elliptic, long and narrowly cuspidate, 
cuneate at the base, glabrous from the beginning on the upper surface 



FLORA OF PERU 743 

(even on the midrib), appressed-hirsute beneath when young, but 
soon quite glabrous; flowering branchlets very short, usually pro- 
duced from the axils of fallen leaves; flowers few (up to 5), on slender 
pedicels 1-2 cm. long; only staminate flowers known; sepals oblong- 
ovate, 7-8 mm. long; petals nearly linear, spreading, 16-20 mm. long 
and 1.5-2 mm. broad. 

Loreto: Iquitos, Tessmann 5387, type. 

2. Diclinanona calycina (Diels) R. E. Fries, Acta Hort. Berg. 
12: 4. 1934. Xylopia calycina Diels, Notizbl. Bot. Gart. Berlin 
10: 175. 1927. 

A low tree; leaves 10-20 cm. long, 3-6 cm. broad, glabrous and 
lustrous above, the midrib with persistent hairs, thinly sericeous 
beneath, oblanceolate-obovate, abruptly cuspidate, cuneate toward 
the base, on each side of the petiole forming a narrow wing (5-8 mm. 
long) which is suddenly contracted to a more or less obtuse base, 
the true petiole 4-6 mm. long; flowers numerous, in dense, freely 
branched, ferruginous-sericeous inflorescences, usually produced 
from the axils of persistent leaves; pedicels 5-8 mm. long; staminate 
and bisexual flowers are known; sepals 5 mm. long; petals erect, 
connivent, linear-oblong, densely sericeous, up to 20 mm. long; 
monocarps (only one noticed in each fruit) sessile, globose, 3-3.5 
cm. in diameter; pericarp ligneous, about 2 mm. thick. Neg. 14250. 

Loreto: Mishuyacu, near Iquitos, Klug 96. La Victoria, on the 
Amazon, Williams 2756. Upper Maranon, near mouth of Rio 
Santiago, Tessmann 4369, type. 

13. XYLOPIA L. 

Shrubs or trees. Inflorescences axillary or sometimes produced 
from old branches or from the trunk, sessile or short-pedunculate; 
flowers few (rarely one) or numerous, usually small, their pedicels 
short. Sepals small, valvate in bud, nearly free or connate into a 
cupular calyx. Petals thick, valvate, linear-oblong or sometimes 
deltoid. Stamens numerous, the anther cells divided by transverse 
membranes into a row of small chambers; connective expanded 
above the anther. Carpels few to numerous, free; ovaries hidden 
in a deep cavity in the center of the torus. Monocarps free, clavate 
to cylindric, containing a row of rather few seeds. A large genus, 
with 45 American species; also in the tropics of the Old World. 

Petals linear-oblong; flower buds elongate, cylindric-trigonous. 
Sepals free, or connate at the base only; carpels few (3-12). 



744 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves appressed-hairy beneath. 
Pedicels and calyx hairy. 
Leaves finely reticulate above; stigmas long-hairy. 

1. X. micans. 
Leaves smooth above; stigmas glabrous. . .2. X. densiflora. 

Pedicels and calyx glabrous 3. X. ligustrifolia. 

Leaves covered with patent hairs beneath .... 4. X. conjungens. 
Sepals connate into a more or less cup-shaped calyx, merely dentic- 
ulate at the margin, and sometimes splitting irregularly. 

Plants with axillary flowers 5. X. aromatica. 

Plants cauliflorous 6. X. Benthamii. 

Outer petals ovate or elongate-deltoid, inner petals elongate-rhombic; 

flower buds ovoid to short-conic. 
Leaves small (up to 11 cm. long), acute at the base. 

7.X. peruviana. 

Leaves large (14-25 cm. long), more or less rounded at the base. 
Leaves rounded-acute at the base, with sparse and short hairs 

beneath 8. X. cuspidata. 

Leaves rounded and often emarginate at the base, covered with 
long, dense hairs beneath 9. X. Poeppigii. 

1. Xylopia micans R. E. Fries, Acta Hort. Berg. 12: 276. 1937. 

A small tree, about 5 meters high; leaves rather small (6-9 cm. 
long and 1.3-2 cm. broad), narrowly lanceolate, very shortly acute 
or nearly truncate at the base, long-caudate-acuminate at the apex, 
glabrous and finely reticulate above, very densely covered with 
appressed, golden brown, long, persistent hairs beneath ; sepals nearly 
free, broadly ovate, 1.5 mm. long; outer petals linear-ligulate, up 
to 18 mm. long, the inner petals a little shorter and narrower; carpels 
about 6; stigmas covered in their upper part with long, patent hairs. 

Loreto: Balsapuerto, 220 meters, Klug 3034, type. 

2. Xylopia densiflora R. E. Fries, sp. nov. 

Arbor ca. 20 m. alta, ramulis novellis adpresse hirsutis. Foliorum 
petiolus 5-6 mm. longus; lamina papyracea, adulta supra glabra et 
laevis, subtus pilis longis adpressis subdense vestita, lanceolata, 
basi sensim acutata, apice longius sensimque angustata, 7-10 cm. 
longa, 2-2.5 cm. lata. Flores numerosi, dense glomerati. Sepala 
ovata, extus sericea, 2-2.5 mm. longa. Alabastra tereti-trigona. 
Petala exteriora plana, oblonga, extus cinereo-sericea, 8-10 mm. 



FLORA OF PERU 745 

longa, interiora lineari-clavata, subtrigona, ca. 9 longa. Stamina 
0.7-0.8 mm. longa, connectivi disco glabro. Pistilla ca. 6; styli basi 
geniculati, una cum stigmatibus glabris 2 mm. longi. 

Loreto: Maucallacta, Rio Paranapura, King 3923 (type in Herb. 
Field Mus.). 

3. Xylopia ligustrifolia Dun. Monogr. Anon. 121. pi. 18. 1817. 
X. ruscifolia Humb. & Bonpl. ex Dun. op. cit. 121. X. usitata Diels, 
Notizbl. Bot. Gart. Berlin 9: 140. 1924. Xylopicrum ligustrifolium 
Kuntze, Rev. Gen. 1: 8. 1891. 

A large tree; leaves small (4-6 cm. long and 1.5-2 cm. broad), 
oblong-elliptic, shortly narrowed to the obtuse apex, acute or more 
or less truncate at the base, glabrous above, strictly appressed-hairy 
and glabrescent beneath; inflorescences 1-2-flowered, axillary; pedi- 
cels glabrous, black; sepals glabrous, nearly free, broadly ovate, 
about 2 mm. long; outer petals linear, finally narrowly ligulate, 15-16 
mm. long and about 3 mm. broad, sericeous; inner petals somewhat 
shorter and narrower; carpels 8-12, the stigmas subulate, penicillate- 
pilose; monocarps clavate, brown, glabrous, 12 mm. long or shorter; 
seeds normally 2, but often only 1 developed. Neg. 14255. 

Loreto: Rio Ucayali, Yarina-cocha, Tessmann 321 7, 5549. Also 
in Colombia and Amazonian Brazil. 

4. Xylopia conjungens R. E. Fries, Acta Hort. Berg. 10: 333. 
1931. 

A tree about 20 meters high; young branchlets covered with dense, 
patent hairs; leaves small (4-6.5 cm. long and 1-2.2 cm. broad), very 
shortly petiolate, elliptic-lanceolate, shortly narrowed to the obtuse 
base, cuspidate at the apex, glabrous and lustrous above, densely 
covered with soft, patent hairs beneath; flowers axillary, solitary; 
calyx deeply lobed, 5-6 mm. across, long-hairy outside; outer 
petals up to 17 mm. long, linear-oblong, clothed with long, dense 
hairs; carpels about 7; styles and stigmas filiform, hirsute; fruits 
unknown. 

Loreto: Mishuyacu, near Iquitos, Klug 970, type. 

Closely related to Xylopia barbata Mart, from the lower Amazon, 
but easily distinguished by its long, acuminate leaves. 

5. Xylopia aromatica (Lam.) Mart. Fl. Bras. 13, pt. 1: 43. 
1841. Cf. R. E. Fries, Acta Hort. Berg. 10: 106. 1930, where 23 syno- 
nyms are quoted. 



746 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tree; leaves large, 8-15 cm. long and 2-4.5 cm. broad, lanceolate 
or elliptic-lanceolate, gradually narrowing at the apex, rounded or 
shortly acute at the base, glabrous above except on the midrib, 
glaucous and hairy beneath; flowers numerous, in axillary inflores- 
cences; sepals almost wholly connate into a cup-shaped calyx; 
flower buds long and narrow, triquetrous, densely sericeous; outer 
petals finally up to 30 mm. long and 2-3 mm. broad; carpels about 25; 
monocarps cylindric, torulose, 2-2.5 cm. long, with about 6, some- 
times less, seeds. 

San Martin: Tarapoto, Ule 6695, Williams 5378, 5850Loreto: 
Florida, Rio Putumayo, Klug 2074- Balsapuerto, Klug 2845. A 
very common species in tropical America, widely distributed from 
the West Indies and Central America (Costa Rica) down to South 
Brazil (Sao Paulo, Matto Grosso) and Paraguay. 

6. Xylopia Benthamii R. E. Fries, var. subnuda R. E. Fries, 
Acta Hort. Berg. 10: 333. 1931. 

A small tree; young branchlets slender, with sparse, patent hairs; 
leaves 8-10 cm. long and 2-2.5 cm. broad, membranaceous, oblong- 
lanceolate, rather abruptly contracted into a long, obtuse cusp, acute 
at the base, glabrous above; young leaves with long, patent hairs 
beneath, soon quite glabrous; flowers clustered on the trunk or old 
branches; calyx a low cup, about 1 cm. across, the teeth hardly 
distinguishable; petals ligneous, glabrous and blackening when dried, 
linear, acute; outer petals 26-28 mm. long and 5-6 mm. broad, the 
inner a little shorter and narrower; monocarps glabrous and obliquely 
striate, oblong, 3.5 cm. long and about 1.5 cm. broad; seeds 5-6. 

Loreto : Mishuyacu, near Iquitos, Klug 557, type of the variety. 
The type of the species occurs in British and Dutch Guiana, Vene- 
zuela, and Amazonian Brazil. The Peruvian variety differs in its 
very sparse hairiness and its larger flowers. 

7. Xylopia peruviana R. E. Fries, Acta Hort. Berg. 10: 117. 
/. 10. 1930. 

Leaves rigid, elliptic, short-cuspidate at the apex, short-acute at 
the base, glabrous above, appressed-hairy beneath when young, 
soon glabrescent and very densely reticulate, 6-11 cm. long and 3-5 
cm. broad; flowers solitary in the axils of the leaves, on pedicels 3-4 
mm. long; sepals connate to the middle, 4 mm. long; flower buds 
conic, densely ferruginous-sericeous; carpels numerous; monocarps 
obliquely cylindric, apiculate, 2 cm. long; seeds 4, often less. 






FLORA OF PERU 747 

San Martin: Tarapoto, Williams 6192, 6225; Spruce 4166, type. 
Yurimaguas, Spruce 4598. 

8. Xylopia cuspidata Diels, Notizbl. Bot. Gart. Berlin 10: 175. 
1927. 

A large tree, the branchlets appressed-pilose, ferruginous; leaves 
large (about 25 cm. long and 9 cm. broad), elliptic-oblong, rounded 
or very shortly acuminate at the base, rounded at the apex, and 
terminating in a narrow cusp 3-4 cm. long, glabrous above, thinly 
and shortly appressed-pilose beneath; flowers solitary on old, leafless 
branches; calyx cup-shaped, 3-4 mm. high, with very short lobes; 
outer petals ovate- triangular, acute, sericeous; inner petals lanceo- 
late, shorter and narrower; carpels numerous, with 4-5 ovules; 
styles and stigmas glabrous, conglutinated ; fruits unknown. Neg. 
14251. 

Loreto: Lower Rio Itaya, Soledad, Tessmann 5323, type. 

9. Xylopia Poeppigii R. E. Fries, Acta Hort. Berg. 10: 121. 
1930. Uvaria ferruginea Poepp. in sched. 

Young branchlets covered with long, more or less patent, ferrugi- 
nous hairs; leaves about 22 cm. long and 7 cm. broad, membrana- 
ceous, rounded and slightly cordate at the base, abruptly contracted 
at the apex into a cusp 2 cm. long or sometimes longer, glabrous 
above, rather densely clothed beneath with long, appressed hairs; 
flowers solitary on short pedicels; sepals long-connate, ferruginous- 
hirsute on the outside, the lobes broadly triangular, acute; petals 
thick, ligneous, the outer ovate- triangular, 20-22 mm. long and about 
1 cm. broad, on the outside densely covered with silver-gray, ap- 
pressed hairs; inner petals a little shorter and narrower, very acute, 
keeled along the back; carpels numerous, with glabrous stigmas; 
ripe fruits not known. Neg. 14258. 

Loreto: Mainas, Poeppig 2219, type. Yurimaguas, Williams 
4105; Killip & Smith 28199, 28732. Balsapuerto, King 3048; 
Killip & Smith 28391, 28612; Kuhlmann (Herb. Rio de Janeiro 
24267}. 

This species is very closely related to Xylopia cuspidata; further 
material is needed to decide whether the two species may be kept 
separate. 

14. ANAXAGOREA St. Hil. 

Small trees or shrubs. Flowers small, axillary, sometimes pro- 
duced on the older, leafless branches; pedicels bracteolate. Sepals 
thin, valvate in bud, sometimes connate at first and irregularly 



748 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

splitting, often deciduous. Petals valvate, all rather alike but the 
inner mostly a little smaller and thicker, all or only the inner keeled 
on the inner surface above the middle. Stamens several, the inner- 
most often sterile; connectives of the fertile stamens not forming a 
disk above the anthers, but elongated into a ligule, sometimes very 
short. Carpels several, with 2 basal, erect ovules. Monocarps free, 
dry, stipitate, club-shaped, opening along the inner side. Seeds 2, 
smooth and lustrous, plane on one side, convex on the other. The 
genus contains 19 species, which occupy a vast region from Central 
America to South Brazil, centering along the Amazon. A few species 
are native of tropical Asia. 
Sepals free even in bud. 

Pedicels 15-25 mm. long, the uppermost bract below their middle; 
outer petals not keeled on the inner surface. 1. A. pachypetala. 

Pedicels about 5 mm. long, the uppermost bract at their top close 
to the calyx; all petals keeled on the inner side. .2. A. minor. 
Sepals connate in bud, irregularly splitting and soon deciduous. 

Outer petals ovate-elliptic, about 10 mm. long. 3. A. megalophylla. 

Outer petals narrowly lanceolate, about 15 mm. long. 

4. A. phaeocarpa. 
Insufficiently known species 5. A. pallida. 

1. Anaxagorea pachypetala (Diels) R. E. Fries, Acta Hort. 
Berg. 10: 175. 1931; 12: 10. 1934. Oxandra pachypetala Diels, 
Notizbl. Bot. Gart. Berlin 10: 173. 1927. Guatteria pachypetala 
Macbr. Field Mus. Bot. 4: 171. 1929. 

A small tree; leaves membranaceous, glabrous, oblong, gradually 
attenuate into a long, narrow cusp, shortly acute at the base, 6-12 
cm. long and 1.5-3 cm. broad; petiole 4-6 mm. long, slender; pedicels 
solitary, slender but rigid, 1.5-2.5 cm. long, bearing below the middle 
a 1.5-2 mm. long bract, amplexicaul; sepals free, ovate, 4-5 mm. 
long; outer petals ovate, rather thick but not keeled on the inner 
side, 8-10 mm. long; inner petals rhombic-obovate, keeled above 
the middle on the inner surface, about 7 mm. long; connective 
scarcely elongated above the anther, rounded-truncate. Neg. 27517. 

Loreto: Rio Maranon, at the mouth of Rio Apaga, Tessmann 
4893, type. 

2. Anaxagorea minor Diels ex R. E. Fries, Acta Hort. Berg. 
12: 22. /. 2, a-e. 1934. 

A small tree with glabrous young branchlets; leaves on petioles 
4-6 mm. long, thin, glabrous, oblong-elliptic, sometimes oblanceolate, 



FLORA OF PERU 749 

acute and decurrent at the base, more or less abruptly cuspidate 
at the apex, 12-16 cm. long and 3-5 cm. broad; inflorescences axillary 
or pseudoterminal, few- (or one-) flowered; pedicels about 5 mm. 
long, rigid, provided with a collar-shaped bract close to the calyx; 
sepals not connate, ovate, 8-10 mm. long; petals up to 15 mm. long, 
lanceolate-oblong, thick, and all keeled on the inner side above 
the middle; connective elongated above the anther to a ligule 
0.6 mm. long. 

Loreto: On the Rio Nanay, near Altura, Williams 849, 854 (type), 
855. Balsapuerto, Killip & Smith 28610. 

3. Anaxagorea megalophylla R. E. Fries, Acta Hort. Berg. 
12: 20. 1934. 

A small tree, 6-9 meters high, the young branchlets ferruginous- 
tomentellous; petioles about 12 mm. long; lamina membranaceous, 
very large, about 35 cm. long and 12-14 cm. broad, obovate and 
abruptly contracted at the apex into a cusp 3 cm. long, longer-attenu- 
ate and decurrent at the base, glabrous on both sides, but minutely 
puberulous beneath when very young, especially on the midrib; 
inflorescences few-flowered; pedicels thick, about 5 mm. long; sepals 
connate in bud, splitting irregularly; petals thick, the outer elliptic 
and not keeled on the inside, the inner rhombic-obovate and keeled 
above the middle on the inner side. 

Peru(?): Monterico, Pearce, type. 

4. Anaxagorea phaeocarpa Mart. Fl. Bras. 13, pt. 1: 40. 1841, 
pro parte; R. E. Fries, Acta Hort. Berg. 12: 21. 1934. 

A shrub or small tree, the young branchlets rufous- tomentellous; 
leaves membranaceous, on petioles 8-15 mm. long, glabrous above, 
when young minutely rufous-puberulous beneath, especially on the 
midrib, finally glabrate, oblong to oblong-oblanceolate, long-cuspi- 
date, rounded-acute at the base and long-decurrent, with the margins 
revolute, up to 25 cm. long and 9 cm. broad; inflorescences 1-4- 
fiowered, produced from old branches; pedicels 1-1.5 cm. long, 
thick, provided with a collar-shaped bract near the top; sepals 
connate in bud, splitting irregularly; petals narrowly lanceolate, long- 
attenuate, the inner keeled on the inside, 14-18 mm. long and 4-6 
mm. broad; connective elongate above the anther into a rounded- 
truncate ligule 0.5-0.6 mm. long. Neg. 6494. 

Loreto: Balsapuerto, Killip & Smith 28529. Also in Amazonian 
Brazil. 



750 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5. Anaxagorea pallida Diels is an incompletely known species 
from Peru, described from only fruiting material (Notizbl. Bot. 
Gart. Berlin 11: 79. 1931). Its leaves and long pedicels are 
reminiscent of Anaxagorea pachypetala, its leaf shape also of 
A. minor. Since the flowers, which afford the best specific characters 
of this genus, are unknown, it is hardly possible to decide the specific 
status of this plant. 

Known only from Loreto: Santa Rosa on the lower Rio Huallaga, 
Williams 4873, type. 

15. GUATTERIOPSIS R. E. Fries 

Trees; leaves medium-sized or large, oblong to oblanceolate. 
Flowers axillary, easily detached; pedicels very short, articulate above 
the middle, the lower part bracteate, the upper without bracts. 
Sepals and petals thick, rigid, valvate in bud, the petals all alike. 
Stamens numerous; connective expanded above the pollen sacs into 
a truncate disk. Carpels numerous; ovule solitary, basal, erect. 
Monocarps free, stipitate, fusiform or elongate-ovoid, dry, one- 
seeded. A small genus of 4 species, centering in Amazonian 
Brazil, one of them entering Peru, another extending south to Minas 
Geraes. 

1. Guatteriopsis sessiliflora (Benth.) R. E. Fries, Acta Hort. 
Berg. 12: 109. 1934. Annona sessiliflora Benth. in Hook. Journ. 
Bot. 5: 8. 1853. Guatteria sessiliflora Saff. Contr. U. S. Nat. Herb. 
18: 6. 1914. 

A small tree, up to 6 meters high, the young branchlets appressed- 
hirsute, strigose; leaves densely verruculose on both sides, from the 
beginning glabrous above, at first densely covered with appressed 
hairs beneath, but finally more or less glabrescent, oblong or oblong- 
lanceolate, short-cuneate and decurrent at the base, abruptly cuspi- 
date at the apex, 15-25 cm. long and 5-8 cm. broad; inflorescences 
axillary, usually 2-flowered; pedicels about 4 mm. long; perianth 
covered with dense, gray hairs; petals 11-15 mm. long; monocarps 
cylindric-obovoid, rounded at the apex, about 17 mm. long and 7-8 
mm. broad. 

Loreto: Along Rio Maranon, near mouth of Rio Tigre, Killip 
& Smith 27522. Also in Amazonian Brazil. 

16. ANNONA L. 

Shrubs or trees with simple or stellate hairs. Flowers solitary 
or in few-flowered inflorescences, terminal or opposite the leaves, or 



FLORA OF PERU 751 

extra-axillary from the internodes. Sepals valvate. Petals 6, free 
or connate at the base, 2-seriate, or the inner rudimentary or absent, 
the outer valvate, the inner imbricate or valvate. Stamens numer- 
ous, the connective generally terminating in a swollen head or 
hoodlike process above the pollen sacs. Carpels numerous; ovule 1, 
basal, erect. Fruit fleshy, formed by the concrescent carpels and 
torus, usually areolate on the surface, the areoles indicating the 
united carpels and often gibbous or acuminate. A large genus of 
about 100 American species; also in tropical Africa. 

Petals free. 

Flower buds globose to ovoid; outer petals round to ovate. 
Petals 6, the inner imbricate, of about the same size as the outer. 
Leaves glabrous beneath, at least when mature, with minute 
pockets or pits in the angles between the midrib and the 
nerves. 
Outer petals abruptly cuspidate; fruit with upward-curved 

spines . . . 1. A. muricata. 

Outer petals gradually narrowed to the apex; spines of the 

fruit patent, straight. 
Leaves 10-20 cm. long, their pits setulose; fruit large, 

up to 15 cm. across 2. A. montana. 

Leaves 6-10 cm. long, the pits glabrous; fruit small, 

3-6 cm. long 3. A. deminuta. 

Leaves softly pubescent beneath, without pits or pockets in 

the axils of the lateral nerves 4. A. excellens. 

Petals 3, the inner absent or, rarely, rudimentary, very minute. 
Leaves softly pubescent beneath with patent hairs. 
Sepals small, much shorter than the petals. 

Leaves medium-sized, 3-6 cm. broad; sepals shortly 
cuspidate; fruits with appressed points on the areoles. 

5. A. scandens. 

Leaves large, 7-12 cm. broad; sepals long-cuspidate; 
fruits with patent, usually recurved points on the 

areoles 6. A. cordifolia. 

Sepals large, broadly ovate and nearly as long as the petals. 

7. A. macrocalyx. 
Leaves appressed-hairy or glabrous beneath. 

Leaves more or less elliptic, with persistent hairs on the 
lower surface. . . .8. A. Tessmannii. 



752 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves oblong, glabrous beneath when mature. 

9. A. iquitensis. 
Flower buds elongate, more or less triquetrous-cylindrical; outer 

petals linear-oblong, the inner rudimentary or absent. 
Leaves appressed-hirsute or finally quite glabrous. 
Leaves lanceolate-elliptic, rounded or obtuse, not acuminate 

at the apex 10. A. squamosa. 

Leaves lanceolate, acuminate 11. A. reticulata. 

Leaves with soft, patent, persistent pubescence beneath. 

12. A. Cherimolia. 
Petals connate at the base. 

Flower buds globose; outer petals round to ovate, up to 7 mm. 

long; leaves less than 12 cm. long. 
Leaves obovate to elliptic, short-acuminate; inner petals nearly 

as long as the outer 13. A. Duckei. 

Leaves lanceolate, long and gradually attenuate at the apex; 
inner petals reduced, about 1 mm. long and much shorter 

than the outer 14. A. neglecta. 

Flower buds elongate-conic; outer petals oblong, 15-25 mm. long; 
leaves 15-25 cm. long 15. A. Ambotay. 

1. Annona muricata L. Sp. PI. 536. 1753. A. Bonplandiana 
HBK. Nov. Gen. & Sp. 5: 58. 1821. A. cearensis Barb. Rodr. PL 
Nuev. Cult. Jard. Bot. Rio Jan. 6: 3. pi. 2. 1898. A. macrocarpa 
Werckle", Tropenpflanzer 428. 1903. A. muricata L. var. borinquensis 
Morales, Fl. Arb. Cub. 60. 1887. Guanabanus muricatus Gomez, Fl. 
Haban. 114. 1897. 

A small tree, up to 8 meters high, with ferruginous-sericeous 
branchlets; leaves rather thin, glabrous and lustrous above, sericeous 
beneath when young, soon glabrate and olivaceous, with small, 
pocket-shaped, hairy pits in the nerve angles, obovate or, rarely, 
oblong, shortly and obtusely acute at the apex, rounded-acute at 
the base, 8-12 cm. long; flowers solitary, on ferruginous-sericeous 
pedicels 1.5-2 cm. long; outer petals cordate at the base, rounded- 
ovate and abruptly cuspidate, 2.5-3.5 cm. long; fruit large, attaining 
15-20 cm., ovoid or oblong-ovoid, dark green, bearing numerous long, 
up-curved spines. 

A cultivated species with edible fruits; probably native of the 
West Indies. Specimens seen from: Loreto, Ule 6522; Caballo-cocha, 
Williams 2399. 



FLORA OF PERU 753 

2. Annona montana Macfad. Fl. Jam. 1: 7. 1837; emend. 
R. E. Fries. A. muricata Veil. Fl. Flum. 5: pi. 126. 1827 (non L.). 
A. muricata L. var. Dun. Monogr. Anon. 63. 1817. A. Marcgravii 
Mart. Fl. Bras. 13, pt. 1: 5. 1841. A. Pisonis Mart. op. cit. 5. A. 
sphaerocarpa Splitg. Tijdsch. Nat. Gesch. 9: 96. 1842. 

A small or medium-sized, glabrous tree; leaves chartaceous, 
lustrous and glabrous above even when young, at first very shortly 
grayish-sericeous beneath, soon glabrate, obovate or elliptic, shortly 
and obtusely cuspidate, acute at the base, 10-20 cm. long and 4-8 cm. 
broad, the pits in the nerve angles of the lower surface rather large, 
more or less rounded and ciliate; outer petals broadly ovate, gradu- 
ally attenuate but not cuspidate, 2-3 cm. long; fruit globose to ovoid- 
globose, up to 15 cm. in diameter, with short, straight prickles. 

Loreto: Caballo-cocha on the Amazon River, Williams 2400. 
Iquitos, Williams 3687. Yurimaguas, Williams 3818, 4450. San 
Martin: Tarapoto, Williams 5438, 5545. Chazuta, Rio Huallaga, 
Klug 3973. "Huanabana" or also "chirimoya" (this name generally 
used for Annona Cherimolia). Distributed over a large area, from 
the West Indies to South Brazil. As it is taken here, "sensu lato," 
it contains several not sharply defined forms, which can hardly be 
regarded as different species. 

3. Annona deminuta R. E. Fries, Acta Hort. Berg. 12: 212. 
/. 14, a-d. 1934. 

A shrub with glabrous branchlets; leaves small, on petioles 5-6 
mm. long, chartaceous, glabrous when mature, glossy above, oliva- 
ceous beneath, with small but deep, round pits without hairs, obovate 
or broadly oblanceolate, cuneate to the acute or rarely rounded- 
acute base, abruptly contracted at the apex into an obtuse cusp 5-10 
mm. long, 6-10 cm. long and 2-4.5 cm. broad; flowers solitary, on 
pedicels 12-15 mm. long; outer petals broadly ovate, gradually 
narrowed to the apex, shortly ferruginous-sericeous outside, about 20 
mm. long; fruit globose-ovoid, 3-6 cm. long and 3-4.5 cm. in diam- 
eter, with straight, conic prickles 3-4 mm. long. 

San Martin: Alto Rio Huallaga, Tarapoto, Williams 5579, type; 
6862. 

4. Annona excellens R. E. Fries, Acta Hort. Berg. 10: 215. 
pi. 10. 1931. 

A small tree; young branchlets densely ferruginous- tomentose; 
petiole about 1 cm. long; lamina coriaceous, elliptic to oblong-elliptic, 
shortly acute or cuspidate at the apex, rounded or very shortly acute 



754 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

at the base, glabrous above and covered with a rufous-gray, persistent 
pubescence beneath, 15-21 cm. long and 6-9 cm. broad; inflores- 
cences few-flowered, sessile, issuing from the middle of the internode 
above the axil; pedicels about 1.5 cm. long; outer petals rounded- 
ovate, acute, pale ferruginous-sericeous outside, 2.5-3 cm. long; 
inner petals obovate, with dorsal keel, rounded at the apex and 
contracted to the base, about 2.5 cm. long; stamens 5 mm. long. 
Loreto: Seringal Auristella, Rio Acre, Ule 9374, type. Also 
in western Amazonian Brazil (Rio Jurua). 

5. Annona scandens Diels, Verh. Bot. Ver. Brandenb. 47: 134. 
1905. 

A climbing shrub with tomentellous branchlets; petiole 5-8 mm. 
long, issuing from the top of a prominent little knob, the lamina 
broadly elliptic to obovate, truncate to rounded at the base, shortly 
acute or rounded at the apex, finally glabrous on the upper side except 
on the midrib, paler and glaucous and softly hirsute beneath, 7-12 cm. 
long and 3-6 cm. broad; flowers solitary, on pedicels 1-2 cm. long; 
petals 3, thick, rigid, ovate, about 2 cm. long, tomentellous out- 
side; fruit elongate to cylindric, gray- tomentellous, up to 5 cm. 
long and 1.5-2 cm. in diameter, the areoles elongate, ending in an 
appressed cusp. 

San Martin: Near Tarapoto, Ule 6521, type; Williams 5431, 
5829, 6737. San Roque, Williams 7395. "Anonilla." 

6. Annona cordifolia Poepp. ex Szyszyl. Diss. Math. Nat. 
Acad. Litt. Cracov. 27: 140. 1894. Rollinia cordifolia Szyszyl. loc. 
cit. Annona scandens Diels var. polychyla Diels, Notizbl. Bot. 
Gart. Berlin 11:86. 1931. 

A shrub, climbing up to 15 meters, the young branchlets densely 
fulvous-tomentose; petiole 10-13 mm. long, rarely shorter, issuing 
from the top of a prominent little knob, the lamina elliptic or nearly 
orbicular, rounded or shortly acute at the apex, rounded and often 
a little emarginate at the base, finally glabrous above except on the 
nerves, thinly and softly tomentose and glaucous beneath, 15-25 
cm. long and 7-12 cm. broad; pedicels 3-4 cm. long; sepals abruptly 
ending in a long, filiform cusp; petals 3, broadly ovate, about 1.5 cm. 
long; fruit ellipsoid to cylindric, grayish-hirsute, 4-5 cm. long and 
2-3 cm. broad, the areoles prominent, tapering into more or less 
recurved cusps. 

Loreto: Mainas, near Yurimaguas, Poeppig 2090, type. Yuri- 
maguas, Killip & Smith 28715; Williams 3810, 3846, 4197, 4757. 



FLORA OF PERU 755 

7. Annona macrocalyx R. E. Fries, Acta Hort. Berg. 10: 246. 
pi. 17. 1931. 

A small tree, up to 5 meters high, the young branchlets pale 
ferruginous- tomentose; leaves elliptic, rounded or shortly acute at 
the base, shortly and broadly cuspidate at the apex, 10-14 cm. long 
and 4-6.5 cm. broad, at first softly hairy but finally glabrescent 
above, gray- tomentose beneath; flowers solitary, issuing from 
about the middle of the internode below a leaf; pedicels 13-20 mm. 
long, bearing near the base (below the articulation) a lanceolate, 
curved bract 1 cm. long, and higher up a second bract of the same 
length; sepals exceptionally large, broadly ovate, acute, tomentose 
outside, 12-15 mm. long and 10-12 mm. broad; petals 3, rounded- 
ovate, 15-18 mm. long. 

Loreto: Masisea, Killip & Smith 26860, type. 

8. Annona Tessmannii Diels, Notizbl. Bot. Gart. Berlin 9: 
140. 1924. 

A shrub or small tree, the young branchlets shortly appressed- 
sericeous; leaves on petioles 7-10 mm. long, elliptic, usually rounded 
at the base, shortly acute, rarely cuspidate, at first soft-hirsute above 
but finally glabrous and lustrous, paler and covered with short, 
appressed hairs on the lower surface, 8-15 cm. long and 4-8 cm. 
broad; inflorescences generally many-flowered, short-pedunculate; 
pedicels 1-2.5 cm. long, rigid; sepals about 2 mm. long; petals 3, 
thick, rigid, broadly ovate, 1.5-2 cm. long; fruit ovoid, 4 cm. long 
(not fully developed?), with numerous conic protuberances on the 
surface. Neg. 27524. 

Loreto: Near Yurimaguas, Rio Paranapura, Mexia 6070. Rio 
Ucayali, Amaquiria, Tessmann 3367, type. La Victoria, Williams 
2853. Caballo-cocha, Williams 2305, 2418. Pebas, Williams 1842. 
Iquitos, Tessmann 5076. Iquitos, Rancho Indiana, on the left 
bank of Rio Maranon, Mexia 6429. Leticia, Wittiams 3048. Rio 
Putumayo, Klug 1621. Also in western Amazonian Brazil. 

9. Annona iquitensis R. E. Fries, Acta Hort. Berg. 10: 251. 
1931. 

A tree; leaves rigid, chartaceous, oblong or narrowly oblong- 
obovate, rarely elliptical-oblong, rounded at the base and abruptly 
contracted at the apex into a blunt cusp about 1 cm. long, pale 
green, glabrous, and smooth above, a little paler beneath and quite 
glabrous, at least when mature, 10-16 cm. long and 3.5-4.5 cm. 
broad; flowers solitary, terminal, on pedicels about 2 cm. long; 



756 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sepals 6-7 mm. long, rounded-triangular, acuminate; petals 3, 
broadly ovate, 13-15 mm. long. 

Loreto: Iquitos, Killip & Smith 2984-8, type. 

10. Annona squamosa L. Sp. PI. 537. 1753. A. cinerea Dun. 
Monogr. Anon. 71. pi. 8. 1817. Xylopia frutescens Sieb. ex Presl in 
Oken, Isis 21: 273. 1828 (non Aubl.). Guanabanus squamosus Gomez, 
Fl. Haban. 114. 1897. 

A small tree, up to 5 meters high; leaves membranaceous, elliptic 
or lanceolate, shortly and obtusely acute, often glaucous beneath 
and finally quite or nearly glabrous, 5-11 cm. long and 2-5 cm. 
broad; inflorescences 1-few-flowered; pedicels glabrous or slightly 
pubescent, 1-2 cm. long; outer petals linear-oblong, obtuse, often 
narrowed to the base, 1.5-3 cm. long; inner petals rudimentary, 
about 1 mm. long; fruit globose-ovoid, glabrous, more or less glaucous 
at first, 8-9 cm. in diameter, the fruiting carpels loosely cohering, 
rounded at the extremities. 

I have not seen any specimen from Peru, but the species is certainly 
cultivated here and there in the country for its edible, delicious fruit. 

11. Annona reticulata L. Sp. PI. 537. 1753. A. Humboldtii 
Dun. Monogr. Anon. 64. pi. 3. 1817. A. Humboldtiana HBK. 
Nov. Gen. & Sp. 5: 56. 1821 (ubique flore excl.). A. laevis HBK. 
op. cit. 60. A. excelsa HBK. op. cit. 59. A. riparia HBK. op. cit. 
59. A. longifolia Sesse" & Moc. Fl. Mex. 146. 1887. 

A small tree, up to 8 meters high; leaves rigidly membranaceous, 
at first appressed-hirsute, especially on the nerves, finally glabrous, 
lanceolate to oblong-lanceolate, gradually long-acuminate, 10-20 cm. 
long, 2-5 cm. broad; inflorescences issuing from the middle of the 
internodes, rarely opposite the leaves, many-flowered; outer petals 
linear-oblong, obtuse, keeled on the inside, 1.5-2, rarely 2.5 cm. long; 
inner petals very small, scale-like; fruit globose-ovoid, 9-10 cm. in 
diameter, smooth, with the surface divided into rhomboid or hexag- 
onal areoles by impressed lines, reddish or reddish brown when ripe. 

Cajamarca(?) : Rio Huancabamba, near Colasai, Humboldt & 
Bonpland 3583. Prov. Jae"n, Pueblo de Perico, on the shore of Rio 
Chinchipe, Raimondi 2247. Bahia Grande, Raimondi 12996. 
Probably native of the West Indies, now cultivated in many tropical 
countries for its edible fruit. 

12. Annona Cherimolia Mill. Gard. Diet. ed. 8. No. 5. 1768. 
A. tripetala Ait. Hort. Kew. 2: 252. 1789. A. pubescens Salisb. 
Prodr. 380. 1796. 



FLORA OF PERU 757 

A small tree, the young branchlets fulvous-tomentose; leaves 
membranaceous, nearly glabrous above, persistently tomentose 
beneath with soft, patent hairs, elliptic, shortly acute at the apex, 
rounded or cuneate at the base, 8-12 cm. long and 4-6 cm. broad, 
sometimes larger; flowers generally issuing from near the base of 
the branchlets, 1-2; pedicels tomentose, 8-12 mm. long; outer 
petals linear, obtuse, rufous-tomentellous outside, 1.5-2.5 mm. long; 
inner petals rudimentary; fruit more or less ovoid, heart-shaped, 
green. 

Without locality: Pavon; Dombey 603; Raimondi 8614, 8620. 
Lima: Lima, Raimondi 9908. Amazonas: Chachapoyas, Mathews. 
Native of the borders of Ecuador and Peru, where it grows at 
an elevation of 1,400-2,000 meters; often cultivated for its delicate 
fruit. 

13. Annona Duckei Diels, Notizbl. Bot. Gart. Berlin 11: 86. 
1931. 

A tree or shrub, the young branchlets slightly sericeous; leaves 
glabrous above, paler and thinly appressed-hairy beneath, obovate 
or elliptic, acute or rounded at the base and shortly decurrent on the 
(5-7 mm. long) petiole, short-acuminate at the apex, 4-10 cm. long 
and 3.5-5.5 cm. broad; flowers issuing a little below a leaf; pedicels 
slender, slightly sericeous, about 1 cm. long; flower buds depressed- 
globose, about 1 cm. in diameter; sepals almost entirely connate; 
outer petals 7 mm. long, connate about half their length, rounded- 
ovate, thinly and shortly sericeous; inner petals nearly as long as the 
outer; fruit depressed-globose, 3-4 cm. in diameter, the areoles flat 
or slightly umbonate-apiculate. Neg. 22519. 

Loreto: Iquitos, Ducke 7609, type. 

14. Annona neglecta R. E. Fries, Acta Hort. Berg. 12: 278. 
1937. A. axillaris R. & P. in sched. 

A tree or shrub, the slender branchlets rufous-sericeous when 
young; petiole 5-8 mm. long, slender; lamina lanceolate, acute 
at the base, longer and rather gradually attenuate to the obtuse 
apex, 7-12 cm. long and 2-3.5 cm. broad, with scattered short 
white hairs on the upper surface when young, finally quite glabrous, 
paler beneath and shortly appressed-hairy; inflorescences 2-4- 
flowered, extra-axillary; pedicels slender, rufous-sericeous, 8-15 mm. 
long; flower buds depressed-globose, about 5 mm. in diameter; 
sepals 1.5 mm. long, nearly free; outer petals thick, rounded, rufous- 



758 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sericeous on the outside, 4 mm. long, the inner petals about 1 mm. 
long; stamens 0.5-0.8 mm. long. Neg. 14239. 

Huanuco: Vitoc, Ruiz, type. 

15. Annona Ambotay Aubl. Hist. PI. Guian. 1: 616. pi. 249. 
1775. 

A small tree, but also stated to be a robust, climbing shrub, 
the young branchlets densely ferruginous-tomentose; leaves on 
petioles 7-10 mm. long, large, membranaceous, when young hairy 
above and sometimes lustrous, finally glabrous, with the midrib 
and nerves tomentose, glaucous beneath when young, and clothed 
with soft, patent, finally more or less evanescent hairs, obovate, 
elliptic, or oblong-lanceolate, short-acute or sometimes rounded at 
the base, cuspidate at the apex, 15-25 cm. long and 5-12 cm. broad; 
inflorescences few- to many-flowered, sessile, opposite the leaves 
or from near the middle of the internodes, often long-persistent on 
leafless branches; pedicels about 5 mm. long, rarely longer; sepals 
2 mm. long; outer petals oblong, obtuse, outside ferruginous-tomen- 
tellous, 15-25 mm. long, connate at the base for about 5 mm.; 
inner petals half as long, linear- triangular; fruit globular, 3-4 cm. 
in diameter, the areoles not obvious. 

Loreto: Mishuyacu, near Iquitos, Klug 958. Also in Amazonian 
Brazil and French Guiana, whence the type. 

The species is here taken in a wide sense, comprising a number 
of forms which may eventually be found to represent different species 
or varieties when more material is accessible. The Peruvian speci- 
men is characterized by its very glossy leaves, with few hairs on the 
lower surface, except on the nerves. 

Annona peruviana Humb. et Bonpl. ex Dun. (Monogr. Anon. 67. 
1817) is a synonym of A. glabra L. (A. palustris L.). The type was 
collected at Guayaquil in Ecuador (Humboldt & Bonpland 3804), 
and the species has never been recorded from Peru. A. glabra grows 
in humid forests near the sea and in mangrove swamps, and is charac- 
terized by large, ovate-elliptic, quite glabrous leaves, without pits 
on the lower surface, by large flowers with 6 free petals, and globose- 
ovoid, smooth fruit 7-12 cm. long with inconspicuous areoles. 

17. ROLLINIA St. Hil. 

Trees or shrubs. Flowers opposite the leaves or extra-axillary 
from the internodes; pedicels articulate, with bracts at the base and 
usually above the articulation. Sepals small, valvate. Petals 



FLORA OF PERU 759 

valvate, connate at the base, the outer produced into a wing or spur. 
Stamens numerous, the connective expanded and truncate above the 
anther. Carpels several to numerous, united in fruit; ovule solitary, 
basal, erect. Closely related to the genus Annona, but differing by 
the presence of wings or spurs on the petals. A large genus of 
56 species, distributed from Central America and the West Indies 
to southern Brazil and northern Argentina. 
Sepals more or less concave, the midrib thickened and keeled on the 

outside. 
Leaves with appressed hairs beneath. 

Petal wings oblong, of the same breadth throughout or slightly 

widened toward the apex. 

Leaves oblong; petal wings straight or only slightly recurved. 
Young branchlets densely ferruginous- tomentellous; petal 
wings obliquely spreading, 4-6 mm. broad. 

2. R. rigidiflora. 
Young branchlets nearly glabrous, with a few appressed 

hairs; petal wings very divergent, about 3 mm. broad. 

3. R. peruviana. 
Leaves more or less elliptic; petal wings recurved. 

4. R. jucunda. 
Petal wings large, from a narrow base cuneate-obovate. 

5. R. papilionella. 

Leaves with patent, soft hairs beneath 6. R. insignis. 

Sepals thin, flat, not keeled. 

Flowers usually 2 or more; pedicels bracteate above the articula- 
tion. 
Fruit large, up to 10 cm. long (edible) ; petal wings more than 

1 cm. long 1. R. mucosa. 

Fruit small (where known, 1-3 cm. in diameter); petal wings 

less than 1 cm. long. 
Petal wings flat, cuneate-obovate or orbicular from a 

narrow base. 
Inflorescences 1-2-flowered; ovaries glabrous. 

7. R. occidentalis. 

Inflorescences 5-6-flowered; ovaries hairy 8. R. Ulei. 

Petal wings shortly rounded to oblong from a broad base, 

horizontal or curved upward. 

Petal wings ferruginous-tomentellous, curving upward from 
a horizontal base 9. R. curvipetala. 



760 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Wings gray-tomentellous, horizontal, straight. 

10. R. pachyptera. 
Wings narrow, spurlike, not narrowed to the base, deflexed. 

11. R. centrantha. 
Flowers solitary; pedicels naked above the articulation. 

Leaves gradually narrowed to the base; petal wings shortly 
oblong 12. R. uniflora. 

Leaves rounded at the base or shortly acute; wings very shortly 

semicircular 13. R. sphaerantha. 

Insufficiently known species (flowers unknown) . . 14. R. microcarpa. 

1. Rollinia mucosa (Jacq.) Baill. Adansonia 8: 268. 1868. 
Anona mucosa Jacq. Obs. 1: 16. 1764. A. reticulata L. /3 mucosa 
Willd. Sp. PI. 2: 1266. 1800. A. obtusiflora Tuss. Fl. Antill. 1: 191. 
1808. A. biflora R. & P. in sched., sec. G. Don, Gen. Syst. 1: 91. 
1831. R. biflora G. Don, op. cit. 91. R. Sieberi A. DC. Me"m. Soc. 
Phys. Geneve 5: 200. 1832. A. reticulata Sieb. ex A. DC. op. cit. 200. 

A small tree with grayish-ferruginous-sericeous branchlets; leaves 
with soft, patent hairs above, glabrescent except on the midrib and 
thicker nerves, densely appressed-hairy beneath, but finally thinly 
sericeous, elliptic to oblong-elliptic or lanceolate, shortly acute or 
rarely rounded at the base, usually longer and gradually narrowed 
at the apex, 12-25 cm. long and 3-9 cm. broad; inflorescences 1-3- 
or, more rarely, many-flowered; pedicels up to 5-6 cm. long, often 
shorter; corolla 2-3.5 cm. across, tomentellous; wings not thickened, 
spreading and often curved upward a little or horizontal, cuneate- 
obovate or oblong; fruit edible, globose, up to 10 cm. in diameter; 
areoles produced into conic tubercles. 

This species is often cultivated for its edible fruit. I have seen 
Peruvian specimens (all cultivated?) from the following localities: 

Without definite locality: Herb. Pawn (type of Annona biflora 
and Rollinia biflora). Sarayaco, Raimondi 987, 12842. San Antonio 
de Tinga Maria, Raimondi 12838. Loreto: Iquitos, cultivated, 
Ducke (Herb. Rio de Janeiro 24178). San Martin: San Roque, 
abandoned land, Williams 7127. 

la. Rollinia mucosa var. neglecta R. E. Fries, comb. nov. 
R. neglecta R. E. Fries, Acta Hort. Berg. 12: 124. 1934. 

An insufficiently known form, characterized by stiff, straight 
and finally reflexed pedicels, by pale grayish-tomentose flowers, and 
strictly horizontal wings, of the same breadth throughout. 

Loreto: Mishuyacu, near Iquitos, in forest, Klug 333. 



FLORA OF PERU 761 

Ib. Rollinia mucosa subsp. aequatorialis R. E. Fries, op. cit. 
123. 

Leaves very thin, membranaceous, on the fertile branchlets more 
or less obovate, with unusually sharp points, cuneate and decurrent 
at the base; petal wings narrow, spreading horizontally. 

Loreto: Iquitos, Killip & Smith 27231. Also in western Amazo- 
nian Brazil. 

Rollinia orthopetala A. DC. (Mem. Soc. Phys. Geneve 5: 200. 
1832) is closely related to R. mucosa and differs chiefly in the erect 
and incurved petal wings. It is native of British Guiana and the 
Brazilian state Para, and has also been collected by Poeppig, 1830-31, 
in the gardens of the Indian missions at Yurimaguas and Tocache 
(Poeppig D2004). 

2. Rollinia rigidiflora R. E. Fries, Acta Hort. Berg. 12: 154. 
/. 9, a. 1934. 

A tree, about 8 meters high, the young branchlets densely 
ferruginous- tomentellous; leaves on petioles 5-8 mm. long, glabrous 
above, sparsely clothed beneath with minute, appressed hairs, oblong 
to oblong-lanceolate or narrowly ovate, rounded at the base, gradually 
narrowed at the apex, 8-16 cm. long and 3-5 cm. broad; inflorescences 
1-2-flowered, short-pedunculate, the pedicels ferruginous-sericeous, 
1.5-3.5 cm. long; sepals 2-3 mm. long; corolla 18-22 mm. across, 
ferruginous-tomentellous; wings obliquely spreading, thick and rigid, 
oblong and sometimes a little widened toward the rounded apex, 
8-11 mm. long and 4-6 mm. broad. 

Loreto: Lower Rio Huallaga, Yurimaguas, Fortaleza, Williams 
3886, type; 444-2. Yurimaguas, El Recreo, Williams 4163. San 
Martin: Pongo de Cainarachi, Rio Cainarachi, tributary of Rio 
Huallaga, King 2716. 

3. Rollinia peruviana Diels, Verh. Bot. Ver. Brandenb. 47: 
135. 1905. 

A small tree or shrub, 3-9 meters high, the young branchlets 
nearly glabrous; leaves on petioles 5-6 mm. long, sparsely white- 
hairy above when young, soon glabrescent, paler and subglaucous 
beneath, with a few short, appressed hairs, lanceolate or lanceolate- 
oblong, very shortly acute or rounded at the base, gradually narrowed 
at the apex, 8-15 cm. long and 2-3.5 cm. broad; inflorescences 
1-2-flowered, the pedicels up to 3 cm. long; corolla 18-20 mm. 
across, yellowish-gray-tomentellous; wings narrowly oblong, not 



762 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

widened toward the apex, spreading and a little recurved, about 10 
mm. long and 3-3.5 mm. broad. Neg. 14279. 

San Martin: Tarapoto, Ule 6426, type. Rio Acre: Monte 
Alegre, Seringal San Francisco (Peru?), Ule 9370. 

4. Roll in ia jucunda Diels, Notizbl. Bot. Gart. Berlin 10: 177. 
1927. 

A tree about 7 meters high; petioles 1-1.3 cm. long; lamina 
glabrous above, cinnamomeous beneath and thinly clothed with 
short appressed lustrous hairs, elliptic or elongate-elliptic, shortly 
acuminate at the apex, short-acute at the base, 10-17 cm. long and 
4-6 cm. broad; inflorescences generally 2-flowered; pedicels straight, 
rigid, ferruginous-sericeous, 1.5-1.8 cm. long; sepals 2.5-3 mm. long; 
corolla ferruginous-tomentellous, the wings oblong, rounded at the 
apex, recurved from the horizontal base, 10-13 mm. long and 4-7 mm. 
broad. Neg. 14270. 

Loreto: Iquitos, Tessmann 5341, type. 

5. Rollinia papilionella Diels, Notizbl. Bot. Gart. Berlin 10: 
176. 1927. 

A large tree, up to 30 meters high, the young branchlets very 
shortly tomentellous; leaves on petioles 12-18 mm. long, rigid, gla- 
brous and lustrous above, densely sericeous beneath with short, 
appressed, cinnamomeous hairs, ovate-elliptic, short-acuminate at 
the apex, rounded at the base, 10-20 cm. long and 5-8 cm. broad; 
inflorescences up to 5-flowered; pedicels 2.5-3 cm. long; sepals very 
short (1.5 mm. long and 3-4 mm. broad); corolla ferruginous-tomen- 
tellous, the wings thin, flat, obliquely spreading, obovate, broadly 
rounded at the apex, 18-23 mm. long and 12-13 mm. broad. Neg. 
14277. 

Loreto: Mouth of Rio Santiago, upper Maranon, Tessmann 
4664, type. 

6. Rollinia insignis R. E. Fries, Acta Hort. Berg. 12: 161. 
pi. 17. 1934. 

A tree, up to 15 meters high, the young branchlets densely 
ferruginous- tomentose; leaves rigid, on petioles 12-15 mm. long, 
glabrous above except on the midrib and larger nerves, glaucous 
beneath and covered with patent, crisp, rufous hairs, broadly elliptic 
to elliptic-obovate, rather abruptly cuspidate, rounded at the base, 
15-25 cm. long and 8-12 cm. broad; inflorescences about 5-flowered; 
pedicels 1.5-2.5 cm. long, thick, rigid; sepals about 3-4 mm. long 



FLORA OF PERU 763 

and 5 mm. broad; corolla ferruginous-tomentellous, the wings spread- 
ing, not thickened, cuneate-obovate, with rounded apex, 17-18 mm. 
long and about 8-10 mm. broad when fully developed. 

Loreto: La Victoria, on the Amazon, in forest, Williams 2685, 
type. Also in Amazonian Brazil. 

7. Rollinia occidentalis R. E. Fries, Acta Hort. Berg. 12: 165. 
pi. 18. 1934. 

A shrub or small tree, up to 8 meters high, the young branchlets 
sericeous; leaves on petioles 6-10 mm. long, membranaceous, quite 
glabrous above even when young, glaucous beneath and clothed 
with golden ferruginous, appressed hairs, soon more or less glabres- 
cent, lanceolate or elliptic, acute at the base and gradually narrowed 
into an obtuse apex, 5-10 cm. long and 1.5-4 cm. broad; inflorescences 
1-2-flowered, the pedicels 5-6 mm. long, in fruiting stage reaching 
8-10 mm.; sepals appressed; corolla about 15 mm. across; wings 
thin, membranaceous, minutely gray-tomentellous, obovate-orbicu- 
lar, narrowed at the base, 7-9 mm. long and 5-8 mm. broad; fruit 
globose, smooth and glabrous, 1.5 cm. in diameter. 

Peru : Santa Ana, in woods, Pearce. Also recorded from northern 
Argentina. 

8. Rollinia Ulei Diels, Verh. Bot. Ver. Brandenb. 47: 136. 
1905. 

A shrub about 4 meters high, the branchlets soon glabrescent; 
petioles slender and relatively long (1-1.5 cm.); lamina lustrous on 
both sides, quite glabrous above, very thinly appressed-hairy beneath 
or nearly glabrous, lanceolate to lanceolate-oblong or narrowly 
elliptic, acute and decurrent at the base and gradually narrowed 
at the apex into an obtuse point, 10-15 cm. long and 4-5 cm. broad; 
inflorescences with many flowers (5-6 or perhaps more); pedicels 
slender, at first sericeous, soon glabrate, 2-3.5 cm. long; sepals 
finally recurved; corolla 1.5-2 mm. across, tomentellous; wings spread- 
ing obliquely, from a narrow base obovate-rounded, 8-10 mm. long 
and 5-7 mm. broad. Neg. 27281. 

San Martin : Tarapoto, Juan Guerra, Ule 6425, type. Tarapoto, 
Wittiams 5710, 5768, 6286. Cuzco: Gay 902. 

9. Rollinia curvipetala R. E. Fries, Acta Hort. Berg. 12: 178. 
f. 11, a-c. 1934. 

A small tree, the young branchlets covered with erect-patent, 
ferruginous hairs; leaves glabrous above except on the midrib and 



764 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

thicker nerves, when young densely, at maturity sparsely, sericeous 
beneath, oblanceolate or oblong-lanceolate, acute at the base, rather 
gradually narrowing into an obtuse apex, 7-11 cm. long and 2.5-3.5 
cm. broad; inflorescences 1-2-flowered; pedicels rufous-strigose, 
8-10 mm. long; sepals appressed, 2-2.5 mm. long; corolla up to 18 
mm. in diameter, ferruginous- tomentellous; wings curved a little up- 
ward from a horizontally spreading base, oblong and obtuse, 8-9 mm. 
long, 3-4 mm. broad, 1 mm. thick. 

Rio Acre: Seringal San Francisco, Ule 9371, type. 

10. Rollinia pachyptera Diels, Notizbl. Bot. Gart. Berlin 10: 
176. 1927. 

A tree about 10 meters high; young branchlets and petioles 
appressed-sericeous-ferruginous; leaves membranaceous, glabrous 
above from the beginning, except for persistent hairs on the impressed 
midrib, sparsely appressed-hairy beneath when young, later glabrous, 
oblong, acute and decurrent at the base, rather abruptly contracted 
at the apex into a cusp 1.5 cm. long, 7-14 cm. long and 2.5-4 cm. 
broad; inflorescences 2-flowered (always?), opposite the leaves; 
pedicels thick, 8-9 mm. long; corolla pale yellow when fresh, fragrant, 
gray- tomentellous when dried; wings horizontally spreading, oblong 
and obtuse at the apex, thick, 7 mm. long. Neg. 14276. 

Loreto: Mouth of Rio Santiago, Tessmann 4704, type. 

11. Rollinia centrantha R. E. Fries, Acta Hort. Berg. 12: 181. 
/. 13, c-f. 1934. 

A tree 6-7 meters high; young branchlets, petioles, and pedicels 
rufous-sericeous; leaves 10-16 cm. long, 5-8 cm. broad, on petioles 
5-10 mm. long, sparsely white-hirsute above, with the thicker nerves 
more densely hairy, probably glabrescent when mature, thinly 
appressed-sericeous beneath, obovate to elliptic or broadly rhombic- 
elliptic, acute at the base, rather abruptly narrowed at the apex 
into a cusp 1.5-2.5 cm. long; inflorescences sessile, 1-2-flowered, 
issuing from the middle of the internode above the axil; pedicels 
very slender, 12-15 mm. long, bracteolate at the middle or below; 
corolla 10-12 mm. across, deep purple when fresh, rufous-sericeous; 
wings linear, up to 5 mm. long and 1.2-1.5 mm. broad, deflexed or 
recurved; fruit globose, 15-18 mm. in diameter, the areoles about 15, 
a little convex and terminating in a short point. 

Loreto: Iquitos, Killip & Smith 27073, type; Williams 8043. 



FLORA OF PERU 765 

12. Rollinia uniflora R. E. Fries, Acta Hort. Berg. 12: 182. 
/. 12, c-d. 1934. 

A small tree with slender, ferruginous-sericeous and soon gla- 
brescent branchlets; leaves on petioles 7-10 mm. long, membrana- 
ceous, glabrous above with scattered hairs more or less persistent 
on the midrib and thicker nerves, paler and nearly glabrous beneath 
with a few appressed hairs, cuneate and long-decurrent at the base, 
abruptly cuspidate at the apex; flowers solitary from the middle of 
the internode, supra-axillary; pedicels very slender, 15-18 mm. long, 
articulate above the basal bract and naked above the articulation; 
sepals reflexed, 2 mm. long; corolla 12-13 mm. across, gray-tomen- 
tellous; wings rounded-oblong, horizontal, laterally compressed, about 
6 mm. long and 5 mm. broad. 

San Martin: Tarapoto, in forest, Spruce, type. 

13. Rollinia sphaerantha R. E. Fries, Acta Hort. Berg. 12: 
184. /. 13, a-b. 1934. 

A tree 3-6 meters high; young branchlets appressed-hirsute, 
ferruginous; leaves on petioles 4-7 mm. long, membranaceous, 
finally glabrate on both sides with a few hairs on the thicker nerves, 
obovate to nearly orbicular, abruptly cuspidate at the apex and 
short-acute at the base, 7-15 cm. long and 4-8 cm. broad; flowers 
solitary, issuing from about the middle of the internode above the 
axil; pedicels slender, 1-2 cm. long, bracteolate at the base; sepals 
rounded, recurved, 2 mm. long; corolla about 12 mm. across, 
ferruginous-sericeous; petals thick, fleshy; wings short, hemispheric; 
ovaries densely hairy. 

Loreto: Yurimaguas, in dense forest, Killip & Smith 27650, type. 

14. Rollinia microcarpa (R. & P.) R. E. Fries, Acta Hort. 
Berg. 12: 187. pi. 20. 1934. Annona microcarpa R. & P. Fl. Peruv. 
5: pi. 487, ined. sec. G. Don, Gen. Syst. 1: 90. 1831. A. pteropetala 
R. & P. in sched.(?). A. pterocarpa R. & P. op. cit. pi. 483 sec. G. 
Don, op. cit. 91(7). R. pterocarpa G. Don, op. cit. 91(7). 

Young branchlets glabrate; leaves 10-20 cm. long and 5-10.5 cm. 
broad, on petioles 10-14 mm. long, glabrous above when mature 
except on the impressed midrib, very sparsely provided with minute, 
appressed hairs beneath, densely and minutely verruculose-punctate, 
elliptic-obovate, rounded or very shortly acute at the base, cuspidate, 
the cusp 0.5-1 cm. long, the veins prominent, especially beneath, 
forming a very dense network; fruit on pedicels 3-5 cm. long, 
elongate-ovate, 2.2-3 cm. long and 1.8-2 cm. broad, covered with 



766 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

numerous hemispheric and obtuse tubercles 2-3 mm. broad; seeds 
oblong-ellipsoid, fuscous, 6 mm. long and 2.5-3 mm. broad. Neg. 
14275. 

Peru: "In Andium montibus," Ruiz, type. 

74. MYRISTICACEAE. Nutmeg Family 
By Albert C. Smith, New York Botanical Garden 

References: Warburg, Monographic der Myristicaceen, Nov. 
Act. Acad. Leop.-Carol. 68: 1-680. pi. 1-25. 1897; A. C. Smith, The 
American species of Myristicaceae, Brittonia 2: 393-510. 1938. 

Trees or shrubs, often aromatic; leaves alternate, simple, entire, 
pinnate-nerved, estipulate; flowers unisexual, usually dioecious, usu- 
ally fascicled, apetalous; perianth small, infundibuliform to rotate, 
3 (rarely 2-5) -lobed, the lobes valvate; stamens (in the Peruvian 
species) 3-10 (rarely 2), the filaments connate into a column, the 
anthers 2-celled, dehiscing longitudinally, free to the base or dorsally 
connate or dorsally adnate to a carnose connective mass; ovary supe- 
rior, sessile, 1-celled, the stigma subsessile, the ovule 1, essentially 
basal; fruit fleshy, 2-valved, the seed erect, covered by an often 
colored aril; endosperm copious; embryo small. A family of tropical 
distribution, the genera well marked and with sharp geographic lim- 
its. Five genera and 73 species in America. 

Tertiary nerves conspicuously subparallel, nearly perpendicular to 
the costa; seed irregularly black- or purple-splotched; anthers 

4-10, free, recurved; pedicels ebracteolate 1. Compsoneura. 

Tertiary nerves not as above; seed uniform in color. 

Hairs of young branchlets and petioles (and sometimes of lower 
surfaces of young leaves) 2-branched, or at least with only 1 
or 2 conspicuous branches. 

Leaf vernation conduplicate; pedicels ebracteolate; anthers free 
(in Peruvian species); fruit usually ellipsoid, the aril con- 
spicuously laciniate 2. Dialyanthera. 

Leaf vernation convolute; pedicels bracteolate at the summit; 
anthers usually dorsally connate, rarely free; fruit usually 
transversely ellipsoid, the aril short-laciniate or nearly en- 
tire 3. Iryanthera. 

Hairs of young branchlets and petioles (and usually of lower 

surfaces of leaves) stellate or many-branched. 
Leaf blades rounded at the apex; pedicels bracteolate at the 
summit; anthers 12 or 14; fruit transversely ellipsoid, the 
aril obscurely laciniate 4. Osteophloem. 



FLORA OF PERU 767 

Leaf blades usually somewhat acute at the apex; pedicels ebrac- 
teolate; anthers usually 3 (2-6); fruit ellipsoid or subglo- 
bose, the aril deeply laciniate 5. Virola. 

1. COMPSONEURA Warb. 

Dioecious shrubs or small trees; leaves glabrous, petiolate, the 
blades entire and often narrowly recurved at the margins, the nerves 
conspicuous, prominulous on both surfaces, the tertiary nerves sub- 
parallel, nearly perpendicular to the costa; inflorescences (in Peruvian 
species) fasciculate-racemose or narrowly paniculate; bracteoles 
none; flowers pedicellate; perianth carnose or thin-carnose, 3 (rarely 
4 or 5) -lobed; filaments connate into a carnose column; anthers 4-8 
(rarely 10), oblong, basifixed and essentially free in Peruvian species, 
often recurved; ovary subglobose or ellipsoid, the style short, the 
stigma 2-cleft; fruit ellipsoid, glabrous, 2-valved, smooth or obscurely 
carinate, the pericarp very thin, the aril essentially entire or minutely 
laciniate at the apex, the seed ellipsoid, the testa conspicuously and 
irregularly splotched with black or purple. 

Inflorescences essentially glabrous; anthers 2-4 times longer than 
the filament column; leaf blades chartaceous or thin-coriaceous, 
often translucent C. Sprucei. 

Inflorescences minutely strigose, at length glabrescent; anthers sub- 
equal to the filament column (rarely up to twice as long); leaf 
blades coriaceous, opaque C. capitellata. 

Compsoneura capitellata (A. DC.) Warb. Nov. Act. Acad. 
Leop.-Carol. 68: 146. pi. 1, f. 1-5. 1897. Myristica capitellata A. 
DC. in DC. Prodr. 14: 697. 1857. C. Tessmannii Markgraf, Notizbl. 
Bot. Gart. Berlin 9: 964. 1926. 

A tree up to 10 meters high; petioles 7-12 (-20) mm. long; leaf 
blades coriaceous, oblong-elliptic, 10-20 (-35) cm. long, 2.5-9 (-12) 
cm. broad, acute to attenuate at the base, obtusely cuspidate or 
short-acuminate at the apex, the secondary nerves 5-12 per side; 
staminate inflorescences minutely strigose on all exterior surfaces, 
including the perianths, at length glabrous, at anthesis 4-15 cm. long; 
flowers in irregular fascicles of 5-25, the pedicels slender, up to 2 
mm. long; perianth often glandular-pellucid-punctate, 1.8-3 mm. 
long, lobed about one-half its length or slightly more; anthers usually 
subequal to the filament column or slightly longer; pistillate inflores- 
cences often more compact than the staminate, the flowers some- 
times 2 or 3 in sessile fascicles, the ovary ellipsoid, minutely strigose. 



768 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Pampayacu, 1,000 meters, Macbride 5065. Junin: 
Pichis Trail, 1,100 meters, Killip & Smith 26053. Loreto: Rio 
Santiago, Tessmann 4108 (type of C. Tessmannii). Vicinity of 
Iquitos, Tessmann 5101; Williams 3786; King 180. Rio Putu- 
mayo, King 2130. San Martin(?): Rio Huallaga, Poeppig 1971 
(type, err. cit. 1791). Amazonian Brazil. "Senimoro-ey" (Huitoto 
Indian name). 

Compsoneura Sprucei (A. DC.) Warb. Nov. Act. Acad. Leop.- 
Carol. 68: 143. pi. 3, f. 1, 2. 1897. Myristica Sprucei A. DC. in 
DC. Prodr. 14: 199. 1856. 

A shrub or small tree, up to 13 meters high; petioles 8-25 mm. 
long; leaf blades chartaceous or thin-coriaceous, elliptic or oblong- 
elliptic, 9-25 (-32) cm. long, 3.5-10 (-15) cm. broad, acute to atten- 
uate at the base, obtusely acuminate or cuspidate at the apex, the 
secondary nerves 4-9 (-12) per side; staminate inflorescences gla- 
brous throughout, 2-8 cm. long; flowers in irregular fascicles of 
3-15, the pedicels slender, up to 2 mm. long; perianth thin-carnose, 
often minutely yellow-glandular, 1.5-3 mm. long, lobed about one- 
half its total length; anthers usually free but sometimes dorsally 
connate up to the middle, usually 2-4 times longer than the filament 
column; pistillate flowers up to 8 per fascicle, the ovary minutely 
glandular; mature fruits 1-4 per inflorescence, 20-37 mm. long, 13-21 
mm. broad, stipitate at the base, obtuse or rounded at the apex, the 
pericarp brittle, 0.3 mm. thick or less, the aril usually red, waxy. 

Loreto: Rio Huallaga, King 2946, 2842; Williams 4202. Rio 
Ucayali, Tessmann 3096. San Martin: Rio Huallaga, King 3982. 
Southern Mexico, Guatemala, and British Honduras to Amazonian 
Brazil and Peru. 

2. DIALYANTHERA Warb. 

Dioecious trees, the inner bark often exuding a watery, reddish 
liquid, the branchlets and petioles usually strigose when young; 
leaves petiolate (petioles winged), the blades chartaceous or thin- 
coriaceous, entire and often slightly thickened at the margins, gla- 
brous above, usually minutely strigose beneath when young, the 
secondary nerves obscurely anastomosing near the margins, the 
veinlets obscure; inflorescences axillary, fasciculate-racemose (in 
Peruvian species); bracteoles none; flowers pedicellate; perianth 
thin-carnose, 3 (rarely 4) -lobed; filaments connate into a carnose 
column; anthers 3, basifixed, free and divergent (in Peruvian species) ; 
ovary subglobose or ellipsoid, the style short, the stigma inconspicu- 



FLORA OF PERU 769 

ously cleft; fruit ellipsoid, glabrous at maturity, 2-valved, the peri- 
carp woody, the aril laciniate more than one-half its total length, 
the seed ellipsoid or subglobose. 

Dialyanthera parvifolia Markgraf, Notizbl. Bot. Gart. Berlin 
9: 964. 1926. Myristica Otoba var. glaucescens A. DC. in Mart. Fl. 
Bras. 5, pt. 1: 122. 1860. 

A tree 5-15 meters high; petioles 1-4 mm. broad, including the 
wings, 10-30 mm. long; leaf blades often minutely papillose or rugose 
on both surfaces, elliptic, 8-20 cm. long, 3.5-8 cm. broad, attenuate 
at the base, obtusely short-acuminate or cuspidate at the apex, the 
secondary nerves 8-14 per side; staminate inflorescences straight, 
slender, simple, strigose on all exterior surfaces, 4-16 cm. long at 
anthesis; fascicles 12-29 per inflorescence, each subtended by a semi- 
orbicular bract 1-1.5 mm. long; flowers 2-13 per fascicle, the pedicels 
slender, 2-4 mm. long; perianth 3-4 mm. long, 3-lobed more than 
one-half its total length; filament column conspicuously swollen at 
the base, the anthers 0.5-0.7 mm. long, about twice as long as the 
filament column; pistillate inflorescences 2-4 cm. long, the flowers 
solitary or paired on the rachis; mature fruits 2 or 3 per inflorescence, 
smooth or faintly carinate, 25-27 mm. long, 20-23 mm. broad, 
short-stipitate at the base, often cuspidate at the apex, the pericarp 
1-2 mm. thick. 

Junin: La Merced, 1,000 meters, Weberbauer 1902. Loreto: 
Rio Santiago, Tessmann 4075. Rio Maranon, Tessmann 3872, 
type. Rio Huallaga Valley, Poeppig 2348B, type of M. Otoba var. 
glaucescens; Killip & Smith 29030, 29272; King 3951. San Martin: 
Rio Huallaga Valley, King 2663. Amazonian Brazil. 

This species has been confused in herbaria and in literature with 
D. Otoba (Humb. & Bonpl.) Warb., which occurs from Costa Rica 
to Colombia. The two are quite distinct in foliage and very differ- 
ent in inflorescence characters. 

3. IRYANTHERA Warb. 

Monoecious (or perhaps sometimes dioecious) shrubs or trees, 
the inner bark frequently exuding a reddish liquid, the branchlets 
ferruginous-strigose when young; leaves petiolate, the blades gla- 
brous, often finely rugose or minutely papillose, fragile when dry, the 
secondary nerves anastomosing toward the margins; inflorescences 
1-3 in the leaf axils or on defoliate branchlets (pistillate often on old 
bark of trunk or branches), minutely strigose on all exterior surfaces, 
fasciculate-racemose or narrowly paniculate; pedicels bracteolate 



770 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

at the summit; perianth usually thin-carnose and strigose without, 

3 (rarely 4) -lobed; filaments connate into a column, the anthers 3 

(rarely 4), connate to the apex or distally divergent or free to the 

base; ovary usually ellipsoid, glabrous, the style short; fruit usually 

transversely ellipsoid, sometimes essentially globose, coriaceous, 

2-valved, the pericarp usually woody, the aril inconspicuously lacini- 

ate distally, the seed transversely ellipsoid or subglobose. "Cumala." 

Perianth campanulate, the lobes less than one-half the total length; 

androecium 1.6-2 mm. long or more; anthers much shorter than 

the filament column, sometimes free. 

Leaf blades thick-coriaceous (17-) 20-45 cm. long; staminate 

inflorescences elongate. 

Leaf blades subcordate to obtuse at the base; staminate inflores- 
cences 2-11 cm. long, the flowers in essentially sessile fasci- 
cles; perianth 2-3 mm. long /. macrophylla. 

Leaf blades acute to cuneate at the base; staminate inflores- 
cences 15-23 cm. long, the flowers borne on short, lateral 

branches; perianth about 4 mm. long I. crassifolia. 

Leaf blades chartaceous or thin-coriaceous, 8-22 cm. long. 

Secondary nerves obvious; leaf apex gradually acuminate; 
branchlets not lenticellate; anthers with inconspicuous 

connectives /. Tessmannii. 

Secondary nerves often immersed, inconspicuous or plane on 
both surfaces; leaf apex obtuse or obtusely short-acuminate; 
branchlets often lenticellate; anthers with conspicuous, 

carnose, fused connectives /. laevis. 

Perianth cupuliform or rotate, the lobes often more than one-half 
the total length; androecium up to 1.6 mm. long; anthers equal- 
ing the filament column or somewhat shorter, connate (rarely 
free at the apex). 
Androecium 0.8-1.6 mm. long; perianth cleft about one-half the 

total length. 

Intramarginal anastomoses not conspicuous, often irregular 
and incomplete; fruit transversely ellipsoid or subglobose, 
inconspicuously carinate, the lateral extremities rounded. 

/. juruemis. 
Intramarginal anastomoses complete and conspicuous; fruit 

transversely ellipsoid. 

Leaf blades minutely papillose or nearly smooth on both 
surfaces; mature fruit up to 11 mm. long and 18 mm. 
broad, conspicuously carinate /. Ulei. 



FLORA OF PERU 771 

Leaf blades very thick, conspicuously papillose on both sur- 
faces; mature fruit 13-17 mm. long, 20-25 mm. broad, 

faintly carinate /. longiflora. 

Androecium 0.5-0.7 mm. long; perianth cleft nearly to the base, 

the lobes at maturity spreading, flattened. 
Flower surfaces densely short-pilose; fruit 25-40 mm. in diame- 
ter, the pericarp 4-5 mm. thick; intramarginal anastomoses 

not conspicuous /. lancifolia. 

Flowers sparsely pilose; fruit less than 13 mm. long and 24 mm. 
broad, the pericarp 0.5-1.5 mm. thick; intramarginal 
anastomoses conspicuous I. paraensis. 

Iryanthera crassifolia A. C. Smith, Brittonia 2: 431. 1938. 

A tree up to 17 meters high; petioles stout, 13-22 mm. long; leaf 
blades thick-coriaceous, oblong or elliptic-oblong, 23-45 cm. long, 
6-13 cm. broad, acute or cuneate at the base, obtusely cuspidate 
or short-acuminate at the apex, the costa prominent, the secondary 
nerves 17-20 per side; staminate inflorescences 15-23 cm. long, nar- 
rowly paniculate, the secondary branches 14-23 per inflorescence, 
up to 5 mm. long; flowers 7-15 per fascicle, the pedicels slender, up 
to 8 mm. long; bracteole cleft to the base, about 1.5 mm. long; peri- 
anth campanulate, flaring distally, about 4 mm. long, 3-lobed about 
one-third its length; filament column 2-2.5 mm. long, the anthers 
about 0.5 mm. long, free to the base; fruiting inflorescences up to 
10 cm. long, the fruits few, slightly transversely ellipsoid, incon- 
spicuously carinate, up to 26 mm. long and 33 mm. broad, the 
pericarp 3-7 mm. thick. 

Loreto: Lower Rio Huallaga, Williams 4817. Rio Amazonas, 
Pebas, Williams 1744, type. Amazonian Brazil. 

Iryanthera juruensis Warb. Verh. Bot. Ver. Brandenb. 47: 
137. 1905. I. trigona Markgraf, Notizbl. Bot. Gart. Berlin 10: 237. 
1928. 

A tree up to 20 meters high; petioles 6-15 mm. long; leaf blades 
chartaceous or thin-coriaceous, usually dull on both surfaces, elliptic- 
obovate or obovate-oblong, 10-20 cm. long, 3-7 cm. broad, acute to 
subattenuate at the base, obtusely short-acuminate, the secondary 
nerves 10-17 per side; staminate inflorescences 2-9 cm. long, the 
fascicles 5-18 per inflorescence, the flowers 3-7 per fascicle, the pedi- 
cels slender, up to 6 mm. long; bracteole one-sided, less than 1 mm. 
long; perianth cupuliform, 2-3.5 (-4) mm. long, 3-lobed more than 
one-half its length; filament column 0.6-1.1 mm. long, usually con- 



772 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

spicuously swollen at the base; anthers 0.3-0.5 mm. long, dorsally 
connate or divergent in the distal half; pistillate inflorescences usu- 
ally aggregated on old bark, 3-6-branched from the base, the flowers 
15^10 near the swollen ends of the branches, the ovary subglobose 
or ellipsoid; mature fruits transversely ellipsoid or subglobose, 
13-20 mm. long, 15-24 mm. broad, inconspicuously carinate, the 
pericarp 1-2 mm. thick. 

Loreto: Rio Huallaga basin, Killip & Smith 28817; Williams 
4823; King 3058. Vicinity of Iquitos, Tessmann 5386, type of I. 
trigona. Rio Itaya, Tessmann 5158; Williams 34-61. Amazonian 
Brazil. 

Iryanthera laevis Markgraf, Notizbl. Bot. Gart. Berlin 9: 
965. 1926. 

A tree up to 25 meters high; branchlets often lenticellate; petioles 
7-15 mm. long, often narrowly winged; leaf blades chartaceous or 
thin-coriaceous, usually shining above, oblong or elliptic-oblong or 
obovate-oblong, 8-17 cm. long, 3-7 cm. broad, obtuse to attenuate 
at the base, obtuse or obtusely short-acuminate at the apex, the 
secondary nerves 8-14 per side; staminate inflorescences 4-10 cm. 
long, the fascicles numerous; flowers 2-9 per fascicle, the pedicels 
slender, up to 12 mm. long; bracteole cupuliform or cleft to the base, 
0.5-1 mm. long; perianth campanulate, flaring distally, 3.5-4.5 
mm. long, 3-lobed about one-third its length; anthers 0.4-0.5 mm. 
long, about 5 times longer than the filament column; pistillate 
flowers resembling the staminate, the ovary ovoid-conic; mature 
fruits 5-10 per inflorescence, transversely ellipsoid, distinctly cari- 
nate, 15-23 mm. long, 18-30 mm. broad, the pericarp 2-5 mm. thick. 

Loreto: Rio Santiago, Tessmann 4139, type. Lower Rio Hua- 
llaga, Killip & Smith 28779. Rio Itaya, Tessmann 5172. Amazo- 
nian Brazil. 

Iryanthera lancifolia Ducke, Journ. Wash. Acad. Sci. 26: 217. 
1936. 

A tree up to 25 meters high; petioles nigrescent, 10-20 mm. long; 
leaf blades coriaceous or thin-coriaceous, narrowly oblong or elliptic- 
oblong, 13-28 cm. long, 4-8 cm. broad, obtuse to acute at the base, 
obtusely acuminate at the apex, the secondary nerves 12-18 per 
side; staminate inflorescences 3-20 cm. long, the fascicles numerous, 
the flowers 5-12 per fascicle, the pedicels slender, up to 6 mm. long; 
bracteole one-sided, 0.5-1 mm. long; perianth cupuliform or subro- 
tate, 1.5-1.8 mm. long, 3-lobed nearly to the base; androecium 0.5- 



FLORA OF PERU 773 

0.7 mm. long, the filament column carnose, the anthers about 0.25 
mm. long; mature fruits slightly transversely ellipsoid or nearly sub- 
globose, 25-34 mm. long, 27-40 mm. broad, inconspicuously carinate, 
the pericarp very rugose, 4-5 mm. thick. 

Loreto: Rio Putumayo, at mouth of Rio Zubineta, Klug 2234. 
Amazonian Brazil. 

Iryanthera longiflora Ducke, Journ. Wash. Acad. Sci. 26: 217. 
1936. 

A tree up to 20 meters high or more; petioles stout, 10-20 mm. 
long, often nigrescent; leaf blades coriaceous, elliptic-oblong, 15-30 
cm. long, 4-10 cm. broad, acute or obtuse at the base, obtusely 
short-acuminate at the apex, the secondary nerves 13-20 per side, 
regularly anastomosing near the margins; staminate inflorescences 
3-8 cm. long, the fascicles 7-12 per inflorescence, the flowers 4-10 
per fascicle, the pedicels slender, up to 5 mm. long; bracteole one- 
sided, 0.5-1.5 mm. long; perianth cupuliform, 2-3 mm. long, 3-lobed 
about one-half its length; anthers 0.4-0.5 mm. long, about twice 
as long as the filament column; mature fruits transversely ellipsoid, 
13-17 mm. long, 20-25 mm. broad, faintly carinate, the pericarp 
less than 1 mm. thick. 

Loreto: Mouth of Rio Santiago, Tessmann 4105, 4605. Amazo- 
nian Brazil. 

Iryanthera macrophylla (Benth.) Warb. Nov. Act. Acad. 
Leop. -Carol. 68: 155. 1897. Myristica macrophylla Spruce in Benth. 
Journ. Bot. & Kew Misc. 5: 6. 1853. 

A tree up to 17 meters high; petioles narrowly winged distally, 
robust, 10-20 mm. long; leaf blades thick-coriaceous, oblong or 
obovate-oblong, 17-35 cm. long, 5-12 cm. broad (occasionally up 
to 40 by 14 cm.), subcordate or rounded or obtuse at the base, 
obtusely cuspidate or short-acuminate at the apex, the costa very 
prominent, the secondary nerves 14-20 per side; staminate inflores- 
cences 3-7 (rarely 2-11) cm. long, the rachis stout, swollen at the 
fascicles; fascicles 6-13 per inflorescence, the flowers 4-12 per fas- 
cicle, the pedicels slender, up to 5 mm. long; bracteole cupuliform or 
cleft to the base, 0.7-1.5 mm. long; perianth 2-3 mm. long, flaring 
distally, 3-lobed about one-third its length; filament column 1.5-2.2 
mm. long, the anthers about 0.5 mm. long, free to the base; mature 
fruits usually 3 or 4 per inflorescence, transversely ellipsoid, carinate, 
8-24 mm. long, 10-26 mm. broad, the pericarp 1.5-6 mm. thick. 



774 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Rio Huallaga basin, 220 meters, Klug 2906, 3045. Ama- 
zonian Brazil and British Guiana. 

Iryanthera paraensis Huber, Bol. Mus. Goeldi 5: 358. 1909. 
/. aeasilis Markgraf, Notizbl. Bot. Gart. Berlin 10: 236. 1928. 

A tree up to 15 meters high; petioles 7-17 mm. long; leaf blades 
chartaceous or thin-coriaceous, oblong or narrowly elliptic-oblong, 
15-40 cm. long, 4.5-11 cm. broad, obtuse to cuneate or attenuate 
at the base, gradually narrowed to an acuminate apex, the secondary 
nerves 17-27 per side, regularly anastomosing near the margins; 
staminate inflorescences 5-22 cm. long, slender, the fascicles numer- 
ous, the flowers 7-20 per fascicle, the pedicels slender, up to 3 mm. 
long; bracteole ovate or minutely peltate, usually less than 0.3 mm. 
long; perianth cupuliform or subrotate, 0.8-1.6 mm. long, lobed 
nearly to the base; androecium 0.5-0.7 mm. long, the anthers 0.2- 
0.3 mm. long; pistillate inflorescences 1-3.5 cm. long, the flowers in 
clusters of 10-20 at the swollen ends of the branches, the ovary sub- 
cylindric; mature fruits transversely ellipsoid, 11-13 mm. long, 
18-24 mm. broad, distinctly carinate, the pericarp 0.5-1.5 mm. 
thick. 

Loreto: Vicinity of Iquitos, Tessmann 5332, type of /. sessilis; 
Klug 91 . Rio Amazonas, La Victoria, Williams 2869. Guiana and 
Amazonian Brazil. 

Iryanthera Tessmannii Markgraf, Notizbl. Bot. Gart. Berlin 
10:236.1928. 

A shrub or low tree, up to 6 meters high; petioles 6-10 mm. long; 
leaf blades elliptic-oblong, 10-22 cm. long, 3-6 cm. broad, acute to 
attenuate at the base, obtusely acuminate at the apex, the secondary 
nerves 9-16 per side; staminate inflorescences 1-4 cm. long, slender, 
the fascicles 3-5 per inflorescence, the flowers 2-5 per fascicle, the 
pedicels slender, up to 5 mm. long; bracteole cupuliform or cleft 
nearly to the base, 0.3-1 mm. long; perianth campanulate, 2.5-3.5 
mm. long, flaring at the apex, 3-lobed about one-third its length; 
filament column 1.8-2.5 mm. long, the anthers 0.3-0.6 mm. long, 
loosely connate to the apex or free to the base; pistillate inflorescences 
less than 1 cm. long, the flowers few; mature fruits usually 2 or 3 
per inflorescence, transversely ellipsoid, carinate, 9-14 mm. long, 
12-20 mm. broad, conspicuously stipitate, the pericarp 0.5-1.5 mm. 
thick. 

Loreto: Lower Rio Huallaga basin, Williams 4815, 4824; Killip 
& Smith 27678, 27704. Rio Maranon, Killip & Smith 27516. Rio 



FLORA OF PERU 775 

Ucayali basin, Tessmann 3396, type. Rio Amazonas region, Williams 
58, 2403, 2535, 2677, 2727, 2815, 2820, 2902, 2952. Amazonian Brazil. 

Iryanthera Ulei Warb. Verb. Bot. Ver. Brandenb. 47: 137. 
1905. I. leptoclada Markgraf, Notizbl. Bot. Gart. Berlin 9: 965. 
1926. /. congestiflora Macbride, Candollea 5: 350. 1934. 

A tree up to 30 meters high; petioles 6-12 mm. long; leaf blades 
coriaceous or thin-coriaceous, oblong-elliptic, 15-30 cm. long, 4-10 
cm. 'broad, obtuse to acute at the base, obtusely cuspidate or acu- 
minate at the apex, the secondary nerves 12-20 per side, conspicu- 
ously and regularly anastomosing near the margins; staminate 
inflorescences 0.5-9 cm. long, the fascicles 3-17 per inflorescence, 
the flowers 5-10 per fascicle, the pedicels slender, up to 5 mm. long; 
bracteole one-sided or subpeltate, 0.5-1 mm. long; perianth cupuli- 
form, 1.5-3 mm. long, 3-lobed about one-half its length or slightly 
more; anthers 0.2-0.6 mm. long, 3-4 times as long as the filament 
column, dorsally connate or- divergent at the apex; pistillate inflores- 
cences often aggregated on old bark, sometimes axillary and much 
contracted, up to 8 cm. long, occasionally less than 1 cm. long, 
the flowers in clusters of 10-40, the ovary cylindric-conic; mature 
fruits transversely ellipsoid, 8-11 mm. long, 13-18 mm. broad, 
conspicuously carinate, the pericarp 0.5-1.5 mm. thick, the seed 
much broader than long. 

Loreto: Rio Santiago, Tessmann 4263. Rio Morona, Tessmann 
3832, type of /. leptoclada. Rio Huallaga basin, Killip & Smith 
28289, type of /. congestiflora; 29096. Rio Amazonas and vicinity 
of Iquitos, Klug 60, 84, 363; Williams 1613. Amazonian Brazil, 
Venezuela, and Colombia; Magdalena Valley of Colombia. 

4. OSTEOPHLOEM Warb. 

Dioecious trees; leaves glabrous at maturity, petiolate, the blades 
coriaceous, entire and narrowly revolute at the margins; inflores- 
cences 1-3 in the leaf axils or on defoliate branchlets, simple or with 
1-4 short, lateral branches; flowers pedicellate, the pedicels bracteo- 
late at the summit, the perianth carnose, 3-lobed; filaments connate 
into a carnose column; anthers usually 12 (sometimes 14), linear, 
dorsally adnate to a carnose connective mass; ovary conic, the stigma 
sessile, oblique; fruit transversely ellipsoid, 2-valved, conspicuously 
carinate, the pericarp woody, the seed about half as long as broad. 

Osteophloem platyspermum (A. DC.) Warb. Nov. Act. Acad. 
Leop.-Carol. 68: 162. pi. 1, f. 1-5; pi. 4, f. 1-4- 1897. Myristica 
platysperma A. DC. in DC. Prodr. 14: 695. 1857. 



776 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tree up to 40 meters high; branchlets minutely and densely 
puberulent with sessile, stellate, 4-6-branched hairs, soon glabrous; 
petioles 15-30 mm. long; leaf blades often shining above, oblong- 
obovate, 8-20 cm. long, 3-6.5 cm. broad, gradually attenuate at the 
base, rounded at the apex and often slightly emarginate, minutely 
punctate and often ceriferous beneath, the secondary nerves 6-12 
per side; staminate inflorescences densely puberulent on all exterior 
surfaces, 1-6 cm. long; flowers solitary or in irregular fascicles of 
2-8, the pedicels up to 5 mm. long; bracteole semiorbicular, about 
2 mm. broad and 1 mm. long; perianth about 4 mm. long, lobed 
nearly to the base; androecium 2.5-3 mm. long, the filament column 
1 mm. long or less; ovary densely and minutely lepidote-tomentel- 
lous; fruiting inflorescences glabrous throughout at maturity, the 
mature fruits 15-25 mm. long, 20-25 mm. broad, the pericarp 0.5- 
1.5 mm. thick. 

Loreto: Mouth of Rio Santiago, Tessmann 4505. Near Iquitos, 
Klug 262. Rio Amazonas, Williams 3140. Amazonian Brazil and 
Colombia. 

5. VIROLA Aubl. 

Dioecious trees or rarely shrubs, the inner bark usually exuding 
a brownish or reddish liquid, the branchlets tomentose or puberulent 
when young; leaves petiolate, the blades usually glabrous above 
and pubescent beneath (hairs stellate or irregularly stellate), the 
secondary nerves anastomosing toward the margins; inflorescences 
solitary, axillary, usually broadly paniculate, pedunculate, the pistil- 
late usually more compact than the staminate; bracts sometimes 
prominent; bracteoles none; flowers usually in fascicles terminating 
the ultimate peduncles, pedicellate or subsessile; perianth usually 
thin-carnose and pubescent without, 3 (sometimes 4, rarely 5) -lobed ; 
filaments connate into a column, the anthers usually 3 (2-6), con- 
nate to the apex or distally divergent; ovary globose or ellipsoid, 
the style short; fruit globose or ellipsoid, 2-valved, the pericarp usu- 
ally woody, the aril deeply laciniate, the seed globose or ellipsoid. 
"Cumala." 
Hairs of lower leaf surfaces stalked, stellate or irregularly branched, 

persistent. 
Secondary nerves close, numerous (40-60 per side). 

Leaf blades 5-11 cm. long, 1.7-4 cm. broad; staminate perianth 
uniformly and sparsely puberulent V. flexuosa. 

Leaf blades 25-60 cm. long, 8-21 cm. broad; staminate perianth 
pubescent only at the base and apex. 



FLORA OF PERU 777 

Upper surface of leaf blades glabrous; anthers shorter than 
the filament column V. multinervia. 

Upper surface of leaf blades pilose with stiff hairs; anthers 

slightly longer than the filament column . . V. decorticans. 

Secondary nerves comparatively spaced, fewer (10-30 per side). 

Fruit persistently tomentose (hairs jointed, at least 1.5 mm. 

long) ; inflorescence branches tomentose (hairs 1-4 mm. long). 

Leaf blades often glaucous beneath; hairs of the inflorescence 
branches and fruit usually without lateral branches, 
those of the fruit 7-9 mm. long; anthers subequal to the 
filament column V. loretensis. 

Leaf blades not conspicuously glaucous; hairs of the inflores- 
cence branches and fruit with conspicuous lateral branches, 
those of the fruit 2-2.5 mm. long; anthers about twice 
as long as the filament column V. mollissima. 

Fruit deciduously tomentose (hairs 0.2-0.8 mm. long); inflores- 
cence branches closely tomentellous or puberulent (hairs 
0.1-0.7 mm. long). 
Leaf blades coriaceous; fruit usually slightly ellipsoid. 

V. sebifera. 
Leaf blades submembranous or chartaceous; fruit essentially 

globose V. cuspidata. 

Hairs of the lower leaf surfaces sessile, stellate, often evanescent. 

Leaf blades 20-55 cm. long, 7-24 cm. broad, the secondary nerves 

comparatively distant (12-27 per side); filament column 

sharply contracted distally; staminate inflorescence 12-30 

cm. long V. calophylla. 

Leaf blades rarely to 35 cm. long and 11 cm. broad, the secondary 
nerves closer; filament column not sharply contracted distally; 
staminate inflorescence not more than 18 cm. long. 
Secondary nerves very close, 40-50 per side; leaf blades less than 

11 by 4 cm V. flexuosa. 

Secondary nerves less than 40 per side. 

Perianth lobed more than one-half its length; anthers sub- 
equal to the filament column or shorter; fruit ellipsoid, 
at maturity 25 mm. long or more. 

Secondary nerves 15-20 per side; leaf blades oblong-elliptic 
or narrowly obovate-elliptic; pericarp not lenticellate. 

V. Pavonis. 



778 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Secondary nerves 23-40 per side; leaf blades oblong or obo- 
vate-oblong; pericarp lenticellate at maturity. 

V. Weberbaueri. 
Perianth lobed distally about one- third its length; anthers 2-3 

times as long as the filament column. 
Secondary nerves 17-30 per side; leaf blades abruptly cus- 
pidate or obtuse at the apex; fruit ellipsoid, 20-28 mm. 

long, soon glabrous V. peruviana. 

Secondary nerves 9-20 per side; leaf blades gradually acu- 
minate or acute at the apex; fruit often subglobose, 
8-16 mm. in diameter, when young densely tomentose. 

V. elongata. 

Virola calophylla Warb. Nov. Act. Acad. Leop.-Carol. 68: 231. 
1897. Myristica calophylla Spruce, Journ. Linn. Soc. 5: 4, nomen. 
1860. 

A tree up to 20 meters high; branchlets and petioles closely puber- 
ulent; petioles 7-20 mm. long; leaf blades coriaceous, oblong or ellip- 
tic-oblong, 20-55 cm. long, 7-24 cm. broad, deeply cordate to broadly 
obtuse at the base, acuminate or cuspidate at the apex, closely and 
minutely puberulent beneath (hairs sessile-stellate), the secondary 
nerves 12-27 per side; staminate inflorescences up to 30 cm. long and 
18 cm. broad, minutely and densely tomentellous on all exterior sur- 
faces; pedicels 1-2 mm. long; perianth infundibuliform, 1.3-2 mm. 
long, 3-lobed about one-third its length; 3 anthers, 0.4-0.7 mm. long, 
exceeding the filament column, connate to the apex; fruiting inflores- 
cences up to 7 cm. long, the mature fruits ellipsoid, 15-25 mm. long, 
12-15 mm. broad, closely and minutely brown-sessile-stellate-tomen- 
tellous, glabrescent, the pericarp 1-5 mm. thick. 

Loreto: Near Iquitos, Kittip & Smith 29933; King 188, 473, 533. 
Rio Itaya, Kittip & Smith 29823; Williams 3291. Rio Nanay, Wil- 
liams 978, 1085. Amazonian Brazil and adjacent Venezuela and 
Colombia. "Cumalablanca." 

Virola cuspidata (Benth.) Warb. Nov. Act. Acad. Leop.-Carol. 
68: 176. 1897. Myristica cuspidata Spruce in Benth. Journ. Bot. & 
KewMisc. 5:5. 1853. 

A shrub or slender tree, up to 10 meters high; branchlets and 
petioles tomentellous, glabrescent; petioles 8-15 mm. long; leaf blades 
thin-coriaceous or submembranous, oblong or narrowly elliptic- 
oblong, 15-35 cm. long, 4-10 cm. broad, obtuse to shallowly cordate 
at the base, acuminate or cuspidate at the apex, tomentose beneath 



FLORA OF PERU 779 

(hairs stalked, 0.2-0.3 mm. long), the secondary nerves 11-25 per 
side; staminate inflorescences up to 22 cm. long and 15 cm. broad, 
closely tomentellous (hairs up to 0.5 mm. long); pedicels up to 3 
mm. long; perianth infundibuliform, 1.5-2.5 mm. long, 3-lobed about 
one-third its length; usually 3 anthers, 0.6-1 mm. long, 1-2 times as 
long as the filament column, usually apiculate and connate nearly 
to the apex; pistillate inflorescences up to 6 cm. long, the ovary 
densely tomentose (hairs about 0.3 mm. long); fruits subglobose, 
10-14 mm. in diameter, at length glabrescent, the pericarp less 
than 0.7 mm. thick. 

Loreto: Vicinity of Iquitos, Tessmann 3615; Killip & Smith 
27110; King 1385, 2502. Amazonian Brazil. 

Virola decorticans Ducke, Journ. Wash. Acad. Sci. 26: 262. 
1936. 

A tree; branchlets and petioles densely tomentose (hairs about 
1 mm. long, irregularly branched) ; petioles stout, 7-20 mm. long; leaf 
blades coriaceous, broadly elliptic or obovate-elliptic, 25-60 cm. long, 
11-21 cm. broad, truncate or shallowly cordate at the base, cuspidate 
or abruptly caudate-acuminate at the apex, pilose above (hairs stiff, 
simple or forked, 0.5-1 mm. long), tomentose beneath (hairs stalked, 
stellate or irregularly branched, about 1 mm. long), the secondary 
nerves 45-60 per side; staminate inflorescences up to 22 cm. long 
and broad, densely tomentose; bracts conspicuous, long-persistent; 
pedicels up to 3 mm. long; perianth 1.5-1.8 mm. long, strigose at the 
base and apex, otherwise glabrous, 3-lobed nearly to the base; anthers 
3, about 0.5 mm. long, slightly longer than the filament column, con- 
nate to the apex; mature fruits ovoid-ellipsoid, 27-35 mm. long, 
17-22 mm. broad, densely and persistently tomentose (hairs 1-1.5 
mm. long). 

Loreto: Rio Amazonas, La Victoria, Williams 3077. Amazonian 
Brazil. 

Virola elongata (Benth.) Warb. Nov. Act. Acad. Leop.-Carol. 
68: 178. pi. 5. 1897. Myristica elongata Benth. Journ. Bot. & Kew 
Misc. 5:5. 1853. 

A shrub or slender tree, occasionally up to 25 meters high; 
branchlets and petioles tomentellous, glabrescent; petioles 5-16 mm. 
long; leaf blades papyraceous or submembranous, narrowly oblong 
to oblong-elliptic, up to 32 cm. long and 11 cm. broad (rarely 7 by 
1.5 cm.), obtuse to subcordate at the base, usually long-acuminate 
at the apex, puberulent beneath (hairs sessile-stellate), the secondary 



780 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nerves 9-20 per side; staminate inflorescences up to 18 cm. long and 
15 cm. broad, closely tomentellous, glabrescent; pedicels up to 3 
mm. long; perianth infundibuliform, 1.8-3 mm. long, 3-4-lobed about 
one-third its length; anthers usually 3 (occasionally as many as 6), 
0.6-1.6 mm. long, 2-3 times as long as the filament column, usually 
apiculate and connate nearly to the apex; pistillate inflorescences 
up to 7 cm. long, the ovary densely tomentellous (hairs about 0.2 
mm. long); fruits ellipsoid or subglobose, 11-16 mm. long, 8-12 mm. 
broad, soon glabrescent, the pericarp usually less than 1 mm. thick. 
Loreto: Rio Putumayo, Klug 2166. Rio Maranon, Tessmann 
4852. Basin of Rio Huallaga, Killip & Smith 28159; Klug 2944; 
Williams 4575. Rio Amazonas and Iquitos, Tessmann 3704; Klug 
356; Williams 2404, 2409, 2526, 2552, 2626, 2660, 2839, 2875. 
Department (?): Ruiz & Pavon. Amazonian Brazil and adjacent 
Venezuela and Colombia. "Ucufe-ey" (Rio Putumayo); "cumala 
caspi" (Rio Huallaga); "cumala blanca." 

Virola flexuosa A. C. Smith, Brittonia 2: 151. 1936. 

A tree up to 30 meters high; young branchlets and petioles 
densely tomentellous; petioles 2-7 mm. long; leaf blades submem- 
branous, oblong or narrowly elliptic, 5-11 cm. long, 1.7-4 cm. broad, 
cordate at the base, obtuse or acute at the apex, tomentellous 
beneath (hairs sessile or short-stalked), the secondary nerves 40-50 
per side; staminate inflorescences up to 9 cm. long and broad, densely 
tomentellous; flowers arranged in small, compact clusters, the pedi- 
cels about 1 mm. long; perianth 1.2-1.6 mm. long, 3-lobed nearly 
to the base; anthers 3, about 0.4 mm. long, subequal to the filament 
column, connate to the apex. 

Loreto: Lower Rio Huallaga, Williams 4423. Rio Ucayali, 
Yarina-cocha, Tessmann 3207. Amazonian Brazil and adjacent 
Bolivia. 

Virola loretensis A. C. Smith, Bull. Torrey Club 58: 95. 1931. 
V. villosa Ducke, Bull. Mus. Paris II. 4: 724. 1932. 

A slender tree, 4-10 meters high ; hairs of the branchlets, petioles, 
lower leaf surfaces, and inflorescence branches ferruginous, multi- 
cellular, 1-4 mm. long; petioles 3-12 mm. long; leaf blades thin- 
coriaceous or papyraceous, oblong, oblong-elliptic, or narrowly obo- 
vate-oblong, 15-35 cm. long, 4-10 cm. broad, cordate or rounded 
at the base, acute or acuminate at the apex, the secondary nerves 
15-26 per side; staminate inflorescences up to 25 cm. long and 15 
cm. broad; pedicels slender, 2-4 mm. long; perianth infundibuliform, 



FLORA OF PERU 781 

1.5-2.5 mm. long, 3-4-lobed nearly one-half its length; anthers 3, 
0.5-0.8 mm. long, equaling or slightly exceeding the filament column, 
apiculate at the apex; ovary densely tomentose; fruits 10-40 per 
inflorescence, subglobose, 13-22 mm. in diameter, densely and per- 
sistently tomentose (hairs 7-9 mm. long), the pericarp usually less 
than 0.5 mm. thick. 

Loreto: Vicinity of Iquitos, Kittip & Smith 27026, 27359, type; 
Ducke 17983, type of V. villosa; Kuhlmann 17894; King 1502, 1543. 
Rio Itaya, Kittip & Smith 29671 ; Williams 3289, 3447. Rio Ama- 
zonas, Williams 2264- Amazonian Brazil. 

Virola mollissima (A. DC.) Warb. Nov. Act. Acad. Leop.- 
Carol. 68: 167. 1897. Myristica mollissima A. DC. in DC. Prodr. 
14:696.1857. 

A slender tree, 5-12 meters high; branchlets, petioles, lower leaf 
surfaces, and inflorescence branches persistently tomentose (hairs 
ferruginous, multicellular, 1-2 mm. long); petioles stout, 8-21 mm. 
long; leaf blades thin-coriaceous, oblong or obovate-oblong, 25-50 
cm. long, 8-19 cm. broad, deeply cordate to rounded at the base, 
acuminate at the apex, the secondary nerves 15-30 per side; stami- 
nate inflorescences up to 25 cm. long and 18 cm. broad; pedicels up 
to 3 mm. long; perianth infundibuliform, about 2 mm. long, 3-lobed 
about one-third its length; anthers 3, about 1 mm. long, twice as 
long as the filament column, apiculate at the apex; fruits 5-10 per 
inflorescence, subglobose, 11-14 mm. in diameter, densely and per- 
sistently tomentose (hairs 2-2.5 mm. long), the pericarp less than 
0.5 mm. thick. 

Loreto: Lower Rio Huallaga basin, Poeppig s. n.; 2195, type; 
Killip & Smith 29044; King 2822, 3128. Rio Ucayali basin, Yarina- 
cocha, Tessmann 3473. 

Virola multinervia Ducke, Journ. Wash. Acad. Sci. 26: 261. 
1936. 

A tree up to 30 meters high; branchlets and petioles densely ferru- 
ginous- tomentose (hairs to 1 mm. long, often irregularly branched) ; 
petioles stout, 4-15 mm. long; leaf blades coriaceous, elliptic-oblong, 
25-45 cm. long, 8-16 cm. broad, rounded or shallowly cordate at the 
base, acute or cuspidate at the apex, tomentose beneath (hairs 
stalked, irregularly branched, 0.3-0.5 mm. long), the secondary 
nerves 40-60 per side; staminate inflorescences up to 20 cm. long 
and broad, densely tomentose; pedicels 1-2 mm. long; perianth 1.2- 
1.5 mm. long, distally strigose, otherwise glabrous, 3-lobed nearly 



782 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to the base; anthers 3, about 0.4 mm. long, slightly shorter than the 
filament column, connate to the apex, obtuse; pistillate inflorescences 
robust, the ovary subglobose, densely tomentellous (hairs about 0.3 
mm. long); mature fruits ellipsoid or obovoid, 20-30 mm. long, 15-25 
mm. broad, densely and persistently setulose (hairs 1-2 mm. long), 
the pericarp 1.5-4 mm. thick. 

Loreto: Lower Rio Huallaga, Yurimaguas, Kuhlmann 24557. 
Amazonian Brazil. 

Virola Pavonis (A. DC.) A. C. Smith, Brittonia 2: 504. 1938. 
Myristica Pavonis A. DC. in DC. Prodr. 14: 697. 1857. V. venosa 
var. Pavonis Warb. Nov. Act. Acad. Leop.-Carol. 68: 225. pi. 7, 
f. 1, 2. 1897. 

A tree up to 23 meters high; young branchlets and petioles 
puberulent, soon glabrous; petioles 4-13 mm. long; leaf blades coria- 
ceous or thin-coriaceous, oblong-elliptic or narrowly obovate-elliptic, 
8-21 cm. long, 2-6.5 cm. broad, attenuate to obtuse or rarely 
rounded at the base, obtusely cuspidate to rounded at the apex, 
sparsely pale-puberulent beneath (hairs sessile-stellate), soon gla- 
brous, the secondary nerves 15-20 per side; staminate inflorescences 
up to 15 cm. long and broad, densely puberulent, soon glabrous; 
pedicels up to 2 mm. long, the perianth 1.5-2 mm. long, 3-lobed more 
than half its length; anthers three, 0.4-0.7 mm. long, shorter than 
the filament column, usually connate to the apex; pistillate inflores- 
cences about as large as the staminate, the ovary densely and closely 
puberulent; mature fruits ellipsoid, 25-50 mm. long, 15-23 mm. broad, 
usually distinctly carinate, glabrous, the pericarp 2-7 mm. thick. 

Huanuco: Chicoplaya, Ruiz & Pavon (type). Loreto: Balsa- 
puerto, Klug 3085. Mouth of Rio Santiago, Tessmann 4047, 4122, 
4146. Near Iquitos, Killip & Smith 27145. Rio Nanay, Williams 
1166. Loreto or San Martin: Rio Huallaga, Poeppig 2058. San 
Martin: Near Moyobamba, 1,100 meters, Klug 3713. Amazonian 
Brazil. 

Virola peruviana (A. DC.) Warb. Nov. Act. Acad. Leop.-Carol. 
68: 188. 1897. Myristica peruviana A. DC. Ann. Sci. Nat. IV. 4: 
30. 1855. 

A tree; branchlets and petioles tomentellous; petioles 7-20 mm. 
long; leaf blades coriaceous or thin-coriaceous, oblong, 16-35 cm. 
long, 6.5-10.5 cm. broad, shallowly cordate or rounded at the base, 
obtuse to cuspidate at the apex, tomentellous beneath (hairs sessile- 
stellate), glabrescent, the secondary nerves 17-30 per side; staminate 



FLORA OF PERU 783 

inflorescences 10-18 cm. long, closely puberulent or glabrous; pedi- 
cels up to 2 mm. long; perianth infundibuliform, 2.3-3.2 mm. long, 
usually densely tomentellous, 3-lobed about one- third its length; 
3 or 4 anthers, 1.1-1.6 mm. long, 2-3 times as long as the filament 
column, connate to the apex and apiculate; pistillate inflorescences 
3-5 cm. long, the ovary ellipsoid, densely tomentose (hairs about 
0.2 mm. long); mature fruits ellipsoid, up to 28 mm. long and 22 
mm. broad, at length glabrescent, conspicuously carinate, the peri- 
carp less than 1.5 mm. thick. 

Huanuco: Pampayacu, Sawada 19. Junin: La Merced, Weber- 
bauer 1861. Loreto: Rio Ucayali, Tessmann 3055, 3259. Depart- 
ment^): Ruiz & Pawn, type. Monterico (Peru?), Pearce. Amazo- 
nian Brazil (upper Rio Purus). 

Virola sebifera Aubl. PL Guian. 2: 904. pi. 345, f. 1-5. 1775. 
Myristica Mocoa A. DC. in DC. Prodr. 14: 195. 1856. V. Mocoa 
Warb. Nov. Act. Acad. Leop.-Carol. 68: 183. 1897. V. peruviana 
var. tomentosa Warb. Nov. Act. Acad. Leop.-Carol. 68: 189. 1897. 

A tree up to 40 meters high; branchlets and petioles tomentose, 
glabrescent; petioles 10-25 mm. long; leaf blades coriaceous, oblong 
or elliptic-oblong, 15-47 cm. long, 6-15 cm. broad, cordate to obtuse 
(occasionally subacute) at the base, acuminate or acute at the apex, 
tomentose beneath (hairs stalked, 0.2-0.3 mm. long), the secondary 
nerves 10-28 per side; staminate inflorescences 8-23 cm. long and 
broad, closely tomentellous on all exterior surfaces (hairs less than 
0.2 mm. long); pedicels slender, up to 3 mm. long; perianth infundi- 
buliform, 1.8-3 mm. long, 3 (occasionally 4 or 5) -lobed about one- 
third its length; anthers 3 (sometimes 4 or 5), 0.7-1.5 mm. long, 
2-4 times as long as the filament column, apiculate and usually 
connate at the apex; pistillate inflorescences 3-7 cm. long, the ovary 
densely tomentose (hairs 0.2-0.3 mm. long); fruits ellipsoid or sub- 
globose, 10-19 mm. long, 7-14 mm. broad, densely tomentose at 
maturity, at length glabrescent, the pericarp usually less than 1 mm. 
thick. 

Huanuco : Chicoplaya, Ruiz & Pawn, type of V. peruviana var. 
tomentosa. Loreto: Rio Huallaga, Kuhlmann 23688, 24546; Killip 
& Smith 27935. Mainas, Poeppig 2390, type of M. Mocoa. San 
Martin: Tarapoto, Spruce 4878. San Roque, 1,350-1,500 meters, 
Williams 6953, 7634. Rio Huallaga, 360-900 meters, Williams 6615. 
Zepelacio, 1,100 meters, Klug 3645. Continental tropical America, 
from Nicaragua to Bolivia and southern Brazil. 



784 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Virola Weberbaueri Markgraf, Notizbl. Bot. Gart. Berlin 
9: 965. 1926. 

A tree up to 30 meters high; branchlets and petioles closely 
puberulent; petioles 3-8 mm. long; leaf blades thin-coriaceous, 
oblong or obovate-oblong, 15-22 cm. long, 4.5-7 cm. broad, obtuse 
or narrowly rounded at the base, cuspidate at the apex, closely 
puberulent beneath (hairs sessile-stellate), glabrescent, the secondary 
nerves 23^40 per side; staminate inflorescences 10-15 cm. long, 
closely and minutely puberulent; pedicels up to 5 mm. long; perianth 
2-3.2 mm. long, 3-lobed nearly to the base; anthers 3, about 1 mm. 
long, subequal to the filament column, minutely apiculate and essen- 
tially connate to the apex; mature fruits ellipsoid, up to 30 mm. long 
and 22 mm. broad, closely brown-puberulent, soon glabrous, the 
pericarp about 2 mm. thick. 

Junin: La Merced, 1,000 meters, Weberbauer 1893, type. Loreto: 
Mouth of Rio Santiago, Tessmann 4005, 4339. 

75. MONIMIACEAE 

Reference: Perkins & Gilg, Pflanzenreich IV. 101. 1901; Nachtr. 
1911. 

Besides the following, Laurelia sempervirens (R. & P.) Tul. has 
been credited by Perkins on the basis of a Pa von specimen without 
definite locality. It is certainly Chilean, the type collection at 
Madrid being labeled as from Conception. 

Anthers large, evalvate; filaments none or very short; sepals 4, 
obvious; petioles and styles mostly short 1. Mollinedia. 

Anthers small; filaments well developed, often broad; sepals 4-7, 
often obsolete; petioles often and styles always long. 

2. Siparuna. 
1. MOLLINEDIA R. & P. 

Tetratame Poepp. & Endl. 

Dioecious shrubs or trees with opposite, rarely entire leaves and 
mostly small flowers, the staminate usually disposed in panicles, 
the pistillate in racemes. Anthers longitudinally dehiscent, but the 
cells mostly confluent. Staminate sepals 4, usually unequal, the 
pistillate equal and forming a calyptrum. Stamens 8-50. Carpels 
6-35. Ovule pendulous. Two keys are appended, the first for 
fruiting material, especially for three forms proposed by me as new 
species which, however, have not been studied in connection with 
the preparation of this account. 



FLORA OF PERU 785 

Mature leaves distinctly but sometimes finely pubescent beneath, 

at least along the midrib (M. simulans). 
Leaves rusty-villous beneath, especially on the midnerve, even 

in age. 
Petioles 12-15 mm. long; blades broadly ovate-elliptic. 

M. repanda. 

Petioles 6-8 (13) mm. long; blades oblong M. pulcherrima. 

Leaves not rusty-villous; petioles 5-8 mm. long, sometimes longer. 
Leaves at least shortly acuminate; branchlets minutely pubes- 
cent. 

Leaves shortly and broadly acuminate, mostly 10-15 cm. long. 

M. latifolia. 
Leaves long-acuminate, 20-30 cm. long. 

Petioles short; sepals entire M. grandifolia. 

Petioles elongate; inner sepals laciniate M. Krukovii. 

Leaves obtuse or rarely acute; branchlets densely hairy. 

M. Casca. 

Mature leaves glabrous beneath or not obviously pubescent. 
Leaves ovate to elliptic, mostly wider than 5 cm. 
Leaves coarsely serrate, the remote teeth 3-4 mm. long. 

M. grosseserrata. 

Leaves indistinctly or repandly and shortly denticulate. 
Leaves caudate-acuminate, the acumen about 1.5 cm. long. 

M. Killipii. 
Leaves acute to acuminate. 

Leaves oblong-elliptic, 5-6.5 cm. wide M. Tessmannii. 

Leaves broadly elliptic, 8-14 cm. wide. 

Inner sepals appendaged; leaves glabrous. . . .M. ovata. 

Inner sepals unappendaged ; leaf nerves minutely puberu- 

lent M. grandifolia. 

Leaves oblong to lanceolate, mostly less than 5 cm. wide. 
Leaves glabrous or only the youngest sparsely pubescent. 
Leaves denticulate or, if acuminate, cuneately so. 

Petioles about 1 cm. long; stamens about 18. .M. lanceolata. 

Petioles 6-8 mm. long; stamens about 24 M. racemosa. 

Leaves entire, caudate-acuminate M. caudata. 

Leaves minutely pubescent beneath in age, entire . . M. simulans. 



786 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Key after Perkins 

(M. Casca, M. caudata, and M. simulans omitted) 
Sepals similar, the inner not appendaged. 
Pubescence conspicuous. 

Leaves broadly ovate-elliptic M. repanda. 

Leaves oblong M. pulcherrima. 

Pubescence minute or wanting. 
Leaves 5-7 cm. wide. 

Stamens few; leaves caudate M. Killipii. 

Stamens many; leaves short-acuminate M. Tessmannii. 

Leaves about 14 cm. wide M. grandifolia. 

Sepals unequal, the inner appendaged. 

Cells of the outer anthers not at all confluent apically . . M. ovata. 
Cells of all anthers confluent apically. 

Stamens about 18; leaves 1.5-3 (-5) cm. wide. . .M. lanceolata. 
Stamens about 25 or more. 

Leaves coarsely dentate M. grosseserrata. 

Leaves entire or finely denticulate. 

Stamens about 25; inflorescence elongate M. racemosa. 

Stamens 30-50; inflorescence short. 

Leaves dentate M. latifolia. 

Leaves entire M. Krukovii. 

Mollinedia Casca Macbr. Candollea 5: 352. 1934. 

Robust, with densely appressed-pilose branchlets; petioles 7-10 
mm. long, glabrate; blades broadly elliptic, rounded at both ends 
or obscurely acute at the apex, about 14 cm. long, half as wide, entire, 
chartaceous, glabrous above, prominently reticulate-veined and 
sparsely puberulent beneath; flowers unknown; carpels about 12, 
glabrous or glabrate, 1.5 cm. long. 

Loreto: Lower Rio Huallaga, 200 meters, Williams 4585, type. 

Mollinedia caudata Macbr. Candollea 5: 351. 1934. 

A glabrous tree, 5 meters high, with slender, terete, remotely 
branched branches; internodes 4-6 cm. long; petioles 5-10 mm. long; 
blades pale green, chartaceous-membranaceous, very densely pellucid- 
puncticulate, obscurely veined, entire, oblong- or ovate-elliptic, 
rounded-acute at the base, subabruptly caudate-acuminate, the acu- 



FLORA OF PERU 787 

men to 2 cm. long, mostly 10-12 cm. long, 3.5-4.5 cm. wide; carpels 
about 6, black, glabrous, 1.5 cm. long. 

Loreto: Pebas, on the Amazon, in forest, Williams 1602, type; 
1724, 1747. Pumayacu, 1,000 meters, in forest, Klug 3215; a tree 
of 10 meters. Junin: Puerto Bermudez, 375 meters, Killip & Smith 
26562. 

Mollinedia grandifolia Perk. Notizbl. Bot. Gart. Berlin 10: 
158. 1927. 

A small, dioecious shrub, only the youngest branchlets, the 
petioles, the leaf nerves beneath, and the receptacle (outside) very 
minutely pubescent; leaves thin, broadly elliptic, 14.5 cm. wide, 
cuneate-rounded at the base, the lateral nerves 6-9; inflorescence 
5-7 cm. long, few-flowered, very lax, the pedicels 5-8 mm. long; 
staminate flowers yellowish green, 6-7 mm. broad; sepals not append- 
aged, the outer ones ovate, entire, larger than the inner ones; stamens 
40-50. Placed by the author near M. salicifolia Perk. Neg. 13434. 

Loreto: Flooded areas, upper Rio Maranon, Tessmann 4581, type. 

Mollinedia grosseserrata Perk. Bot. Jahrb. 27: 673. 1900; 44. 

Leaves about 10 cm. long and 4-7.5 cm. wide, rounded or more 
or less cuneate at the base, shortly, broadly, and acutely acuminate, 
glabrous, deeply serrate on the upper margins, chartaceous; inflores- 
cence 4-5 cm. long, the pedicels 6 mm. long; staminate flowers 5 mm. 
broad; receptacles ashy-pilose, longer than the rigid, subequal sepals; 
inner sepals with a long, deeply incised, incurved appendage; sta- 
mens 24, the anther cells confluent. The Williams (sterile) specimen 
has similar leaves but the caudate tips are 1 cm. long. Neg. 13435. 

Peru: Without locality, Lobb, type. Loreto : Near Altura, Rio 
Nanay, in forest, Williams 857(?). 

Mollinedia Killipii Macbr. Candollea 5: 351. 1934. 

A tree of 4.5-6 meters, with slender, glabrous branchlets, the 
internodes about 5 cm. long; petioles 8-10 mm. long; blades elliptic, 
about 12 cm. long, 5-7 cm. wide, rounded and acute or broadly 
cuneate at the base, narrowly caudate-acuminate, the acumination 
1.5 cm. long, entire, membranous, noticeably reticulate-veined only 
beneath, glabrous or very minutely and sparsely puberulent beneath ; 
inflorescence few-flowered, glabrous; pedicels slender, strongly divari- 
cate, 2.5 cm. long; receptacles obconic, 4 mm. long, 3.5 mm. wide; 
sepals subequal, entire, inflexed, truncate, 2 mm. long; stamens 



788 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

about 15, nearly 1.5 mm. broad, the cells confluent, the connective 
scarcely or not produced. 

Loreto: Soledad, on Rio Itaya, 110 meters, in forest, Killip & 
Smith 29751, type. 

Mollinedia Krukovii A. C. Smith, Phytologia 1: 114. 1935. 

Younger branchlets, petioles, and leaves beneath minutely pilose; 
petioles 1.5-3.5 cm. long; blades chartaceous, ovate, 20-30 cm. long, 
11-17 cm. wide, acute at the base, the apical acumination 1 cm. 
long, entire, becoming glabrescent beneath, obscurely but abun- 
dantly pellucid-punctulate, the lateral nerves 7-9 pairs, elevated on 
both sides, the many reticulate veins prominent beneath; pedicels 
8-10 mm. long, finely pilose; staminate flowers yellow, the fleshy 
receptacle glabrous within; outer sepals broadly ovate, to 10 mm. 
long, nearly as broad, the inner ones oblong-orbicular, about 8 mm. 
long and broad, with a thin, laciniate margin; stamens about 50, 
the glabrous, fleshy filaments 1 mm. long, the oblong anthers twice 
as long. Compared by the author with M. latifolia; cf. Klug 3215, 
which perhaps belongs here, cited under M . caudata. 

Peru: Probably. Amazonian Brazil. 

Mollinedia lanceolata R. & P. Syst. 1: 143. 1798; 43. 

Leaves 12-17 cm. long, 1.5-3 (-5) cm. wide, gradually narrowed 
to the base and to the acuminate apex, thin but rigid, glabrous, irregu- 
larly denticulate above the middle; staminate flowers 3.5 mm. 
broad, the papery receptacle plane, sparsely yellowish-pilose, the 
outer sepals much larger than the fimbriate-appendaged inner ones; 
stamens sessile, the anther cells confluent. Neg. 13441. 

Peru: Without locality, Ruiz & Pavon, type. Santa Cruz, Cerro, 
1,800 meters (Pearce; Peru?). 

Mollinedia latifolia (Poepp. & Endl.) Tul. Monogr. 402. 1855; 
49. Tetratome latifolia Poepp. & Endl. Nov. Gen. & Sp. 2: 47. 1838. 

Leaves broadly ovate-elliptic, 15-20 cm. long, rounded at the 
base, papyraceous, regularly but remotely and inconspicuously den- 
ticulate toward the acute apex, sparsely and minutely pubescent 
beneath; inflorescence 2.5 cm. long, the pedicels 5 mm. long; flowers 
3.5 mm. broad; receptacles papery, densely yellowish-pilose, longer 
than the very unequal sepals, the inner sepals deeply fimbriate- 
appendaged; stamens about 30, equal; pistillate flowers ashy-pilose, 
like the numerous carpels; style very short. A shrub or tree 6 
meters high. The pubescence is typically minute. Neg. 13442. 



FLORA OF PERU 789 

Loreto: Mainas, Poeppig 2216, type. Balsapuerto, 150-350 
meters, in forest, Kittip & Smith 28587, 20691, 28409, 28671. Ama- 
zonian Brazil. 

Mollinedia ovata R. & P. Syst. 1: 143. 1798; 41. 

Leaves ovate-oblong, rounded at the base, acutely acuminate, 
chartaceous, indistinctly denticulate toward the apex, glabrous; 
inflorescence 5.5 cm. long, 4-5-flowered, the peduncles 2 cm. long, 
the pedicels half as long; staminate flowers 4-8 mm. broad; recep- 
tacles papyraceous, nearly glabrous, equaled by the minutely dentic- 
ulate, subequal sepals; stamens 33-35; anther connective produced; 
pistillate flowers yellow, 4-5 mm. broad, yellowish-pilose like the 
receptacles within and the 35 carpels. Fruit of the Killip and Smith 
specimen 1 cm. thick, obtuse, green. Neg. 13450. 

Without locality: Ruiz & Pavon. Rio Acre: Ule 9395. Aya- 
cucho: Rio Apurimac Valley, 400 meters, Killip & Smith 22862. 
San Martin: San Roque, in forest, 1,400 meters, Williams 7187. 

Mollinedia pulcherrima Sleumer, Repert. Sp. Nov. 39: 275. 
1936. 

Except for the leaves above (these at maturity glabrous) densely 
clothed throughout, even to the sepals without, with a brown, velu- 
tinous indument; leaves oblong, gradually acuminate, the tip 
subacute, the base mostly subrotund, rarely broadly attenuate, char- 
taceous, entire or minutely and remotely denticulate above the 
middle, 11-19 cm. long, 4-6.5 cm. wide, the lateral nerves slightly 
impressed above, strongly developed beneath, the veins distinctly 
reticulate; inflorescence 4-6 cm. long; pedicels about 1.5 cm. long; 
receptacle plane, chartaceous, the staminate 8-9 mm. broad; outer 
sepals broadly ovate, 4-5 mm. long, glabrous within; stamens many, 
more than 20, sessile, glabrous, the anthers nearly hippocrepous; 
pistillate flowers unknown; drupes (young) sessile, crowded, pilose. 
Separated from M. caloneura Perk., 39, of Bolivia, with the nerves 
strongly impressed above, the lateral ones more numerous. 

Cajamarca: Tambillo, Cutervo (Raimondi, type). 

Mollinedia racemosa (Schlecht.) Tul. Monogr. 382. 1885; 45. 
Tetratome racemosa Schlecht. Linnaea 20 : 114. 1847. T. tri flora Poepp. 
& Endl. Nov. Gen. & Sp. 2: 46. pi. 163. 1838. 

Leaves often more or less fascicled, 10-15 cm. long, 3-5.5 cm. 
wide, cuneate at the base, remotely serrate-denticulate toward the 
attenuate, acute apex, finally glabrous; inflorescence long-pseudo- 



790 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

racemose, to 15 cm. long, many-flowered, the slender, flexuous 
peduncle to 1.5 cm. long, the pedicels 5-8 mm. long; receptacles 
obconic, appressed-sericeous; outer sepals suborbicular, the longer 
inner ones repand-lacerate. Illustrated, Pflanzenreich IV. 101 : /. 6. 
Neg. 13457. 

Loreto: Poeppig 1577, type. 

Mollinedia repanda R. & P. Syst. 1: 142. 1798; 34. 

Branchlets, petioles, leaves beneath, and inflorescences rusty- 
tomentose-pilose; leaves broadly ovate-elliptic, 15-20 cm. long, 7-10 
cm. wide, shortly or not at all acute, rounded or shortly cuneate at 
the base, remotely and obscurely sinuate, coriaceous, sometimes 
rigid, rather rugose-nervose, lustrous and glabrous above; fruiting 
racemes solitary, axillary, the peduncle 1.5 cm., the pedicels 2 cm. 
long; receptacles orbicular, the margins completely reflexed, 15-25- 
carpellate; drupes ovate, subsessile, 1.5-2 cm. long. Neg. 13458. 

Huanuco: Chinchao, Pillao, Ruiz & Pavon. Upper Amazon, 
2,100 meters, Kanehira 13. 

Mollinedia simulans Macbr. Candollea 5: 352. 1934. 

A shrub; branches glabrous, the internodes 3-6 cm. long; petioles 
6-7 mm. long; blades entire, oblong-lanceolate, cuneate or broadly 
cuneate at the base, shortly or long-acuminate, about 12 cm. long, 
3.5-4.5 cm. wide, chartaceous-membranous, glabrous and little 
lustrous above, beneath paler, minutely and sparsely puberulent and 
somewhat reticulate-veined; carpels about 10, appressed-strigose, 
2.5 cm. long. 

San Martin: San Roque, 1,400 meters, in forest, Williams 6977, 
type. Florida, Rio Putumayo, 180 meters, in forest, Klug 2226; a 
shrub 1 meter high. "Amoqui-ey" (Huitoto name). 

Mollinedia Tessmannii Perk. Notizbl. Bot. Gart. Berlin 10: 
159. 1927. 

A small, dioecious tree, the growing parts soon glabrous; leaves 
usually oblong-elliptic, cuneate-rounded at the base, shortly and 
broadly acuminate, the tip acute, minutely and remotely denticulate, 
the lateral nerves 6-7; staminate inflorescence 3-9 cm. long, many- 
flowered, the pedicels 3-5 mm. long; flowers 6-9 mm. long,' about 
half as broad, the pedicels 4-5 mm. long, gray-pilose; sepals unap- 
pendaged, the 2 outer ones broadly, the inner ones narrowly, ovate; 
stamens 50; pistillate inflorescence 4 cm. long, the flowers smaller 
than the staminate ones; sepals minute; receptacles and carpels 



FLORA OF PERU 791 

(about 30) pilose. Allied to M. macrantha Tul., 40, with shorter 
staminate inflorescences and smaller flowers. Neg. 13465. 

Loreto: Cumaria, Rio Ucayali, 225 meters, in lowland forest, 
Tessmann 3349. Sacarita de Masisea, between Cumaria and Iparya, 
215 meters, Tessmann 3364- 

2. SIPARUNA Aubl. 

Citrosma R. & P. 

Shrubs or trees with subentire or denticulate leaves, the petioles 
often unequal, opposite or sometimes ternate. Inflorescence axillary, 
often cymose. Receptacles nearly always campanulate or globose- 
urceolate, the sepals 4-7, sometimes obsolete, not rarely connate 
into an entire or lobed ring. Stamens 1-60, often unequal. Anthers 
dehiscing within. Styles filiform or ligulate, free or connate. Car- 
pels 4-20. Ovule basal, erect. The name of Ruiz and Pavon has 
been spelled by later authorities Citriosma. Since a large number 
of the Peruvian species are imperfectly known, a key based on char- 
acters of foliage and pubescence has been attempted, which, to be 
followed, requires, at least sometimes, material for comparison. 
The technical key is after Miss Perkins' but modified to group a 
majority of the species with few stamens together, because the genus 
is probably divisible into two groups, one with many, the other 
with few stamens. The several species proposed by me have been 
placed in the technical key, but their validity remains doubtful. 
The Peruvian forms, so far as known, are dioecious except as indi- 
cated, but the character, at least for some species, seems to be feeble. 
Ruiz and Pavon, referring to insect deposits frequent on these shrubs, 
observed in their journal that the branches appeared as though 
sprinkled with intent by saliva. 

Artificial key 
Petioles all short, the longest 10-15 mm., most of them shorter. 

Pubescence mostly or entirely simple, hispid, or practically lacking 
on the leaves. 

Pubescence present. 

Hairs spreading; leaves of medium to large size. 

Leaves auricled at the somewhat narrowed base. 

S. auriculata. 

Leaves not at all auricled, cuneate at the base. S. hispida. 
Hairs appressed, the leaves uniformly small (4-9 cm.). 



792 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves entire S. parvifolia. 

Leaves denticulate. 

Leaves slightly pubescent beneath S. ovalis. 

Leaves glabrate S. Mathewsii. 

Pubescence none on the leaves or sometimes 2-3 fascicles of 

hairs on the leaves beneath S. subinodora. 

Pubescence stellate or lepidote, sometimes sparse. 
Branches and inflorescence more or less densely lepidote, the 
scales sometimes stellate, but the scales larger than the 
marginal hairs. 

Scales very minutely or not at all stellate. 
Leaves acute or broadly short-acuminate. 

Leaves sparsely lepidote beneath S. lepidantha. 

Leaves glabrous or sparsely stellate-pilose beneath. 

S. guianensis. 

Leaves acuminate, with a distinct acumen (cf. some forms 
of S. guianensis) . 

Leaves mostly shorter than 9 cm.; stamens 6. 

S. crassi flora. 
Leaves mostly longer than 9 cm.; stamens 4-5. 

S. ucayaliensis. 

Scales rather obviously stellate (cf. also S. guianensis). 
Leaves obovate, with a very abrupt, linear-caudate tip; 

stamens 4-5 S. lepidiflora. 

Leaves elliptic, acuminate; stamens about 12. S. cervicornis. 

Branches and inflorescence more or less stellate-pubescent, some- 
times minutely or densely so but not definitely lepidote. 

Leaves obviously very pubescent, especially beneath, even if 
finely so, the hairs not confined to the nerves. 

Pubescence fine, pale, scabrous-stellate or pilose on the 
leaves above, stellate-puberulent beneath. 

Leaves subentire S. aiperula. 

Leaves closely mucronulate-denticulate . .S. saurauiifolia. 
Pubescence long and soft, often fulvous. 

Leaves closely mucronulate-denticulate . . S. saurauiifolia. 
Leaves remotely or unevenly denticulate or subentire. 
Leaf veins reticulately impressed above . S. Weberbaueri. 



FLORA OF PERU 793 

Leaf veins not impressed above. 

Petioles 4-5 mm. long S. tomentosa. 

Petioles 10 mm. long or longer S. fulva. 

Leaves green, even beneath, though sometimes distinctly 
pubescent on the veins. 

Inflorescence mostly 2-4 cm. long, densely and finely stel- 
late-tomentose. 

Pubescence extraordinarily minute and compact, espe- 
cially on the inflorescence. 

Leaves 6.5-7.5 cm. wide; pedicels 1 mm. long. 

S. cervicornis. 
Leaves 4-5.5 cm. wide; pedicels 1.5-2.5 mm. long. 

S. obstipa. 
Pubescence somewhat spreading, the branches obvious 

(cf . some forms of S. guianensis) S. bifida. 

Inflorescence mostly about 1-1.5 cm. long, often openly 

pubescent, especially when larger. 
Leaves thin, slightly pubescent on the nerves above. 
Leaves entire, mostly under 5 cm. wide. .S. loretensis. 
Leaves more or less dentate, mostly more than 5 cm. 

wide. 
Pubescence minute, appressed; leaf reticulation 

faint S. thecaphora. 

Pubescence spreading; leaf reticulation conspicuous. 

S. harongifolia. 

Leaves firm or almost coriaceous, glabrous or nearly so 
above. 

Inflorescence greenish. 
Leaves of medium size. 

Pistillate sepals conspicuous; leaves conspicuously 

acuminate S. Poeppigii. 

Pistillate sepals minute or none; leaves acute or 

short-pointed 5. guianensis. 

Leaves small, 2-3 cm. wide S. calignosa. 

Inflorescences compactly yellow-gray-pubescent. 

S. obstipa. 
Petioles mostly longer, or at least half of them 1.5 cm. long or longer. 



794 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaf veins more or less reticulately bullate-impressed above; pubes- 
cence often abundant, even in immature leaves the character 
reasonably well marked (cf. S. harongifolia). 

Leaves to 7 (-10) cm. wide, mostly narrower or the pubescence 
beneath soft. 

Leaves glabrous S. umbelliflora. 

Leaves pubescent, at least beneath. 

Pubescence pale; leaves scarcely reticulate- veined beneath. 

S. Tulasnei. 
Pubescence often more or less distinctly fulvous; leaves 

always conspicuously reticulate-veined beneath. 
Pubescence soft and dense beneath. 

S. muricata, S. pyricarpa. 
Pubescence soon chiefly confined to the veins beneath. 

Pubescence distinctly fulvous S. neglecta. 

Pubescence not at all fulvous S. suaveolens. 

Leaves mostly (except the younger) 1 to several decimeters wide, 
the pubescence beneath never soft, but in S. pyricarpa 
dense. 

Leaves mostly 8-12 cm. wide. 
Stem pubescence hispid-stellate or fasciculate. 

Pistillate inflorescences short S. aspera. 

Pistillate inflorescences elongate S. exsculpta. 

Stem pubescence stellate-fasciculate but denser and shorter, 

thus tomentose S. pyricarpa. 

Leaves mostly 15-30 cm. wide. 
Leaves conspicuously serrate. 

Leaves cuneate at the base S. podocarpa. 

Leaves cordate at the base S. gigantophylla. 

Leaves entire or nearly so S. plana. 

Leaf veins not very obviously if at all impressed above, even in 

mature leaves, unless in S. Uleana, a nearly glabrous plant. 
Pubescence of the leaves beneath simple or rarely hispid-stellate 
or lepidote (except in S. Gilgiana), often extremely minute 
and mostly or entirely confined to the nerves, or even 
lacking. 

Leaves glabrous beneath or sometimes with a few (4-5) remote 
hairs. 



FLORA OF PERU 795 

Leaf width rarely 4.5 cm. 

Inflorescence dense, umbelliform S. umbelliflora. 

Inflorescence open, cymose S. Tayloriana. 

Leaf width mostly 6-12 cm. 

Leaves oblong-elliptic, about 6-7 cm. wide. .S. cuzcoana. 
Leaves broadly elliptic, about 10 cm. wide. 

Inflorescence glabrate; leaf margins denticulate. 

S. Uleana. 
Inflorescence rusty-stellate-lepidote; leaves entire. 

S. Williamsii. 
Leaves, at least on the midnerve, somewhat pubescent 

beneath. 
Pubescence not lepidote. 

Midrib and nerves minutely puberulent or hispidulous. 
Pubescence minute. 

Trichomes stellate S. Gilgiana. 

Trichomes simple S. macrotepala. 

Pubescence laxly hispid-stellate, partly simple. 

S. suaveolens. 
Midrib (especially) long-hispid. 

Petioles 2-5 cm. long. 

Inflorescence glabrate S. pseudoumbellata. 

Inflorescence pubescent S. lyrata. 

Petioles 1-1.5 cm. long S. hispida. 

Pubescence entirely lepidote S. decipiens. 

Pubescence of the leaves beneath stellate, dense, or at least dis- 
tributed over the surface. 

Pubescence of mature leaves gray or scarcely fulvous, fine, 
the stellate or fasciculate hairs distinct or merely approxi- 
mate; leaves never cordate. 
Leaves cuneate at the base. 

Leaf margins distinctly and closely dentate. 

S. harongifolia. 
Leaf margins entire or remotely or undulately denticulate. 

Leaves obviously pubescent above, at least on the 
nerves; inflorescence few-flowered. 



796 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves medium in size, to 15 cm. long and about 6 

cm. wide S. dasyantha. 

Leaves large, 30 cm. long and 15 cm. wide. 
Petioles densely fulvous-tomentose . .S. pauciflora. 
Petioles rather loosely gray-stellate-pubescent. 

S. tabacifolia. 
Leaves glabrate above, very minutely and sparsely 

puberulent; flowers many S. heteropoda. 

Leaves rounded-cuneate at the base. 
Leaf width exceeding 10 cm. 

Leaves dentate; stamens 6 S. eriocalyx. 

Leaves entire; stamens about 20 S. plana. 

Leaf width about 7 cm S. galbina. 

Pubescence of the leaves fulvous beneath and dense, i. e., the 
hairs more or less interwoven, or the leaves cordate at 
the base. 
Leaves minutely and equally or subequally dentate, or 

subentire. 

Leaves short-cuneate at the obtuse base. 
Petioles to about 2 cm. long, often shorter. . .S. fulva. 

Petioles to 3-4 cm. long S. radiata. 

Leaves rounded and somewhat cordate at the base. 

Leaves subentire, the pubescence becoming sparse. 

S. magnified. 
Leaves closely denticulate, the pubescence dense. 

S. muricata. 
Leaves coarsely and unequally dentate S. eriocalyx. 

Key after Perkins 

Flowers characteristically monoecious; styles more or less coherent 

in a column (so far as known). 
Petioles to 3 cm. long; leaves, or at least the sepals, densely 

lepidote S. decipiens. 

Petioles all short, rarely to 1.5 cm. long. 

Receptacles, often also the branchlets, lepidote-pubescent. 
Leaves large, at least some of them longer than 15 cm., 

short-acuminate. 
Leaves sparsely lepidote beneath S. lepidantha. 



FLORA OF PERU 797 

Leaves, at least on the nerves, stellate-puberulent. 

S. cervicornis. 
Leaves 8-15 cm. long, long-acuminate. 

Leaves mostly shorter than 9 cm. ; stamens 6 . S. crassiflora. 
Leaves mostly longer than 9 cm.; stamens 4-5. 

S. ucayaliensis. 
Receptacles or sepals not lepidote, at least not typically so 

(cf. S. cervicornis). 

Sepals (pistillate) well developed, the receptacle tomentose. 

S. Poeppigii. 
Sepals minute or wanting. 

Style column exserted S. guianensis. 

Style column short, included S. bifida. 

Flowers characteristically dioecious; styles free or very rarely 

coherent (so far as known). 
Stamens mostly 5, 4 of them more or less connate. 
Leaves rounded to cordate at the base; inflorescence short. 

S. magnified. 
Leaves narrowed at the base; inflorescence 1.5-3 cm. long. 

Leaves tomentose beneath S. radiata. 

Leaves lepidote-stellate beneath S. lepidiflora. 

Stamens often more numerous, always free. 

A. Stamens few, rarely as many as 20 (the number is unknown 
for a few species, apparently allied to the following). 

Stamens only 1 or 2 S. Gilgiana. 

Stamens 5-10, rarely to 20. 

Leaves soon glabrous or subglabrous beneath. 

Leaf margin entire, undulate, or obscurely denticulate. 
Leaves membranous. 

Leaves oblong-lanceolate, long-cuneate to the base. 
Petioles shorter than 1 cm. 

Leaves to 6 cm. wide; stamens 5-6. 

S. subinodora. 
Leaves to 3 cm. wide; stamens 17-18. 

S. parviflora. 

Petioles 3-6 cm. long S. cuzcoana. 

Leaves broader, rounded-cuneate at the base. 



798 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles 2-4 cm. long; leaves abruptly acute. 
Petioles 2-2.5 cm. long; blades with 10-12 lateral 

nerves, 10 cm. wide S. Williamsii. 

Petioles 2-4 cm. long; blades with 7-8 lateral 

nerves, 6-10 cm. wide S. macrotepala. 

Petioles 1-2 cm. long; blades about 7 cm. wide, 

acuminate S. thecaphora. 

Leaves coriaceous S. Tayloriana. 

Leaf margin obviously denticulate. 

Leaf texture coriaceous S. Tayloriana. 

Leaf texture scarcely firm, sometimes papyraceous. 
Leaves subcordate at the base, short-acuminate. 

S. Uleana. 
Leaves somewhat cuneate at the base, acuminate. 

S. thecaphora. 
Leaves, at least beneath, more or less pubescent. 

B. Pubescence hirsute-pilose, the hairs simple, branched, 

or long-stellate, or rarely lepidote-stellate. 
Leaves rather manifestly auricled at the base. 

Petioles 5-8 mm. long S. auriculata. 

Petioles 2-3 cm. long S. lyrata. 

Leaves not manifestly auricled. 
Leaves definitely rounded or cordate at the base. 

Leaves mostly only 6 cm. long S. ovalis. 

Leaves uniformly larger. 
Sepals large, 3-4 mm. long. 

Petioles 1.5-2.5 cm. long; pedicels 1 cm. long. 

S. harongifolia. 

Petioles 2-4 cm. long; pedicels 2-6 mm. long. 
Leaves not bullate above by the impressed 

veins. 
Pubescence minute; leaves not at all 

lyrate S. macrotepala. 

Pubescence pilose; leaves somewhat ly- 
rate at the base S. lyrata. 

Leaves bullate above by the impressed 
veins S. aspera. 



FLORA OF PERU 799 

Sepals minute or obsolete. 

Inflorescence characteristically short, 1-2 cm. 
long. 

Leaves ternate or scattered; receptacles 
subsessile S. asperula. 

Leaves opposite; pedicels 2-3 mm. long. 

S. obstipa. 

Inflorescence typically several centimeters 

long. 

Leaves more or less denticulate; stamens 
few. 

Leaves to 30 cm. wide, 21-26-nerved. 

S. gigantifolia. 

Leaves to 15 cm. wide, 16-18-nerved. 

S. exsculpta. 

Leaves entire; stamens about 20. .S. plana. 
Leaves cuneate to the base. 

Petioles mostly or all shorter than 15 mm. 
Pubescence mostly or entirely simple. 

Hairs spreading S. hispida. 

Hairs appressed S. parviflora. 

Pubescence more or less stellate or lepidote. 

Mature leaves glabrous above, merely lepidote 
beneath S. ucayaliensis. 

Mature leaves, if glabrate above, not merely 

lepidote beneath. 

Leaves membranous, slightly pubescent on 
the often many nerves above. 

Leaves entire, mostly narrower than 5 

cm S. loretensis. 

Leaves denticulate, usually wider. 

S. thecaphora. 

Leaves firm, chartaceous, glabrous or nearly 
so above, the lateral nerves few (7-9). 

Leaves to 7 cm. long S. calignosa. 

Leaves 9-16 cm. long S. obstipa. 

Petioles, at least part of them, longer than 15 mm. 



800 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves more or less bullate above by the im- 
pressed veins. 

Leaves ternate or scattered S. suaveolens. 

Leaves opposite S. Tulasnei. 

Leaves not bullate above, the veins little if at 

all impressed. 
Sepals connate or none. 
Leaves of medium size, glabrate, denticu- 
late S. thecaphora. 

Leaves large, stellate-puberulent, entire. 

S. plana. 
Sepals free. 
Leaves and receptacles minutely pilose. 

S. macrotepala. 

Leaves and receptacles more or less stellate- 
tomentose. 

Leaves glabrate above S. heteropoda. 

Leaves pubescent above. . .S. tabacifolia. 
B. Pubescence tomentose in quality, at least that of 
leaves and stems (this key contrast, after Perkins, 
not always easily interpreted). 

Leaves more or less coriaceous and noticeably bullate 
above by the impressed veins. 

Petioles to 1 cm. long S. Weberbaueri. 

Petioles, at least some of them, 1.5 cm. long, often 

longer. 
Leaves mostly 8-12 cm. wide. 

S. exsculpta, S. pyrocarpa. 
Leaves mostly 15-30 cm. wide. 

Leaves cuneate at the base S. podocarpa. 

Leaves cordate at the base S. gigantophylla. 

Leaves membranous, the veins little if at all impressed 

above. 
Leaves ternate. 

Petioles to 1 cm. long. 

Leaves sparsely tomentose S. asperula. 

Leaves densely tomentose S. tomentosa. 

Petioles to 3 cm. long S. galbina. 



FLORA OF PERU 801 

Leaves opposite. 
Leaves unequally and coarsely dentate. 

S. eriocalyx. 

Leaves minutely dentate or entire. 
Petioles to 2 cm. long; sepals connate, tomentose. 

S. fulva. 
Petioles to 3 cm. long; sepals free, glabrate. 

S. galbina. 
A. Stamens many (20) 30-60 (cf. S. plana). 

Glabrous shrub with umbellate inflorescence. .S. umbelliflora. 
Pubescent shrubs; fruit often long-muricate. 
Stamens about 30. 

Petioles to 1.5 cm. long S. saurauiifolia. 

Petioles 2-5 cm. long. 

Leaves about 10 cm. wide; sepals connate. S. pauciflora. 
Leaves 4-7.5 cm. wide; sepals free. .S. pseudoumbellata. 
Stamens 40-60. 
Pubescence soon confined to the leaf nerves beneath. 

S. neglecta. 

Pubescence dense on the leaves beneath. . . .S. muricata. 
Pubescence almost none; leaves small S. Mathewsii. 

Siparuna aspera (R. & P.) A. DC. in DC. Prodr. 16, pt. 2: 645. 
1868. S. calocarpa Perk. Repert. Nov. Sp. 1: 153. 1905. Citrosma 
aspera R. & P. ex Tul. Monogr. 325. 1855. 

A stout and heavy-leaved, rather coarsely rusty-pubescent shrub; 
petioles 1-2 cm. long; leaves opposite, to 25 cm. long, mostly sub- 
cordate at the rounded base, the margins undulate, finally glabrous 
above and more or less conspicuously bullate by the impressed nerves 
and veins; staminate inflorescence to 2.5 cm. long, the peduncles 1 
cm. long, the pedicels 3-6 mm. long; flowers 4 mm. broad; receptacles 
papery, glabrous within, twice as long as the 4 ovate, acute sepals; 
stamens 9; pistillate inflorescence 2-4-flowered, the peduncle 5-6 
mm. long, the pedicels half as long, strongly enlarged in fruit; flowers 

5 mm. broad, the sepals rotund; fruit 1-1.5 cm. thick, obovoid, 
unequally sericeous. Sleumer has indicated in herb. Madrid the 
identity of S. calocarpa and S. aspera. Weberbauer 115 was a tree 

6 meters high. 

Junin: Below Huacapistana, 1,400 meters, Weberbauer 2317, type 
of S. calocarpa. Colonia Peren (Weberbauer 115). Huanuco: Chin- 



802 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

chao, Muna, and Pillao, Ruiz & Pavon. Huanuco to Pampayacu, 
Kanehira 235. Without locality, Ruiz & Pavon 24-40. Pampayacu, 
Kanehira 58; a shrub. Cueva Grande, 1,050 meters, sunny, brushy 
slopes, 4760; a shrub or small tree; flowers greenish yellow; fruits 
reddish. 

Siparuna asperula (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 647. 
1868; 100. Citrosma asperula Tul. Ann. Sci. Nat. III. 4: 35. 1855. 
C. amazonum Tul. Monogr. 332. 1855. 

A tree, the ternate or scattered leaves slightly asperous; branch- 
lets and petioles (about 1 cm. long) rather densely pale rusty-pubes- 
cent; blades often rounded and unequal at the base, nearly 10 cm. 
long and half as wide; staminate and pistillate receptacles fasciculate, 
subsessile, the velum ashy-pubescent, the 5-6 minute sepals unequal ; 
stamens 5-8; styles many, exserted. Neg. 13479. 

Junin: Tocache, Poeppig 1961. Loreto: Rio Putumayo, in forest, 
Klug 1641; a shrub of 3 meters; flowers white; fruit red. 

Siparuna auriculata A. DC. Journ. Bot. 3: 219. 1865; 94. 

Well marked by the basally auriculate leaves, abruptly narrowed 
to the base, remotely denticulate and, especially beneath, densely 
long-hispid, the branches similarly pubescent; petioles 5-8 mm. 
long, the blades about 20 cm. long and half as wide; pistillate in- 
florescence very short; fruit rusty-hispid, crowned by the unequal, 
oblong sepals. Compare S. Uleana and S. hispida. S. grandiflora 
(HBK.) Perk., Colombian, lacks, apparently, the leaf auricles. 
Neg. 13480. 

San Martin: Tarapoto, Spruce 4369, type. San Roque, 1,400 
meters, in forest, Williams 7076, 7573; a large or small shrub. 

Siparuna bifida (Poepp. & Endl.) A. DC. in DC. Prodr. 16, 
pt. 2: 652. 1868; 114. Citrosma bifida Poepp. & Endl. Nov. Gen. 
& Sp. 2: 48. 1838. 

Similar to S. guianensis, but the longer inflorescences racemose 
or the branches racemose and, particularly, the styles free or united 
below and reflexed at the tip. The leaves may be as long as 16 cm., 
as wide as 5.5 cm., their abrupt, caudate acuminations to 1.5 cm. 
long; fruit loosely villous, subechinate. Perkins, from Bolivian 
material, has described the staminate inflorescences as 2.5 cm. long, 
the pedicels 1-2 mm. long, the flowers to 1 mm. wide, papyraceous; 
stamens about 12, free, little exserted, the plane filaments glabrous. 
Neg. 8087. 



FLORA OF PERU 803 

Loreto: Yurimaguas, Poeppig 2131B, type; in or at edge of forest, 
200 meters, Williams 4719, 7829; a shrub. Lower Rio Nanay, in 
forest, Williams 642. Iquitos, 120 meters, edge of forest, Williams 
3539; Ule 6905 (det. Perkins). Yarina-cocha, Tessmann 5479. 
Bolivia; Amazonian Brazil. 

Siparuna calignosa Macbr. Candollea 5: 353. 1934. 

A tree of 4.5-5.5 meters, the somewhat angulate branchlets 
at first slightly scabrous-stellate; petioles 4-7 mm. long, densely 
short-pubescent with brownish green, stellate hairs; blades elliptic, 
becoming 7 cm. long and 3.5 cm. wide, acute at the base, very short- 
acuminate, minutely denticulate or entire, glabrous above except 
the obscurely strigillose nerves, sparsely and minutely stellate- 
pubescent on the nerves beneath, the lateral nerves about 7, the veins 
scarcely obvious; racemes slender, axillary, to 15 cm. long; pedicels 
1-2 mm. long; receptacles depressed-conic, 3 mm. thick, 2 mm. 
high, the mouth small; sepals minute, glabrous; stamens 4, free. 
Apparently resembling S. hylophila Perk., with much longer peti- 
oles, or S. guianensis Aubl., with larger leaves and many stamens. 

Loreto: Iquitos, in forest, 100 meters, Killip & Smith 27069, 
type. 

Siparuna cervicornis Perk. Verh. Bot. Ver. Brandenb. 47: 
145. 1905. 

A small tree, about 8 meters high, the pubescence of the branches 
and leaf nerves minutely and compactly stellate; petioles 1.5 cm. 
long; blades 15-20 cm. long, 6.5-7.5 cm. wide, chartaceous, cuneate 
at the base, acute at the apex, the lateral nerves 10-14; inflorescence 
racemose, many-flowered, to 4 cm. long; staminate flowers 1-1.5 mm. 
broad, the pedicels 1 mm. long; receptacles papery, broadly open, 
the sepals and velum lacking; stamens 11 or 12, slightly exserted. 
The type has lepidote-stellate pubescence. The Brazilian S. sarmen- 
tosa Perk. Nachtr. 56, is glabrate, the broader staminate flowers with 
about 15 exserted stamens, the pedicels 1-2 cm. long. Neg. 13487. 

Loreto: La Victoria, in forest, Williams 2706. Mouth of Rio 
Santiago, Tessmann 4395. Rio Acre: Krukoff 5270; Ule 9391. 
Brazil. 

Siparuna crassiflora Perk. Bot. Jahrb. 28: 702. 1901; 112. 

Doubtfully distinct from S. ucayaliensis, but typically differing 
by the somewhat smaller leaves (6.5-9 cm. long, 2.5 cm. wide), the 
flowers in pseudo-umbels less than 1 cm. long, the pedicels 7 mm. 



804 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, the stamens 6. Illustrated, Pflanzenreich IV. 101: /. 23. 
Brazilian specimens referred here have longer leaves and more stel- 
late scales; compare also the apparently similar S. lepidiflora. 
Neg. 13492. 

San Martin: Tarapoto, Spruce 4907. Loreto: Tessmann 3191. 
Amazonian Brazil. 

Siparuna cuzcoana Perk. Bot. Jahrb. 45: 461. 1911; Nachtr. 47. 

A green shrub, the sparse pubescence obvious only on the young 
parts; leaves ternate or verticillate, 5-20 cm. long, 2-6.5 cm. wide, 
cuneately attenuate to the (3-6 cm. long) petiole, acute or short- 
acuminate at the apex, thin, obscurely undulate-dentate, sometimes 
with a few stellate hairs beneath, even in age, on the laxly reticulate 
veins; staminate inflorescence 1-1.5 cm. long, cymose, the peduncle 
about 5 mm. long, the pedicels 6 mm. long; receptacles fleshy, 
sparsely stellate-pubescent outside, the subequal sepals glabrous; 
stamens 15, unequal, glabrous. A shrub of 4 meters with citrus- 
yellow flowers, at first greenish, 6 mm. wide. Neg. 13493. 

Cuzco: Provincia de Convencion, 1,400 meters, Weberbauer 5042 , 
type. 

Siparuna dasyantha Perk. Notizbl. Bot. Gart. Berlin 6: 134. 
1914. 

Related to S. loretensis Perk., but the branchlets, petioles (1-2.5 
cm. long), and blades, the latter especially on the nerves of both sides, 
rusty-stellate-pilose; blades thin, ovate-oblong, 11-14.5 cm. long, 
4-5.5 cm. wide, attenuate to the base, shortly and broadly acuminate 
at the apex, the tip obtuse, the lateral nerves 8-9; inflorescence 
subsessile, axillary, few-flowered, cymose; staminate flowers 2 mm. 
broad, pedicellate; receptacle thin, pilose only outside; stamens 6. 
A shrub of 2-7 meters. Neg. 13494. 

Rio Acre: Ule 9392, type. 

Siparuna decipiens (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 643. 
1868; 110. Citrosma decipiens Tul. Monogr. 368. 1855. 

Closely related to S. lepidantha, but more or less yellow-lepidote 
throughout, only the older leaves glabrate above; petioles to 3 cm. 
long; inflorescence 4-6 cm. long; stamens 4-6, included; receptacles 
ventricose below, constricted beneath the throat, the sepals connate 
into a tube; velum a cylindric tube, the aperture nearly a slit; styles 
united. Illustrated, Pflanzenreich IV. 101 :/. 26. Neg. 27573. 

Rio Acre: Ule 9623. Brazil; French Guiana. 



FLORA OF PERU 805 

Siparuna eriocalyx (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 646. 
1868; 102. Citrosma eriocalyx Tul. Ann. Sci. Nat. III. 4: 35. 1855. 

Branches, petioles, and inflorescence densely yellowish-stellate- 
tomentose; petioles 3-6 cm. long; blades almost caudate-acuminate, 
conspicuously and more or less irregularly dentate, the teeth often 
glandular-calloused, sparsely hispidulous above and on the midnerve 
stellate-tomentose, . beneath evenly stellate-hispidulous, scarcely 
venose above, slightly so beneath, 10-20 cm. long, 10-12 cm. wide 
(the younger leaves half as large) ; inflorescence about 1 cm. long, the 
pedicels 4-5 mm. long; receptacle globose-urceolate; sepals 4-6, 
large, rotund, connate at the base, in the pistillate flowers reflexing; 
stamens 6, exserted, the styles 10-12, free. The Ecuadorean S. apici- 
fera (Tul.) A. DC., 105, may have a longer inflorescence, and the 
fruit is said to be calloused or aculeate; compare also S. pyricarpa. 
Very near, also, is S. gesnerioides (HBK.) A. DC., 102, of Ecuador, 
with minutely sinuate-denticulate leaves. Neg. 34793. 

Huanuco: Chinchao, Dombey, type; River o. 

Siparuna exsculpta Perk. Verh. Bot. Ver. Brandenb. 47: 144. 
1905; Nachtr. 54. 

A shrub or tree of 2.5-4.5 meters; resembling S. aspera and S. pyri- 
carpa, but the leaves sometimes thinner and crenulate-denticulate, 
the pistillate inflorescence to 3 cm. long, the pedicels to 1 cm. long. 
Negs. 13498, 27575. 

San Martin: Tarapoto, Ule 6831, type. Junin: Colonia Perene", 
in forest, 680 meters, Killip & Smith 25027, 25017. 

Siparuna fulva A. DC. Journ. Bot. 3: 219. 1865; 103. 

Said to be related to S. eriocalyx; whole plant velvety- tomentose, 
even the leaves above, the pubescence, especially on the branches, 
bright brown; leaves 9-15 cm. long, 5-7 cm. wide, acutely short- 
acuminate, venose only beneath, the lateral nerves 10-13, obscurely 
reticulate-veined, entire or remotely calloused; sepals 4-5, connate 
at the base, ovate-rotund; styles exserted; flowers 3-4, on pedicels 
5 mm. long, these in fruit to 2 cm. long, the fruit 1 cm. thick. 

San Martin: Tarapoto, Spruce 4361, type. 

Siparuna galbina Macbr. Candollea 5: 354. 1934. 

A shrub of 3-4.5 meters, the young branchlets, petioles, and leaves 
beneath softly stellate-pubescent; petioles opposite or sometimes 
ternate, to 3 cm. long; blades pale yellowish green, chartaceous, 
sparsely stellate-scabrous above on the prominent nerves, these 



806 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

about 8, reticulate-veined beneath, marginally undulate or minutely 
and remotely denticulate, broadly ovate-elliptic, rounded or broadly 
cuneate-rounded at the base, subacute at the apex, finally about 
13 cm. long and 7 cm. wide; inflorescence axillary, densely stellate- 
pubescent, about 1 cm. long; pedicels 3-5 mm. long; receptacle ovoid, 
nearly 4 mm. thick; sepals 5-6, glabrate, 1 mm. thick, 2 mm. broad; 
styles many, conspicuously exserted, free, filiform, revolute. 
Flowers orange-green. 

Ayacucho: Carrapa, 1,000 meters, wooded hillside, Killip & 
Smith 22453, type. 

Siparuna gigantophylla Perk. Bot. Jahrb. 54: Beibl. 118: 2. 
1916. 

Like S. podocarpa, but the petioles 4-13 cm. long; blades 40 cm. 
long or larger and 35 cm. wide, cordate at the base, densely hispid 
above, unequally serrate, 21-26-nerved; staminate flowers 5-10 mm. 
broad, the pedicels 2-3 mm. long, densely stellate-pilose, in panicles 
7.5 cm. long; sepals ovate, unequal; stamens 6-7. 

Cajamarca: Tabaconas, 1,900 meters, Weberbauer 6242, type. 

Siparuna Gilgiana Perk. Notizbl. Bot. Gart. Berlin 10: 161. 
1927. 

A small tree, the young branchlets, petioles (1.5-4 cm. long), 
and staminate receptacles outside stellate-pilose; leaves usually 
elliptic, 7.5-15.5 cm. long, 3-8 cm. wide, acuminate, cuneate-rounded 
at the base, undulate-dentate, papyraceous, glabrous except the 
nerves (7-8), which are minutely pilose beneath; cymes 1-1.5 cm. 
long, 3-6-flowered, the pedicels 1 mm. long; stamens 2, free. 
Related to S. Mouraei Perk., 86. Compare S. thecaphora, the number 
of its stamens unknown, to which part of the following material may 
belong. The Ecuadorean S. Eggersii Hieron., 90, has broader leaves, 
staminate pedicels 5 mm. long, and 7-8 nearly included stamens. 
Neg. 13500. 

Loreto : Pongo de Manseriche, Tessmann 4464, type. La Victoria, 
Williams 2798. Yurimaguas, Williams 4262. Iquitos, Williams 
1405; Killip & Smith 27243, 27227. Santa Rosa, Williams 4946. 
Puerto Arturo, Williams 5363. Junin: Vitoc, Ruiz & Pavon (det. 
Sleumer). Without locality: Ruiz & Pavon 24-32. 

Siparuna guianensis Aubl. PI. Guian. 2: 865. pi. 333. 1775; 113. 

Related to S. Poeppigii, but sepals of the pistillate flowers reduced 

and the styles long-connate, long-exserted ; often reddish-tomentose 



FLORA OF PERU 807 

on the younger parts, but glabrate in age; pedicels 3-6 mm. long; 
sepals 4-6, minute, suborbicular; stamens 10-12, long-exserted, un- 
equal ; fruit yellow. Illustrated, Pflanzenreich IV. 101 : /. 25, 27. 
A shrub or small tree. 

San Martin: Tarapoto, 400-900 meters, Williams 5626, 6674, 
5385. Moyobamba (Weberbauer, 289). San Roque, Williams 7309. 
Lamas, 840 meters, Williams 6524- Loreto: Iquitos, open land, 100 
meters, Williams 1510, 7949; Killip & Smith 26097. Mishuyacu, 
100 meters, Killip & Smith 29874; King 85, 2546. Trinidad to 
Colombia, Bolivia, and northern Brazil. "Isula micuna," "isula 
caspi," "curuinsi-sacha." 

Siparuna harongifolia Perk. Bot. Jahrb. 28: 686. 1901; 96. 

Resembling S. thecaphora, but the leaves pubescent beneath, 
sometimes 25 cm. long and 15 cm. wide, and the inflorescence to 
2.5 cm. long; staminate flowers 5 mm. broad; sepals 4, large, rusty- 
pilose, especially on the margins; stamens 6, exserted, unequal, the 
medial ones much smaller than the outer; fruit about 12 mm. thick. 
Neg. 13504. 

San Martin: Tarapoto, Spruce 4553. Puno: Sangaban, Lechler 
2362. Vitoc, Ruiz & Pavon 24-31 (det. Sleumer). Cuzco: Machu- 
picchu, 2,200 meters, Herrera 3206, 3211, 3205. 

Siparuna heteropoda Perk. Notizbl. Bot. Gart. Berlin 6: 
135. 1914. 

Branches 4-sided, the youngest brown-stellate-pilose as also the 
very unequal petioles (1.5-7 cm. long), and to a less extent the thin, 
obscurely undulate-dentate blades; blades oblong-obovate, cuneateat 
the base, obtusely acuminate at the apex, 12.5-22.5 cm. long, 4.5-9 
cm. wide, the lateral nerves 11-12; cymes 3 cm. long, subsessile; 
staminate flowers reddish yellow, pedicellate; receptacles stellate- 
pilose outside, fleshy; stamens 9. As much as 10 meters high. 
Compared by the author with S. riparia (Tul.) A. DC., with fewer 
flowers and dentate leaves. Neg. 13505. 

Rio Acre: Ule 9393, type. 

Siparuna hispida A. DC. Journ. Bot. 3: 219. 1865; 98. 

Branches and foliage densely long-hispid with brownish, mostly 
simple hairs; blades 5-13 cm. long, 3-5 cm. wide, narrowly acuminate, 
crenulate-dentate; petioles 10-15 cm. long; pistillate inflorescence 
very short, subsessile, the pedicels 3 mm. long, the flowers little 
longer; receptacle densely tomentose, the 5 ovate sepals glabrate 



808 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

within; styles 6-8, evidently exceeding the fleshy velum; fruit hispid, 
1 cm. long. Neg. 8090. 

San Martin: Tarapoto, Spruce 4243, type. Bolivia. 

Siparuna lepidantha Perk. Bot. Jahrb. 28: 701. 1901; 111. 

A shrub or tree, sometimes 12 meters high, monoecious, the 
young branchlets, petioles, leaf nerves beneath, and flowers more 
or less densely lepidote; leaves opposite, the blades obovate-oblong, 
10-20 cm. long, 5.5-9 cm. broad, broadly short-acuminate, rounded- 
cuneate at the base, entire, chartaceous, glabrous above; petioles 
1-1.5 cm. long. Neg. 13509. 

Loreto: Mainas, Poeppig 2424, type. Yurimaguas, 135 meters, 
in forest, Killip & Smith 28242. Iquitos, 100 meters, in forest, 
King 838, 846, 237. 

Siparuna lepidiflora Perk. Notizbl. Bot. Gart. Berlin 10: 
166. 1927. 

Youngest branchlets, petioles, leaves beneath, and inflorescence 
gray-lepidote; leaves opposite, the blades oblong-obovate, 7.5-15.5 
cm. long, 3.5-6 cm. wide, papyraceous, entire, at maturity glabrous, 
the lateral nerves 5-6; inflorescence 1.5-2.5 cm. long, cymose, sub- 
sessile, the pedicels 5-10 mm. long; staminate flowers 4-5 mm. broad, 
lepidote; receptacle campanulate, broadly open; stamens 4-5. 
As much as 10 meters high, the trunk 13 cm. in diameter. Neg. 
13510. 

Loreto: Soledad, lower Rio Itaya, in upland forest, Tessmann 
5283, 5326. Mishuyacu, 100 meters, in forest, King 1504; a tree of 
6 meters. 

Siparuna loretensis Perk. Verh. Bot. Ver. Brandenb. 47: 143. 
1905; Nachtr. 51. 

A green shrub, the branches and leaves very sparsely rusty- 
stellate-pubescent; petioles 6-7 mm. long, densely pubescent; blades 
oblong-lanceolate, to 15 cm. long, acuminate, acute at the base, 
with about 30 nerves; inflorescence rarely 1 cm. long; flowers brown- 
ish, 3 mm. broad, the pedicels to 3 mm. long; receptacle membranous, 
glabrous within; sepals connate; stamens 6, subequal. Neg. 13511. 

Loreto: Cerro de Escalera, 1,200 meters, Ule 6829, type. 

Siparuna lyrata Perk. Verh. Bot. Ver. Brandenb. 47: 142. 1905; 
Nachtr. 50. 

A slightly pubescent shrub, sometimes 8 meters high, the 
branches, petioles, leaves beneath, and receptacles pale fasciculate- 



FLORA OF PERU 809 

hispid-stellate; petioles 2-3 cm. long; blades gradually attenuate 
toward the somewhat lyrately enlarged base, to 30 cm. long and 10 
cm. wide, the lateral nerves 18-20; inflorescence short-paniculate, 
to 2 cm. long; staminate flowers straw-colored, 2-6 mm. broad, the 
pedicels to 5 mm. long; receptacle glabrous within, papery, hirsute, 
one-third longer than the (3-4 mm. long) punctate sepals; stamens 
6. Resembling S. auriculata, from which it differs in its branched 
pubescence and longer petioles, but nearly related to the Colombian 
S. grandiflora (HBK.) Perk., 95. Neg. 13512. 

Loreto: Pampas de Ponasa, 1,100 meters, Ule 6832, type. 

Siparuna macrotepala Perk. Verh. Bot. Ver. Brandenb. 47: 
140. 1905; Nachtr. 48. 

A citrous-scented shrub with rusty-pilose branches and sparsely 
and minutely pilose leaves and receptacles; leaves opposite, the 
blades elliptic or broadly oblong, cuneate-rounded at the base, 
broadly acuminate at the apex, 10-20 cm. long, 6.5-10 cm. wide, 
thin, subentire or very minutely serrulate, the lateral nerves 7-8; 
petioles 2-4 cm. long; inflorescence cymose, 3-3.5 cm. long; pistillate 
flowers 2 mm. broad, the pedicels 3-4 mm. long; receptacle coria- 
ceous, papery, the lanceolate sepals over 4 mm. long, glabrous above; 
styles about 7. A shrub 2.5 meters high, with yellow flowers. 
Neg. 13513. 

Loreto: Pampas de Ponasa, Ule 6833, type. La Victoria, edge of 
forest, Williams 2763. Balsapuerto, 220 meters, in forest, King 2924. 

Siparuna magnifica Perk. Bot. Jahrb. 28: 699. 1901; 108. 

A shrub or tree of 5 meters or less, the mature branchlets glabrate; 
petioles 1-8 cm. long; leaves opposite, chartaceous, the blades obo- 
vate-oblong, 20-30 cm. long, 10-15 cm. wide, shortly and broadly 
acuminate, faintly undulate or dentate, at first rusty-tomentose 
on both sides, the pubescence persisting on the nerves; flowers 
crowded, the pedicels only 2 mm. long, the flowers 3 mm. broad, 
the broadly ovate receptacle loosely stellate-pubescent; sepals 
reduced to a ring; stamens 5, the outer connate into a tube. The 
Colombian S. mollis (HBK.) DC. has denser, more grayish pubes- 
cence, especially on the receptacles, and if not distinct specifically, 
is the earlier name. Neg. 13514. 

Loreto: Yurimaguas, Poeppig 2145, type; Killip & Smith 28742, 
28363. Pebas, in forest, Williams 1783, 1715. Mishuyacu, 100 
meters, Klug 531. San Martin: Zepelacio, 1,100 meters, in forest, 
Klug 3760; flowers violet and white. 



810 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 



Siparuna Mathewsii Perk. Bot. Jahrb. 28: 680. 1901; 91. 

Glabrous except the mostly alternate leaves, these slightly pubes- 
cent beneath with simple hairs; petioles 1-1.5 cm. long; blades ovate- 
oblong, acutish, cuneate-rounded at the base, 2.5-3 cm. wide, 
minutely denticulate above the middle; inflorescence 2 cm. long, 
the peduncle 1 cm. long, the pedicels half as long; staminate flowers 
5 mm. broad; receptacle subglobose, papery, glabrous, the sepals 
obsolete; stamens 40-50, the outer much larger. Illustrated, 
Pflanzenreich IV. 101 :/. 23 (receptacle). Neg. 13515. 

Peru: Without locality (Mathews 1560, type). 

Siparuna muricata (R. & P.) A. DC. in DC. Prodr. 16, pt. 2: 
651. 1868; 106. Citrosma muricata R. & P. Prodr. 123. pi. 29. 1794; 
Syst. 1: 265. 1798. 

A shrub or tree as much as 9 meters high; marked among species 
with leaves bullate-nervose at maturity by the abundant and soft, 
fulvous pubescence on the branches, petioles, and leaves beneath; 
petioles 2.5-3 cm. long, leaves 3-4-verticillate, the blades oblong- 
lanceolate, to 30 cm. long and 3-5 (-8) cm. wide, acute, very mi- 
nutely but densely serrate; staminate inflorescence crowded; recep- 
tacles at first cup-shaped, later ruptured and spreading, the 4 sepals 
becoming obsolete; stamens 60; pistillate inflorescence 2-3 cm. long, 
the pedicels 5-8 mm. long, densely fleshy-tuberculate, the 4-6 sepals 
as long; styles few. Illustrated, Pflanzenreich IV. 101: /. 23, 24- 
The younger leaves are not bullate. The Ecuadorean S. buddleiifolia 
(Benth.) A. DC., 106, has opposite leaves and about 20 stamens. 
Neg. 27576. 

Huanuco: Macora, Muna, and Acomayo, also Pillao, Ruiz & 
Pavon, type. Cochero, Ruiz & Pavon. Without locality, Ruiz & 
Pawn 24-36, 24-33, 24-35. Panao, 2,700 meters, in thicket, 3623. 
Twelve miles south of Panao, 3,000 meters, in dense thicket, 2202. 
Pampayacu, Kanehira 234, 65. Without locality: Dombey. Junin: 
Chanchamayo, Isern 2384. 

Siparuna neglecta (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 647. 
1868; 105. Citrosma neglecta Tul. Ann. Sci. Nat. III. 4: 36. 1855. 

Resembling S. muricata, but much less pubescent, the thick, 
often opposite leaves glabrous above; petioles 1-2 cm. long; inflores- 
cence 2.5-4 cm. long, axillary, the peduncle 2 cm. long, the pedicels 
1 cm. long; staminate sepals 4-6, scarcely equal, spreading; stamens 
40-50, not at all exserted; velum thick, the aperture small; drupes 
1 cm. thick, densely long-muricate. Perkins, Notizbl. Bot. Gart. 



FLORA OF PERU 811 

Berlin 10: 166, placed this species between numbers 62-66 of her 
monograph. Neg. 13523. 

Junin: Lateral valley of Rio Comas, 2,800 meters, Weberbauer 
6620, 6608. Ecuador. 

Siparuna obstipa Macbr. Candollea 5: 354. 1934. 

A shrub, or a tree of 5 meters; branchlets at first minutely and 
sparsely stellate-pubescent, compressed at the nodes, 4 mm. broad; 
petioles opposite, 5-7 mm. long; blades thin-chartaceous, glabrous 
and little if at all lustrous above, minutely stellate-pubescent on 
the nerves beneath or glabrate, elliptic or oblong-elliptic, rounded- 
acute at the base, subcaudate-acuminate and acute at the apex, 
9-16 cm. long, 3-5 cm. wide, the lateral nerves about 9, inconspicu- 
ous on both sides, the veins obscure; staminate racemes axillary, 
1-1.5 cm. long, solitary or geminate, densely but minutely grayish- 
yellow-stellate-pubescent; pedicels to 2 (2.5) mm. long; receptacle 
ovoid, 2 mm. thick; sepals and velum none; stamens 15, very unequal, 
the 10 outer ones well exserted and apparently bent to one side, the 
inner (sterile?) much shorter. The exact nature of the 5 inner sta- 
mens(?) is not clear to me. A small tree with the aspect of S. cervi- 
cornis Perk. 

Loreto: Iquitos, in forest, 120 meters, Williams 3790, type. 
Mishuyacu, 100 meters, in forest, Klug 320. 

Siparuna ovalis (R. & P.) A. DC. in DC. Prodr. 16, pt. 2: 650. 
1868; 95. Citrosma ovalis R. & P. Syst. 1: 266. 1798. 

Branches rusty-tomentose; leaves mostly ternate, the blades 
oblong-elliptic, mostly to 6 cm. long and 3 cm. wide, scarcely acute, 
often subcordate at the base, minutely hispidulous on the nerves, 
especially beneath, where also minutely puberulent, soon glabrate 
above; petioles about 5 mm. long; staminate inflorescence to 1 cm. 
long, usually sessile, the pedicels 4-8 mm. long; flowers 2.5 mm. 
broad; receptacle little pubescent, about as long as the 4-6 suberect 
sepals; stamens 6-8. Neg. 8542. 

Huanuco: Mufia, Sarriapata, Playa, Tambo Nuevo, Ruiz & 
Pawn 906, type. Without locality, Ruiz & Pawn 24-38. 

Siparuna parviflora Perk. Verh. Bot. Ver. Brandenb. 47: 141. 
1905; Nachtr. 48. 

A low or tall, sparsely rusty-pilose or glabrate shrub; leaves 
opposite, the blades suboblong, 2.5-3 cm. wide, slightly narrowed 
to the obtuse apex, but more so to the (4-6 mm. long) petioles, thin, 



812 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

entire, the lateral nerves 5-6; inflorescence to 1.5 cm. long, 1-2- 
flowered, the flowers 3 mm. broad, on pedicels 3-4 mm. long; recep- 
tacles slightly pilose outside only; sepals glabrous, connate; stamens 
17-18. 12a. Ule 6584a, presumably the pistillate plant, has flowers 
4 mm. wide, minute, oval sepals, exserted, free styles, and a yellow- 
ish, subglobose drupe 8 mm. thick. The name was published both 
as parvifolia and as above (in the remarks), and maintained as "parvi- 
folia" in the Nachtrdge. Neg. 13525. 

Loreto: Cerro de Escalera, 1,100 meters, Ule 6584, type; 6584a. 

Siparuna pauciflora (Beurl.) A. DC. in DC. Prodr. 16, pt. 2: 
656. 1868; 105. Citrosma pauciflora Beurl. Vet. Akad. Handl. Stockh. 
1854; 144. 1856. 

Closely related to S. neglecta, but the nerves of the much larger 
leaves not at all or not obviously impressed above, sparsely pubes- 
cent, the scarcely exserted stamens fewer (20-30). Illustrated, 
Pflanzenreich IV. 101: /. 28. Neg. 13526. 

Loreto: Balsapuerto, Killip & Smith 28527, 28485(1). Leticia, 
Ule 6217 (det. Perkins). Extending to Colombia and Costa Rica. 

Siparuna Perkinsiana Standl., sp. nov. 

Arbuscula 4-metralis, ramulis crassis subteretibus in sicco fuscis 
fere glabris, internodiis ut videtur brevibus; folia breviter petiolata 
magna opposita herbacea, in sicco fusca, utrinque plus minusve lucida, 
petiolo crassiusculo fere glabro; lamina obovato-elliptica 20-26 cm. 
longa 9.5-11 cm. lata, abrupte longiuscule acuminata, infra medium 
subabrupte angustata et basin versus attenuata, basi ipsa sublyrate 
dilatata et utroque latere subauriculata, saltern e medio usque apicem 
grosse remote sinuato-serrata, versus basin vulgo integra, supra 
glabra, subbullata, nervis perspicuis crassis, venis laxe reticulatis 
subimpressis, subtus concolor glabra, costa gracili elevata, nervis late- 
ralibus utroque latere ca. 11 angulo lato divergentibus arcuatis; inflo- 
rescentia laxe pauciflora vix petiolo longior glabra vel glabrata. The 
leaves are similar to those of S. lyrata and S. auriculata, to which 
the plant undoubtedly is closely related, but both those species have 
abundant pubescence on all or most parts of the plant. The type 
specimen is in very unsatisfactory condition as regards the flowers, 
which are so much deteriorated that they cannot be dissected 
satisfactorily. 

San Martin: Zepelacio, near Moyobamba, 1,100-1,200 meters, 
in forest, King 3352 (type in Herb. Field Mus.) ; flowers red-brown. 



FLORA OF PERU 813 

Siparuna plana Macbr. Candollea 5: 355. 1934. 

Monoecious, the branchlets and petioles at first densely rusty- 
stellate-puberulent, the branchlets compressed at the nodes and 
1 cm. thick; petioles opposite, 12-22 mm. long; blades large, often 15 
cm. wide, nearly 30 cm. long, broadly ovate-elliptic, rounded or 
broadly cuneate-rounded at the base, acuminate at the apex, entire 
or somewhat undulate, glabrous and slightly lustrous above, or the 
impressed nerves stellate-pubescent, the nerves and laxly reticulate 
veins prominent and sparsely stellate-pubescent beneath; panicles 
axillary, about 5 cm. long, densely stellate-pubescent; staminate 
receptacles mostly terminal and fasciculate, flat, cupular, about 
4 mm. broad, the pedicels 5-7 mm. long; sepals and velum none; 
stamens about 20; pistillate receptacle sessile, the sepals apparently 
minute. Young and perhaps not correctly interpreted ; the staminate 
receptacles are extremely shallow, and the relationship apparently 
is with S. guianensis Aubl. 

Loreto: San Ramon, Yurimaguas, 200 meters, in forest, Williams 
4584, type. "Isula micunan." 

Siparuna podocarpa Perk. Bot. Jahrb. 54: Beibl. 118: 2. 1916. 

A shrub with yellowish-stellate-pilose branchlets (4-sided), recep- 
tacles, and fruits; petioles (3.5-6.5 cm. long), pedicels, and blades 
above sparsely pilose, the blades velvety-tomentose beneath, coria- 
ceous, crenulate-dentate, 25-35 cm. long and 10-15 cm. wide, the 
nerves (lateral 16-19) and veins impressed above, the leaf surface 
bullate; pistillate flowers 4-7 mm. wide, on pedicels 4-12 mm. long, 
in many-flowered, subumbellate cymes, the peduncles 1-1.5 cm. long; 
receptacle coriaceous; styles about 10; fruit obpyriform, the pedicels 
3.5-5.5 cm. long. Neg. 13528. 

Cajamarca: Tabaconas, 1,900 meters, Weberbauer 6240, type. 

Siparuna Poeppigii (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 653. 
1868; 112. Citrosma Poeppigii Tul. Ann. Sci. Nat. III. 4: 39. 1855. 

A shrub or small tree, probably monoecious, the young branch- 
lets sparsely lepidote-stellate; leaves opposite, about 20 cm. long 
and 6-9 cm. wide, acutely acuminate, rounded at the base, entire, 
sparsely appressed-stellate-pubescent beneath, glabrous above; pistil- 
late inflorescence shorter than the (1-1.5 cm. long) petioles, more 
or less densely yellowish-stellate-pubescent, the 4-8 flowers sub- 
sessile; receptacle obconic, tomentose, equaled by the 4 erect, broadly 
ovate sepals; velum conic; styles exserted in a column. Illustrated, 
Pflanzenreich IV. 101 : /. 24. Neg. 8086. 



814 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Yurimaguas, Poeppig D2060, 2066, type; at 200 meters, 
Williams 4721, 4141, 3833; Killip & Smith 27988, 28826, 28696, 
27943. Iquitos, 100 meters, in forest, Killip & Smith 27215 
San Martin: Juanjui, 400-800 meters, in forest, King 4400, 3777; 
flowers green. 

Siparuna pseudoumbellata Perk. Bot. Jahrb. 54: Beibl. 118: 
1. 1916. 

A shrub 3 meters high, the branchlets and foliage more or less 
laxly pubescent throughout with long, yellowish hairs; petioles 2-5 
cm. long; blades opposite, oblong-obovate, 9-18.5 cm. long, 4-7.5 
cm. wide, short-acuminate, cuneate at the base, unequally serrate, 
the lateral nerves 11-13; staminate flowers 7 mm. broad, greenish, 
in subumbellate, many-flowered cymes, the pedicels 3-8 mm. long, 
nearly glabrous; receptacle usually glabrous; sepals 4-6, glabrous, 
equal; stamens about 30. "Near S. Tonduziana Perk." S. lyrata 
might be sought here. 

Piura: Above Palambla, 1,450 meters, Weberbauer 6031, type. 

Siparuna pyricarpa (R. & P.) Perk. Bot. Jahrb. 28: 690. 1901; 
100. Citrosma pyricarpa R. & P. Syst. 1: 264. 1798. C. limoniodora 
R. & P. ex Tul. Monogr. 330. 1855. 

A large-leafed, dioecious tree, the younger branches softly and 
thickly pubescent, as also the petioles and blades beneath, the petioles 
to 2 cm. long; blades abruptly short-acuminate, to 25 cm. long and 
10 cm. wide, coarsely erose-dentate or subentire, the upper surface 
more or less bullate by the impressed veins; staminate inflorescence 
2-3 cm. long, the peduncle as long; receptacle obconic, the 3-6 sepals 
broadly ovate-acutish ; stamens 5-6. According to Sleumer in Herb. 
Madrid, C. dentata Poepp. & Endl. Nov. Gen. & Sp. 2: 48. 1838, 
vix R. & P., belongs here. Negs. 8541, 27580. 

Huanuco: Cochero, Chinchao, Posuso, Muna, and Pillao (Ruiz 
& Pavon). Macora, Ruiz & Pavdn, type. Pampayacu, Poeppig 
D1228. Monzon, 1,500 meters, Weberbauer 3454; 284 (det. Perkins). 

Siparuna radiata (Poepp. & Endl.) A. DC. in DC. Prodr. 16, 
pt. 2: 656. 1868; 109. Citrosma radiata Poepp. & Endl. Nov. Gen. & 
Sp. 2: 48. 1838. 

Similar to S. magnifica; leaves often smaller, always cuneate at 
the base and rather rough-pubescent above; cymes 2-3 cm. long; 
pedicels 2-3 cm. long; receptacle densely rusty-tomentose. Included 
by Perkins in the synonymy of S. cristata (Poepp. & Endl.) A. DC., 



FLORA OF PERU 815 

III, but not cited as to collection; the latter species is Brazilian, 
monoecious, with 12-15 free stamens. Illustrated, Pflanzenreich 

IV. 101: /. 23. Neg. 13529. 

Loreto: Yurimaguas, Poeppig D2145, type; edge of forest, 200 
meters, Williams 4740. "Isula micunan." 

Siparuna saurauiifolia Perk. Nov. Sp. Repert. 1: 155. 1905; 
Nachtr. 55. 

A low shrub, densely yellowish-brown-pubescent nearly through- 
out; petioles to 1.5 cm. long; leaves opposite or ternate, the blades 
lanceolate or suboblong, to 8-15 cm. long and 3.5-5 cm. wide, rounded 
at the base, acute at the apex, minutely serrate, coriaceous, in age 
sparsely pilose above, the lateral nerves 14-17, prominent only 
beneath; staminate inflorescence to 3.5 cm. long, the peduncle to 
2 cm. long, the pedicels 1.5 cm. long; flowers greenish, 6.5 mm. broad; 
receptacle papery, the sepals at maturity obscure; stamens 30, 
unequal; pistillate inflorescence shorter, the pedicels nearly 2 mm. 
long, the flowers 3 mm. broad, the 3-5 sepals stellate- tomentose on 
both sides, nearly as long as the coriaceous receptacle; styles 4-5; 
fruit 6 mm. thick, densely long-muricate and slightly pubescent. 
Compare S. tomentosa (R. & P.) Perk. Negs. 27579, 13537. 

Amazonas: Chachapoyas, 2,000 meters, Weberbauer 4370, 4371; 
262. 

Siparuna suaveolens (Tul.) A. DC. in DC. Prodr. 16, pt. 2: 
647. 1868; Nachtr. 51. Citrosma suaveolens Tul. Ann. Sci. Nat. III. 
4:36. 1855. 

A scandent or clambering shrub with spreading branches, the 
younger ones pubescent with few fasciculate hairs; leaves ternate or 
verticillate, the blades obovate-lanceolate, to 30 cm. long, the pubes- 
cence on the nerves beneath sparse at maturity, mostly simple; peti- 
oles 1-3 cm. long; staminate inflorescence to 3.5 cm. long, soon gla- 
brate, the peduncles 6 mm. long, the pedicels 4-5 mm. long; receptacle 
papery; sepals 4, obtuse or acute, unequal, glabrous within; stamens 
8, the filaments petaloid, very unequal; pistillate inflorescence 3-5 
cm. long, the pedicels 6-10 mm. long; sepals 5-6, rotund, the larger 
to 4 mm. long, finally reflexed. A tree 3-4 meters tall, with agree- 
able odor, the fruits reddish (Weberbauer). Negs. 13533, 34802. 

Cuzco: Above Lucumayo, 1,500 meters, Weberbauer 4985. Santa 
Ana, Weddell 4760, type. 



816 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Siparuna subinodora (R. & P.) A. DC. in DC. Prodr. 16, pt. 
2: 649. 1868; 88. Citrosma subinodora R. & P. Syst. 1: 265. 1798. 

Branches trigonous; leaves ternate, the blades obovate-oblong- 
lanceolate, to 20 cm. long and 6 cm. wide, subacute to acuminate, 
attenuate at the base, more or less obsoletely undulate-crenate, 
papyraceous, glabrous; petioles 3-8 mm. long; inflorescence short, 
the staminate flowers shortly-pedicellate; receptacle hispidulous, 
finally glabrate; sepals 5 or 6, suborbicular, reflexing; velum glabrous, 
finally suberect; stamens 5-6, included. Negs. 13534, 8540. 

Junin: Chacahuassi, Ruiz & Pavon, type. Without locality, 
Ruiz & Pavon 24-34. 

Siparuna tabacifolia Perk. Verh. Bot. Ver. Brandenb. 47: 144. 
1905; Nachtr. 52. 

A shrub with large, thin leaves, rather laxly gray-pubescent or 
pale-rusty; petioles 3-6 cm. long; blades obovate, short-acuminate, 
obscurely denticulate, the lateral nerves 11-14; pistillate flowers 
solitary, yellowish, 4 mm. broad, the pedicels to 8 mm. long; recep- 
tacle coriaceous, densely stellate- tomentose; sepals 4-5, glabrous 
above, 2 mm. long; styles about 12; fruit obpyriform, sparsely pilose, 
the pedicels to 4 cm. long. 41b. Neg. 13535. 

Loreto: Cerro de Ponasa, 1,300 meters, Ule 6670, type. 

Siparuna Tayloriana Perk. Notizbl. Bot. Gart. Berlin 10: 162. 
1927. 

A shrub with climbing branches, opposite or sometimes ternate, 
glabrous leaves, slender petioles 1-2 cm. long, and few-flowered cymes 
3-6 cm. long; blades usually oblong, 6.5-12.5 cm. long, 2.5 cm. wide, 
cuneate or rounded-cuneate at the base, acuminate at the apex, 
obsoletely crenate-serrate, coriaceous, with 9-10 lateral nerves; 
peduncle 1-4 cm. long, the glabrous pedicels 4-5 mm. long; pistillate 
flowers 2-3 mm. broad, the coriaceous receptacle and ovate-acute 
sepals glabrous, as also the globose fruit, this about 6 mm. thick. 
Assigned No. 25a in the monograph. 

Huanuco: Rio Posuso, 2,000 meters, in forest, Weberbauer 6739, 
type. 

Siparuna thecaphora (Poepp. & Endl.) A. DC. in DC. Prodr. 
16, pt. 2: 657. 1868; 90. Citrosma thecaphora Poepp. & Endl. Nov. 
Gen. & Sp. 2: 48. 1838. 

A shrub or small tree, nearly glabrous, the thin, minutely den- 
ticulate leaves sparsely pubescent only on the nerves; blades obovate- 



FLORA OF PERU 817 

oblong, about 15 cm. long and nearly half as wide, acuminate, the 
petioles 1-2 cm. long; pistillate inflorescence short, the subsessile 
flowers pseudofasciculate; flowers to 2.5 mm. broad; receptacle 
obconic, chartaceous, slightly stellate-puberulent; sepals connate, 
glabrous, like the flat velum with a narrow throat; styles 10-11, 
little exserted, recurving. Neg. 13541. 

Junin: San Ramon, 1,100 meters, in forest, Killip & Smith 26005. 
Loreto: Yurimaguas, Poeppig D2157b, type. Iquitos, 100 meters, 
Killip & Smith 27239; King 75; Williams 3544, 1367; Ule 6248 (det. 
Perkins). San Antonio, 110 meters, Killip & Smith 29479. Caballo- 
cocha, in forest, Williams 2142, 2094, 2024. Rio Itaya, Williams 
3283, 3411. Rio Nanay, Williams 639. Pinto-cocha, Rio Nanay, 
Williams 826. "Curuinsi-sacha," "sacha-limon," "macusaro," "isula 
caspi." 

Siparuna tomentosa (R. & P.) Perk. Bot. Jahrb. 28: 691. 
1901; 101. Citrosma tomentosa R. & P. Syst. 1: 265. 1798. 

A shrub, softly and densely pale-rusty-pubescent nearly through- 
out; blades to 15 cm. long, 4-6 cm. wide, obtusely acuminate, the 
petioles 4-5 mm. long; pistillate inflorescence on a peduncle 4-7 mm. 
long, the pedicels 8-10 mm. long; receptacle suburceolate; sepals 
5-7, unequal; styles free, scarcely exserted. According to Weber- 
bauer, clambering in habit. Neg. 8372. 

Amazonas: Moyobamba, Weberbauer 4509; 291 (det. Perkins). 
San Martin: San Roque, 1,400 meters, Williams 7457. Rumizapa, 
400-900 meters, Williams 6755. Huanuco: Chinchao, Muiia, Pillao 
to Chacahuassi, and Cochero, Ruiz & Pavon 902, type. Without 
locality, Ruiz & Pavdn 24-37. Muna, 2,100 meters, 3936. Bolivia. 

Siparuna Tulasnei Perk. Bot. Jahrb. 28: 688. 1901; 98. 

More or less yellowish-pubescent, but the leaves finally glabrous 
above; petioles 1.5-2 cm. long; blades 10-20 cm. long, 5-7 cm. wide, 
sometimes narrowly acuminate, cuneate to the rounded base, mi- 
nutely denticulate or undulate; pistillate flowers apparently 5 mm. 
broad; receptacle subglobose, the 4 triangular sepals glabrous inside; 
styles about 10, exceeding the glabrous velum, the aperture narrow. 
-Neg. 13543. 

Huanuco: Tocache, Poeppig 1818, type. 

Siparuna ucayaliensis Perk. Notizbl. Bot. Gart. Berlin 10: 
167. 1927. 

A monoecious or dioecious shrub, more or less lepidote-pubescent 
except the mature leaves above; leaves opposite, oblong-obovate, 



818 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cuneate at the base, rounded at the apex and extended into a cau- 
date mucro, chartaceous, entire, 9.5-15.5 cm. long, 2.5-6 cm. wide, 
the lateral nerves 7-9; inflorescence cymose, 1-2 cm. long; flowers 
minute; stamens 4-5; staminate pedicels 2-4 mm. long, the pistillate 
1 mm. ; styles 2-3, little exserted. Differing from S. lepidantha Perk, 
in its pointed leaves, not lepidote on the midrib above. A shrub 
about 5 meters high. Compare also 5. crassiflora. Neg. 13544. 

Loreto: Yarina-cocha, Rio Ucayali, 155 meters, in forest, Tess- 
mann 5409, type. 

Siparuna Uleana Perk. Verh. Bot. Ver. Brandenb. 47: 142. 
1905;Nachtr. 49. 

A small, glabrous or essentially glabrous shrub with opposite, 
short (1-2 cm.) petioled, suboblong leaves, these 15-30 cm. long, 
7.5-13 cm. wide, subcordate at the base, thin, minutely denticulate; 
inflorescence 1.5-3 cm. long; flowers reddish, 3 mm. broad, the pedi- 
cels 2-3 mm. long; receptacle membranous, the 4 longer, lanceolate 
sepals 3.5 mm. long; stamens 6, unequal, the flat filaments very 
broad. Mature leaves are somewhat bullate above, sometimes 
more or less enlarged, toothed, and auricled at the base, slightly so 
in the type collection. Neg. 13545. 

Loreto: Cerro de Ponasa, 1,300 meters, Ule 6830, type. San 
Martin: Moyobamba, King 3352 (distributed as S. auriculata DC.?). 

Siparuna umbelliflora Perk. Nov. Sp. Repert. 1: 153. 1905; 
Nachtr. 49. 

A glabrous shrub; leaves ternate or verticillate, the blades more 
or less lanceolate, less than 15 cm. long, short-acuminate, elongate- 
narrowed to the slender petioles, some of which are shorter than 1.5 
cm.; leaf margins unequally and conspicuously serrate, the lateral 
nerves 11-14; inflorescence umbellate, the peduncle to 1.5 cm. long, 
the pedicels to 6 mm. long; staminate flowers 4 mm. broad, the recep- 
tacle obconic, 3 times exceeding the 4 minute sepals; stamens 52, 
short-exserted, unequal. Neg. 13546. 

Cajamarca: Above San Miguel, 2,600 meters, Weberbauer 3905, 
type; 258. 

Siparuna Weberbaueri Perk. Nov. Sp. Repert. 1: 154. 1905; 
Nachtr. 53. 

A shrub or small tree, densely and softly fulvous-tomentose, or 
the mature leaves glabrous and bullate above; petioles to 1 cm. long; 
leaves mostly ternate or verticillate, the blades to 7.5 cm. long, 3.5 



FLORA OF PERU 819 

cm. wide, rounded or broadly acute at each end, coriaceous, the 
lateral nerves 10-12; inflorescence about 1 cm. long, the peduncles 
and pedicels 5 mm. long; staminate flowers greenish, 3.5 mm. broad; 
receptacle papery, glabrous within; sepals connate- tubular; stamens 
10, unequal. 

Libertad: Angon, valley of Rio Mixiollo, Weberbauer 7063. 
Junin: Above Huacapistana, 2,000 meters, Weberbauer 2161, type; 
246. Huacapistana, 1,000-2,400 meters, Kittip & Smith 24136 
Huanuco: Valley of Rio Posuso, 1,300 meters, Weberbauer 6742. 

Siparuna Williamsii Macbr. Candollea 5: 353. 1934. 

A tree, reported to attain a height of 15 meters, the branchlets 
terete, at first slightly rusty-stellate-puberulent, compressed at the 
nodes and about 1 cm. thick; petioles opposite, glabrate, acutely 
sulcate above, 2.2-5 cm. long; blades elliptic, rounded to the acute 
base, abruptly and very shortly acuminate, 20-30 cm. long, 10-12 
cm. wide, chartaceous, glabrous or sometimes sparsely and minutely 
stellate-puberulent on the nerves beneath, the lateral nerves 10-12, 
not at all impressed above, the veins moderately prominent beneath, 
laxly reticulate; staminate racemes simple or 1-2-branched, mostly 
5-6 cm. long, densely stellate-pubescent with minute hairs; pedicels 
slender, to almost 1 cm. long; receptacle ovoid, to 2 cm. thick; sepals 
and velum none; stamens 12-15, the extremely broad, flattish fila- 
ments not or scarcely exserted. In the broad filaments, resembling 
S. pauciflora (Beurl.) A. DC., but vegetatively suggesting S. Uleana 
Perk. 

Loreto: Alto Rio Itaya, 145 meters, in forest, Williams 3371, 
type. San Antonio, Rio Itaya, 110 meters, Kittip & Smith 29467. 
Soledad, Rio Itaya, 110 meters, in forest, Kittip & Smith 29818. 

76. LAURACEAE. Laurel Family 
Reference: Mez, Jahrb. Bot. Gart. Berlin 5: 1-556. 1889. 

The majority of the many Peruvian species of this family are 
trees, often of great beauty, not rarely of great utility, and generally 
distributed in the montana. In the higher and drier regions there 
are a number of shrubs which, however, never ascend, according to 
Weberbauer, above 3,200 meters, and on the western ranges only to 
2,000 meters. The following remarks were written before Koster- 
mans' recent revisions. 

Few families afford a greater fascination in their study, for the 
characters, largely floral, are of a neatness and variation that seem 



820 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to be almost infinite in the number of combinations in which they 
have developed. Closely knit, therefore, the family has always 
been subjected to diverse interpretation regarding the lines to be 
drawn between groups of more or less obviously related species. It 
seems to me a great pity to divide a family of such economic and gen- 
eral interest into many narrowly limited genera, however logical 
this may seem, and I have written the following paragraphs hoping 
to call attention to the possibility of strengthening the generic segre- 
gation proposed by Mez, but not by further segregation, which, of 
course, is always the easiest way to deal with a family having many 
aberrant elements, thus isolating its larger and better defined groups. 

The relative value of characters used to indicate generic lines in 
the Lauraceae in Field Mus. Bot. 11: 19-20. 1931, I published a 
general comment has been too little considered, and a reconsidera- 
tion of at least some generic limits seems necessary. For instance, 
if treated relatively as other groups, Aniba should be defined to 
include Acrodiclidium (Licaria) and Endlicheria as regards Peru, as 
well as Silvia, Misanteca, Huberodaphne, Systemonodaphne, Phylloste- 
monodaphne, and Chanekea, so that species more nearly related to 
each other than to those of any other genus might conveniently have 
one generic name. It seems to me that the characters upon which 
these genera rest cannot logically be accepted as indicative of generic 
lines. For instance, the degree of filament development and glandu- 
losity is highly variable throughout the family. 

Dismissing the argument that no character in the family is con- 
stant which is true, for even the usually fundamental one of 2- and 
4-celled anthers must be qualified with exceptions evidently ten- 
dencies must rule in determining probable natural lines of develop- 
ment. One is faced with the fact that regularity of development 
of staminodia as foliaceous or reduced in series 4 suggests its 
reliability in delineating genera. In contrast, the variation of stami- 
nodia in the outer series within a group of admittedly related species 
seems to point to its lack of value as a generic character. If one 
does not reason thus as regards characters in the Lauraceae, that is, 
consider their relative or major validity in the family as a whole, one 
is soon forced to establish many new "genera" on negative char- 
acters. The result of the latter method is to lose in a multitude of 
generic names, all of which purport to have equal value, any com- 
prehensive view of the natural grouping of species within the family. 

Aniba sens, lat., as suggested, would strengthen the whole generic 
system of the family and form a group of no more variation than 



FLORA OF PERU 821 

Ocotea or Nectandra, and one even better contrasted with its nearest 
relative, Ajouea. Acrodiclidium is logically only a section of Aniba, 
with the 2 outer series of stamens modified or aborted, a character 
not of generic value, as shown clearly by its variability in Ajouea 
and elsewhere. Further, Endlicheria cannot be regarded as a group 
distinct from Aniba on the character of dioecious flowers, since both 
perfect and dioecious species exist in other genera, and some species 
may even be either (Ocotea minarum, for example). Other char- 
acters purporting to define Endlicheria exist in every degree of 
development in most if not all accepted genera. (But Kostermans 
has shown recently that there may be a partially constant fruit 
character and a generally constant difference in anther shape facts 
that strengthen the argument. It seems to me that only one genus 
is involved, composed of several fairly well marked groups of species.) 
The unreliability of the anther form as a generic character, at least 
here, is shown in Endlicheria anomala, whose second pair of anthers 
usually has the more circular shape that normally characterizes 
anthers of Acrodiclidium and Aniba. The form of the anthers else- 
where has not been regarded by Mez himself as alone significant in 
drawing generic lines. 

Urbanodendron Mez would not necessarily be included in Aniba 
sens. lat. because its position in the 2-celled anther series of genera 
is about analogous to that of Pleurothyrium in the 4-celled series. 
Urbanodendron could be maintained, although, except for its fila- 
ments and style, it is essentially the same as the Peruvian Endlicheria 
Sprucei. 

In accord with the above remarks, there may be found in the 
following synopsis of the Peruvian laurels a vegetative and technical 
key to Aniba, sens. lat. For convenience an artificial key has been 
attempted for all the species, disregarding generic lines, in hope that 
it may lead one to at least the better marked forms, even when 
flowers are lacking, but of course it requires herbarium material for 
comparison. 

Besides the native Lauraceae listed below, the European laurel, 
a beautiful, evergreen tree,Laurus nobilis L., is sometimes cultivated. 
Its crowded, coriaceous, lanceolate leaves are aromatic, dark green, 
and smooth, usually paler and the midrib reddish beneath; its inflo- 
rescences are short, few-flowered, and subtended by brown bracts 
that fall with the opening of the flowers, these 4-parted, the stami- 
nate usually with 10-12 stamens, the pistillate with 2-4 staminodia. 



822 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Although Kostermans' work is cited in so far as published at this 
writing, at the beginning of generic descriptions, I must take occa- 
sion to record here my deep indebtedness to him. He not only 
made available all his manuscripts before publication but had the 
kindness to make many suggestions and many insertions in my 
manuscript, which had already (for several years, based on Mez) 
been prepared. 

Generic key after Mez (after Kostermans as to Aniba group) 
Anthers of the exterior series 2-celled or aborted. 

Fruit on a naked, not enlarged pedicel; staminodia of series 4 
foliaceous .... 1. Beilschmiedia. 

Fruit subtended by a cup; staminodia often minute or none. 

Staminodia of series 4 large, foliaceous 2. Ajouea. 

Staminodia of series 4 minute or none. 

Flowers dioecious; pedicel and cup typically fleshy, the former 
enlarged 3. Endlicheria. 

Flowers perfect; pedicel and cup (so far as known) woody, 
the former not elongate, the cup usually deep. 

Outer stamens (6 or 9) fertile, free 4. Aniba. 

Outer stamens (6) none or modified, only the inner 3 fertile, 
connate or free 5. Licaria. 

Anthers of 2 outer series 4-celled or very rarely all 2-celled. 

Staminodia (series 4) large, sagittate, stipitate; perianth lobes, 
with few exceptions, persistent. 

Outer perianth lobes usually shorter than the inner; perianth 
not at all enlarging above the globose fruit 6. Persea. 

Outer perianth lobes rarely shorter than the inner, strongly 
enlarged about the usually ellipsoid fruit, or deciduous, but 
the pedicel then enlarged 7. Phoebe. 

Staminodia (series 4) none or small and stipitiform. 

Outer stamens eglandular (normally); each pair of anther cells 
in juxtaposition. 

Anther cells in pairs, one pair more or less exactly above the 
other 8. Ocotea. 

Anther cells placed arcuately in line 9. Nectandra. 

Stamens all basally glandular; each pair of anther cells at right 
angles to the other 10. Pleurothyrium. 



FLORA OF PERU 823 

Artificial key to Peruvian Lauraceae 

Based largely on foliage; the leaf size is taken from herbarium 
specimens which usually are branchlets bearing inflorescences. 

A. Leaves small, i.e., mostly less than 10 cm. long, few, if any, a 
little longer, or in this case only 2.5 cm. wide (see forms of Persea 
caerulea). 

B. Leaves obviously pubescent beneath. 
Pubescence elegantly sericeous. 
Inflorescence many-flowered. 
Leaves silvery-pubescent beneath, acute at the base. 

Ocotea guianensis. 
Leaves densely coppery-pubescent beneath, rounded at the 

base Persea Haenkeana. 

Leaves slightly coppery-pubescent beneath, acute at the 

base Ocotea cuprea. 

Inflorescence few-flowered. 

Leaves about 8 cm. long Persea stricta. 

Leaves 5-6 cm. long Persea Weberbaueri. 

Pubescence villous-tomentose, sometimes finely so, usually rusty 
or reddish (see Persea stricta and Ocotea macropoda). 

Leaves (especially beneath) shortly and densely tomentulose. 
Leaves oblong-lanceolate, mostly 10-12 cm. long. 

Leaves densely rusty-pubescent; petioles short. 

Ocotea Jelskii. 

Leaves cinereous- tomentulose beneath; petioles to 1.5 
cm. long Nectandra Mathewsii. 

Leaves ovate-elliptic, often shorter. 

Pubescence very reddish; leaves about 10 cm. long. 

Ocotea cardinalis. 

Pubescence rusty or rusty reddish; leaves usually 5-9 
cm. long. 

Inflorescence corymbose-panicled. 

Inflorescence many-flowered; style much shorter 
than the ovary Persea corymbosa. 

Inflorescence few-flowered; style and ovary subequal. 

Persea ferruginea. 
Inflorescence racemose Ocotea ferruginea. 



824 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves rather laxly long-villous or finely pilose, at least 
beneath. 

Branchlets soon glabrous Ocotea aciphylla. 

Branchlets villous or pilose. 

Leaves subcordate at the base Persea Ruizii. 

Leaves acute at the base. 

Pubescence villous Persea hirta. 

Pubescence fine, pilose or tomentulose. 

Flowers glabrous Ocotea tenella. 

Flowers tomentose .Licaria multiflora,L.Quirirafuina. 
B. Leaves glabrous or with minute or obscure pubescence, or bar- 
bellate in the nerve axils beneath (small-leaved forms of 
Ocotea leptobotra might be sought here). 
Leaves 3 to rarely 5 cm. long. 
Inflorescence pilose. 
Inflorescence sparsely pilose, about equaling the leaves. 

Ocotea Mandonii. 
Inflorescence densely pilose, exceeding the leaves. 

Persea boldufolia. 

Inflorescence glabrous Ocotea jumbillensis. 

Leaves all or mostly longer. 

C. Inflorescences small, mostly only 1-5 cm. long; see also 

forms of Nectandra Pichurim. 

D. Inflorescence glabrate or, in any case, not cinereous- 
pubescent. 
Flowers 7-8 mm. wide. 

Leaves narrowly lanceolate Nectandra microcarpa. 

Leaves elliptic Ocotea munacensis. 

Flowers tiny or small. 

Flowers 3 mm. wide, the inflorescence crowded- thyrsoid. 

Phoebe heterantha. 

Flowers often smaller, the inflorescence not crowded- 

thyrsoid. 
Inflorescence glabrate; flowers 2 mm. long. 

Leaves obscurely veined Ocotea minutiflora. 

Leaves distinctly veined. 
Petioles 4-7 mm. long. 



FLORA OF PERU 825 

Leaves broadly elliptic, caudate. 

Ocotea tarapotana. 
Leaves suboblong, acuminate. 

Endlicheria mishuyacensis. 

Petioles to 1 cm. long Ocotea Killipii. 

Inflorescence pubescent; flowers 2-5 mm. long. 
Shrubs with rigid, concolorous leaves. 

Flowers subracemose Ocotea Weberbaueri. 

Flowers subcorymbose. 

Branches glabrate. . . .Ocotea Schomburgkiana. 

Branches tomentose Ocotea monzonensis. 

Trees with thinner, flexible leaves, paler beneath. 
Leaves rounded at the base . . . Phoebe maynensis. 
Leaves acute at the base . . . Phoebe pichichensis. 
D. Inflorescence cinereous-pubescent. 

Leaves elliptic Persea americana. 

Leaves suboblong Phoebe Poeppigii. 

C. Inflorescence longer, often 2-4 times longer. 
D. Inflorescence shorter than or about equaling the leaves. 

Leaves coppery-pubescent beneath Ocotea cuprea. 

Leaves glabrous or glabrate (at least the adult ones). 
Leaves obtuse or subcordate at the base. 
Leaves rigid; petioles thickish, about 1 cm. long. 

Leaves glaucous Persea Mutisii. 

Leaves lustrous above, comparatively few but 

crowded Ocotea piurensis. 

Leaves flexible; petioles slender, 1.5-3 cm. long. 
Leaves subcordate; petioles 1.5-3 cm. long. 

Persea cordata. 
Leaves subobtuse; petioles to 1.5 cm. long. 

Acrodiclidium armeniacum. 
Leaves acute at the base, or flexible and the petioles 

short. 
Inflorescence glabrous. 

Plants entirely glabrous Ocotea subrutilans. 

Plants with the leaf nerves beneath minutely 
strigose. 



826 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels 5-20 mm. long. 

Acrodiclidium armeniacum. 

Pedicels 2-3 mm. long Nectandra Sawadai. 

Inflorescence at least minutely pubescent. 

Branches and inflorescence sericeous- tomentulose. 

Per sea durifolia. 
Branches and inflorescence glabrate or minutely 

pubescent. 
Leaves thin, lance-elliptic. 

Leaves with barbellate pubescence. 

Ocotea minarum. 
Leaves with no pubescence. 

Nectandra Williamsii. 
Leaves subcordate, oblong-elliptic. 
Petioles 5-8 mm. long; leaves mostly 3 cm. 

wide Persea scoparia. 

Petioles 10-12 mm. long; leaves 3.5 cm. wide. 
Inflorescence more or less panicled; leaves 

3-nerved Phoebe Poeppigii. 

Inflorescence subracemose; leaves pinnate- 
nerved Phoebe Mathewsii. 

D. Inflorescence 2-3 times longer than the leaves. 

Branchlets pilose Ajouea tambillensis. 

Branchlets glabrous Ocotea piurensis. 

A. Leaves of medium to large size, mostly 12 cm. to several deci- 
meters long. 

E. Leaves of average or medium size, mostly 12-15 cm. long or, 
if larger, only (3) 4.5 to rarely 8 cm. wide or, if wider, only 
12-15 cm. long. 

F. Flowers conspicuous, 10-15 mm. wide. 
Leaves glabrous or glabrate beneath. 

Pedicels usually 3-5 mm. long, the leaves usually long-acu- 
minate Nectandra lucida, N. globosa. 

Pedicels usually very short, the leaves acute or short-acu- 
minate. 
Leaves acuminate at the base; perianth tube obvious. 

Nectandra Jelskii. 
Leaves merely acute at the base. 



^ FLORA OF PERU 827 

Inflorescence rusty- tomentose; perianth tube none. 

Nectandra furcata. 
Inflorescence glabrate; perianth tube obvious. 

. Nectandra maranonensis. 
Leaves pubescent beneath even in age, or at least densely bar- 

bellate in the nerve axils. 
Leaf pubescence pale or confined to the nerve axils; perianth 

tube conspicuous, except in Endlicheria bullata. 
Leaves broad. 

Inflorescence and leaves subequal . Nectandra acutifolia. 
Inflorescence often shorter than leaves. 

Nectandra globosa. 
Leaves oblong-linear, about 3 cm. wide. 

Endlicheria bullata. 
Leaf pubescence rusty, evenly distributed; perianth tube 

short. 
Leaves more or less pilose on both sides, at least on the 

nerves above, rarely only puberulent beneath. 
Leaves rigid-coriaceous or nearly so, subobtuse at the 

base; pedicels 1-2 mm. long Nectandra Laurel. 

Leaves more or less flexible, acute at the base; pedicels 

3-8 mm. long Nectandra reticulata. 

Leaves glabrate above and merely puberulent-tomentose 

beneath Nectandra longifolia. 

F. Flowers minute to medium-sized, often inconspicuous, 3 to 

rarely 8 mm. wide, or smaller. 

G. Mature leaves obviously pubescent beneath or at least 
densely barbellate in the nerve axils, the pubescence 
sometimes very minute and confined to the veins. 
H. Pubescence sparse or minute, fine, always inconspicu- 
ous, or merely barbellate. 

Leaves barbellate in the nerve axils beneath (sometimes 
obscurely so in Phoebe maynensis), or slightly pubes- 
cent on the costa. 
Scandent shrubs; petioles to 5 mm. long. 

Ocotea tarapotana. 

Erect shrubs or trees; petioles 1-2 cm. long. 
Leaves more or less distinctly trinerved. 

Phoebe maynensis. 



828 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. 

Leaves pinnate-nerved. 
Leaves thin, longer than the inflorescence. 
Petioles to 18 mm. long; flowers dioecious. 

Endlicheria dysodantha. 
Petioles to 11 mm. long; flowers perfect. 

Nectandra Sawadai. 
Leaves thick, about equaling the inflorescence. 

Ocotea cuneifolia. 
Leaves not barbellate beneath or, if slightly so, otherwise 

pubescent also. 
Petioles elongate, most or some of them more than 2.5 

cm. long (to 6 cm.). 
Leaves obscurely reticulate-veined beneath; stami- 

nodia small. 

Inflorescence tomentose; pedicels 1-4 mm. long. 

Nectandra lineatifolia. 
Inflorescence minutely pubescent; pedicels 5-8 

mm. long Nectandra myriantha. 

Leaves obviously reticulate- veined beneath; stami- 

nodia large, sagittate. 

Ovary densely pilose; leaves rather thin; flowers 
6-7 mm. long; perianth segments subequal. 

Per sea americana. 
Ovary glabrous; perianth lobes very unequal, 

except in Nectandra elongata. 
Leaves short-acuminate, slightly reticulate- 
veined. 

Leaves obtuse or rounded at the base (usu- 
ally), rather thin; flowers 4-6 mm. long; 
pedicels 2-5 mm. long or longer. 

Persea caerulea. 

Leaves acute at the base, coriaceous; flowers 
7-8 mm. long; pedicels 2 mm. long. 

Persea Raimondii. 

Leaves long-acuminate, prominently reticulate. 

Nectandra elongata. 
Petioles short, mostly less than 2.5 cm. long (5-25 mm. 

long), that is, those of the branchlets. 
I. Some petioles 1.5-2.5 cm. long. 



FLORA OF PERU 829 

Pubescence, at least at first, dense. 
Leaves glabrate in age; filaments glabrous. 

Ocotea Rusbyana. 
Leaves often minutely pilose beneath in age; 

filaments pilose. 
Leaves microscopically papillose beneath. 

Aniba rosaeodora. 
Leaves not papillose. 
Pubescence beneath obvious only under a 

lens Ocotea Dielsiana. 

Pubescence beneath obvious (see Ocotea 

opifera) Ocotea Trianae. 

Pubescence, from the first, sparse or inconspicuous. 
Leaves subtriplinerved, shorter than or sub- 
equaling the inflorescence. 

Phoebe cinnamomifolia. 

Leaves pinnate-nerved, longer than the inflores- 
cence. 

Leaves subobtuse Bellota sulcata. 

Leaves acuminate. 
Leaves obscurely tomentose beneath. 

Aniba Coto, A. Muca. 

Leaves evenly puberulent-strigillose be- 
neath . Nectandraelongata,N.Macbridei. 
Leaves glabrous except the minutely stri- 

gose costa or nerves. 

Leaves narrowly acuminate; pedicels elon- 
gate .... Acrodiclidium armeniacum. 
Leaves gradually acuminate; pedicels 
1-2 mm. long. . .Ocotea Ottoschmidtii. 
I. Some petioles 5-10 mm. long, none much longer. 
Petioles only about 5 mm. long. 

A shrub or tree Ocotea rubrinervis. 

A scandent shrub Ocotea tarapotana. 

Petioles about 10 mm. long (see A. rosaeodora). 
Leaves obovate, very cuneate at the base. 

Ocotea cuneifolia. 
Leaves not obovate . . . . J. 



830 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

J. Leaves rigid, elliptic, 5 cm. wide, much shorter than the inflores- 
cence. 

Leaves acute at each end Per sea crassifolia. 

Leaves subobtuse Ocotea micans. 

Leaves lanceolate or elliptic, not rigid or subequaling the inflores- 
cence. 

Pubescence a minute puberulence. 

Leaves membranous, acuminate or caudate (see Ocotea marmel- 

lensis, rarely Phoebe Poeppigii). 

Flowers large, 8 mm. wide Nectandra acutifolia. 

Flowers small. 

Leaves acuminate, mostly 4.5 cm. wide. 

Leaves ovate-acuminate; perianth tube obvious. 

Endlicheria anomala. 

Leaves elliptic-lanceolate; perianth tube none (see End- 
licheria debilis). 
Flowers dioecious; leaves mostly 4.5 cm. wide. 

Ocotea puberula. 
Flowers perfect; leaves rarely 4.5 cm. wide. 

Nectandra Sawadai. 
Leaves caudate, mostly 6-7 cm. wide. 

Acrodiclidium latifolium. 
Leaves rigid-coriaceous or coriaceous. 
Leaves rigid; branchlets more or less angled. 

Leaves rarely acuminate Ocotea aurantiodora. 

Leaves sharply long-acuminate Ocotea aciphylla. 

Leaves coriaceous; branchlets terete Aniba cylindri flora. 

Pubescence a sparse pilosity. 
Leaves about 10 cm. long. 

Petioles 1.5-2.5 cm. long Ocotea Trianae. 

Petioles usually 1 cm. long. 

Leaves pubescent beneath ; perianth tube obsolete. 

Ocotea petalanthera. 
Leaves becoming glabrous; perianth tube conspicuous. 

Ocotea otuzensis. 
Leaves about 15 cm. long or longer Endlicheria longifolia. 



FLORA OF PERU 831 

H. Pubescence obvious or even conspicuous, rather coarse 
or, if fine, dense enough to color the leaf surface (see 
also Ocotea Raimondii and Aniba rosaeodora). 
K. Pubescence cinereous or pale rusty, usually pilose- 
hirsute or sericeous or, if tomentose, laxly so (see 
Endlicheria bullata). 

Leaves rounded, obtuse, or acute at the base; petioles 
often 2-3 cm. long. 

Petioles scarcely 5 mm. long Ocotea licanioides. 

Petioles mostly much longer. 

Leaves acuminate; petioles 1.5-3 cm. long. 

Inflorescence open, often little shorter than the 

leaves, these lustrous above. 
Peduncles elongate; perianth lobes unequal. 

Persea cordata. 
Peduncles short; perianth lobes equal. 

Nectandra superba. 

Inflorescence dense, much shorter than the 
leaves, these dull above. Nectandra dioica. 
Leaves subobtuse; petioles 1.5 cm. long. 

Bellota sulcata. 

Leaves basally acute, only above the base, if at all, 
more or less rounded; petioles usually shorter (see 
Nectandra superba). 
Leaves thin-coriaceous; outer anthers 2-celled (except 

in Nectandra spp.). 
Leaves acuminate, glabrous above. 

Petioles to 14 mm. long; flowers obscurely stri- 

gose Endlicheria anomala. 

Petioles often longer; flowers densely sericeous. 
Endlicheria Lhotzkyi, E. sericea. 
Leaves acute, the costa above pubescent. 

Nectandra Herrerae. 

Leaves coriaceous, except in Ocotea amazonica with 

the leaves pilose beneath; outer anthers 4-celled. 

Leaves mostly 5-8 cm. wide (see also Ocotea Rus- 

byana) . 

Branchlets and inflorescence glabrate or cinere- 
ous-puberulent Ocotea Trianae. 



832 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branchlets and inflorescence rusty-puberulent 
or shortly yellow-villous or tomentose. 

Branchlets and inflorescence rusty-puberu- 
lent. 
Leaves obovate; inflorescence ample. 

Leaves very rigid-coriaceous. 

Pleurothyrium Poeppigii. 
Leaves flexible-coriaceous. 

Pleurothyrium cuneifolium. 

Leaves not at all obovate; inflorescence 
few-flowered Ocotea micans. 

Branchlets and inflorescence yellow-villous or 

velvety-tomentose. 

Pubescence villous; petioles to 25 mm. long. 

Nectandra cissiflora. 

Pubescence tomentose; petioles to 15 mm. 

long Nectandra comasensis. 

Leaves mostly 3.5-4.5 cm. wide. 
Leaves tomentulose beneath; inflorescence and 
leaves subequal. 

Leaves mostly narrower than 4 cm. 

Nectandra Pichurim. 
Leaves mostly wider. 

Nectandra Mathewsii, N. Macbridei. 

Leaves pilose-puberulent beneath; inflorescence 
manifestly shorter than the leaves. 

Pubescence a minute puberulence. 

Ocotea puberula. 

Pubescence a more or less sparse pilosity. 

Inflorescence slightly pubescent. 

Ocotea petalanthera. 
Inflorescence densely pubescent. 

Leaves apparently glabrous beneath. 

Ocotea Rusbyana. 

Leaves obviously pilose beneath. 

Ocotea amazonica. 

K. Pubescence definitely rusty or reddish-tomentose- 
villous. 



FLORA OF PERU 833 

Leaves rigid-coriaceous, densely reddish-tomentose, at 
least beneath, and attenuate-acuminate. 

Endlicheria Szyszylowiczii. 

Leaves somewhat flexible, soon glabrate above or 
tomentose only on the nerves, not elongate-acu- 
minate. 

Petioles 2.5-5 cm. long; leaves rounded-subcordate 
at the base (see Per sea peruviand). 

Persea subcordata. 

Petioles 2.5 cm. long or often shorter; leaves not sub- 
cordate at the base. 

Branchlets villous-hirsute Endlicheria hirsuta. 

Branchlets not villous-hirsute. 

Leaves reticulate-veined, if acuminate, not 

narrowly so; outer anthers 4-celled. 
Leaves about 5 cm. wide. . .Ocotea macropoda. 
Leaves about 8 cm. wide. 
Inflorescence narrow, dense. 

Ocotea ovalifolia. 
Inflorescence broad, open. 

Endlicheria anomala. 

Leaves obscurely reticulate-veined, or long and 

narrowly acuminate; outer anthers 2-celled. 

Inflorescence tomentose, subequaling the 

leaves Endlicheria tomentella. 

Inflorescence pilose, much shorter than the 
leaves Endlicheria Sprucei. 

G. Mature leaves glabrous or essentially so, even be- 
neath Ocotea otuzcensis. 

L. Leaves mostly under 5 cm. wide (inflorescence leaves 
2 to about 5 cm. wide). 

M. Leaves more or less abruptly subcaudate-acuminate, 
the acumination usually 1 cm. long or longer. 

Perianth tube none or extremely short. 

Inflorescence glabrous . . . Acrodiclidium armeniacum, 
Inflorescence pubescent. 

Inflorescence racemose-paniculate. 

Ocotea maynensis. 



834 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Inflorescence fasciculate-paniculate. 

Ocotea Schomburgkiana. 
Perianth tube obvious. 
Leaves coriaceous, very lustrous above. 

Ocotea leptobotra. 

Leaves papyraceous or rarely coriaceous, dull or 
little lustrous, except in Endlicheria mishuya- 
censis. 

Inflorescence often 2-4 cm. long, glabrate, or gla- 
brous if elongate (Ocotea marmellensis). 

Leaves caudate; perianth tube short. . . .Ocotea 
Killippii, 0. caudata, 0. marmellensis, Nectandra 

citrifolia. 

Leaves long-acuminate; perianth tube elongate. 

Endlicheria mishuyacensis. 

Inflorescence often 6-10 cm. long, more or less 

tomentose. 

Petioles almost 1.5 cm. long or longer. 
Petioles to 1.5 cm. long; inflorescence lax. 

Leaves thin, the reticulation obscure, fine. 

Aniba Tessmannii. 

Leaves thin, the reticulation beneath coarse. 

Aniba firmula. 

Petioles to 2 cm. long; inflorescence dense. 

Nectandra maynensis. 

Petioles rarely 8 mm. long Ocotea Keriana. 

M. Leaves obtuse, acute, or, if acuminate, gradually 
or shortly so (see also Ocotea Keriana). 

Leaves mostly less than 2 cm. wide. 

Nectandra microcarpa. 
Leaves much wider. 

Perianth tube none, or extremely short (see Acrodi- 
clidium Pucheri); leaves rarely conspicuously 
reticulate- veined . 

Leaves obscurely reticulate- veined. 

Ocotea multiglandulosa. 

Leaves obviously reticulate-veined, or at least 
beneath. 



FLORA OF PERU 835 

Leaves densely white-puncticulate; inflorescence 

glabrate Ocotea albopuncticulata. 

Leaves not white-puncticulate; inflorescence 

more or less pubescent. 

Flowers 2-2.5 mm. long. . . .Ocotea puberula. 
Flowers 3.2-3.5 mm. long. Ocotea Tessmannii. 

Perianth tube conspicuous; reticulate venation often 
prominent. 

Inflorescence glabrous or essentially so. 
Branchlets pilose, at least toward the tip. 

Flowers 2.5 mm. long Ajouea Jelskii. 

Flowers 3-4 mm. long .... Ajouea tambillensis. 
Branchlets glabrous . Pleurothyrium densiflorum. 
Inflorescence pubescent. 

Leaves exceptionally densely and prominently 
reticulate-veined on both sides. 

Acrodiclidium Pucheri. 

Leaves laxly and usually lightly reticulate- 
veined above. 

Petioles about 20 mm. long. 

Nectandra intermedia. 

Petioles about 10-15 (-20) mm. long. 

Bark often aromatic; inflorescences shorter 
than the leaves. 

Leaves firm-coriaceous, rather obscurely 
reticulate- veined . 

Leaves densely reticulate-veined be- 
neath; outer stamens modified. 
Acrodiclidium Pucheri. 

Leaves laxly reticulate- veined beneath; 
stamens all fertile. 

Aniba Muca, A. Canelilla. 

Leaves very thin, conspicuously reticu- 
late-veined . . . Endlicheria rubriflora. 

Bark not aromatic; inflorescence usually 
about equaling the leaves, or longer 
(see Endlicheria rubriflora). 



836 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves closely reticulate- veined beneath, 

obtuse at the base .Ocoteapichurensis. 

Leaves laxly reticulate-veined, acute at 

the base. 
Leaves subtriplinerved. 

Phoebe cinnamomifolia. 
Leaves pinnate-nerved. 
Leaves subobtuse . Phoebe Mathewsii. 

Leaves acuminate Nectandra 

Pichurim, Aniba perutilis. 

L. Leaves mostly over 5 cm. wide (about 5-10 cm. wide). 
Petioles 3-6 cm. long. 

Ovary densely pilose; flowers 6-7 mm. long. 

Persea americana. 

Ovary glabrous; flowers 4-6 mm. long. Persea caerulea. 
Petioles mostly 1-2 cm. long. 

N. Pedicels mostly 4-5 mm. long or longer. 
Inflorescence glabrous or essentially so. 
Leaves obsoletely reticulate-veined above. 

Ajouea tambillensis. 

Leaves more or less reticulate-veined even above. 
Leaves thin, very densely reticulate- veined, 
often 8 cm. wide or wider. 

Bellota sulcata, Endlicheria rubriflora. 
Leaves thick, rather laxly reticulate- veined, 

often narrower. 
Inflorescence longer than the leaves. 

Ajouea tambillensis. 
Inflorescence shorter than the leaves. 
Petioles scarcely 1 cm. long. 

Pleurothyrium densiflorum. 
Petioles about 1.5 cm. long .Ocotea leptobotra. 
Petioles about 2.5 cm. long.OcoZea obovata. 
Inflorescence more or less pubescent. 

Flowers umbellate Ocotea piurensis. 

Flowers not evidently umbellate. 

Leaves thin, very densely reticulate-veined, 8-9 
cm. wide. . . .Bellota sulcata. 



FLORA OF PERU 837 

Leaves not as above in all respects. 
Perianth tube short; leaves not broadly 

rounded-obovate. 
Leaves subobtuse or acute, 6-12 cm. wide; 

anthers 2-celled Bellota sulcata. 

Leaves acuminate, 4.5-9 cm. wide; anthers 

4-celled. 

Lateral nerves of the leaves 9-11; flowers 

strigose. . .Nectandra capanahuensis. 

Lateral nerves of the leaves 5-8; flowers 

puberulent. 

Leaves 6-8 cm. wide; inflorescence 10 
cm. long . Nectandra breaparinensis. 
Leaves about 5 cm. wide; inflorescence 
about 5 cm. long . Nectandra lucida. 
Perianth tube conspicuous; leaves more or 
less broadly rounded-obovate, conspicu- 
ously cuneate at the base. 

Inflorescence glabrous or slightly strigose; 

flowers dioecious Ocotea obovata. 

Inflorescence tomentulose; flowers perfect. 

Ocotea compacta. 
N. Pedicels mostly much shorter. 

0. Perianth tube conspicuous, except in Endlicheria 

arunciflora, where short. 
Leaves long-acuminate or caudate. 
Leaves acuminate or subcaudate. 
Leaves narrowed at both ends. 

Reticulation of the leaves obscure. 

Aniba Puchury-minor. 
Reticulation of the leaves conspicuous. 

Endlicheria mishuyacensis. 

Leaves rounded at the base or at both ends. 

Inflorescence small . . . .Ocotea marowynensis. 

Inflorescence ample to very ample. 

Reticulation of the leaves very coarse, 

lax Endlicheria arunciflora. 



838 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Reticulation of the leaves very fine, 
dense Aniba citrifolia. 

Leaves conspicuously caudate. .Ocotea laxiflora. 

Leaves rounded or short-acuminate (see Ocotea 
compactd). 

Leaves rounded at the apex, conspicuously obo- 
vate Ocotea obovata. 

Leaves short-acuminate, little or not at all obo- 
vate. 

Reticulation of the leaves conspicuous on 
both sides Endlicheria rubriflora. 

Reticulation of the leaves inconspicuous, at 
least above. 

Leaves subcoriaceous, dull, at least be- 
neath. 

Inflorescence 5-8 cm. long, the flowers 
clustered. 

Perianth segments unequal. 

Aniba Canelilla. 

Perianth segments equal. 

Aniba Puchury-minor. 

Inflorescence often shorter, the flowers 
not clustered Ocotea leptobotra. 

Leaves papyraceous, lustrous beneath. 

Ocotea Ottoschmidtii. 
0. Perianth tube none or short. 

Leaves lustrous or very lustrous above. 

Leaf reticulation prominent, at least beneath. 

Leaves acuminate; flowers to 4 mm. wide or 
wider. 

Inflorescence pubescent. 

Leaves about two-thirds longer than 
broad Nectandra yarinensis. 

Leaves scarcely one-third longer than 
broad Nectandra latifolia. 

Inflorescence glabrous or glabrate. 

Nectandra maranonensis. 



FLORA OF PERU 839 

Leaves subobtuse; flowers 7-8 mm. wide. 

Nectandra purpurea. 
Leaf reticulation obscure. 

Inflorescence short, cinereous-tomentulose. 

Ocotea architectorum. 

Inflorescence ample, glabrate. 

Nectandra Raimondii. 
Leaves little if at all lustrous. 
Petioles to 1 cm. long. 
Leaves elliptic, merely acute at each end. 
Flowers 4 mm. wide . Nectandra viburnoides. 

Flowers 7 mm. wide. 

Nectandra mamnonensis. 

Leaves cuneately narrowed to the base. 

Ocotea Tessmannii. 

Petioles to 2 cm. long . . . Ocotea albopunctulata. 

E. Leaves ample, mostly 2 to several decimeters long, or 8 cm. 
wide or wider. 

P. Leaves about twice longer than broad, often broadly elliptic 
or obovate, or at any rate mostly less than 3 times longer 
than broad. 

Q. Leaves pubescent beneath, obviously but sometimes 
finely so. 

Leaves subcordate at the base; petioles to 4 cm. long. 

Persea subcordata. 

Leaves acute to acuminate at the base or, if rounded, 
the petioles mostly under 2.5 cm. long. 

Petioles mostly 2.5 cm. long or longer. 

Branchlets more or less acutely angled or distinctly 
ribbed Persea ferruginea* 

Branchlets not as above. 

Inflorescence large, about half as long as the leaves 
or longer. 

Inflorescence long-peduncled, subequaling the 
leaves Nectandra Arnottiana. 

Inflorescence short-peduncled, much shorter 
than the leaves. . .Persea boliviensis. 



840 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Inflorescence only about 3 cm. long. 

Persea peruviana. 
Petioles mostly 2 cm. long or shorter. 

Leaves rounded at the apex. 

Pleurothyrium chrysophyllum. 
Leaves acute or acuminate. 

Branchlets rusty- or yellow-pubescent. 
Leaves mostly 10 cm. wide or narrower. 

Reticulation of the leaves not conspicuous. 

Pleurothyrium cuneifolium. 
Reticulation of the leaves conspicuous. 

Nectandra reticulata. 
Leaves mostly 7-8 cm. wide. 

Nectandra myriantha. 
Branchlets minutely gray-tomentose. 

Ocotea ovalifolia. 

Q. Leaves glabrous beneath, or the pubescence minute and 
very obscure. 

Base of the leaves cuneate. 

Petioles grooved above; fruits (so far as known) soon 
exserted. 

Flowers 3-4 mm. long, perfect Ocotea cuneifolia. 

Flowers 2-3 mm. long, dioecious Ocotea obovata. 

Petioles plane above; fruits (young) enclosed in the peri- 
anth Endlicheria robusta. 

Base of leaves rounded or shortly acute. 
Leaves thin, more or less acuminate. 

Branchlets or inflorescence velvety-puberulent. 

Endlicheria Tessmannii. 
Branchlets glabrous or glabrate. 

Leaves prominently reticulate-veined on both sides. 

Endlicheria glaberrima. 
Leaves not prominently reticulate-veined. 

Inflorescence pubescent . Nectandra capanahuensis. 

Inflorescence glabrous Ocotea olivacea. 

Leaves coriaceous or subcoriaceous, or obtuse. 



FLORA OF PERU 841 

Inflorescence glabrate or pubescent, but not densely 

sericeous. 
Leaves finely reticulate- veined. 

Ovary densely pilose; leaves rather thin; flowers 
6-7 mm. long Persea americana. 

Ovary glabrous. 

Leaves obtuse or rounded at the base (usu- 
ally), rather thin; flowers 4-6 mm. long; 
pedicels 2-5 mm. long or longer. 

Persea caerulea. 

Leaves acute at the base; flowers 7-8 mm. 
long; pedicels 2 mm. long. 

Persea Raimondii. 

Leaves scarcely reticulate-veined or laxly so. 

Acrodiclidium macrophyllum. 

Inflorescence densely sericeous-pilose or tomentulose. 
Leaves rigid; perianth tube short. . .Ocotea caniflora. 

Leaves flexible; perianth tube fleshy, conspicuous. 

Endlicheria paradoxa. 

P. Leaves about 3 times longer than broad, often lance-elliptic. 

R. Leaves glabrous beneath or glabrate (cf. Endlicheria gla- 
berrima and Cryptocarya robusta). 

Branchlets and inflorescence velvety brown- or red-tomen- 
tose. 

Inflorescence several decimeters long; perianth tube 2 
mm. long Pleurothyrium maximum. 

Inflorescence about 1 cm. long; perianth tube 1 mm. long. 

Pleurothyrium Williamsii. 

Branchlets and inflorescence puberulent or glabrous. 
Branchlets terete, or at least neither sulcate nor angled. 
Branchlets tomentulose. 

Petioles 2 cm. long Nectandra Macbridei. 

Petioles 1 cm. long Nectandra yarinensis. 

Branchlets glabrous. 

Leaves narrowed to or acute at the base. 

Lateral nerves of the leaves few. 

Nectandra Raimondii. 



842 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Lateral nerves of the leaves many. 

Pleurothyrium Krukovii. 
Leaves rounded at the base. 

Acrodididium macrophyllum. 

Branchlets, at least the young ones, angled or sulcate. 
Pedicels mostly 2-4 mm. long. 
Leaves glabrous Ocotea ucayaliensis. 

Leaves obscurely and minutely pubescent on the 
nerves Ocotea aurantiodora. 

Pedicels very short or wanting. 

Inflorescence dense; leaves obscurely reticulate- 
veined Ocotea amplissima. 

Inflorescence very open; leaves conspicuously reticu- 
late-veined Ocotea grandifolia. 

R. Leaves pubescent beneath, the hairs sometimes minute. 

Leaves subsessile Per sea grandis. 

Leaves petioled. 

Inflorescence persistently bracteate. 

Endlicheria bracteata. 
Inflorescence not persistently bracteate. 

Inflorescence extremely narrow, elongate, and lax; all 
filaments glandular Pleurothyrium bifidum. 

Inflorescence not extremely narrow, elongate, and lax; 
filaments never all glandular. 

Pedicels 1 mm. long or shorter. 

Inflorescence dense Aniba gigantifolia. 

Inflorescence very open. 

Branchlets ferruginous-velvety. 

Nectandra Macbridei, Ocotea magnifica. 

Branchlets hirsute Endlicheria verticillata. 

Branchlets puberulent Ocotea amplissima. 

Pedicels 2 (1.5) -4 mm. long. 

Petioles about 2 cm. long or longer; perianth not 
fleshy. 

Branchlets slightly sulcate; leaf veins obscure 
or very fine above. 



FLORA OF PERU 843 

Petioles to 3.5 cm. long; anthers sessile. 

Nectandra longifolia. 

Petioles about 2 cm. long; anthers pedicellate. 

Endlicheria Williamsii. 

Branchlets angled; leaves reticulate- veined 

above Ocotea aurantiodora. 

Petioles about 1 cm. long. 

Perianth fleshy, persistent; leaf nerves 4-5. 

Cryptocarya robusta. 
Perianth not persistent; leaf nerves 12-16. 

Endlicheria loretensis. 

1. BEILSCHMIEDIA Nees 

Hufelandia Nees; Bellota Gay. 

A genus well marked in fruit by the lack of a cup. Perianth tube 
shallow. Staminodia of series 4 foliaceous. Ovary subglobose, 
scarcely or not at all distinct from the broad, conic style that is stig- 
matic laterally at the tip. Cryptocarya R. Br., by error on Dombey 
labels sometimes accredited to Peru, but so far as known the speci- 
mens Chilean, has slenderer, urceolate flowers, the ovary immersed 
and in fruit included in the ligneous perianth tube. 

Beilschmiedia sulcata (R. & P.) Kosterm., comb. nov. Laurus 
sulcata R. & P. Fl. Peruv. 4: pi. 356. 1802; Laurogr. pi. 11. Persea 
sulcata Meissn. in DC. Prodr. 15, pt. 1: 54. 1864. Hufelandia sul- 
cata Nees, Linnaea 21: 494. 1848. H. latifolia Nees, Syst. Laur. 
674. 1836. 

A tree or shrub with slender, angled, at first silky-strigose branch- 
lets and rather coriaceous, finally glabrate, orbicular-elliptic, sub- 
obtuse leaves 8-18 cm. long and 6-12 cm. wide, reticulate-veined, 
especially above; petioles 1-2 cm. long; inflorescence tomentulose, 
subequaling the leaves; flowers 2.5-3 mm. long, on pedicels to 3 mm. 
long; anthers (outer) sometimes emarginate, the connective elongate; 
fruit to 3.5 cm. long and nearly 2 cm. in diameter, the pedicels 3-5 
mm. long. Kostermans notes that the drawing of Ruiz and Pavon 
shows an anther with 4 cells, probably in error. 

Huanuco: Muna, Ruiz & Pavon, type. Cajamarca: Tambillo, 
Raimondi. Cutervo (Jelski 187). San Carlos (Mathews 1433, type 
of H. latifolia). San Martin: San Roque, Williams 7077 (det. Koster- 
mans). North to Costa Rica. "Tashango Colorado," "ushun 
muena." 



844 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

2. AJOUEA Aubl. 

Like Beilschmiedia, but the perianth tube conspicuous and only 
the 2 outer series of stamens fertile, the 2 inner modified to stamino- 
dia, or rarely only 1 row, or less rarely the first 3 rows, fertile. Sta- 
minodia of series 4 foliaceous, rarely almost stipitiform. Cup 
usually little developed, borne on a thick, fleshy pedicel, the margin 
entire or with 6 large teeth. 

Ajouea tambillensis Mez, Jahrb. Bot. Gart. Berlin 5: 33. 1889. 
A. Jelskii Mez, loc. cit. 

Branchlets densely yellowish-tomentose, finally glabrous; petioles 
glabrous, 1-1.5 cm. long; leaves glabrous except for a few hairs on 
the midnerve beneath, coriaceous, elliptic or elliptic-lanceolate, acute 
at the base, shortly obtuse-acuminate or obtuse, 7-15 cm. long, 
3.5-6 cm. wide, the lateral nerves 6-12, scarcely prominent, the veins 
reticulate beneath; panicles axillary, pyramidal, many-flowered, gla- 
brous even to the flowers, these 2.5-3 mm. long, the equal sepals 
pilose within; outer 6 stamens pilose, the ovate anthers with a con- 
spicuous connective; staminodia to 1 mm. long, stipitate; ovary gla- 
brous, 1 mm. long, the style somewhat shorter . Ajouea rubra A. C. 
Smith, Phytologia 1: 115. 1935, of Amazonian Brazil, has broad peri- 
anth lobes, filiform staminodia, the leaves 4-6 cm. wide, and only the 
2 outer series of stamens fertile. Thus it resembles A . scandens Ducke, 
a vine, with long-petioled, broad leaves (Smith). 

Cajamarca: Tambillo, Prov. Cutervo, Jelski 185, type; 184, 195, 
type collections of A. Jelskii; Raimondi 4097 (det. Schmidt). 
Loreto: Manfinfa, upper Rio Nanay, Williams 1090. "Ciruelillo," 
"yakumuena." 

3. ENDLICHERIA Nees 

References: 0. C. Schmidt, Repert. Sp. Nov. 31: 170-179. 1933; 
Kostermans, Recueil Trav. Bot. Ne"erl. 34: 500-557. 1937. 

Resembling Aniba except that the flowers are dioecious, and, 
according to Kostermans, with differently shaped anthers and usually 
a shallow, fleshy fruit cup, the pedicel fleshy and enlarged. Third 
series of stamens rarely 4-celled. Style seldom as long as the usually 
glabrous ovary. It seems desirable at present to maintain these 
established groups, but it may be remarked that they may be shown 
eventually to constitute one genus, for the distinctions may be rela- 
tive. In Ocotea the flowers may be perfect or dioecious, and at least 
one species of Aniba has anthers resembling those of Endlicheria. 



FLORA OF PERU 845 

If the fruit character is constant, it may prove to be the funda- 
mental distinction. 

For other keys compare the two joint keys under Aniba to Aniba, 
Endlicheria, and Acrodiclidium. 

Branchlets and leaves glabrate, or the latter sparsely pubescent be- 
neath, especially on the nerves, the former puberulent (see 
also E. Tessmannii, with long petioles). 

Inflorescence branches glabrous or glabrate; leaves ample, often 
25-30 cm. long or longer. 

Leaves membranous; filaments pilose E. pyriformis. 

Leaves coriaceous; filaments nearly suppressed E. paradoxa- 

Inflorescence branches more or less puberulent; leaves often me- 
dium in size (see also E. paradoxa}. 

Leaves glabrous or essentially so, or minutely sericeous at first. 
Branchlets and leaves beneath minutely sericeous. 

E. robusta. 

Branchlets and leaves not at all sericeous-pubescent. 
Nerve axils beneath somewhat tufted-pubescent. 

E. dysodantha. 
Nerve axils beneath not pubescent. 

Pedicels 1-2 mm. long E. mishuyacensis. 

Pedicels 3-5 mm. long E. rubriflora. 

Leaves more or less pubescent beneath on the nerves. 
Base of the leaves cuneate; third series of anthers 2-celled. 
Leaves alternate, hirsutulous on the nerves beneath. 

E. hirsuta. 
Leaves subverticillate, puberulent on the nerves beneath. 

E. Cocuirey. 
Base of the leaves acute; third series of anthers 4-celled. 

E. anomala. 
Branchlets, and usually also the leaves beneath, conspicuously and 

softly pubescent. 

Pubescence, at least on the branchlets, distinctly tomentose or 
rarely villous and nearly always, apparently typically, reddish 
in color. 
Petioles short, 6-20 mm. long. 

Flowers glabrous, at least the staminate ones. 

Petioles 1-2 cm. long; flowers red-brown. . . .E. arunciflora. 



846 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles rarely 1 cm. long; flowers yellow-green. 

E. verticillata. 
Flowers pubescent. 

Leaves coriaceous or rigid, ovate, with an obtuse or acute base. 

E. Szyszylowiczii. 
Leaves chartaceous-elliptic, contracted at the base. 

Leaves rarely 20 cm. long E. Sprucei. 

Leaves mostly longer than 20 cm E. Cocuirey. 

Petioles 2-5 cm. long. 
Leaves densely tomentulose beneath. 

Leaves rigid-coriaceous E. tomentella. 

Leaves chartaceous E. Williamsii. 

Leaves soon glabrous beneath E. Tessmannii. 

Pubescence grayish rusty, usually rather loose or, if appressed, 

sericeous. 

Pubescence often closely appressed, the leaves thus lustrous- 
sericeous beneath E. Lhotzkyi. 

Pubescence of the leaves loose, hirsute or villous or, if tomentu- 
lose, the leaves not sericeous beneath. 
Leaves broader than oblong-linear. 
Leaves more or less caudate-acuminate. 
Leaf veins conspicuously reticulate. 

Lateral nerves of the leaves 15-17 E. bracteata. 

Lateral nerves of the leaves 6-8 E. debilis. 

Leaf veins scarcely reticulate above E. Poeppigii. 

Leaves acute or obtusely acuminate E. hirsuta. 

Leaves oblong-linear, conspicuously bullate E. bullata. 

Endlicheria anomala Nees ex Meissn. in DC. Prodr. 15, pt. 1: 
173. 1864; 508. Geoppertia anomala Nees, Syst. Laur. 370. 1836. 

Branchlets lightly tomentose; petioles to 1.5 cm. long; blades 
papyraceous, somewhat sericeous-hirsute beneath, elliptic, acumi- 
nate, 3-8 cm. wide, reticulate-veined on both sides; flowers obscurely 
strigose, 2 mm. long; perianth tube very short; stamens of the third 
series conglutinate below the ovate, truncate, 4-celled anthers, these 
equaled by the glabrous filaments; ovary of the staminate flower 
none, of the pistillate flower glabrous, the style short; fruit to 13 mm. 
long; cup nearly flat, 4 mm. broad, the obconic pedicel 5 mm. long, 






FLORA OF PERU 847 

3 mm. thick at the summit. A tree, sometimes 15 meters high. 
Williams states that the bark, when incised, yields a small amount 
of sweet, brown resin. Negs. 7304, 7305. 

Loreto: Yarina-cocha, Tessmann 3439 (det. Schmidt). Near 
Iquitos, 100 meters, Tessmann 5126; Williams 1494, 1495, 1500, 3670, 
8144 (det. Kostermans); Klug 728; Killip & Smith 27192, 29959 
(det. Kostermans). Pebas, Williams 1877 (det. Schmidt). Rio 
Itaya, Williams 15, 176, 3307 (det. Kostermans), 176 (det. Schmidt). 
Rio Nanay, Williams 609 (det. Schmidt). Brazil; Guiana. "Nana," 
'muena del agua," "canela," "canela-muena." 

Endlicheria arunciflora (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 
5: 131. 1889; 517. Ampelodaphne arunciflora Meissn. in DC. Prodr. 
15, pt. 1:81. 1864. 

Branchlets stout, reddish-brown-villous; petioles 1-2 cm. long; 
blades rigid, pilose on the nerves, even in age, pubescent beneath, 
the nerves immersed above, beneath prominently ribbed and loosely 
reticulate, lanceolate, acuminate, to about 18 (-25) cm. long and 4 
(-9) cm. wide; staminate inflorescence subtended by persistent bracts 
5-10 mm. long, pilose only in the axils, diffusely pyramidate-panicu- 
late, pendulous, longer than the leaves; flowers subsessile, yellowish 
red, glabrous, 1 mm. long, 2.5 mm. wide; perianth tube conspicuous; 
filaments glabrous, those of the third series (glands small) subequal- 
ing the suborbicular anthers, the connective not produced. Fruit 
and pistillate flowers unknown. A tree of 20 meters. Illustrated, 
Mart. Fl. Bras. 5, pt. 2: pi. 57. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 1403, 1264 
(det. Schmidt). Venezuela; Brazil. 

Endlicheria bracteata Mez, Repert. Sp. Nov. 16: 306. 
1920; 520. 

A small tree with rusty-hispid-villous branches and inflorescence, 
the branches terete, the inflorescence apparently pendent and about 
equaling the thin, obovate, caudate-acuminate leaves; leaves 
subverticillate, to 30 cm. long and 10 cm. broad, conspicuously reticu- 
late-veined, glabrate above, softly pilose beneath; bracts and bract- 
lets persistent; perianth tube subglobose; flowers 2 mm. long, 3.5 
mm. wide, cream-colored or white, gray-hirsute, subsessile; filaments 
and anthers equal, glabrous, the anthers depressed-orbicular, broadly 
emarginate, the connective not produced; basal glands rather large, 
short-stipitate; ovary glabrous. Kostermans observes that the type 
is staminate, pistillate flowers and fruit being unknown. A tree of 



848 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

6 meters. Illustrated, Kostermans, Recueil Trav. Bot. Ne'er!. 34: 
519. Neg. 3813. 

Amazonas: Moyabamba, Weberbauer 4680, type. San Martin: 
Zepelacio, 1,100 meters, Klug 3745 (det. Kostermans). Loreto: 
Pumayacu, 600-1,200 meters, Klug 3187. 

Endlicheria bullata Ducke, Archiv. Jard. Bot. Rio Janeiro 4: 
190. 1925; 511. 

A striking shrub because of its strongly bullate, linear-oblong, 
acuminate leaves, these 20-30 cm. long, about 3 cm. wide, lustrous 
above, nearly dull beneath and conspicuously spreading-hispid on 
the reticulate veins; petioles to 1 cm. long, mostly shorter, hispidu- 
lous, like the young branches; panicle usually shorter than the leaves, 
reddish-tomentose; pedicels scarcely 2 (-4) mm. long, with small, 
subpersistent bractlets; staminate flowers about 5 mm. broad, the 
.perianth tube very short, the segments equal, subobtuse, glabrous 
within, spreading; outer dilated filaments and introrse, truncate an- 
thers subequal, the glands of series 3 large, the extrorse anthers much 
smaller than the inner; filaments glabrous, the outer suppressed. A 
shrub about 5 meters high (Krukoff). Illustrated, Kostermans, 
Recueil Trav. Bot. Ne"erl. 34: 510. Neg. 3814. 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5780 (det. Koster- 
mans). Amazonian Brazil. 

Endlicheria Cocuirey Kosterm. Recueil Trav. Bot. Ne'er!. 34: 
522. 1937. 

Branchlets minutely but densely reddish- tomentulose; petioles 
to 10 mm. long; leaves subverticillate, chartaceous, glabrous above 
except for the costa, beneath minutely and sparsely pubescent on 
the 12-17 prominent nerves, densely reticulate-veined on both sides, 
elliptic, 18-32 cm. long, 6.5-11 cm. wide, attenuate at the base, 
acuminate; staminate panicles many-flowered, densely tomentulose, 
to 9 cm. long; pedicels hirsute, 1 mm. long; flowers hirsute, white, 
3 mm. wide, 1.5 mm. long, the tube glabrous within, shorter than 
the equal sepals; stamens glabrous, the outer anthers as long as the 
filaments; glands small; sterile ovary 1 mm. long. Fruit and pis- 
tillate flowers unknown. A tree of 4 meters (Klug). Differs from 
E. bracteata in its larger leaves and different indument (Kostermans). 

Loreto: Florida, 180 meters, Klug 2253, type. "Hioma cocuir-ey." 

Endlicheria debilis Kosterm. Recueil Trav. Bot. Ne'er!. 34: 
555. 1937. 



FLORA OF PERU 849 

A tree of 6-7.5 meters; known only in fruit, but in pubescence 
and in leaf form resembling E. bracteata; allied, according to the 
author, to E. mishuyacensis ; leaves alternate, 12-17 cm. long, 3.5-5 
cm. wide; fruit cup smooth, to 12 mm. high, 18 mm. wide, decurrent 
into the fleshy pedicel, this 3 cm. long, 5 mm. broad at the top. 

Loreto: Balsapuerto, 150-350 meters, Kittip & Smith 28400, type. 

Endlicheria dysodantha (R. & P.) Mez, Jahrb. Bot. Gart. 
Berlin 5: 118. 1889; 537. Laurus dysodantha R. & P. Fl. Peruv. 4: 
pi. 355. 1802; Laurogr. pi. 14- Geoppertia dysodantha Nees, Linnaea 
21:514. 1848. 

A shrub or small tree, essentially glabrous except for tufts of 
hairs beneath in the nerve axils of the leaves and a very short, white 
pilosity on the young flowers; petioles 10-18 mm. long; blades papy- 
raceous, elliptic-lanceolate, acuminate, 3.5-7 cm. wide, reticulate- 
veined on both sides; inflorescences much shorter than the leaves; 
flowers 2 mm. long, the tube very short; outer anthers subsessile; 
connective acuminately produced; inner filaments broad; glands 
large, conglutinate. A small tree, to 12 meters high, with a trunk 
diameter of about 6 cm. Neg. 27587. 

Loreto: Mouth of Rio Santiago, Tessmann 3953, 3999, 4248 (det. 
Schmidt). Middle Rio Blanco, Tessmann 3058a. Huanuco(?): 
Macora, Ruiz, type. Puno: Sangaban, Lechler 2317. Bolivia; Ama- 
zonian Brazil. 

Endlicheria Lhotzkyi (Nees) Mez, Jahrb. Bot. Gart. Berlin 
5:122. 1889; 544. Ocotea Lhotzkyi Nees, Syst. Laur. 475. 1836. 

A tree with gray-green, coriaceous leaves, these becoming gla- 
brate above, permanently silvery- or brown-lustrous-sericeous 
beneath; branchlets angled, densely reddish-pubescent; petioles 1.5- 
2.5 cm. long; leaves ovate-lanceolate, rather obtuse at the base, short- 
acuminate, 11-17 cm. long, 4-6.5 cm. wide, the lateral nerves often 
very prominent beneath, the veins laxly reticulate; inflorescences 
loosely thyrsoid-paniculate, much shorter than the leaves, densely 
pilose; flowers 2.5 mm. long; perianth tube campanulate-conic, the 
lance-ovate lobes equal; pedicels obsolete, or to 1 mm. long; outer 
stamens fertile, the anthers ovate, longer than broad, obtuse, 2-celled, 
subequaling the sparsely pilose filaments; connective little produced. 
Probably here belong the Peruvian records for the typically Antil- 
lean E. sericea Nees, with inflorescence and leaves subequal, and 
pedicellate flowers. A tree almost 30 meters high (Krukoff). Neg. 
3817. 



850 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rio Acre: Mouth Rio Macauhan, Krukoff 5279 (det. Koster- 
mans). Brazil. 

Endlicheria mishuyacensis A. C. Smith, Bull. Torrey Club 
58: 102. 1931; 538. 

Branchlets and foliage evanescently strigose; petioles rugose, 
slender, 3-10 mm. long; blades oblong-lanceolate or narrowly obo- 
vate, attenuate at the base, obtusely acuminate, 7-16 cm. long, 2-2.5 
cm. broad, chartaceous, soon glabrous, densely and prominently 
reticulate-veined on both sides; inflorescence 10-20-flowered, 3-6 cm. 
long, strigillose, the flowers 2-2.5 mm. long, white; pedicels to 1.5 
mm. long; perianth tube obconic; outer anthers sessile, the connec- 
tive slightly protruding, the inner ones nearly as large, on broad 
filaments, these eglandular; ovary glabrous, longer than the style; 
cup thick, with a thin margin, 3 cm. wide, 1 cm. high, the pedicel to 
1 cm. long. Compared by the author with E. dysodantha. A tree 
of 6-10 meters. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 204, type; 
411, 621, 703; Killip & Smith 29870. Upper Rio Nanay, Williams 
1124- Balsapuerto, Killip & Smith 28400. Amazonian Brazil. 

Endlicheria paradoxa Mez, Jahrb. Bot. Gart. Berlin 5: 114. 
1889; 534. 

A glabrous, large-leaved tree, to 25 meters high ; branchlets rusty- 
pilose at the tips; leaves lustrous above, beautifully and closely reticu- 
late-veined on both sides, elliptic-lanceolate; inflorescence at first 
sericeous-tomentulose, paniculate, much shorter than the leaves; 
pedicels 1-4 mm. long; staminate flowers 3-4 mm. long, the subglo- 
bose, very fleshy tube strongly accrescent after an thesis; stamens 
fertile; anthers introrse, elongate- triangular-ovate, with produced 
connective, simulating those of Cryptocarya; glands small, congluti- 
nate; ovary glabrous, the style short. The stamens of the inner row 
are fertile, the valves developed (Kostermans). The blades, accord- 
ing to Schmidt, are as much as 38 cm. long and 15.5 cm. wide, with 
thick petioles to 4 cm. long. The type gives the impression of being 
abnormal ( Kostermans) . 

Lima: Santa Cruz, 2,100 meters (Pearce). Guiana(?). 

Endlicheria paniculata (Spreng.) Macbr., comb. nov. Citrosma 
paniculata Spreng. Syst. 2: 545. 1825. E. hirsuta Nees, Linnaea 8: 
38. 1833; 549. Cryptocarya hirsuta Schott in Spreng. Syst. 4, pt. 2: 
405. 1827. Geoppertia longifolia Nees, Syst. Laur. 368. 1836. E. longi- 



FLORA OF PERU 851 

folia Mez, Jahrb. Bot. Gart. Berlin 5: 119. 1889. Aniba hirsuta Pax 
ex Sampaio, Comm. Linh. Telegr. Matto Grosso, Annexe 5, Bot. 10: 
9. pi. 8. 1917. 

A shrub or a large tree, with rusty or grayish-tomentose branch- 
lets and panicles, the leaves chartaceous to coriaceous, ovate-elliptic- 
lanceolate, little longer than the panicles, acute at both ends, 
glabrate above, usually pilose-hirsute beneath, even in age; petioles 
to 1.5 cm. long; blades usually about 15 cm. long, 3.5-5 cm. wide, 
laxly if at all reticulate- veined above; pedicels 1.5-4 mm. long, the 
long-pilose flowers 2-3 mm. long, the tube very short; filaments gla- 
brous or pilose; connective little produced; glands large, basal or 
suprabasal; cup shallow, 4-7 mm. high, 10-15 mm. broad, the pedi- 
cels 5-12 mm. long. According to Kostermans, the glands may be 
attached only slightly above the base or one-third above (E. longi- 
folia) ; variable also as to foliage, this even glabrous. E. Klugii 0. C. 
Schmidt, of adjacent Colombia, is similar to E. tomentella (which see 
below), but the larger leaves (20-40 cm. long) are only short-pilose 
beneath, the perianth tube narrowly obconic, 2 mm. long, the pilose 
outer filaments S-curved. E. Krukovii (A. C. Smith) Kosterm., of the 
upper Amazon, has less pubescent, caudate-acuminate leaves. Negs. 
19260, 3818. 

Huanuco: Cochero, Poeppig, type of E. longifolia. Loreto: Pongo 
de Manseriche (Mexia 6329; det. Kostermans). Brazil; Paraguay. 

Endlicheria Poeppigii Kosterm. Recueil Trav. Bot. Ne'er!. 34: 
555. 1937. Geoppertia hirsuta Nees, var. hirsutior Mejssn. in DC. 
Prodr. 15, pt. 1: 172. 1864. 

Branchlets rusty-villous with long, somewhat spreading hairs, 
the branches glabrate; petioles 5-8 mm. long; blades elliptic, 15 cm. 
long, 6 cm. wide, mostly about 10 cm. long and half as wide, more or 
less narrowed to the rounded or acutish base, nearly caudate-acumi- 
nate, chartaceous, glabrous above or the impressed midnerve puberu- 
lent, pilose-hirsute beneath on the prominent nerves and reticulate 
veins, the nerves about 5 on each side, impressed above; inflorescence 
2-4 (-15) cm. long, the rachis rusty-villous, the calyx ashy-strigose, 
perhaps 1 mm. long (immature) ; staminate flowers 4 mm. broad. 
Neg. 35000. 

Loreto: Yurimaguas, Poeppig 2298, type; (King 2833; det. 
Kostermans). 

Endlicheria pyriformis (Nees) Mez, Jahrb. Bot. Gart. Berlin 
5: 116. 1889; 540. Cryptocarya pyriformis Nees, Syst. Laur. 220.1836. 



852 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

E. glaberrima Mez, Bull. Herb. Boiss. II. 5: 236. 1905. Aniba flexuosa 
A. C. Smith, Phytologia 1: 117. 1935. 

Glabrous, with ample, thin, elliptic, short-acuminate leaves, 
conspicuously reticulate- veined on both sides; inflorescence few- 
flowered, laxly squarrose-paniculate; flowers green or reddish, to 4 
mm. long, the tube infundibuliform, 2 mm. long, twice longer than 
the segments; pedicels 2-4 mm. long; outer anthers 3 times longer 
than the pilose filaments; connective slightly produced; glands of 
series 3 minute, hirsutely stipitate, attached above the filament base; 
style obvious; cup 5-10 mm. high, 7-15 mm. wide, the fleshy, red 
pedicel 7-14 mm. long. E. formosa A. C. Smith, 535, has white, 
subglobose flowers 2 mm. long, with large, sessile glands; Amazonian 
and to be expected. E. pyriformis is said to attain a height of 
30 meters. Neg. 3820. 

Loreto: Between Yurimaguas and Balsapuerto (Killip & Smith 
28291; det. Kostermans). Pongo de Manseriche, Tessmann 4736 
(det. Kostermans). Yurimaguas, Vie 6296, type of E. glaberrima. 
Bolivia to the Guianas. 

Endlicheria robusta (A. C. Smith) Kosterm. Recueil Trav. 
Bot. Ne"erl. 34: 556. 1937. Cryptocarya robusta A. C. Smith, Bull. 
Torrey Club 58: 97. 1931. 

Essentially glabrous except that the branchlets and leaves beneath 
at first are minutely sericeous with a dense pubescence; petioles 10-12 
mm. long; blades oblong or elliptic, cuneate at the base, obtuse- 
acuminate, 17-22 cm. long, 6-9 cm. wide, subcoriaceous, olivaceous, 
the lateral nerves 4-6, rather prominently reticulate- veined beneath ; 
panicles few-flowered, to 7 cm. long, sericeous; fruiting pedicels 3-7 
mm. long; remnants of sepals and stamens on young fruits show the 
connective slightly protruding, the outer anthers ovate-triangular, 
merging with pilose filaments as long; glands rather small; ovary 
glabrous. The shape of leaves and the reticulation point to a rela- 
tionship with E. debilis and E. hirsuta, but the shape of the anthers 
to the group of E. dysodantha ( Kostermans). 

Junin: San Nicolas, Pichis Trail, 1,100 meters (Killip & Smith 
26077, type). 

Endlicheria rubriflora Mez, Jahrb. Bot. Gart. Berlin 5: 
494. 1889; 512. Aniba reticulata A. C. Smith, Bull. Torrey Club 
58: 99. 1931. 

A tree; branchlets soon glabrous; petioles 6-15 mm. long; blades 
oblong-ovate, cuneate at the base, obtusely and abruptly acuminate 



FLORA OF PERU 853 

(acumen to 2.5 cm. long), 10-28 cm. long, 3.5-9 cm. wide, charta- 
ceous, glabrous or minutely and sparsely pilose beneath, abundantly 
reticulate-veined on both sides; panicle many-flowered, in bud slightly 
pilose but soon glabrous, to 12 cm. long; flowers at first slightly pilose 
outside, the staminate with the tube glabrous within, scarcely 1 mm. 
long, the pistillate similar but the sepals caducously tomentulose 
within; pedicels 2-3 mm. the pistillate 5 mm. long; stamens all fertile, 
the anthers shorter than the glabrous filaments; basal glands very 
large, conglutinate; style obsolete, the ovary glabrous; cup and pedi- 
cel fleshy, the cup 2.5 cm. wide, the pedicel 1 cm. long. 

Loreto: Yurimaguas, 135 meters, Killip & Smith 28050, type of 
A. reticulata. Pumayacu, Klug 3178(1}. Colombia. 

Endlicheria sericea Nees, Linnaea 8: 38. 1833; 122; 545. 

A small to tall (35 meters) tree, sericeous-tomentulose on the 
branchlets, leaves beneath, and inflorescence, even to the flowers; 
leaves chartaceous-coriaceous, finally lustrous and glabrous above, 
reticulate-veined on both sides, ovate-elliptic, acuminate, sometimes 
20 cm. long and half as wide; pedicels 3-7 mm. long; staminate 
inflorescence pyramidate-paniculate, subequaling the leaves, the 
squarrose pistillate inflorescence shorter and few-flowered; staminate 
flowers 2.5 mm. the pistillate 4 mm. long; perianth tube and segments 
subequal; anthers of third series biglandular; connective produced; 
ovary included in the tube, the half -included fruit ovoid, the perianth 
lobes subpersistent. The wood is highly valued for manufacture of 
fine furniture (Herrera). 

Ancash: Moro (Pearce). Cuzco: Provincias Convention and 
Paucartambo (Herrera). Bolivia to Venezuela and West Indies. 
"Laurel bianco." 

Endlicheria Sprucei (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
125. 1889; 514. Geoppertia Sprucei Meissn. in DC. Prodr. 15, pt. 1: 
172. 1864. 

A shrub or tree, the slender branchlets almost velvety-red-tomen- 
tose; petioles to 8 mm. long; blades chartaceous, the primary nerves 
at least tomentose, usually on both sides, especially beneath reticu- 
late-veined, elliptic, narrowly long-acuminate at the apex, rarely 18 
cm. long and 7 cm. wide; inflorescences crowded, paniculate, shorter 
than the leaves, the long-pilose flowers 1.5 mm. long, 5 mm. wide, 
the pistillate 2 mm. long; perianth tube very short, the segments and 
limb subequal; filaments short, united with the sepals, with 2 small 
basal glands within; glands of the inner stamens large; anther con- 



854 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nective not produced. A shrub of 4-6 meters, the flowers red or red- 
brown (King). Neg. 3821. 

Loreto: Mishuyacu, 100 meters, King 272, 273 (det. Schmidt). 
Colombia; Brazil. 

Endlicheria Szyszylowiczii Mez, Jahrb. Bot. Gart. Berlin 5: 
121. 1889; 548. 

A shrub or tree with thick, velvety-pubescent branchlets and 
rigid-coriaceous, ovate leaves, rusty-tomentose on both sides, 
becoming glabrate above; blades obtuse at the base, acuminate, 
reticulate-veined beneath, 4-7.5 cm. wide; inflorescence thyrsoid- 
paniculate, shorter than the leaves; flowers subsessile, or the pedicels 
to 3 mm. long, densely villous, 2.5 mm. long, the tube and lobes 
subequal; filaments hirsute; otherwise likeE". hirsuta; fruit and pistil- 
late flowers unknown. Falsely called "lanche pardo" (Raimondi) . 

Cajamarca: Tambillo, Raimondi (det. Schmidt). Cutervo (J el- 
ski 165, type; 191, 195}. Bolivia. "Ishpingo rufo," "oberillo." 

Endlicheria Tessmannii 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 227. 1928; 521. 

Young branches and panicles subvelutinous; petioles 3.5-5 cm. 
long; blades broadly elliptic or obovate-elliptic, thin, glabrous or 
minutely pilose beneath on the nerves, more or less acuminate, 9-15 
cm. wide; inflorescence about 20 cm. long; staminate flowers 2 mm. 
long, white, on pedicels 1.5-2 mm. long; tube and lobes subequal; 
stamens glabrous, the connective not protruding, the outer, broadly 
ovate anthers on pilose filaments as long; basal glands rather large; 
fruit and pistillate flowers unknown. Allied to E. bracteata; the 
type, according to the collector, a tree of 8 meters, branching at 2.5 
meters, the trunk 15 cm. in diameter. E. macrophylla (Meissn.) Mez, 
523, has leaves densely hirsute beneath. Neg. 3822. 

Loreto: Near Iquitos, 100 meters, Tessmann 5146, type; Klug 
161, 26 (det. Kostermans). 

Endlicheria tomentella Mez, Jahrb. Bot. Gart. Berlin 5: 115. 
1889; 533. 

A large tree with angled, rufous-tomentulose branchlets and 
coriaceous, elliptic leaves, glabrous above, red-tomentose beneath, 
obscurely reticulate- veined ; blades acuminate, about 10 cm. (10-18 
cm.) long and half as wide; inflorescence cinereous- tomentulose, 
shorter than or equaling the leaves; pistillate flowers white, 2-2.5 
mm. long; broadly obconic tube and lobes subequal; filaments of the 



FLORA OF PERU 855 

outer series shorter than the anthers, of the inner longer; glands 
minute; anthers all 2-celled, the connective produced; cup verrucose, 
1 cm. high, 1.5 cm. broad, the pedicel 3 mm. long. A tree of 20-35 
meters (Pearce). Pubescence sometimes grayish. 
Ancash: Near Moro (Pearce, type). Bolivia. 

Endlicheria verticillata Mez, Bull. Herb. Boiss. II. 5: 235. 
1905; 518. 

Branches stout, densely reddish-yellow-hirsute; petioles to 10 
mm. long; blades elliptic, acuminate, to 25 cm. long, chartaceous, 
prominently reticulate-veined and loosely hirsute-pilose on the 
nerves on both sides; staminate inflorescence pyramidate-paniculate, 
often shorter than the leaves; bracts and bractlets persistent; flowers 
greenish yellow, glabrous, short-pedicellate, 1.5 mm. long, 3 mm. 
wide, the slender tube and lobes subequal; basal glands small, sessile; 
outer anthers broader than long, the elliptic inner ones on filaments 
as long; fruit and pistillate flowers unknown; ovary glabrous. A 
tree of 20 meters ( Ule). Neg. 3823. 

Loreto: Rio Itaya, Tessmann 5288. Amazonian Brazil. 

Endlicheria Williamsii 0. C. Schmidt, Repert. Sp. Nov. 31: 
177. 1933; 529. 

Branches and leaves beneath rather softly tomentulose, the leaves 
becoming glabrate above and dull, chartaceous, lanceolate or elliptic, 
20-25 cm. long, 6-9 cm. wide, the acumen to 2 cm. long, narrowed 
to the petiole, this 2 cm. long, the lateral nerves 8-10, reticulate; 
staminate inflorescence pyramidate-paniculate, 6-14 cm. long, lax, 
many-flowered, the slender pedicels to 2.5 mm. long, villous-pilose, 
the axils tomentose; pistillate inflorescence thyrsoid, few-flowered; 
staminate flowers slightly pilose outside, densely pilose within, the 
obconic tube 1 mm. long, the lobes twice as long, the similar pistillate 
flowers smaller; stamens glabrous; glands large, sessile; connective 
not produced; ovary glabrous, longer than the style. A tree of 23 
meters, with wide-spreading crown; fruit ovoid, with adherent, red 
calyx (Williams). E. Klugii 0. C. Schmidt, 530, has petioles 5-6.5 
cm. long and broader leaves. 

Loreto: Timbuchi, Rio Nanay, Williams 1002, 1003, 1004- Maqui- 
sapa, upper Rio Nanay, Williams 1193, type; 1203. "Isma muena," 
"pampa-muena," "muena blanca." 

4. ANIBA Aubl. 

Leaves alternate or verticillately crowded. Flowers perfect. 
Stamens 9, included, all fertile or rarely those of the third row sterile, 



856 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

this series usually with sessile glands. Fruit finally exserted from 
the usually simple-margined and verruculose cup. Preceding two 
keys to Aniba is a joint key to the genus withLicaria and Endlicheria, 
which could conveniently be treated as sections of Aniba. 

Vegetative key to Aniba sens. lat. (including Licaria and Endlicheria) 
A. Leaves large, 25-60 cm. long or longer. 

Leaves pilose or tomentose beneath, sometimes finely so. 
Leaves broader than oblong-linear. 
Flowers subsessile, the pedicels rarely 1 mm. long. 
Inflorescence tomentose, the bractlets deciduous. 

A. gigantifolia. 

Inflorescence villous, the bractlets persistent .E. bracteata. 
Flowers pedicellate, the bractlets 1.5 mm. long or longer. 
Pedicels mostly longer than 2.5 mm. ; flowers dioecious. 

E. loretensis. 
Pedicels rarely 2.5 mm. long; flowers perfect. 

L. macrophylla. 

Leaves oblong-linear, conspicuously bullate E. bullata. 

Leaves glabrous beneath or essentially so. 
Inflorescence branches glabrous. 

Stamens all fertile; leaves thin E. glaberrima. 

Stamens in part (outer or series 3) reduced to staminodia; 
leaves coriaceous. 

Petioles 4 cm. long; outer stamens fertile E. paradoxa. 

Petioles 7 cm. long; outer stamens reduced to staminodia. 

L. endlicheriaefolia. 
Inflorescence branches pubescent. 

Pedicels 1-4 mm. long. 
Staminate flowers 2-3 mm. long; stamens of series 3 fertile. 

Petioles 4-5 cm. long E. Tessmannii. 

Petioles to 1 cm. long E. Cocuirey. 

Staminate flowers 4 mm. long; stamens of series 3 reduced 

to staminodia E. paradoxa. 

Pedicels 6 mm. long L. macrophylla. 

A. Leaves small to medium-sized, mostly 10-20 cm. long, sometimes 

to 25 cm. long. 
B. Branchlets tomentose, villous, or pilose (cf. L. multiflora). 



FLORA OF PERU 857 

Branchlets densely gray-pilose; leaves pilose on both sides. 

E. verticillata. 

Branchlets villous, tomentose, or hirsute, the pubescence often 
reddish. 

Leaves long-acuminate or caudate-acuminate (see also E. 
debilis). 

Branchlets often red-tomentose; petioles 5-10 mm. long. 

Leaves thinnish, epapillose, with 5-6 lateral nerves. 

E. Sprucei. 

Leaves coriaceous, papillose beneath, with 10-12 nerves. 

A. cylindriflora. 

Branchlets rusty-sericeous; petioles longer. . . .L. latifolia. 

Branchlets rusty- villous; petioles 3-5 mm. long. 

E. Poeppigii. 
Leaves short-acuminate. 

Inflorescence much longer than the leaves . . . .E. arunciflora. 
Inflorescence equaling or shorter than the leaves. 

Leaves lustrous-sericeous beneath . .E. sericea,E.Lhotzkyi. 

Leaves not so pubescent. 
Leaves mostly shorter than 15 cm. 

Leaves pubescent on both sides, at least on the 
nerves E. Szyszylowiczii. 

Leaves soon glabrous above. 
Pubescence distinctly reddish. 

Trichomes obvious beneath, the vein reticula- 
tion obscure E. tomentella. 

Trichomes obscure beneath, the vein reticula- 
tion prominent A. firmula. 

Pubescence grayish-rusty. 

Pubescence tomentulose E. anomala. 

Pubescence villous E. Poeppigii. 

Leaves mostly 20 cm. long. 
Petioles about 2 cm. long. 

Leaves chartaceous E. Williamsii. 

Leaves rigid-coriaceous E. rosaeodora. 



858 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles to 1.5 cm. long. 
Leaf reticulation prominent on both sides. 

E. longifolia. 
Leaf reticulation prominent only beneath. 

Branchlets villous-hirsute E. hirsuta. 

Branchlets red-tomentulose A. firmula. 

B. Branchlets glabrous, or more or less puberulent-tomentulose, 

but the hairs usually deciduous. 
Leaves prominently reticulate-veined on both sides. 
Leaves thin; branchlets slightly tomentose; anthers of series 

three 4-celled E. anomala. 

Leaves and other characters otherwise, at least in part. 
Leaves glabrous except for axillary tufts of hairs beneath. 

E. dysodantha. 

Leaves more or less pubescent, or the axillary tufts lacking. 
Flowers many, often glabrous. 
Panicles pubescent. 

Pedicels 3-4 mm. long E. paniculata. 

Pedicels 1 mm. long L. Pucheri, L. limbosa. 

Panicles soon glabrous E. rubriflora. 

Flowers pilose, few E. mishuyacensis. 

Leaves prominently reticulate-veined only beneath, if at all. 
Reticulation prominent beneath, coarse or dense. 
Leaves mostly 4.5 cm. wide and shorter than 15 cm. 
Outer stamens fertile; branchlets soon glabrous. A. Muca. 
Outer stamens reduced to staminodia; branchlets puberu- 

lent A. Puchury-minor. 

Leaves mostly wider than 4.5 cm. and longer than 15 cm. 
Leaves usually 6 cm. wide or less; ovary glabrous. 

Branchlets smooth A. salicifolia. 

Branchlets verruculose A. Puchury-minor. 

Leaves 7-9.5 cm. wide; ovary pilose. . . .A. gigantifolia. 
Reticulation beneath obscure or fine. 

Leaves mostly 7-9 cm. wide L. latifolia. 

Leaves mostly 3-6 cm. wide. 

Pubescence reddish . . . .E. tomentella. 



FLORA OF PERU 859 

Pubescence not at all reddish. 

Inflorescence glabrous or slightly pilose. .L. armeniaca. 
Inflorescence tomentulose. 

Perianth lobes definitely unequal A. Canelilla. 

Perianth lobes equal. 

Leaves short-acuminate; filaments short. 

A. perutilis. 
Leaves conspicuously acuminate; filaments long. 

Leaves 2.5-3 (-4) cm. wide L. multi flora. 

Leaves 4.5-10 cm. wide. 
Leaves not minutely papillose beneath. 

Branchlets slender, lustrous; petioles gla- 
brous A. salicifolia. 

Branchlets thick, verruculose; petioles pu- 
bescent A. Puchury-minor. 

Leaves papillose beneath. 

Bractlets persistent; ovary glabrous. 

A. cylindrifolia. 
Bractlets deciduous; ovary pubescent. 

A. Goto. 

Technical key to Aniba sens. lat. (including Licaria and Endlicheria) 
Third series of stamens fertile, the outer modified or lacking. 
Perianth lobes reflexing and longer than the obsolete tube. 

L. armeniaca. 

Perianth tube developed, often subequaling or longer than the 
lobes. 

Leaves soon glabrous, or slightly tomentulose beneath; pedicels 
5-6 mm. long. 

Inflorescence short, few-flowered; staminodia of series 4 
wanting L. Pucheri. 

Inflorescence elongate, 20-40-flowered ; staminodia of series 4 

present. 

Leaves cordate at the base; flower tube glabrous within. 

L. macrophylla. 
Leaves not cordate; flower tube sericeous within. 

L. endlicheriaefolia. 



860 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves pubescent beneath; pedicels short. 
Leaves ovate-lanceolate. 

Outer staminodia present L. multi flora. 

Outer staminodia wanting L. Quirirafuina. 

Leaves broadly elliptic L. latifolia. 

All series of stamens fertile or only the inner 3 sterile. 
Flowers perfect (see technical key to Aniba). 
Flowers dioecious. 

Stamens of the 2 outer series fertile E. paradoxa. 

Stamens of the 3 outer series fertile. 
Anthers of series three 4-celled. 

Leaves mostly shorter than 15 cm., somewhat 3-nerved. 

E. anomala. 

Leaves much larger, pinnate-nerved E. loretensis. 

Anthers of series three 2-celled. 
Third series of stamens biglandular above the base. 

E. longifolia. 

Third series of stamens eglandular, or all the stamens 
biglandular at the base. 

Stamens all biglandular E. Sprucei. 

Stamens only of series 3 biglandular, or these eglandular. 
A. Stamen connective more or less produced. 

Leaves at maturity at least minutely pubescent 

beneath. 

Pubescence a dense pilosity or tomentum. 
Leaves mostly about 20 cm. long; pubescence 
of the branchlets spreading. 

Leaves acute E. hirsuta. 

Leaves more or less caudate E. bracteata. 

Leaves mostly 10-15 cm. long; pubescence of 

the branchlets tomentose. 
Pubescence more or less persisting on the leaf 

veins above E. Szyszylowiczii. 

Pubescence on leaves above soon deciduous. 
Leaves pilose-sericeous beneath. 

E. Lhotzkyi, E. sericea. 



FLORA OF PERU 861 

Leaves red-tomentose beneath. 

E. tomentella. 
Leaves beneath with minute or sparse pubescence. 

Bractlets persistent E. bracteata. 

Bractlets deciduous E. robusta. 

Leaves at maturity glabrous beneath, unless for 

axillary tufts of hairs. 

Stamens of third series eglandular (see E. debilis) . 

E. mishuyacensis. 
Stamens of third series biglandular. 

Leaves with axillary tufts of hairs beneath. 

E. dysodantha. 
Leaves glabrous. 
Staminate flowers 3 mm. wide; cells of the 

outer anthers large E. pyriformis. 

Staminate flowers 5-6 mm. wide; cells of the 

outer anthers small E. rubriflora. 

A. Stamen connective not produced (seeE. rubriflora). 
Leaves oblong-linear, conspicuously bullate. 

E. bullata. 

Leaves broader than linear-oblong. 
Leaves large to very large, mostly longer than 
20cm. 

Petioles 4-5 cm. long E. Tessmannii. 

Petioles to 1 cm. long E. Cocuirey. 

Leaves medium in size, mostly shorter than 20 cm. 
Leaves more or less pubescent, not typically 

lanceolate. 
Leaf pubescence beneath dense in age; sta- 

minodia none E. Williamsii. 

Leaf pubescence sparse; staminodia obvious. 

E. verticillata. 
Leaves glabrous at maturity, lanceolate. 

E. arunciflora. 
Vegetative key to Aniba sens, strict. 

Perianth lobes definitely unequal A. Canelilla. 

Perianth lobes equal or subequal. 

Leaves not minutely papillose beneath. 



862 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves alternate; stamens of the third series sterile. A. perutilis. 
Leaves usually subverticillate; stamens all fertile. 

Leaves often 25 cm. long or longer; stamens of the third series 

longer than the outer ones A. gigantifolia. 

Leaves usually smaller; stamens of the third series smaller 

than the outer ones. 
Branchlets slender, lustrous; petioles slender, glabrous. 

A. salicifolia. 
Branchlets thick, verruculose; petioles thick, pubescent. 

A. Puchury-minor. 
Leaves papillose beneath. 

Bractlets persistent; ovary glabrous A. cylindri flora. 

Bractlets deciduous; ovary pubescent. 

Leaves abruptly rounded at the base A. rosaeodora. 

Leaves acute at the base. 

Leaves rigid-coriaceous A. Goto. 

Leaves chartaceous or at least flexible. A. Muca, A.firmula. 

Technical key to Aniba (after Kostermans) 

Stamens of the third row large but sterile A. perutilis. 

Stamens of the third row fertile. 

Stamens of series 3 larger than the outer, the subquadrate anthers 
with cells protruding; connective of the outer stamens well 

extended beyond the small cells A. gigantifolia. 

Stamens of series 3 smaller, or rarely as large as the outer, the 
anthers suborbicular, the cells not protruding; connective of 
the outer stamens not at all or obscurely extended. 
Leaves not papillose beneath, sometimes pilose. 

Sepals distinctly unequal A. Canelilla. 

Sepals equal or subequal. 

Branchlets and petioles slender, glabrous. . . .A. salicifolia. 
Branchlets and petioles thick, the former verrucose, the 

petioles pubescent A. Puchury-minor. 

Leaves minutely yellowish-papillose beneath. 

Ovary glabrous A. cylindriflora. 

Ovary pubescent (at least minutely). 

Leaves abruptly rounded at the base A. rosaeodora. 

Leaves acute at the base. 



FLORA OF PERU 863 

Flowers 1.5-3 mm. long; leaves chartaceous to coriaceous. 

A. Muca, A. firmula. 
Flowers 1.5 mm. long; leaves rigid-coriaceous. . .A. Goto. 

Aniba Canelilla (HBK.) Mez, Jahrb. Bot. Gart. Berlin 5: 53. 
1889. Cryptocarya Canelilla HBK. Nov. Gen. & Sp. 7: 192. pi. 645. 
1825. A. elliptica A. C. Smith, Phytologia 1: 115. 1935. 

Glabrous except the cinereous-tomentulose branchlets of the 
inflorescence and the flowers; petioles 8-20 mm. long; blades sub- 
coriaceous, the 6-8 (-10) nerves and abundant reticulation more or 
less prominent beneath, elliptic, cuneate at the base, acute or obtuse 
at the apex, 13-20 cm. long, usually less than half as wide; pedicels 
1.5-2.5 mm. long; flowers 2.5-3 mm. long, 2.5 mm. wide, the broadly 
ovate outer segments scarcely 1 mm. long, the broader inner ones half 
longer; outer filaments fleshy, pilose, twice as long as the obtuse 
anthers, the inner filaments thrice as long; connective not protrud- 
ing, the anthers of the third series with suborbicular pores; basal 
glands large, sessile; style as long as the glabrous ovary. The Rio 
Acre tree was 12 meters high, the trunk 6 cm. in diameter (Krukoff). 
The species is said to be well known on the Rio Orinoco for its cinna- 
mon-flavored and scented bark. This is sometimes used for making 
a stimulating tea (Ducke). The species has been confused with 
Ocotea Quixos (Lam.) Kosterm., which has fruit cups with a broad, 
double margin. 

Rio Acre: Rio Macauhan, Krukoff 5601, type of A. elliptica. 
Amazonian Brazil ; Venezuela ; Guiana. ' 'Louro precioso, ' ' ' 'canelilla. ' ' 

Aniba Goto (Rusby) Kosterm. in mss., comb. nov. Nectandra 
Goto Rusby, Bull. Torrey Club 49: 260. 1922. 

In general characters similar to A. Muca; leaves often persistently 
pilose on the costa beneath, rigid, 2.5-5 cm. wide; cup thick, roughly 
tuberculate, obscurely double, to 2.2 cm. broad, 2 cm. high, 12 mm. 
deep, with a thick, verrucose pedicel. According to collectors, a 
tree 15-20 meters tall, with cinnamon-brown bark, pungent in odor 
and taste. 

Peru: Probably. Bolivia; Colombia. "Goto." 

Aniba cylindriflora Kosterm., ined. 

Branchlets thick, terete, like the few-flowered, broadly pyramidal 
panicles densely but minutely tomentulose; petioles 1 cm. long; 
blades coriaceous, glabrous and lustrous above, dull and (under a 
lens) densely papillose and minutely pilose beneath, finally glabrate, 



864 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lanceolate or elliptic-lanceolate, contracted to the petiole at the 
subrotund base, the slender apical acumen to 1.5 cm. long, the 
lateral nerves 10-12, prominent and reticulate on both sides; pani- 
cles to 5 cm. long; flowers sessile, 3-3.5 mm. long, 4-5 mm. broad, 
the cylindric tube glabrous within; sepals subequal; outer anthers 
about twice exceeded by the slender, hirsute filaments, the filaments 
of the orbicular inner anthers glabrous on the upper portion; ovary 
glabrous, well exceeded by the style; cup rugulose, minutely rusty- 
verruculose, to 10 mm. high, 18 mm. broad. Near the Amazonian 
A. parviflora (Meissn.) Mez, 56, with very unequal sepals. 

Junin: Colonia Perene", Killip & Smith 25061 (det. Kostermans). 

Aniba firmula (Nees & Mart.) Mez, Jahrb. Bot. Gart. Berlin 5: 
58. 1889. Aydendron firmulum Nees & Mart. Linnaea 8: 36. 1833. 
Aydendron panurense Meissn. in DC. Prodr. 15, pt. 1: 89. 1864. 
Aniba panurensis Mez, loc. cit. 

Similar to A. Muca, but the leaves usually obovate-elliptic, acu- 
minate, 7-22 cm. long, 2-3 cm. wide, sometimes coriaceous, soon 
glabrous; flowers 1.5-2.5 mm. long. According to Ducke, the entire 
tree is fragrant. It is not clear to me that the species is separable 
from A. Muca, even varietally; Schmidt referred the Raimondi col- 
lection to the latter. Neg. 22064. 

Cajamarca: Cutervo, Jelski (det. Kostermans). Tambillo, Rai- 
mondi (det. Kostermans). Loreto: Mishuyacu, Klug 409 (det. 
Schmidt). Brazil; Venezuela; Colombia. "Ishpingo chico." 

Aniba gigantifolia 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10: 225. 1928. A. Williamsii 0. C. Schmidt, Repert. Sp. Nov. 31: 
169. 1933. 

Branches and ample inflorescence, including the flowers, tomentu- 
lose; blades mostly subverticillate, obovate-elliptic, about 20 (10-35) 
cm. wide, narrowed to a petiole 1-2 cm. long, glabrous above, very 
minutely tomentulose beneath or sparsely pilose only on the 10-30 
prominent nerves; inflorescence densely paniculate, 15-20 cm. long; 
flowers orange-yellow, 2-2.5 mm. long, the obconic tube attenuate to 
a pedicel scarcely 1 mm. long; ovary pilose, attenuate to the some- 
what longer, pilose style; stamens all fertile, those of series 3 with 
large, subquadrate anthers, the pores crescent-shaped, the connective 
of the outer stamens well exserted; glands subbasal, rather small; 
cup rather thin, with small, rusty warts. Said to be a tree of 10 
meters. It often is recognizable by its enormous leaves but, accord- 
ing to Kostermans, it is separable from A. Hostmanniana (Nees) 



FLORA OF PERU 865 

Mez of Surinam and the lower Amazon only by its pilose ovary; it 
should be treated, perhaps, as merely a geographic variety. Koster- 
mans reduces A. Williamsii with some hesitation, remarking that 
its leaves become glabrous, as in Venezuelan specimens, the branches 
being more slender than usual, and the flowers slightly smaller. 
The similar Amazonian A. affinis (Meissn.) Mez, 63, has smaller, 
alternate leaves, as also the related A. Burchellii Kosterm., but this 
has leaves minutely papillose beneath. Neg. 3800. 

Loreto: Iquitos, Tessmann 3628, type; Killip & Smith 27217 (det. 
Schmidt). Alto Rio Itaya, Williams 3495, type of A. Williamsii. 
"Muena amarillo." 

Aniba Muca (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 57. 1889. 
Laurus Muca R. & P. Fl. Peruv. 4: pi. 360. 1802; Laurogr. pi. 18. 
Aydendron Muca Nees, Linnaea 21: 498. 1848. 

A medium-sized tree with aromatic bark, minutely tomentulose 
on the angular young branchlets, inflorescence, and sometimes on 
the leaves beneath; leaves oblong or elliptic-lanceolate, chartaceous, 
minutely papillose beneath, more or less acuminate, acute at the 
base, usually about 10-12 (-22) cm. long, the nerves 7-10, prominent 
and reticulate beneath; inflorescence shorter than the leaves; flowers 
1.5 mm. long; pedicels 1 mm. long; perianth tube abruptly enlarging, 
pilose within ; sepals subequal ; outer filaments densely pilose, as long 
as or longer than the anthers, the smaller inner anthers with filaments 
glabrous above, the large glands subbasal; ovary small, pilose, sub- 
equaled by the style; cup rather thin, to 1.5 cm. high and broad, 
rugulose, with minute, rusty warts. A. parviflora (Meissn.) Mez, 
56, Amazonian, may be distinguished by its very unequal sepals, in 
this respect resembling A. Canelilla. Neg. 27592. 

Junin: Vitoc, Raimondi (det. Kostermans). La Merced, 1,000 
meters, Weberbauer 1919 (det. Mez). Huanuco: Posuso, Cochero, 
Ruiz & Pavon, type. Cuzco: Cosnipata, 700 meters, Weberbauer 
6961 (det. Schmidt). Bolivia. "Muca-muca." 

Aniba perutilis Hemsl. Kew Bull. 7, 197. 1894. A. compacta 
A. C. Smith, Bull. Torrey Club 58: 97. 1931. 

Twigs and dense foliage deciduously puberulent, the leaves gla- 
brous, except toward the base on both sides, pulverulent beneath 
when young; petioles 1-2 cm. long; blades often more or less obovate, 
attenuate-acuminate at the base, shortly obtuse-acuminate at the 
apex, 10-15 cm. long, 3.5-6.5 cm. wide, coriaceous, the costa promi- 



866 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nent, at least beneath, like the 7-12 lateral nerves, the closely reticu- 
late veins obscure; panicle densely yellowish-tomentulose, to 13 cm. 
long, many-flowered; pedicels 2-3 (-6) mm. long; perianth tube 
broad, 2 mm. long, the lobes fleshy, rounded; outer stamens fertile, 
those of the third series large but sterile; filaments pilose, stout, 
broader than long but equaling the deltoid, obtuse anthers; stami- 
nodia minute, stipitiform, pilose, sometimes wanting; ovary glabrous, 
1 mm. long, the style slightly shorter; fruit large, to nearly 3 cm. 
long and 2 cm. thick, the irregularly thickened cup 8-15 mm. high, 
with a few minute, rusty spots. In Colombia furnishing a valuable 
timber for cabinetwork and construction; see Kew Bull. loc. cit. 
Illustrated, Hook. Icon. pi. 2485. 

Loreto: Mishuyacu, 100 meters (Klug 608, type of A. compacta; 
755). Colombia; Bolivia. "Comino," "muena negro." 

Aniba Puchury-minor (Mart.) Mez, Jahrb. Bot. Gart. Berlin 
5: 70. 1889. Ocotea Puchury-minor Mart, in Buchner, Repert. Pharm. 
35: 172. 1830. Aydendron amazonicum Meissn. in DC. Prodr. 15, pt. 
1 : 89. 1864. Aniba amazonica Mez, op. cit. 69. A. Tessmannii 0. C. 
Schmidt, Notizbl. Bot. Gart. Berlin 10: 226. 1928. 

Branchlets angled, densely but minutely ferruginous-tomentu- 
lose; petioles to nearly 1.5 cm. long; blades mostly elliptic, 10-25 
cm. long, 3-6 (-10) cm. wide, narrowed to each end, the apical acu- 
mination sometimes 2 cm. long, glabrous or minutely tomentulose 
beneath, papyraceous; panicles usually 6-10 cm. long, slender, lax, 
tomentulose; flowers yellow, 2.5-3 mm. long, 2.5 mm. broad, the 
tube 1.5 mm. long; pedicels thick, 1 mm. long; stamens all fertile, 
the filaments at least 3 times longer than the anthers; ovary glabrous; 
fruit subglobose, 7-8 mm. long. A tree 7-20 meters high, with a 
trunk diameter of 40 cm. (Tessmann); bark dark purplish brown 
(Williams}. Var. intermedia (Meissn.) Kosterm., comb. nov. (Ay- 
dendron intermedium Meissn. in DC. Prodr. 15, pt. 1: 89. 1864), has 
panicles to 17 cm. long and slightly larger flowers. The seeds are 
sold as "pichurin" beans and are used in treatment of dysentery 
(Kostermans). Neg. 3799. 

Loreto: Soledad, Tessmann 5222, type of A. Tessmannii. Yuri- 
maguas, Poeppig 2478, type of A. amazonica; Williams 5897, 4287; 
Killip & Smith 27658. Puno: Isilluma, Raimondi (det. Schmidt; 
the var., fide Kostermans). Rio Acre: Ule 9407. Bolivia to Brazil 
and Colombia. "Muena amarillo," "miena amarillo." 



FLORA OF PERU 867 

Aniba rosaeodora Ducke, Rev. Bot. Appl. 8:845. 1928. 

Branchlets angulate-sulcate, densely puberulent-tomentulose; 
petioles 10-17 mm. long; blades elliptic to ovate, at maturity rounded 
at the base, the margin incurved, broadly and obtusely acuminate, 
the acumen to 1 cm. long, 15-20 (-30) cm. long, 4-6 (-13) cm. wide, 
rigid-coriaceous, glabrous above, dull, yellow, densely and micro- 
scopically papillose and pulverulent-tomentulose beneath, the pubes- 
cence disappearing, the nerves 9-11, prominent, like the areolate 
veins, especially beneath; panicles rusty-tomentulose, to 14 cm. 
long; pedicels thick, 0.5-1 mm. long; flowers 1.5-2 mm. long, the 
sulcate tube 1 mm. long; sepals subequal; outer stamens with broad, 
lanuginous filaments, the connective minutely protruding, the inner 
filaments with small anthers, glabrous above the large glands; ovary 
small, minutely pilose, the style slightly longer; cup rather shallow, 
with few rusty spots, to 2 cm. high, 3 cm. broad, the obscure outer 
margin formed by the thick tubercles, the pedicels to 1.5 cm. long. 
The Peru collection was referred by A. C. Smith, Bull. Torrey Club 
58: 100. 1931 (and also by Schmidt) to A. citrifclia (Nees) Mez, 74, 
a Brazilian species with the connective produced, the stamens of the 
third row with large anthers. Said to attain a height of 30 meters. 
Rose oil, used especially in the making of perfumes, is distilled from 
the bark, which formerly was shipped to France for this purpose, but 
is said now to be more economically treated by distillation at Cayenne 
(Kostermans). Illustrated, Arch. Jard. Rio Jan. 5: 109. pi. 5. Neg. 
3808. 

Loreto: Mishuyacu, 100 meters, Klug 1313. Brazil; Guiana; 
Colombia. 

Aniba salicifolia (Nees) Mez, Jahrb. Bot. Gart. Berlin 5: 71. 
1889. Aydendron salicifolium Nees, Syst. Laur. 252. 1836. 

A tree with lustrous, slender, terete branchlets, soon glabrous; 
petioles 5-10 mm. long; blades obovate-elliptic, narrowed to the 
slightly decurrent base, broadly and obtusely acuminate, charta- 
ceous, glabrous, dull, mostly 10-15 cm. long, 3-4.5 cm. wide, the lat- 
eral nerves 6-8, rather prominent beneath; panicles few-flowered, 
ferruginous-tomentulose, 4-6 cm. long; pedicels 1 mm. long; flowers 
2-3 mm. long, 2 mm. wide, the narrowly obconic tube abruptly 
enlarging, glabrous within; outer stamens elliptic, with broad, villous 
filaments and depressed-ovate anthers, the connective scarcely pro- 
truding, the outer filaments glabrous on the narrowed upper portion; 
glands slightly suprabasal; ovary glabrous, 1 mm. long, the style 



868 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to 1.5 mm. long; cup about 1 cm. high and broad, with rather large, 
rusty marks like spots, obscurely stipitate.- A tree 13 meters high 
(Krukoff). Neg. 22065. 

Loreto: Yurimaguas, 200 meters, Williams 4638 (det. Koster- 
mans). Brazil; Guiana. "Louro," "siuchi caspi." 

5. LICARIA Aubl. 

Acrodiclidium Nees. 

Reference: Kostermans, Recueil Trav. Bot. Ne"erl. 33: 719-754. 
1936; 34: 575-604. 1937. 

Like Aniba, but usually only the third series of stamens fertile, 
rarely also the second, the latter mostly and the first always modi- 
fied into leafy staminodia, or these rarely lacking; fourth series of 
stamens usually aborted. Perianth tube sometimes obsolete. Style 
shorter than or subequaling the ovary. Cup usually double-mar- 
gined. Kostermans, op. cit. 754, segregates Phyllostemonodaphne 
Kosterm. to provide for a Brazilian species with only 1 row of 
stamens reduced to staminodia. Besides the following, Ducke, 
Archiv. Jard. Bot. Rio Janeiro 4: 187. 1925, has recorded L. Canella 
Kosterm. as one of the more common species on the upper Ama- 
zonian tributaries, where it is known as "casca preciosa." Its bark, 
with very agreeable aroma similar to that of cinnamon but milder, 
is used, powdered, to scent linen and sometimes as a stimulating tea. 

Leaves medium-sized, 10-20 cm. long; stamens of series 4 aborted. 
Leaves ovate-lanceolate, about 5 cm. wide, often narrower. 

Leaves narrowly long-acuminate, subobtuse at the base; flowers 
globose or urceolate, short. 

Leaves 3-6 cm. wide; flowers glabrescent L. armeniaca. 

Leaves 2.5-3 cm. wide; flowers densely tomentose. 

Outer staminodia present L. multiflora. 

Outer staminodia wanting L. Quirirafuina. 

Leaves merely short-acuminate, the base acute; flowers infundi- 

buliform, 2-3 mm. long L. Pucheri. 

Leaves broadly elliptic, about 8 cm. wide L. latifolia. 

Leaves large, about 30 cm. long; staminodia of series 4 present. 
Leaves cordate at the base; perianth tube glabrous within. 

L. macrophylla. 

Leaves rounded-acute at the base; perianth tube sericeous within. 

L. endlicheriaefolia. 



FLORA OF PERU 869 

Licaria armeniaca (Poepp.) Kosterm. Recueil Trav. Bot. Ne"erl. 
34: 584. 1937. Acrodiclidium armeniacum Mez, Jahrb. Bot. Gart. 
Berlin 5: 85. 1889. Evonymodaphne armeniaca Nees, Syst. Laur. 264. 
1836. Oreodaphne Evonymodaphne Meissn. in DC. Prodr. 15, pt. 1: 
139. 1864. 

A shrub or tree, the branchlets 4-angled, soon glabrous, the 
leaves thin, soft, glabrous except the minutely strigose costa beneath; 
petioles to 1.5 cm. long; blades ovate-lanceolate, narrowly long- 
acuminate, the base subobtuse, about 10 (-18) cm. long and (3-) 4- 
6 cm. wide, scarcely lustrous, somewhat reticulate- veined beneath; 
inflorescence about as long as the leaves, few-flowered, extremely 
lax, soon glabrous; flowers 1.5-2 mm. long, on pedicels 5 mm. long, 
to 20 mm. long in fruit; perianth tube obsolete, sericeous within; 
outer staminodia elongate-stipitate, the inner wanting; basal glands 
large, often sepaloid. Neg. 3835. 

San Martin: Tocache, Poeppig 1787, type. Loreto: Rio Itaya, 
Williams 64, 104- (det. Schmidt); Killip & Smith 29288 (det. Koster- 
mans). Mishuyacu, 100 meters, King 1267, 1379 (det. Schmidt). 
La Victoria, Williams 2728 (det. Kostermans). Brazil. 

Licaria endlicheriaefolia Kosterm. Recueil Trav. Bot. Ne"erl. 
34: 583. 1937. Acrodiclidium endlicheriaefolium Kosterm. Med. Bot. 
Rijks Mus. Utrecht 37: 724. 1936. 

A glabrous tree; petioles 7 cm. long; blades broadly elliptic, 
rounded or subcordate at the base, the veins prominent on both 
sides; perianth tube glabrous within; outer staminodia spatulate, 
fimbriate marginally at the apex; fertile stamens glabrous, with 
rather large cells; staminodia of series 4 short, glabrous. Otherwise 
similar to L. macrophylla. A tree of 7-8 meters. 

Junin: San Ramon, 1,000 meters (Killip & Smith 24901, type). 

Licaria latifolia (A. C. Smith) Kosterm. Recueil Trav. Bot. 
Ne"erl. 34: 601. 1937. Acrodiclidium latifolium A. C. Smith, Bull. 
Torrey Club 58: 100. 1931. 

A small tree, except for the leaves above more or less densely 
ferruginous-pilose; petioles 12-15 mm. long, slender; blades oblong 
or broadly ovate-oblong, the base broadly cuneate, the apex abruptly 
caudate-acuminate, glabrous or sparsely pilose above, 15-19 cm. long, 
7-9 cm. broad, the veinlets subimmersed; inflorescence lax, 7-9 cm. 
long, solitary in the axils, many-flowered; flowers perfect, the tube 
to 1.5 mm. long, glabrous within; outer stamens sterile; filaments 
of the fertile third series 2-3 times longer than the anthers; basal 



870 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glands minute; ovary and style subequal, glabrous. I believe this 
species to be only a variety of L. brasiliensis Nees, with smaller, less 
acuminate leaves, denser panicles, and larger glands ( Kostermans) . 
Loreto: Mishuyacu, 100 meters (Klug 30, type). 

Licaria limbosa (R. & P.) Kosterm. Recueil Trav. Bot. Ne"erl. 
34: 585. 1937. Acrodiclidium limbosum Mez, Jahrb. Bot. Gart. Berlin 
5: 89. 1889. Laurus limbosa R. & P. Fl. Peruv. 4: pi. 361. 1802. 
Nectandra limbata Nees, -Linnaea 21: 509. 1848. A. limbatum Macbr. 
Field Mus. Bot. 11:18. 1931. 

Flowering branchlets densely and minutely ferruginous-tomen- 
tulose but soon glabrous; leaves alternate, glabrous, somewhat 
lustrous, obviously and densely areolate-reticulate on both sides, evan- 
escently so above, ovate-elliptic to elliptic, 11-19 cm. long, 3-7 cm. 
wide, cuneately acute at the base, acuminate at the apex, the costa 
and lateral nerves filiform and prominent above, the costa flattened 
beneath; petioles slender, glabrous, 1-2 cm. long; panicles axillary 
and more or less clustered on the young branchlets, densely ferrugi- 
nous- tomentulose, many-flowered, 4-7 cm. long; bracts and bractlets 
deciduous; pedicels 1 mm. long, glabrous like the flowers outside, 
these 2-2.5 mm. long, the obconic tube 1 mm. long; sepals erect, 
fleshy, acute, 1.25 mm. long, villous at the base within; filaments 
densely but laxly villous, as long as the ovate anthers; basal glands 
stalked; ovary glabrous, 1 mm. long, the style 1.5 mm. long; fruit 
to 2.5 cm. long, the cup verruculose, red, with small, rusty dots, to 
1.5 cm. high, 2.5 cm. broad, 1 cm. deep, the outer margin spreading, 
to 4 mm. wide, the thinner inner one erect, to 2.5 mm. high. Koster- 
mans, from whom this description is copied, originally, op. cit. 736, 
included this species with L. Pucheri, which see. Perhaps the Tacna 
reference is a mistake for Tarma. 

San Martin: San Roque, 1,400 meters, Williams 6978. Tacna: 
Tafalla. Junin: Tarma, Ruiz. Bolivia to Costa Rica and Venezuela. 

Licaria macrophylla (A. C. Smith) Kosterm. Recueil Trav. 
Bot. Ne"erl. 34: 583. 1937. Acrodiclidium macrophyllum A. C. Smith, 
Bull. Torrey Club 58: 101. 1931. 

Soon glabrous except for the inflorescence branches; petioles stout, 
2-3 cm. (5-7 according to Kostermans) long; blades oblong-ovate, 
cuneate at the base, long-acuminate at the apex, 25-40 cm. long, 
10-16 cm. wide, coriaceous, the veinlets slightly elevated beneath, 
the finer ones obscure; inflorescence racemose-paniculate, 20-40- 
flowered; flowers perfect, glabrous, 3 mm. long, the tube sericeous 



FLORA OF PERU 871 

within, to 1.5 mm. long, fleshy; outer stamens sterile, large, auricled, 
those of the third, fertile series with pilose, glandular filaments; 
anthers auricled, the cells minute; staminodia of series 4 large, densely 
sericeous; ovary semi-immersed, the style subequal. 

Loreto: Mishuyacu, 100 meters (King 80, type); also 246 (det. 
Schmidt). 

Licaria multiflora Kosterm. Recueil Trav. Bot. Ne*erl. 34: 584. 
1937. Acrodiclidium multiflorum Kosterm. op. cit. 33: 735. 1936. 

Branchlets slender, densely ferruginous- tomentose; leaves char- 
taceous, elliptic, 7-10 cm. long, 2.5-3 cm. wide, narrowed at the base, 
distinctly and broadly acuminate at the apex, the obovate acumen 
to 1.5 cm. long, lustrous and glabrous above, the veins more or less 
evidently and densely reticulate, scarcely obvious, or at least not 
prominent, beneath, dull, sparsely tomentose, the nerves 4-5, the 
veins scarcely conspicuous, areolate-reticulate; petioles 10-15 mm. 
long; panicles finely tomentulose, the branchlets to 2 cm. long; 
pedicels 2-3 mm. long; flowers urceolate, densely ferruginous-tomen- 
tulose, 1-1.5 mm. long, 0.75 mm. broad, the tube urceolate, 1 mm. 
long and broad; perianth segments glabrous within, to 1 mm. long; 
outer staminodia glabrous, 1.5 mm. long; fertile stamens subexserted, 
glabrous, with distinct, slender filaments; ovary glabrous, equaled 
by the style. A tree 25 meters high (Krukoff). Simulating this 
species is L. Appelii (Mez) Kosterm., found as near as Rio Livra- 
mento; its leaves are 6-7 by 2-2.5 cm., and the flowers 3 mm. long. 

Rio Acre: Rio Purus, near mouth of Rio Macauhan, Krukoff 
5324, type. Amazonian Brazil. 

Licaria Pucheri (R. & P.) Kosterm. Recueil Trav. Bot. Ne"erl. 
34: 585. 1937. Laurus Pucheri R. & P. Fl. Peruv. 4: pi 352. 1802. 
Aniba foeniculacea Mez, Repert. Nov. Sp. 3: 65. 1906. Acrodiclidium 
Pucheri Kosterm. op. cit. 33: 736. 1936. 

Similar toL. limbosa; blades 12-16 cm. long, 3-6 cm. wide; peti- 
oles to 1 cm. long; panicles axillary, subterminal, lax, sparsely and 
minutely sericeous; pedicels to 9 mm. long; filaments puberulous at 
the base; ovary and style 1 mm. long; cup 12 mm. high, 2 cm. wide, 
the inner margin 1 mm. high. 

Junin: Vitoc, Ruiz & Pawn, type. La Merced, Schunke 262; 
Weberbauer 1890, type of Aniba foeniculacea. "Puchery," "cucherin." 

Licaria Quirirafuina Kosterm. Recueil Trav. Bot. Ne*erl. 34: 
584. 1937. 



872 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Similar to L. multiflora; leaves 9-12 cm. long, 3-4 cm. wide, the 
acumen slender; pedicels 1.5 mm. long; perianth tube less than 1 mm. 
long, 0.5 mm. broad; outer staminodia wanting; glands large; style 
longer than the ovary, exserted. A tree 8 meters high (Klug). 

Loreto: Florida, Rio Putumayo (Klug 2131, type). "Efuina qui- 
rirafuina" (Huitotoname). 

6. PERSEA Mill. 

Flowers perfect, the tube none, the segments persisting, the 
outer ones shorter than the inner or rarely subequal. Third series 
of stamens rarely staminodial. Filaments usually longer than the 
anthers, the anthers of the third series sessile or stipitate-glandular. 
Staminodia most often barbate. Ovary pilose or glabrous, sub- 
globose, the style and perianth lobes scarcely at all accrescent. 
Genus noteworthy as the source of the popular fruit "palta," Persea 
americana Mill. Ruiz and Pavon wrote in their journal with evident 
appreciation of the fruit of this beautiful tree. Weberbauer found 
the tree cultivated to 2,500 meters in the inter-Andean valleys of 
central Peru. 

Key based on leaf characters 

Leaves large, mostly 15 cm. to several decimeters long. 
Pubescence of the leaves beneath dense, conspicuous. 

Leaves subsessile P. grandis. 

Leaves evidently petiolate, the petioles 1-5 cm. long. 
Leaves suBobtuse or subcordate at the base ... P. subcordata. 
Leaves acute at the base. 
Petioles 3-5 cm. long. 

Inflorescence about 5 cm. long P. peruviana. 

Inflorescence ample P. boliviensis. 

Petioles about 1.5 cm. long P. crassifolia. 

Pubescence of the leaves minute, obscure, or none. 

Ovary densely pilose; leaves rather thin; flowers 6-7 mm. long, 

the segments subequal P. americana. 

Ovary glabrous; outer perianth segments distinctly shorter 

than the inner. 

Leaves usually obtuse or rounded at the base, rather thin; 
flowers 4-6 mm. long; pedicels 2-5 mm. long or longer; 
style little longer than the ovary P. laevigata. 






FLORA OF PERU 873 

Leaves acute at the base; flowers 7-8 mm. long; pedicels 
2 mm. long; style slender, nearly 3 times longer than 

the ovary P. Raimondii. 

Leaves medium-sized, mostly 6-9 cm. long (rarely 13 cm.). 
Leaves definitely pubescent beneath. 
Pubescence sericeous. 

Inflorescence many-flowered; leaves rounded at the base. 

P. Haenkeana. 

Inflorescence often few-flowered; leaves acute at the base. 
Leaves usually 8-10 cm. long; ovary glabrous. 
Leaves about 3 cm. wide; flowers subsessile. .P. stricta. 
Leaves about 6 cm. wide; flowers pediceled. 

P. crassifolia. 
Leaves about 5-6 cm. long; ovary pubescent. 

P. Weberbaueri. 

Pubescence villous-tomentose, sometimes finely so (see P. stricta). 
Pubescence a short, dense tomentum. 
Pubescence short, close; style 2-3 times longer than the 

ovary P. corymbosa. 

Pubescence longer and softer; style and ovary subequal. 

P. ferruginea. 

Pubescence rather long and loose, villous or pilose. 
Leaves subcordate at the base. 

Inflorescence not long-peduncled P. Ruizii. 

Inflorescence very long-peduncled P. subcordata. 

Leaves acute at the base P. hirta. 

Leaves glabrous or minutely and obscurely pubescent. 

Leaves 3 to rarely 5 cm. long P. boldufolia. 

Leaves mostly 8-9 cm. long or longer. 
Leaves obtuse or subcordate at the base. 

Leaves rigid; petioles thick, about 1 cm. long. . P. Mutisii. 
Leaves flexible; petioles slender, 1.5-3 cm. long. 

P. pyrifolia. 

Leaves acute at the base, or flexible and the petioles only 
1 cm. long or shorter. 

Branchlets and inflorescence glabrate P. scoparia. 

Branchlets and inflorescence sericeous P. durifolia. 



874 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Technical key 

Third series of anthers sterile, the 2 outer series 4-celled. 
Flowers long-pediceled, the pedicels to 4 mm. long . . P. crassifolia. 
Flowers sessile or short-pedicellate. 

Branchlets soon glabrate P. Raimondii. 

Branchlets densely ferruginous-pubescent. 

Leaves somewhat acuminate P. peruviana. 

Leaves more or less rounded at both ends P. boliviensis. 

Third series of anthers 2-celled, usually fertile. 

All the anthers 2-celled P. durifolia. 

Outer anthers 4-celled. 
Anthers of the third series 2-celled. 

Leaves sericeous beneath with a copper-colored pubescence. 

P. Haenkeana. 
Leaves not so pubescent. 

Filaments longer than the anthers; style longer than the 

ovary P. stricta. 

Filaments and anthers, like the style and ovary, subequal. 

P. subcordata. 

Anthers of the third series 4-celled. 
Ovary glabrous. 

Perianth segments subequal P. Ruizii. 

Perianth segments unequal, the outer ones distinctly 

shorter. 
Floral glands at least short-stipitate. 

Inflorescence long-peduncled P. hirta. 

Inflorescence short-peduncled P. corymbosa. 

Floral glands sessile. 

Leaves subsessile, long P. grandis. 

Leaves petiolate. 

Mature leaves pubescent beneath. 

Pubescence coppery-sericeous P. Haenkeana. 

Pubescence whitish or reddish, tomentose. 

P. ferruginea. 
Mature leaves glabrate or obscurely pubescent. 

Branchlet tips tomentose P. pyrifolia. 

Branchlet tips glabrate or puberulent. 



FLORA OF PERU 875 

Leaves 4 cm. long P. boldufolia. 

Leaves larger. 

Inflorescence crowded, subsessile . . P. Mutisii. 
Inflorescence open, peduncled. 

P. scoparia, P. laevigata. 
Ovary pilose. 

Leaves sericeous beneath, small P. Weberbaueri. 

Leaves glabrate, 10-20 cm. long P. americana. 

Persea americana Mill. Gard. Diet. ed. 8. 1768. P. gratissima 
Gaertn. Fruct. 3: 222. 1807. 

The well known "palta" ("avocado" of the United States) is some- 
times cultivated. Its rather thin, glabrate, broadly elliptic leaves, 
the lateral nerves rugose-reticulate beneath, mostly 10-20 cm. long 
and half as wide, distinguish it from all native species except P. 
laevigata, which, however, has much less prominent nervation and 
a glabrous ovary. The palta is a large, beautiful tree. The fruits 
are known also as "aguacates." 

Junin: La Merced, 700 meters, Kittip & Smith 23422. Dense 
forest, Schunke Hacienda, Killip & Smith 24630. Loreto: Cachi- 
puerto, King 3118. Lower Rio Nanay, Williams 325, 324. Mishu- 
yacu, near Iquitos, 100 meters, in clearing, Klug 1426, 756. Iquitos, 
in forest, 100 meters, Killip & Smith 29841- Lima: Lima, culti- 
vated, Ruiz & Pavon. Probably native of Mexico and Central 
America. "Palto," "palta," "huira palta," "paltai." 

Persea boldufolia Mez, Repert. Nov. Sp. 3: 66. 1906. 

A shrub of 2 meters, readily recognized by its numerous small 
(mostly less than 4 cm. long) leaves, obtuse at the base and rounded 
at the apex; inflorescence densely ferruginous-pilose, few-flowered, 
to 6 cm. long; flowers subsessile, 4 mm. long; filaments pilose, little 
longer than the 4-celled (all fertile) anthers; glands sessile; ovary 
glabrous. Compared by Mez with P. brevipes Meissn., of Ecuador, 
with acute leaves about twice as long and pedicellate flowers 5 mm. 
long. Neg. 3560. 

Amazonas: Molinopampa, east of Chachapoyas, 2,400 meters, 
Weberbauer 4379, type; 263. 

Persea boliviensis Mez & Rusby, Mem. Torrey Club 6: 113. 
1896. 

A shrub or tree, densely ferruginous-tomentulose on the branch- 
lets and more finely or less obviously so in the thyrsoid inflorescences 



876 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and on the leaves beneath; blades elliptic, subrotund at both ends, 
as much as 20 cm. long and 8 cm. wide, coriaceous, reticulate- 
veined, especially beneath; petioles to 4 cm. long, nearly as long as 
the subsessile, few-flowered inflorescences; pedicels 1-2 mm. long; 
flowers 7 mm. long, the outer segments much shorter than the 
inner; outer anthers 4-celled, those of the third series 2-celled but 
sterile; staminodia pilose. Compared by the author with P. peru- 
viana. P. Benthamiana Meissn. has flowers 4-5 mm. long with 
3 series of 4-celled, fertile anthers. 

Junin: Hacienda Schunke, 1,400-1,700 meters, Killip & Smith 
24831; Schunke 205, 206 (det. Schmidt). Bolivia. 

Persea corymbosa Mez, Jahrb. Bot. Gart. Berlin 5: 158. 1889. 

A rather densely leafy shrub or small tree, to 5 meters high, 
conspicuously ferruginous-tomentose on the branchlets, inflorescence, 
and lower leaf surface; petioles fleshy, to 9 mm. long; blades rigid, 
elliptic or ovate, rounded or subacute at each end, 3-5 cm. wide 
and twice as long; inflorescence subsessile, corymbose, shorter than 
the leaves; flowers 5 mm. long; filaments of the third series glandular 
and a little exceeding the 4-celled anthers, the others shorter. 

Cajamarca: Tambillo and Cutervo (Jelski 173; 177, type); 
Raimondi (det. Schmidt). Huambos, 3,100 meters, Weberbauer 4159 
(det. Mez); 260. 

Persea crassifolia Mez, Repert. Nov. Sp. 3: 67. 1906. 

A shrub or tree of 6 meters, with very rigid, elliptic leaves, 
acute at each end; young parts and leaves beneath sericeous-pilose; 
blades about 10 cm. long and 6 cm. wide; inflorescence ferruginous- 
tomentose, much exceeding the leaves, the stout, angled pedicels 
4-5 mm. long; flowers green, 7 mm. long; filaments pilose, only the 
2 outer series with fertile, 4-celled anthers; ovary glabrous, half 
as long as the slender style. Related to P. perumana Nees. Neg. 
3744. 

Huanuco: Monzon, 2,000-2,500 meters, Weberbauer 3525, type; 
255. 

Persea durifolia Mez, Repert. Nov. Sp. 3: 65. 1906. 

A tree of 8 meters, glabrous or glabrate except the sericeous 
and brownish-tomentulose branchlets and inflorescence; petioles very 
stout, to 1 cm. long; blades rigid, distinctly glaucous beneath, 
elliptic, acute at each end or the base subrounded, usually 8-10 
cm. long and half as wide, sometimes 13 cm. long; inflorescence 



FLORA OF PERU 877 

subequaling the leaves, corymbose, long-peduncled ; flowers sub- 
sessile, 3 mm. long; filaments long-pilose, twice longer than the 
2-celled (all fertile) anthers; ovary glabrous, exceeded by the style. 
Aberrant, with P. cuneata Meissn. of Colombia, in having all 
the anthers 2-celled. Neg. 3564. 

Cuzco: Santa Ana, 2,100 meters, Weberbauer 5010, type. 

Persea ferruginea HBK. Nov. Gen. & Sp. 2: 159. 1817; 158. 
P. Humboldtii Mez, Jahrb. Bot. Gart. Berlin 5: 158. 1889. 

Except for the much softer pubescence, similar to P. corymbosa; 
glabrous ovary and style subequal. Compare also P. vestita Mez, 
in a note under P. subcordata, and P. sericea HBK. Neg. 3565. 

Piura: Huancabamba, 2,500 meters, Weberbauer 6311. Huai- 
cani (Pearce). Ecuador; Colombia. 

Persea grandis Mez, Jahrb. Bot. Gart. Berlin 5: 160. 1889. 

The only Peruvian species with subsessile leaves, these about 
30 cm. long, long-cuneate-oblong from the base, about 8 cm. wide; 
filaments pilose, half as long as the 4-celled anthers; ovary glabrous, 
exceeding the style. Related to P. hirta Nees. A beautiful tree of 
20-25 meters. The type locality may be Bolivian. 

Peru(?): Sandillani (Pearce, type). 

Persea Haenkeana Mez, Jahrb. Bot. Gart. Berlin 5: 174. 1889. 

Blades elliptic-lanceolate, about 6 cm. long and 2 cm. wide, 
with the inflorescence beautifully pubescent beneath with sericeous, 
copper-colored hairs; petioles to 6 mm. long; inflorescence long- 
peduncled, about equaling the leaves; flowers many, white, on 
pedicels 1-1.5 mm. long, the segments unequal; filaments some- 
what pilose, the anthers 4-celled (all?); style and glabrous ovary 
slightly unequal. A tree of 3-6 meters (Pearce}. Neg. 19281. 

Peru: Haenke, type. Santa Cruz (Pearce}. 

Persea hirta Nees, Syst. Laur. 673. 1836; 159. P. rugosa 
Meissn. in DC. Prodr. 15, pt. 1: 47. 1864. 

Similar in pubescence to P. Ruizii; petioles about 5 mm. long; 
blades obovate or subelliptic, about 8.5 cm. long and 4 cm. wide; 
inflorescence as long, broadly pyramidal; flowers 5 mm. long, sub- 
sessile; filaments glabrous; glands long-stipitate; ovary and style 
subequal. 

Amazonas: Yambrasbamba (Mathews 1428, type). 



878 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Persea laevigata HBK. Nov. Gen. & Sp. 2: 157. 1817; 171. 
Laurus caerulea R. & P. Fl. Peruv. 4: pi. 350. 1802, nomen; Laurogr. 
pi. 2. P. caerulea Mez, Jahrb. Bot. Gart. Berlin 5: 171. 1889. 

A shrub or a tree to 18 meters high, with much the aspect of 
the cultivated P. americana, but with tomentulose inflorescence, 
slightly smaller flowers, and glabrous ovary; blades very broadly 
ovate or elliptic, often 15-25 cm. long and 7-10 cm. wide. P. 
Buchtienii Schmidt, of Bolivia, has glabrous pedicels and glabrate 
flowers. Negs. 3561, 3562. 

Huanuco: Below Mufia, 1,900 meters, Weberbauer 6791 (?; det. 
Schumann), 6821; Ruiz, type. Puyash, Sawada 96. Villcabamba, 
1,800 meters, 5008 (det. Schmidt). Cajamarca: Nancho and Chiri- 
nos, Raimondi. San Martin: Cerro Morro, Moyobamba, 1,200 
meters, Weberbauer 4588 (det. Mez); 289. Tarapoto, Spruce 4919 
(det. Schumann and Mez, but leaves acute at the base). Zepelacio, 
1,100 meters, King 3693. Without locality: Ruiz & Pavdn 1322, 
1324- North to Costa Rica and Venezuela. "Aguacate," "agua- 
cate cimarron," "piria," "palton." 

Persea Mutisii HBK. Nov. Gen. & Sp. 2: 158. 1817; 155. 

A very densely leafy shrub or small tree with stout, soon gla- 
brate, angled branchlets; leaves glaucous, glabrous or nearly so, 
rigid, broadly elliptic-obovate, mostly less than 10 cm. long, half 
as wide; panicles dense, subsessile, umbellate, terminal and axillary, 
ferruginous- tomentose, shorter than the leaves; pedicels often short, 
sometimes 7-8 mm. long; flowers 6-7 mm. long; anthers 4-celled, 
all fertile, the glabrous filaments of the outer series twice as long, the 
inner sparsely pilose, the glands sessile; style much longer than the 
glabrous ovary. Neg. 3589. 

Cajamarca: Cutervo (Jelski 197; det. Mez). Ecuador; Colombia; 
Venezuela. 

Persea peruviana Nees, Syst. Laur. 155. 1836; 140. 

Branchlets, inflorescence, and leaves beneath softly and densely 
ferruginous-pubescent with fine, crisped hairs; blades elliptic-oblong, 
subacuminate, subrigid, 15-25 cm. long and half as wide; petioles 
3-5 cm. long; inflorescence dense, thyrsoid, only 3 cm. long; flowers 
6-8 mm. long, sessile or nearly so; anthers 4-celled, the 2 inner 
series sterile, about equaling the densely yellow-lanate filaments; 
style much longer than the glabrous ovary. Probably a large tree. 
Neg. 3574. 

Huanuco: Cochero, Poeppig 1310, type. 



FLORA OF PERU 879 

Persea pyrifolia Nees, Syst. Laur. 145. 1836. Laurus cordata 
Veil. Fl. Flum. 4: pi. 50; text 163. 1825. P. cordata Mez, Jahrb. 
Bot. Gart. Berlin 5: 165. 1889, non Meissn. 

Branchlets densely leafy, more or less yellowish-tomentulose, 
like the short but long-peduncled inflorescences; leaves glabrous 
or minutely and finely pubescent beneath, ovate or elliptic-obovate, 
mostly acute or acuminate, usually about 10 cm. long and 4-6 cm. 
wide (narrower in the Peruvian form); flowers ferruginous-tomen- 
tose, 5 mm. long, on pedicels 1-3 mm. long; outer perianth segments 
much shorter than the inner; filaments pilose, half longer than the 
4-celled anthers; style 2.5 times as long as the glabrous ovary. 
A tree of 12 meters, said to be valuable for its fine wood. The 
Spruce specimen may be rather P. Raimondii. 

San Martin: Tarapoto, Spruce 4919 (det. Mez). Brazil. "Ca- 
nella rosa." 

Persea Raimondii 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10: 725. 1929. 

Branches glabrate; petioles 2-3.5 cm. long; blades mostly elliptic, 
acute at the base, narrowed and short-acuminate at the apex, 
12-19 cm. long, 5.5-9 cm. wide, short-strigillose or pilose beneath, 
or glabrate in age; inflorescence very sparsely short-pilose, some- 
times 9.5 cm. long; stamens 4.5 mm. long, the densely pilose fila- 
ments 3 mm. long, the anthers 4-celled, those of the third series 
2-celled and apparently sterile; style 3 times as long as the glabrous 
ovary. Probably a tree. Related to the reddish-tomentose P. 
boliviensis Mez & Rusby. See also P. cordata, at least as to speci- 
men cited. Neg. 3576. 

Cajamarca: Tambillo, Raimondi, type. "Paltojera." 

Persea Ruizii Macbr. Field Mus. Bot. 11: 18. 1931. Laurus 
ferruginea R. & P. Fl. Peruv. 4: pi. 372. 1802; Laurogr. pi. 5. P. 
ferruginea Mez, Jahrb. Bot. Gart. Berlin 5: 154. 1889, non HBK. 

A shrub or tree, the fleshy branchlets densely red-villous at 
first, like the inflorescence and rugose- veined leaves beneath; peti- 
oles thick, 1 cm. long; blades rigid, finely pilose on the nerves above, 
even in age, cordate-ovate, obtuse, 6-10 cm. long, 4-6 cm. wide; 
inflorescence much shorter, subracemose, few-flowered; flowers 7-10 
mm. long, the pedicels 1-4 mm. long; filaments long-pilose, slightly 
longer than the 4-celled anthers, the glands of series 3 sessile; style 
a little longer than the glabrous ovary. Neg. 3578. 

Huanuco: Pillao, Ruiz & Pawn, type. 



880 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Persea scoparia Mez, Spic. Laur. 115. 1892. 

A tree or shrub with slender, glabrous branchlets, pruinose at 
the tip; petioles mostly 5-8 mm. long; blades oblong-elliptic, sub- 
obtuse or somewhat acute at both ends, coriaceous, glabrous, the 
larger 9 cm. long and 3 cm. wide; inflorescence a little shorter, 
slenderly long-peduncled, minutely ferruginous-sericeous-tomentu- 
lose or glabrous; flowers few, on pedicels scarcely 1 mm. long, the 
outer segments much shorter than the ovate inner ones, these nearly 
2 mm. long, tomentulose. Compared by Mez with P. Mutisii. 
Similar, too, is P. brevipes Meissn. of Ecuador, with subsessile, 
pilose inflorescences, pedicels 2-5 mm. long, and flowers 5 mm. long. 

Peru: Probably. Bolivia. 

Persea stricta Mez, Jahrb. Bot. Gart. Berlin 5: 141. 1889. 

A tree with the anthers of P. subcordata, but the blades mostly 
about 8 cm. long and 3 cm. wide, rigid, sericeous-tomentulose 
beneath, acute or acutish at each end; petioles to 6 mm. long; 
inflorescence yellowish- tomentulose, the flowers 3.5-4 mm. long, sub- 
sessile; filaments twice as long as the anthers; style half longer 
than the glabrous ovary. P. sericea HBK., of Ecuador, with 
petioles to 15 mm. long and flowers 5-6 mm. long, which might be 
sought here, has essentially the stamen characters of P. ferruginea 
HBK. Pearce's locality may be Bolivian. 

Peru(?): Sandillani (Pearce, type). 

Persea subcordata (R. & P.) Nees, Linnaea 21: 492. 1848; 
142. Laurus subcordata R. & P. Fl. Peruv. 4: pi. 369. 1802; Laurogr. 
pi. 6. 

A tall tree with ample, cordate-ovate-subelliptic, subobtuse 
leaves; branchlets, inflorescence, and leaves beneath, especially 
on the nerves, ferruginous- villous or pilose; blades 15-20 cm. long 
and 11-12 cm. wide, the reticulate nervation conspicuous beneath; 
petioles to 4 cm. long; inflorescence long-peduncled, about equaling 
the leaves; flowers 5 mm. long; anthers in 3 fertile series, those 
of the third series 2-celled; filaments scarcely as long, pilose, the 
glands subsessile; glabrous ovary and style subequal. A tree 25-30 
meters high (Pearce). P. vestita Mez, 154, of Bolivia probably 
occurs in southeastern Peru; the anthers of the third series are 
4-celled; petioles to 2 cm. long; leaves narrower, scarcely cordate. 
Neg. 3578. 



FLORA OF PERU 881 

Huanuco: Chinchao, Ruiz & Pavon, type. Pan de Azucar, 
Sawada 76 (det. Schmidt). Amazonas: Chachapoyas (Mathews 
179}. Department uncertain: Sandillani (Pearce). 

Persea Weberbaueri Mez, Repert. Nov. Sp. 3: 66. 1906. 

A shrub, well characterized by its small, elliptic-rounded leaves, 
glabrous above but sericeous beneath; petioles to 8 mm. long; 
inflorescence subequaling the leaves, densely ferruginous-tomentu- 
lose, like the young branchlets; pedicels to 3 mm. long; flowers 
4^4.5 mm. long, the outer segments only half as long as the inner; 
filaments densely long-pilose, thrice longer than the 4-celled anthers 
(all fertile); glands short-stipitate; style twice as long as the densely 
pilose ovary. Neg. 3580. 

Puno: Below Tambo Cachicachi, 1,800 meters, Weberbauer 1161, 
type; 242. 

7. PHOEBE Nees 

Similar to Persea, but the perianth tube sometimes obvious, the 
lobes often equal, and the lower anther cells of the 4-celled anthers 
little broader than the broadest part of the upper 2. Perianth 
lobes (when persistent) and pedicel obviously accrescent in fruit. 
The difference in anther cells used by Mez to distinguish this genus 
from Persea is often slight or not clear. 

Leaves mostly of an ovate-elliptic type, rounded, obtuse, oblique, 

or short-acute at the base; anthers partly 2-celled. 
Leaves shortly obtuse-acuminate, pinnate-nerved; anthers of the 

third series 2-celled P. heteranthera. 

Leaves subacutely caudate-acuminate, somewhat 3-nerved; an- 
thers of the third series 4-celled P. maynensis. 

Leaves mostly more oblong- or lance-elliptic, acute at the base; 

anthers of the third series unknown (4-celled in P. Mathewsii). 

Inflorescence subracemose; leaves pinnate-nerved ... P. Mathewsii. 

Inflorescence more or less panicled; leaves usually 3-nerved from 

near the base. 
Style and ovary subequal. 

Leaves 3-nerved P. Poeppigii. 

Leaves pinnate-nerved P. pichisensis. 

Style elongate P. cinnamomifolia. 

Phoebe cinnamomifolia (HBK.) Nees, Linnaea 21: 488. 1848; 
212. Persea cinnamomifolia HBK. Nov. Gen. & Sp. 2: 160. 1817. 



882 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tree with oblong-lanceolate or elliptic-lanceolate leaves, acute 
at both ends, mostly 10-15 cm. long and 4 cm. wide, and narrow, 
panicled, more or less white-pilose inflorescences about equaling 
the leaves, of few to many flowers; blades glabrous or obscurely 
puberulent beneath, somewhat 3-nerved, the nerves and reticu- 
lation fine, prominent only beneath; flowers slightly to densely 
pilose, 2.5 mm. long; filaments and 4-celled anthers glabrous, sub- 
equal; glands foliaceous, staminodia large, cordate, slightly pilose. 
-Neg. 3590. 

Cajamarca: Cutervo (Jelski 166}. Colombia; Venezuela. 

Phoebe heteranthera (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 
5: 185. 1889. P. Ruiziana Nees, Linnaea 21: 489. 1848. Laurus 
heteranthera R. & P. Fl. Peruv. 4: pi. 364. 1802; Laurogr. pi. 4. 

A shrub or tree with yellowish-puberulent branchlets, soon gla- 
brate, and crowded, thyrsoid inflorescences much shorter than the 
leaves; blades coriaceous, glabrous but barbellate beneath in the 
nerve axils, mostly 7-8 cm. long and 3-4 cm. wide, the nerves im- 
mersed above, the reticulation beneath fine; flowers sparsely puberu- 
lent, about 3 mm. long; stamens pilose, the outer anthers 4-celled, 
those of the third series 2-celled; glands and staminodia large, the 
former punctate, the latter pilose; style and ovary subequal. Neg. 
3594. 

Peru: According to a Ruiz specimen in Herb. Berlin. Loja, 
Ecuador, according to Mez, loc. cit., but without data at Madrid. 

Phoebe Mathewsii (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
217. 1889. Persea punctata Meissn. var. Mathewsii Meissn. in DC. 
Prodr. 15, pt. 1: 505. 1864. 

A shrub or tree, minutely yellowish-tomentose on the young 
branchlet tips; blades narrowly elliptic, obovate or oblong, acute 
at the base, subobtuse and mucronulate at the apex, about 9 cm. 
long and 3.5 cm. wide, lustrous above, sparsely pilose on the nerves 
above and beneath, becoming glabrate, rather thick, conspicuously 
and finely reticulate- veined on both sides; inflorescence exceeding the 
leaves, at least in fruit; perianth lobes enlarged at the base of the 
ellipsoid fruit, this 11 mm. long, 7 mm. thick; pedicels very short, 
fleshy-turbinate. Referred to Phoebe with doubt, the flowers being 
unknown (Mez). Neg. 27633. 

Amazonas: Chachapoyas, Mathews, type. 

Phoebe maynensis Nees, Syst. Laur. 118. 1836; 214. P. peruvi- 
ana Meissn. var. glabriflora Meissn. in DC. Prodr. 15, pt. 1: 32. 1864. 



FLORA OF PERU 883 

A tall, glabrate tree, sometimes 40 meters high; branchlet tips 
and inflorescence minutely and sparsely puberulent; blades char- 
taceous, undulate-margined, glabrous but barbellate beneath in the 
nerve axils, somewhat 3-nerved, mostly 15 cm. long and 5 cm. wide, 
the acumination often 1 cm. long; filaments and sometimes the 4- 
celled anthers sparsely long-pilose, subequal ; staminodia subcordate, 
glabrous; style slightly shorter than the ovary. Negs. 359, 3595. 

Loreto: Yurimaguas, Poeppig 2305, type; at 200 meters, Williams 
4774 (det. Schmidt). Junin: La Merced, 600 meters, 5364 (det. 
Schmidt). San Martin: San Roque, 1,350 meters, Williams 7406, 
Juanjui, 400 meters, King 3768. "Muena." 

Phoebe pichisensis A. C. Smith, Bull. Torrey Club 58: 
103. 1931. 

A tree of 10 meters; branchlets sub terete, soon glabrous; petioles 
slender, puberulent, 6-10 mm. long; blades ovate-oblong, cuneate 
at the base, short-acuminate, '8-12 cm. long, 2.5-4 cm. wide, nearly 
glabrous above, pilose beneath, chartaceous, the lateral nerves pale, 
conspicuous; panicle many-flowered, minutely puberulent; flowers 
3-3.5 mm. long; staminodia glabrous, 0.5 mm. long. Very similar 
to P. brasiliensis Mez, with pilose staminodia. 

Junin: Pichis Trail, 900 meters, Killip & Smith 25430, type. 
San Martin: San Roque, 1,400 meters, Williams 7626, 7085 (det. 
Schmidt). "Pishcu muena," "muena." 

Phoebe Poeppigii Meissn. in DC. Prodr. 15, pt. 1: 33. 1864. 
Laurus triplinervis R. & P. Fl. Peruv. 4: pi. 363. 1802; Laurogr. pi. 9. 
P. triplinervis Mez, Jahrb. Bot. Gart. Berlin 5: 211. 1889, non 
Griseb. ; P. peruviana Meissn. op. cit. 32. P. heterotepala Mez, Repert. 
Nov. Sp. 3: 67. 1906. 

Very similar to P. cinnamomifolia (HBK.) Nees, but the leaves 
minutely pilose beneath, the flowers a little larger, their segments 
often unequal, the ovary and style subequal; glands subglobose; 
staminodia sagittate. A shrub or tree, sometimes 10-12 meters 
high (Weberbauer}. The unequal perianth segments, upon which 
character Mez establishes P. heterotepala, seem to me to be equally 
evident in the specimens of Ruiz and Poeppig. Negs. 27632, 7269, 
3606. 

Huanuco: Monzon, 1,200 meters, Weberbauer 3487, 3496; 284. 
Cochero, Ruiz & Pavon, type; Poeppig 1311. 



884 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

8. OCOTEAAubl. 

Scarcely recognizable among the genera having 4-celled anthers, 
except as indicated in the key, many characters being shared with 
other genera; the flowers, most often dioecious, may be perfect, the 
perianth tube conspicuous or none, the lobes persistent or deciduous, 
the filaments longer or shorter than the anthers, or none, at least 
those of series 3 glandular. Ovary various, the style longer or shorter. 
Cup of the fruit simple or double-margined. The anther cells are 
placed as in Persea and Phoebe, with which groups it could most 
readily be united if the perhaps arbitrary acceptance of the develop- 
ment of staminodia as a generic character should be discarded. In 
the following descriptions the flowers may be understood to be dioe- 
cious unless otherwise stated. 0. balanocarpa and 0. Trianae, 
described from fruiting material, are omitted from the following 
keys; the latter may suggest 0. guyanensis. 

Key based on leaf characters 

A. Leaves mostly less, often much less, than 10 cm. long, or at any 
rate about 4.5 cm. wide or narrower. 

Leaves conspicuously pubescent beneath (see 0. cuprea). 

Pubescence silvery, the trichomes long, closely appressed. 

0. guyanensis. 

Pubescence softly pilose on the leaves beneath 0. tenella. 

Pubescence a rusty or reddish tomentum. 

Leaves oblong-lanceolate, mostly 10-12 cm. long. . .0. Jelskii. 
Leaves ovate-elliptic, often shorter. 

Pubescence reddish; blades about 10 cm. long . .0. cardinalis. 
Pubescence rusty; blades about 6 cm. long. . .0. ferruginea. 
Leaves glabrous or more or less obscurely pubescent beneath. 
Leaves, at least the younger ones, sericeous-puberulent beneath. 

Leaves smooth above, not foveolate 0. cuprea. 

Leaves foveolate- veined 0. aciphylla. 

Leaves glabrous or glabrate. 

Flowers small, sometimes 2 mm. long; inflorescence glabrous 
or soon glabrate. 

Leaves firm, obscurely veined, heavy or fleshy-coriaceous 
and dull. 



FLORA OF PERU 885 

Leaves acute; filaments pilose 0. minutiflora. 

Leaves rounded at the apex; filaments glabrous. 

0. subrutilans. 

Leaves distinctly veined or thin, more or less lustrous. 
Petioles short, 3-7 mm. long. 

Leaves acute at both ends 0. minarum. 

Leaves rounded at the apex 0. jumbillensis. 

Leaves caudate-acuminate 0. tarapotensis. 

Petioles long and often slender, to 11 mm. long or longer. 

Inflorescence usually only 2-5 cm. long. 

0. leptobotra, 0. caudata. 
Inflorescences mostly elongate. 
Flowers usually dioecious. 
Leaves acuminate, often wider than 3.5 cm. 

Leaves 3.5-6.5 cm. wide; perianth lobes unequal. 

0. Killipii. 
Leaves 3-4 cm. wide; perianth lobes equal. 

0. marmellensis. 

Leaves acute, 2.5-3.5 cm. wide 0. minarum. 

Flowers perfect 0. Keriana. 

Flowers usually 2-2.5 mm. long; inflorescence pubescent or 
tomentulose. 

Leaves about 2 cm. wide. 

Flowers perfect; leaves acuminate 0. Weberbaueri. 

Flowers dioecious; leaves subobtuse 0. Mandoni. 

Leaves often 4 cm. wide or wider. 

Flowers racemose or subracemose 0. munacensis. 

Flowers most often subcorymbose. 

Branchlets more or less tomentulose; leaves acute. 

Leaves about 4 cm. wide; pedicels to 5 mm. long. 

0. monzonensis. 
Leaves 2.5-3.5 cm. wide; pedicels to 3 mm. long. 

0. minarum. 
Branchlets glabrate; leaves acuminate. 

Leaves usually much longer than the inflorescence; 
flowers dioecious. 



886 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves chartaceous; flowers racemose-paniculate. 

0. maynensis. 
Leaves rigid-coriaceous; flowers fastigiate-panic- 

ulate 0. Schomburgkiana. 

Leaves and inflorescence often subequal; flowers 

perfect. 
Leaves cuneate at the base, long-acuminate. 

0. Keriana. 

Leaves obtuse at the base, rounded or broadly 
acuminate at the apex 0. piurensis. 

A. Leaves mostly longer than 10 cm., or most of them definitely 

wider than 4.5 cm. 
B. Leaves medium-sized, mostly 12-15 cm. long and 4.5-7.5 cm. 

wide. 
C. Leaves pubescent, usually very minutely so, at least on the 

veins beneath. 

Leaves broadly elliptic or obovate, usually 5.5-8 cm. wide 
(see 0. aciphylla). 

Pubescence rusty, rather conspicuous; petioles about 1 cm. 

long. 
Inflorescence much shorter than the leaves; flowers gla- 

brate 0. macropoda. 

Inflorescence and leaves subequal (unless in 0. amazo- 
nica). 

Flowers puberulent 0. ovalifolia. 

Flowers pilose 0. amazonica. 

Pubescence pale, minute; petioles often only 5 mm. long. 

Petioles 12-18 mm. long; branchlets glabrous. 

0. Ottoschmidtii. 

Petioles all or most of them shorter than 10 mm. ; branch- 
lets, at least at first, pubescent. 

Branchlets terete, glabrate 0. rubrinervis. 

Branchlets sharply angled, slightly pubescent. 

Branchlets tomentulose; leaves acute-attenuate at 
the base 0. aurantiodora. 

Branchlets pilose, not tomentulose; leaf blades 
rounded at the base. . . .0. otuzensis. 



FLORA OF PERU 887 

Leaves lance-elliptic, usually narrower (see also 0. aurantio- 

dora and 0. amazonica). 

Petioles, except in 0. Raimondii, 1.5-2.5 cm. long; pubes- 
cence, at least at first, dense. 
Leaves more or less glabrate in age, the hairs loose or 

coarse; filaments glabrous 0. Rusbyana. 

Leaves densely and minutely pilose beneath, even in age; 

filaments pilose. 
Pubescence beneath obvious only under a lens, but 

the leaves bicolorous 0. Dielsiana. 

Pubescence on the leaves beneath obvious. 

0. Trianae, 0. Raimondii. 
Petioles usually about 1 cm. long; pubescence from the 

first sparse, except in 0. aciphylla. 
Pubescence a minute puberulence. 
Leaves acuminate. 

Pubescence sparse 0. puberula. 

Pubescence dense 0. aciphylla. 

Leaves obtuse or obscurely acute 0. micans. 

Pubescence a sparse pilosity 0. petalanthera. 

C. Leaves (mature) glabrous, or with some pubescence in the 

axils beneath, at the leaf base, or on the costa. 
Leaves more or less abruptly subcaudate-acuminate, the acu- 

mination 1 cm. long or longer (seeO. Killipii). 
Perianth tube none or extremely short. 

Inflorescence glabrous 0. tarapotana. 

Inflorescence at least minutely pubescent ... 0. maynensis. 
Perianth tube obvious. 

Leaves papyraceous, dull or little lustrous 0. caudata. 

Leaves coriaceous, very lustrous above. 

Leaves densely reticulate- veined beneath. .0. laxiflora. 
Leaves subobscurely and laxly reticulate-veined be- 
neath 0. leptobotra. 

Leaves obtuse, acute, or, if long-acuminate, gradually so. 
Perianth tube conspicuous. 

Inflorescence densely tomentulose 0. compacta. 

Inflorescence glabrous or glabrate. 



FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves conspicuously obovate, obtuse or rounded at 

the apex 0. obovata. 

Leaves scarcely obovate, or only the larger ones, acute. 

0. cuneifolia. 

Perianth tube obsolete or very short. 
Reticulate venation of the leaves obscure, even beneath. 
Inflorescence rather lax, the branchlets slender. 

0. multiglandulosa. 
Inflorescence very dense, the branchlets thick. 

0. architectorum. 

Reticulation of the leaves more or less prominent. 
Leaves densely white-puncticulate beneath; inflores- 
cence glabrate 0. albopunctulata. 

Leaves not white-punctate; inflorescence more or less 

pubescent. 
Flowers 2-2.5 mm. long. 

Perianth tube none 0. puberula. 

Perianth tube 1 mm. long 0. Killipii. 

Flowers 3.2-3.5 mm. long 0. Tessmannii. 

B. Leaves ample, mostly 20-35 cm. long, or at least 8 cm. wide 

(see 0. cuneifolia, 0. aurantiodora). 

Leaves broadly elliptic or obovate, about twice as long as wide 
(see sometimes 0. aurantiodora). 

Leaves pubescent beneath, the pubescence ferruginous. 

Petioles developed 0. ovalifolia. 

Petioles almost obsolete 0. licanioides. 

Leaves glabrous or glabrate beneath. 
Flowers pubescent. 
Leaves rigid-coriaceous; flowers sericeous, subsessile. 

0. caniflora. 
Leaves somewhat flexible; flowers moderately pubescent, 

pedicellate 0. obovata. 

Flowers glabrous. 

Leaves suboblong; branchlets angulate 0. olivacea. 

Leaves oval-elliptic; branchlets terete. .0. marowynensis. 
Leaves lance-elliptic, 3 or more times as long as wide. 
Pedicels 2-4 (1-5) mm. long. 



FLORA OF PERU 889 

Leaves thin; petioles mostly 2.5-3 cm. long. .0. ucayalensis. 
Leaves thick; petioles mostly 2 cm. long or shorter. 

0. aurantiodora. 
Pedicels very short or obsolete. 

Inflorescence dense; leaves rigid-coriaceous in age, obscurely 

reticulate- veined 0. amplissima. 

Inflorescence very large and open; leaves rather thin or, if 

rigid, strongly reticulate- veined. 
Leaves densely pubescent beneath 0. magnified. 

Leaves glabrous or minutely puberulent beneath. 

0. grandifolia. 

Technical key 
A. Flowers perfect. 

Leaves definitely reddish-pubescent beneath. 

Perianth tube obsolete 0. ovalifolia. 

Perianth tube well developed. 

Anthers elliptic or subovate 0. Jelskii. 

Anthers quadrate. 

Flowers 2-2.5 mm. long 0. cardinalis. 

Flowers 4-5 mm. long 0. ferruginea. 

Leaves glabrous or more or less obscurely pubescent beneath. 

B. Anthers of series 1 and 2 subsessile, or longer than the fila- 
ments. 
Leaves 20 cm. long or longer. 

Leaves chartaceous; inflorescence glabrous 0. olivacea. 

Leaves fleshy-rigid ; inflorescence slightly tomentulose. 

0. amplissima. 
Leaves smaller, usually much smaller. 

Leaves obovate, at least the larger ones. 

Perianth tube obsolete 0. jumbillensis. 

Perianth tube and lobes subequal. 

Flowers 2-3.5 (-4) mm. long; outer anthers ovate; 
style obvious 0. cuneifolia. 

Flowers 2 mm. long; anthers subquadrate; stigma sub- 
sessile 0. minutiflora. 

Leaves not obovate, sometimes oblanceolate, acuminate. 



890 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves densely foveolate 0. aciphylla. 

Leaves smooth. 

Inflorescence 5-14 cm. long. 
Leaves coriaceous, conspicuously reticulate-veined 

beneath 0. piurensis. 

Leaves chartaceous, less conspicuously reticulate- 
veined. 

Flowers 3.5-4 mm. wide 0. Raimondii. 

Flowers smaller. 
Leaves acuminate, 10 cm. long or longer. 

Petioles to 11 mm. long; inflorescence puber- 

ulent 0. Keriana. 

Petioles to 18 mm. long; inflorescence gla- 
brous 0. Ottoschmidtii. 

Leaves acute, rarely if ever 10 cm. long. 

0. minarum. 
Inflorescence 2-3 (-4) cm. long. 

Leaves glabrous 0. Weberbaueri. 

Leaves soft-pubescent beneath 0. tenella. 

B. Anthers of series 1 and 2 shorter than or barely equaling the 

well developed filaments. 

Perianth tube conspicuous; pedicels 3-6 mm. long. 
Ovary and style subequal; leaves obtuse at the base. 

0. piurensis. 
Ovary shorter than the slender style; leaves cuneate at the 

base 0. compacta. 

Perianth tube obsolete; pedicels 2-3 mm. long. .0. Tessmannii. 
A. Flowers dioecious or rarely monoecious. 
Leaves ample, mostly 20-35 cm. long and 8 cm. wide or wider. 
Leaves pubescent beneath or glabrate in age. 
Leaves densely villous beneath. 

Leaves subsessile, the short petioles stout. . . .0. licanioides. 
Leaves conspicuously petiolate, the petioles slender. 

0. magnified. 

Leaves sparsely pilose or becoming glabrate. .0. aurantiodora. 
Leaves glabrous, even beneath, or essentially so. 
Flowers white-pubescent. 



FLORA OF PERU 891 

Leaves very rigid, elliptic; style pilose 0. caniflora. 

Leaves coriaceous, obovate; style glabrous. . .0. grandifolia. 
Flowers glabrous or nearly so. 
Perianth tube conspicuous, at least 1 mm. long. .0. caudata. 

Perianth tube obsolete 0. ucayalensis. 

Leaves small to medium-sized, rarely longer than 15 cm. 
C. Outer filaments pubescent, sometimes very short. 
Leaves (at least the younger ones) beautifully sericeous; fila- 
ments very short. 
Pubescence of the younger leaves coppery; perianth tube 

obsolete 0. cuprea. 

Pubescence of the leaves silvery beneath; perianth tube 

conspicuous 0. guyanensis. 

Leaves less pubescent; filaments often well developed. 

Flowers mostly or all conspicuously pedicellate, the pedi- 
cels 3-7 mm. long, or only a few of them shorter. 

Leaves obovate; flowers white 0. obovata. 

Leaves elliptic; flowers greenish yellow. . .0. monzonensis. 
Flowers mostly or all subsessile, the longest pedicels 2 (-3) 
mm. long. 

Perianth tube well developed 0. otuzensis. 

Perianth tube obsolete. 

Inflorescence ample, longer than the leaves. 

0. Dielsiana. 
Inflorescence shorter than the leaves (see, rarely, 0. 

cuprea) . 
Flowers strigose; leaves sparsely strigose beneath 

on the nerves 0. petalanthera. 

Flowers glabrous or puberulent, like the leaves 
beneath, or the leaves barbellate in the axils of 
the nerves. 

Blades about 15 cm. long 0. puberula. 

Blades 3-4 cm. long 0. Mandonii. 

C. Outer filaments glabrous, sometimes very short or even 

lacking. 

Inflorescence conspicuously shorter than the leaves (some- 
times terminal). 



892 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves lustrous-pubescent beneath 0. micans. 

Leaves, if pubescent, not lustrous. 
Leaves densely white-puncticulate beneath. 

0. albopuncticulata. 
Leaves less or not at all puncticulate. 
Inflorescence glabrous. 
Perianth tube evident; anthers quadrate, obtuse, 

sessile or subsessile 0. caudata. 

Perianth tube short or obsolete; anthers ovate or 

elliptic and acute 0. tarapotana. 

Inflorescence pubescent. 
Leaves tomentose or pilose beneath. 

Flowers glabrate 0. macropoda. 

Flowers pilose 0. amazonica. 

Leaves glabrous, or the nerves pilose beneath. 
Leaves pilose, at least on the nerves. 
Inflorescence puberulent, racemose. 

0. munacensis. 

Inflorescence pilose, pyramidal. . .0. amazonica. 
Leaves glabrous. 
Leaves rigid; style well developed. 

Flowers subsessile 0. architectorum. 

Flowers pedicellate 0. Schomburgkiana. 

Leaves thin or flexible; style obsolete. 

0. maynensis. 

Inflorescence more or less approximating the leaves in length, 
usually terminal or sub terminal. 

Inflorescence glabrous or essentially so (seeO. marmellensis). 

Leaves fleshy-coriaceous, rounded at the apex. 

0. subrutilans. 
Leaves chartaceous or finally coriaceous, more or less 

acuminate. 

Leaves more or less, sometimes obscurely, pubescent 
beneath. 

Leaves caudate-acuminate 0. caudata. 

Leaves obtusely acuminate 0. Killipii. 

Leaves glabrous. 



FLORA OF PERU 893 

Anthers subquadrate, obtuse; inflorescence narrowly 

paniculate 0. laxiflora. 

Anthers subovate, acutish; inflorescence squarrose- 

paniculate 0. leptobotra. 

Inflorescence at least finely pubescent. 
Petioles short, 5-8 mm. long. 

Leaves to 10 cm. long 0. rubrinervis. 

Leaves 20 cm. long or longer 0. licanioides. 

Petioles all or mostly longer than 8 mm. 

Perianth tube conspicuous 0. aurantiodora. 

Perianth tube none or short. 
Anthers more or less emarginate; leaves obscurely 

short-acuminate. 

Inflorescence lax, paniculate, pilose; ovary pilose. 

0. multiglandulosa. 

Inflorescence thyrsoid, tomentulose; ovary gla- 
brous 0. minarum. 

Anthers ovate, acute; leaves acuminate .0. Rusbyana. 
Anthers rectangular; leaves caudate. 

0. marmellensis. 

Ocotea aciphylla (Nees) Mez, Jahrb. Bot. Gart. Berlin 5: 243. 
1889. Oreodaphne aciphylla Nees, Linnaea 8: 43. 1833. 

A tree 20 meters high, the bark aromatic; petioles to 13 mm. long; 
blades coriaceous only in age, glabrous and typically very lustrous 
above, more or less sericeous beneath, densely punctulate-foveolate- 
venulose on both sides, ovate-lanceolate or lanceolate, scarcely or 
shortly acute at the base, long-acuminate, about 10 cm. long and 
2.5-3 cm. wide; inflorescence laxly pyramidal, subhirsute, shorter 
than the leaves; flowers perfect, tomentose, the tube conspicuous; 
filaments densely pilose; anthers ovate, very acute; ovary glabrous. 
Neg. 22071. 

Loreto: Mishuyacu, 100 meters, Klug 891, 608, 755. Florida, 
180 meters, Klug 2315. Brazil. "Muena negra." 

Ocotea albopunctulata Mez, Repert. Sp. Nov. 16: 307. 1920. 

A tree of 30 meters, with terete, soon glabrate branchlets and 
coriaceous, elliptic, broadly acuminate leaves, usually a little more 
than 10 cm. long and less than half as wide; petioles often 2 cm. long; 
blades glaucescent green above, dull and densely white-punctulate 



894 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

beneath, obscurely pilose, at least on the nerves; inflorescences much 
shorter than the leaves; pedicels to 3 mm. long; flowers 2.5 mm. long, 
the tube very short; filaments glabrous; anthers large, subovate, 
rounded at the apex. Neg. 3618. 

Rio Acre: Seringal San Francisco, Ule 9398, type. Near mouth 
of Rio Macauhan, Krukoff 5279. 

Ocotea amazonica (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
359. 1889. Oreodaphne amazonica Meissn. in "DC. Prodr. 15, pt. 1: 
123. 1864. 

Younger branchlets ferruginous- tomentose, angled; petioles to 
13 mm. long, tomentose; blades chartaceous or rather membranous, 
at maturity lustrous above and glabrous except for the somewhat 
tomentulose, impressed nerves, pilose and prominently nerved be- 
neath, reticulate-veined on both sides, elliptic or obovate, acute at 
the base, mucronate-acuminate, about 13 cm. long and 3.5 cm. wide; 
inflorescence few-flowered, typically much shorter than the leaves, 
ferruginous- tomentose; pedicels 1-3 mm. long; staminate flowers 
2-3 mm. long, tomentose-pilose, the tube obsolete, the segments 
reflexed, acute; filaments glabrous, longer than the anthers, these 
subrectangular, obtuse; staminodia none. Neg. 3619. 

Peru: Possibly. Amazonian Brazil. 

Ocotea amplissima Mez, Repert. Nov. Sp. 3: 68. 1906. 

A well marked shrub because of its thick, angled, glabrate branch- 
lets and its large (about 30 cm. long and 10 cm. wide), extremely 
fleshy-rigid, elliptic leaves, obtuse or acute at the base, beautifully 
acuminate; inflorescence slightly tomentulose; flowers perfect; peri- 
anth tube elongate, equaling the lobes; pedicels to 1.5 mm. long; 
anthers suborbicular, obviously contracted at the base; ovary gla- 
brous, slightly shorter than the style. A shrub of 3-4.5 meters, with 
green flowers, allied to 0. Weberbaueri Mez and 0. Stuebelii Mez, 
the latter of Colombia, with leaves a third smaller, the pedicels to 
3 mm. long. Neg. 3620. 

Huanuco: Monzon, 2,000 meters, Weberbauer 3550, type; 255. 
Pan de Azucar, Sawada 80 (det. Schmidt). 

Ocotea architectorum Mez, Repert. Nov. Sp. 3: 70. 1906. 

A tall tree, 20 meters high, with minutely puberulent, subangu- 
late branchlets, rigid, glabrous, elliptic blades, and short, terminal, 
cinereous- tomentulose inflorescences of subsessile, white flowers; 
petioles about 1 cm. long; blades short-acuminate, about 15 cm. long 



FLORA OF PERU - 895 

and 6-7 cm. wide; perianth tube short; filaments glabrous, shorter 
than the subrectangular anthers; ovary glabrous, elliptic, equaling 
the style. Wood used for construction. Bark of the branches 
aromatic (Weberbauer). Compared by Mez with the West Indian 
0. leucoxylon (Sw.) Mez. 0. ceanothifolia (Nees) Mez, 365, similar 
but with leaves 10 cm. long and 3 cm. wide, and conspicuous perianth 
tube, may extend from Brazil. Neg. 3622. 

Cajamarca: Chugur, 2,700 meters, Weberbauer 4079, type; 258, 
259. "Roble bianco." 

Ocotea aurantiodora (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 
5: 295. 1889. Laurus aurantiodora R. & P. Fl. Peruv. 4: pi. 349. 1802; 
Laurogr. pi. 15. Mespilodaphne aurantiodora Meissn. in DC. Prodr. 
15, pt. 1:101. 1864. 

Branches acutely angled, tomentulose; petioles 2-3 cm. long; 
blades acutely attenuate at the base, more or less abruptly acute 
or acuminate at the apex, sometimes very broadly elliptic, often 15- 
20 cm. long and 10 cm. wide, or even larger, lustrous above, sparsely 
pilose-strigillose beneath or finally glabrate, rigid-coriaceous, promi- 
nently reticulate- veined on both sides; flowers pubescent, the peri- 
anth tube conspicuous; filaments glabrous, those of the outer series a 
third as long as the minutely emarginate, ovate-rectangular anthers. 
A shrub or tree with loose, many-flowered inflorescences about 
equaling the leaves. The type at Madrid, in fruit, has only upper 
leaves, the largest 4.5 cm. wide and 13 cm. long. Comparable is 
0. opifera Mart., 291, of the upper Amazon, with tomentulose in- 
florescence, very short perianth tube, and ovate anthers. 0. Sprucei 
(Meissn.) Mez, 323, and 0. viburnoides (Meissn.) Mez, 322, have 
narrower leaves, 4-5 cm. wide; the former has leaves pruinose- 
glaucous beneath and obscurely reticulate- veined ; the latter has 
leaves scarcely paler beneath but densely and minutely reticulate- 
veined on both sides. Neg. 29399. 

Huanuco: Cochero and Chinchao, Ruiz & Pavon, type. Chin- 
chao, Raimondi. Cajamarca: Tambillo, Raimondi. Colombia. 
"Negrito." 

Ocotea balanocarpa (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 
388. 1889. Laurus balanocarpa R. & P. Fl. Peruv. 4: pi. 354. 1802; 
Laurogr. pi. 16. Mespilodaphne balanocarpa Nees, Linnaea 21: 
495. 1848. 

A shrub or tree, known only in fruit, with minutely strigillose, 
subangulate young branches and coriaceous, glabrous, ovate-elliptic 



896 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves, densely and obviously foveolate-reticulate, especially beneath ; 
petioles to 7 mm. long; blades acute at the base, more or less acumi- 
nate at the apex, about 10 cm. long and 5 cm. wide; inflorescence 
apparently few-flowered; cup of the fruit subglobose, entire-margined, 
the smooth, subglobose fruit nearly 2 cm. long, mucronulate. Neg. 
3627. 

Peru: Ruiz & Pavon, type. 

Ocotea caniflora Mez, Repert. Nov. Sp. 3: 69. 1906. 

A tree as much as 20 meters high, with exceptionally rigid, ellip- 
tic, short-acuminate leaves about 20 cm. long and 8 cm. wide; 
branchlets subangulate, obscurely ferruginous-tomentulose, like the 
crowded inflorescences; flowers white, white-pubescent, the perianth 
tube short; filaments glabrous, equaling the subrectangular anthers; 
ovary glabrous, the shorter style densely pilose toward the tip. 
Neg. 3636. 

Cuzco: Santa Ana, 1,700 meters, Weberbauer 5036, type. 

Ocotea cardinalis Mez, Repert. Nov. Sp. 3: 69. 1906. 

A well marked shrub because of its distinctly red-tomentose 
pubescence, especially on the leaves beneath and the inflorescence; 
petioles about 1 cm. long; blades acute at the base, broadly short- 
acuminate, about 4.5 cm. wide; flowers green, perfect, 2-2.5 mm. long, 
with a short tube, the pedicels 1 mm. long; filaments slightly pilose, 
about as long as the quadrate anthers; style shorter than the gla- 
brous ovary. Related to 0. discolor Mez, 263, of Colombia, with 
much shorter petioles, narrower leaves, and flowers 3-5 mm. long. 
Neg. 3637. 

Huanuco: Monzon, 2,000 meters, Weberbauer 3545, type; 255. 

Ocotea caudata (Nees) Mez, Jahrb. Bot. Gart. Berlin 5: 378. 
1889. Oreodaphne caudata Nees, Linnaea 21: 519. 1848. Ocotea 
marowynensis Mez, op. cit. 380. 

A shrub or small tree of 2-6 meters, well marked by its thin (in 
age subcoriaceous), elliptic-lanceolate, definitely caudate-acuminate 
leaves, mostly about 10 cm. long and 4-5 cm. wide; petioles to 18 
mm. long; reticulation of the leaves conspicuous; inflorescence often 
only 2-4 cm. long, with widely spreading branchlets, glabrate; 
flowers glabrous, 1-1.5 mm. long, on pedicels at least twice as long; 
perianth tube rather conspicuous; anthers of the outer series quad- 
rate, sessile, the filaments pilose; style much exceeded by the 
glabrous ovary. Flowers brownish yellow. 



FLORA OF PERU 897 

Loreto: Iquitos, 100 meters, Tessmann 5148. Mishuyacu, 100 
meters, King 123. San Martin: Tarapoto, Ule 6503. Amazonian 
Brazil; British Guiana. 

Ocotea cuneifolia (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 
259. 1889. Laurus cuneifolia R. & P. Fl. Peruv. 4: pi. 359. 1802; 
Laurogr. pi. 28. Oreodaphne cuneata Nees, Syst. Laur. 385. 1836. 
Ocotea cuneata Macbr. Field Mus. Bot. 11: 18. 1931, non Gomez, 1894. 

A tree of 13 meters, the branchlets subterete, minutely puberulent; 
petioles slender, 1-1.5 (-2) cm. long; blades glabrous, or barbellate 
in the nerve axils beneath, subcoriaceous, elliptic-lanceolate or some- 
what obovate, short-acuminate or acute, mostly about 10 cm. long 
and 5 cm. wide but sometimes much larger; inflorescence shorter than 
the leaves, dense, glabrate or puberulent; flowers perfect, greenish, 
2-3 (-4) mm. long, on pedicels as long or longer; filaments very short, 
densely pilose, the ovate anthers minutely emarginate; ovary gla- 
brous, the style as long. Neg. 27612. 

Huanuco: Cochero, Ruiz & Pavdn, type; Dombey. Without local- 
ity, Raimondi. Monzon, 900 meters, Weberbauer 3445; 285. Without 
locality: Ruiz & Pavon 13-34. Bolivia. 

Ocotea cuprea (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 299. 
1889. Oreodaphne cuprea Meissn. in DC. Prodr. 15, pt. 1: 114. 1864. 

Readily recognized by the oblong-lanceolate leaves, about 7 cm. 
long and 2-3 (-5) cm. wide, copper-colored beneath with a fine, 
sericeous pubescence, at least when young, sometimes glabrate; 
inflorescence few-many-flowered, thyrsoid; pedicels 1-2 mm. long; 
flowers yellowish, 2 mm. long; perianth tube obsolete; filaments 
pilose, very short, the anthers somewhat emarginate; ovary glabrous, 
attenuate into a style twice as long. A tree 8-17 meters high, with 
the aspect of some Myristicaceae. Neg. 3648. 

San Martin: Tarapoto, Spruce 4844, type. Loreto: Iquitos, 
100 meters, Tessmann 5349; Killip & Smith 27302 (det. Schmidt). 
Balsapuerto, 220 meters, Klug 2952. Pebas, Williams 1884. Puma- 
yacu, 600-1,200 meters, Klug 3240. Bolivia. 

Ocotea Dielsiana 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 10: 
232. 1928. 

Young branches slightly tomentulose; blades elliptic or obovate- 
elliptic, narrowed at the base, subrotund and short-acuminate at 
the apex, 12-16 cm. long, 3.6-5.8 cm. wide, thin, glabrous above 
and apparently so beneath, but under a lens with a fine, close, 



898 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

appressed pubescence apparent; inflorescence ample, to almost 20 cm. 
long; flowers dioecious, yellowish, 3.5 mm. long, on pedicels 1 mm. 
long; perianth tube obsolete; filaments pilose, one- third as long as the 
subrectangular, not emarginate but attenuate anthers (of series 1); 
ovary glabrous. Allied to 0. opifera Mart., which see under 0. auran- 
tiodora and 0. grandifolia. According to the collector, a tree of 
6 meters, with a trunk diameter of 17 cm. Neg. 3650. 

Loreto: Iquitos, Tessmann 5097, type; 3629. "Muena," "muena 
blanca." 

Ocotea ferruginea (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
265. 1889. Pleurothyrium ferrugineum Meissn. in DC. Prodr. 15, 
pt. 1:169. 1864. 

Branchlets, leaves beneath, and inflorescences densely ferrugi- 
nous- tomentose; petioles 3-8 mm. long; blades very rigid, in age 
glabrous above, prominently reticulate-veined only beneath, ovate 
or broadly elliptic, mostly about 5 cm. long and 3 cm. wide, rounded 
or barely acute at each end; inflorescences few-flowered, shorter than 
the leaves; perianth tube elongate; flowers 4-5 mm. long, perfect; 
filaments pilose, the anthers quadrate; staminodia stipitiform; ovary 
glabrous. A shrub of 1-2 meters with greenish yellow flowers (Weber- 
bauer). Neg. 27615. 

Cajamarca: Cutervo (Jelski 178, 193, 199}; Raimondi. Yam- 
brasbamba, Mathews 1429, type. Molinopampa, 2,000 meters, 
Weberbauer 4343 (det. Mez). Amazonas: Cuelap, 2,600 meters, 
Weberbauer 4323; 263. 

Ocotea grandifolia (Nees) Mez, Jahrb. Bot. Gart. Berlin 5: 
290. 1889. Oreodaphne grandifolia Nees, Linnaea 21: 517. 1848. 

A tree or tall shrub of 5-25 meters, easily recognized by its gla- 
brous, deeply and acutely angled branchlets; leaves obovate, acute, 
often 30 cm. long and 10 cm. wide, granulate above, conspicuously 
reticulate-veined on both sides, glabrous or nearly so; inflorescence 
sparsely strigillose; pedicels to 1.5 mm. long; flowers pilose, 2 mm. 
long; perianth tube obsolete or in the pistillate flowers conspicuous; 
filaments glabrous, the anthers subovate; ovary glabrous, the style 
slightly shorter. Flowers yellow, turning yellowish brown. 0. opi- 
fera Mart., 291, is similar, if, indeed, it is not the same, or a variety 
with branchlets less definitely wing-angled; leaves acuminate, more 
pubescent beneath, the inflorescence more or less hirsute. Neg. 3664. 



FLORA OF PERU 899 

Loreto: Yurimaguas, Poeppig 2069, type; Ule 6291 (det. Mez); 
at 200 meters, Williams 4609, 7833 (det. Schmidt); Kittip & Smith 
27554, 29056. Mouth of Rio Santiago, Tessmann 4702. Florida, 180 
meters, King 2194. San Martin: Tarapoto, 400-900 meters, Wil- 
liams 6618 (det. Schmidt). Brazil. "Sipra muena," "muenablanca," 
"maraco-f uina. " 

Ocotea guyanensis Aubl. PL Guian. 781. pi. 310. 1775; 296. 

Recognizable by the beautiful luster of the sericeous pubescence 
that densely clothes the under side of the leaves; blades glabrous 
above, reticulate-veined, mostly 10-12 cm. long and 2.5-3 cm. wide, 
gradually attenuate to the acuminate apex; flowers 3-5 mm. long; 
anthers ovate, the short filaments tomentulose. A tree of 5-20 
meters, with pyramidal, many-flowered inflorescences about equal- 
ing the leaves. 

Loreto: Cerro de Escalera, 1,100 meters, Ule 44p (det. Mez). 
Bolivia to the Guianas. 

Ocotea Jelskii Mez, Jahrb. Bot. Gart. Berlin 5: 261. 1889. 

Similar to 0. ferruginea (Meissn.) Mez, but the leaves less reticu- 
late-veined beneath, less rigid, and twice as long; inflorescences 
equaling or exceeding the leaves; filaments pilose; anthers elliptic 
or subovate, acutish or obtuse. A tree of 8-12 meters (Jelski). 0. 
Benthamiana Mez, of Ecuador, related, has leaves more than 20 cm. 
long, about 10 cm. wide, densely reticulate-veined beneath, gla- 
brous filaments, apically rounded anthers, and no staminodia. 

Cajamarca: Tambillo (Jelskii 167, 190, 1205); Raimondi (det. 
Schmidt). Bolivia(?). "Ishpingo." 

Ocotea jumbillensis 0. C. Schmidt, Repert. Sp. Nov. 31: 
182. 1933. 

A small tree, 5 meters high, the glabrous, terete branches with 
many large lenticels; petioles 3-5 mm. long, glabrous; blades elliptic 
or narrowly obovate-elliptic, narrowed and rounded at the base, or 
sometimes decurrent to the petiole, rounded at the apex, 2-4 cm. long, 
about 1.5 (-2) cm. wide, coriaceous, glabrous except for the nerves, 
these 7-10, the slender reticulation more or less prominent on both 
sides; inflorescences glabrous, racemose-paniculate, 2.5-5.5 cm. long; 
flowers perfect, greenish, barely 4 mm. wide, tomentulose, like the 
pedicels, these 1-3 mm. long; perianth tube obscurely developed; 
outer anthers ovate, rectangular, broadly truncate, 1 mm. long, the 



900 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sparsely pilose filaments 0.3 mm. long; ovary narrowly ellipsoid, 
1 mm. long, the style half as long. Compared by the author with 
0. vaccinioides (Meissn.) Mez, 252, of eastern Brazil. 

Amazonas: Between Jumbilla and San Carlos, 2,900 meters, 
Weberbauer 7155, type. 

Ocotea Keriana A. C. Smith, Bull. Torrey Club 58: 104. 1931. 

A tree of 9 meters; twigs and leaves soon glabrous, or the latter 
minutely tomentulose on the nerves beneath; petioles slender, 6-11 
mm. long; blades oblong-ovate-lanceolate, cuneate at the base, long- 
acuminate at the apex, 11-17 cm. long, 3.5-5.5 cm. wide, chartaceous; 
panicles many-flowered, minutely tomentulose, to 14 cm. long; 
flowers perfect, pale yellow, the pedicels 1-3 mm. long; perianth tube 
1 mm. long, the lobes 1.8 mm. long; stamens all fertile, the glands of 
the third series 0.3 mm. thick; ovary glabrous, the style subequal. 

Loreto: Iquitos, 100 meters, Killip & Smith 27111, type. 

Ocotea Killipii A. C. Smith, Bull. Torrey Club 58: 108. 1931. 

A tree of 6-9 meters, the young twigs, petioles, and blades 
deciduously pilose; petioles slender, to 15 mm. long; blades ovate- 
oblong, cuneate at the base, obtuse-acuminate at the apex, 10-16 cm. 
long, 3.5-6.5 cm. wide, finally subcoriaceous, abundantly reticulate- 
veined; staminate inflorescences many-flowered, essentially glabrous; 
pedicels to 1.5 mm. long; perianth tube 1 mm. long, the ovate-oblong 
outer lobes broader than the inner ones; stamens all fertile, the 
filaments only of series 3 (eglandular) equaling the anthers; lower 
anther cells obvious, the connective not produced. Related to 0. 
laxiflora (Meissn.) Mez; see also 0. leptobotra. 

Junin: Rio Pinedo, La Merced, 900 meters, Killip & Smith 23648, 
type. Loreto: Masisea, Rio Ucayali, 275 meters, Killip & Smith 
26861. San Martin: Tarapoto, 400-900 meters, Williams 6708. 

Ocotea laxiflora (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
371. 1889. Mespilodaphne laxiflora Meissn. in DC. Prodr. 15, pt. 1: 
107. 1864. 

A shrub or small tree, glabrous except the tips of the young 
branchlets; petioles to 1 cm. long; blades subcoriaceous, strongly 
reticulate-veined on both sides, lustrous above, elliptic-lanceolate, 
acute at the base, acuminate at the apex, 10-17 cm. long, 4 cm. wide 
or wider; inflorescences laxly and narrowly paniculate or subpanicu- 
late, racemose, somewhat shorter than the leaves; perianth tube con- 
spicuous; anthers sessile or nearly so, obtuse or minutely emarginate; 



FLORA OF PERU 901 

style much shorter than the glabrous ovary. 0. debilis Mez, 370, 
related, with obsolete perianth tube and exceptionally lax inflores- 
cences, may extend from Brazil; also 0. pauciflora (Nees) Mez, 370, 
like 0. laxiflora, but the leaves conspicuously acuminate, only 3 cm. 
wide, the filaments pilose. Illustrated, Mart. Fl. Bras. 5, pt. 2: pi. 75. 
Negs. 7278, 7286. 

Loreto: Mishuyacu, 100 meters, Klug 486 (det. Schmidt). Brazil 
to Colombia and the Guianas. 

Ocotea leptobotra (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 
376. 1889. Laurus leptobotra R. & P. Fl. Peruv. 4: pi. 368. 1802; 
Laurogr. pi. 21. L. nitida R. & P. Fl. Peruv. 4: pi. 353. 1802; Laurogr. 
pi. 13. Oreodaphne leptobotra Nees, Linnaea 21: 520. 1848. 0. ter- 
minalis Nees, Syst. Laur. 426. 1836. 

Similar to 0. caudata, but rather readily distinguished by the 
lustrous, more coriaceous leaves, reticulate-veined beneath, less 
abruptly caudate-acuminate, and often 7-8 cm. wide (sometimes only 
half as wide); anthers sessile, subovate, acutish. Also resembling 
0. laxiflora, but the leaves thinner, less densely reticulate- veined, and 
the inflorescence laxer. Neg. 27619. 

Huanuco: Chacahuassi, Ruiz & Pawn. Cochero, Ruiz & Pavon. 
, San Martin: Tocache, Poeppig 1861. Tarapoto (Spruce 3918). 

Ocotea licanioides A. C. Smith, Bull. Torrey Club 58: 107. 1931. 

A shrub of 3 meters, with drooping branches; twigs, leaves 
beneath and inflorescences throughout densely brownish-villous- 
tomentose; petioles stout, 5-8 mm. long; blades oblong, truncate or 
broadly cuneate at the base, subacute(?) at the apex, 20-25 cm. long, 
8-10 cm. wide, thick-coriaceous, finally glabrous above; staminate 
inflorescences many-flowered; pedicels 1 mm. long or shorter; peri- 
anth tube obsolete; stamens all fertile, the filaments glabrous, the 
stamens of the fourth series and the gynecium lacking. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 1506, type. 
Upper Rio Nanay, Williams 1118. "Muena negra." 

Ocotea macropoda (HBK.) Mez, Jahrb. Bot. Gart. Berlin 5: 
348. 1889. Persea macropoda HBK. Nov. Gen. & Sp. 2: 160. 1817. 

A shrub or tree, villous-tomentose on the angled branchlets and 
the leaf veins beneath; petioles to 17 mm. long; blades elliptic, acute, 
mostly 10-12 cm. long, subcoriaceous, rather prominently reticulate- 
veined, becoming glabrate and lustrous above; inflorescences laxly 



902 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

thyrsoid, much shorter than the leaves; flowers subsessile, subgla- 
brous, 2.5-3 mm. long; perianth tube obsolete; filaments glabrous, 
the subquadrate anthers sub truncate; glabrous ovary and style sub- 
equal. Neg. 35023. 

Cajamarca: Tambillo, Raimondi; (Jelski 169). Colombia; Brazil. 
"Tashango corazon negro." 

Ocotea magnifica 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10:233. 1928. 

Branches thick or fleshy, angled, densely ferruginous-velutinous, 
like the leaves beneath and on the nerves above; blades lanceolate, 
narrowed to the base, acuminate, 25-35 cm. long, 8-9 cm. wide; 
inflorescence to 50 cm. long, few-flowered; flowers greenish yellow, 
3.5 mm. long; perianth tube obsolete; outer anthers elliptic-rec- 
tangular, slightly longer than the pilose filaments; ovary glabrous. 
A tree related to 0. opifera Mart., which see under 0. aurantiodora. 
Trunk 18 cm. in diameter (Tessmann). 

Loreto: Yarina-cocha, 150 meters, Tessmann 3477, type. 

Ocotea Mandonii Mez, Jahrb. Bot. Gart. Berlin 5: 311. 1889. 

A glabrate shrub or small tree, the rigid, broadly elliptic leaves 
barbellate in the nerve axils beneath, the inflorescence more or less 
pilose; petioles about 7 mm. long; blades broadly elliptic, cuneate- 
narrowed to the base, more or less obtuse, conspicuously reticulate- 
veined on both sides, the upper surface somewhat lustrous, about 
3.5 cm. long and 2 cm. wide; inflorescence as long as the leaves, sub- 
corymbose-paniculate; pedicels 1-3 mm. long; perianth tube none; 
filaments sparsely pilose, slightly shorter than the rectangular, 
obtuse anthers; ovary subglobose, about equaled by the style; fruit 
ellipsoid, 2.5 cm. long, 1 cm. thick, attenuate at the base, the cup 
plane. Neg. 3682. 

Peru: Probably. Northern Bolivia. 

Ocotea marmellensis Mez, Bull. Herb. Boiss. II. 5: 238. 1905. 

Branchlets slender, obscurely angled, minutely tomentulose at 
the tips; petioles slender, about 1 cm. long; blades oblong, rather 
abruptly caudate-acuminate, slightly lustrous, finely reticulate- 
veined on both sides, chartaceous, typically puberulent-pilose 
beneath or glabrous, 10-13 cm. long, 3-4 cm. wide; flowers loosely 
paniculate, the inflorescence shorter than or equaling the leaves, 
typically puberulent but often glabrous; perianth slightly puberulent 
or glabrous, the tube obvious, with the equal segments 2 mm. long, 



FLORA OF PERU 903 

these exceeding the glabrous stamens; filaments and rectangular, 
obtuse anthers subequal; staminodia none; basal glands of series 3 
large, sessile. The Rio Acre material is glabrous or essentially so, 
with more abruptly acuminate leaves longer than the inflorescences, 
and may be known as var. acrensis Macbr., var. nov. Glabra vel 
subglabra; foliis abrupte acuminatis inflorescentiis plus minusve 
longioribus. A tree about 30 meters high (Krukoff). Neg. 3683. 

Rio Acre: Rio Macauhan, Krukoff 5262 (type of the variety); 
Krukoff 5529. Amazonian Brazil. 

Ocotea maynensis (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
359. 1889. Oreodaphne maynensis Meissn. in DC. Prodr. 15, pt. 1: 
129. 1864. 

A medium-sized tree with glabrate, terete branchlets and thin 
or rather thick, ovate-elliptic, glabrous, caudate-acuminate leaves 
about 10 cm. long and 3.5-4 cm. wide; leaf reticulation prominent on 
both sides; inflorescence slightly pilose-hirsutulous, much shorter 
than the leaves; pedicels 1-2 mm. long; flowers 2-2.5 mm. long; 
perianth tube obsolete; filaments glabrous, slightly longer than the 
subquadrate anthers; ovary glabrous, the stigma subsessile. A tree 
of 8-12 meters, the trunk 20-40 cm. in diameter, branching at about 
4 meters (Tessmanri). Flowers white, the wood soft, yellow (Poep- 
pig~). Flowers yellowish green (Tessmann). Neg. 3686. 

Loreto: Yurimaguas, King 2889, 2834, 2829; Poeppig 2364, type; 
2305; Williams 4043, 4574 (det. Schmidt). Iquitos, 100 meters, 
Tessmann 3536, 3627, 5113, Balsapuerto, Killip & Smith 28401, 
28604, 28390, 28597; King 3010. Mishuyacu, 100 meters, King 
1397, 943, 895, 401, 883, 1597, 2889. San Martin: Tarapoto, 400- 
900 meters, Williams 6599. "Muena." 

Ocotea micans Mez, Jahrb. Bot. Gart. Berlin 5: 297. 1889. 

Similar to 0. guyanensis, but the leaves somewhat larger, espe- 
cially broader, and definitely longer than the few-flowered inflores- 
cences; flowers 4-5 mm. long; filaments glabrous; anthers subrec- 
tangular. Neg. 3688. 

Huanuco: Punta de Esperanza, 3,000 meters, Sawada 60 (det. 
Schmidt). Colombia. 

Ocotea minarum Mart, ex Mez, Jahrb. Bot. Gart. Berlin 5: 
305. 1889. 

A tree of medium size (about 10 meters), with somewhat angled, 
minutely tomentose branchlets and thin, lanceolate or elliptic leaves, 
acute at each end, mostly 7-8 cm. long and 2.5-3.5 cm. wide, very 



904 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

minutely and sparsely pubescent beneath; inflorescence about equal- 
ing the leaves, slightly puberulent; pedicels 1-3 mm. long; flowers 
rarely perfect, pilose or tomentulose; filaments glabrous or nearly so, 
very short, the anthers broadly rectangular, minutely emarginate; 
glabrous ovary and style subequal. Description of flowers from 
Mez, otherwise from Weberbauer specimens referred here by Mez, 
which, Schmidt has noted, have perfect (5014) or monoecious (4.987) 
flowers. Flowers yellowish or greenish (Weberbauer). 

Cuzco: Santa Ana, 1,300 meters, Weberbauer 5014- Lucumayo, 
2,100 meters, Weberbauer 4987; 280. Cajamarca: Tambillo (Jelski 
202, 207, 209). Brazil. 

Ocotea minutiflora 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10: 231. 1928. 

Branches glabrous or somewhat pilose at first; petioles slender, 
7-10 mm. long; blades obovate, narrowed to the base, somewhat 
acuminate, 5.5-8.5 cm. long, 2.5-3.5 cm. wide, glabrous, papyra- 
ceous, brownish yellow beneath; flowers perfect, pilose within, on 
slender pedicels 1.5-2 mm. long; perianth lobes subacuminate, about 
equaling the tube; anthers (series 1) subquadrate, their very short 
filaments pilose, those of series 3 sublanceolate, broadly rounded at 
the apex; ovary ovoid, glabrous, 1 mm. long, the stigma subsessile. 
A tree with yellow flowers, about 7 meters high, the trunk 18 cm. 
in diameter (Tessmann). Neg. 3690. 

Loreto: Iquitos, 100 meters, Tessmann 5366, type; Williams 
3667 (det. Schmidt). 

Ocotea monzonensis Mez, Repert. Nov. Sp. 3: 71. 1906. 

A shrub of 3 meters, with terete, ferruginous-tomentose branches 
and rather rigid, glabrate, elliptic leaves acute at each end; petioles 
1 cm. long; blades about 8 cm. long and 4 cm. wide; pedicels slender, 
squarrose-spreading, often 4-5 mm. long; flowers 2-2.5 mm. long; 
filaments pilose, the anthers subrectangular; ovary glabrous, twice 
as long as the thick style. Flowers greenish yellow. Neg. 3692. 

Huanuco: Monzon, 2,000 meters, Weberbauer 3712, type; 255. 

Ocotea multiglandulosa (R. & P.) Mez, Jahrb. Bot. Gart. 
Berlin 5: 280. 1889. Laurus multiglandulosa R. & P. Fl. Peruv. 4: pi. 
365. 1802; Laurogr. pi. 22. Oreodaphne Poeppigiana Nees, Syst. Laur. 
404. 1836. Ocotea Poeppigiana Macbr. Field Mus. Bot. 11: 18. 1931. 

A tree of 20 meters, the branches slender, slightly angulate, 
obscurely puberulent; petioles 1-2 cm. long; blades glabrate, sub- 



FLORA OF PERU 905 

coriaceous, obovate, acute at the base, short-acute at the apex, 
mostly about 12 cm. long and 5 cm. wide, the reticulation very fine; 
inflorescence lax, shorter than the leaves; flowers green, tomentulose, 
on pedicels 1-2 mm. long; perianth tube none or obsolete; filaments 
glabrous, twice as long as the subovate-quadrate, emarginate anthers; 
pilose ovary and style subequal. Wood soft, yellow (Poeppig}. Neg. 
27620. 

Loreto: Yurimaguas, Poeppig 2435. Chicoplaya, Ruiz & Pawn, 
type. 

Ocotea niunacensis 0. C. Schmidt, Repert. Sp. Nov. 31: 185. 
1933. 

A glabrate shrub, the young parts, like the green flowers, slightly 
puberulent; petioles 6-8 (-10) mm. long; blades usually elliptic, 
slightly narrowed to the base and to the short-acuminate apex, 7.5-10 
cm. long, 4-5.5 cm. wide, glabrous and lustrous above, and beneath 
except for the puberulent nerves and the barbellate axils of the nerves, 
thin-coriaceous, the lateral nerves 8-10, the reticulation evident only 
beneath; inflorescences axillary, racemose, 5-7-flowered, to 3.5 cm. 
long, the flowers often crowded, 7 mm. broad, the tube scarcely 
1 mm. long, the narrowly elliptic lobes 4 mm. long; anthers subquad- 
rate, 0.8 mm. long, the glabrous filaments barely half as long; basal 
glands of the stamens of series 3 very large; ovary globose, 1 mm. long, 
the style half longer. Similar to 0. monzonensis, with paniculate 
inflorescences. 

Huanuco: Between Chaclla and Muna, 2,500 meters, Weberbauer 
6794, type. 

Ocotea obovata (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 366. 
1889. Laurus obovata R. & P. Fl. Peruv. 4: pi. 357. 1802; Laurogr. 
pi. 27. Oreodaphne obovata Meissn. in DC. Prodr. 15, pt. 1 : 119. 1864. 

An essentially glabrous tree, 7-25 meters high, with obtusely 
angled branches and chartaceous, obscurely reticulate- veined, broadly 
obovate leaves, long-cuneate at the base, rounded at the apex, mostly 
15 cm. long and 8-9 cm. wide; inflorescence much shorter than the 
leaves, the widely spreading branches several centimeters long, 
slightly puberulent; flowers white; filaments sparsely pilose, the 
anthers ovate, rounded; glabrous ovary and style subequal. Neg. 
27621. 

Huanuco: Chinchao, Ruiz & Pavon, type. Without locality: 
Ruiz & Pavon 13-31. Venezuela. 



906 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ocotea olivacea A. C. Smith, Bull. Torrey Club 58: 105. 1931. 

A tree of 10-15 meters, with glabrous, sulcate-angled branchlets 
and glabrous, chartaceous, olivaceous leaves; petioles stout, 3-6 mm. 
long; blades oblong or oblong-elliptic, abruptly cuneate at the base, 
long-acuminate at the apex, 25-28 cm. long, 8.5-9.5 cm. wide; 
panicles 4-7 cm. long, the branches glabrous; flowers perfect, mi- 
nutely tomentose inside, the tube less than 1 mm. long; stamens all 
fertile, the anthers longer than the short, glabrous filaments; ovary 
and style subequal. Compared by the author with 0. minutiflora. 

Loreto: Iquitos, 100 meters, Killip & Smith 29843, type. 

Ocotea Ottoschmidtii Macbr. Candollea 5: 355. 1934. 0. 
Weberbaueri 0. C. Schmidt, Repert. Sp. Nov. 31: 181. 1933, non Mez. 

A shrub of 3 meters, glabrous except the leaf nerves and the 
flowers within; petioles 12-18 mm. long; blades ovate-elliptic, acute 
at the base, acuminate, 10-18 cm. long, 5-9 cm. wide, dull above, 
lustrous beneath, papyraceous, the lateral nerves mostly 8, prominent 
on both sides; inflorescence 3-9 cm. long, narrowly paniculate; 
flowers many, perfect, greenish, nearly 3 mm. broad, the pedicels 
1-2 mm. long; perianth tube 1.5 mm. long, the ovate segments 
shorter; anthers subsessile, subquadrate; ovary obo void-globose, 
1.5 mm. long, the style 0.2-0.3 mm. long. Compared by the author 
with 0. pretiosa (Nees) Mez, of southern Brazil, but in some respects 
similar to 0. piurensis Mez. 

Libertad: Valley of Rio Mixiollo, 1,500 meters, Weberbauer 7053, 
type. 

Ocotea otuzcensis 0. C. Schmidt, Repert. Sp.Nov.31: 183. 1933. 

A tree of 8 meters, the younger parts, except the tomentulose 
inflorescence and yellowish flowers, more or less pilose, the leaves 
glabrous in age; petioles 7-10 mm. long, to 2.5 mm. thick; blades 
broadly elliptic, rounded at the base and sometimes slightly decur- 
rent, short-acuminate, 9-14 cm. long, 5-8 cm. wide, lustrous above, 
coriaceous, the costa and the 7-10 lateral nerves more or less im- 
pressed above but prominent beneath; inflorescence to 15 cm. long, 
the flowers crowded at the tips of the branches; pedicels 2 mm. 
long; perianth tube 3 mm. long, pilose; stamens 1.6 mm. long, the 
subquadrate anthers contracted to the sparsely pilose filament, this 
0.6 mm. long; fruit subglobose, 2.5 cm. long, the perianth tube 
becoming much enlarged. 

Libertad: Prov. Otuzco, above Suchabamba, 2,700 meters, 
Weberbauer 6998, type. 



FLORA OF PERU 907 

Ocotea ovalifolia (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 5: 261. 
1889. Laurus ovalifolia R. & P. Fl. Peruv. 4 : pi. 346. 1802. Oreodaphne 
sublanuginosa Nees, Linnaea 21: 515. 1848. Ocotea sublanuginosa 
Macbr. Field Mus. Bot. 11: 19. 1931. 

A tree, related to 0. cuneifolia but in general characters similar 
to 0. macropoda; blades elliptic, merely acute at each end, rather 
thin, conspicuously ferruginous-pilose beneath, especially on the 
nerves, often 20 cm. long and half as wide; inflorescence narrow, 
15 cm. long, the rachis evanescently but densely ferruginous-villous, 
the flowers 3 mm. long, puberulent; pedicels 1.5 mm. long or less, 
anthers subtrapeziform. Neg. 27623. 

Huanuco: Chinchao, Ruiz & Pav6n, type. 

Ocotea petalanthera (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 
5: 358. 1889. Oreodaphne petalanthera Meissn. in DC. Prodr. 15, 
pt. 1: 125. 1864. 

A shrub or tree, similar to 0. maynensis, but the oblong-elliptic, 
acuminate leaves sparsely pilose-strigose beneath on the nerves; 
inflorescence densely strigose, nearly half as long as the leaves; fila- 
ments densely pilose, the anthers ovate, obtuse. According to Kos- 
termans, the filaments are glabrous. Neg. 3702. 

Loreto: Yurimaguas, Poeppig 2426, type. Brazil; Guianas. 

Ocotea piurensis Mez, Repert. Sp. Nov. 16: 308. 1920. 

A sparsely leafy shrub, glabrous almost throughout; blades 
elliptic, coriaceous, about 10 cm. long and 4 cm. wide, obtuse at 
the base, broadly acuminate or rounded at the apex, closely and con- 
spicuously reticulate-veined beneath; inflorescence at first ferrugi- 
nous-tomentulose, fasciculate, terminal, exceeding or shorter than 
the leaves; pedicels elongate, as much as 5 mm. long; flowers dull 
yellow, perfect, 3 mm. long; filaments glabrous (those of series 3 
with 2 sessile glands), the slightly longer anthers rectangular, broadly 
truncate and somewhat emarginate; glabrous ovary and style 
subequal. 

Piura: Quiros Valley, Weberbauer 6348, 6352, 6428. 

Ocotea puberula (Nees & Mart.) Nees, Syst. Laur. 472. 1836; 
343. Strychnodaphne puberula Nees & Mart. Linnaea 8: 39. 1833. 

A more or less puberulent or essentially glabrous shrub or small 
tree with thin or subcoriaceous, elliptic-lanceolate, acuminate leaves, 
the blades mostly 15 cm. long and 4.5 cm. wide, acute at the base, 
reticulate-veined on both sides, longer than the many-flowered, 



908 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pyramidal inflorescences; petioles sometimes 3 cm. long; perianth 
tube none; flowers 2-2.5 mm. long, on pedicels 1-2 (-4) mm. long, 
white or greenish, glabrate; filaments somewhat pilose, the subovate 
anthers acutish or obtuse; ovary and style subequal, glabrous. 
Known in Brazil as "canella babosa" and "louro bacato." Neg. 
22085. 

Cuzco: Santa Ana, 1,300 meters, Weberbauer 5013 (det. Mez); 
280. Without locality: (Mathews 1220). Argentina and Paraguay 
to British Guiana. 

Ocotea Raimondii 0. C. Schmidt, Repert. Sp. Nov. 31: 184. 
1933. 

A tree, the green parts puberulent or short-pilose except the 
mature leaves above, the inflorescence and perfect flowers tomentu- 
lose; petioles 5-15 mm. long; blades lanceolate, elliptic or oblanceo- 
late, narrowed to the base and apex, or rounded at the apex and 
acuminate, the lateral nerves mostly 9-12, rather prominent beneath, 
like the fine reticulation; inflorescence axillary, racemose-paniculate, 
to 11.5 cm. long; flowers 3.5-4 mm. long, on pedicels 2-3 mm. long, 
the tube inconspicuous; anthers subrectangular, 0.4-0.5 mm. long, 
rounded at the base and apex, the sparsely pilose filaments 0.3-0.4 
mm. long; ovary subovate, 1 mm. long, the style but slightly longer. 

Cajamarca: Tambillo, Prov. Cutervo, Raimondi 6736, type. 

Ocotea rubrinervis Mez, Jahrb. Bot. Gart. Berlin 5: 351. 1889. 

A tree of 7 meters; branches soon glabrate, terete; leaves sub- 
coriaceous, broadly elliptic, subrotund or acute at the base, very 
shortly obtuse-acuminate, minutely and sparsely pilose beneath, 
densely foveolate above, mostly about 10 cm. long and 6-7 cm. wide, 
the nervation prominently reticulate beneath; inflorescence ferru- 
ginous-pilose-puberulent, subequaling or shorter than the leaves; 
perianth tube nearly obsolete; filaments glabrous, about as long as 
the ovate, sub truncate anthers; pistillate flowers unknown; pedicels 
and persistent perianth strongly accrescent in fruit, the fruit ellip- 
soid, 12 mm. long, 7.5 mm. in diameter. Neg. 3713. 

San Martin: Tarapoto, Spruce 4580, type; at 750 meters, Wil- 
liams 6102 (det. Schmidt). Panama. "Muena blanca." 

Ocotea Rusbyana Mez, Jahrb. Bot. Gart. Berlin 5: 303. 1889. 

Apparently very similar to 0. Dielsiana 0. C. Schmidt, but the 

leaves areolate-granulate above, slenderly reticulate-veined beneath, 



FLORA OF PERU 909 

the pedicels 1-3 mm. long, the filaments glabrous, the anthers ovate. 
A tree of 7-12 meters, with rusty-cinereous-tomentose, angulate 
branchlets. 

Huanuco: Chaupagrapata, Sawada 98 (det. Schmidt). Santa 
Cruz (Peru?; Pearce, type). Bolivia. 

Ocotea Schomburgkiana (Nees) Benth. & Hook. f. Gen. PI. 3: 
158. 1880; 337. Oreodaphne Schomburgkiana Nees, Linnaea 21: 269. 
1848. 

Branchlets minutely tomentulose at the apex, glabrate, terete; 
petioles to 7 mm. long; blades coriaceous or almost rigid, promi- 
nently reticulate-veined on both sides, glabrous, ovate or elliptic, 
rather acute at the base, definitely acuminate, about 8.5 cm. long 
and 4.5 cm. wide; inflorescence shorter than the leaves, pilose; pedi- 
cels to 1.5 mm. long; perianth tube very short, the segments ovate; 
filaments glabrous, the glands of series 3 globose; anthers equaling 
the filaments, rectangular, sub truncate; staminodia usually none; 
ovary glabrous; fruit ovoid, 5 mm. long, conspicuously exserted, the 
cup fleshy. It is more than probable that the Peruvian material 
represents a different species. Neg. 27627. 

Junin: La Merced, 1,200 meters, 5791, 5793 (det. Schmidt). 
Guiana. 

Ocotea subrutilans Mez, Repert. Nov. Sp. 3: 70. 1906. 

A shrub 3 meters high, glabrous even to the filaments, with 
fleshy, coriaceous, elliptic leaves, acute at the base and rounded at 
the apex, rarely as long as 12 cm. and as wide as 4 cm. ; inflorescence 
about as long as the leaves; flowers whitish, 1.5 mm. long, on pedi- 
cels to 2 mm. long; perianth tube obconic; anthers quadrate-orbicu- 
lar; filaments glabrous; ovary globose, the style very short. Neg. 
3726. 

Loreto: Moyobamba, 1,300 meters, Weberbauer 4757, type; 267. 

Ocotea tarapotana (Meissn.) Mez, Jahrb. Bot. Gart. Berlin 5: 
304. 1889. Oreodaphne tarapotana Meissn. in DC. Prodr. 15, pt. 1: 
129. 1864. 

A climbing shrub, 4-8 meters long, the trunk 2.5 cm. in diameter 
(Spruce), with very slender, glabrous or glabrate branchlets, the 
foliage nearly that of 0. caudata, but with the floral characters rather 
of 0. maynensis, except that the elliptic anthers are acute; fruits 
ellipsoid, 12 mm. long, 9 mm. thick. Flowers yellowish. Neg. 3727. 



910 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Tarapoto, Spruce 4226, type; Williams 5811, 5866, 
6250, 6914, 6279, 5391, 6224, 5604, 6229 (det. Schmidt). Loreto: 
La Victoria, Williams 2579 (det. Schmidt). Tira Doble, Williams 
905. "Muena aguaras." 

Ocotea tenella A. C. Smith, Phytologia 1: 119. 1935. 

A tree of 13 meters, the branchlets terete, at first densely cinere- 
ous-tomentulose, like the leaves beneath and the slender petioles, 
these 4-7 mm. long; blades glabrous and impressed-punctate above, 
chartaceous, elliptic-oblong, 6-9 cm. long, 3-5 cm. wide, acute at 
the base, shortly obtuse-acuminate, the lateral nerves 3 or 4, with 
the reticulate veins prominent beneath; inflorescences solitary, 
axillary, slender, 2-4 cm. long, 10-20-flowered, glabrous except for 
the sparsely tomentose rachis; pedicels 1 mm. long; flowers glabrous, 
1.5-2 mm. long, the short tube obconic, the oblong, obtuse lobes 0.8 
mm. long; outer anthers sessile, oblong-orbicular, obtuse or minutely 
apiculate, those of series 3 similar but with short filaments; ovary 
0.8 mm. long, the style very short. The leaves are soft in texture 
as well as softly pubescent beneath. 

Rio Acre: Rio Macauhan, Krukoff 5406, type. 

Ocotea Tessmannii O. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10: 233. 1928. 

A tree of 15 meters, the trunk 20 cm. in diameter, with sulcate 
branchlets and thin, glabrous leaves mostly 8-15 cm. long and 3.5-5 
(-6.5) cm. wide; blades gradually narrowed to a slender petiole 1-1.5 
cm. long, obtuse-acuminate; inflorescence paniculate, to 8.5 cm. long; 
flowers pale yellowish, on pedicels 2-3 mm. long, the tube obsolete, 
the narrowly elliptic lobes spreading or recurved; anthers subquad- 
rate or ovate-rectangular, slightly emarginate, shorter than the 
glabrous filaments, these 1.5 mm. long (series 1); ovary ovoid, 1.2 
mm. long, the thick style slightly longer. Neg. 3729. 

Loreto : Soledad, Tessmann 5264, type. Yurimaguas, 200 meters, 
Williams 4728 (det. Schmidt). San Martin: Tarapoto, 750 meters, 
Williams 6594 (det. Schmidt). "Muena." 

Ocotea Trianae Rusby, Bull. N. Y. Bot. Gard. 6: 506. 1910. 

Branchlets and leaves beneath finely and closely sericeous-pilose; 
blades oblong, mostly somewhat inequilateral, acuminate at each 
end, the base decurrent into the short, broad, sulcate petiole, lustrous 
and glabrous above, the 12-15 nerves impressed above, very promi- 



FLORA OF PERU 911 

nent and reticulate beneath, 7-15 cm. long, 3-5 cm. wide; panicles 
slightly longer than the leaves, arising from the upper axils, broad 
and lax, at least in fruit, but the pedicels none or thick; fruiting calyx 
5-7 mm. long, 6-7 mm. wide, the margin beset with minute, pilose 
tufts; fruit about one- third exserted. 

Junin: La Merced, Schunke 278 (vel aff.; det. Schmidt). 
Loreto: Caballo-cocha, Williams 2153. Iquitos, 100 meters, Williams 
8083, 7974- Bolivia; Colombia. "Muena blanca," "pampa muena." 

Ocotea ucayalensis 0. C. Schmidt, Notizbl. Bot. Gart. Berlin 
10: 234. 1928. 

A tree, the trunk 50 cm. in diameter; branches terete, sulcate, 
the younger ones angled, glabrous; blades 12-20 cm. long, 5.5-6.5 
cm. wide, narrowed at each end, acuminate, glabrous; inflorescence 
to 9.5 cm. long; flowers yellowish green, 4-4.5 mm. long; perianth 
tube very short; anthers subrectangular, rounded at the base and 
apex, the slightly longer filaments subglabrous; ovary glabrous. 
Related to 0. maynensis (Meissn.) Mez. Neg. 3731. 

Loreto: Mouth of Rio Utiquenea, middle Ucayali, Tessmann 
3291, type. 

Ocotea Weberbaueri Mez, Bull. Herb. Boiss. II. 5: 236. 1905. 

A low shrub of 2 meters, with glabrate branchlets, rigid, glabrous, 
narrowly elliptic, acuminate leaves about 7 cm. long and 2 cm. 
wide, and somewhat paniculate, tomentulose, inflorescences only 
2-3 cm. long; petioles 5 mm. long; flowers yellowish white, cernuous, 
2.5 mm. long, perfect, the pedicels less than 2 mm. long, the tube 
conspicuous; filaments pilose, much shorter than the elliptic-orbicular 
anthers; ovary glabrous, shorter than the style. Compared by the 
author with 0. Benthamiana, which see under 0. Jelskii. Neg. 3742. 

Puno: Tambo Cachicachi, 1,800 meters, Weberbauer 1309, 
type; 242. 

9. NECTANDRA Rottb. 

Exhibiting the variation in many essential characters recorded 
for Ocotea, and possibly only artificially separated by the difference 
in the position of the anther cells in series 1. Filaments rarely 
developed. Style usually shorter than the ovary. Cup simple- 
margined, the perianth lobes perhaps always deciduous. N. cras- 
sipes and N. punctata are not included in the following key, based 
on that of Mez. 



912 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers conspicuous, 10 (9) -15 mm. wide. 

Leaves glabrous or glabrate beneath, rarely sparsely barbellate 

in the axils of the nerves. 

Pedicels usually 3-5 mm. long; leaves usually long-acuminate. 
Inflorescences usually puberulent; flowers 10-11 mm. wide. 

N. globosa. 

Inflorescences glabrate; flowers 9 mm. wide N. lucida. 

Pedicels usually very short; leaves acute or short-acuminate. 
Leaves acuminate at the base; perianth tube obvious. 

N. Jelskii. 
Leaves merely acute at the base. 

Perianth tube none; inflorescence ferruginous- tomentose. 

N. furcata. 
Perianth tube obvious. 

Inflorescence glabrate N. maranonensis. 

Inflorescence puberulent N. maynensis. 

Leaves pubescent beneath, even in age, or at least distinctly 

barbate in the axils of the nerves. 
Leaf pubescence pale or confined to the nerve axils; perianth 

tube conspicuous. 
Pubescence pilose, mostly or entirely on the nerves or in the 

nerve axils, ferruginous N. acutifolia. 

Pubescence a minute puberulence on the whole leaf surface 

beneath, pale N. lineatifolia. 

Leaf pubescence ferruginous, evenly distributed; perianth 

tube short. 
Leaves more or less pilose on both sides, at least on the 

nerves above, rarely only puberulent beneath. 
Leaves rigid-coriaceous or nearly so, subobtuse at the base; 

pedicels obsolete N. Laurel. 

Leaves more or less flexible, acute at the base; pedicels 

obvious N. reticulata. 

Leaves glabrate above, merely puberulent-tomentulose be- 
neath N. longifolia. 

Flowers small, 3-7 (8) mm. wide. 
A. Leaves pubescent beneath (see N. cissiflora). 

Flowers dioecious; leaves rounded-obtuse at the base (see also 
N. longifolia). 



FLORA OF PERU 913 

Pubescence pale, pilose; blades about 10 cm. long. .N. dioica. 
Pubescence ferruginous, velutinous; blades larger. 

N. Arnottiana. 

Flowers perfect; leaves acute at the base, except in N. longifolia. 
Leaves oblong-lanceolate, mostly narrower than 4 cm. 

Flowers 3-4 mm. wide N. Pichurim. 

Flowers 6-8 mm. wide N. globosa. 

Leaves ovate-elliptic-lanceolate or obovate, mostly 5 cm. 
wide or wider. 

Pedicels 5-8 mm. long N. myriantha. 

Pedicels 1-4 mm. long. 

Leaves strigose or pilose-tomentulose beneath (see N. 
longifolia). 

Leaves obovate, short-acuminate N. Pearcei. 

Leaves not obovate. 

Leaves narrowly long-acuminate. 
Leaves about one- third as wide as long; flowers 

7.5 mm. wide or larger N. globosa. 

Leaves nearly half as wide as long; flowers 6 mm. 

wide N. comasensis. 

Leaves short-acuminate or acute. 

Flowers 3-4 mm. broad; filaments longer than 

the anthers N. Pichurim. 

Flowers 4-5 mm. broad; filaments and anthers 

subequal N. Herrerae. 

Leaves very minutely puberulent or sericeous-puberulent 

beneath. 
Flowers 3.5-4 mm. wide, the tube conspicuous. 

N. Matthewsii, N. Macbridei. 
Flowers 6-7 mm. wide. 

Branchlets strongly and acutely angled-sulcate; 

leaves sericeous beneath N. longifolia. 

Branchlets angled or subterete; mature leaves 

merely sericeous-puberulent beneath. 
Leaves not at all reticulate-veined above. 

N. lineatifolia. 
Leaves obviously reticulate-veined above. 



914 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves elegantly acuminate N. acutifolia. 

Leaves short-acute N. superba. 

A. Leaves glabrous beneath or essentially so. 
Flowers 6-8 mm. broad. 

Leaves mostly narrower than 2 cm N. microcarpa. 

Leaves much wider. 
Leaves rounded at the base (see N. purpurea). 

Petioles about 1.5 cm. long N. maranonensis. 

Petioles about 1 cm. long N. lucida. 

Leaves acute or acutish at the base. 

Leaves thin; pedicels 4-5 mm. long. . . N. breaparinensis. 
Leaves rather thick; pedicels mostly much shorter. 
Leaves to 40 cm. long; inflorescence ample, the flowers 
cymosely crowded on divaricate branchlets. 

N. Macbridei. 

Leaves often medium in size, mostly shorter than 

20 cm.; inflorescence often not ample, never with 

divaricate ultimate branchlets of crowded cymes. 

Leaves conspicuously reticulate-veined on both 

sides N. purpurea. 

Leaves very obscurely reticulate- veined, at least 

above. 
Inflorescence typically cinereous-puberulent. 

N. globosa. 
Inflorescence glabrate. 

Leaves obscurely immersed-reticulate-veined 

beneath N. maynensis. 

Leaves prominently reticulate- veined beneath. 

N. lucida. 

Flowers 3.5-5 mm. broad, except in N. Sawadae (5.5 mm.). 
Mature leaves thick, the lateral nerves beneath very promi- 
nent. 

Inflorescence cinereous- or yellowish-tomentose or puberu- 
lent. 

Leaves 6-9 cm. wide N. cissiflora. 

Leaves 3-5 cm. wide N. Pichurim. 

Inflorescence glabrate N. Raimondii. 



FLORA OF PERU 915 

Mature leaves thin-subcoriaceous, not conspicuously nerved 
beneath. 

Petioles mostly 1.5-2.5 cm. long. 
Pedicels 4-5 mm. long; leaves 4.5-8 cm. wide. 

N. capanahuensis. 

Pedicels mostly 3 mm. long or shorter; leaves often less 
than 5 cm. wide. 

Leaves very shortly acuminate N. intermedia. 

Leaves gradually long-acuminate N. Pichurim. 

Petioles mostly about 1 cm. long, often thick. 
Leaves scarcely one-third longer than broad. 

Inflorescence cinereous-tomentulose; leaves lustrous 

above N. latifolia. 

Inflorescence greenish, glabrate; leaves dull. 

N. viburnoides. 

Leaves about two-thirds longer than wide. 
Petioles conspicuously thickened and deeply sulcate 

above, with sharp edges N. yarinensis. 

Petioles normal to slender, not sulcate above. 
Perianth tube conspicuous. 

N. laevis, N. capanahuensis. 
Perianth tube obsolete. 

Inflorescence much shorter than the caudate- 
acuminate leaves N. citrifolia. 

Inflorescence subequaling the acuminate leaves. 

Leaves dull above, lanceolate, 2-3 cm. wide. 

N. Williamsii. 
Leaves lustrous above, elliptic, to 4.5 cm. wide. 

N. Sawadae. 

Nectandra acutifolia (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 
5: 409. 1889. N. acuminata Macbr. Field Mus. Bot. 11: 19. 1931. 
Laurus acutifolia R. & P. Laurogr. pi. 17. Persea acuminata Nees 
& Mart, ex Nees, Syst. Laur. 170. 1836. N. berchemiae/olia Meissn. 
in DC. Prodr. 15, pt. 1: 154. 1864. 

A tree of 6-12 meters; branchlets soon glabrate, but the blades, 
even in age, ferruginous-barbate, at least in the nerve axils beneath, 
often somewhat pilose, and above somewhat tomentulose; petioles 
sometimes 3.5 cm. long; blades coriaceous, often reddish on the 



916 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

strongly veined under surface, ovate-elliptic or ovate-lanceolate, 
acuminate at each end, 10-20 cm. long, 4-8 cm. wide; inflorescence 
sparsely pubescent, about as long as the leaves; pedicels 1-5 mm. 
long; flowers white, tomentulose; glands prominent; anthers sessile, 
subobtuse. Neg. 29396. 

Huanuco: Haenke. Without locality, Ruiz & Pavon. Monzon, 
1,500 meters, Weberbauer 3555, 3456 (det. Mez); 256, 284. Caja- 
marca: Tambillo, Raimondi (det. Schmidt). Loreto: Caballo- 
cocha, Williams 2088 (det. Schmidt). Bolivia to Colombia. "Mund- 
shuy," "muena amarilla." 

Nectandra Arnottiana Nees, Syst. Laur. 289. 1836; 402. 
Pleurothyrium chrysothyrsus Meissn. in DC. Prodr. 15, pt. 1 : 169. 1864. 

Apparently similar to N. dioica Mez, but the pubescence fer- 
ruginous-velutinous-tomentose, the inflorescence long-peduncled, 
many-flowered, little shorter than the large (30-40 cm. long, 15 
cm. wide), ovate-elliptic leaves, the perianth tube conspicuous, 
the filaments very broad, the glands of series 3 large. According 
to the collector, a large tree. Pistillate flowers unknown. 

Amazonas: Chachapoyas (Mathews 3031, type). 

Nectandra breaparinensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 227. 1928. 

Similar to N. capanahuensis, but the thin, elliptic leaves with 
fewer (5-8) lateral nerves, mostly 12-15 cm. long and 6-8 cm. 
wide, tomentulose. Neg. 3749. 

Loreto: Brea Parina, Tessmann 5522, type. "Muena." 

Nectandra capanahuensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 228. 1928. 

A tree with obovate-elliptic, more or less acuminate, dull leaves 
10-25 cm. long and 4.5-9 cm. wide, the lateral nerves 9-11; 
inflorescence to 17 cm. long, paniculate, somewhat pilose; flowers 
white, 5 mm. wide, densely strigose outside, densely pilose within, 
the tube short. Related to N. lucida Nees, but the flowers smaller 
and the inflorescence much larger. Neg. 4299. 

Loreto: Middle Rio Blanco, Tessmann 3058, type. Florida, 
King 2133. "Marogofuina." 

Nectandra cissiflora Nees, Syst. Laur. 296. 1836; 453. 
A tree of 8-24 meters, resembling N. intermedia Mez, but the 
leaves broader (15-20 cm. long, 6-9 cm. wide), sometimes sparsely 



FLORA OF PERU 917 

pilose beneath, the filaments very much shorter than the depressed- 
ellipsoid anthers, or the anthers subsessile; flowers white. Neg. 
3751. 

Huanuco: Cochero, Poeppig 3061, type. Cuzco: Santa Ana, 
1,300 meters, Weberbauer 5027 (det. Mez); 280. Chinche, 1,700 
meters, Weberbauer 4988 (det. Mez). Amazonian Brazil. 

Nectandra citrifolia Mez & Rusby, Mem. Torrey Club 6: 
115. 1896. 

A shrub or tree, glabrous except the somewhat puberulent- 
tomentose young branchlets and flowers, the flowers perfect, scarcely 
5 mm. wide; blades glaucous green above, brownish beneath, finely 
and faintly reticulate-veined, the lateral nerves rather prominent 
beneath, the blades chartaceous, 9-12 cm. long, 4-5 cm. wide, 
caudate-acuminate; inflorescence subracemose, few-flowered, the 
pedicels 2-5 mm. long; perianth tube broadly obconic, short, the 
segments elliptic; outer filaments none, the anthers reniform, rotund 
at the apex, papillose; staminodia minute. The Rio Acre locality 
is near the Peruvian boundary, according to the map of the Geo- 
graphic Society of Peru. 

Rio Acre: Rio Macauhan, Krukoff 5467, 5244. Bolivia. 

Nectandra comasensis 0. C. Schmidt, Repert. Sp. Nov. 31: 
185. 1933. 

A tree with terete branches, the tips tomentulose; petioles 12-15 
mm. long; blades mostly ovate-elliptic, the rounded base narrowed 
to the petiole, the acuminate apex to 1.5 cm. long, coriaceous, 
soon glabrous above, densely pilose beneath, the lateral nerves 
11-12, prominent beneath; inflorescence axillary, 11-13 cm. long, 
many-flowered; perfect flowers densely crowded at the tips of the 
branches, white, 6 mm. wide, short-pilose, like the pedicels, these 
2-3 mm. long; perianth tube scarcely 1 mm. long; outer anthers 
sessile, the inner longer than the pilose filaments; ovary elliptic, 
glabrous, 1 mm. long, equaled by the style. Related to N. reticulata 
(R. & P.) Mez, with larger flowers and leaves (Schmidt). 

Junin: Rio de Comas, 2,500 meters, Weberbauer 6616. 

Nectandra crassipes Meissn. in DC. Prodr. 15, pt. 1: 150. 
1864; 423. 

Known only in fruit, but presumably related to N. longifolia, 
from which its obtuse-based leaves, subsericeous beneath, seem 



918 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to distinguish it; fruit globose, 7 mm. in diameter. If there are 
further distinctive characters, they must be in the flowers; included 
by Ruiz and Pavon in their Laurus longifolia ( Nectandra longifolia) . 
Peru: Ruiz & Pav6n, type. 

Nectandra dioica Mez, Repert. Sp. Nov. 16: 308. 1920. 

A tall tree, as much as 40 meters high, the young branchlets, 
petioles, leaves beneath, and inflorescences rather densely whitish 
or yellowish pilose- tomentulose, the hairs appressed; petioles over 
2 cm. long; blades coriaceous, finally glabrous and dull above, 
elliptic, short-acute or rounded at the base, short-acuminate at the 
apex, conspicuously reticulate-veined on both sides; inflorescence 
thyrsoid-paniculate, dense, much shorter than the leaves; flowers 
dioecious, 3 mm. long, the pedicels scarcely 2 mm. long; bractlets 
deciduous; perianth tube short; anthers sessile, suborbicular, the 
glands of series 3 minute. Kostermans thinks this may be the 
same as N. grandis (Mez) Kosterm. Neg. 3753. 

Rio Acre: Vie 9402, type. Amazonian Brazil. 

Nectandra elongata Mez, Jahrb. Bot. Gart. Berlin 5: 444. 1889. 

A shrub or tree; blades long-acuminate, short-acute at the 
base, about 20 cm. long, little more than 5 cm. wide, coriaceous, 
puberulent beneath, strongly reticulate- veined on both sides; 
petioles to 2.5 cm. long; inflorescence broad and lax, with divaricate 
branches, about equaling the leaves; flowers 6-8 mm. wide, strigose- 
tomentose, sessile or the pedicels 2 mm. long, the tube short; fila- 
ments glabrous, the outer ones 2-3 times shorter than the obscurely 
tricuspidate anthers; ovary and style glabrous, equal in length. 
Neg. 31209. 

Huanuco: Cochero, Poeppig, type. 

Nectandra furcata (R. & P.) Nees, Linnaea 21: 501. 1848; 430. 
Laurus furcata R. & P. Fl. Peruv. 4: pi. 367. 1802; Laurogr. pi. 8. 

A shrub or tree, glabrous or essentially so except for the fer- 
ruginous- tomentulose inflorescence; petioles rarely 12 mm. long; 
blades ovate-lanceolate or elliptic-lanceolate, 10-15 cm. long and 
3-6 cm. wide, or larger, only the lateral nerves at all prominent 
beneath; inflorescence somewhat squarrosely paniculate, about 
equaling the leaves; flowers white, on pedicels 1-2 mm. long; anthers 
(outer) sessile; stigma subsessile, or the style obvious. The Weber- 
bauer material was referred by Mez to N. magnoliifolia Meissn. 
Neg. 29397. 



FLORA OF PERU 919 

Huanuco: Cochero, Chinchao, etc., Vitoc, Ruiz & Pavon, type. 
Junin: Huacapistana, 1,700 meters, a shrub of 5 meters, Weber- 
bauer 1982; 246. Cuzco: Urubamba Valley, 1,800 meters, a tree 
of 15 meters, Weberbauer 5059; 245. 

Nectandra globosa (Aubl.) Mez, Jahrb. Bot. Gart. Berlin 5: 
415. 1889. Laurus globosa Aubl. PI. Guian. 1: 364. 1775. N. pul- 
verulenta Nees, Syst. Laur. 283. 1836. N. Tessmannii 0. C. Schmidt, 
Notizbl. Bot. Gart. Berlin 10: 229. 1928. 

Often a beautiful tree, 25 meters high; branchlets more or less 
puberulent-tomentulose; blades ovate-lanceolate, acute at the base, 
mostly 15-20 cm. long and 5-8 cm. wide, or the lower twice as 
large, subcoriaceous, finely reticulate- veined beneath; petioles to 
2 cm. long; inflorescence shorter than the leaves, more or less pubes- 
cent; pedicels 1-5 mm. long; flowers white, fragrant, 10-11 mm. 
wide; perianth tube conspicuous; anthers sessile; ovary and style 
subequal or very unequal (N. pulverulenta). The specimens cited 
appear specifically unseparable unless by proportionate differences 
in length of the style. See also N. lucida. 0. C. Schmidt, op. cit. 
230, gives a table of comparative measurements for N. lucida, N. 
capanahuensis, N. Tessmannii, and N. breaparinensis that may be 
found, when more collections are available, to illustrate the range 
of variation of a single or at most two somewhat variable species. 

Cajamarca: Cutervo (Jelski 174, 179, 181, 188, 201, 204). 
Huanuco: Cochero (Poeppig 1427, type of N. pulverulenta) . Mon- 
zon, 600 meters, Weberbauer 3608 (det. Mez); 285. Junin: La 
Merced, 570.4. Chanchamayo, Schunke 5, 419 (N. pulverulenta, 
according to Schmidt); at 1,000 meters, Weberbauer 1900 (N. 
pulverulenta, according to Mez); 282; 570.4 (det. Schmidt). San 
Martin: Juanjui, Klug 3795. Moyobamba, 800 meters, Weberbauer 
4574 (det. Mez); 291. Loreto: Middle Rio Blanco, Tessmann 3070 
(det. Schmidt). Lower Rio Maranon, Tessmann 3758 (det. Schmidt). 
Soledad, Tessmann 5289. Lower Rio Huallaga, Williams 3814, 
4015, 4913. Caballo-cocha, Williams 2268 (det. Schmidt). Pam- 
payacu, Klug 3166. La Victoria, Williams 2863. Leticia, Williams 
3045. Florida, Klug 2264- Pumayacu, Klug 3227. Mouth of 
Rio Santiago, Tessmann 3942 (type of N. Tessmannii). Brazil 
to Colombia and Central America. "Muena," "muena amarilla," 
"muena blanca," "nomebe fuina." 

Nectandra Herrerae 0. C. Schmidt, Repert. Sp. Nov. 31: 
188. 1933. 



920 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branches velvety-tomentose, the leaves beneath, pedicels, and 
flowers yellowish-pilose, the flowers perfect, 4-5 mm. broad; petioles 
7-12 mm. long; blades lance-elliptic, narrowed and acute at the 
base, 10-20 cm. long, 3-6.5 cm. wide, papyraceous, the costa per- 
manently pubescent above, the lateral nerves 7-9, the prominent 
reticulation fine; inflorescence 8-13 cm. long, the peduncle 2-3.5 cm. 
long, many-flowered, the pedicels 1-1.5 mm. long; perianth tube 
scarcely 2 mm. long; outer anthers mostly broadly transverse- 
elliptic, 0.4 mm. long, equaling the glabrous filaments; ovary globose, 
glabrous, slightly exceeded by the style. Here might be sought 
N. japurensis Nees, 440, from adjacent Brazil, with smoother, 
wide leaves, ferruginous-pubescent inflorescence, and subquadrate 
anthers; also N. viburnoides Meissn., 453, with sparsely pilose, 
roundish-elliptic leaves and pedicels 2-4 mm. long. 

Cuzco: Valle del Urubamba, 2,050 meters, Hen era 2662, type; 
2669, 2673. 

Nectandra intermedia Mez, Repert. Sp. Nov. 16: 308. 1920. 

Branchlets subangulate, glabrate; petioles to 2 cm. long, slender; 
blades coriaceous, glabrous, dull, oblong-elliptic, short-acuminate, 
about 15 cm. long and 4.5 cm. wide; inflorescence about equaling 
the leaves, lax, cinereous-pilose; flowers greenish yellow, 5 mm. 
broad, on pedicels 2 mm. long, the bractlets deciduous; perianth 
tube short; filaments glabrous, slightly shorter than the anthers; 
glands (series 3) large; anthers all sublaterally dehiscent, obtuse. 
Very near N. Pichurim (HBK.) Mez. Neg. 3756. 

Rio Acre: Ule 9408, type. 

Nectandra Jelskii Mez, Jahrb. Bot. Gart. Berlin 5: 429. 1889. 

Very similar to N. furcata, but the rather smaller, elliptic leaves 
on somewhat longer petioles that are wing-margined above by the 
decurrent-acuminate leaf base. 

Cajamarca: Tambillo, Jelski 172, type; Raimondi. 

Nectandra laevis Mez, Jahrb. Bot. Gart. Berlin 5: 451. 1889. 

A shrub or tree, with the general characters of N. yarinensis 
O. C. Schmidt, but the petioles slender, the inflorescence often 
longer than the leaves, and the perianth tube prominent. Probably 
a form or variety of N. Pichurim (HBK.) Mez. A tree 25 meters 
high (Krukoff). 



FLORA OF PERU 921 

Cajamarca: Tambillo, Jelski 186, type. Junin: Cahuapanas, 
340 meters, Kittip & Smith 26744- Rio Acre: Rio Macauhan, 
Krukoff 5339. Bolivia. 

Nectandra latifolia (HBK.) Mez, Jahrb. Bot. Gart. Berlin 
5: 454. 1889. Ocotea latifolia HBK. Nov. Gen. & Sp. 2: 165. 1817. 

Usually readily recognizable among the small-flowered, glabrous 
species by the broadly elliptic (10-20 cm. long, 5-8 cm. wide) leaves, 
conspicuously lustrous above, and, to a less extent, by the reticulate- 
veined under surface; petioles 1 cm. long; flowers 4 mm. wide, 
on pedicels 1-4 mm. long; perianth tube obsolete; filaments glabrous 
or nearly so, subequaling the depressed-orbicular anthers; style 
obvious. A tree of 10-12 meters, with a trunk diameter of 15-25 
cm.; flowers yellowish. 

Loreto: Mouth of Rio Santiago, Tessmann 4169, 4243, 4475 
(det. Schmidt). Rio Acre: Ule 9397 (det. Mez). Brazil to northern 
South America. "Muena." 

Nectandra Laurel Klotzsch ex Nees, Linnaea 21: 505. 1848; 403. 

A shrub or small tree with conspicuously ferruginous-pubescent 
foliage, and villous inflorescences of white, honey-scented flowers; 
petioles to 2.5 cm. long; blades ovate-elliptic, narrow-acuminate, 
laxly reticulate-veined beneath, about 10 (-30) cm. long and 4-8 
cm. wide, well exceeding the short inflorescences; flowers 13-15 
mm. wide, anthers sessile, truncate, the connective triangularly 
acuminate-produced; style and ovary subequal. Neg. 27602. 

Cajamarca: Tambillo, Raimondi (det. Schmidt); Jelski 189. 
Chugur, 2,200 meters, Weberbauer 4150 (det. Mez). Bolivia to 
Venezuela and Panama. "Roble," "mundshuy." 

Nectandra lineatifolia (R. & P.) Mez, Jahrb. Bot. Gart. 
Berlin 5: 424. 1889. Laurus lineatifolia R. & P. Fl. Peruv. 4: pi. 371. 
1802; Laurogr. pi. 26. 

The more or less angled branchlets, inflorescence, and leaves 
beneath pale ferruginous with a minute puberulence, this not con- 
cealing the leaf surface; petioles 1.5-2 cm. long; blades elliptic, 
about 15 cm. long, 6-10 cm. wide, acuminate, chartaceous-cori- 
aceous, the veins obvious only beneath and there minutely pulveru- 
lent; flowers 6-7 mm. wide, numerous; perianth tube well developed; 
anthers sessile, acute or acuminate; glands of series 3 conspicuous; 
style short. Neg. 27603. 



922 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Moyobamba, Mathews. Huanuco: Chinchao, Ruiz 
& Pavdn. Ecuador. 

Nectandra longifolia (R. & P.) Nees, Linnaea 21: 502. 1848; 
423. Laurus longifolia R. & P. Fl. Peruv. 4: pi. 370. 1802; Laurogr. 
pi. 24. 

A shrub or tree with almost sulcately acute-angled branchlets; 
petioles to 3.5 cm. long; blades coriaceous, densely sericeous beneath, 
even in age, with a pale yellowish pubescence, broadly elliptic- 
lanceolate, acuminate, rounded or subacute at the base, 20 cm. 
long, or longer, 6-9.5 cm. wide, thick-coriaceous, obscurely reticu- 
late-veined beneath; flowers 12-14 mm. wide, on pedicels 3-6 mm. 
long; anthers sessile, acute or subobtuse; glands of series 3 small; 
cup about 7 mm. wide, 4 mm. deep, the fruit half exserted. The 
Madrid material is in fruit only. Neg. 29398. 

Peru: Without locality, Ruiz & Pavon, type. 

Nectandra lucida Nees, Syst. Laur. 334. 1836; 429. 

A tree or shrub, the branchlets glabrate; petioles 1.5 cm. long; 
blades coriaceous, glabrous, lustrous above, above scarcely at all 
but beneath prominently and densely reticulate- veined, elliptic or 
obovate-elliptic, acute or subobtuse at the base, acuminate at the 
apex, 15-20 cm. long, 5-7 cm. wide; inflorescence short, subthyrsoid, 
glabrate, much shorter than the leaves; flowers perfect, minutely 
tomentulose, 8-9 mm. broad; perianth tube conspicuous; anthers 
sessile, rotund; ovary glabrous, the style slightly shorter. Doubt- 
fully distinct from N. globosa. Neg. 27604. 

Loreto: Yurimaguas, 135 meters, Killip & Smith 27709 (det. 
Schmidt). Rio Acre: Rio Macauhan, Krukoff 5257, 5760. 

Nectandra Macbridei 0. C. Schmidt ex Macbr. Field Mus. 
Bot. 8: 81. 1930. 

Branchlets tomentulose; petioles thick, about 2 cm. long; blades 
narrowly elliptic or lanceolate, narrowed to each end, acuminate, 
6-16 cm. wide, lustrous above, prominently nerved and dull beneath; 
inflorescence ample, many-flowered, the flowers crowded in cymes, 
tomentulose; perianth tube short; filaments very short; ovary and 
style subequal. A small tree, about 7 meters high, with few branches. 
Compared by the author with N. pulverulenta Nees (N. globosa 
in a broad sense), but with very differently developed and much 
more abundantly flowered inflorescences, and with flowers about a 
third smaller. 

Huanuco: Rio Huallaga Canyon, 1,200 meters, 4228, type. 



FLORA OF PERU 923 

Nectandra maranonensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 229. 1928. 

A glabrous tree, 10 meters high, with thick, ovate-elliptic, sharply 
acuminate blades lustrous on both sides, about 10 cm. long and half 
as wide; petioles 1.5 cm. long; inflorescence to 7.5 cm. long, very 
shortly pilose in the axils; flowers white, 6 mm. wide, on slender pedi- 
cels 2 mm. long; perianth tube obsolete; style half as long (0.5 mm.) 
as the ovary. Neg. 3760. 

Loreto: Mouth of Rio Santiago, Tessmann 4833, type. Caballo- 
cocha, Williams 2447 (det. Schmidt). La Victoria, Williams 2858 
(det. Schmidt). "Hihuha," "tihuha," "muena amarilla." 

Nectandra Matthewsii Meissn. in DC. Prodr. 15, pt. 1: 159. 

1864; 448. 

A tree with unusually small flowers and papyraceous or subchar- 
taceous, lanceolate leaves that are very closely and finely puberulent 
beneath or at first cinereous- tomentulose; blades oblong-lanceolate, 
acute at the base, acutely acuminate, 10-25 cm. long, 4-5.5 cm. wide, 
slightly reticulate- veined beneath; petioles to 1.5 cm. long; inflores- 
cence cinereous-tomentulose, narrow, the widely divaricate branches 
shorter than the leaves; flowers tomentulose; glabrous filaments and 
suborbicular anthers subequal, the glands of series 3 large. See also 
N. japurensis Nees, under N. Herrerae, with glabrous flowers 5 mm. 
wide. Neg. 27605. 

Peru: Without data, Mathews 1431, type. 

Nectandra maynensis Mez, Jahrb. Bot. Gart. Berlin 5: 428. 
1889. N. amazonum (Mart.) Nees, var. glabrata Meissn. in DC. 
Prodr. 15, pt. 1: 150. 1864. 

A tall tree, 25 meters high, with glabrous foliage and branchlets, 
only the short inflorescence very minutely puberulent; petioles slen- 
der, mostly 2 cm. long; blades ovate-elliptic, subacute at the base, 
long-acuminate, mostly 10 cm. long and 4-5 cm. wide (7-15 cm. 
long, 3-5 cm. wide); pedicels 2-4 mm. long; flowers 7 mm. wide; 
anthers sessile, subacute; perianth tube conspicuous; glands of series 
3 small; stigma subsessile. Wood yellowish, soft (Poeppig). Appar- 
ently N. leucantha Nees, var. peruviana Meissn. op. cit. 151 is the 
same (Laurus Hihuha R. & P. Laurogr. pi. 19}. It is known only 
from "Guayaquil," Ruiz, and from Brazil. Neg. 3761. 

Loreto: Yurimaguas, Poeppig 2343, type. "Tihua." 



924 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Nectandra microcarpa Meissn. in DC. Prodr. 15, pt. 1: 162. 
1864; 441. 

The very narrow, linear-lanceolate, densely reticulate-veined 
leaves, 8-15 cm. long, readily distinguish this species, which is 
related to N. elongata Mez; inflorescence glabrous, subrectangular; 
truncate anthers and glabrous filaments subequal. A tree 6-9 meters 
high, with many spreading branches. Negs. 22068, 3777. 

San Martin : Tarapoto, Spruce 41 91 , type. Chazuta, Rio Huallaga, 
Klug 4054, 4012; a tree of 4 meters; flowers white and yellow. 

Nectandra myriantha Meissn. in DC. Prodr. 15, pt. 1: 163. 
1864; 452. 

A tree of 30 meters; blades coriaceous, the lateral nerves promi- 
nent beneath, but the veins scarcely if at all reticulate, 15-20 
cm. long, 7-8 cm. wide, short-acuminate, acute at the base; inflores- 
cence open, about equaling the leaves; flowers subglabrous or pilose, 
yellowish, 4-6 mm. wide, the tube obsolete; filaments shorter than 
the depressed-oval, subtomentose-papillose anthers; glands of series 
3 large. The wood is unpleasantly scented. Mature leaves of the 
Ule specimen are pilose beneath, suggesting that A 7 . Pearcei Mez, 
which see, may not be specifically distinct. Negs. 22069, 19267, 
22075. 

Rio Acre: Ule 9401 (det. Mez). Brazil. 

Nectandra Pearcei Mez, Jahrb. Bot. Gart. Berlin 5: 444. 1889. 

Apparently very similar to N. myriantha Meissn., the blades 
more or less obovate, about 15 cm. long and 6 cm. wide, minutely 
sericeous above, the nerves sulcate-immersed, strigose beneath, the 
lateral nerves prominent; inflorescence, including the flowers, tomen- 
tulose, the flowers 6-7 mm. wide; anthers of series 1 and 2 subsessile, 
depressed-suborbicular, broader than long, papillose. The type 
locality is unknown to me, but is given as Peruvian by Mez. 

Peru(?): Chailla (Pearce, type). 

Nectandra Pichurim (HBK.) Mez, Jahrb. Bot. Gart. Berlin 
5: 449. 1889. Ocotea Pichurim HBK. Nov. Gen. & Sp. 2: 166. 1817. 

A white-barked tree of 10-20 meters, usually well marked by 
the lanceolate, narrowly long-acuminate (typically) leaves, often 
only 3 cm. wide and 10 cm. long; blades somewhat sericeous beneath, 
or in var. cuprea Mez (op. cit. 450) densely so and copper-colored, 
rather thin and faintly reticulate-veined; inflorescence ample, open, 
somewhat shorter than the leaves; flowers 3-4 mm. wide, on pedicels 



FLORA OF PERU 925 

1-4 mm. long; depressed, suborbicular anthers much shorter than 
the filaments; glands of series 3 large. According to Kostermans, 
Fl. Suriname 2: 290. 1936, the HBK. tree is different and, if so, 
apparently the correct name for the following material is N. cuspi- 
data Nees, Syst. Laur. 330. 1836. The native Loreto name, "isula 
micuna," refers to the large ants that live on the tree. The bark is 
used as a poultice for the bites of ants (Tessmann). Neg. 35014. 

Junin: Chanchamayo Valley, Schunke 4-98, 284. (det. Schmidt). 
Rio Pichis, Killip & Smith 26697 (det. Schmidt). San Martin: San 
Roque, Williams 7730 (det. Schmidt). Moyobamba, 800 meters, 
flowers yellowish, Weberbauer 4501 (det. Mez); 284, 290. Huanuco: 
Monzon, 1,500 meters, a tree of 8 meters; flowers yellowish, Weber- 
bauer 3455 (var. cuprea; det. Mez). Pampayacu, Sawada 8 (det. 
Schmidt). Loreto: Pebas, Williams 1705. Balsapuerto, Killip & 
Smith 29007. Iquitos, Tessmann 3588; flowers greenish white. 
Rio Acre: Ule 9396 (det. Mez). Argentina and Brazil to Panama. 
"Isula micuna," "pishcu nahu muena." 

Nectandra punctata (R. & P.) Klotzsch ex Nees, Linnaea 21: 
509. 1848. Laurus punctata R. & P. Fl. Peruv. 4: pi. 347. 1802. 
Gymnobalanus punctata Meissn. in DC. Prodr. 15, pt. 1: 141. 1864. 
G. punctata var. elongata Meissn. loc. cit. N. punctata var. elongata 
Meissn. ex Mez, Jahrb. Bot. Gart. Berlin 5: 463. 1889. 

Known only from foliage, which is quite lacking in distinctive 
characters, and from the fruit, this name could well be discarded. 
It is based on a Ruiz specimen from Mufia, and the pubescent variety 
elongata on a Pavon specimen from Vitoc. Neg. 27608. 

Nectandra purpurea (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 
5: 443. 1889. Laurus purpurea R. & P. Fl. Peruv. 4: pi. 351. 1802; 
Laurogr. pi. 7. 

A shrub or tree, rather similar in foliage to N. latifolia (HBK.) 
Mez, but the flowers said to be nearly twice as broad, the glabrous 
filaments and truncate, depressed-oval or subquadrate anthers very 
unequal in length. Only a leaf specimen seen by me. 

Huanuco: Cochero and Chinchao, Ruiz, type. 

Nectandra Raimondii 0. C. Schmidt, Notizbl. Bot. Gart. Ber- 
lin 10: 726. 1929. 

A glabrous tree, only the axillary inflorescences sparsely tomentu- 
lose; petioles 5-15 mm. long; blades mostly elliptic, 11-24 cm. long, 
4.5-8.5 cm. wide, narrowed at each end, the acumination to 3.2 cm. 



926 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, the lateral nerves 5-6; flowers perfect, 3.5 mm. broad, on slen- 
der pedicels 2 (-3) mm. long; perianth lobes (tube very short) 1.5-1.8 
mm. long; fertile stamens 9, the outer ones leafy-dilated, sessile, 
0.5-0.6 mm. long; ovary ellipsoid, glabrous, 0.8-1 mm. long, con- 
tracted to a slightly shorter style. Related to N. capanahuensis 
0. C. Schmidt (group of N. lucida Nees). Neg. 3764. 

Cajamarca: Tambillo, Raimondi 3524; type. Huanuco: Chin- 
chao, Sawada 91 (det. Schmidt). "Mundshuy gateado." 

Nectandra reticulata (R. & P.) Mez, Jahrb. Bot. Gart. Berlin 
5: 404. 1889. Laurus reticulata R. & P. Fl. Peruv. 4: pi. 348. 1802; 
Laurogr. pi. 3. N. rigida Nees, Syst. Laur. 284. 1836. Ocotea rigida 
HBK. Nov. Gen. & Sp. 2: 163. 1817. 

Very similar to N. Laurel Klotzsch & Karst., but the leaves 
usually rather thinner and narrowed or acute at the base, the well 
developed inflorescence nearly as long; outer anthers foliaceous, 
acute. A tree as much as 40 meters high. N. rigida is a form or 
variety with the style shorter than the ovary. 

San Martin: Rio Cainarachi, 230 meters, Klug 2696. Caja- 
marca: Cutervo, Raimondi (det. Schmidt). Huanuco: Cochero 
and Chinchao, Vitoc, Ruiz & Pawn, type. Cochero, Poeppig 1233. 
Puyash, Sawada 93. Yanano, 1,800 meters, 3788 (det. Schmidt). 
Muna, 2,100 meters, 3907 (det. Schmidt). Cuzco: Santa Ana, 1,300 
meters, Weberbauer 5016 (det. Mez). Cosnipata, 700 meters, Weber- 
bauer 6963 (det. Schmidt). Puno: Rio Chaupimayo, Soukup 505. 
Loreto: Mouth of Rio Santiago, Tessmann 3936 (det. Schmidt). 
Amazonas: Rio Utcubamba, 1,900 meters, Weberbauer 4303 (N. 
rigida, det. Mez; but Schmidt has noticed that the style is elongate). 
Without locality, Poeppig. Brazil to Mexico. "Muena," "huarme 
tashango." 

Nectandra Sawadai 0. C. Schmidt, Repert. Sp. Nov. 31: 
187. 1933. 

Branchlets terete, glabrous like the entire plant, except the leaf 
nerves beneath and the flowers, these perfect, 5.5 mm. broad; peti- 
oles 7-11 mm. long; blades elliptic, narrowed at each end, acuminate, 
10-13 cm. long, 3.3-4.5 cm. wide, thin, somewhat lustrous above, 
the lateral nerves 6-8, prominently reticulate- veined on both sides; 
inflorescence 5-9 cm. long; pedicels 2-3 mm. long; perianth tube 
obsolete; outer anthers subsessile, rotund-rectangular, the inner 
filaments and staminodia glabrous, the glands large; ovary glabrous, 



FLORA OF PERU 927 

1 mm. long, the style 0.6 mm. long. Related to N. nitidula Nees, 
436, with larger flowers and pubescent pedicels. 

Huanuco: Pampayacu, Sawada 10, type. 

Nectandra superba A. C. Smith, Phytologia 1: 120. 1935. 

A tree 35 meters high, the trunk 70 cm. in diameter; subterete 
branchlets, petioles (15-25 mm. long), and inflorescence brownish- 
tomentose; blades coriaceous, drying olivaceous, glabrous and 
lustrous above except on the costa, beneath densely appressed- 
pilose, acute at each end, elliptic-oblong, 15-20 cm. long, 5-8 cm. 
wide, the lateral nerves 5-8, prominent beneath, the reticulate 
veins conspicuous on both sides; inflorescences to 30-flowered, 5-12 
cm. long, the lateral branchlets 4-10 mm. long; flowers subsessile, 
in fascicles of 2-4; perianth tube 1.5-2 mm. long, equaled by the 
ovate lobes; outer anthers subsessile, suborbicular, obtuse, bearing 
dorsally 2-4 minute, black glands; anthers and filaments of series 
3 subequal, glandular. Separated by the author from N. lineati- 
folia because of its prominent leaf reticulation and subsessile flowers 
with blunt anthers. 

Rio Acre: Rio Macauhan, Krukoff 5757, type. 

Nectandra viburnoides Meissn. in DC. Prodr. 15, pt. 1: 162. 
1864. 453. 

Distinguishable from N. latifolia (HBK.) Mez by the dull, often 
somewhat narrower leaves, usually less than 6 cm. wide, and the 
subsessile stigma. A shrub, or a tree to 20 meters high, with white 
or straw-colored flowers. Neg. 3771. 

Rio Acre: Ule 9472 (det. Mez). Brazil. 

Nectandra Williamsii 0. C. Schmidt, Repert. Sp. Nov. 31: 
188. 1933. 

A shrub, glabrous except for the slightly strigillose flowers and 
tomentulose pedicels, the flowers 4 mm. broad, perfect, the pedicels 
2-3 mm. long, slender; petioles 4-8 mm. long; blades lance-elliptic, 
6-11 cm. long, 2-3.5 cm. wide, acute at the base, acuminate, dull 
above, papyraceous, the lateral nerves 7-8, rather conspicuous and 
reticulate beneath; inflorescence 6-10 cm. long; perianth tube obso- 
lete; outer anthers rotund-trapezoid, 0.4-0.5 mm. long, longer than 
the glabrous filaments; staminodia stipitiform, pilose; ovary and 
style glabrous, the ovary 0.8 mm. long, the style half as long. 

San Martin: Tarapoto, 750 meters, Williams 6001, type; 5950. 



928 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Nectandra yarinensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 230. 1928. 

A small tree, the trunk 6 cm. in diameter, with short-acuminate, 
narrowly elliptic-oblong, pergamentaceous leaves; blades 15-30 cm. 
long, 5-10 cm. wide, cuneately narrowed to the very thick petiole; 
inflorescence to 13 cm. long; flowers whitish, 4 mm. wide, minutely 
and sparsely puberulent, the tube obsolete. Compared by the 
author with the smaller-leaved N. surinamensis Mez, 454. Neg. 
3773. 

Loreto: Middle Rio Ucayali, Boca del Yarina, Tessmann 3376, 
type. 

10. PLEUROTHYRIUM Nees 

Flowers perfect; perianth tube rarely well defined, the segments 
equal. Stamens of the 3 outer series fertile, the fourth always 
aborted. Filaments glabrous or papillose, all 9 basally glandular, 
the glands usually confluent. Upper anther cells introrse, the lower 
extrorse. Style usually conspicuous. Fruit apparently unknown. 
Doubtfully separable naturally from Ocotea, which may have, excep- 
tionally, all the filaments glandular, but in that genus the lower 
anther cells of the outer series are never exactly extrorse, nor is the 
androecium so compact by the fusion of the glands. 

Leaves glabrous or glabrate beneath. 

Leaves medium-sized, 10-20 cm. long P. densiflorum. 

Leaves very large, several decimeters long. 

Perianth tube 2 mm. long P. maximum. 

Perianth tube scarcely 1 mm. long. 

Leaves membranous, with 20 or more nerves . . P. Williamsii. 

Leaves coriaceous, with 16 or fewer nerves P. Krukovii. 

Leaves with cinereous or ferruginous pubescence beneath. 

Leaves very rigid P. Poeppigii. 

Leaves chartaceous-coriaceous. 

Leaves short-acute; inflorescence branched. 

Pubescence lax, ferruginous on the leaves beneath. 

P. chrysophyllum. 

Pubescence close, cinereous on the leaves beneath. 

P. cuneifolium. 

Leaves caudate-acuminate; inflorescence subsimple. .P. bifidum. 



FLORA OF PERU 929 

Pleurothyrium bifidum Nees, Syst. Laur. 351. 1836; 469. 

A tree of 20-30 meters, the trunk 40 cm. in diameter; with the 
characters of P. chrysophyllum Nees except that the often larger 
leaves are definitely acute or acuminate and are about equaled by 
the inflorescence; flowers golden brown; style very short or obsolete. 
Resembles also P. panurense Mez, 468, in foliage, but the inflores- 
cence of that Amazonian species is much shorter than the leaves and 
the style equals the ovary. In the type some of the leaves are more 
than 30 cm. long. Neg. 31202. Pleurothyrium nobile A. C. Smith, 
Phytologia 1 : 120. 1935, Amazonian, was separated from P. bifidum 
by its glabrous or only minutely tomentulose inflorescence, and large 
flowers, on pedicels 6-10 mm. long; from P. Poeppigii by its gla- 
brous leaves. 

Loreto: Mouth of Rio Santiago, Tessmann 4634 (det. Schmidt). 
Yurimaguas, Poeppig 2398, type. 

Pleurothyrium chrysophyllum Nees, Syst. Laur. 351. 1836; 
471. 

Similar to P. Poeppigii Nees, but the thinner leaves rounded at 
the apex, 15-25 cm. long, 10-15 cm. wide, ferruginous-tomentose 
beneath, with 20-24 sometimes forked lateral nerves; panicles 
oblong, 10-15 cm. long, the branches 2-5 cm. long, the pedicels 
obsolete or 3 mm. long; filaments definitely shorter than the anthers. 
A beautiful tree 20-25 meters high, with yellow wood and cinna- 
mon-colored flowers (Poeppig). 

Huanuco: Cochero, Poeppig 1718 (according to Nees; in herb. 
Berlin as 1719, and as from Chile); also 1432, fide Nees. 

Pleurothyrium cuneifolium Nees, Syst. Laur. 352. 1836; 470. 

A tree of 12 meters, the trunk 20 cm. in diameter; blades oblong- 
oval or obovate, gradually attenuate to a cuneate base, 15-20 cm. 
long, 6-8 cm. wide, abruptly acute or acuminate, finely but softly 
pale-ferruginous-tomentose beneath, like the subterminal panicles, 
the nerves 14-15; petioles 8-10 mm. long; panicle branches com- 
pressed, about 2 cm. long, bifid, the branchlets 1-3-flowered; flowers 
yellow-orange, 5-7 mm. long, the outer segments elliptic, the inner 
oblong; filaments shorter than the anthers; style longer than the 
ovary. The fistulose branchlets are reported to be inhabited by 
ants. Neg. 3582. 

Loreto: Mouth of Rio Santiago, Tessmann 4526 (det. Schmidt). 
Balsapuerto, 220 meters, Klug 2931; a tree of 5 meters; flowers 



930 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

yellow. Cachipuerto, King 3116. Fortaleza, 140 meters, King 2779; 
a tree of 14 meters. Pumayacu, 600-1,200 meters, King 3195, 3228. 
San Martin: Zepelacio, Klug 3567. Brazil. 

Pleurothyrium densiflorum A. C. Smith, Bull. Torrey Club 
58: 109. 1931. 

A tree of 10 meters, essentially glabrous, or the young parts 
pale-puberulent, like the ample inflorescence and the flowers; peti- 
oles slender, 7-15 mm. long; blades oblong or elliptic-oblong, cuneate 
at the base, obtuse or subacute at the apex, 13-18 cm. long, 4.5-6.5 
cm. wide, dull, chartaceous; flowers 3-5 mm. broad, nearly 4 mm. 
long, the segments oblong, obtuse; pedicels 3-6 mm. long; outer 
filaments 0.4-0.5 mm. long, little shorter than the anthers. 

Loreto: Mishuyacu, 100 meters, Klug 1372, type; 1301. Balsa- 
puerto, 220 meters, Klug 2908; a tree of 6 meters; flowers white. 
Caballo-cocha, Williams 2273. "Canela muena." 

Pleurothyrium Krukovii A. C. Smith, Phytologia 1: 121. 1935. 

Branchlets terete, fistulose, soon glabrate; petioles rugose, 
canaliculate, 1.5-3 cm. long; blades coriaceous, glabrous, the 14-16 
nerves and reticulate veins prominent beneath, obscure above, 
acute or short-acuminate, acute at the base, 25-45 cm. long, 7-12 
cm. wide; inflorescences solitary, axillary, 7-12 cm. long, many- 
flowered, the branchlets slender, 4-10 mm. long, like the flowers 
minutely cinereous- tomentulose; flowers in fascicles of 3-6; mature 
pedicels 3-4 mm. long; perianth tube 1 mm. long, the oblong, obtuse 
lobes 2.5 mm. long; stamens 1 mm. long, the anthers oblong, emar- 
ginate, equaled by the slender filaments, all 9 glandular, the glands 
not confluent; ovary glabrous, equaled by the tomentulose style. 
Compared by the author with P. parviflorum Ducke, with much 
smaller leaves and inflorescences. 

Rio Acre: Rio Macauhan, Krukoff 5563, type; 5255, 5722. 

Pleurothyrium maximum 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 235. 1928. 

A small tree, 4-6 meters high, the trunk 7-8 cm. in diameter, 
with velvety, red-pubescent branchlets and inflorescence; blades 
obovate-lanceolate, 30-40 cm. long, 7.5-14 cm. wide, cordate at 
the base, more or less acuminate, papyraceous, glabrous; flowers 
green, 7-9 mm. long, the tube 2 mm. long, in narrow, thyrsoid 
panicles sometimes 65 cm. long; pedicels slender, 5-10 mm. long; 
stamens 1.6 mm. long, the anthers and filaments subequal; ovary 



FLORA OF PERU 931 

1.8 mm. long, nearly twice as long as the style. A well marked 
species because of its cordate-based, large leaves, greatly elongate, 
slender inflorescence, and green flowers. Neg. 3583. 

Loreto: Mouth of Rio Santiago, Tessmann 4040, type; 4529. 

Pleurothyrium Poeppigii Nees, Syst. Laur. 349. 1836; 471. 

A shrub or tree with fleshy, ferruginous-tomentulose branch- 
lets and inflorescence, and obovate, rigid-coriaceous leaves, gla- 
brous above but very finely and densely pulverulent beneath; blades 
about 15 cm. long, 5-8 cm. wide, subobtuse, acute at the base; peti- 
oles about 2 cm. long; inflorescence many-flowered, the branches 
5-10 cm. long, the pedicels 3-6 mm. long; filaments and anthers, 
like the style and ovary, subequal. Neg. 3586. 

Loreto: Yurimaguas, Poeppig, type. 

Pleurothyrium Williamsii 0. C. Schmidt, Repert. Sp. Nov. 31: 
189. 1933. 

Branches quadrangular; petioles and inflorescence, including the 
flowers, velvety-brown-tomentose; petioles 5 mm. long, 2.5 mm. 
thick; blades narrowly oblanceolate, 30-40 cm. long, 7.5-9.5 cm. 
wide, truncate-cordate at the base, narrowed at the apex into an 
acumination 3 cm. long, membranaceous, glabrous except on the 
nerves, these rather prominent, especially the 22-24 lateral ones 
beneath, finely reticulate- veined ; inflorescence 8-12 cm. long, the 
peduncle 4-6 cm. long; flowers 7-8 mm. broad, on pedicels 4-6 mm. 
long, the tube scarcely 1 mm. long; stamens 0.7-0.9 mm. long; ovary 
1 mm. long, the style only slightly shorter. 

Loreto: Pebas, Williams 1766, type. 

77. HERNANDIACEAE. Hernandia Family 
Reference: Meissner in Mart. Fl. Bras. 5, pt. 2: 291-294. 1866. 

The Peruvian representatives are probably always scandent or 
subscandent shrubs; leaves simple, with 3 (-5) principal nerves 
from the broad base; flowers almost minute, in axillary, corymbose 
or paniculate cymes. Calyx with 3-5 valvate, subequal segments 
in 2 series, the same number of stamens with 2-celled anthers in 
1 row, and 1-2 rows of glandular staminodia. Fruit dry, somewhat 
costate-angulate, oblong or ovoid, the surrounding receptacle more 
or less inflated. The family has been included in the Lauraceae. 



932 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. SPARATTANTHELIUM Mart. 
Characters as above. 

Branches glabrous or glabrate, the leaves sometimes puberulent 
beneath. 

Sepals 4; leaves somewhat lustrous. S. amazonum. 

Sepals 5-7; leaves dull or nearly so S. atrum. 

Branches conspicuously pubescent, sometimes shortly so. 

Branches densely and shortly puberulent- villous S. acreanum. 

Branches villous with long, spreading hairs S. tarapotanum. 

Sparattanthelium acreanum Pilger, Notizbl. Bot. Gart. Ber- 
lin 6: 295. 1915. 

Branchlets, petioles, inflorescence, and leaves beneath some- 
what villous- tomentose; blades chartaceous, ovate-elliptic, rounded 
or slightly subacute at the base, 7-10 cm. long, 4-5.5 cm. wide, 
glabrate above except on the nerves, these 3 and, with the veins, 
prominent beneath; petioles 1.5-2.5 cm. long; panicles axillary and 
terminal, to 15 cm. long, with divaricate branches; flowers densely 
aggregate, densely rusty- tomentose, short-pedicellate; sepals 5, 
lanceolate, villous within, 2-2.5 mm. long. Said to differ in leaf 
form from S. tarapotanum; leaves less pubescent and sepals 4 in the 
similar S. tupiniquinorum Mart., 291. Neg. 13547. 

Rio Acre: Cobija (Bolivia), Ule 9405, type. 

Sparattanthelium amazonum Mart. Denkschr. Bot. Ges. 
Regensb. 3: 303. pi. 11. 1841; 293. 

Petioles 10-20 mm. long; blades concolorous, rounded or emar- 
ginate at the base, acuminate, ovate or oblong, herbaceous, gla- 
brous like the branchlets and peduncles, 5-12 cm. long, 2.5-3.5 cm. 
wide, 3-nerved, the veins lax, slender; panicle corymbiform, lax, 
about equaling the leaves, the white-tomentulose pedicels and 
flowers 1 mm. long, the perianth 4-parted; filaments and anthers 
subequal. Neg. 7308. 

Loreto: Fortaleza, 140 meters, Klug 2771. Region of Iquitos, 
King 599; Killip & Smith 29835, 27281; Williams 1953. Brazil. 

Sparattanthelium atrum Pilger, Notizbl. Bot. Gart. Berlin 
6:295.1915. 

Younger branches glabrous; leaves thin, not at all lustrous, ellip- 
tic, rounded or subcordate at the base, short- or long-acuminate, 
10-15 cm. long, 5.5-7.5 cm. wide, the acumen 1-2 cm. long, glabrous, 



FLORA OF PERU 933 

the 3 nerves, like the veins, prominent only beneath; peduncles gla- 
brous, 3-4.5 cm. long; panicles axillary, to 8 cm. long, lax; flowers 
puberulent, densely aggregate, the slender, puberulent pedicels as 
long; sepals 6 (5-7), more or less connate, obtuse, 1.5-1.7 mm. long. 
Differs from S. amazonum by its dull leaves and more numerous 
sepals (Pilger) . How constant these characters are for this and other 
species remains to be shown by more abundant collections. Neg. 
13549. 

Rio Acre: Seringal Guanabara (Brazil), Ule 9406, type. 

Sparattanthelium tarapotanum Meissn. in Mart. Fl. Bras. 
5, pt. 2: 294. 1866. S. Sprucei Rusby, Descr. N. Sp. S. Amer. PL 
22. 1920. 

Apparently similar to S. amazonum, but the branches and leaves 
villous, the leaves becoming glabrate above; panicles dense; pedicels 
about 2 mm. long; perianth often 5-parted; filaments very short. 
Neg. 13550. 

San Martin: Tarapoto, Spruce 4222, type; Spruce 422a, type of 
S. Sprucei; Williams 6280, 5471, 6171, 6252. Juanjui, 400-800 
meters, river bank, Klug 4233; flowers greenish brown. 

78. PAPAVERACEAE. Poppy Family 
Reference: Fedde, Pflanzenfam. ed. 2. 17b: 5-145. 1936. 

Peruvian members of the family are of some economic impor- 
tance. The poppy-like Argemone mexicana is the source of thistle 
oil, a clear, yellow oil extracted from the seeds, suitable for fuel and 
use as an ointment; in Peru (Hen era) as in Brazil this oil is used as 
a purge and for colic. Further, in Peru the flowers are employed as a 
narcotic, and the plant furnishes a yellow dye (Herrera). 

Herbs. 

Plants prickly 1. Argemone. 

Plants unarmed 2. Chelidonium. 

Shrubs or small trees 3. Bocconia. 

1. ARGEMONE L. Prickly poppy 

Coarse, mostly bushy-branched and prickly plants with sub- 
pinnately lobed leaves and showy, poppy-like flowers. Stigmas not 
united, at least basally, to form a flat disk as in the true poppy, 
Papaver. 



934 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Argemone mexieana L. Sp. PI. 508. 1753. 

Capsules prickly; flowers typically yellow or orange; stigma lobes 
suberect; style obsolete. Var. ochroleuca (Sweet) Lindl. has lighter- 
colored, more loosely disposed flowers, the stigma lobes divergent, 
the style developed. A striking variation is var. gynophora (Fedde) 
Macbr., comb. nov. (A. mexieana var. ochroleuca subvar. gynophora 
Fedde, Pflanzenreich IV. 104: 278. 1909), with slightly double flowers 
that persist about the base of a stipitate capsule, the stipe 2-3 cm. 
long. Illustrated, Gray, Gen. 1: pi. 47; Pflanzenreich IV. 104: 274 
(var. gynophora). 

Lima: Lurin, Ruiz & Pavon; Gaudichaud; (WilkesExped.). Ma- 
tucana, 2,400 meters, 401. Bed of Rio Rimac, Wawra 451 (type of 
var. gynophora). Cuzco: In grain and other cultivated fields (He- 
rrera). Oropeza Valley, 3,150 meters, Herrera 2169. Piura: Parinas 
Valley, Haught 272; growing in watercourse among the Amotape 
Hills. San Martin: Tarapoto, in pastures, 750 meters, Williams 
5420, 6606. Tacna: Tacora, Isern 2276. Arequipa: Cachendo 
(Gunther & Buchtien 289). Presumably native of tropical America; 
now nearly cosmopolitan. "Ccarhuinchu," "ccarhuinchunca." 

2. CHELIDONIUM L. 

Unarmed, round-stemmed herbs with irregularly lobed or sub- 
pinnate leaves, long-pediceled, yellow flowers," and linear pods on 
slender pedicels. Petals 4. 

Chelidonium ma jus L. Sp. PI. 505. 1753. 

Leaves gray-green, irregularly divided or lobed; pedicels bracte- 
ate at the base; pods slightly torulose. Illustrated, Pflanzenreich 
IV. 104: 213. 

Lima: Adventive on rock walls at Matucana and elsewhere. 
Native of the Old World. "Celidefia." 

3. BOCCONIA L. 

By Paul C. Standley 

Reference: Hutchinson, Kew Bull. 275-282. 1920. 
Tall shrubs or small trees, simple or sparsely branched, the large 
leaves clustered at the ends of the branches; sap orange; flowers 
small, apetalous, in large, terminal panicles; ovary long-stipitate, 
with a single ovule; valves of the small fruit fleshy, opening from be- 
low upward; seeds surrounded by a large aril. From South America 
only one other species is known, B. pubibractea Hutchinson, for which 



FLORA OF PERU 935 

its author cites five collections from "Colombia." The type was 
collected by Pearce on slopes of the mountains about "Mufia," which 
I suspect is Mufia, Peru. However, I have seen no Peruvian material 
agreeing with the description. Pearce's labels usually bear no name 
of the country in which the specimens were collected, and since many 
of the localities are obscure, often it is difficult to refer them defi- 
nitely to any of the Andean countries. 

Leaves deeply lobate, with elongate, narrow, acuminate lobes, abun- 
dantly tomentose beneath or rarely glabrate B. Pearcei. 

Leaves entire or merely coarsely crenate. 

Leaves entire or practically so, glabrous beneath . .B. Macbrideana. 

Leaves coarsely crenate, densely tomentose beneath .B. integrifolia. 

Bocconia integrifolia Humb. & Bonpl. PI. Aequin. 1: 119. 
pi. 35. 180-7. B. frutescens var. integrifolia Kuntze, Rev. Gen. 3, 
pt. 2: 4. 1898. 

A shrub or small tree, usually 3-4.5 meters high; leaves short- 
petiolate, elongate-elliptic, the larger ones more than 30 cm. long, 
obtuse or acute, serrate to shallowly crenate-lobate, the low lobes, 
when present, rounded or very obtuse, glabrous above, usually pale 
beneath and brownish- tomentose; panicles as much as 35 cm. long; 
sepals obovate-elliptic, 1 cm. long, glabrous; stamens about. 10; 
fruit ellipsoid, 12 mm. long. 

Cajamarca: Near Cascas (Bonpland, type). Without locality: 
Gay 675. Huanuco: Between Huanuco and Pampayacu, Kanehira 
233. Chaclla, 2,700 meters, 3644; a shrub 4.5 meters high, leafless 
except at the top; inflorescence more or less drooping, pale lavender. 
Twelve miles south of Panao, 3,000 meters, in creek bottom and open 
woods, 2226. Junin: Huacapistana, 1,800-2,400 meters, Killip & 
Smith 24149; a tree of 3-4.5 meters. Bolivia to Venezuela. 

Bocconia Macbrideana Stand!., sp. nov. 

Frutex ut videtur ramosus, omnino glaber; folia parva petiolata 
crasse membranacea, petiolo gracili 2-2.5 cm. longo; lamina angus- 
tissime oblonga, fere lineari-oblonga, 9-15 cm. longa 1.5-2.5 cm. 
lata acutiuscula apiculata, basi attenuato-acutata, integra vel per- 
obscure remote serrulata, subtus viridis, non glauca; panicula bre- 
viter pedunculata thyrsiformis vel anguste pyramidalis 16-20 cm. 
longa 7-10 cm. lata laxe multiflora; sepala 4-5 mm. longa elliptica 
apice abrupte contracta glabra; styli rami minute sparse puberuli; 
fructus longe stipitatus ellipsoideo-oblongus ca. 7 mm. longus longis- 
sime rostratus. 



936 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Junin: Rio de Comas, 2,300-2,400 meters, Weberbauer 6619 (type 
in Herb. Field Mus.). 

The species of Bocconia have been separated chiefly by the form 
of their leaves, and in its leaves this plant is certainly unlike any of the 
species known heretofore. 

Bocconia Pearcei Hutchinson, Kew Bull. 278. 1920. 

A shrub or small tree, 1-3 meters high, the branches glaucous; 
leaves petiolate, as much as 45 cm. long, deeply pinnate-lobate, the 
lobes acute or acuminate, serrate, glabrous above, pale beneath and 
brownish-tomentose, rarely glabrate; panicles as much as 40 cm. 
long, or probably larger; sepals 10-12 mm. long, glabrous; stamens 
about 20; fruit glaucous, 1 cm. long. 

Huanuco: Huasahuasi, Dombey; Ruiz & Pavon. Without local- 
ity: Poeppig 1089. Junin: Chanchamayo Valley, 1,200 meters, 
Schunke 415. Chanchamayo, Isern 2239. La Merced, 5712. 
Sandia: (Weberbauer, 89)', Gay. Cuzco: Machupicchu, Soukup 312. 
San Martin: San Roque, in forest, 1,400 meters, Williams 7139. 
Foot of Cerro Campana, Spruce 4301. Bolivia to Ecuador. "Palo 
de tinta," "palo amarillo," "haiuna." 

This plant has been referred generally toB. frutescens L., to which, 
indeed, it is closely related, but, as pointed out by Hutchinson, that 
species is restricted to Mexico, Central America, and the West Indies. 

78. FUMARIACEAE. Fumitory Family 
By Julian A. Steyermark 

1. FUMARIA (Tourn.) L. 

Slender, leafy-stemmed annuals with finely dissected, compound 
leaves. Flowers small, in racemes or spikes. Pedicels 2-bracted. 
Corolla irregular, with 1 petal spurred at the base, the 4 petals in 2 
pairs. Stamens 6, in two sets of 3 each. Style deciduous. Fruit 
small, 1-seeded, globular, indehiscent. Seeds crestless. All the 
species of Fumaria found in Peru are natives of Europe. 

Corolla 9-12 mm. long F. capreolata. 

Corolla 3-7 mm. long. 

Sepals very small, 0.5-1 mm. long, less than one-fourth the length 
of the corolla F. parviflora. 

Sepals 1.5-2 mm. long, one-third or more the length of the corolla. 

F. densiflora. 



FLORA OF PERU 937 

Fumaria capreolata L. Sp. PI. 701. 1753. 

Plants 15-40 cm. tall; ultimate leaf segments relatively large, 
1-4 mm. broad, oblong-cuneate, obtuse; peduncles 3.5-5.5 cm. long, 
many-flowered; corolla greenish white with purplish tips; sepals half 
as long as the corolla; fruit rounded at the tip. 

Lima: San Geronimo, 150 meters, 5900. Atocongo, 250-500 

meters, Pennell 14774- 

Fumaria densiflora DC. Cat. Hort. Monsp. 113. 1813. 

Plants up to 40 cm. tall; ultimate leaf segments short and narrow, 
2-3 mm. long, 0.5-1 mm. broad, acute; peduncles about equaling 
the petioles, 0.5-1.5 cm. long, many-flowered; corolla rosy purple, the 
tips dark purple, 5-6 mm. long; fruit slightly short-pointed at the apex. 

Lima: Matucana, 2,400 meters, 379. 

Fumaria parviflora Lam. Encycl. 2: 567. 1788. 

Plants up to 30 cm. tall; ultimate leaf segments long and slender, 
2.5-7 mm. long, 0.5-1 mm. broad, acute; peduncles about equaling 
the petioles, less than 1 cm. long, many-flowered; corolla pinkish 
with dark purple tips, 3-5 mm. long; fruit slightly short-pointed 
at the apex. 

Lima: Matucana, 2,400 meters, 307. 

79. CRUCIFERAE. Mustard Family 
Reference: Engl. & Prantl, Pflanzenfam. ed. 2. 17b: 227-658. 1936. 

Herbs, sometimes woody at the base in some of the alpine species, 
with watery, often acrid sap. Leaves alternate or basal, simple or 
pinnately divided. Flowers racemose or corymbose. Sepals 4, 
deciduous or persistent. Petals 4, hypogynous, nearly equal, usually 
narrowed at the base into a claw. Stamens 6, hypogynous, 2 shorter 
than the other 4. Pistil 1, composed of 2 united carpels. Fruit a 
silique or silicle, 1-2-celled, when 2-celled, the cells separated by a 
thin partition stretched between the 2 marginal placentae, from 
which the valves separate when ripe, sometimes indehiscent or 
separating into 1-seeded joints. In this flora, where simplicity in 
classification is sought, generic lines in the Cruciferae sometimes 
have been drawn less finely than by the most recent monographer. 
Nevertheless, the following key, as regards major details, is taken 
directly from his. 



938 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pods with a gynophore, and divided into 2 parts (in Cremolobus the 
gynophore is rarely absent). 

Pod parts provided with a cross wing 1. Loxoptera. 

Pod parts without a cross wing. 
Pod spectacle-shaped, winged or at least marginate. 

2. Cremolobus. 
Pod not spectacle-shaped, not at all margined, keeled. 

3. Urbanodoxa. 

Pods not divided into 2 parts, rarely with a gynophore. 
Pods consisting of a lower and upper segment, the latter often 
beaklike; cotyledon leaves nearly always folded lengthwise; 
pubescence simple or none. 

Segments of the fruit not sharply defined; petals usually yellow. 

4. Brassica. 

Segments of the fruit sharply defined; petals usually white or 
purple 5. Raphanus. 

Pods not divided into a valvular and stylar portion, sometimes 
sectionally parted; cotyledons never folded lengthwise; 
pubescence various. 

Pods laterally compressed, the septum linear or nearly so. 
Pods not distinctly 2-parted. 
Pods evidently flattened. 

Cotyledon leaves petiolate; pods circular to elliptic; 
leaves indistinctly if at all auricled .... 6. Lepidium. 
Cotyledon leaves sessile; pods more or less obcordate, 
except in Isatis; leaves auricled or hastate. 

Plants glabrous, or the trichomes simple 7. Isatis. 

Plants pubescent, the trichomes somewhat branched. 

8. Capsella. 

Pod valves rounded, the pods elliptic 9. Mancoa. 

Pods distinctly 2-parted; cotyledon leaves petiolate. 

10. Coronopus. 
Pods not laterally compressed, the septum broad. 

Pods, if longer than broad, only a few times longer; Peruvian 

species sometimes suffrutescent 11. Draba. 

Pods narrow, usually and often many times longer than broad, 
except in Sarcodraba, Eudema, and Alyssum. 



FLORA OF PERU 939 

A. Leaves simple to variously divided but not bipinnately 

dissected, or the moistened seeds not mucilaginous. 
B. Pods short, usually thick, sometimes linear but the 

plants then tufted perennials. 

Plants rather loosely growing, often rhizomatous or 
annual (Alyssum sometimes suffrutescent). 

Pods nerveless 12. Cardamine. 

Pods nerved or net- veined. 

Plants glabrous or nearly so, or spreading-hispid- 

ulous. 

Leaves often divided; septum cells not cross- 
partitioned 13. Rorippa. 

Leaves subentire; septum cells cross-partitioned. 

14. Dictyophragmus. 

Plants appressed-strigose, suffrutescent, flowering 

as an annual (Peruvian species) . 15. Alyssum. 

Plants tufted perennials, often caudex-forming or 

suffrutescent. 

Nectar glands distinct; stigma large, sessile; a suf- 
frutescent plant with showy flowers. 

16. Mathewsia. 
Nectar glands confluent; flowers rather small. 

Plants suffrutescent, glabrous 17. Eremodraba. 

Plants not suffrutescent but often with a woody 

caudex, often pubescent. 

Leaves rosulate, apparently sessile; sepals usu- 
ally persistent; styles 1.5-2 mm. long. 
Flowers conspicuously pedicellate; pods sub- 
oblong 18. Englerocharis. 

Flowers subsessile; pods ovoid-elliptic. 

19. Catadysia. 

Leaves partly cauline, at least the lower ones 
narrowed basally or petioled; sepals cadu- 
cous; style often shorter than 1 mm. 

20. Weberbauera. 

B. Pods elongate-linear; nectar glands confluent; plants 
often annual or, if suffrutescent, tall and loosely 

branched 21. Sisymbrium. 

A. Leaves bipinnately parted, often dissected, the divisions 
fine; moistened seeds mucilaginous. . .22. Descurainia. 



940 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. LOXOPTERA 0. E. Schulz 

Slender annuals with narrow, slightly dentate leaves. Outer 
sepals narrowed toward the base, the inner somewhat cucullate. 
Filaments filiform, the anthers cordate. Medial nectar glands 
absent. Pod valves roundish, with a medial as well as a radial wing. 
Floral characters nearly those of Urbanodoxa, but the fruit, except 
for the medial wing, nearly that of Cremolobus. 

Loxoptera stenophylla (Muschl.) 0. E. Schulz, Bot. Jahrb. 66: 
93. 1933. Cremolobus stenophyllus Muschl. Bot. Jahrb. 50: Beibl. 
111:7. 1913. 

Glabrous, much branched, 20-30 cm. high; leaves narrowly 
lanceolate, all but the upper ones coarsely dentate, to 4 cm. long and 
1 cm. wide; racemes terminal, lax, elongate; pedicels 1 cm. long; 
flowers 3-3.5 mm. broad; petals 2-2.5 mm. long; glands obsolete; 
pods broadly winged. Illustrated, Engl. & Prantl, Pflanzenfam. ed. 
2. 17b:314. 

Lima: On stony outcrops, 1,400 meters, Chosica, Weberbauer 
5335, tyve;Esposto 17. 

2. CREMOLOBUS DC. 

Glabrous or pubescent herbs or small, suffrutescent plants with 
alternate or opposite, entire to pinnate leaves and racemose, slen- 
der-pedicellate, ebracteate flowers. Sepals equal at the base. Pod 
stipitate, 2-celled, dehiscent, the valves compressed-carinate or 
winged, more or less pendulous. Seeds solitary, pendulous, emargi- 
nate. Style thickened at the base. See also the segregate Urbanodoxa. 

Plants annual. 

Pods pilose and ciliate; petals and sepals subequal. .C. Paysonii. 
Pods glabrous or puberulent; petals longer than the sepals. 

Leaves pinnatifid, with linear segments C. Weberbaueri. 

Leaves subentire to subpinnately lobed. 

Leaves ovate-subrhombic-elliptic, dentate to subentire. 

Valve wings of the pods deeply lobed C. sinuatus. 

Valve wings or margins subentire to irregularly crenate. 

Valves wing-margined, more or less crenate. 

C. parviflorus. 
Valves merely marginate, subentire C. humilis. 



FLORA OF PERU 941 

Leaves pinnately dentate or lobate or lyrate-pinnatifid, often 
suboblong. 

Valve wings shallowly crenate; leaves suboblong, pinnately 
dentate. 

Valves rugulose- veined C. pinnatifidus. 

Valves not rugulose C. aphanopterus. 

Valves entire-margined, not winged, rugulose; leaves 
broadly lyrate-lobed C. Benoistii. 

Plants perennial, more or less ligneous toward the base, often 
scandent. 

Style none C. subscandens. 

Style developed. 

Plants glabrous or nearly so. 
Pods 4-6 mm. wide; style elongate; leaves oblong-ovate. 

C. suffruticosus. 

Pods often wider; style short; leaves elliptic. . .C. peruvianus. 
Plants pubescent; style elongate C. pubescens. 

Cremolobus aphanopterus Gray, U. S. Expl. Exped. Bot. 55. 
pi. 3. 1854. 

Simple, or branching above, glabrous or nearly so; leaves 12-16 
mm. long, deeply pinnatifid, the oblong divisions very obtuse; 
racemes many-flowered; pedicels 4-6 mm. long; flowers very small, 
the obovate petals sessile; stipe as long as the ovary and long style; 
pod minutely puberulent, not rugose or reticulate, wingless but 
with an undulate or interrupted margin, not longer than the stipe, 
scarcely 2 mm. wide; seeds suborbicular. 

Lima: Near Obrajillo (Wilkes Expedition, type). 

Cremolobus Benoistii Macbr., sp. nov. 

Annua, ad 30 cm. alta; caulibus striato-angulatis sparse hispi- 
dulis; foliis (radicalibus ignotis) late oblongo-ellipticis, valde basin 
versus cuneato-angustatis, superne profunde irregulariter pinnato- 
lobatis, supra sparsissime hispidulis vel fere glabris, ad 5 cm. longis 
atque 2.5 cm. latis, lobis oblongis obtusis plerumque 3-4 mm. latis; 
racemis sat dense hispidulis ad 15 cm. longis; pedicellis fructiferis ca. 
7 mm. longis recurvo-divaricatis; sepalis obtusis fere 2 mm. longis; 
petalis ca. 3 mm. longis; siliculae valvis suborbiculatis reticulato- 
venosis 1 mm. latis integerrimis emarginatis. Similar to the gla- 



942 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

brous C. aphanopterus and C. pinnatifidus, with crenate-margined 
pods; named for Raymond Benoist, who found the material in the 
Paris herbarium. Neg. 34245. 

Provincia de Conduces, 3,000 meters, Raimondi (type in herb. 
Paris). 

Cremolobus humilis Muschl. Bot. Jahrb. 40: 269. 1908. 

Stems mostly simple, 3-10 cm. high; basal leaves ovate or obovate- 
oblong to subrotund, subacute, narrowed to the base, entire or den- 
tate, 5-10 mm. long, 3-8 mm. wide; cauline leaves sessile, the middle 
ones oblong, to 7 mm. long; racemes 2-6 cm. long; pedicels about 4 
mm. long, slender; petals white, oblong, 1.5-3 mm. long; pods nar- 
rowly winged, minutely crenulate, obscurely reticulate, 1 mm. wide. 
Neg. 13836. 

Ancash: Huaraz, 4,300 meters, Weberbauer 3097, type; 227. 
Junin: Shallow, stony soil, Rio Blanco, 3,600 meters, 697. Lima: 
In shallow soil on rock, Viso, 2,700 meters, 601. Canta, 2,000-2,900 
meters, Pennell 14588. San Buenaventura, 2,700-2,900 meters, 
Pennett 14533. Above Obrajillo, 3,300-3,400 meters, Pennell 13223, 
14398. Arequipa: Volcan de Misti, Pennell 13223. 

Cremolobus Paysonii 0. E. Schulz ex Macbr. Field Mus. Bot. 
8:80.1930. 

An erect annual, to 20 cm. high, branching above, the branches 
erect-spreading; lower leaves unknown, the upper oblanceolate, acute, 
cuneate at the base, coarsely dentate or subpinnatifid, with 2-4 
unequal teeth on each side; racemes 20-40-flowered ; sepals 2 mm. 
long, the white petals a little longer; fruiting pedicels spreading, 
4-8 mm. long; stipe and style 1.2 mm. long; pods 4 mm. broad, cen- 
trally pilose, the broad wing ciliate, irregularly and obtusely dentate 
or sinuate-repand. Similar to C. chilensis DC., reported from Peru 
but apparently by error, but that has smaller pods with narrower, 
entire wings. 

Lima: Matucana, 2,400 meters, 211, type. 

Cremolobus parviflorus Wedd. Ann. Sci. Nat. V. 1: 283. 1864. 
C. pinnatifidus Hook. var. integrifolius Gray, U. S. Expl. Exped. 
Bot. 56. 1854. 

A small, glabrous annual with oblong to subrhombic-lanceolate, 
acute leaves, attenuate to a short petiole, coarsely few-serrate; 
fruiting raceme lax, elongate; pods much shorter than the slender 
pedicels, with narrow, crenate wings. Type, Mandon 905, from 
Bolivia. Neg. 27293. 



FLORA OF PERU 943 

Lima: Chicla Pass, 4,000 meters (Ball, fide Gilg & Muschl.). 
Obrajillo (Wilkes Expedition). Bolivia. 

Cremolobus peruvianus (Lam.) DC. Syst. 2: 419. 1821. Bis- 
cutella peruviana Lam. Diet. 3: 620. 1791. 

A sprawling, suffrutescent plant with slender, branching stems; 
leaves ovate-elliptic, to 3 cm. wide, subcuneate at the base, short- 
petiolate, acute, serrate to subentire, 5-8 cm. long; racemes panicu- 
late, to 10 cm. long; pedicels filiform, 6 mm. long; pod valves 6-8 
mm. broad. Type from Ecuador. Neg. 27294. 

Junin: Jauja, 2,200-2,300 meters, Weberbauer 6673. Without 
locality: Jussieu; Ruiz & Pawn. Ecuador. 

Cremolobus pinnatifidus Hook. Icon. 1: pi. 100. 1837. 

Plants about 10 cm. high, with one or two branches, glabrous; 
leaves sessile, oblong, sublyrate-pinnatifid; racemes finally longer 
than the branches; sepals ovate; pods rugose, the broad wings crenate. 
The commonest species in Peru. C. chilensis DC., of Chile, has 
pods with entire, narrow wings; reported from Peru but, so far as 
known, on misdetermined specimens. 

Moquehua: Carumas, Weberbauer 7281. Torata, 2,700 meters, 
Weberbauer 7416. Lima: Huaros, 3,200-3,400 meters, Pennell 14702. 
Huamantanga (Mathews 590) . Matucana, 84, 112, 428, 467, 557. Rio 
Blanco, in clumps in slide rock, or in stony, shallow soil on rocks, 696. 

Cremolobus pubescens Hook. Icon. 1: pi. 81. 1837. 

A sprawling or clambering, pubescent plant with ovate, short- 
petiolate, subentire leaves 4 cm. long or longer, and paniculate, leafy 
racemes of minute flowers; petals and sepals equal; pods broadly 
winged, repand-denticulate. Probably a variety of C. peruvianus. 

Peru: Almirante (Mathews 1606). Ecuador. 

Cremolobus sinuatus Hook. Icon. 1: pi. 81. 1837. 

Slender and often simple, with few petiolate, coarsely serrate 
leaves about 1 cm. long, and short, subpaniculate racemes of small 
flowers; pods broadly winged, the wings deeply and very obtusely 
lobed, the lobes sometimes even subretuse. 

Cajamarca: Cascas, Raimondi. Lima(?): Cuesta de Puruchuco 
(Mathews, type). 

Cremolobus subscandens Kuntze, Rev. Gen. 3, pt. 2: 4. 3898. 
Clambering in shrubs, glabrous or finely pubescent; leaves oppo- 
site, ovate, remotely serrate, 4 cm. long; racemes loosely panicled; 



944 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

floral glands large; style none; one valve of the pods usually aborted, 
the other one reticulate, the margin apiculate-serrate or dentate- 
fimbriate. 

Amazonas: Chachapoyas, Weberbauer 4438. Bolivia. 

Cremolobus suffruticosus DC. Syst. 2: 419. 1821. 

Similar in habit to C. peruvianus, but the leaves ovate-oblong, 
about 1 cm. wide, often more closely serrate, and the much smaller 
pods with an elongate style. Negs. 27295, 13832. 

Huanuco: Raimondi. Lima: Cheuchin, Ruiz & Pawn. Without 
locality: Dombey. Ancash: Raimondi. Chile; Ecuador. 

Cremolobus Weberbaueri Muschl. Bot. Jahrb. 40: 270. 1908. 

Stems erect, sometimes laxly branched, to 20 cm. high; leaves 
pinnate, the remote, linear divisions mostly entire; racemes 3-6 cm. 
long, few-flowered; pedicels 4-5 times longer than the white or yellow- 
ish flowers; sepals broadly lanceolate, 2 mm. long; petals ovate, 3-4 
mm. long; pod wings crenate, the valves about 1 mm. wide. 

Ancash: Ocros, 3,200 meters, Weberbauer 2660, (171, type). 
Moquehua: Carumas, 3,200 meters, Weberbauer 7281 a. Lima: Ma- 
tucana, 2,400 meters, Weberbauer 5256. Between Matucana and 
Tambo de Viso, 2,400-2,700 meters, Weberbauer 5209. 

3. URBANODOXA Muschl. 

Slender annuals, with opposite lower leaves. Racemes mostly 
terminal and soon lax. Petals distinctly clawed. Glands at the base 
of the filaments conspicuous, the medial ones absent. Pod valves 
laterally compressed, not at all margined, broadly obovate, rugulose. 
A weak segregate from Cremolobus, which also may have opposite 
leaves and large glands, but in the petals and pods there is a possibly 
fundamental difference. 

Urbanodoxa rhomboidea (Hook.) Muschl. Bot. Jahrb. 40: 271. 
1908. Cremolobus rhomboideus Hook. Icon. 1 : pi. 32. 1837. 

Glabrous; leaves rhombic-ovate, angulate-serrate, narrowed to 
a petiole 0.5-1 mm. long, the blades 3-4 cm. long, 1.5-2.5 cm. wide; 
flowers 4-5 mm. broad; sepals lanceolate, 2-2.5 mm. long; petals 
obovate-cuneate, 3-4.5 mm. long; pedicels 2.5-3 cm. long, 3-4 times 
longer than the pods. Illustrated, Engl. & Prantl, Pflanzenfam. 
ed. 2. 17b: 315. Neg. 27328. 



FLORA OF PERU 945 

Ancash: Hacienda Cajabamba, 3,000 meters, Weberbauer 3130. 
Lima: Cuesta de Puruchuco (Mathews 1061, type). Viso, 2,700 
meters, 572. Moquehua: Carumas, 3,200 meters, Weberbauer 7296. 

4. BRASSICAL. 

In this genus, which includes cabbage (col), mustard (mostaza), 
etc., several species are to be expected as weeds. They are all char- 
acterized by yellow flowers, hispid pubescence, or none, and slender 
pods tipped by a beaklike style. 

Brassica campestris L. Sp. PI. 666. 1753. 

Cauline leaves clasping; flowers exceeding the buds; pods erect, 
3.8-8 cm. long, with a conic beak one-third as long or longer. The 
related B. Napus L. (nabo) bears the opened flowers lower than the 
buds. Herrera lists as cultivated in Peru B. oleracea L., cabbage 
(col, or repollo), and its races, cauliflower (coliflor), and Brussels 
sprouts (colinabo). 

Cuzco: Frequent in fields (Herrera). Lima: Lima, along street, 
150 meters, 71. Junin: La Oroya, 3,600 meters, 969. La Quinua, 
3,600 meters, 2012. Loreto: Lower Rio Huallaga, 155-210 meters, 
Williams 4066. Huanuco: Chavinillo, 2,250 meters, 1979. Wheat 
field, Huanuco, 2,100 meters, 2367. Native of the Old World. 

Brassica nigra (L.) Koch in Rohling, Deutschl. Fl. ed. 3. 4: 713. 
1833. Sinapis nigra L. Sp. PI. 668. 1753. 

Similar to B. campestris, but the flowers borne below the buds, 
the cauline leaves petioled, not clasping, and the pods only 1-2 cm. 
long, with a beak 1.5-2.5 mm. long. 

Lima: Along railroad and near a farm, 2,400 meters, 241. 
Loreto: Lower Rio Huallaga, 155-210 meters, Williams 4488. 
Europe and Asia. "Mostaza." 

Brassica juncea (L.) Cosson, Bull. Soc. Bot. France 6: 609. 1859. 
Sinapis juncea L. Sp. PI. 668. 1753. 

Glabrous, somewhat glaucous, the leaves not clasping, narrowed 
at the base, the upper ones oblong and subentire; pods 3.5 cm. long, 
the beak about one-third as long. 

Loreto: Lower Rio Huallaga, 155-210 meters, Williams 4076. 
Iquitos, 120 meters, Williams 3587. San Martin: San Roque, 1,350- 
1,500 meters, Williams 7089. Zepelacio, near Moyobamba, 1,200- 
1,600 meters, Klug 3309. Native of the Old World. 



946 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5. RAPHANUS L. 

This well known genus scarcely needs description here, except 
to note that it is marked by its definitely biarticulate pods, the 
upper segment always fertile, and by its depressed-capitate stigma. 
In general, too, it may be known by its coarse, erect habit and 
rather showy, white or purplish flowers. 

Pods slender, to 4 mm. thick; petals white or purplish, with violet 
veins R. Raphanistrum. 

Pods stout, much thicker; petals often purplish, with yellowish or 
obscurely violet veins R. sativus. 

Raphanus Raphanistrum L. Sp. PI. 669. 1753. 

Annual; pods very fragile, 3.5-4 mm. thick, the beak 6-20 mm. 
long; ovules 4-11. There are many named forms, the following 
collection belonging to f. carneus (Schweigg. & Koerte) Thell. (var. 
purpurascens Dum.). In 1922 the plant colored purple the grain 
fields about Huanuco. 

Cajamarca: Celendin, Woytkowski 8. Huanuco: Near Huanuco, 
2,100 meters, 2368. Without locality: Escomel Native of the Old 
World. 

Raphanus sativus L. Sp. PI. 669. 1753. 

Annual, the pods at least 8 mm. thick, the beak conic; ovules 
10-12; petals rarely to 22 mm. long and the pods to 9 cm. long. 
The radish (r&bano) of cultivation, and wherever cultivated, an 
occasional escape. 

Lima: Matucana, 2,400 meters, 172. Cuzco: Hacienda Tancac 
(Herrera). Native of the Old World. 

6. LEPIDIUM L. Peppergrass 

Reference: Thellung, Mitt. Bot. Mus. Zurich 28: 1-340. 1906. 
Annuals or perennials, sometimes suffruticose toward the base, 
the basal, rosulate leaves evanescent. Pubescence always simple. 
Leaves often pinnately parted, but various. Flowers white, in 
elongating racemes, sometimes showy, but often not, the petals 
even rudimentary. Glands 4-6. Pods laterally compressed, dehis- 
cent, the septum lanceolate to elliptic, emarginate and often more or 
less winged, ovate to orbicular. Nearly all species vary at times 
and are difficult to separate; the key does not presume to provide 
for all variations, but perhaps holds for many of them. 



FLORA OF PERU 947 

Style at least equaling the emarginate pod apex; middle cauline 

leaves simply incised to entire, not auriculate at the base. 
Petals longer than the calyx; pods to 5.5 mm. long, 3-5 mm. wide. 
Calyx 2-3 mm. long, fully equaling the pod margins. .L. Meyenii. 
Calyx 1.5 mm. long; style scarcely equaling the pod margins. 

L. Weddellii. 
Petals and calyx subequal; pods suborbicular, to 4.5 mm. long and 

broad L. cyclocarpum. 

Petals often shorter than the calyx; pods to 3.5 mm. long, 2.5 mm. 

wide L. Walpersii. 

Style much shorter than the pod margins; cauline leaves various, but 

often divided or auriculate at the base. 
Cauline leaves attenuate to the base, not dilated-auriculate. 
Plants annual or biennial; cauline leaves more or less serrate 
or pinnatifid. 

Pods 2.5-3 mm. wide; leaves serrate L. virginicum. 

Pods 4 mm. wide; leaves pinnatifid L. Raimondii. 

Plants perennial, subligneous below; leaves all more or less 

divided L. abrotanifolium. 

Cauline leaves evidently dilated or auriculate at the base (see a 

form of L. aletes} . 
Annuals or short-lived perennials; pods rarely as narrow as 

2.5 mm.; cauline leaves sublanceolate. 
Petals reduced; glands half (or less) as long as the calyx. 

L. subvaginatum. 
Petals and calyx subequal; glands only one-sixth as long as 

the calyx L. aletes. 

Perennials with some leaf remains about the caudex; pods 
rarely, or barely, 2.5 mm. wide; cauline leaves typically 
subobovate. 
Sepals persistent; leaves mostly pinnatifid. 

L. bipinnatifidum. 
Sepals caducous; upper leaves, at least, mostly undivided. 

L. Chichicara. 

Lepidium abrotanifolium Turcz. Bull. Soc. Nat. Mosc. 27: 
308. 1854; 246. 

Root fleshy, the crown clothed with old leaf fibers; stems many, 
subvelutinous-pubescent above, 5-10 cm. long; basal leaves pin- 



948 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nately parted, long-petioled, the sessile cauline ones less divided or 
only pinnate-incised; petals oblong, about equaling the calyx, or 
setaceous and much shorter; calyx 1 mm. long; pedicels slightly 
compressed, pubescent; pods narrowly obovate, shallowly or rather 
deeply emarginate, 3.5 mm. long, 2-2.5 mm. wide. Var. Steinmanni 
Thell. of southern Bolivia (to be expected) has nearly elliptic pods, 
3 by 2.25 mm., and a promptly deciduous calyx. L. pubescens Desv., 
reported by Thellung from Peru on the basis of a Dombey specimen 
without locality, is scarcely woody, hirsutulous, and has minutely 
ciliate pods 4 mm. long and 3.5-3.7 mm. wide. The Dombey speci- 
men, as so often with his material labeled merely "Peru," is doubt- 
less Chilean, since the species otherwise is unknown outside Chile 
and California. Neg. 27314. 

Lima: Chosica, 2,400 meters, 484- Viso, 2,700 meters, 569. 
Near Matucana, Weberbauer 160, 178, 169, 180. Junin: La Oroya, 
3,700 meters, Weberbauer 2567. Bolivia; Ecuador. 

Lepidium aletes Macbr. Candollea 5: 357. 1934. L. calycinum 
Godron, Me"m. Acad. Montpel. 1: 416. 1853, non Steph. 1800. 

Not unlike the preceding, but the stems often many, the pubes- 
cence below subappressed, and the leaf rachis less dilated; calyx to 
1 mm. long, the petals setaceous, about as long; pods broadly obovate 
or elliptic, distinctly emarginate, 3 mm. long, 2.25-2.75 mm. broad. 
Here Thellung has referred as a variety a depauperate plant with 
the leaf base not dilated, the rachis linear, L. calycinum var. gracile 
(Chod. & Hassl.) Thell.; to it Schulz has assigned a collection made 
in Peru. 

Arequipa: Pampa de Arrieros, 3,750 meters, Pennell 13330. 
Tacna: Candarave, 3,000 meters, Weberbauer 7380 (the variety). 
Southern South America; adventive in France. 

Lepidium bipinnatifidum Desv. Journ. Bot. 3: 165, 177. 1814; 
241. L. Humboldtii DC. Syst. 2: 532. 1821. 

Often perennial, the arcuately ascending stems 10-30 cm. high, 
hirsutulous with slender, spreading hairs; cauline leaves sometimes 
5 cm. long and 3 cm. wide, strongly dilated at the base and auricu- 
late-amplexicaul; sepals 0.75 mm. long, half longer than the linear 
petals and much longer than the conspicuous glands; pods obovate, 
broadly emarginate, 2.5-3 mm. long, 1.75-2.25 mm. broad, equaling 
or shorter than the angled, pubescent pedicels. Employed in popular 
medicine as a styptic (Herrera). Neg. 34547. 



FLORA OF PERU 949 

Puno: Altos de Toledo, 3,000 meters, Meyen; Raimondi. Are- 
quipa: Pampa de Arrieros, 3,750 meters, Pennell 13331. Huanuco: 
A dooryard weed, Huanuco, 2,700 meters, 1782. Cochero, Poeppig 
1687. Cuzco: About Cuzco (Herrera). Lima: Chicla (Ball). 
Cheuchin, Ruiz & Pavon. Obrajillo (Wilkes Expedition). Without 
locality (Dombey, type). Cajamarca: Tambillo, Jelski 234- Cutervo 
and Nancho, Raimondi. Bolivia to Colombia. "Chichicara," "mosta- 
cilla," "chicchi," "mayo-mostaza," "chichira," "anuccara." 

Lepidium Chichicara Desv. Journ. Bot. 3: 165, 179. 1814; 220. 

With much the appearance and habit of the preceding, but the 
upper leaves often merely incisely serrate, and the pods rather ellip- 
tic than obovate and slightly angulate-emarginate, equaled by the 
pedicels, mostly about 3 mm. long and over 2 (2.5) mm. wide. 
See also L. Walpersii. Often a weed in waste places. Neg. 34549. 

Puno: Meyen. Arequipa: North of Arequipa, 2,400 meters, 
Hinkley & Hinkley 62. Lima: Viso, Weberbauer 140. Matucana, 
2,400 meters, 437, 483 (glabrescent form), 659, 660. Chicla (Ball). 
Rio Blanco, 3,000-3,500 meters, Killip & Smith 21544 (det. Schulz). 
Cajamarca: Cutervo (Jelski 235, in part). Bolivia. "Chichicara," 
"chichiaccara." 

Lepidium cyclocarpum Thell. Mitt. Bot. Mus. Zurich 28: 
214. 1906. 

A slender, somewhat branched, erect annual, sparsely pubescent 
with spreading trichomes; basal leaves unknown; lower cauline leaves 
long-petiolate, irregularly dentate, the upper ones oblong-oblanceo- 
late, attenuate to the sessile base, remotely dentate; petals narrowly 
spatulate; sepals 1.3 mm. long; pedicels nearly filiform, to 1.5 times 
longer than the rotund, shortly emarginate pod. The Colombian 
and Bolivian L. Trianae Thell. is perennial, with pods only 2.5 mm. 
broad. Negs. 13857, 30124. 

Lima: Amancaes Hill, Weberbauer 1614,' (Mathews 752, type). 

Lepidium Meyenii Walp. Nov. Act. Acad. Leop.-Carol. 19: 
Suppl. 1:249. 1843; 202. L. gelidum Wedd. Ann. Sci. Nat. V. 1: 
283. 1864. 

A matlike perennial, the mostly decumbent stems sparsely 
branched and seldom more than a few centimeters long; basal leaves 
densely rosulate, vaginately dilated at the base of the long petiole, 
mostly pinnate-parted, 2-3 cm. long and 5 mm. wide; cauline leaves 
reduced, entire, lobed or deeply crenate; sepals 2-3 mm. long, usu- 



950 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ally caducous; petals longer than the sepals, sessile; lower pedicels 
often twice as long as the more or less roundish (variable), slightly 
emarginate pod, its apical margins barely equaling or shorter than 
the style. The mats are formed by the persistent, crowded petioles. 
Highlands, at 3,000^,500 meters. Subsp. gelidum (Wedd.) Thell. is 
the typical form. Negs. 13859, 34550. 

Puno: Pisacoma, Meyen 33, type. Chuquibambilla, Pennell 
13396. Moquehua: Carumas, near Volcan Ticsani, 4,000 meters, 
Weberbauer 7321. Cordillera above Torata, 4,100 meters, Weber- 
bauer 7473. 

Lepidium Raimondii O. E. Schulz, Notizbl. Bot. Gart. Berlin 
10: 727. 1929. 

Annual, branching from the base, short-pilose with curved tri- 
chomes, even on the pedicels, these to 4 mm. long, spreading; lower 
cauline leaves unknown, the upper pinnatifid, with 4-5 linear, entire 
or nearly entire divisions, ciliate, broadly sessile but not auriculate; 
fruiting racemes 30-60-flowered; pods suborbicular, 4-5 mm. long, 
rounded at the base, the short style included in the deeply emar- 
ginate apex. Certainly, according to the author, related to L. vir- 
ginicum, but the pods larger and the leaves dissected. 

Cajamarca: Cascas, 1,800 meters, Raimondi 7538, type. 

Lepidium subvaginatum Thell. Mitt. Bot. Mus. Zurich 28: 
249. 1906. L. subvaginatum Steud. Nomencl. ed. 2. 2: 28. 1841, 
nomen. 

A perennial, or becoming so, with usually a solitary stem, some- 
times corymbosely branched, the pubescence widely spreading, the 
trichomes slender; basal leaves bipinnate or tripinnate, the cauline 
less divided, the lower half of the rachis strongly dilated, sheathlike, 
the base itself cordate- or sagittate-auriculate; sepals usually persis- 
tent, less than 1 mm. long; petals rudimentary, filiform; pods ovate, 
deeply emarginate, to 3.5 mm. long, 2.5 mm. wide. 

Cajamarca: Cutervo, Raimondi. Chile. 

Lepidium virginicum L. Sp. PI. 645. 1753; 222. 

Stems mostly solitary, corymbosely branched above, the pubes- 
cence arcuately subappressed or nearly lacking; basal leaves lyrate- 
pinnatifid; sepals caducous, 0.75 mm. long, the petals usually about 
twice as long; pedicels slender, often half again as long as the sub- 
orbicular, broadly emarginate pods, these 3-4 mm. long, 2.5-3 mm. 
wide. A widely distributed or adventive, variable species, the Peru- 
vian form with pedicels and obovate or elliptic pods subequal, the 



FLORA OF PERU 951 

petioles shorter than the sepals, probably a variety (subsp. centrali- 
americanum Thell. Mitt. Bot. Mus. Zurich 28: 225. 1906). 

Lima: Gravelly stream flat, 2,400 meters, 140; Ruiz & Pav6n 
Moquehua: Carumas, 3,100 meters, Weberbauer 7312. Cajamarca: 
Cutervo, Raimondi. The variety ranging to Mexico; Venezuela. 

Lepidium Walpersii Macbr. Candollea 5: 357. 1934. L. lance- 
olatum Walp. Nov. Act. Acad. Leop.-Carol. 19: Suppl. 1: 250. 1843, 
non Presl, 1826. L. Chichicara Desv. var. lanceolatum Thell. Mitt. 
Bot. Mus. Zurich 28: 221. 1906. 

Allied, perhaps, toL. Chichicara, but smaller and notably different 
in its merely sessile cauline leaves, not at all auriculate and often 
subentire, and in its somewhat smaller pods, the style quite equaling 
the pod margins. L. depressum Thell., 201, Bolivian, has similar pods 
but the style is exserted and the leaves pinnatifid; L. quitense Turcz., 
213, is tall, strict, with entire, linear leaves, or these pinnate, with 
remote, linear lobes. 

Puno: Lake Titicaca, Meyen, type. Pisacoma, Meyen, less typi- 
cal. Bolivia. 

Lepidium Weddellii 0. E. Schulz, Notizbl. Bot. Gart. Berlin 
11: 391. 1932. L. affine Wedd. Ann. Sci. Nat. V. 1: 284. 1864, non 
Ledeb. L. Meyenii Walp. subsp. affine Thell. Mitt. Bot. Mus. 
Zurich 28: 204. 1906. 

A depauperate, depressed plant, nearly L. Meyenii but with 
smaller calyx and the style obsolete, merely equaling the apical mar- 
gins of the orbicular pod. See Macbride, Candollea 5: 357. 1934. 

Ancash : Pelagatos Mountain, 4,400 meters, Weberbauer 7232 (det. 
Schulz). Bolivia. 

7. ISATIS L. Woad 

A strict annual, or more enduring, with narrow, hastate-based, 
entire leaves, and with bractless, at first crowded racemes of rather 
small, yellow flowers. Style none, the stigma large. Pods usually 
1-seeded, flat, broadly winged, the pedicels capillary toward the base. 

Isatis tinctoria L. Sp. PI. 670. 1753. 

Plants several decimeters high, often densely leafy below, the 
narrowly oblong, sessile, obtuse leaves several centimeters long; 
petals 3.5 mm. long; pods 1.5 cm. long, 4 mm. wide, oblong-elliptic. 
Native of Europe and formerly cultivated as a dye plant, woad 
has become widely established. 

Arequipa: Rocky river bank, Tiabaya, 2,100-2,200 meters, 
Pennell 13087. 



952 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

8. CAPSELLA Medic. Shepherd's purse 

In general, similar to Thlaspi, but the petals obovate, the seeds 
often more numerous, and the plants somewhat pubescent. 

Capsella Bursa-pastoris (L.) Medic. Pflanzengatt. 1: 85. 1792. 
Thlaspi Bursa-pastoris L. Sp. PL 647. 1753. 

A simple or sometimes branched annual, with a rosette of some- 
what pinnate-dentate leaves, and clasping, lanceolate cauline ones, 
these usually 2-3 cm. long; flowers small, white; pods 5-8 mm. long, 
obcompressed, obtriangular. Usually in cultivated or waste ground. 
Illustrated, Fawc. & Rendle, Fl. Jamaica 3, pt. 1:241. Abundant 
throughout the Department of Cuzco; leaves used as poultices for 
bruises (Herrera}. 

Cuzco: Valle del Huatanai, Oropesa to Cuzco (Herrera). Lima: 
Amancaes and Lima, Martinet.- Junin: Chanchamayo Valley, 1,500 
meters, Schunke 483. Puno: Orurillo, Soukup 541- Native of the 
Old World, but widely naturalized as a weed in America. 

9. MANCOAWedd. 

Low, spreading, much branched, hispid-stellate or glabrous herbs 
with white, corymbose, ebracteate flowers. Sepals equal, persistent. 
Stamens glabrous. Pods elliptic, subvesicular, the valves almost 
rounded dorsally, the septum thin, the cells many-seeded. Coty- 
ledons incumbent. 

Mancoa hispida Wedd. Ann. Sci. Nat. V. 1: 285. 1864. 

Hispid throughout; leaves oblong-lanceolate, coarsely serrate or 
subpinnatifid, rather obtuse; sepals broadly elliptic; pods stellate- 
hispid. M. laevis Wedd., of Bolivia, is glabrous, the leaves lyrate- 
pinnatifid. M. foliosa (Wedd.) 0. E. Schulz has pinnate leaves, but 
only the stems are pubescent. Both these species of northern Bolivia 
are to be expected. Neg. 27313. 

Tacna: Werdermann 1131. Arequipa: Open, gravelly slopes, 
Nevada de Chachani, 3,500-3,600 meters, Pennell 13257. Prov. 
Condesuyos, 4,000 meters, Weberbauer 6850. Moquehua: Tola 
heath, cordillera above Torata, 3,800-3,900 meters, Weberbauer 7475. 
Moho, 3,900 meters, Weddell, type. Bolivia. 

10. CORONOPUS Zinn 

Annuals or short-lived perennials, branched from the base, gla- 
brous or usually more or less papillose-pubescent with simple tri- 
chomes. Leaves petiolate, oblong, more or less pinnate-dentate. 



FLORA OF PERU 953 

Flowers small, in short racemes that lengthen with the maturing 
of the pedicellate, didymous pods, each nutlike part of which is 
indehiscent. 

Coronopus didymus (L.) Smith, Fl. Brit. 2: 691. 1800-1804. 
Lepidium didymum L. Mant. 1 : 92. 1767. 

Leaves all pinnatifid ; pods at both top and bottom slightly emar- 
ginate, net- veined; stamens 2 (-6). The plant, crushed and applied 
hot to the affected parts, is used to clean and cure mat del voile; 
macerated with butter, it is placed on the abdomen in the treatment 
of swelling (Ruiz & Pavdn). 

Lima: Raimondi. Lurin, Ruiz & Pavdn. Lima, roadsides, 150 
meters, 69. Callao, 24 meters, 5886. Native of Europe; becoming 
cosmopolitan. "Pichiccara," "huanucara," "mastuerzo silvestre." 

11. DRABA L. 

Reference: 0. E. Schulz, Pflanzenreich IV. 105: 16-343. 1927. 

Small, rarely annual, often tufted or cespitose plants with scapose 
stems, or these sparsely leafy. Leaves simple, only the rosulate basal 
ones petioled. Flowers usually yellow or white. Nectary glands 
various. Pods short, usually ovoid or lanceolate, 2-celled, the lateral 
nerves of the septum anastomosing. A number of species probably 
growing in Peru but apparently not yet found there are mentioned, 
mostly under D. Pickeringii, but not all of them would key to the 
species with which for convenience they are associated. The speci- 
mens cited were, with few exceptions, determined by both Ekmann 
and Schulz. 

Annuals, the stems leafy below D. araboides. 

Perennials; stems scapose, or, if leafy, suffruticose. 
Stems scapose; plants cespitose or pulvinate. 

Pubescence mostly simple; scapes 10-30 cm. high. 

Pods obtuse; petals 4.5 mm. long; leaves to 2.5 cm. long. 

D. scopulorum. 

Pods acute; petals 6.5 mm. long; leaves to 4 cm. long. 

D. Schusteri. 

Pubescence mostly stellate; scapes shorter. 
Petals longer than the sepals; scapes obvious, even in flower; 

leaves narrowed at the base. 
Stem pubescence partly or entirely simple. 



954 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Fruiting racemes dense; petals 4 mm. long, white. 

D. Pickeringii. 
Fruiting racemes open; petals shorter, yellowish. 

D. Brackenridgei. 

Stems and leaves, at least the latter, stellate-tomentose. 
Petals 2.5 mm. long; pods densely hirsute. 

D. atacamensis. 
Petals 3^4 mm. long; pods glabrous or nearly so. 

D. Macleanii. 

Petals shorter than or equaling the sepals; scapes often obso- 
lete and in fruit barely rising above the leaves, these 
often minute, with a petiolar base. 

Sepals 2.5 mm. long; scapes, if obvious, not floccose-tomen- 
tose. 

Stems stellate-tomentose; sepals 2.5 mm. long; petals 
2 mm. long D. argentea. 

Stems villous; sepals and petals 2.5 mm. long. 

D. cryptantha. 

Sepals 2 mm. long; scapes flocculose-tomentose. 

Petals shorter than the sepals; style obsolete. 

D. chlorantha. 

Petals and sepals equal; style manifest. . D. alchemilloides. 
Stems leafy (sometimes only at the base) but suffruticose. 

Leaves cinereous with a dense, stellate pubescence. 

D. matthioloides. 
Leaves greenish, or the pubescence in part simple. 

Plants sparsely and minutely stellate-pubescent . D. peruviana. 
Plants with pubescence 'of stellate and simple hairs. 
Plants suffruticose, not tufted, leafy. 
Pubescence mostly stellate. 

Petals yellow; branches many D. ochropetala. 

Petals ochroleucous; branches few or none.Z). solitaria. 

Pubescence, especially on the leaves above, simple. 

D. alyssoides. 

Plants tufted, with crowded, scaly caudex branches. 

D. cuzcoensis. 



FLORA OF PERU 955 

Draba alchemilloides Gilg, Bot. Jahrb. 42: 473. 1909; 143. 

A pulvinate perennial, the leaves densely rosulate at the tips 
of the branches; blades obovate-oblong, nearly rotund at the apex, 
gradually narrowed to the base, entire, 4-6 mm. long, about 3 mm. 
wide, white- tomentose with short, stellate hairs; flowers yellowish, 
in a capitate corymb; peduncle and pedicels (2-3 mm. long) densely 
stellate-pubescent; sepals oblong, rounded at the apex, stellate- 
pilose only outside, the clawed petals scarcely longer; pod ovoid, 
3-4 mm. long, 1.5-1.8 mm. wide, densely stellate- tomentose; style 
slender, 0.4-0.75 mm. long. According to the original description, 
the leaves are sometimes longer, the style very short and fleshy. 
The above diagnosis (as for all other species) is from 0. E. Schulz. 
Neg. 13894. 

Ancash: Above Piscapacha, 4,600 meters, Weberbauer 2902; 
224 . Huanuco : ( Pearce) . 

Draba alyssoides Humb. & Bonpl. ex DC. Syst. 2: 355. 
1821; 148. 

Stems ordinarily simple, 10-30 cm. high, more or less ligneous 
below and densely leafy, closely hirsute, even on the sepals, with 
spreading, simple trichomes to 1 mm. long and shorter, stipitate, 
furcate ones; leaves entire or with 1-2 teeth on each side, narrowed 
to the base, sessile, the midnerve strongly thickened at the base, 
7-12 mm. long, with stiff, simple trichomes on both sides, beneath 
also with shorter, stellate ones; racemes elongating, to 30-flowered, 
bracteate below; pedicels to 10 mm. long; sepals 4 mm. long, not 
saccate, rounded at the apex, broadly oblong-elliptic, the inner 
slightly larger; petals white, sparsely veined, 7 mm. long, emarginate; 
filaments somewhat dilated at the base; ovules 16-24; style 1-4 
mm. long; pods 8-11 mm. long, 4-5 mm. wide, hirsute with mostly 
simple (a few bifurcate) hairs; seeds brown, 1.2-1.5 mm. long. 
Var. thlaspiformis (Gilg & Brandt) 0. E. Schulz, 150, is smaller, 
branching, the 12-ovulate pods shorter and narrower. The species 
is typical of a group of closely related species, or varieties (?), and is 
therefore no doubt in Peru in one form or another. D. Hemsleyana 
Gilg, 150, is similar, but the pods are glabrous, about 10 mm. long, 
2 mm. wide, the style 2 mm. long; D. splendens Gilg, 150, also has 
narrow pods, but these, like the leaves on both sides, have simple 
pubescence; both are Ecuadorean. Neg. 13895. 

Peru: Probably. Ecuador; Colombia. 



956 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Draba araboides Wedd. Ann. Sci. Nat. V. 1: 287. 1864; 334. 
D. soratensis Gilg & Muschi. Bot. Jahrb. 42: 480. 1909, pro parte, 
non Wedd. 

Plants 10 cm. high in fruit, the slender, erect stems sometimes 
branched from the base, leafy below and more or less hispid, the 
crowded leaves rather obovate-cuneate, with 2-4 small teeth on 
each side; pedicels 2-4 mm. long; flowers heteromorphous, the 
larger, perfect ones with sepals 1.8 mm. and petals 3 mm. long, the 
smaller ones with 4 minute petals, the sepals 1.5 mm. long; pods 
broadly linear, to 1 cm. long. The larger leaves are about 2 cm. 
long, and 1 cm. wide above the middle. Neg. 34581. 

Lima: Mt. San Agustin, near Lima, 300-400 meters, Weberbauer 
5244- Matucana, 2,400 meters, Weberbauer 5254- Cuzco: Without 
locality, Gay, type. 

Draba argentea 0. E. Schulz, Pflanzenreich IV. 105: 134. 1927. 

Similar to D. Pickeringii, but only 1-2 cm. high, even in fruit, 
the pedicels then to 3 mm. long; leaves 3-6 mm. long, white- tomen- 
tose; sepals oblong; petals 1-nerved; pods capitately crowded, 
oblong-ellipsoid, 3-4 mm. long, 2 mm. wide, more or less densely 
pubescent with furcate and substellate hairs. Neg. 13896. 

Junin: Alpamina, 4,500 meters, Weberbauer 5114, type. Pace- 
chac, 4,600 meters (Hill 13). 

Draba atacamensis Gilg, Bot. Jahrb. 42: 475. 1909; 134. 

Densely pulvinate, the stems only 1-1.5 cm. long; leaves obovate- 
oblong, 12-16 mm. long, densely rosulate, white-tomentose with 
minute, substellate hairs, but often toward the base also with some 
simple hairs; sepals 1.8 mm. long; petals 2.5 mm. long; ovules 32; 
pedicels 1-3 mm. long, the oblong-ellipsoid pods 6 mm. long, 2.5 
mm. wide, densely hirsute with bifurcate and substellate hairs. 
Illustrated, Pflanzenreich IV. 105: 133. 

Tacna: Volcan Tacora, 4,500 meters, Werdermann 1509 (det. 
Schulz). Chile. 

Draba Brackenridgei Gray, U. S. Expl. Exped. Bot. 53. 1854; 
136. D. cephalantha Gilg, Bot. Jahrb. 42: 477. 1909. 

Mature plants more than 10 cm. high; leaves densely rosulate, 
ovate or ovate-oblong, acute, entire, 5-7 (-12) mm. long, 3-5.5 mm. 
wide, more or less densely stellate-tomentose beneath, above with 
rather long and mostly simple hairs; scape 2-3 (-8) cm. high, the 
pedicels finally to 3 mm. long; sepals oblong, about 2.5 mm. long, 



FLORA OF PERU 957 

green, the pubescence partly furcate; petals clawed, obovate, retuse; 
pods oblong-lanceolate, 7-9 mm. long, scarcely 2 mm. wide, pubes- 
cent with simple and furcate hairs; style less than 1 mm. long; seeds 
0.75 mm. long. The fully mature pods of my No. 1054 are 8-9 mm. 
long, those of cotype material at Paris 6-7 mm. long. The species 
does not seem to be sharply separated from D. Macleanii. 

Junin: In limestone rocks between La Oroya and Tarma, 4,000 
meters, Weberbauer 2551; 223; type of D. cephalantha. La Oroya, 
4,300 meters, Weberbauer 2604- Tarma, limestone ledges, 3,900 
meters, 1054- Banos and Obrajillo (Wilkes Expedition, type). 

Draba chlorantha 0. E. Schulz, Pflanzenreich IV. 105: 143. 
1927. 

Closely allied to D. alchemilloides, but only 1.5-3 cm. high, 
the scapes scarcely 1 cm. long; petals 1.2-1.5 mm. long, pale greenish 
yellow; style very short. Neg. 13898. 

Ancash: On limestone, Mt. Huacchara, 4,600 meters, Weberbauer 
7245, type. Ayacucho: Mt. Rasuvilca,4,500 meters, Weberbauer 7497, 
in large part (det. Ekmann and Schulz). 

Draba cryptantha Hook. f. Fl. Ant. 2: 234. 1847; 142. D. 
Weberbaueri Gilg & Muschl. Bot. Jahrb. 42: 474. 1909. 

Pulvinate, the leaves densely rosulate at the tips of the very 
short branches; blades obovate, acutish, gradually attenuate to the 
base, entire, 3-4 mm. long, 2-3 mm. wide, above white-pilose- 
tomentose, beneath white-tomentose with rather short, stellate 
hairs, like the pedicels, sepals, and pods; corymb dense, subcapitate, 
scarcely exceeding the leaves; sepals obovate-oblong; pods ovoid, 
3 mm. long, 1.5-2 mm. broad, the style very short; ovary 9-ovulate. 
D. Weberbaueri, according to description, has stellate-pubescent 
leaves, sepals only 1.5 mm. long, and yellowish, very long-clawed 
petals. The plant may be diminutive, the peduncle barely exserted 
from the leaves even in mature fruit. 

Junin: Above Lima, 4,500 meters, Weberbauer 5190; 221 (type 
of D. Weberbaueri). Culnai to Casacancha (Wilkes Expedition). 
Pacechac (Hill 12). Cerro de Pasco, Mathews 942. Ayacucho: 
Mt. Rasuvilca, 4,500 meters, Weberbauer 7497 (in part; det. Ekman 
and Schulz). Cuzco: Cordillera de Pachahusan, 4,400 meters, 
Herrera 2577b. 

Draba cryptantha var. laxistellata (Gilg & Brandt) 0. E. 
Schulz, Pflanzenreich IV. 105: 143. 1927. D. laxistellata Gilg & 
Brandt ex 0. E. Schulz, loc. cit. 



958 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Fruiting scapes 2 cm. long; leaves 6-9 mm. long; ovary 16- 
ovulate; pedicels to 3 mm. long; pods 4 mm. long, 2 mm. broad, 
sparsely pilose; style 0.5 mm. long. Probably a distinct species. 

Ayacucho: Santa Ine"s, 4,600 meters, Weberbauer 5428, type. 
Junin: Huancayo, 4,700 meters, Kittip & Smith 22043. 

Draba cuzcoensis 0. E. Schulz, Kew Bull. 490. 1936. 

Perennial, from a branching root, the caudex branches crowded, 
short, densely scaly with the persistent petioles of old leaves, in 
flower 3-7 cm. high, in fruit to 12 cm.; upper leaves forming dense, 
cespitose rosettes 1.5-4 cm. wide; stems 1-3, simple, sparsely leafy, 
hirsutulous, even to the sepals, with stipitate, bifurcate and branched 
hairs or rarely some of the hairs simple; basal leaves oblong-obovate, 
subobtuse, entire, narrowed to the petiole, to 5 mm. wide, 1-2.5 cm. 
long; cauline leaves sessile, smaller, ovate or linear, all hispid and 
ciliate with simple and branched hairs; racemes 10-20-flowered, 
elongating, in fruit with pedicels to 8 mm. long; sepals 2 mm. long, 
rounded, hyaline-margined; petals yellow, 3.2 mm. long, minutely 
emarginate; nectary glands binate; ovules 18; pods oblong-ellipsoid, 
compressed, 6-9 mm. long, 3 mm. broad, acute, the style 5 mm. 
long, glabrous, dull green or black-violet; seeds 1 mm. long, 0.75 
mm. broad, tuberculate. 

Cuzco: Near Cuzco, 4,000 meters (Dora B. Stafford 264, type). 

Draba Macleanii Hook. f. FL Ant. 2: 235. 1847; 135. D. 
siliquosa Ball, Journ. Linn. Soc. Bot. 22: 29. 1885, non Hook. f. 

Cespitose, in fruit to 6 cm. high, the elongating racemes 12-15- 
flowered; leaves rosulate, oblong or narrowly obovate, obtuse, entire, 
scarcely petioled, 8-12 mm. long; pedicels to 4 mm. long in fruit; 
sepals 2.5 mm. long; petals 3-4 mm. long; ovary 16-20-ovulate; 
pods oblong or subellipsoid, 6-7 mm. long, 1-2 mm. wide, the style 
0.5 mm. long. Var. stylosa Gray, U. S. Expl. Exped. Bot. 53. 1854, 
has laxer corymbs and finally glabrous pods with a longer style. 

Junin: Yauli, on igneous rock, 4,400 meters, Weberbauer 269. 
Rio Blanco, 5,000 meters, 3033. Lima: Casapalca (Ball). Moque- 
hua: Tacora, Weddell. Without locality (Maclean). Bolivia. 

Draba matthioloides Gilg & Schulz, Pflanzenreich IV. 105: 
151. 1927. 

A small, suffrutescent plant, a few decimeters high, much 
branched at the base; stems very leafy above, very densely and 
shortly white-stellate-tomentose, like the sepals, these 5-7 mm. 



FLORA OF PERU 959 

long, broadly lanceolate; leaves lance-oblong, acutish, entire, sessile, 
1.5-2.5 cm. long; pedicels filiform, to 1.5 cm. long, or longer in fruit; 
petals 7-9 mm. long, orange or reddish; ovules 28; style slender, 
4-5 mm. long; pods ellipsoid or lanceolate, 8-15 mm. long, 3^4 mm. 
wide, stellate-scabrous. Probably only a variety or form of D. 
alyssoides. 

Ancash: Between Cajamarca and Hualgayoc, 4,200 meters, 
Weberbauer 4231, type; 272. On limestone, Prov. Pallasca, 4,200 
meters, Weberbauer 7019. 

Draba ochropetala 0. E. Schulz, Pflanzenreich IV. 105: 154. 
1927. 

Suffruticose, to 20 cm. high, the many ascending, leafy stems 
minutely cinereous-stellate-pubescent; leaves oblong or narrowly 
ovate, subobtuse, sometimes obscurely dentate, sessile, 7-10 mm. 
long, white- tomentose with simple and furcate hairs; racemes 
scarcely elongating, bracteate, 5-12-flowered; pedicels 2-4 mm. 
long; sepals 4 mm. long, substellate- tomentose; petals 6 mm. long; 
ovules 18-24; pods broadly ovoid, toward the margins short-pubes- 
cent with simple hairs, the style 1-1.5 mm. long. Neg. 13913. 

Ancash: Mt. Huacchara, on limestone, 4,600 meters, Weberbauer 
7233, type. 

Draba peruviana (DC.) 0. E. Schulz, Pflanzenreich IV. 105: 
151. /. 17. 1927. Berteroa peruviana DC. Syst. 2: 293. 1821. D. 
Pavonii Gilg, Bot. Jahrb. 42: 479. 1909. 

Suffrutescent, to about 30 cm. high, the suberect branches, as 
in the related species, clothed with the remains of the crowded 
leaves; leaves oblong-lanceolate, acute, narrowed to the base, 2-4- 
dentate, 10-16 mm. long, 3-4 mm. wide, laxly and minutely stellate- 
pubescent; flowers in dense, subcorymbose racemes at the tips of 
the leafy-bracteate branchlets; pedicels 7-10 mm. long; sepals nearly 
glabrous, 6 mm. long; petals 8-10 mm. long; pods sparsely hispid, 
5-6 mm. long, 2.5-3.5 mm. wide, the style 2.5-3 mm. long. 

Huanuco: Muna and Tambo Nuevo, Ruiz & Pavon, type; 
3,800-3,900 meters, Weberbauer 6727. Tambo de Vaca, 3,900 
meters, 4440. Argentina. 

Draba Pickeringii Gray, U. S. Expl. Exped. Bot. 54. 1854; 134. 

Cespitose, with many rosulate, crowded, obovate-cuneate leaves 
at the tips of the slender stems; leaves entire, 6-8 mm. long, the 
dense pubescence above mostly simple, that beneath stellate-tomen- 



930 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

tose, at the petiolar base simply ciliate; racemes little elongating, 
ebracteate, 10-25-flowered ; pedicels to 6 mm. long; sepals 2.8 mm. 
long, stellate-pubescent; pods broadly ovoid, 4-6 mm. long, 2.5-3 
mm. broad, pubescent with simple and furcate hairs, the style 
slender, 0.5 mm. long. Var. Pearcei 0. E. Schulz (as yet known 
only from Bolivia) has densely furcate-stellate-pubescent pods. D. 
Hookeri Walp., D. affinis Hook, f., and D. atacamensis Gilg are 
closely related species to be expected, especially the last two, the 
former from both Bolivia and Ecuador and the latter from northern 
Chile. They all have shorter petals (to 2.5 mm.) and shorter pedicels. 
D. Hookeri, of Ecuador, has pods barely 1 mm. wide, typically 
glabrous or the hairs in part simple. D. affinis and D. atacamensis 
have pods 1.5-2.5 mm. broad, the former with mostly simple hairs, 
the latter with branched and stellate ones. Illustrated, Pflanzen- 
reich IV. 105: /. 12. 1927. A cotype in the Paris herbarium has 
mature pods 7 mm. long, 3.5 mm. wide, the style 1 mm. long. 

Junin: Yauli, in igneous rock, Weberbauer 269a; (195, 203, 221}. 
Huaron, on ledges, 4,200 meters, 1124- Lima: Casapalca, 5,000 
meters, loose, igneous slopes, 824- Between Culnai and Obrajillo 
(Wilkes Expedition, type). 

Draba Schuster! 0. E. Schulz, Pflanzenreich IV. 105: 137. 1927. 

Stems many, in fruit to 30 cm. high, minutely cinereous-stellate- 
pubescent; leaves oblong-elliptic, obscurely callous-denticulate, 
acute, with the petiolar base 3-4 cm. long, more or less pubescent 
and ciliate with simple, lustrous hairs; racemes lax, to 15 cm. long, 
20-40-flowered; sepals 3 mm. long, the pubescence simple and 
furcate; pedicels in fruit to 1 cm. long or longer; pods 7-11 mm. 
long, 2.5-4 mm. broad, the filiform style 1.5 mm. long; ovules 12-16. 

Piura: Southeast of Piura, in grass steppe, 2,700 meters, Weber- 
bauer 6033, type. 

Draba scopulorum Wedd. Ann. Sci. Nat. V. 1: 286. 1864; 137. 

Except as indicated in the key, very like D. Schusteri, but smaller 
and the ovules 20-28. D. soratensis Wedd., of Bolivia, has petals 
3 mm. long, an oblong pod, and obsolete style. Neg. 13919. 

Cuzco: At 4,500 meters, Herrera 2312 (f. elongata Wedd.; fruiting 
pedicels to 2 cm. long). Bolivia. "Sanchalli." 

Draba solitaria 0. E. Schulz, Pflanzenreich IV. 105: 155. 1927. 

Closely allied to D. ochropetala but smaller, to 10 cm. high; 
leaves 5-8 mm. long, simply pilose-strigose above, stellate-tomentose 



FLORA OF PERU 961 

beneath; pedicels 1-2 mm. long; petals 5 mm. long; ovules 14-16; 
style 1 mm. long. Type collected 50 meters below a glacier, on 
soil detritus devoid of other plants. Neg. 13920. 

Ancash: Prov. Pallasca, north end of Cordillera de Pelagatos, 
4,450 meters, Weberbauer 7235, type. 

12. CARDAMINE L. 

Reference: 0. E. Schulz, Bot. Jahrb. 32: 280-623. 1903. 

Annual and often weak-stemmed, or rhizomatous and usually 
more enduring herbs with rarely a few simple trichomes. Leaves 
usually alternate and petiolate, sometimes auricled, simple or 
variously pinnatifid. Racemes corymbiform, mostly ebracteate. 
Sepals erect, spreading, the lateral ones sometimes cucullate at the 
base. Petals sessile, obovate. Nectar glands usually all present, 
the stamens 6, the oblong anthers sagittate at the base. Pod linear, 
attenuate into the often filiform style, the more or less bilobate 
stigma minute. Compare under Sisymbrium the cordate-leaved 
perennial, Ivania, if sought here. Species plastic, apparently not 
stabilized. 

Racemes usually more or less bracted; pods 1.3-3 cm. long; flowers 
2-4 mm. long. 

Pods usually 2-3 cm. long; seeds 1 mm. long C. bonariensis. 

Pods 13-16 mm. long; seeds 0.75 mm. long C. Killipii. 

Racemes ebracteate; pods 2.5-5 mm. long; flowers 3-8 mm. long. 

Plants typically glabrous; flowers often 6.5-8 mm. long. 

C. Jamesonii. 

Plants typically more or less pubescent; flowers rarely longer 

than 6 mm. 

Fruiting pedicels 1 cm. long or longer; flowers 4.5 mm. long 
or longer C. ovata. 

Fruiting pedicels mostly 6 mm. long; flowers 3-4 mm. long. 

C. africana. 

Cardamine africana L. Sp. PL 655. 1753; 414. 

Similar to C. ovata; leaves often crowded at the base of the 
weak stems, trifoliolate, the terminal leaflet ovate, more or less 
acuminate, rounded at the base, the similar lateral ones oblique, 
all 4-8-crenate-serrate or somewhat 3-lobed, rarely subentire, 
minutely ciliate and usually sparsely pilose; racemes dense, 6-16- 



962 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

flowered; fruiting pedicels mostly 6 mm. long; flowers 3-4 mm. long; 
pods 2.5-4.5 cm. long, nearly 2 mm. broad. Usually well marked 
by its large, broad, merely dentate leaflets and broad pods. 

Huanuco: Chinchao, Poeppig 1845. Without locality: Vidal- 
Senege. Tropical mountains, Africa, East Indies, Andes, and Brazil. 

Cardamine bonariensis Pers. Syn. PI. 2: 195. 1806; 450. C. 
flaccida Cham. & Schlecht. subsp. bonariensis 0. E. Schulz, Bot. 
Jahrb. 32: 450. 1903. C. laxa Benth. var. pumila Gray, U. S. Expl. 
Exped. Bot. 50. 1854. 

Rhizomes much branched; stems decumbent-ascending, often 
20 cm. long, rooting at the nodes; basal leaves 2.5-6.5 cm. long, 
the scarcely smaller cauline ones long-petiolate, the uppermost 
often simple; leaflets all subequal, orbicular or short-ovate, entire 
or merely crenate, 6-14 mm. long, 6-19 mm. wide; flowering pedicels 
2-3 mm., in fruit 5-10 mm. long; flowers usually small, the sepals 
1.5-2 mm. long; ovules 20-28; pods mostly 15-18 mm. long, 1 mm. 
wide, the style obsolete or 1 mm. long; seeds emarginate. Highly 
variable, the most common form being var. flaccida (Cham. & 
Schlecht.) Macbr., comb. nov. (C. flaccida Cham. & Schlecht. 
Linnaea 1: 21. 1826), typically more robust, more erect, the leaflets 
larger, angled-crenate, the flowers 3-4 mm. long, the pods mostly 
2 cm. long, 1.5 mm. wide, the seeds margined; but none of these 
characters apparently concomitantly holding. A reduced state is 
var. minima (Steud.) Macbr., var. nov. (C. minima Steud. Flora 
39: 410. 1856; C. flaccida subsp. minima 0. E. Schulz, op. cit. 451), 
with leaves mostly trifoliolate, the leaflets 1.5-5 mm. long or smaller, 
to about 10 mm. wide; pedicels filiform, 5-30 mm. long; flowers 3^4 
mm. long; pods 8-14 mm. long. See Schulz for other named forms 
under, however, a later name, C. flaccida. Sometimes terrestrial, 
sometimes half immersed in brooks or along their banks, and from 
the ocean to the snows, according to Weberbauer. Illustrated, 
Bot. Jahrb. 32: pi 10. 

Cajamarca: Cajamarca, Osgood & Anderson 58. Nancho, Rai- 
mondi. Ocros, 3,500 meters (Weberbauer 171). Ancash: Martinet 
715. Culnai to Casacancha (Wilkes Expedition). Huanuco: Ambo, 
2,550 meters, 2431. Lima: Obrajillo (Wilkes Expedition). Prov. 
Cajatambo, Weberbauer 2688. Junin: Pampas, Raimondi. Ama- 
zonas: Lamud, Raimondi. Puno: San Antonio, Lechler 1811 (var. 
minima). Arequipa: Raimondi. Cuzco: Vilcabamba, Raimondi. 
Paucartambo, Herrera 2331. La Raya, in Distichia cushions, 4,300 



FLORA OF PERU 963 

meters, Pennell 131*98 (var. minima). Cerro de Cusilluyoc, 3,300- 
3,500 meters, Pennell 14164- Chile and Argentina to Mexico. 
"Berro." 

Cardamine Jamesonii Hook. Lond. Journ. Bot. 6: 293. 1847; 
421. 

Plants glabrous, simple or with long branches; lower leaves 
2-3-pinnate; terminal leaflet suborbicular, subcordate at the base, 
coarsely 2-4-crenate-lobed or subincised, 12-16 mm. long, nearly 
as broad, the lateral ones similar, oblique, usually manifestly petio- 
lulate; upper cauline leaves with one pair of broadly ovate or oblong- 
ovate, smaller leaflets; racemes lax from the first, sometimes 
trifoliolate-bracted at the base, 8-15-flowered, the flowering pedicels 
6-8 mm. long, scarcely longer in fruit; flowers 6.5-8 mm. long; 
sepals ovate; petals rose-colored, broadly obovate; pods nearly 
4 cm. long, 2 mm. broad, attenuate to the style, this 3-5 mm. long; 
seeds 1.75 mm. long. 

Peru: Probably. Bolivia to Central America and Haiti. 

Cardamine Killipii 0. E. Schulz, Notizbl. Bot. Gart. Berlin 
10: 341. 1928. 

Nearly C. bonariensis var. flaccida, but the leaves simple or 
subsimple, the cordate-orbicular, repand-angulate terminal leaflet 
sometimes accompanied by one much smaller lateral one, this ovate, 
sessile; racemes several decimeters long, equaling the stems; pods 
13-16 mm. long; seeds only 0.75 mm. long. 

Lima: Rio Blanco, 3,000-3,500 meters, Kittip & Smith 21712. 
Colombia. 

Cardamine ovata Benth. PI. Hartw. 158. 1845; 412. 

Stems 1 to several decimeters long, ascending-erect, branching 
above, regularly 4-8-1 eaved, white-hirsute or glabrescent; leaves 
mostly with 2 pairs of 3-5-crenate-serrate, rarely subentire, strigu- 
lose or nearly glabrous leaflets, the lower ones ovate, roundish or 
acutish at the apex, the upper ones oblong-ovate or lanceolate, the 
terminal one always petiolulate, the lateral leaflets oblique and 
subsessile, or sometimes subdecurrent, all varying greatly in size 
from medium to almost minute; racemes to 30-flowered, usually 
trifoliolate-bracted at the base; fruiting pedicels 1 cm. long or longer; 
flowers white, about 4.5 mm., rarely 6.5 mm. long; pods more or 
less recurving, 4-5 cm. long, 1.8 mm. broad. Var. unijuga 0. E. 
Schulz no doubt occurs. 



964 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cuzco: Pillahuata, 3,000-3,300 meters, Pennell 14108 (det. 
Schulz). Without locality, Ruiz & Pav6n.Puno: Sandia, 2,600 
meters (Weberbauer 240). Huanuco: Tambo de Vaca, 3,600 meters, 
4462 (det. Schulz). Bolivia to Colombia and Venezuela. 

13. RORIPPA Scop. 
Nasturtium R. Br.; Kardamoglyphos Schlecht. 

Reference: 0. E. Schulz, Repert. Sp. Nov. 33: 273-285. 1934; 
Engl. & Prantl, Pflanzenfam. ed. 2. 17b: 554. 1936. 

The berro or watercress that may be said to typify this group 
is cosmopolitan in distribution and in popularity as a salad plant, 
particularly in Latin countries like Peru. Herrera has recorded 
local Peruvian names as "mostacilla," "mayu-mostaza," "chicchi," 
"chijchi," and "occoruro." Botanists have at one time or another 
given it more than twice as many scientific designations; see Sprague, 
Journ. Bot. 62: 225-228. 1924; but now, according to the Inter- 
national Rules, only two are to be regarded as legally correct, 
Rorippa Nasturtium-aquaticum (L.) Hayek, and Nasturtium offi- 
cinale R. Br., the latter name being conserved for those who restrict 
the genus Nasturtium to the watercress; see Marie- Victorin, Contr. 
Bot. Univ. Montreal 17: 17. 1930. In Rorippa I include Kardamo- 
glyphos, notwithstanding the lack of nectar glands, for the degree 
of development of these within the group is variable. Admittedly 
the character is important in classification of the family elsewhere. 

Petals white, conspicuous, well exceeding the sepals; medial nectar 
glands wanting R. Nasturtium-aquaticum. 

Petals slightly if at all longer than the sepals, or yellow, sometimes 

none. 

Stems leafy, ascending to erect; seeds in 2 rows. 
Annuals or short-lived plants with ample leaves or leaflets; 

nectar glands present. 
Leaves pinnate, the leaflets remote, narrow . . . R. bonariensis. 

Leaves lyrate-pinnate, the terminal leaflet much larger than 
the lateral ones R. clandestina. 

Leaves obovate, only the basal ones pinnate R. Sinapis. 

Perennials, depressed-ascending, the leaflets small; nectar 

glands none R. nana. 

Stems nearly leafless, creeping; seeds in 1 row R. sarmentosa. 



FLORA OF PERU 965 

Rorippa bonariensis (Poir.) Macloskey, Kept. Princeton Univ. 
Exped. Patag. 8, pt. 1: 434. 1905. Nasturtium bonariense DC. 
Syst. 2: 193. 1821. Sisymbrium bonariense Poir. in Lam. Encycl. 
7: 205. 1806. S. Mandonii Fourn. Recherch. Crucif. 109. 1865. N. 
erectum Trev. ex Sweet, Hort. Brit. ed. 3. 26. 1839, nomen. 

Leaves deeply pinnatifid or dissected, the terminal lobe elliptic- 
oblong, 3-6 mm. wide, entire or sparsely dentate, the lateral lobes 
2-4 on each side, mostly entire; sepals 1.5 mm. long, the petals 
2 mm. long; pedicels 2-8 mm. long; ovules 64-72; pods 10-18 mm. 
long, 1-1.5 mm. wide, the style often nearly 0.5 mm. long. 0. E. 
Schulz makes a variety of the plant of Treviranus; it is merely the 
most vigorous form, to 70 cm. high, the lateral leaf lobes 4 mm. 
wide, the dentate terminal one 6-8 mm. wide. R. Eggersii (Schulz) 
Macbr., comb. nov. (N. Eggersii 0. E. Schulz, Repert. Sp. Nov. 
33: 285. 1934), known as near as Ecuador, has terminal leaflets 
10 mm. wide, the elliptic lateral ones much smaller; petals white, 
scarcely exceeding the sepals; ovules 40; pedicels 1-2 mm. long, 
the pod 6-10 mm. long. Illustrated, Mart. Fl. Bras. 13, pt. 1 : pi. 66. 

Huanuco: Cochero, Poeppig 1454- Ancash: Martinet 718. 
Loreto: Cerro de Escalera, Ule 6697. Cuzco: Valle de Paucartambo, 
H err era 1258. Chile and Argentina, northward to Ecuador. 
' 'Llamppuchichira. ' ' 

Rorippa clandestina (Spreng.) Macbr., comb. nov. Nasturtium 
clandestinum Spreng. Nov. Prov. 29. 1819. N. clandestinum var. 
brevistylum 0. E. Schulz, Repert. Sp. Nov. 33: 278. 1934. 

Glabrous; all the leaves lyrate-pinnatifid, the terminal lobe 
obovate-reniform, often 2 cm. wide, the 3-7 lateral lobes obliquely 
obovate, repand-dentate; sepals and petals equal, 2 mm. long, the 
stamens longer; ovules 90-116; fruiting pedicels 3-8 mm. long, 
the pods 2-3 cm. long, 1.5-2 mm. wide, the style (0.5-) 1-2 mm. 
long. The Raimondi plant is the short-styled form, with a style 
0.5-1 mm. long. Var. Sodiroi (0. E. Schulz) Macbr., comb. nov. 
(N. clandestinum var. Sodiroi Schulz, Repert. Sp. Nov. 33: 277. 
1934), Ecuadorean, has pedicels 1-1.5 mm. long. 

Libertad: Prov. Huamanchuco, Raimondi. Extending to Para- 
guay and Venezuela. 

Rorippa nana (Schlecht.) Macbr., comb. nov. Kardamoglyphos 
nana Schlecht. Linnaea 28: 472. 1856. Nasturtium nanum Wedd. 
Ann. Sci. Nat. IV. 5: 290. 1864. 



966 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowering when still diminutive and tufted, but developing 
ascending stems a decimeter long or longer; leaves subpinnate to 
pinnate, 1.5-2.5 cm. long, the leaflets about 7, entire or obtusely 
1-2-dentate; flowers at first pedicellate in the leaf rosettes, later 
borne in racemes, the pedicels then about 1.5 cm. long; pods 5-7 
mm. long, 2.5-3 mm. thick at the base, tapering slightly to the 
style, this 0.5 mm. long. Illustrated, Engl. & Prantl, Pflanzenfam. 
ed. 2. 17b: 553. Neg. 34605. 

Loreto: Cerro de Ponasa, Ule 6697. Puno: Agapata, Lechler 
2583, type. Argentina to Ecuador. 

Rorippa Nasturtium-aquaticum (L.) Hayek, Sched. Fl. Stir. 
Exs. Lief. 3-4: 22. 1905. Sisymbrium Nasturtium-aquaticum L. 
Sp. PI. 657. 1753. Nasturtium officinale R. Br. in Ait. Hort. 
Kew. ed. 2. 4: 110. 1812. N. fontanum Aschers. Fl. Brandenb. 1: 
32. 1864. 

Glabrous except for an occasional hair, leafy, the leaflets oval, 
obtuse, 1-several pairs; medial nectar glands none; flowers white; 
stamens shorter than the petals; stigma broad; pedicels divaricate; 
ovules 28-48; pods linear, the small, reticulate seeds in 2 rows, 1 mm. 
long. From sea strands to snow (Weberbauer). Illustrated, Engl. 
& Prantl, Pflanzenfam. ed. 2. 17b: 553. Hayek did not hyphenate the 
species name, but gave the Linnaean reference. 

Lima: In limewater pockets, on seashore, Barranco Miraflores 
(Weberbauer, 148). Junin: La Oroya (Weberbauer, 180). Tarma, 
Weberbauer 25P-4- Cuzco : Pisac, 3,000-3,200 meters, Pennell 13718. 
Prov. Quispicanchi, 3,200 meters (Herrera). Valle del Paucartambo, 
3,000 meters, Herrera 3360. Puno: Lake Titicaca (Weberbauer, 
185). Cosmopolitan; in America presumably introduced from the 
Old World. 

Rorippa sarmentosa (Soland.) Macbr., comb. nov. Cardamine 
sarmentosa Soland. ex Forst. Fl. Ins. Austr. Prodr. Append. 91. 1786. 
Nasturtium sarmentosum 0. E. Schulz in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 17b: 555. 1936. 

A creeping, rooting, nearly scapose, glabrous plant with small 
flowers; basal leaves long-petiolate, pinnate, the 3 pairs of leaflets 
about 1 cm. long, sinuate, round, petiolulate; sepals linear, acute, 
3 mm. long; petals 3.5 mm. long, purplish white; nectar glands 
narrowly confluent; seeds large, in 1 row, reticulate, slightly mucila- 
ginous when wet. An introduction in Peru, if the record is correct. 

Lima: (Wilkes Expedition; det. Gray). Polynesian. 



FLORA OF PERU 967 

Rorippa Sinapis (Burm. f.) Macbr., comb. nov. Sisymbrium 
Sinapis Burm. f. Fl. Ind. 140. 1768. Nasturtium Sinapis O. E. 
Schulz, Repert. Sp. Nov. 33: 278. 1934. 

Well marked among Peruvian species by its simple or subsimple, 
glabrous leaves, all except the radical ones with 1 or 2 leaflets, the 
upper leaves oblong, serrate, acuminate at each end; petals often 
none; pedicels 2-7 mm. long; pods 1-3.2 cm. long, to 1.2 mm. wide; 
style 0.5-1 mm. long. Confused with this has been R. indica (L.) 
L. H. Bailey, which species, according to Schulz, op. cit. 279, is simi- 
lar to R. clandestine but is usually pubescent and has mostly shorter 
pedicels and pods, the latter 1-2 cm. long, 1.5-3 mm. wide. 

Rio Acre: Ule 153. India, and widely naturalized elsewhere. 

14. DICTYOPHRAGMUS 0. E. Schulz 

A small, glabrous, bluish green, branching annual with dense, 
short racemes of fragrant, white flowers and relatively large, sub- 
entire leaves, these narrowed to the auricled base. Sepals not cucul- 
late. Petals broadly obovate. Nectar glands forming a ring, thickest 
on the inner side. Stigma subsessile, capitate. Pods oblong, acute. 
Seeds winged. 

Dictyophragmus Englerianus (Muschl.) 0. E. Schulz, Bot. 
Jahrb. 66: 92. 1933. Streptanthus Englerianus Muschl. Bot. Jahrb. 
40: 269. 1908. 

Plants 10 cm. high or less; leaves clasping, remotely denticulate 
or subentire, 2-4 cm. long, about 1 cm. wide, oblong-obovate, obtuse 
or acutish; pedicels 2-3 mm. long; petals 6 mm. long, conspicuously 
clawed; pods 2 cm. long, 4 mm. wide, net- veined. Illustrated, Engl. 
& Prantl, Pflanzenfam. ed. 2. 17b: 534. 

Arequipa: On the lower edge of the loma, 100 meters, Mollendo, 
Weberbauer 1453, type; 144. 

15. ALYSSUM L. 

Lobularia Desv. 

The well known garden plant, sweet alyssum or "espumilla," is 
often cultivated. It is surprising to find this plant maintained by 
Schulz as a generic type on the basis of a nectar gland character, a 
character which he himself has discounted in the case of Rorippa 
and which Briquet showed to be of no value here as a generic criterion. 

Alyssum maritimum (L.) Lam. Encycl. 1: 98. 1783. Clypeola 
maritima L. Sp. PI. 652. 1753. Lobularia maritima Desv. Journ. Bot. 
3:169.1814. 



968 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A low, spreading or bushy, rather light green but often cinereous- 
pubescent annual, becoming woody below or persisting as a half 
shrub; leaves lanceolate, tapering to the base; flowers honey-scented, 
white, at first crowded, the racemes greatly elongating in fruit; pods 
spheroid, 1-seeded, reticulate- veined. 

Lima: Along Rio Rimac, Martinet 1024- Arequipa: Isern 2113. 
Cuzco: (Herrera). Native of the Mediterranean region. 

16. MATHEWSIA Hook. & Am. 

Machaerophorus Schlecht. 

More or less woody, glabrous or stellate-pubescent, leafy plants 
with obtuse, lanceolate cauline leaves and ebracteate racemes. 
Calyx closed. Petals long-clawed, about equaled by the linear an- 
thers, these on slender filaments. Pods dorsally compressed, broadly 
lanceolate, reticulate- veined. Stigma subsessile, large. The genus 
was named for Andrew Mathews, whose collections in Peru from 1833 
to his death at Chachapoyas, November 24, 1841, supplemented in 
many cases those of Ruiz and Pavon, from their type localities. See 
Journ. Bot. 27: 371. 1889. 

Mathewsia matthioloides (Schlecht.) C. Muell. in Walp. Ann. 
Bot. 7: 144. 1868. Machaerophorus matthioloides Schlecht. Linnaea 
28: 469. 1856. 

Nearly glabrous, blue-green, the leaves with minute divisions, 
and thus at once distinct from M. peruviana; flowers yellow; sepals 
narrow, about 15 mm. long, the petals 2.5 cm. long; pods to 7 cm. 
long, 3.5 mm. wide or wider. A caducous lanosity is more or less 
evident on some parts. The stems and elongate fruiting racemes 
areflexuous. Neg. 27330. 

Puno: Cordillera de Puno (Lechler 1702, type). 

Mathewsia peruviana 0. E. Schulz, Notizbl. Bot. Gart. Berlin 
10: 562. 1929. 

Woody and densely leafy below, 10-20 cm. high, softly gray- 
tomentose, even to the pods, these oblong, 4-5 mm. wide, 2-2.3 cm. 
long, rounded at the base, narrowed to the short style; leaves sinu- 
ate-pinnatifid, with about 5 triangular lobes on each side, clasping 
at the base, 3.5 cm. long; racemes finally 20-25-flowered ; pedicels 
3-5 mm., in fruit to 7 mm. long; sepals 7 mm. long, the white petals 
twice as long, elliptic above the narrow claw; stamens equal, nearly 
10 mm. long. M. laciniata Phil., of northern Chile, is said to be 
similar but is described as having white-woolly pods only 3 mm. broad. 

Arequipa: Lomas de Atico, Raimondi, type. 



FLORA OF PERU 969 

17. EREMODRABA 0. E. Schulz 

Low, glabrous, suffrutescent plants, the Peruvian species with 
the aspect of Sisymbrium. Petals yellow. Anthers oblong; filaments 
slender; medial nectar glands small. Ovules typically 6-8. Stigma 
capitate-flattened, subsessile. Pods narrow, flat, indistinctly nerved, 
with a thin septum. Seeds in 1 row, on short, filiform funicles, 
lustrous. 

Eremodraba Hinkleyana 0. E. Schulz in Engl. & Prantl, 
Pflanzenfam. ed. 2. 17b: 617. 1936. 

Leaves linear, remotely dentate or almost laciniate, sagittate- 
amplexicaul, longer than the flowering raceme; ovary with 16-18 
ovules (Schulz). The author remarks, "I place this plant here with 
some doubt, the fruit being unknown." The type species is E. intri- 
catissima (Phil.) O. E. Schulz (Philippi called it a Draba), much 
branched and with linear, fleshy leaves, known only from the deserts 
of northern Chile. 

Arequipa: Chacchani Mt., Hinkley, type. 

18. ENGLEROCHARIS Muschl. 
Brayopsis Gilg & Muschl. 

Small, cespitose, simple-haired perennials, the ovate leaves 
crowded-rosulate about the congested inflorescence of scapose pedi- 
cels that elongate in fruit. Sepals more or less persistent, nearly 
as long as the greenish yellow, white, or purplish petals. Nectar 
glands confluent, the medial reduced or wanting. Style and ovary 
subequal. Style short or nearly obsolete. Stigma depressed. Pods 
short-oblong, the septum complete. Illustrated, Weberbauer 198. 
The genus Brayopsis differs in no fundamental respect except 
that the medial nectar glands are wanting, a variation paralleled 
in Nasturtium and Eudema. 

Leaves and ovary glabrous, or the leaves sparsely pilose. 

Petals and sepals subequal, about 4 mm. long E. calycina. 

Petals definitely longer than the sepals, to 5 mm. long. 

E. Weberbaueri. 
Leaves and usually the ovary very pubescent. 

Petals white or purplish; ovules 12-16 E. peruviana. 

Petals greenish yellow; ovules many E. alpaminae. 



970 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Englerocharis alpaminae (Gilg & Muschl.) Baehni & Macbr., 
comb. nov. Brayopsis alpaminae Gilg & Muschl. Bot. Jahrb. 42: 
483. 1909. 

Tap root fibrous below; leaves obovate-lanceolate, acute, long- 
cuneate at the base, entire, 7-8 mm. long, 2 mm. wide above, densely 
long-ciliate; flowers solitary, the peduncles 7-8 mm. long; pods 
acute at both ends, densely pubescent with simple hairs, 1 cm. long, 
scarcely 2 mm. wide, the peduncle to 1.5 cm. long. My collections 
were from plants growing among rocks; flowers of No. 3074 greenish 
yellow. Var. argentea (Gilg & Muschl.) Baehni & Macbr., comb, 
nov. (B. argentea Gilg & Muschl. Bot. Jahrb. 42: 483. 1909), has 
leaves to 2.5 mm. wide; peduncle 4-10 mm. long; petals 6 mm. long. 
Illustrated, Engl. & Prantl, Pflanzenfam. ed. 2. 17b: 631. 

Ancash: Piscapacha, Huaraz, 4,500 meters, Weberbauer 2901, 
type of the variety. Lima: Alpamina, 4,500 meters, Weberbauer 
5123, type. Junin: Cerro de Pasco, 4,200 meters, 30? '4- Moro- 
cocha, 4,500 meters, 887. Huancavelica, 4,200 meters, 1119. 

Englerocharis calycina (Desv.) Baehni & Macbr., comb. nov. 
Draba calycina Desv. Journ. Bot. 3: 185. 1814. Braya calycina 
Wedd. Chlor. And. 2: pi. 85a. 1857. Sisymbrium calycinum Wedd. 
Ann. Sci. Nat.- V. 1: 289. 1864. Brayopsis calycina Gilg & Muschl. 
Bot. Jahrb. 42: 484. 1909. B. pycnophylla Gilg & Muschl. loc. cit. 

Similar to E. Weberbaueri but the leaves stiffly ciliate-pilose 
and the peduncles often longer, in fruit even to 3.5 cm. long; petals 
whitish; pods 8-11 mm. long, the style obsolete. Var. filiformis 
(0. E. Schulz) Baehni & Macbr., comb. nov. (B. calycina var. fili- 
formis 0. E. Schulz, Pflanzenreich IV. 105: 240. 1924) has linear 
leaves and slender peduncles, in fruit to 2 cm. long. There are also 
named glabrate and cinereous-pubescent forms found with the usual 
state. Illustrated, Engl. & Prantl, Pflanzenfam. ed. 2. 17b: 630. 

Ancash: Huaura, 4,400 meters, Weberbauer 7247a. Above Pisca- 
pacha, 4,500 meters, Weberbauer 2901. Lima: Matucana, 2,700 
meters, 619. Junin: Near Yauli, Weberbauer 321. Ayacucho: 
Huanta region, Weberbauer 7494, type of var. filiformis; 7499. 
Cuzco: Quispicanchi, 4,500 meters, Weberbauer 7770 (glabrous- 
leaved form). Moquehua: Volcan Ticsani, 4,000 meters, Weber- 
bauer 7329. 

Englerocharis peruviana Muschl. Bot. Jahrb. 40: 276. 1908. 
Leaves entire, 1.5-2 cm. long, 5-9 mm. wide, densely pilose- 
strigose; fruiting scapes 2-4 cm. long; sepals 4-5 mm. long; fruiting 



FLORA OF PERU 971 

pedicels 1-1.5 cm. long; pods 7.5-9 mm. long, 2-2.5 mm. wide, the 
style 1.5-2 mm. long. A variant with glabrous ovary is var. leio- 
carpa Schulz, Pflanzenreich IV. 105: 197. 1924. The plant grows in 
shallow, shifting detritus. 

Lima: Casapalca, 3,650 meters, 869. Junin: Yauli, 4,600 meters, 
Weberbauer 331, type; 221. Above Huancayo, 4,900 meters, Weber- 
bauer 6525. Ancash: Cordillera de Pelagatos, 4,450 meters, Weber- 
bauer 7231 (the variety). 

Englerocharis Weberbaueri (Gilg & Muschl.) Baehni & Macbr., 
comb. nov. Brayopsis Weberbaueri Gilg & Muschl. Bot. Jahrb. 42: 
483. 1909. 

Caudex branches conspicuously clothed with remains of the 
glabrous, oblong leaves, or these sometimes ciliate on the long- 
cuneate base; scapes glabrous, to 1.5 cm. long in fruit; ovary 18- 
ovulate; pods almost 1 cm. long, 2 mm. wide. Neg. 13871. 

Ancash: Near Huaraz, 4,400 meters, Weberbauer 2971, type. 
Junin: Rio Blanco, 4,500 meters, 2995. 

19. CATADYSIA 0. E. Schulz 

A tufted, depressed perennial, each plant a rosette of spreading, 
broadly obovate, shallowly dentate leaves, from which rise the 
many short stems of leafy-bracted, crowded, subsessile, white 
flowers. Petals obovate, clawed, emarginate, twice as long as the 
sepals. Anthers oval. Pods reticulate- veined, narrowly ovate, 
tipped by the slender style, the stigma flat. 

Catadysia rosulans 0. E. Schulz, Notizbl. Bot. Gart. Berlin 
10: 559. 1929. 

Basal leaves 5-10 mm. wide, on petioles 1-1.5 cm. long, often 
violet-tinged beneath; peduncles 5-8 mm. long; sepals 3 mm., the 
petals 5.5 mm. long, pale lilac; stamens 4 mm. long; pod 5-6 mm. 
long, 2 mm. wide, the style 1.2 mm. long. 

Ancash: Mt. Huacchara, in limestone, 4,300 meters, Weberbauer 
7236, type. 

20. WEBERBAUERA Gilg & Muschl. 
Alpaminia and Pelagatia 0. E. Schulz. 

Low, rather loosely branched herbs with laxly rosulate leaves 
at the tips of the branchlets. Cauline leaves deeply incised to sub- 
entire. Flowers white or rose, in racemes that are much elongate 



972 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

in fruit. Pods linear, erect, the valves convex. Style short. Seeds 
in 1 series, brown, short-funiculate. Cotyledons incumbent. Funi- 
cles slender or in part dilated (in the segregate genera), a variation 
found also in Cardamine. 

Whole plant (including the flowers) densely pubescent. 

W. trichocarpa. 
Plants glabrous or somewhat pubescent. 

Leaves lyrate-pinnatifid W. bracteata. 

Leaves entire or repand-dentate. 

Stems pilose W. spathulifolia. 

Stems glabrate W. pusilla. 

Weberbauera bracteata (0. E. Schulz) Macbr. Candollea 5: 
356. 1934. Pelagatia bracteata 0. E. Schulz, Pflanzenreich IV. 105: 
192. 1924. 

Stems depressed, 3-5 cm. long, equaled by the lower leaves; 
cauline leaves much smaller, dentate or entire; racemes about 10- 
flowered; pedicels 2-5 mm. long; sepals glabrous, 2.2 mm. long, the 
ochroleucous petals slightly longer; ovary 12-ovulate; pods 7.5 mm. 
long, 2 mm. broad; seeds nearly uniseriate. Well marked by the 
simply ciliate leaves and the malvaceous habit. 

Ancash: Cordillera de Pelagatos, in limestone, 4,600 meters, 
Weberbauer 7234, type. 

Weberbauera pusilla (Gillies) 0. E. Schulz, Pflanzenreich IV. 
105: 194. 1924. Erysimum pusillum Gillies ex Hook. & Arn. in Hook. 
Bot. Misc. 3: 140. 1833. Arabis spathulata Walp. sensu Meyen, Obs. 
Bot. 248. 1843. Weberbauera densiflora Gilg & Muschl. Bot. Jahrb. 
42: 481. 1909, pro parte. 

Similar to W. spathulifolia, but the stems little pubescent, or 
glabrous, the petioles and blades of the basal leaves subequal, and 
the petals broadly obovate. Perhaps better treated as a variety, 
but it has a distinct range. Neg. 34580. 

Puno: Altos de Toledo, 4,000 meters, Meyen. Chile; Bolivia; 
Argentina. 

Weberbauera spathulifolia (Gray) 0. E. Schulz, Pflanzenreich 
IV. 105: 193. 1924. Sisymbrium spathulifolium Gray, U. S. Expl. 
Exped. Bot. 60. 1854. S. ciliolatum Fourn. Recherch. Crucif. 129. 
1865. Braya densiflora Muschl. Bot. Jahrb. 40: 275. 1908. Weber- 
bauera densiflora Gilg & Muschl. Bot. Jahrb. 42: 481. 1909, pro parte. 



FLORA OF PERU 973 

Tufted, the spreading-ascending stems 10 cm. long or less; leaves 
narrowly lanceolate, long-petiolate, the upper and also the subses- 
sile, finally bractlike cauline leaves repand-dentate; racemes 10-15- 
flowered; pedicels to 2 mm. long; sepals with a few simple or forked 
hairs, a third shorter than the (3 mm. long) white or reddish, nar- 
rowly obovate-cuneate petals; fruiting pedicels erect-spreading, to 
6 mm. long; pods 6-19 mm. long. Var. integrifolia 0. E. Schulz, 
Field Mus. Bot. 8: 80. 1930, has nearly or quite entire leaves. The 
leaves of the Gay plant are entire and simply ciliate. Illustrated, 
Pflanzenreich, loc. cit. 

Junin: Obrajillo (Wilkes Expedition). Yauli, 4,400 meters, 
Weberbauer 304, type of B. densiflora. Near La Oroya, 4,000 meters, 
Weberbauer 2550. Above Lima, Weberbauer 3758. Cerro de Pasco, 
4,200 meters, 3065 (var.); Gay 2556 (herb. Paris). Lima: Rio 
Blanco, 4,500 meters 2991, 811 (var.). Viso, 2,700 meters, 590 
Moquehua: Above Torata, 3,800-3,900 meters, Weberbauer 7471. 
Without locality: Ruiz & Pavdn. Puno: Puno, 4,000 meters, Sou- 
kup 106. 

Weberbauera trichocarpa (Muschl.) Macbr. Candollea 5: 356. 
1934. Eudema trichocarpa Muschl. Bot. Jahrb. 40: 276. 1908. Bray- 
opsis trichocarpa Gilg & Muschl. Bot. Jahrb. 42: 484. 1909. Alpa- 
minia trichocarpa 0. E. Schulz, Pflanzenreich IV. 105: 191. 1924. 

Stems mostly procumbent or finally somewhat ascending, only 
3-4 cm. long, even in fruit; leaves fleshy, entire, congested, broadly 
linear; flowers about 4, crowded, reddish or pink, or the petal mar- 
gins yellowish; pedicels to 1 cm. long; sepals 4 mm. long, strigose- 
pilose as also the longer, oblong petals; ovary multiovulate; pods 
to 1 cm. long and 2.5 mm. wide. Illustrated, Pflanzenreich, loc. cit. 

Junin: Alpamina, 4,500 meters, Weberbauer 5119, type. Rio 
Blanco, 4,500 meters, 2990. Cerro de Pasco, 4,200 meters, 3073 
Ancash: In stony puna mats on limestone, Conchucos, 4,500 meters, 
Weberbauer 7229, 7229a. 

21. SISYMBRIUM L. 

Halimolobus Tausch.; Phlebiophragmus 0. E. Schulz; Neuonto- 
botrys 0. E. Schulz; Pennellia Nieuwl. 

Reference: 0. E. Schulz, Pflanzenreich IV. 105: 46-157. 1924. 

Erect or diffuse annuals, or the plants enduring and suffrutes- 
cent at the base, glabrous, or sparsely pubescent with simple or 
rarely branched hairs. Leaves usually lyrate-pinnatifid. Pods linear 



974 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and often greatly elongate, the style obsolete or short to well devel- 
oped. Septum of the pods usually 1-2- , rarely 2-4-nerved, or nerve- 
less. Seeds not mucilaginous when wet except as stated, being 
variable in this respect, as in Rorippa. Here might be sought the 
north Chilean Ivania 0. E. Schulz, Repert. Sp. Nov. 33: 188. 1933, 
with entire, cordate leaves, short, dense racemes, and stipitate 
ovary. In addition to the following, there is a specimen in Herb. 
Madrid, Isern 2115, from Camino de Palca, Dept. Junin, apparently 
new, but which is so exceptional in character that I have not wished 
to give it a name now. Dr. Charles Baehni has kindly studied it 
and determined that the primary root is lateral. It is an almost 
perfect match for the North American Thelypodium lasiophyllum 
(Hook. & Arn.) Greene in general aspect, but the pods are not 
recurved and the primary root is clearly lateral. It may be described 
as follows: 

Slender, apparently annual or biennial, to at least 30 cm. high, 
the terete stems below and the leaves sparsely hispidulous with 
spreading, simple hairs or with an occasional furcate one; lower 
leaves unknown; middle cauline leaves oblong, pinnate-lobed or 
dentate, to 10 cm. long and 3 cm. wide, mostly somewhat shorter 
and about 1.5 cm. wide, long-attenuate to the petiolate base, the 
lobes acute, to 2 mm. wide, oblong-linear, remote, about 4 on each 
side of the oblong-linear portion of the leaf; uppermost leaves den- 
ticulate, linear; racemes ebracteate; pedicels glabrous or sparsely 
hispidulous, divaricate-ascending, 5 mm. long; pods 1-nerved, 
arcuate-ascending, torulose, 4 cm. long, 0.75 mm. wide; style 1 mm. 
long, the stigma depressed-capitate, not lobed; septum nerveless; 
seeds not mucilaginous when wet; petals pale, filiform-clawed, about 
4 mm. long; anthers blunt; style short, thick, the stigma depressed, 
not obviously if at all lobed. Neg. 34289. 

Leaves distinctly amplexicaul, at least the upper ones. 

Leaves glabrous; seeds not mucilaginous when wet. .S. Grayanum. 
Leaves more or less pubescent. 

Seeds not mucilaginous when wet; flowers purple or white. 
Leaves entire to coarsely dentate. 

Leaves subentire; pedicels 10-20 mm. long S. effusum. 

Leaves coarsely dentate; pedicels 4-8 mm. long . .S. lanatum. 

Leaves 3-4-pinnate S. Weberbaueri. 

Seeds mucilaginous when wet; flowers yellowish. .S. hispidulum. 



FLORA OF PERU 975 

Leaves not at all, or at least most of them not obviously amplexicaul. 
Style narrower than the ovary; leaves linear, entire, about 1 mm. 

wide S. Berningeri. 

Style thick, short; leaves broader than 1 mm. 

Seeds mucilaginous; leaves hispidulous, remotely denticulate, 

oblong-linear, the upper ones linear, entire S. gracile. 

Seeds not mucilaginous when wet; leaves otherwise. 
Leaves merely coarsely dentate. 

Leaves glabrous S. oleraceum. 

Leaves hispidulous S. peruvianum. 

Leaves, at least the middle ones, pinnatifid. 
Septum of the pod nerveless; pedicels 1-1.5 mm. long. 

S. officinale. 
Septum 2-4-nerved; pedicels 6-15 mm. long. 

S. macrorrhizum. 

Sisymbrium Berningeri (0. E. Schulz) Baehni & Macbr. 
Candollea 7: 295. 1937. Neuontobotrys Berningeri 0. E. Schulz, 
Notizbl. Bot. Gart. Berlin 11: 392. 1932. 

Plants bluish green but more or less pubescent, even to the ovary, 
with simple hairs, becoming bushy-branched and ligneous below; 
leaves linear, entire, fleshy; racemes 20-40-flowered, the yellow 
(fading to reddish) petals 7 mm. long, twice as long as the sepals; 
pedicels 3-6 mm. long, more or less recurving, contorted in fruit; 
pods slender, the middle nerve not prominent, the style 1 mm. long, 
the stigma bilobed; ovules 16-32; seeds in 1 row. The similar 
Polypsecadium Harmsianum (Muschl.) 0. E. Schulz, of Bolivia, has 
longer, many-seeded pods, the entire leaves lanceolate. Technically 
closely related but probably a convenient segregate is Werdermannia 
pinnata (Barne"oud) 0. E. Schulz (S. pinnatum Barn.), with finely 
pinnate leaves, the leaflets divaricate, and strict racemes of whitish 
flowers. S. Berningeri was based on specimens from Pampa Ossa, 
Arica, and so may be expected in Tacna. 

Peru: Probably. Chile. 

Sisymbrium effusum 0. E. Schulz, Pflanzenreich IV. 105: 

72. 1924. 

A tall (2 meters), much branched shrub, the erect-spreading, 
flexuous branches pubescent with spreading hairs; lower leaves lan- 
ceolate, short-acuminate to the petiole, the upper ones clasping and 
deeply sagittate, with elongate, acuminate auricles, all green above 



976 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

but strigose-hispid, softly white-pubescent beneath; racemes 30-50- 
flowered; pedicels spreading or somewhat recurving, 1-2 cm. long; 
sepals 4 mm. long, glabrous; petals violet, 5 mm. long; ovules 32; 
young pods nearly 5 cm. long, with a style 1 mm. long, pubescent 
with long, weak hairs. 

Huancavelica: Surcubamba, 2,800 meters, Weberbauer 6499, type. 

Sisymbrium gracile Wedd. Ann. Sci. Nat. V. 1: 288. 1864. 
Heterothrix gracilis 0. E. Schulz, Pflanzenreich IV. 105: 297. 1924. 
Pennellia gracilis 0. E. Schulz in Engl. & Prantl, Pflanzenfam. ed. 2. 
17b: 644. 1936. 

Biennial, branching above, more or less hispid below with bifur- 
cate hairs; basal leaves oblong, obtuse, remotely denticulate, cune- 
ately narrowed to the sessile or petiolate base, hispid like the stems; 
cauline leaves narrower, the lower ones broadly linear, sparsely and 
remotely denticulate, the linear upper ones acuminate, entire; ra- 
cemes lax; pedicels 3.5-9 mm. long; sepals 3 mm. long, the inner some- 
what cucullate; petals 4 mm. long, with a broad claw; filaments 
dilated at the base; pods 1.2-2.8 cm. long, 1.5 mm. wide, the style 
scarcely 0.5 mm. long, the valves slenderly 1-nerved, the septum 
nerveless; seeds in 1 row, mucilaginous in water. Neg. 34562. 

Tacna: Tacora, Weddell, type. Bolivia. 

Sisymbrium Grayanum Baehni & Macbr. Candollea 7: 295. 
1937. S. amplexicaule Gray, U. S. Expl. Exped. Bot. 61. 1854; 58; 
non Desv. 

Glaucescent and glabrous, or nearly so, with divaricately branched, 
leafy stems woody below, often 60-70 cm. high; lower leaves obovate- 
elliptic, sinuate-dentate, 6 cm. long, the upper smaller and entire; 
racemes in bud corymbose-congested, elongating, the pedicels fili- 
form, 5-6 mm. long; sepals 4-4.5 mm. long, the pale lilac, narrowly 
obovate-cuneate petals 7-8 mm. long; pods 2-2.5 cm. long, the style 
obsolete, the valves 1-nerved, irregularly spreading from suberect 
pedicels; ovules 36-40; seeds 1-seriate. Var. tenuicaule (0. E. Schulz) 
Baehni & Macbr. loc. cit., has leaves only 4 cm. long, sepals 4 mm. 
long, and narrower, white petals. 

Lima: Rock outcrops, 1,200 meters, Weberbauer 5328. Cabal and 
Obrajillo (Wilkes Expedition, type). Without locality (Gay 291). 
Ancash: Above Pisco, 1,000 meters, Weberbauer 5378 (the variety). 

Sisymbrium hispidulum (DC.) Triana & Planch. Ann. Sci. 
Nat. IV. 17: 63. 1862. Halimolobus hispidulus 0. E. Schulz, Pflan- 
zenreich IV. 105: 291. 1924. H. Weddellii 0. E. Schulz, op. cit. 292. 



FLORA OF PERU 977 

Greggia arabioides Muschl. Bot. Jahrb. 40: 274. 1908. Turritis hispi- 
dula DC. Syst. Nat. 2: 213. 1821. Arabia andicola Walp. ex Meyen, 
Obs. Bot. 248. 1843, as to the Meyen plant. 

A subsimple biennial with flocculose, branched pubescence through- 
out; lower cauline leaves subentire and subsessile, the upper more 
manifestly dentate and clasping-sagittate, or the uppermost entire; 
racemes finally about 40-flowered; petals 3.5 mm. long, drying ochro- 
leucous; sepals 2.5 mm. long; fruiting pedicels to 11 mm. long, the 
pods 2-2.6 cm. long, sparsely stellate-pilose. Seeds mucilaginous 
when wet. Var. Weddellii (Fourn.) Baehni & Macbr. Candollea 
7: 296. 1937, is often stouter and more canescent; sepals to 4.5 mm. 
long; pods more or less hirsute, 1-2.2 cm. long. Var. Herrerae (0. 
E. Schulz) Baehni & Macbr., comb. nov. (Halimolobus Weddellii var. 
Herrerae 0. E. Schulz, Notizbl. Bot. Gart. Berlin 11: 391. 1932) has 
lower leaves deeply sinuate-dentate, pods to 2.8 cm. long, with a 
style 2-2.5 mm. long (cf. Notizbl. Bot. Gart. Berlin 11: 391. 1932). 
Greggia camporum Gray, as to Weberbauer 130, 313, is no doubt this 
plant. 

Huanuco: Canyon ledges and slopes, Llata, 2,100 meters, 2252. 
Ayacucho: Brook edges, Huanta, 3,200 meters, Weberbauer 7518. 
Puno: Titicaca, Meyen (var.). Cuzco: Urubamba Valley (Herrera; 
var.). Valle del Paucartambo and Sacsahuaman, Herrera 1041 
(var.), 2268. Lima: Rio Rimac (Ball). Chicla (Ball); Weberbauer 
254. Rio Blanco, Kittip & Smith 21562 (var.). Ancash: Between 
Samanco and Huaraz, 3,000 meters, Weberbauer 3157 (var.). Cu- 
tervo, Raimondi (var.). lea: Raimondi (var.). Bolivia to Mexico. 

Sisymbrium lanatum (Walp.) 0. E. Schulz, Notizbl. Bot. 
Gart. Berlin 11: 642. 1932. Arabis lanata Walp. in Meyen, Obs. 
Bot. 248. 1843. 

Suffruticose, with ascending or diffuse, much branched, cinereous- 
pubescent stems; leaves lanceolate, acuminate, deeply clasping, 
dentate, pubescent like the stems; racemes dense, 50-120-flowered ; 
pedicels 4-8 cm. long; sepals 4.5 mm. long; petals rose-red, 7 mm. long; 
ovules 28-32; pods flexuous-ascending, 10-22 mm. long, the style 
0.5 mm. long; seeds biseriate. Illustrated, Engl. & Prantl, Pflan- 
zenreich IV. 105: /. 16. Var. fragile (Wedd.) 0. E. Schulz, loc. cit. 
(S. fragile Wedd. Ann. Sci. Nat. V. 1: 288. 1864) is less densely pubes- 
cent and has white flowers; the typical form is known only from 
the type. 

Tacna: Candarave, 3,500 meters, Weberbauer 7570. Tacora, 1,500 
meters (Meyen, type); Raimondi; Isern 2116; Weddell (type of the 



978 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

variety). Arequipa: Yura, 2,700 meters, Weberbauer 6839. Airam- 
pal, 3,200 meters, Weberbauer 1 406 .Without locality: Ruiz & 
Pawn. Lima: Viso, 2,700 meters, 570. Bolivia. 

Sisymbrium macrorrhizum (Muschl.) Macbr. Candollea 5: 
355. 1934. Thelypodium macrorrhizum Muschl. Bot. Jahrb. 40: 268. 
1908. Phlebiophragmus macrorrhizus 0. E. Schulz, Pflanzenreich 
IV. 105: 165. /. 4- 1924. 

Nearly glabrous, suffruticose, branching from the base, the assur- 
gent branches to about 50 cm. high; lowest leaves simple, narrowed 
to an evident petiole, repand-dentate; middle leaves deeply pinnatifid, 
with about 4 remote lobes on each side; upper leaves linear; racemes 
loosely corymbose, 25-40-flowered ; pedicels 6-10, in fruit to 15 mm. 
long, filiform; sepals 3 mm., the petals 5 mm. long; ovules 40-52; 
style obsolete; pods to 4.5 cm. long, the septum 2-4-nerved, spread- 
ing and more or less recurved-ascending. In rocky wastes and out- 
crops. The genus Phlebiophragmus rests on the short style and the 
2-4-nerved septum. The rigid, sparse pubescence of the leaves is 
mostly simple. Var. ramosissimum (Muschl.) Macbr., comb. nov. 
(Erysimum ramosissimum Muschl. Bot. Jahrb. 40: 273. 1908; P. 
macrorrhizus var. ramosissimus 0. E. Schulz, op. cit. 166) has pinnati- 
sect leaves, the terminal lobe linear, shorter pedicels, and petals 4 
mm. long. Schulz suggests that S. leptocarpum as to Wilkes Expedi- 
tion material (det. Gray) may belong here. Var. Meyenii (0. E. 
Schulz) Macbr., comb. nov. (S. Meyenii 0. E. Schulz, Pflanzenreich 
IV. 105: 76. 1924; Phlebiophragmus macrorrhizus var. Meyenii 0. E. 
Schulz, Bot. Jahrb. 66: 98. 1933), has 3-4-pinnate lower leaves and 
rather densely pubescent stems. 

Arequipa: Pampa de Arrieros, 3,200 meters, Weberbauer 1395, 
4847 (var.). Above Arequipa, 2,800 meters, Weberbauer 6843. Are- 
quipa, 2,600-2,700 meters, Pennell 13207, 13234. Lima: Matucana, 
2,400 meters, 417, 634; Weberbauer 5271. Ancash: Ocros, 3,000 
meters, Weberbauer 2720. Moquehua: Carumas, Weberbauer 7290, 
7462 (var.). Puno: Pisacoma, Meyen (var. Meyenii). 

Sisymbrium officinale (L.) Scop. Fl. Cam. ed. 2. 2: 26. 1772; 
139. Erysimum officinale L. Sp. PI. 660. 1753. 

Erect, finally widely branched above, pubescent with stiff hairs; 
leaves pinnatisect, strigose-pilose on both sides and ciliate; pedicels, 
even in fruit, only 1-1.5 mm. long, erect; petals yellow, 3-4 mm. long; 
pods 10-14 mm. long, typically hirsute. 



FLORA OF PERU 979 

Moquehua: Carumas, 3,100 meters, Weberbauer 7287. Lima: 
Viso, on railway embankment, 2,700 meters, 636. Native of Europe. 

Sisymbrium oleraceum 0. E. Schulz, Pflanzenreich IV. 105: 
74. 1924. 

Glabrous, woody at the base, erect, to 55 cm. high, branched, 
with short, leafy branchlets from the axils of the leaves; leaves 
obviously petiolate, ovate, acute, unequally and coarsely dentate 
except at the entire, cuneate base; racemes lax, 20-30-flowered; pedi- 
cels 4-5 mm. long, to 15 mm. in fruit; sepals 4 mm. long, obtuse; 
petals yellow, 6.5 mm. long; ovules 28; style short, thick, finally 
1 mm. long; stigma capitate, subbilobate; pods 4-4.5 cm. long, 1.5 
mm. wide, the valves 3-nerved; seeds 1-seriate. Neg. 13851. 

Cuzco: Huancalli, 3,400-3,500 meters, Pennett 13714 (det. 
Schulz). Without locality: Tafalla, type. 

Martinet 257, from Tamboraque (herb. Paris), seems to belong 
here, but the upper middle leaves are few-pinnate, the divisions nar- 
row, and the stigma is not evidently lobed. 

Sisymbrium peruvianum DC. Syst. 2: 477. 1821; 55. Arabis 
bracteata Wedd. Ann. Sci. Nat. V. 1: 291. 1864. S. anomalum Wedd. 
op. cit. 289. 

Annual, the several to many stems branching from the base and 
procumbent or ascending, 10 cm. to several decimeters long, his- 
pidulous; leaves mostly petioled, the lower ones remotely, the upper 
ones coarsely dentate; racemes 12-60-flowered ; pedicels 1.5-3 mm. 
(in fruit 5 mm.) long; sepals glabrous, oblong, obtuse, less than 2 
mm. long, the white or pinkish petals 2.5-3 mm. long; ovules 16-40; 
style none; stigma minute; pods recurved or pendent, 1-2.5 cm. long, 
1.5 mm. wide, more or less densely spreading-pubescent; valves 1- 
nerved. On rocks. Neg. 13852. 

Lima: Viso, 3,000 meters, 597. Junin: Morococha, 4,500 meters, 
879. Puno: Altos de Toledo, 4,400 meters, Meyen. Azangaro, 
Lechler 1 770. Moquehua : Carumas, 2,700-2,800 meters, Weber- 
bauer 7352. Without locality: Ruiz & Pavdn, type. Bolivia. 

Sisymbrium Weberbaueri 0. E. Schulz, Field Mus. Bot. 8: 
80. 1930. 

Sparsely hispidulous, suffruticose, 30 cm. high; lower and middle 
leaves 3-4-pinnate, subclasping, the linear lobes entire, the terminal 
one elongate, the lateral ones remote and unequal; upper leaves 



980 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

linear, dentate or entire; racemes 10-20-flowered; pedicels 7 mm. 
long; sepals nearly 5 mm. long, obtuse, broadly hyaline-margined; 
petals white, 8 mm. long; ovules about 60; style 1 mm. long, the 
stigma capitate, subbilobate. 

22. DESCURAINIA Webb & Berth. 
Reference: 0. E. Schulz, Pflanzenreich IV. 105: 305-346. 1924. 

Often weedy or ragged plants of varying duration, with mostly 
bipinnatifid, usually finely dissected leaves. Pubescence mostly 
branched, sometimes floccose, frequently glandular. Flowers small. 
Pedicels in fruit filiform. Sepals narrow. Petals spatulate. Nectary 
glands confluent. Pods linear. Pistil cylindric. Seeds 0.5-1.2 mm. 
long. The generic name should be accepted as interpreted by 0. E. 
Schulz and others. "D. canescens" of Hicken, "very frequent" at 
Sicuani, Prov. Canchis, Cuzco, must be a misdetermination, since it 
is a North American species. 

Pods capitately congested, 4-5 mm. long D. athrocarpa. 

Pods in more or less elongate racemes, the flowers rarely capitate- 
congested. 
Leaves simply pinnate-dissected; pods 9-12 mm. long. 

D. titicacensis. 

Leaves bipinnately dissected. 
Racemes at first subcapitate; pods 4-10 mm. long. 

Diffuse or cespitose plants; racemes bracted, 10-40-flowered. 

D. depressa. 
Erect plants; racemes ebracteate, to 130-flowered. 

D. Macbridei. 
Racemes soon elongate; pods 10-19 mm. long. 

Upper leaves not markedly smaller than the lower, the ulti- 
mate lobes acutish. 

Cauline leaves 3-10 cm. long (some always longer than 
5 cm.). 

Lateral leaf lobes crowded D. myriophylla. 

Lateral leaf lobes remote D. Perkinsiana. 

Cauline leaves 1.5-5 cm. long (some always shorter than 
5 cm.) D. leptodada. 

Upper leaves notably smaller than the lower, the ultimate 
lobes acute D. pulcherrima. 



FLORA OF PERU 981 

Descurainia athrocarpa (Gray) 0. E. Schulz, Pflanzenreich 
IV. 105: 340. 1924. Sisymbrium athrocarpum Gray, U. S. Expl. 
Exped. Bot. 59. 1854. D. Urbaniana Muschl. Bot. Jahrb. 40: 271. 
1908. D. Gilgiana Muschl. op. cit. 272. 

A cinereous-stellate-pubescent perennial, 10-20 .cm. high, with 
rosulate, petiolate, bipinnate basal leaves and headlike inflores- 
cences; leaf lobes 6-8 pairs; pedicels 3-5 mm. long in fruit; sepals 
purple at the tip, 2 mm. long; petals 2.5 mm. long, pale yellow; pods 
4-5 mm. long, to 1 mm. wide; seeds biseriate. Schulz treats D. 
Gilgiana as a variety (op. cit. 341), as also var. macrorrhiza (Muschl.) 
0. E. Schulz, loc. cit., both of which seem to be individual variations 
due to age or habitat, the stems many, the leaf lobes smaller, the 
root to even 2 cm. in diameter and woody. Illustrated, Pflanzenreich 
op. cit. 340. 

Lima: At 3,000 meters, Raimondi.Jumn: Near Yauli, 4,400 
meters, Weberbauer 310, 222, 223. Above Lima, 4,000-4,700 meters, 
5181. Obrajillo (MacLean, type). Cuzco: Region of Cuzco, Herrera 
2576. Moyabamba: Chachapoyas, Raimondi. Ancash: Mount 
Huacchara, Weberbauer 7237. Tacna: Volcan Tacora, 4,300 meters, 
Werdermann 1508. Bolivia; Chile. "Alcalde-ccora," "celendin." 

Descurainia depressa (Phil.) Reiche, Fl. Chile 1: 118. 1896; 339. 
Sisymbrium depressum Phil. Verzeich. Pfl. Antofag. 4. 1891. S. 
myriophyllum Walp. ex Meyen, Obs. Bot. 248. 1843, pro parte 
(Meyen plant), non Willd. 

A biennial or perennial with often many prostrate stems, several 
centimeters or even decimeters long, floccose, even to the sepals; 
leaves cinereous-lanate, to 2 cm. long, with about 5 pairs of minute 
leaflets; pedicels to 4 mm. long; petals 2-3 mm. long, yellow or often 
reddish; pods erect, sometimes capitately congested, 4-10 mm. long, 
glabrous. Var. Pflanzii (Muschl.) Schulz, loc. cit., has the pods 
stellate-tomentose in some degree. It grows with the typical state 
and is S. titicacense Wedd. Chlor. And. 2: pi. 85c. 1857, non Walp. 
Neg. 13928. 

Puno: Altos de Toledo and Pisacoma, Meyen (var. and type). 
Lima: Casapalca, 5,000 meters, 827 (in part). Junin: Cerro de 
Pasco, 4,200 meters, 2398. Rio Blanco, 4,500 meters, 3023. Extend- 
ing to northern Chile and Argentina. 

Descurainia leptoclada Muschl. Bot. Jahrb. 40: 272. 1908; 338. 

An erect, often very tall (sometimes branched from the base), 

cinereous-pubescent biennial, the pubescence substellate; leaves 1-3 



982 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cm. long or longer, with 4-6 pairs of small leaflets, these with 3-5 
oblong, acutish lobes; sepals 2 mm. long, the pale yellow petals a 
third longer; ovules 50-62; pods 9-13 mm. long, the pedicels about 
4 mm. long; otherwise similar to D. myriophylla. Var. dasycarpa 
0. E. Schulz, Pflanzenreich IV. 105: 338. 1924, is the form with 
somewhat hairy pods. A glandular variant (var. glandulosa 0. E. 
Schulz, loc. cit.) is known from Ecuador. Neg. 13930. 

Arequipa: Nevado de Chachani, 4,300-4,600 meters, Pennell 
13304, 13288. Puno: Altos de Toledo, Meyen. San Antonio (Lech- 
ler 1799). Lima: Chicla, 3,720 meters, Weberbauer 245, type; 169. 
Matucana, Martinet 195. Cuzco: Pampa de Anta (Herrera 675). 
Bolivia; Chile. 

Descurainia Macbridei 0. E. Schulz, Field Mus. Bot. 8: 
81. 1930. 

A biennial, sometimes tall, often several decimeters high, with 
long, very leafy branches from the base, cinereous-pubescent with 
short, substellate hairs; leaves 1.5-6 cm. long, otherwise similar to 
those of D. myriophylla; inflorescence at an thesis capitate, then 
greatly elongating, the pedicels 2-4 (-8 in fruit) mm. long; sepals 
2 mm. long, the petals 3 mm. long, pale yellow, narrowly obovate- 
cuneate; pods 6-7 mm. long, 1.5-1.8 mm. thick; seeds 0.6 mm. long. 

Junin: In rock crevices and on bunch grass slopes, Huaron, 4,000 
meters, 1121, type. 

Descurainia myriophylla (Willd.) R. E. Fries, Nov. Act. Soc. 
Upsala IV. No. 1: 143. 1905; 335. Sisymbrium myriophyllum Willd. 
exDC.Syst. 2:477. 1821. 

Biennial, often tall, densely leafy, shortly substellate-canescent, 
the branches ascending; leaves 3-10 cm. long, with 6-10 pairs of 
pinnate-lobate leaflets; racemes at first dense, corymbiform, elon- 
gating to 120 flowers with pedicels 2-3 mm. long, these in fruit to 
7 mm.; sepals 2.2 mm. long, the petals 3 mm. long; ovules 75-85; 
pods 10-14 mm. long, 1.5 mm. thick; seeds 0.6-0.75 mm. long. 
Var. glandulosa Schulz, op. cit. 336, is known only from the "middle 
Andes," collected by Ball; it is the glandular state. Negs. 13931, 
34566. 

Cuzco: Wet slope in open canyon, La Raya, 4,400-4,500 meters, 
Pennell 13512. Huanuco: River canyon, 2,100 meters, Llata, 2266. 
Lima: Obrajillo (Wilkes Expedition). Puno: Azangaro, Lechler 
1729. Puno, 4,000 meters, Soukup 216; Isern 2114. Ayacucho: 
Coracora, grass steppe, Weberbauer 5805; 186. Arequipa: Isern 
2112. Colombia; Ecuador. 



FLORA OF PERU 983 

Descurainia Perkinsiana Muschl. Bot. Jahrb. 49: 199. 1913; 336. 

Like D. myriophylla but, so far as known, glandular, and the 
greenish leaflets with only about 4 lobes; pedicels 3-4 mm. long; 
petals 3-4 mm. long, drying whitish violet; pods often of a beautiful 
violet color. Illustrated, Pflanzenreich IV. 105: 337. 

Puno: Pisaloma and Titicaca, Meyen. Puno, 4,000 meters, 
Soukup 101. Have, Soukup 559. Isla del Sol, Titicaca, Seler. 
Cuzco: Huasao (Herrera). Junin: Near Tarma, 3,900 meters, 1055; 
petals dark yellow; sepals red. Arequipa: Nevado de Chachani, 
3,700 meters, Pennell 13283. Arequipa, 2,600-2,700 meters, Pennell 
13208. North of Arequipa, Pampa, 3,355 meters, Hinkley & Hink- 
ley 14. 

Descurainia pulcherrima Muschl. Bot. Jahrb. 49 : 200. 1913 ; 336. 

Similar to D. myriophylla, but typically more or less glandular; 
lower leaves (soon caducous) about 8 cm. long, the ultimate lobes 
oblong; upper leaves only 1-2 cm. long, with minute, acute lobes, all 
the leaves cinereous-tomentose, the upper often also glandular. 
Sometimes 2.5 meters high. Apparently restricted to rocky or partly 
shrubby slopes. Neg. 13933. 

Moquehua: Carumas, 3,000 meters, Weberbauer 7286. Lima: 
Matucana, 2,400 meters, 165. Viso, 2,700 meters, 635. Huanuco: 
Chavinillo, 2,400 meters, 1974. Yanahuanca, 3,000 meters, 1191. 
Bolivia; Argentina. 

Descurainia titicacensis (Walp.) Lillo (Fitogeogr. Tucuman 
228. 1919) ex Hauman & Irigoyen, Anal. Mus. Nac. Hist. Nat. B. A. 
32: 260. 1923; 338. Sisymbrium titicdcense Walp. Nov. Act. Acad. 
Leop.-Carol. 19: Suppl. 1: 249. 1843. 

Plants 10-20 cm. high, sometimes perennial; leaves and assur- 
gent branches cinereous-tomentose; leaves 1-3 cm. long, with 4-5 
pinnae, their oblong lobes 4-8 mm. long; racemes 20-30-flowered ; 
petals 3-4 mm. long, yellow; ovules 38-56. Schulz has designated 
var. dasycarpa and var. glandulosa, to care for these variations, 
apparently usual for each species. Neg. 13936. 

Puno: Pisacoma and Titicaca, Meyen, type; Weddell 4450. 
Cuzco: Pucara, 3,700 meters, Weberbauer (glandular). Prov. Can- 
chis, Sicuani (Hicken). Junin: Tarma region, 4,000 meters, Weber- 
bauer 2545. La Quinua, Poeppig 1830. Ayacucho: At 4,600 meters, 
back of Pisco, Weberbauer 5443. Lima: Obrajillo, Poeppig 1829. 
Casapalca, 4,800 meters, 827. Tacna: Tatora, Weddell. Argentina. 
"Occe-ccora." 



984 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

80. CAPPARIDACEAE Lindl. Caper Family 

Reference: Eichler in Mart. Fl. Bras. 13, pt. 1: 237-292. 1865; 
Pax & Hoffmann, Pflanzenfam. ed. 2. 17b: 146-223. 1936. 

Best known product of the family is the capers of commerce, 
obtained from Capparis spinosa L. of the Mediterranean region. 
Capers (alcaparras) are the pickled fruits or flower buds, used as a 
condiment. 

Fruit capsular; plants usually herbaceous or subherbaceous. 

Filaments free only from about the middle of the stalklike gyno- 

phore. 

Leaves usually compound; pods valvate-margined, not con- 
torted 1. Gynandropsis. 

Leaves simple; pods not valvate-margined, contorted after 

dehiscence 2. Podandrogyne. 

Filaments entirely free 3. Cleome. 

Fruit baccate; shrubs or trees. 
Leaves simple. 

Sepals free or partially connate; stamens 4 to many. 

Calyx lobes free or the 2 outer ones joined; fruit rarely globu- 
lar 4. Capparis. 

Calyx lobes united, parting irregularly at an thesis; fruit globu- 
lar 5. Morisonia. 

Sepals connate into a tubular-campanulate calyx; stamens 6; 

flowers reddish-tomentose 6. Steriphoma. 

Leaves compound 7. Crataeva. 

1. GYNANDROPSIS DC. 

Pedicellaria Schrank. 

Herbaceous or more or less shrubby plants with 3-7-foliolate 
leaves and rather showy flowers with conspicuously exserted stamens 
and pistil. Lower portion of the 6 filaments united to the stalklike 
gynophore. Fruit a narrow or oval capsule. The name Gynandropsis 
has been conserved. 

Bracts simple, sometimes caducous. 

Plant a shrub with glabrous capsule G. brachycarpa. 

Plants suffrutescent or herbaceous, the capsule usually pubescent. 
Plants pubescent, often woody below, with oval or club-shaped 
capsules. 



FLORA OF PERU 985 

Pubescence of the stems rather dense, the hairs often unequal; 

leaflets usually about 2 cm. wide. 

Bracts present, at least at anthesis G. densiflora. 

Bracts caducous. 

Pubescence rather dense, all the hairs spreading. .G. Ulei. 

Pubescence moderate, only the longer hairs widely spread- 
ing G. Mathewsii. 

Pubescence sparse, the remote, weak hairs subequal; leaflets 

narrower G. hispidula. 

Plants glabrous or glabrate, annual, with linear capsules. 

Petioles 10-12 mm. long; capsule short-stipitate . .G.Herrerae. 
Petioles 10-30 mm. long; capsule long-stipitate . .G. speciosa. 
Bracts trifoliolate G. triphylla. 

Gynandropsis brachycarpa (Vahl) DC. Prodr. 1: 238. 1824. 
Cleome br achy car pa V ah} ex DC. loc. cit. in syn.; Triana & Planch. 
Ann. Sci. Nat. IV. 17: 73. 1862. 

A shrubby, glandular-hirsute plant with 5-7-foliolate leaves, 
ebracteate racemes, and broadly oblong or club-shaped, glabrous cap- 
sules 3 cm. long; pedicels about 2 cm. long; calyx segments narrow, 
acute; petals broader, twice longer, rose-colored; stipes 3-4 cm. long. 

Huanuco: Pampayacu, Kanehira 291. Without locality (Jus- 
sieu). Colombia. 

Gynandropsis densiflora Benth. PL Hartw. 160. 1845. Cleome 
densiflora Triana & Planch. Ann. Sci. Nat. IV. 17: 72. 1862. 

Plants slender, suffrutescent, the stems viscid-hirsute, with stri- 
gillose hairs intermixed; capsule ovate-oblong, glandular-hirsute, 
becoming glabrate, the stipe about 2 cm. long. See also G. Ulei and 
G. Mathewsii, to which segregates the following specimens may be 
referable. 

Junin: Region of Huacapistana, Weberbauer 2337; 252. Rio 
Masamerich, Prov. Jauja, 1,600 meters, Weberbauer 6653. Hua- 
nuco: Chinchao, Ruiz & Pavon (det. Sleumer). Colombia; Ecuador; 
Venezuela. 

Gynandropsis Herrerae Macbr. Field Mus. Bot. 4: 168. 1929. 

An annual herb, very distinct from other Peruvian species by its 
linear, short-stipitate capsule, about 5 cm. long, and its almost com- 
plete lack of pubescence; upper leaves all 3-foliolate, the leaflets 
6-8 cm. long; bracts absent or caducous; petioles 10-12 mm. long. 



986 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Related, perhaps, to G. gynandra and G. triphylla, but the upper 
leaves 3-foliolate, the bracts entire and caducous, and the capsules 
subsessile. 

Cuzco: Near Rio Yanamayo, below Pillahuata, 2,000-2,300 
meters, in forest, Pennell 14073, type; petals "flesh ocher." 

Gynandropsis hispidula DC. Prodr. 1: 238. 1824. Cleome 
hirsuta R. & P. ex DC. loc. cit. in syn. 

Subherbaceous or perhaps slightly shrubby, like the related G. 
Mathewsii; pubescence soon rather sparse; leaflets about 3 cm. long, 
12 mm. wide; bracts persistent; flowers reddish yellow; capsule his- 
pidulous. Negs. 8461, 5812. Weberbauer 4450 (Neg. 5812) was indi- 
cated by Gilg as a new species of Pedicellaria, still unpublished. 

Peru: Mountains of Huanuco, Ruiz & Pavon. Pueblo Nuevo, 
Ruiz & Pavdn. Amazonas: Chachapoyas, Weberbauer 4450(1}. 

Gynandropsis Mathewsii Briq. Ann. Cons. Jard. Bot. Geneve 
17: 387. 1914. 

Similar to the last preceding species but with a fine, crisped pu- 
bescence mixed with longer, spreading hairs, and distinguished from 
G. Ulei by the pubescence and the caudate-acuminate sepals, longer 
than the pink petals. The glabrous or minutely granular-glandular, 
oval capsules are mostly about 2.5 cm. long, and 8 mm. wide, slightly 
longer than the stipe, this 2 cm. long. Neg. 8464. 

Huanuco: Rio Huallaga canyon, below Rio Santo Domingo, 1,200 
meters, 4213. Without locality, Mathews 193, type (in flower; co- 
type, Paris, in fruit). Pampayacu, Poeppig 1530. Junin: Chancha- 
mayo Valley, 1,500 meters, Schunke 266. Cuzco: Cerca al puente de 
Machupicchu, 1,800 meters, Vargas 524. 

Gynandropsis speciosa (HBK.) DC. Prodr. 1: 238. 1824. 
Cleome speciosa HBK. Nov. Gen. & Sp. 5: 84. pi. 436. 1821. 

Readily known by its entire, roundish or one-sided, cordate-based 
bracts (or the 1 or 2 lowest trifid), its showy, pink flowers (petals 
2.5-3 cm. long), and slender capsules 3-8 cm. long, borne on stipes 
about as long. Glabrous or somewhat pubescent; type Colombian; 
now adventive in the Orient and elsewhere. 

Loreto: Fortaleza, Yurimaguas, 200 meters, Williams 4459. Iqui- 
tos, 100 meters, Williams 8136. Caballo-cocha, Williams 2391 . Para- 
iso, Williams 3236. San Martin: San Roque, a weed and also in 



FLORA OF PERU 987 

cultivation, 1,400 meters, Williams 7052, 7175. La Victoria, Williams 
2600. Pebas, Williams 18SO. Widely distributed or adventive. 
"Plumaje de la reina," "flor de la viuda." 

Gynandropsis triphylla (L.) DC. Prodr. 1: 237. 1824. Cleome 
triphylla L. Sp. PI. ed. 2. 938. 1763. 

A glabrous or nearly glabrous annual with trifoliolate leaves and 
bracts; leaflets 2-3 cm. long; petals 6 mm. long; pedicels about 1 cm. 
long; capsule oblong-linear, about 4 cm. long, the stipe half as long. 
Near the widely distributed G. gynandra (L.) Briq. (G. pentaphylla} , 
but all the leaves 3-foliolate. The latter species, to be expected, may 
be known by its larger (about 1 cm. long) flowers. It is illustrated in 
Fawc. & Rendle, Fl. Jamaica 3, pt. 1: 229. 

Peru(?): Pawn. West Indies(?). 

Gynandropsis Ulei Briq. Ann. Cons. Jard. Bot. Geneve 17: 
385. 1914. 

Distinguishable, perhaps, from G. Mathewsii by the more abun- 
dant, soft, spreading, glandular indument, but not clearly distinct 
from G. densiflora Benth., which typically is more hirsute. As sug- 
gested by Briquet, op. cit. 388, these three species may be forms of 
a single variable one. Negs. 1184, 5818. 

San Martin: Pongo de Chilcayo, Tarapoto, Ule 64-30, type. 

2. PODANDROGYNE Ducke 

Tall herbs with simple leaves and dense racemes of monoecious 
flowers, the lower ones pistillate. Genus anomalous, connecting the 
capsular and baccate groups, the fruit capsule-like, contorted after 
the dehiscence of the 2 valves. Replum lacking. Sepals almost free. 

Podandrogyne glabra Ducke, Arch. Jard. Bot. Rio Jan. 5: 115. 
pi. 7. 1930. 

Glabrous, unbranched, unarmed, a meter high, with thin, ovate- 
oblong-elliptic, acuminate leaves 15 to nearly 30 cm. long and 6-12 
cm. wide; racemes to 20 cm. long; sepals 4-5 mm. long, acuminate; 
petals red, obovate-oblong, to 1 cm. long; stipe 3.8-4 cm. long; 
style extremely short; seeds black, lustrous, 3-4 mm. broad. Negs. 
5796, 1185. 

Loreto: Hacienda Indiana, mouth of Rio Napo (Ducke 19701, 
type). Soledad, Rio Itaya, 110 meters, in forest, Killip & Smith 
29594; a shrub 1 meter high. San Antonio, Killip & Smith 29480; 
herb 60-120 cm. high. Ecuador. 



988 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Podandrogyne orba Macbr., comb. nov. Gynandropsis orba 
Macbr. Candollea 5: 359. 1934. 

Perhaps distinct from the type of the genus, P. glabra, the ovary 
with a style about 0.5 mm. long; one stamen lower than the others; 
sepals narrowly lance-acuminate, 3 mm. long. Apparently the leaf 
nerves are fewer, about 6 or 7, and mostly 2 cm. apart. 

Junin: Rio Mayo, Weberbauer 6760, type. Pichis Trail, 650-900 
meters, Killip & Smith 26125. 

3. CLEOME L. 

Annual herbs or sometimes shrubs or even small trees, with 3-11- 
foliolate leaves, or these rarely simple, and often showy flowers 
with long, usually much exserted stamens. Stamens 4-6, the fila- 
ments free. Fruit a slender or short and thick capsule, often borne 
on a stipe. Indebtedness is acknowledged to Heilborn, Arkiv Bot. 
23, No. 10: 1-19. 1930, for his helpful treatment of a number of the 
shrubby and half-shrubby species centering about C. gigantea and 
C. glandulosa. 

Leaves all or mostly 3-5-foliolate; petals clawed; annuals or biennials, 

rarely more enduring and woody below. 
Plants provided with at least stipular prickles. 

Capsule sessile; leaflets 3 C. aculeata. 

Capsule more or less stipitate. 

Stipes and capsules 10-15 cm. long; plants subscandent. 

C. longipes. 
Stipes and capsules much shorter. 

Petals about 2 cm. long C. spinosa. 

Petals much shorter. 

Plants somewhat glandular-pubescent. 

Stipes 1 cm. long, the capsule much longer. 

C. psoraleaefolia. 
Stipes 3 cm. long, the capsule 1-2 cm. long. 

C. brachypoda. 

Plants glabrate C. virens. 

Plants unarmed, even stipular prickles lacking. 

Leaflets linear; capsule sessile, deflexed C. deflexa. 

Leaflets usually, perhaps always, broader than linear (C. deflexa 

imperfectly known). 
Bracts present. 
Bracts ternate. 



FLORA OF PERU 989 

Flowers about 10 mm. long C. chilensis. 

Flowers 3.5-6 mm. long. 

Flowers 5-6 mm. long; stipe of the capsule about 1 cm. 

long C. limoneolens. 

Flowers 3.5 mm. long; stipe about 4 mm. long. 

C. Mathewsii. 
Bracts all or nearly all entire. 

Flowers about 5 mm. long C. pachystigma. 

Flowers (7) 10 mm. long or longer. 
Flowers scarcely 1 cm. long. 

Uppermost leaves entire C. purpurea. 

Uppermost leaves 3-foliolate C. rosea. 

Flowers about 1.5 cm. long or longer. 

Stems more or less long-pubescent C. pubescens. 

Stems glabrous or glabrate C. longifolia. 

Bracts caducous at an thesis C. virens. 

Leaves, at least some of them, 7-foliolate; petals attenuate at the 

base; mostly shrubby plants, at least below, rarely prickly 

(C. limoneolens, C. chilensis, and C. pubescens might be sought here) . 

Racemes ebracteate in flower, at least above, the small bracts 

promptly deciduous. 

Stems hirsute- villous C. hirsuticaulis. 

Stems glabrous or short-pubescent. 

Style well developed, to 1 cm. long in fruit. . .C. monochroma. 
Style short or obsolete. 
Leaflets pubescent beneath; capsule pubescent, 17-20 cm. 

long C. gigantea. 

Leaflets sparsely glandular beneath, obscurely pubescent 
or glabrate; capsule 10-12 cm. long. . .C. amazonica. 
Racemes bracteate, the bracts conspicuous. 
Plants eglandular or nearly so, often subherbaceous, at least 

above; capsule linear-cylindric, glabrous. 
Petals 2-2.5 cm. long. 

Capsule 6 cm. long or longer; sepals rotund-ovate. 

C. Figueroae. 
Capsule 4 cm. long; sepals lanceolate, acuminate. 

C. longifolia. 
Petals 10-17 mm. long. 



990 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Bracts suborbicular, obtuse C. Herrerae. 

Bracts ovate, acuminate C. potamophila. 

Plants more or less viscid-glandular; shrubs or small trees; 

capsule often glandular. 

Sepals narrowly lanceolate, much shorter than the petals; 
pubescence conspicuously stipitate-glandular. 

C. glandulosa. 

Sepals ovate-lanceolate, shorter than the petals, pubescent 
but scarcely glandular C. Lechleri. 

Cleome aculeata L. Syst. ed. 12. 2: 232. 1768. 

A diffuse, glandular-puberulent herb with 3-foliolate leaves, or 
the uppermost leaves entire, and with stipular prickles; sepals 2-3 
mm. long, the petals about twice as long, white; pedicels 1-2.5 cm. 
long; capsule 2.5-5 cm. long. This is the usual application of the 
Linnean name; but Sleumer in herb. Madrid has referred the Ruiz 
and Pavon specimens to C. spinosa L., which must mean a change 
in interpretation, unfortunate considering how well established both 
names are. Neg. 5774. 

Huanuco: Posuso, 600 meters, 4-648; Ruiz & Pavdn. San Martin: 
Tarapoto, 400-900 meters, Williams 5583. Without locality: Ruiz 
& Pavon 25-52. Tropical America and Africa. 

Cleome amazonica Heilborn, Arkiv Bot. 23, No. 10: 9. 1930. 

A more or less glandular, suffrutescent plant, the glands obscure 
and sessile; leaflets sessile or subsessile, very sparsely pilose, acumi- 
nate; sepals and petals linear-oblong, subequal; stigma subsessile, 
or the style in fruit 2-4 mm. long; stipe about 5 cm. long; capsule 
glabrous or nearly so, 10-12 cm. long, 7-9 mm. thick. Neg. 18200. 

Rio Acre: Monte Mo, Ule 9411, type. 

Cleome brachypoda Briq. Ann. Cons. Jard. Bot. Geneve 17: 
378. 1914. 

Similar to C. psoraleaefolia, but petals only a few millimeters long, 
and the capsule thick-cylindric, 1-3 cm. long, nearly 5 mm. thick, the 
stipe scarcely 3 mm. long. Neg. 5765. 

Loreto: Pebas, in forest, Williams 1956. Brazil. 

Cleome chilensis DC. Prodr. 1: 238. 1824. 

This herbaceous annual, sometimes confused with C. aculeata, 
which has sessile capsules, is well marked by its 3-foliolate bracts. 
The author distinguished two varieties: glabra, with 7-foliolate leaves, 



FLORA OF PERU 991 

and pubescens, with 3-5-foliolate leaves. One of the type sheets 
(Paris) of the glabrate plant is labeled "Cheuchin," and the mature 
pods and stipes are equal in length, about 2 cm. long; the single speci- 
men of the more viscid-pubescent variety has pods 2.5-3 cm. long, 
equaled by the stipe, and purports to come from Chile, although the 
locality "Cheuchin" has been written on the label; none of the mate- 
rial shows stipular spines. Illustrated, Deless. Icon. 3: pi. 2. Neg. 
34611. 

Lima: Amancaes, Weberbauer 1616; Ruiz & Pawn. Cheuchin, 
Dombey, type. San Mateo, Isern 2189. San Buenaventura, 2,700 
meters, open, rocky slopes, Pennell 14512. Atocongo, 250-500 
meters, open, sandy hill, Pennell 14781. Matucana, 2,400 meters, 
175, 559; Sauvatier 366 (nearly var. pubescens!}; Weberbauer 55. 
Cajamarca: San Miguel, Weberbauer 3920. Arequipa: Mollendo, 
Hitchcock 22376. Cuzco: Valle del Urubamba, Ollantaitambo, 2,800 
meters, Herrera 3422. Bolivia. "Inca-lacma." 

Cleome deflexa DC. Prodr. 1: 240. 1824. 

A glandular herb with 3-foliolate leaves, the upper leaves simple, 
like the leaflets oblong-linear; capsule glabrous, acuminate, sessile, 
deflexed. By Robert Brown said to be scarcely distinct from 
C. violacea L., with the suggestion that it may not be Peruvian. 

Peru(?): Without locality, in Herb. Lambert (Ruiz & Pavdri). 

Cleome Figueroae Macbr. Field Mus. Bot. 4: 168. 1929. 

Plants slender, suffrutescent, 1 meter high or more, with minutely 
pubescent, 7-foliolate leaves, rotund bracts, pale yellow flowers with 
glabrous, rotund-ovate, acute sepals to 5 mm. long, and glabrous, 
linear capsules 6 cm. long, exceeded by their slender stipes. 
Related to C. longifolia Presl. Named for Reyes Figueroa, capable 
guide and friend. 

Huanuco: Huacachi, near Muna, 1,950 meters, in old clearing, 
4173, type. 

Cleome gigantea L. Mant. 2: 430. 1771. 

A slender, strict, somewhat glandular-pubescent shrub 0.5-2 
meters high, with ebracteate racemes and glabrous capsules; leaves 
7-foliolate, densely pilose-hirsute beneath; flowers only 3 cm. long, 
brown. In Peru according to Heilborn, but no specimens cited by 
him ; see C. monochroma. 

Huanuco: Chinchao, Ruiz. Without locality, Ruiz & Pavon 
25-56. Cuzco: Urubamba Valley, 1,000 meters, Weberbauer 7955. 



992 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Juanjui, 400 meters, in forest, Klug 3874- Tropical 
South America. 

Cleome glandulosa R. & P. ex DC. Prodr. 1: 238. 1824. 

A shrub or small tree with very glandular-hispid branches, leaves, 
and capsules; leaves 7-9-foliolate, the leaflets ovate-lanceolate; 
sepals narrowly ovate-lanceolate, about half as long as the petals, 
these becoming 2 cm. long; pedicels 1.5 cm. long; capsule and stipe 
each 4-5 cm. long.- My specimens were from small trees with 
smooth trunks about 1.5 meters high and short crowns of few, very 
leafy branches. The abundant, yellow glands of foliage and branches 
were pungently and disagreeably scented. The Quechua name 
seemed to be "ancas tucapinq." C. Jamesonii Briq., of Ecuador, 
has narrowly lanceolate leaflets and slender stipes more than twice 
as long as the pods. Weberbauer 3203 from Ancash is similar to 
C. glandulosa but the pubescence is almost tomentose and the cap- 
sules are glabrous. It is indicated in herb. Berlin as a new species. 
Neg. 5772. 

Without locality: (Ruiz; Dombey). Huanuco: Mito, 2,700 
meters, 1692. Junin: Tarma, Ruiz & Pavdn; Dombey; Isern 2007. 
Huariaca, 2,850 meters, 3094. Chanchamayo Valley, between 
Huacapistana and Palca, Weberbauer 1760; 246. 

Cleome Herrerae Macbr. Field Mus. Bot. 4: 169. 1929. 

Similar, perhaps, to C. longifolia Presl, but petals about 17 mm. 
long, sepals 6-7 mm. long, not attenuate at the base, capsules about 
5 cm. long, stipes 3.5 cm. long, and pedicels only 1.5 cm. long. 
C. tunariensis Kuntze, Bolivian, little known, has bracts to 4 cm. 
wide and green petals (ex char.). C. Bangiana Gilg ex Heilborn, 
Bolivian, has subequal sepals and petals. 

Cuzco : Valle de San Miguel, Cedrobamba, 2,260 meters, Herrera 
1992, type. Puno: Churumayo, Soukup 547. "Facma." 

Cleome hirsuticaulis Macbr. Candollea 5: 360. 1934. 

An erect, strict, unarmed shrub, 2.5-3 meters high; stems and 
pedicels densely glandular-pubescent with firm, spreading hairs; 
leaflets oblong-lanceolate or oblong-elliptic, almost caudate-acumi- 
nate, cuneate at the base, to 14 cm. long, 4 cm. wide, very thin, 
sparsely hirsutulous above, glabrous beneath except on the costa; 
pedicels 4-6 cm. long, about equaled by the stipes; capsule 14-17 
cm. long, to 8 mm. in diameter, narrowly fusiform, attenuate at 
the apex, short-pilose; style 3 mm. long. 



FLORA OF PERU 993 

Junin: Hacienda Schunke, above San Ramon, 1,400-1,700 
meters, in forest, Killip & Smith 24812, type. 

Cleome Lechleri Eichl. in Mart. Fl. Bras. 13, pt. 1: 250. 1865. 

A viscid, glandular-tomentose plant, related to C. glandulosa, 
but with oblong or ovate-lanceolate sepals one-half to two-thirds 
as long as the petals; bracts rotund; capsules and stipes subequal, 
or the latter longer than the pedicels, sparsely glandular-setulose 
or glabrous. The pubescence is more viscid than glandular. C. 
ecuadorica Heilborn and C. chrysogyna Gilg, also Ecuadorean, 
apparently too near C. Lechleri, have glabrous pods, those of the 
former 3 times as long as broad, exceeded by the stipe; of the latter 
5 times as long as broad, and nearly as long as the stipe. The 
Weberbauer collections 4909 and 5652 are annotated by Gilg in 
herb. Berlin as new, a disposition apparently vouched for by Heil- 
born, but he does not supply descriptions nor distinguish them in 
his key. The presence or absence of glands, unless supported by 
other characters, is open to question as a specific character. Neg. 
8453. 

Ayacucho: Prov. Huanta, 3,200 meters, Weberbauer 5652(1). 
Cuzco: Urubamba, Weberbauer 4909(1). Puno: Tabina, Lechler 
1859, type. 

Cleome limoneolens Macbr. Field Mus. Bot. 4: 169. 1929. 

Similar to C. chilensis, but woody at the base, the flowers only 5-6 
mm. long. The foliage is lemon-scented. 

Junin: San Rafael, 2,550 meters, 3145, type. Huanuco: Hua- 
cachi, near Muna, 1,950 meters, 4088. 

Cleome longifolia Presl, Rel. Haenk. 2: 84. 1835. 

An unarmed herb or shrub, apparently related to C. spinosa 
Jacq.; leaflets 5-7, to 10 cm. long, petiolate, pubescent on both sides, 
narrowly lanceolate, long-acuminate; petals 2 cm. long, 4 times as 
long as the lanceolate, acuminate sepals, these 6 mm. long; stigma 
sessile; capsule glabrous, linear, 4 cm. long. 

Huanuco: In the mountains (Haenke, type). 

Cleome longipes DC. Prodr. 1: 239. 1824. 

An almost glabrous, prickly, woody-stemmed (but wholly herba- 
ceous above), more or less scandent plant, to 3 meters long, with 
3-foliolate leaves, a crowded inflorescence, and very slender, long- 
stipitate, pendulous capsules as much as 15 cm. long, little exceeding 



994 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the filiform stipes; flowers scarcely 2 mm. long, creamy white, with 
dull purplish brown stamens. Negs. 7965, 5774. 

Loreto: Prov. Mainas, Poeppig. Puerto Arturo, in forest, 200 
meters, Williams 5142. Junin: Puerto Bermudez, 375 meters, in 
thicket, Kittip & Smith 26629, 26681. Ecuador to Costa Rica. 

Cleome Mathewsii Briq. Ann. Cons. Jard. Bot. Geneve 17: 374. 
1914. 

A somewhat viscid-pilose herb with 5-foliolate leaves, small 
flowers, and very shortly stipitate, sparsely glandular or asperous, 
pendent pods 2.5-3 cm. long, the spreading pedicels to 1.5 cm. long. 
-Neg. 8454. 

Without locality: Mathews 3044, type. 

Cleome monochroma Macbr. Field Mus. Bot. 4: 169. 1929. 

A slender, subherbaceous annual or biennial several decimeters 
tall, with large, 7-foliolate, sparsely pubescent leaves, villous and 
glandular, greenish white flowers 3.5-4 cm. long (the nearly linear 
sepals and petals of the same color), and long-acuminate capsules 
10-14 cm. long, nearly 1 cm. wide at base, borne on glabrous stipes 
and pedicels about half as long; leaflets sessile, attenuate at each 
end, softly subappressed-strigillose on both sides. Probably C. 
gigantea L. as to Peru, but perhaps with C. Kerberi Briq. and 
C. amazonica Heilborn races of one species; but the Mexican plant is 
less pubescent, the style on the ripe pod 1 cm. long or longer, the 
stipes pubescent; and the Linnean plant has a sessile stigma. Pax 
and Hoffmann list C. monochroma among the Herbaceae, apparently 
overlooking my own reduction of it at one time to C. gigantea. 

Huanuco: Chinchao, Ruiz & Pawn. Cochero, Poeppig 1242. 
Junin: La Merced, 600 meters, 5386, type. Cuzco : Urubamba Valley, 
Weberbauer 7955. Ecuador. 

Cleome pachystigma Briq. Ann. Cons. Jard. Bot. Geneve 17: 
377. 1914. 

A glandular-puberulent or almost glabrous herb with 5-foliolate 
leaves, rose-colored flowers 5 mm. long, and glabrous, linear capsules 
about 3 cm. long, tipped with a caplike stigma 2 mm. broad; stipes 
1-1.5 cm. long. Neg. 27282. 

Peru(?): (Pavon, type). 

Cleome potamophila Gilg in herb., sp. nov. 
Eglandulosa vel fere eglandulosa, praesertim ad caules plus 
minusve dense furfuraceo-tomentosa; foliola supra glabrata, ca. 9 cm. 



FLORA OF PERU 995 

longa, utrinque acuminata; bracteae ovatae acuminatae; sepala 6 
mm. longa 4 mm. lata; petala 13-14 mm. longa; capsula glabra; 
aliter C. Lechleri similis; frutex 3-metralis, floribus viridi-flavescenti- 
bus. "Difficult to classify, but recognized by Gilg as new" (Heil- 
born). Neg. 5776. 

Ancash: Pampa Romas, Weberbauer 3203 (type in herb. Berlin). 

Cleome psoraleaefolia DC. Prodr. 1: 239. 1824. 

Allied to C. spinosa; leaflets mostly 5; pilose pubescence very 
sparse or typically rather long, dense, and viscid; flowers pinkish, 
5-7 mm. long; pedicels about 2 cm. long, the stipes to 2.5 cm. long, 
the thick-linear, strongly verruculose, finally glabrous capsule 
twice as long or longer. In the type the stipes are 2.5 cm. long, 
but in most specimens they are much shorter. 

Loreto: Iquitos, 100 meters, Williams 1397, 14.68. San Martin: 
Juanjui, 400-800 meters, in clearing, Klug J+232; plants 1 meter 
high; flowers white and violet. Brazil; Uruguay. 

Cleome pubescens Sims, Bot. Mag. pi. 1857. 1816. 

A vigorous, more or less glandular-pubescent herb with 5-7- 
foliolate leaves and white flowers 15-17 mm. long; capsule linear- 
cylindric, 7-10 cm. long, the stipe to 6 cm. long. Reported from 
Peru, but the material so named seen by me is referable to other 
species. Described from garden plants of unknown origin. Seems 
to be C. spinosa without prickles. 

Peru: Doubtful. Eastern South America; Panama; Hispaniola. 

Cleome purpurea DC. Prodr. 1: 239. 1824. 

A glabrous herb with 3-5-foliolate leaves, the floral and upper 
ones simple, ciliate; capsule unknown, the ovary glabrous, twice the 
length of the stipe. Possibly referable to C. rosea Vahl. 

Peru: (Herb. Lambert, type). 

Cleome rosea Vahl ex DC. Prodr. 1: 239. 1824. 

Similar to C. purpurea, but even the uppermost leaves 3-5- 
foliolate; petals pink, to 1 cm. long; ovary somewhat pubescent, a 
little shorter than the stipe. Mature material referred here by 
Eichler is glabrate, the pedicels about 2 cm. long; stipes as long or 
longer; capsule 4-5 cm. long, 4 mm. wide, short-acuminate or 
beaked, not torulose, the stigma small. 

Loreto: Iquitos, 100 meters, Williams 3585. Brazil. 

Cleome spinosa Jacq. Enum. PI. Carib. 26. 1760. 



996 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A tall, more or less glandular herb with at least stipular prickles 
and mostly 5-7 leaflets (the middle leaves) ; flowers about 2 cm. long; 
bracts broad; capsule usually 5-10 cm. long, linear-cylindric, the 
stipe 2.5 cm. long; pedicels 2-3 cm. long. Illustrated, Gard. Chron. 
III. 45: 115; Fawc. & Rendle, Fl. Jamaica 3, pt. 1: 225. 

Piura: Negritos, Haught F18. Junin: Chanchamayo, Isern2186. 
Without locality, Ruiz & Pavon. Widely distributed in tropical 
America. 

Cleome virens Macbr. Candollea 5: 360. 1934. 

A shrub, but herbaceous above, 1-1.5 meters high, glabrous; 
stems and petioles (to 8 cm. long) aculeate or unarmed; leaflets 3, 
sometimes 5, ovate-elliptic, attenuate to the base, short-acuminate, 
the lateral ones 5-10 cm. long, 3-4 cm. wide, the slightly larger ter- 
minal one conspicuously petiolulate, all very thin; racemes finally 
20 cm. long; bracts caducous; pedicels nearly filiform, about 1 cm. 
long, in fruit to twice as long, widely divaricate; sepals narrowly 
lanceolate, acute, 2.5 mm. long; petals 4 mm. long, the blade elliptic- 
spatulate, the claw 1 mm. long; stamens 7 mm. long; capsule gla- 
brous, more or less torulose, linear-cylindric, 10-17 cm. long, the 
stipe scarcely 1 cm. long. Flowers grass-green and yellow, according 
to Klug. Apparently near C. serrata Jacq., with minutely serrulate- 
ciliate leaves and subsessile capsules; also related to C. latifolia 
Vahl, with elongate stipes. 

Loreto: La Victoria, Williams 2596, type; 2842. Iquitos, 120 
meters, edge of forest, Williams 8183 (unarmed form) ; Klug 1163, 487. 
Yurimaguas, 135 meters, in forest, Killip & Smith 27653. Mouth 
of Rio Santiago, 200 meters, old clearing, Mexia 6150; a vine with 
greenish flowers. Meuna-cano, near Iquitos, 105 meters, bank of 
creek in dense shade, Mexia 6502a; an herb 2.5 meters high; flowers 
greenish. Above Pongo de Manseriche, 200 meters, beside a stream- 
let, Mexia 6249; an herb 2 meters high with a woody base; leaflets 
2-5; flowers greenish. San Martin: Juanjui, 400-800 meters, river 
bank, Klug 4397; a vine; flowers green and yellow. 

4. CAPPARIS L. 

Shrubs or trees with simple, petiolate, often thick leaves. Pubes- 
cence, if present, usually stellate or lepidote. Flowers frequently 
large and white, with several to many stamens. Sepals free or 
partly connate, or more rarely the calyx 4-lobed or crenate. Pax 
and Hoffmann record C. cynophallophora L. for Peru, probably on 
the basis of material interpreted here as C. flexuosa or C. odora- 



FLORA OF PERU 997 

tissima. C. didymobotrys R. & P. Fl. Peruv. 4: pi. 423. 1802, also 
DC. Prodr. 1: 253. 1824, said to be from Guayaquil, may well prove 
to be an earlier name for a species listed here, but no material has 
been seen. 

Leaves glabrous or, if finely pubescent beneath, the hairs simple. 
Sepals small, in a single series. 

Leaves glabrous, firm or coriaceous. 

Stamens longer than the petals; gynophore soon longer than 

the stamens. 
Leaves retuse, obtuse, or obtusely acute, rarely abruptly 

acute. 
Petioles all or mostly elongate, 2 to several centimeters 

long. 

Leaves obtuse, rounded, or retuse at the apex, those 
of flowering branches rarely if ever wider than 
10 cm. 

Leaves somewhat narrowed at the rounded base. 

C. laurina. 
Leaves not at all narrowed at the broadly rounded 

base C. macrocarpa, C. Sprucei. 

Leaves abruptly short-acuminate, often wider than 

10 cm C. magnified. 

Petioles all short, 10-14 mm. long C. Quina. 

Leaves narrowly and acutely acuminate C. sola. 

Stamens barely as long as the petals; gynophore 5-8 mm. long. 

C. Baducca. 

Leaves finely pubescent beneath, thin, soft C. prisca. 

Sepals ample, more or less in 2 series, that is, imbricate. 
Leaves lance-oblong or oblong-obovate, 1-3.5 cm. wide. 

Leaves glabrous C. eucalyptifolia. 

Leaves puberulent-pilose beneath. 

Leaves lance-oblong, acutish C. mollis. 

Leaves oblong-obovate, emarginate C. prisca. 

Leaves elliptic, 2-4 cm. wide or wider. 

Leaves about 5 cm. long; calyx obtuse, open, about 7 mm. 

wide C. tarapotensis. 

Leaves often longer than 5 cm.; calyx always wider than 
10 mm. 



998 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves thin, drying dark, at least some of them acutely 

acuminate C. nitida. 

Leaves coriaceous, drying yellow or brown, usually obtuse. 

C. flexuosa. 
Leaves usually lepidote or stellate-pubescent beneath, sometimes 

sparsely and minutely so, as in C. Schunkei. 
Calyx small, crenate-lobed or deeply dentate. 

Calyx crenate-lobed C. ovalifolia. 

Calyx acutely dentate C. indica. 

Calyx ample, valvate in bud. 

Leaves distinctly longer than broad. 

Leaves lepidote beneath C. odoratissima. 

Leaves stellate-pubescent. 

Pubescence dense; leaves rigid, suboblong. 

Fruit ovoid-oblong C. angulata. 

Fruit subglobose C. Gaudichaudiana. 

Pubescence sparse; leaves thin, elliptic C. Schunkei. 

Leaves subrotund C. cordata. 

Capparis angulata R. & P. Fl. Peruv. 4: pi. 431. 1802; DC. 
Prodr. 1: 253. 1824. C. scabrida HBK. Nov. Gen. & Sp. 5: 95. 1821. 

Branchlets, leaves beneath, sepals, and petals ashy-stellate- 
tomentose; blades lustrous and somewhat rough above, coriaceous, 
subelliptic-oblong, obtuse at both ends, cuspidate, often 11 cm. 
long, 4-5 cm. wide; petioles to 15 mm. long; pedicels 2.5-5 cm. long; 
stipe 7 cm. long or longer; petals twice as long as the deeply lobed 
calyx, this nearly 1 cm. long; stamens elongate; fruit ovoid-oblong. 
The type, collected by Ruiz and Pa von, is from Guayaquil. 
Illustrated, Weberbauer, 152. Negs. 342, 29270. 

Piura: Negritos, HaughtF69; a shrub or small tree of 2-4 meters. 
Paita, d'Urville; Gaudichaud. Lambayeque (Gaudichaud) ; Weber- 
bauer 3777. Ecuador. "Sapote." 

Capparis Baducca L. Sp. PI. 504. 1753. C. frondosa Jacq. 
Enum. PL Carib. 24. 1760. 

A shrub or small tree; blades usually elliptic-oblong and some- 
what obovate, obtuse or slightly narrowed to an acutish apex, 
sometimes abruptly acuminate, minutely cordate at the base, 10-30 
cm. long, 3.5-10 cm. wide, not very lustrous above, prominently 
reticulate-veined on both sides, the lateral nerves conspicuous 



FLORA OF PERU 999 

beneath, chartaceous-coriaceous; petioles 0.5-5 cm. long; racemes 
scarcely equaling the leaves, even in fruit; calyx lobes roundish, 
barely 1.5 mm. long; petals greenish white or purplish, obovate, 
fugacious, 9-10 mm. long, equaled by the many stamens, these 
white, pink at the base; fruit 3-7.5 cm. long, purplish brown, some- 
what nodose, the stipe only 5-8 mm. long. Description adapted 
from Fawcett and Rendle. 

Loreto: Balsapuerto, 220 meters, in forest, Klug 3013; a tree 
of 5 meters; flowers white or rose. Extending to northern South 
America; West Indies; Central America and Mexico. 

Capparis cordata R. & P. Fl. Peruv. 4: pi. 432. 1802; DC. 
Prodr. 1: 251. 1824. C. crotonoides HBK. Nov. Gen. & Sp. 5: 95. 
pi. 437. 1821. 

Leaves ovate to suborbicular-cordate, 3-4 (-6) cm. long, nearly 
as wide, net-veined and densely stellate-pubescent, especially be- 
neath; flowers few, the densely stellate-pubescent sepals 5-6 mm. 
long, the sparsely stellate-pubescent petals twice as long; fruit 
globose, stellate-pubescent, 7 mm. thick. Flowers cream-colored, 
the sepals in one series, valvate. The Ruiz and Pavon plant came 
from Guayaquil. 

Piura: Paita, d'Urmlle. Truxillo, Humboldt. Quebrada Ancha, 
10 miles east of Talara, Haught F89; a shrub of 2 meters, common. 
Cape Parinas, Haught 196. Ancash: Yautan, 600 meters, gravelly 
river valley, 2568; a shrub of 3.5 meters; flowers pale cream. Ecua- 
dor; Bolivia. "Vichaya," "bichaya." 

Capparis eucalyptifolia Haught, Trop. Woods 20: 30. 1929. 

A small tree, closely related to C. mollis with imbricate sepals, 
but it is entirely glabrous from the first and the petals and stamens, 
when developed, are nearly twice as long. The Ecuadorean C. 
heterophylla R. & P. has elliptic, obtuse or bluntly pointed leaves 
5-7 cm. long and 2-3.5 cm. wide. 

Piura: Amotape Hills, Haught F96, type. Libertad: Near 
Chepe'n, Weberbauer 4814, in part. 

Capparis flexuosa L. Sp. PI. ed. 2. 722. 1762. C. cynophallophora 
L. pro parte; cf. Fawc. & Rendle, Journ. Bot. 52: 142. 1914. C. 
lanceolata R. & P. ex DC. Prodr. 1: 249. 1824. 

Leaves glabrous, very variable in shape, but usually elliptic, 
4-10 cm. long (1-) 2-4.5 cm. wide, on petioles about 5 (-10) cm. 
long and with an oblong, axillary gland; stamens about 3 times as 



1000 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long as the petals, these 1.5-2 cm. long; sepals 4-7 mm. long; fruit 
linear, 4-10 (-20) cm. long, the gynophore half as long. The Madrid 
Ruiz and Pavon specimen of C. lanceolata is from Guayaquil, and is 
referred by Sleumer to C. cynophallophora L. var. angustifolia Eichl. 
Peru: Probably. Ranging north to Mexico and Florida. 

Capparis Ga.udichaudiana Eichl. in Mart. Fl. Bras. 13, pt. 1: 
273. 1865. 

Very doubtfully distinct from C. angulata, since a series of 
Gaudichaud specimens exhibit leaves rounded to cuneate at the 
base and pedicels to 3 cm. long, even in flower, the only remaining 
distinctive character being the supposed difference in fruits. 
Neg. 8472. 

Piura: Paita, Gaudichaud, type. 

Capparis indica (L.) Fawc. & Rendle, Journ. Bot. 52: 143. 
1914. Breynia indica L. Sp. PI. 503. 1753. C. Breynia L. Syst. 
ed. 10. 1071. 1759. 

Leaves thin, oblong-elliptic, 5-10 cm. long, 2-4.5 cm. wide, pale 
beneath, with numerous scales; flowers about 1 cm. long, densely 
stellate- tomentose; calyx lobes open in bud, 2 to 3 mm. long; petals 
white, 10-12 mm. long; fruit linear, 5 cm. to several decimeters 
long. The herbarium record may be an error. 

Peru(?): Without locality, Pav6n(1}; det. Briquet. Ranging to 
Mexico and West Indies. 

Capparis laurina HBK. Nov. Gen. & Sp. 5: 93. 1821. 

Supposed to be related to C. Sprucei, from which it apparently 
differs in the globose fruit and in the leaves, these somewhat nar- 
rowed at the base, flexible-coriaceous, to. 15 cm. long, 6.5 cm. wide, 
often smaller, on petioles 2-3 cm. long. The type seems to me to 
be a mature specimen of C. macrocarpa. The lateral nerves and 
netted venation are the same, but more pronounced. The flowers 
are said to be violet, the reflexed calyx green, the tree handsome, 
with a rounded crown. Neg. 34628. 

Piura: Huancabamba, Humboldt & Bonpland, type. 

Capparis macrocarpa R. & P. Fl. Peruv. 4: pi, 540. 1802; 
DC. Prodr. 1: 250. 1824. C. petiolaris HBK. Nov. Gen. & Sp. 5: 
91. 1821. 

A shrub or small tree; at once recognizable by the rather thin 
but firm, glabrous, broadly elliptic (varying to elliptic-oblong) 



FLORA OF PERU 1001 

leaves, on slender petioles 2 to several centimeters long; fruit to 
8 cm. long and 2.5 cm. thick, but often shorter and about 2 cm. 
thick. In the HBK. type, in flower, the leaves are thin, but a 
netted venation is appearing; they are 10-12 cm. long and 4.5-5 cm. 
wide, on petioles 2-7 cm. long; petals 1.5 cm. long; calyx glabrous, 
the roundish divisions obtuse, reflexed; stamens several centimeters 
long. The Ruiz and Pavon type is from Guayaquil; its leaves are 
oblong-elliptic. It is not clear to me that C. laurina and C. Sprucei 
are distinct. Neg. 29272. 

San Martin: Tarapoto, Ule 6428; Williams -6889, 6903, 6786, 
6281, 6660, 6903, 6904. Ecuador. "Mango micuna," "nina caspi." 

Capparis magnified Gilg, Bot. Jahrb. 40: 140. 1907, nomen. 

Petioles obscurely pulverulent, to 15 cm. long; blades to 30 cm. 
long, 17 cm. wide, broadly elliptic, abruptly short-acuminate, the 
broad acumen less than 1 cm. long, acute at the broadly rounded 
base, membranous, drying pale green, slightly lustrous, concolorous, 
finely reticulate-veined on both sides, the 6-7 lateral nerves equally 
prominent on both surfaces. The leaves resemble those of Morisonia 
oblongifolia, but the petioles are not lepidote. 

Rio Acre: Mouth of Rio Macauhan, Krukoff 5788. Amazonian 
Brazil. 

Capparis mollis HBK. Nov. Gen. & Sp. 5: 88. 1821. 

Branchlets and under surfaces of the lanceolate leaves finely 
white-pubescent; blades tapering to an acutish apex, 5-8 cm. long, 
1-2.5 cm. wide; sepals in 2 imbricate series; petals about 1 cm. 
long, the stamens 2 cm. long. Negs. 344, 34630. 

Piura: Near Olleras, between Rio Macara and Huancabamba 
(Humboldt & Bonpland, type). Lambayeque: (Weberbauer, 153). 
Libertad: Near Chepe"n, 400 meters, Weberbauer 4814, in part. 
"Margarito." 

Capparis nitida R. & P. ex DC. Prodr. 1: 252. 1824. 

Related to C. mollis, but with glabrous, lustrous, usually acutely 
long-acuminate blades, these 10 cm. long, 3-4 cm. wide; petioles 
7 mm. long; flowers subsolitary in the upper axils or in few-flowered, 
terminal racemes; pedicels 15 mm. long; ovary ovoid, truncate; 
fruit subglobose, about 2 cm. thick, the stipe to 10 cm. long. 
Williams 4683 from the lower Rio Huallaga has obtusely short- 
acuminate leaves to 17 cm. long and 10 cm. wide, slightly stellate- 
pubescent beneath near the base; it doubtfully belongs here, but 



1002 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

there are no flowers. The Madrid type is without data. Negs. 
27289, 343. 

San Martin: Tarapoto, Ruiz & Pavdn; Spruce 4294; Williams 
5725, 5567. Juanjui, 400-800 meters, forest and river bank, King 
4369, 4352; a shrub or tree of 3-4.5 meters; flowers white or greenish 
white. Loreto: Yurimaguas, in forest, Williams 4683. "Yututo 
caspi." 

Capparis odoratissima Jacq. Hort. Schonbr. 1: 57. pi. 110. 1797. 

Leaves lustrous, coriaceous, oblong-elliptic, densely rusty-lepi- 
dote-pubescent beneath, like the inflorescences and the angulate 
fruits; flowers small, somewhat purplish, the petals little longer than 
the valvate sepals, the stamens only shortly exserted. The following 
herbarium record (det. Briquet) may be an error; see C. indica. 
C. cynophallophora L., similar, might occur; it has long-exserted 
stamens and stipitate fruits. C. Fiebrigii Gilg, known from Mexico 
and Bolivia, is a gnarled shrub with small, oblong leaves. 

Peru(?): Pavon (det. Briquet). Ranging to Central America 
and Venezuela. 

Capparis ovalifolia R. & P. Fl. Peruv. 4: pi. 432. 1802; DC. 
Prodr. 1: 253. 1824. C. avicennifolia HBK. Nov. Gen. & Sp. 5: 
94. 1821. 

A well marked species because of its unique calyx, this campan- 
ulate and merely crenate-lobate; leaves lustrous above, pale beneath, 
with a very close, stellate pubescence, usually 4-5 cm. long, 2-2.5 
cm. wide; fruit about 2.5 cm. thick. The Ruiz and Pavon type 
came from Guayaquil and was actually collected by Tafalla, the 
artist of the expedition. Fruit edible (Martinet). Neg. 27284. 

lea: Near lea, 350 meters, Weberbauer 7196. Libertad: Coastal 
sands between Trujillo and Cascas, Humboldt. Cajamarca: Near 
Chepe"n, Weberbauer 3778. Lambayeque: (Weberbauer, 153). 
Piura: Paita, Gaudichaud; d'Urville. Talara, 200 meters, Pennell 
12201; Johnston 3507. Farinas Valley, Haught F157; a shrub of 
2-3 meters. Ecuador. "Bichayo," "vichaya." 

Capparis prisca Macbr. Candollea 5: 358. 1934. 

Apparently a shrub, the branches thick; branchlets, petioles, 
pedicels, and blades beneath crisp-puberulent; petioles slender, 2.5 
cm. long; blades nearly oblong or oblong-obovate, minutely cordate 
at the base, emarginate, 7-8 cm. long, 2.5-3 cm. wide, dull, char- 
taceous, the nerves and veins scarcely obvious; pedicels 3 cm. long; 



FLORA OF PERU 1003 

sepals pulverulent, oval, reflexed, 3.5 mm. long; petals irregularly 
oblong-elliptic, 12 mm. long; scales thick, 2 mm. long; stamens 
about 4 cm. long. 

Lambayeque: Between the haciendas Sangana and Batan Grande, 
Prov. Lambayeque, 1,700 meters, Weberbauer 7167, type. 

Capparis Qiiina Macbr. Candollea 5: 357. 1934. 

A shrub, glabrous except for the minutely pulverulent inflores- 
cence; petioles 10-14 mm. long; blades elliptic, rounded at the base, 
slightly emarginate at the apex, to 12 cm. long, 7 cm. wide, mostly 
somewhat smaller, coriaceous, lustrous and strongly reticulate- 
veined on both sides; pedicels 6-7 mm. long; calyx lobate, the 
segments scarcely 2.5 mm. long; petals rounded-obovate, 7 mm. 
long; stamens about 20, long-exserted ; stipe 3 cm. long; fruit nearly 
globose, 3 cm. thick, punctate-tuberculate. 

San Martin: Rio Mayo, Tarapoto, 400-900 meters, in forest, 
Williams 6282, type. Cumbasa, Tarapoto, Williams 5753. "Quina- 
quina." 

Capparis Schunkei Macbr. Field Mus. Bot. 4: 170. 1929. 

Differing from other Peruvian species with similar calyx in its 
lustrous, nearly glabrous foliage (blades mostly 10-15 cm. long, 
4.5-6 cm. wide), and in its branched inflorescence; pedicels 7-10 
mm. long; sepals closed in bud, oblong-lanceolate, 3 mm. long, 
reflexed; petals 5 mm. long. Williams 6692 is rather more pubes- 
cent; it has a fruit 3 cm. long, 2.5 cm. thick, on a stipe 3 cm. long. 
The similar C. detonsa Triana & Planch., Colombian, has somewhat 
larger leaves and flowers twice as large. C. hypoleuca Presl, from 
Guayaquil, has leaves sparsely stellate-pubescent above, densely 
white-tomentose beneath; calyx open in bud, 4-dentate; pedicels 
about 3 mm. long. Named for the naturalist Carlos Schunke, of 
La Merced. 

Junin: Chanchamayo Valley, 1,500 meters, Schunke 418, type; 
514. Hacienda Schunke, La Merced, 1,200 meters, 5701. San 
Martin: Santa Rosa, 200 meters, Williams 4820. Tarapoto, 400- 
900 meters, in forest, Williams 6692. Loreto: Yurimaguas, Williams 
4118; King 2820. Mishuyacu, 100 meters, King 225. 

Capparis sola Macbr. Candollea 5: 359. 1934. C. acutifolia 
Macbr. op. cit. 358, non Sweet. 

A glabrous shrub or small tree; petioles 1.5-3 cm. long; blades 
nearly membranous, reticulate-veined, lustrous, especially above, 



1004 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ovate-lanceolate or ovate-elliptic, broadly cuneate at the base, 
gradually and acutely acuminate at the apex, to 15 cm. long, 6 cm. 
wide, or often smaller; racemes to about 10 cm. long, few-flowered; 
pedicels 10-12 mm. long; sepals rounded-elliptic, scarcely 2 mm. 
long; petals white, rounded-obovate, 5-7 mm. long; stamens 1.5-2 
cm. long; stipe 2.5 cm. long, the ovary oblong-cylindric. Inad- 
vertently described under two names, to which carelessness A. C. 
Smith has called attention. A. C. Smith and Krukoff, Bull. Torrey 
Club 64: 409. 1937, have found C. sola an important ingredient of 
the curare poison of the Javas. 

Loreto: Mishuyacu, near Iquitos, Klug 961, type. San Martin: 
Chazuta, 260 meters, Klug 3966. 

Capparis Sprucei Eichl. in Mart. Fl. Bras. 13, pt. 1: 281. 1865. 

Completely glabrous; leaves coriaceous, ovate-elliptic, rounded 
at each end or emarginate at the apex, 3-7 cm. wide, lustrous above, 
paler and dull beneath, prominently reticulate-veined on both 
surfaces; racemes many-flowered, the flowers about 18 mm. broad, 
on pedicels 3.5 cm. long; calyx lobes suborbicular, erose-ciliate, 
open in bud. Mature fruit unknown. It seems probable that this 
is the same as C. macrocarpa. 

San Martin: Near Tarapoto (Spruce 3986, type). 

Capparis tarapotensis Eichl. in Mart. Fl. Bras. 13, pt. 1: 
284. 1865. 

A densely leafy shrub 2.5 meters high; leaves elliptic, obtuse 
at each end, about 5 cm. long, 2.5-3 cm. wide, subsessile; flowers 
about 1.5 cm. wide; fruit beaked, linear, 2-3 cm. long, the stipe as 
long or longer; pedicels 1 cm. long or shorter. C. heterophylla R. & P., 
from Guayaquil, has a very shortly stipitate ovary. The Brazilian 
specimen is from limestone in Matto Grosso (Malme). Neg. 8475. 

San Martin: Tarapoto, Spruce 4406; Ule 6619, 6698; Williams 
6654. Apurimac: At 1,100 meters, Weberbauer 5872. Brazil. 

5. MORISONIA L. 

Similar to Capparis, but the calyx rupturing irregularly at 
anthesis; stamens 6-20, shorter than the petals; the Peruvian species 
with long, slender, furfuraceous petioles and broad, glabrous, char- 
taceous, finely reticulate-veined leaves. 

Morisonia oblongifolia Britton, Bull. Torrey Club 16: 17. 1889. 

A shrub or small tree as much as 7 meters high, with glabrous, 

densely reticulate-veined, roundish or elliptic-ovate blades to 25 



FLORA OF PERU 1005 

cm. long and 15 cm. wide, or smaller, rounded at the apex or rarely 
acutish, the base slightly cordate; petioles several centimeters long, 
furfuraceous like the branchlets and the few-flowered inflorescence; 
pedicels about 1 cm. long; calyx obconic, 8 mm. long; petals pale- 
puberulent, 8-10 mm. long; fruits subglobose, furfuraceous, 3 cm. 
thick. Neg. 5797. 

San Martin: Region of Tarapoto, 400-900 meters, in forest, 
Williams 6215, 6870, 5715, 6650; Spruce 4529. Juanjui, 400 meters, 
in forest, Klug 3773, 4399; shrub or tree of 3-5 meters; flowers 
yellow-green and white. Loreto: Iquitos, Williams 8161. Rio 
Acre: Mouth of Rio Macauhan, Krukoff 5702, 5508, 5431. Bolivia 
and Amazonian Brazil. 

At least as regards the Peruvian and Brazilian material, the 
specific name is a poor one, since the leaves are in no sense oblong. 
Negative 5797 is of Ule 5799b, from Jurua Miry, Brazil, which is 
indicated by Gilg as a new species, but is still unpublished. 
Paul C. Standley. 

6. STERIPHOMA Spreng. 

Shrubs, similar in appearance to Capparis, but conspicuously 
red-tomentose; calyx calyptriform; stamens always 6. 

Steriphoma peruvianum Spruce ex Eichl. in Mart. Fl. Bras. 
13, pt. 1:267. 1865. 

Branchlets, petioles, and calyx rufous- or orange- tomentose; 
leaf blades lanceolate, acuminate, some of them often peltate at 
the base. A straggling or clambering shrub or small tree, with a 
terminal inflorescence of orange flowers. Neg. 5800. 

San Martin: Tarapoto, Spruce 4128; Ule 6429; Williams 
5732. Morales, 400-900 meters, in forest, Williams 5647. Juan 
Guerra, 720 meters, Williams 6859. Juanjui, 400 meters, in forest, 
Klug 3772. 

7. CRATAEVAL. 

Shrubs or trees with trifoliolate leaves and racemose or corym- 
bose flowers. Petals 4. Stamens long-exserted, 8-50. Fruit large, 
long-stipitate. The fruit and wood have a garlic-like odor. 

Crataeva Benthamii Eichl. in Mart. Fl. Bras. 13, pt. 1: 
265. 1865. 

Leaflets ovate-lanceolate, 4-8 cm. wide or narrower, glabrous; 
stamens 20-50 and 3-4 times as long as the corolla; petals lanceolate 



1006 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

to almost linear (var. leptopetala Eichl.), about 1 cm. long; fruit glo- 
bose, nearly 3 cm. in diameter, brownish, with white dots. C. radia- 
tiflora DC. Prodr. 1: 243. 1824, to which the Peruvian plant may 
be referable, is known from as near as Guayaquil; it has broadly 
ovate-elliptic leaflets and somewhat longer petals. Eichler saw a 
further difference in the white-verruculose markings of the branch- 
lets, a character that seems to be highly variable. The widely dis- 
tributed C. Tapia L. is with difficulty, if at all, distinguishable from 
the two preceding species by its more strictly perfect flowers with 
fewer stamens, these only twice as long as the petals; fruit ovoid. It 
seems questionable if all the names should not apply to one variable 
species. Negs. 8466, 8467. 

Loreto: Sarayaco (Huber). Huallaga, Yurimaguas, 200 meters, 
Williams 4655. Pebas, in forest, Williams 1578, 1845. Caballo- 
cocha, in forest, Williams 2450, 2219, 2270, 2411, 2443. La Victoria, 
Williams 3122, 2854, 2665. Rio Masana, Williams 93, 8161, 90. 
Rio Acre: Rio Macauhdn, Krukoff 5665. Brazil. "Palo de candela," 
"tamara," "insira mashan," "tapia," "nina caspi." 

81. TOVARIACEAE 

Reference: Pax & Hoffmann, Pflanzenfam. ed. 2. 17b: 224-226. 
1936. 

A well marked family, consisting of one genus, related to the Cap- 
paridaceae, but the flower parts usually 8, the ovary incompletely 
(usually) 6-celled, the fruit a green berry, the aspect similar to that 
of certain Phytolaccaceae, to which group Bentham and Hooker 
referred it. 

1. TOVARIA R. & P. 

Slender, sparsely and virgately branched, green-barked shrubs, 
often flowering as annuals. Leaves alternate, trifoliolate, glabrous. 
Stipules minute, triangular. Racemes terminal, elongating, lax. 
Filaments dilated toward the base. Flowers perfect. 

Tovaria pendula R. & P. Syst. Veg. 85. 1798; Fl. Peruv. 3: 73. 
pi. 306. 1802. 

Leaflets lanceolate, acuminate, often 8 cm. long and chiefly 
2-3 cm. wide; racemes drooping, the laxly disposed, pale yellowish 
green or greenish white flowers about 7 mm. broad; petals obtuse, 
exceeding the acute sepals, about 4 mm. long; stamens exserted. 
According to Weberbauer, in Peru at middle elevations on the eastern 
ranges. The plant, both growing and in the herbarium, has a strong 



FLORA OF PERU 1007 

and characteristic odor which has been variously described as that of 
Apium graveolens, Cestrum, and cumarin. In the field it recalled to 
me the odor of preparations used as supplementary foods for stock, 
but the reason was not obvious until Dr. Charles Baehni identified 
it as that of species of Trigonella. Illustrated, Pflanzenfam. op. cit. 225. 
Huanuco: Chinchao and Chacahuassi, Ruiz & Pavdn. Rio Po- 
suso, 2,200 meters, Weberbauer 6784- Junin: Huacapistana, Weber- 
bauer 2326; 252; at 1,800-2,400 meters, Kittip & Smith 24272, 24173; 
shrub 2 meters high or less; perianth pale green; filaments greenish 
white; anthers brown or yellow; fruit green. La Merced, 600 meters, 
5566. Bolivia to Venezuela. 

82. PODOSTEMACEAE 

Reference: Engler, Pflanzenfam. ed. 2. 18a: 3-68. 1930. 

These aquatic plants that usually grow attached to rocks in swift 
streams, often in waterfalls, probably are more numerous in Peru 
than is indicated below, because few collectors find them. Resem- 
bling algae or mosses in general appearance, they are readily recog- 
nized. The small flowers consist of 3-5 free, or more or less united 
sepals, or these may be reduced to scales. The stamens are one to 
several, and then whorled, the anthers usually introrse, 4-celled. 
Ovary 2-3-celled, free, with a thick, central placenta. Capsule usu- 
ally septicidal, many-seeded. 

1. APINAGIA Tul. 

Plants thalloid, or floating and with fibrous roots. Flowers borne 
in a dichotomous or sickle-like inflorescence. Stamens usually 2-4, 
free or united. 

Apinagia peruviana Wedd. ex Engler, Pflanzenfam. ed. 2. 18a: 
38. 1930. Neolacis peruviana Wedd. in DC. Prodr. 17: 64. 1873. 

Leaves oblong-lanceolate, 1-3 cm. long, rather broad, irregularly 
2-3-pinnatifid; flowers axillary; pedicels finally 1.5-2 cm. long, con- 
gested in fruit; capsules 8-nerved, 2.5-3.5 mm. long. 

Puno: Sangaban (Lechler 2298, type). Piura: Frias, 900-1,000 
meters, Weberbauer 6426. 

83. CRASSULACEAE. Orpine Family 

Reference: Berger & Harms in Engl. & Prantl, Pflanzenfam. ed. 2. 
18a: 352-483. 1930. 

A number of the Peruvian species are highly decorative, and not 
at all uncommon on rock ledges at middle elevations. One or more 



1008 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

appear to be known already in cultivation. Besides the following, 
Wawra collected (No. 2564) at Lima the treelike, cultivated Aeonium 
arboreum (L.) Webb. & Berth., of the Mediterranean region, with 
spatulate leaves 7-8 cm. long and 2 cm. wide, the golden yellow flowers 
8-12-parted. 

Leaves simple. 

Leaves small, a few millimeters to rarely 1 cm. long. 

Stamens as many as the sepals, in 1 series; a diminutive annual. 

1. Crassula. 
Stamens twice as many as the sepals, in 2 series; plants becoming 

perennial 2. Villadia. 

Leaves larger, often several centimeters long. Flowers more or 

less tubular 3. Echeveria. 

Leaves pinnate 4. Bryophyllum. 

1. GRASSULA L. 

Tillaea L. 

Small, branched, somewhat succulent herbs with minute, solitary, 
axillary, pedicellate flowers. Commonly many plants grow laxly 
matted together, in loose or sandy soils and often under shrubs, 
especially in the loma regions (Weberbauer, 145). 

Flowers few and segregated on long, slender pedicels much exceed- 
ing the subtending leaves; carpels 10-15-seeded . .C. Macbridei. 

Flowers many and crowded, appearing verticillate, sessile or on very 
short pedicels much shorter than the subtending leaves; carpels 
2-seeded C. connata. 

Crassula connata (R. & P.) Berger in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 18a: 389. 1930. Tillaea connata R. & P. Fl. Peruv. 1: 70. 
pi. 106. 1798. T. rubescens HBK. Nov. Gen. & Sp. 6: 43. 1823. 

More or less rubescent; leaves connate, lanceolate, mucronate, 
2-4 mm. long; pedicels 2-4 mm. long; sepals 4, longer than the 4 
petals. The native name refers to the similarity in odor to musk 
(Ruiz & Pavon). 

Lima: Chicla (Batt). Lurin, 60 meters, 5970. Matucana, 2,400 
meters, 182. Chancai, Ruiz & Pavon. Puno: Limestone ledges, 
3,900 meters, Chuquibambilla, Pennell 13043. Arequipa: Mejia 
and Cachendo (Gunther & Buchtien 285, 285a). Junin: Yauli, 919. 
Cuzco: Valle del Apurimac, Herrera. Extending to Lower Cali- 
fornia. "Almizclillo." 



FLORA OF PERU 1009 

Crassula Macbridei Steyermark, sp. nov. 

Herba pusilla furcata omnino glabra, 2.5-6 cm. alta; caule tenui; 
foliis oppositis, anguste oblanceolatis, acutis, basi connata attenua- 
tis, 4-8 mm. longis, 1.5-2 mm. latis; pedicellis axillaribus filiformibus 
solitariis 8-18 mm. longis foliis multo longioribus; sepalis 4 oblongo- 
lanceolatis 1.8-2 mm. longis, apice aristato recurvato-patentibus, 
petalis ovatis subacutis longioribus; staminibus 2, filamentis fili- 
formibus, antheris ovalibus; carpellis 3 ovoideis sepalis paullo 
brevioribus, seminibus in quoque carpello 10-15 late oblongis cas- 
taneis 0.2 mm. longis, fortiter papillatis. 

Lima: Sandy lomas along the sea, Lurin, 60 meters, Sept. 23, 
1923, Macbride 5971 (type in Herb. Field Mus.). 

This new species is placed in Crassula following Berger's treat- 
ment of the family. In general appearance it resembles the South 
African Crassula expansa Ait., from which it differs in having the 
petals shorter than the sepals, in the diminutive stems, and in the 
4-merous flowers. It is related to the South American Crassula pedun- 
cularis (Smith) Schonl. (Tillaea peduncularis Smith) of Uruguay and 
Argentina, from which it differs in its larger flowers, oblanceolate 
instead of linear-lanceolate leaves, forking branches, and more numer- 
ously seeded carpels. 

With 4-merous sepals and petals it seems properly to belong to 
the section Vaillantii Schonl. as treated in Berger's evaluation of 
the family, although in its 10-15-seeded carpels it diverges from 
this section. 

2. VILLADIA Rose 

Altamiranoa Rose. 

Reference: Berger in Engl. & Prantl, Pflanzenfam. ed. 2. 18a: 
468-470. 1930. 

Small-leaved perennials with small flowers, and therefore with 
the aspect and habit of Sedum, but resembling Echeveria in having 
the flower parts constantly 5 and the petals to a greater or less 
degree united into a tube. In the Peruvian species the inflorescence 
is presumably always one-sided, on which basis the generic name 
Altamiranoa has been proposed, but the inflorescence in Echeveria 
is variable. 

Petals shorter than or subequaling the sepals, greenish .V. Dyvrandae. 
Petals distinctly longer than the sepals. 
Flowers dark red. 



1010 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petals subobtuse V. andina. 

Petals subacute, mucronate-carinate V, Berillonana. 

Flowers white, yellowish, or pale reddish. 
Leaves (5) 6-9 mm. long; flowers white. 
Petals 5-10 mm. long; style 1-3 mm. long. 
Style 1 mm. long; inflorescence diffuse; petals to 10 mm. 

long V . Weberbaueri. 

Style 2-3 mm. long; inflorescence strict; petals 5-8 mm. 

long V. Dielsii. 

Petals 4-5 mm. long; style 0.5 mm. long V. virgata. 

Leaves 2-5 mm. long. 

Flowers 8-12 in a short raceme, yellowish or reddish. . 

V. incarum. 
Flowers in a cyme (or sometimes racemose). 

Leaves 2-3 mm. long, strongly imbricated .... V. imbricata. 
Leaves 3.5-5 mm. long, not or slightly imbricated. 
Stems creeping or ascending; leaves 2-3 mm. wide. 

V. Berillonana. 
Stems erect; leaves 3-4.5 mm. wide V. Grandyi. 

Villadia andina (Ball) Baehni & Macbr. Candollea 7: 285. 
1937. Sedum andinum Ball, Journ. Linn. Soc. 22: 38. 1885. 

Cespitose, with many slender branches from the base, these 5-8 
cm. long, the flowering stems ascending; leaves minute, hemispheric; 
cyme 4-flowered, the terminal flower sessile, the lateral ones short- 
pedicellate; petals coalescent at the base, dark red, oblong-lanceo- 
late, subobtuse, twice as long as the ovate-elliptic sepals; stamens 10. 

Lima: Chicla (Ball, type). 

Villadia Berillonana (Hamet) Baehni & Macbr. Candollea 7: 
285. 1937. Sedum Berillonanum Hamet, Bot. Jahrb. 50: Beibl. 112: 
9. 1913. Altamiranoa Berillonana Berger in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 18a: 470. 1930. 

Flowering stems creeping and branching at the base, finally 
erect and simple, to 8 cm. long; leaves ovate, entire, sessile, 3.5-5 
mm. long, less than 3 mm. wide; inflorescence biparted but not 
branched, few-flowered, 8-12 cm. long; flowers subsessile; sepals 
entire, acutish, elongate-ovate, 3.5-5 mm. long; corolla a little 
longer, the entire, suboblong segments acutish-mucronate, united 
to about the middle; scales broader than long, 1 mm. broad; stamens 
to 6 mm. long; styles 2 mm. long. 



FLORA OF PERU 1011 

Ayacucho: Shrubby rock outcrop, 3,000 meters, Weberbauer 5501, 
type. 

Villadia Dielsii Baehni & Macbr. Candollea 7: 285. 1937. 
Cotyledon stricta Diels, Bot. Jahrb. 37: 410. 1906, non V. stricta 
Rose. Altamiranoa stricta Berger in Engl. & Prantl, Pflanzenfam. 
ed. 2. 18a: 470. 1930. 

Principal stems erect, much branched, the very strict branches 
15-20 cm. high; leaves dense but not rosulate, erect, triangular- 
ovate, semiclasping, 6-8 mm. long, 1-1.5 mm. wide; branches of the 
inflorescence scorpioid; flowers subsessile; sepals oblong-ovate, 3.5 
mm. long; petals subundulate, 5-8 mm. long, recurved at the apex, 
white; stamens 5 mm. long; styles 2-3 mm. long. 

Ancash : Rocky, grassy fields, 2,200 meters, near Huaraz (Weber- 
bauer 3000, type). Lima: Open hillside, Rio Blanco, Kittip & Smith 
21062. 

Villadia Dyvrandae (Hamet) Baehni & Macbr. Candollea 7: 
286. 1937. Sedum Dyvrandae Hamet, Bot. Jahrb. 50: Beibl. 112: 10. 
1913. Altamiranoa Dyvrandae Berger in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 18a: 470. 1930. 

Stems ascending, often branched above the middle, to about 15 
cm. high; leaves elongate-ovate, subobtuse, 2.5-6 mm. long, less 
than 3 mm. wide; inflorescence to 2.5 cm. long, spikelike, lax, few- 
flowered; sepals ovate-oblong, entire, subobtuse, to 4.5 mm. long; 
corolla less than 4 mm. long, the very broadly ovate segments about 
one-third united; stamens 1.5 mm. long; scales longer than broad, 
1.5 mm. long; styles 0.6 mm. long. Flowers greenish. Neg. 18247. 

Lima: In rocks, Matucana, 2,400 meters, Weberbauer 5278, type; 
Macbride 445. 

Villadia Grandyi (Hamet) Baehni & Macbr. Candollea 7: 
286. 1937. Sedum Grandyi Hamet, Bot. Jahrb. 50: Beibl. 112: 11. 
1913. Altamiranoa Grandyi Berger in Engl. & Prantl, Pflanzenfam. 
ed. 2. 18a: 470. 1930. 

Similar to V. Berillonana, but erect; inflorescence laxer, 1.5 cm. 
long, biparted, not branched; leaves ovate-suborbicular; pedicels 
0.5 mm. long; sepals unequal, 4 of them subovate, subobtuse, 3-3.5 
mm. long, 1 of them long-ovate, 3 mm. long; petals broadly ovate, 
united one-fourth their length; stamens to 4 mm. long; styles 1.5 
mm. long; scales broader than long, 1 mm. wide. Neg. 34154. 

Amazonas: Chachapoyas, Mathews, type. 



1012 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Villadia imbricata (Diels) Baehni & Macbr. Candollea 7: 
286. 1937. Cotyledon imbricata Diels, Bot. Jahrb. 37: 411. 1906. 
Altamiranoa imbricata Berger in Engl. & Prantl, Pflanzenfam. ed. 2. 
18a: 470. 1930. 

Much branched, subcespitose, ascending, the branches 5-10 cm. 
long; leaves broadly sessile, triangular or nearly reniform, 2-3 mm. 
long, 3-4 mm. wide; branches of the inflorescence scorpioid, divari- 
cate; sepals 3 mm. long, 1 mm. wide; petals 4 mm. long, 1.5-2 mm. 
wide, the acute apex recurved, greenish white; stamens 3.5 mm. 
long; carpels 2.5 mm. long. 

Cajamarca: La Tahona, 2,600 meters (Weberbauer 4053, type). 
Huanuco: Llata, 2,100 meters, 2262 (det. Rose). Arequipa: Alca, 
Weberbauer 6877. 

Villadia incarum (Ball) Baehni & Macbr. Candollea 7: 286. 
1937. Cotyledon incarum Ball, Journ. Linn. Soc. 22: 37. 1885. 
Altamiranoa incarum Berger in Engl. & Prantl, Pflanzenfam. ed. 2. 
18a: 469. 1930. 

Plants erect, branched below, to 20 cm. high, the lateral branches 
short; leaves ovate-cylindric; flowers ochroleucous or reddish, erect, 
8-12 in a short raceme, the uppermost sessile; sepals ovate-lanceolate; 
petals linear-lanceolate, acute, half longer than the sepals, one-third 
to one-half united. The spelling "incanum" (Ball) was corrected 
in the errata. 

Lima: Chicla (Ball, type). Rio Blanco, Killip & Smith 21556(1). 

Villadia virgata (Diels) Baehni & Macbr. Candollea 7: 286. 
1937. Cotyledon virgata Diels, Bot. Jahrb. 37: 410. 1906. Alta- 
miranoa virgata Berger in Engl. & Prantl, Pflanzenfam. ed. 2. 18a: 
470. 1930. 

Principal stems decumbent or erect, much branched, the strict 
branches curved-erect, to 25 cm. high; leaves nearly appressed, or 
erect, ovate, 5-8 mm. long, 2-3 mm. wide; inflorescence spikelike; 
sepals narrowly triangular, 3 mm. long; petals ovate, white, 4-5 
mm. long; stamens about 4 mm. long; scales conspicuous, oblong- 
spatulate, truncate, 1.5-2 mm. long; styles 0.5 mm. long. 

Cuzco: Herrera 3155. Ancash: Near Tallenga, 3,300 meters 
(Weberbauer 2853, type). Huanuco: Llata, 2,100 meters, 2265 (det. 
Rose). 

Villadia Weberbaueri (Diels) Baehni & Macbr. Candollea 7: 
286. 1937. Cotyledon Weberbaueri Diels, Bot. Jahrb. 37: 411. 1906. 



FLORA OF PERU 1013 

Altamiranoa Weberbaueri Berger in Engl. & Prantl, Pflanzenfam. ed. 
2. 18a: 470. 1930. 

Principal stems decumbent, the ascending branches 10-15 cm. 
long; leaves very broadly ovate, subacute; inflorescence fastigiate- 
cymose, the branchlets many-flowered; sepals 4 mm. long, 1.5 mm. 
wide; petals white, the elongate apex finally recurved, 2 mm. wide; 
stamens 6 mm. long; styles 1 mm. long. 

Amazonas: Above Balsas, 2,300 meters (Weberbauer 4282, type). 
Rio Maranon, below Chavanillo, 2,250 meters, 1984 (det. Rose). 

3. ECHEVERIA DC. 

Leaves spiraled in loose or dense rosettes, large in comparison 
with those of other groups. Flowers 5-parted, the corolla more or 
less tubular. A convenient segregate from the Old World Cotyledon. 
Cf. Poellnitz, Repert. Sp. Nov. 39: 193-270. 1936, to whose work I 
am indebted for preparation of the following account. 

Flowers conspicuously pedicellate, the racemes soon open, the 

plants caulescent. 
Plants acaulescent or nearly so. 

Leaves spatulate, smooth, glabrous E. peruviana. 

Leaves lanceolate, subasperous, papillose E. Backebergii. 

Plants caulescent; leaves lanceolate E. chiclensis. 

Flowers sessile or subsessile, crowded in an thesis; plants acaulescent. 
Bracts broadly ovate or roundish; flowers flesh-colored. 

E. eurychlamys. 
Bracts oblong-lanceolate; flowers scarlet E. excelsa. 

Echeveria Backebergii Poellnitz, Repert. Sp. Nov. 38: 185. 
1935. 

Leaves densely rosulate, often minutely white-mucronulate, to 
6 cm. long; stems to 30 cm. tall, 6 mm. thick at the base; cauline 
leaves many; flowers yellowish, reddish toward the base, on pedicels 
as much as 2.5 cm. long; sepals unequal, the longer ones 8 mm. long; 
corolla about 14 mm. long, the tube 3-4 mm. long, the lobes dis- 
tinctly extrorse-curved at the apex. In cultivation by the collector 
at Kiel, and by the author. 

Lima: Matucana, 2,600 meters (Kurt Backeberg S2). Matucana, 
2,400 meters, 279. 

Echeveria chiclensis (Ball) Berger in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 18a: 473. 1930. Cotyledon chiclensis Ball, Journ. Linn. 
Soc. 22: 38. 1885. 



1014 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Stems erect, 1 to several decimeters tall, very leafy, the cauline 
leaves 2.5-5 cm. long; basal leaves linear-lanceolate, glabrous, acute, 
10-15 (-20) cm. long; lowest pedicels 12-20 mm. long, about equaled 
by the bracts, erect or finally nodding; calyx segments ovate; 
petals 13-20 mm. long, bright red, linear-oblong, acuminate, about 
one-fourth united, the tips recurving; carpels acute, subrecurved- 
divaricate. 

Lima: Chicla, 4,000-4,300 meters (Ball, type). Huanuco: Hua- 
nuco, Ruiz & Pavdn. La Oroya, 3,700-4,300 meters (Kalenborn 104). 
Junin: Tarma, 2,900 meters, 1061. Huariaca, 2,850 meters, 3254. 

Echeveria eurychlamys (Diels) Berger in Engl. & Prantl, 
Pflanzenfam. ed. 2. 18a: 473. 1930. Cotyledon eurychlamys Diels, 
Bot. Jahrb. 37: 411. 1906. 

Outer leaves of the dense rosettes short and broad, the inner 
ones broadly ovate-elliptic, acute, 3-3.5 cm. long, to 2 cm. wide; 
scapes ascending, with the finally elongate inflorescence 25-30 cm. 
long, the leaves appressed; bractlets rotund-ovate or suborbicular, 
13-15 mm. long, 12-13 mm. wide; pedicels 2-3 (-5) mm. long; 
sepals ovate-oblong, 6 mm. long; petals lanceolate, flesh-colored, 15- 
17 mm. long, recurved at the tips; stamens 10 mm. long; styles 4-5 
mm. long. Neg. 18249. 

Cajamarca: La Tahona, 3,100 meters, Weberbauer 4056, type. 

Echeveria excelsa (Diels) Berger in Engl. & Prantl, Pflanzen- 
fam. ed. 2. 18a: 473. 1930. Cotyledon excelsa Diels, Bot. Jahrb. 37: 
412. 1906. 

Scapes 3-5 cm. thick; leaves subrosulate, oblong-obovate, nar- 
rowed to the base, acute, 12-15 cm. long, 3.5-4 cm. wide; cauline 
leaves few, to 4 cm. long; scapes to over a meter tall; spikes narrowly 
cylindric, the remote flowers short-pedicellate; bracts oblong-lanceo- 
late, 15 mm. long, 3 mm. wide; sepals ovate, 7 mm. long, 3 mm. 
wide; petals lanceolate, scarlet, with paler, spreading tips, acute, 16- 
20 mm. long, 5-7 mm. wide. Neg. 18250. 

Ancash: Between Samanco and Huaraz, 3,300 meters, Weberbauer 
3149, type. 

Echeveria peruviana Meyen, Reise 1: 448. 1834. Cotyledon 
peruviana Baker in Saunders, Ref . Bot. 1 : under pi. 58. 1869. 

Plants glabrous, acaulescent or nearly so; leaves obovate-spatu- 
late, 5-7.5 cm. long, those of the scape shorter, few; lower flowers 
solitary; pedicels 3-12 mm. long; calyx lobes spreading or widely 



/r 



FLORA OF PERU 1015 

divaricate; corolla red, 12-15 mm. long, the tube 1.5-4 mm. long. 
The 5-angled flowers are lighter toward the spreading tips. Neg. 
18248. 

Tacna: Meyen. Above Supe (Weberbauer, 169). Junin: Tarma 
(Weberbauer, 176). Argentina; Chile. 

4. BRYOPHYLLUM Salisb. 

This introduced and naturalized plant is readily recognizable as 
a sprawling, succulent herb with pinnate leaves and ample, loose 
inflorescence of nodding, bronze-green flowers. 

Bryophyllum pinnatum (Lam.) Kurz, Journ. Asiat. Soc. Bengal 
40, pt. 2: 52. 1871. Cotyledon pinnata Lam. Encycl. 2: 141. 1786. 
B. calycinum Salisb. Bot. Mag. pi. 1409. 1811. 

Leaflets 3-5, oval, crenate, 7-13 cm. long; calyx 2-3 cm. long, 
inflated, the deltoid teeth 7-10 mm. long; corolla tube to 3.5 cm. 
long, the lobes 12 mm. long, triangular. 

Huanuco: Ambo, in rocky stream bed, 2,450 meters, 2420. 
Loreto: Lower Rio Huallaga, 155-210 meters, Williams 5154- Pebas, 
Williams 1837, 2382. Near Iquitos, Williams 1396.- San Martin: 
Moyobamba, 1,100 meters, King 3752. Native of the Old World, 
often naturalized in tropical America. 

84. SAXIFRAGACEAE. Saxifrage Family 

A small rock saxifrage, several hydrangeas, and a number of 
species of wild currants, some with beautiful, elongate racemes of 
roseate flowers, are elements of this family or group of families 
as it occurs in Peru, that seem pleasantly familiar to a botanist com- 
ing from the north. Novelties to him will be the Escallonias, and the 
curious Phyllonomas with their short inflorescences of small flowers 
arising directly from the midnerve of the leaf. Such a diversity in 
character in one family has naturally suggested to a number of botan- 
ists notably Hutchinson, Families of Flowering Plants 1 : 120. 1926 
their disposition as separate families, but here Engler's at least con- 
venient arrangement in one family is followed, and acknowledgment 
is made of the use of his work: Pflanzenfam. ed. 2. 18a: 74-226. 1928 
(1930). 

Alpine herbs 1. Saxifraga. 

Trees, shrubs, or vines. 

Inflorescence not epiphyllous. 
Flowers racemose; leaves alternate. 



1016 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves not elongate, palmate-nerved; ovary 1-celled; fruit a 

berry 2. Ribes. 

Leaves elongate; ovary 2-3-celled; fruit capsular. 

3. Escallonia. 

Flowers cymose-umbellate; leaves often opposite . 4. Hydrangea. 
Inflorescence arising from the leaf costa 5. Phyllonoma. 

1. SAXIFRAGA L. Saxifrage 

Reference: Engler & Irmscher, Pflanzenreich IV. 117: 378-382. 
1916. 

This large genus, of wide distribution, is represented in the high 
Andes of Peru by only one polymorphic species. It is common on 
rocks between, according to Weberbauer, 2,800 and 4,500 meters, 
but is mentioned by Ruiz and Pavon in their journal only once, when 
they found it above Tarma and noted the common names "puchup- 
pus" and "siempreviva." 

Saxifraga magellanica Poir. in Lam. Encycl. 6: 686. 1804; 378. 
S. cordillerarum Presl, Rel. Haenk. 2: 55. 1831. 

Compactly cespitose, more or less densely glandular-pilose, the 
subligneous or herbaceous stems 1-5 cm. long, often columnar in 
form because clothed by the imbricate, dead leaves; scapes 1-10 cm. 
high, 1-several-flowered; leaves 5-8 cm. long, 3-5-lobed or bifid, the 
cauline ones sessile, cuneate, mostly 3-lobed; sepals ovate, obtuse, 
shorter than the subcampanulate receptacle, the 3-nerved petals 
2-3 times as long; stamens scarcely equaling the sepals; capsule 
ovoid, truncate. The typical form, with petals not more than 3 mm. 
long, is confined to the antarctic region. This the monographers have 
designated subsp. Poiretii; the Andean form, with petals as much as 
9 mm. long, subsp. peruviana, dividing this into three varieties and 
many forms. Since these smaller divisions are of doubtful taxonomic 
significance, there being no proof that they are more than ecological 
states, they may most practically be treated as constituting one vari- 
able variety, this weak, but separated geographically. Illustrated 
(the many leaf forms), Pflanzenreich IV. 117: 380. 

Saxifraga magellanica Poir. var. peruviana (Sternb.) Macbr., 
comb. nov. S. peruviana Sternb. Rev. Saxifr. 55. pi. 22. 1810; Suppl. 
2: 74. pi. 20. 1831. S. cordillerarum Presl, vars. Haenkei, laxa, and 
Bonplandii Presl, Rel. Haenk. 2: 55. 1831. For other synonymy and 
named forms see the monograph. 



FLORA OF PERU 1017 

Petals usually 5-9 mm. long. Stature and form of the plant 
highly variable, also the leaves, these especially in depth of lobing, 
variations probably associated with local conditions and seasons. 
5. Boussingaultii of Chimborazo (Ecuador) has ovate petals shorter 
than the gland-toothed sepals (but see Weberbauer 5159, fide Engl. 
& Irmsch.). 

Ancash: Above Samanco, 3,700 meters (Weberbauer, 171). Rio 
Chiquian, 3,000 meters (Weberbauer, 178). Huaraz, 4,600 meters, 
Weberbauer 2980; 225. Hualgayoc, 3,700 meters (Weberbauer, 272) . 
Lima: Chicla, 3,720 meters, Weberbauer 240; (Ball). Cheuchin, Ruiz 
& Pavon. Matucana, Sawada 482. Tambo de Viso, 2,650 meters, 
Weberbauer 128. Above Lima, 4,800 meters, Weberbauer 5159. Rio 
Blanco, 3,600 meters, rock slide, 717; petals white, filaments red. 
Matucana, 2,400 meters, in mats on rock ledges, 444- Rio Blanco, 
3,000-3,500 meters, open hillside, Kittip & Smith 21583. Junin: 
Above Tarma, Ruiz & Pavon; Weberbauer; Isern. Yauli, 4,500 meters 
(Weberbauer, 223). Mt. La Juntai, 4,700 meters, Kittip & Smith 
22106. La Oroya, Isern 2347. Huaron, 4,200 meters, rocky slopes, 
1123. La Quinua, limestone canyon walls, 3088. Huanuco: Chas- 
qui, 3,150 meters, shaded canyon ledges, 1764> Quina, Sawada P63. 
Tambo de Vaca, 3,900 meters, exposed, rocky knolls, 4394. Cuzco : 
Valle de Paucartambo, 3,900 meters, Herrera 1466. Puno: Sandia, 
3,000 meters (Weberbauer, 183) ; Soukup 535. San Antonio, Lechler 
1797. Department unknown: Tongos to Pampas, 4,300 meters, 
Raimondi. Without locality (Haenke). Bolivia to Colombia. "Hua- 
manripa," "puchuppus." 

2. RIBES L. 

Reference: E. de Janczewski, Me"m. Soc. Phys. Hist. Nat. Geneve 
35:199-517.1907. 

Various species of wild currants grow in Peru at 2,800-4,500 
meters, generally along brooks at lower altitudes, in rock crevices 
at higher elevations (Weberbauer). There they are often gnarled 
shrubs half concealed by the protecting rocks, half prostrate in slides, 
or deformed and stunted, suggesting a precarious existence. When 
the species are better known, the number recognized will probably 
be fewer. 

Racemes, at least the staminate, mostly (3-) 4 to many centimeters 

long, often pendent. 

Flowers, at least the staminate, campanulate-turbinate (cf. R. 
albifolium~) ; petals more or less conchiform, except in R. elegans. 



1018 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves distinctly paler beneath, medium in size; flowers and 

rachis eglandular-pilose, or with only a few stipitate glands. 

Petals obovate, about as long as the stamens . . R. bolivianum. 

Petals roundish, about equaling the filaments R. elegans. 

Leaves concolorous or only slightly paler beneath; flowers 

usually more or less glandular. 
Leaves ample, 10 cm. long and broad; flowers densely pilose 

(only the pistillate known) R. macrostachyum. 

Leaves medium-sized; flowers sparsely pubescent. 
Stamens and petals equally inserted; northern Peru. 

R. viscosum, R. peruvianum. 
Stamens and petals unequally inserted; southern Peru. 

R. incarnatum. 
Flowers tubular, often 5-9 mm. long; petals rounded or elongate, 

except in R. albifolium. 

Petals conchiform; leaves more or less pale-pubescent beneath. 

R. albifolium. 

Petals not conchiform; leaves usually concolorous (cf. R. 
macrobotrys). 

Branches almost shaggy-red-hirsute R. hirticaule. 

Branches less pubescent. 
Leaves rarely 2 cm. wide; stipitate glands dense. 

R. Weberbaueri. 
Leaves much larger. 

Leaves deeply cordate at the base; racemes lax, the pedi- 
cels soon longer than the bracts R. macrobotrys. 

Leaves more or less cordate; pedicels shorter than the 

bracts. 
Stamens and petals subequal, unequally inserted. 

R. incarnatum. 
Stamens and petals unequal, equally inserted. 

R. elegans. 
Racemes few-flowered, often shorter than 2.5 cm.; flowers cam- 

panulate. 
Leaves broadly elliptic to rotund. 

Leaves more or less lobed; stamens and petals equal. 

Branchlets glandular R. brachybotrys. 

Branchlets puberulent, soon glabrous R. praecox. 



FLORA OF PERU 1019 

Leaves oval, not lobed; stamens and petals unequal. 

R. ovalifolium. 
Leaves oblong or cuneate. 

Leaves cuneate R. cuneifolium. 

Leaves oblong R. incertum. 

Ribes albifolium R. & P. Fl. Peruv. 3: 12. pi. 232. 1802; 418. 
Rebis discolor Spach, Ann. Sci. Nat. II. 4: 28. 1835. 

Apparently very similar to R. macrobotrys; leaves smaller, less 
lobed, more regularly and sharply serrate, and more abundantly 
pilose-strigillose beneath; racemes 4-7 cm. long in flower; pedicels 
3-4 mm. long; flowers 5-6 mm. long, scarcely glandular; petals 
roundish but clawed, about as long as the filaments of the fertile 
anthers. R. hirtum HBK., 419, found as near as Ecuador, has more 
tubular flowers and the petals are nearly as long as the stamens. 
Neg. 29708. 

Huanuco: Tambo Nuevo, near Mufia, and at Marainiyoc, near 
Vitoc, Ruiz & Pavdn. 

Ribes bolivianum Jancz. Bull. Acad. Cracov. 759. 1905; 409. 

Branchlets subglabrous except for short-stipitate glands; leaves 
entire or obscurely 3-lobed, to 6 cm. long, 4 cm. wide, commonly 
deeply cordate at the base, white-pubescent beneath; petioles 3 cm. 
long, tomentose and subsessile-glandular; staminate racemes to 10 
cm. long, tomentose, the pedicels scarcely 1 mm. long, the bractlets 
2-3 times as long; flowers minute, shallow-campanulate; petals 
oblong-obovate, little exceeded by the equally inserted stamens, the 
anthers nectar-bearing; pistillate flowers similar, but the smaller 
petals more obovate; fruit to 9 mm. long, distinctly pedicellate. A 
vigorous shrub to 2 meters high. The Gay specimen has leaves 
softly white-pilose beneath. R. andicola Jancz., 411, of the northern 
Andes, scarcely differs unless in the often glandular ovary and the 
smaller leaves with more than 3 lobes; the Peruvian plant is perhaps 
better treated as a variety. Neg. 27342. 

Puno: Cuyoengo, Prov. Sandia, 3,100 meters, Weberbauer. 
Nahuelbuta, Gay. Cuzco: Paso de Tres Cruces, 3,700 meters, 
Pennell 13910. Bolivia. 

Ribes brachybotrys (Wedd.) Jancz. Bull. Acad. Cracov. 759. 
1905; 404. R. viscosum R. & P. var. brachybotrys Wedd. Chlor. 
And. 2: 216. 1857. R. sucheziense Jancz. op. cit. 8. 1906(?). 

Branchlets reddish gray, sparsely roughened with short-stipitate 
glands; leaves crowded, paler beneath, rotund, 1.5-3 cm. wide, 3- 



1020 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lobed, finely serrate, bullate-rugulose and glabrous above or with a 
few evanescent, scattered, stipitate glands, beneath minutely pilose 
and slightly glandular, the nerves and veins prominent; racemes 
about 1 cm. long, little glandular but densely pilose; flowers sub- 
sessile, few, small; stamens and petals subequal; style merely bifid. 
R. Pentlandi Britton, 403, Bolivian, has leaves broadly flabellate, 
obscurely if at all trilobed. The flowers, at least sometimes, are 
perfect, on which basis Janczewski proposed R. sucheziense, but 
apparently the type of R. brachybotrys may have fertile stamens, 
persisting on the young fruits. The type of the former is Weberbauer 
1006, in rocks at 4,500 meters, at Suchez, Bolivia, near Puno, Peru. 
Negs. 4161 (R. sucheziense), 34648. 

Puno: Peninsula Copacabana, Lake Titicaca, 3,900 meters, 
Weddell, type. Near Puno, very common, 4,000 meters, Soukup 65. 
Orurillo, Soukup 560. Have, Soukup 561. Cuzco: La Raya, 4,400 
meters, dry cliff, Pennell 13518. Bolivia. 

Ribes cuneifolium R. & P. Fl. Peruv. 3: 13. pi. 233. 1802; 425. 

A small, densely branched shrub, more or less densely or usually 
sparsely stipitate- or subsessile-glandular; leaves glabrous, lustrous, 
broadly cuneate or deltoid, obscurely 3-lobed, crenate, rarely 12 
mm. long, sometimes 10 mm. wide at the subtruncate apex; flowers 
few, minute, reddish; petals subcuneate, slightly exceeding the sub- 
sessile stamens; fruit reddish, glabrous, about 5 mm. thick, sessile; 
bractlets ciliate. Neg. 27343. 

Junin: La Juntai, 4,700 meters, Killip & Smith 22070. La Oroya, 
Isern 2242. Diezmo, Ruiz & Pavdn, type. Rio Blanco, 4,500 meters, 
among rocks on subalpine slopes, 2982. Apurimac; Above Sanaica, 
3,800 meters, Weberbauer 7176. 

Ribes elegans Jancz. Bull. Acad. Cracov. 761. 1905; 420. 

Rather similar to R. albifolium; leaves about 4 cm. long, 3.5 cm. 
wide, 3-lobed, subcordate at the base, pubescent beneath on the 
nerves, the densely pubescent petioles 1.5 cm. long; staminate ra- 
cemes pendent, 6-20 cm. long, sparsely stipitate-glandular; pedicels 
3-6 mm. long, exceeding the linear bractlets; flowers tubular, scarlet, 
yellow within, the sepals twice as long as the receptacle; petals nearly 
rotund, scarcely longer than the filaments and inserted at the same 
height; anthers without nectaries. The leaves at flowering time 
are much smaller; the cinereous-pubescent, pendent racemes are 
essentially eglandular. Neg. 4151. 



FLORA OF PERU 1021 

Huanuco: Southwest of Monzon, 3,500 meters, Weberbauer 
3306, type. 

Ribes hirticaule Macbr. Field Mus. Bot. 8: 115. 1930. 

Stems long-rusty-hirsute, scarcely glandular, with minute, white 
hairs intermixed; stipules conspicuous, long-ciliate, about 1 cm. long; 
petioles to 4 cm. long; blades cordate-ovate, 3-lobed and irregularly 
serrate, mostly about 10 cm. long and slightly wider, glabrate above 
or sparsely pubescent with minute, white hairs and a few longer, 
rusty ones, paler beneath and cinereous-pubescent, especially on the 
nerves, the lobes acute or subacuminate; racemes 10-15 cm. long, 
conspicuously bracted; pedicels 3-5 mm. long; bractlets about 3 mm. 
long; flowers fiery red, like the pedicels cinereous-pubescent, 5-6 mm. 
long, tubular-campanulate; sepals acute; petals oblong, 1 mm. long, 
in the pistillate flower, inserted with and equaling the stamens; ovary 
densely glandular-hirsute. Perhaps only an extraordinarily reddish- 
hirsute variety of R. macrobotrys, but it lacks the stipitate glands, 
and the petals are larger. 

Ayacucho: Evergreen shrub-wood, Putis, Choimacota Valley, 
3,400 meters, Weberbauer 7529, type. 

Ribes incarnatum Wedd. Chlor. And. 2: 215. 1857; 432. 

Nearly glabrous except for the sparsely pubescent racemes; leaves 
crowded, ovate or oblong, 3-lobed or subentire, unequally and acutely 
serrate, 3-5 cm. long, the nerves beneath and petioles pubescent; 
racemes 5-12 cm. long, pendent, pubescent and sparsely glandular- 
pilose; bracts lanceolate, 5 mm. long, glandular-ciliate like the (half 
as long) pedicels; flowers campanulate, short-pedicellate, carmine; 
stamens inserted below the narrowly obovate, minute petals; recep- 
tacle 5 mm. long, puberulent; fruit glandular-pilose. 

Cuzco : Cordillera de Santa Ana, Weddell, type. 

Ribes incertum Macbr. Field Mus. Bot. 8: 116. 1930. 

Nearly glabrous throughout, the branchlets short, numerous; 
leaves terminally congested, broadly oblong, apparently sessile, 
scarcely or not at all narrowed at the apex, shortly and obtusely 
denticulate, obscurely 3-nerved beneath; racemes greatly reduced, 
the 1-3 greenish yellow, campanulate, rotate, short-pedicellate flowers 
not at all stipitate-glandular, minutely pulverulent; sepals scarcely 
acutish; petals oblong or oblong-spatulate, almost 1 mm. long, un- 
equally (always?) inserted, not exceeded by the conspicuous anthers. 



1022 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ancash: In slide rock, 4,200 meters, Pomopampa, east of Huaraz, 
2493, type. Cordillera de Pelagatos, 4,600 meters, Weberbauer 7226, 

Ribes macrobotrys R. & P. Fl. Peruv. 3: 12. pi. 232. 1802; 437. 
Rebis macrobotrys Spach, Ann. Sci. Nat. II. 4: 27. 1835. 

Branchlets flexuous, lustrous, glabrous; petioles 3-4.5 cm. long, 
setose-glandular, at least at the dilated base; blades deeply cordate, 
broadly ovate, 3-lobed, the middle lobe conspicuously longer than 
the spreading lateral ones, all incised or coarsely toothed, the obtuse 
teeth apiculate-calloused-serrate, paler beneath, slightly pilose only 
on the nerves, glabrous and lustrous above, to 10 cm. long, 7-8 cm. 
wide; racemes stipitate-glandular and pilose, to 20 cm. long; pedicels 
4 mm. long, typically exceeding the nearly linear bracts, soon much 
longer; ovary densely glandular; calyx sparsely glandular and pilose; 
petals oblong, only equaling the filaments of the fertile anthers. 
Flowers 8-9 mm. long. Negs. 4157, 29710, 29711. 

Huanuco: Tambo de Vaca, 3,900 meters, among bushes, 4418; 
slender shrub of 1-3 meters; flowers red outside, yellow within. 
Huasa-huasi, Dombey; Ruiz & Pawn, type. Junin: Tarma, Isern 
2349. Camino de Vitoc a los altos de Palca, Isern 2156 (more 
glandular). 

Ribes macrostachyum Jancz. Bull. Acad. Cracov. 292. 1906; 413. 

Branches and petioles more or less glandular-hispid ; leaves large, 
to 12 cm. wide, more or less 5-lobed, coarsely dentate and denticu- 
late, cordate at the base, glabrous or nearly so except for minute 
hairs or glands; pistillate racemes to 15 cm. long, glandular-pubes- 
cent; bracts linear, 6 mm. long; pedicels 1.5-2 mm. long, equaled 
by the linear bractlets; flowers small, apparently turbinate, very 
pubescent; sepals 2.5 times as long as broad; petals apparently con- 
chiform, exceeding the small, nectariferous anthers; ovary densely 
pubescent. A shrub with large, thin leaves; perhaps a shade form 
of R. viscosum. Neg. 27348. 

Amazonas: Chachapoyas, Mathews, type. 

Ribes ovalifolium Jancz. Bull. Acad. Cracov. 761. 1905; 426. 

Young shoots eglandular; leaves ovate or subelliptic, almost un- 
divided or with 2 obscurely developed lateral lobes, truncate or 
rounded at the base, subglabrous, not glandular, to 3.5 cm. long, 2.5 
cm. wide, often smaller; petioles to 1 cm. long; pistillate racemes 
2-3-flowered, the rachis pubescent; bractlets lanceolate, 2-3 mm. 



FLORA OF PERU 1023 

long; flowers subsessile, often tetramerous or hexamerous, reddish 
brown, slightly pubescent; petals narrow, twice as long as the minute 
anthers; berry globose, puberulent. Compared by the author with 
R. cuneifolium, but lacking the glands of that species. Neg. 4159. 
Ancash: Above Ocros, 3,600 meters, Weberbauer 2771, type. 

Ribes Pentlandi Britton, Mem. Torrey Club 3, pt. 3: 26. 
1893; 403. 

Leaf blades coriaceous, glabrous but densely glandular with ses- 
sile glands, obovate, obscurely if at all 3-lobed, rounded or subcuneate 
at the base, about 2.5 cm. long and wide; petioles to 1 cm. long; 
flowers (staminate) about 10; bracts 3 mm. long; bractlets 1 mm. 
long; pedicels obsolete; sepals 1.5 times as long as broad, ligulate; 
petals conchiform, equaling the stamens; style biparted; ovary 
glandular, the glabrous berry 4 mm. thick. Near R. brachybotrys, 
but differing in the parted style and in the form of the leaves. The 
Peruvian locality is not given by Janczewski. Neg. 27351. 

Lima: San Mateo (Mathews, fide Britton). Bolivia. 

Ribes peruvianum Jancz. Bull. Acad. Cracov. 759. 1905; 407. 

Apparently scarcely distinguishable from R. Dombeyanum but, 
according to the author, the leaves truncate or subcordate at the 
base and verruculose, the bracts narrower, 3-4 mm. long, 0.75 mm. 
wide, the racemes (staminate) denser, the pedicels only 1.5 mm. long. 
Two varieties were distinguished : var. maius, leaves large, anthers 
oblong; and var. minus, leaves 2.5 cm. wide and long, scarcely lobed, 
the anthers roundish. 

Ancash: Prov. Huari, Weberbauer 3777. Tallenga, Weberbauer 
2851 e. Huanuco: Tantamayo, Prov. Huamalies, Weberbauer 3728; 
Mathews. Huariaca, 2,850 meters, 3092, a shrub of 1.5 meters; Sa- 
wada P80. Lima: Near San Marcos, 3,000 meters, Raimondi. 
Cajamarca: Between Magdalena and Contumaya, 3,100 meters, 
Weberbauer 7221. 

Ribes praecox Macbr. Field Mus. Bot. 8: 116. 1930. 

Branches tortuous, obscurely pulverulent, finally glabrous and 
lustrous; leaves few, appearing with the flowers, more or less densely 
glandular with yellow, sessile glands, especially beneath, there also 
minutely puberulent, rugulose-reticulate above, denticulate, more 
or less broadly 3-lobed, ovate-rotund, apparently about 1 cm. 
long; racemes ashy-pulverulent, minutely glandular with sessile 
glands, 1-1.5 cm. long; bracts 2 mm. long, scarcely equaled 



1024 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

by the pedicels, the bractlets 1 mm. long; flowers greenish 
yellow, short-campanulate, 3-3.5 mm. long, the sepals strongly 
reflexed; petals distinctly obovate, 1.5 mm. long, equaled by the 
equally inserted stamens (staminate flowers). R. Weddellianum 
Jancz., 402, of Ecuador, similar to this and R. brachybotrys, has 
eglandular leaves. 

Huanuco: Shrubby trees on rocks, fences, and cliffs, 2,100 meters, 
Llata, 2251, type. 

Ribes viscosum R. & P. Fl. Peruv. 3: 13. 1802. R. glandulosum 
R. & P. op. cit. 13. pL 233, non Grauer, 1784. Coreosma Dombeyana 
Spach, Ann. Sci. Nat. II. 4: 25. 1835. R. Dombeyanum Spach ex 
Jancz. Me"m. Soc. Phys. Hist. Nat. Geneve 35: 408. 1907. 

Younger branches lustrous, at first sparsely stipitate-glandular; 
petioles and rachis densely stipitate-glandular and puberulent, the 
petioles 2-4 cm. long, the rachis 5-8 cm. long, laxly flowered; blades 
rotund-cordate, about 4 cm. wide and long or larger, distinctly 3- 
lobed or rarely with 2 additional, obscurely developed, spreading 
lobes near the base, doubly serrulate, the secondary teeth fine, 
sparsely glandular above, soon glabrous, beneath paler, asperulous, 
especially on the 3 principal nerves and rather prominent veins; 
racemes pendent, the yellowish flowers soon recurving on the elon- 
gating pedicels, these finally about equaled by the glandular-dentate 
bractlets, those 5-6 mm. long; ovary sparsely stipitate-glandular; 
calyx slightly pilose, 4 mm. long; stamens and obovate-roundish 
petals subequal. Description from the type of R. Dombeyanum. R. 
andicolum Jancz., 411, widely distributed north of Peru, if sought 
here, is mentioned under R. bolivianum. No specimen of R. viscosum 
R. & P. was found at Madrid. Spach interpreted the Dombey mate- 
rial at Paris, from the type locality, Tarma, as viscosum, which he 
cited in synonymy and wrote the name on the sheet. In spite of the 
fact that R. viscosum is described as having 5-lobed, strongly veined 
leaves, it seems probable that it is mature material of R. glandulosum. 
Ruiz and Pavon described the fruit as small and purplish, and stated 
that R. glandulosum came from Tarma. The Madrid sheet of R. 
glandulosum purports to be from "Chili," but, as Janczewski remarks, 
it has not been found there since. Its racemes are as much as 5 cm. 
long; leaves obscurely 3-lobed; petioles stipitate-glandular; branch- 
lets glabrate. Neg. 27344. 

Junin: Tarma, Dombey; Ruiz & Pavdn. Camino de Tarma a La 
Oroya, Isern 2259. Lima: Between San Marcos and Hupioc, 3,000 
meters, Raimondi. 



FLORA OF PERU 1025 

Ribes Weberbaueri Jancz. Bull. Acad. Cracov. 763. 1905; 435. 

Branches glandular-setose; blades 3-5-divided, deeply dentate, 
glandular-setulose, only 1.5 cm. long and broad, cordate at the base; 
petioles 1 cm. long; pistillate racemes pendent, 4-5 cm. long, lax, 
glandular-pubescent; bracts persistent, ovate-lanceolate, 6 mm. 
long; pedicels 1-2 mm. long; bractlets 1-1.5 mm. long; flowers 
tubular, reddish, glandular; petals narrowly obovate, inserted above 
the subsessile, nectariferous anthers; berry red, glandular-hispid. 
R. Lehmannii Jancz., of Ecuador, 436, similar, has even smaller, 
obovate-cuneate leaves that are undivided. Neg. 4163. 

Cajamarca: In rocks northwest of Hualgayoc, 3,700 meters, 
Weberbauer 4064, type. 

3. ESCALLONIA Mutis 
Stereoxylon R. & P. 

Flowers 5-parted; ovary 2-3-celled, with 4-6 parietal placentae; 
style 1; capsule completely septicidal. Shrubs and trees of some 
local importance, especially E. pendula and the "chachacoma," E. 
resinosa, commonly planted even to an altitude of 3,400 meters 
(Weberbauer). The red heartwood yields a crimson and violet dye 
(Ruiz & Pawn); the trunks are used for fuel (Hen era}. The latter 
authority records that the tree is attacked by the larva of a lepi- 
dopterous insect called "huaittampu," which is eaten toasted and is 
sold in the markets of Cuzco. Ruiz and Pavon wrote the native 
name as above, but others have varied the vowels, notably as "cha- 
chacuma." Beside the following, E. pulverulenta (R. & P.) Pers., 
Chilean, marked by a parted style with reniform stigmas, has been 
recorded, probably in error; cf. E. pendula. 

Flowers few to many, terminal. 
Flowers racemose. 

Racemes 2-4-flowered E. salicifolia. 

Racemes several-many-flowered. 

Flowers about 6 mm. long, or longer; filaments elongate. 

Branchlets glabrous or pulverulent E. pendula. 

Branchlets densely glandular-hirsute E. Herrerae. 

Flowers 4-5 mm. long; filaments very short. .E. Atahuallpae. 
Flowers panicled. 

Disk of the flower conic. 

Pedicels 6-10 mm. long E. piurensis. 

Pedicels short E. cuneifolia. 



1026 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Disk pulvinate; pedicels short. 

Leaves evanescently glandular toward the subsessile base, 
about 1 cm. wide, often narrower. 

E. resinosa, E. Mandoni, E. angustifolia. 
Leaves obscurely or not at all glandular, about 2 cm. wide, 

the petiole well defined. 
Inflorescence glabrous or glabrate. 

Panicle branches ascending; leaves firm. . .E. paniculata. 
Panicle branches widely divaricate; leaves chartaceous. 

E. Pilgeriana. 

Inflorescence tomentulose E. micrantha. 

Flowers solitary. 

Leaves glabrous or essentially so. 

Leaves mostly 1 cm. long or longer, obovate; branchlets often 

short, stout. 
Branchlets little or not at all tortuous. 

E. patens, E. myrtilloides. 

Branchlets elongating, tortuous E. tortuosa. 

Leaves mostly shorter than 1 cm., scarcely obovate; branchlets 

slender E. corymbosa. 

Leaves glandular-pubescent, linear-revolute E. poliifolia. 

Escallonia angustifolia Presl, Rel. Haenk. 2: 47. 1831. 

Young branches spreading, angled, covered by exuding resin; 
leaves subcoriaceous, glabrous, concolorous, glandular-punctate, 
reticulate-veined, linear-lanceolate, obtuse, callous-serrulate, nar- 
rowed at the base to the short, glandular petiole, 4-6 cm. long, 7 
mm. wide; panicle compound, many-flowered, little longer than the 
leaves; pedicels spreading, angulate, glabrous; capsule nodding, ovoid, 
glabrous, tipped by the elongate style, the stigma capitate-peltate. 
Compare E. resinosa, but apparently distinguished by Mattfeld. 

Tacna: Tacna and Arica, Raimondi (det. Mattfeld). Arequipa: 
Banos de Yura, Raimondi (det. Mattfeld). Without locality: 
Haenke (photo, of type, Mus. Nat. Pragae 991). 

Escallonia Atahuallpae Macbr. Field Mus. Bot. 11: 23. 1931. 

A tree 10-12 meters high, with conspicuously exfoliating branches, 
the branchlets granular- tomentose; leaves lanceolate, cuneate at each 
end, acute, minutely and densely denticulate, glabrate or glabrous, 
sessile or subsessile, to about 10 cm. long, 2.5 cm. wide; racemes pend- 
ent, terminal, simple, 10 cm. long, sparsely glandular; bracts setaceous, 



FLORA OF PERU 1027 

3-5 mm. long; pedicels 4 mm. long; calyx lobes triangular, acute, 
scarcely 2 mm. long; petals erect, oblong, 3 mm. long, equaled by 
the style, this with a peltate stigma; anthers nearly 2.5 mm. long, 
much exceeding the nearly obsolete filaments. Aberrant in the 
genus by reason of the reduced filaments, but, as Dr. Jose" Cuatre- 
casas has suggested to me, perhaps a heterostylous form of E. pendula. 
Ancash: Gravelly river valley, 900 meters, Tambo de Paricota, 
2541, type. 

Escallonia corymbosa (R. & P.) Pers. Syn. 1: 234. 1805. 
Stereoxylon corymbosum R. & P. Fl. Peruv. 3: 14. pi. 234. 1802. E. 
hypsophila Diels, Bot. Jahrb. 37: 412. 1906(7). E. adscendens Rusby, 
Mem. Torrey Club 6: 32. 1896. 

A compact, densely branched, spreading shrub or small tree, the 
branchlets many, appressed-ascending, little tortuous, exfoliating 
and scarred below the tips; leaves subentire or obscurely denticulate, 
glabrous, lustrous, evenose above, dull and paler beneath, somewhat 
petiolate by the cuneate base, acutish, elliptic or slightly obovate, 
5-8 mm. long, 3-4 mm. wide; pedicels 2 mm. long; receptacle sub- 
glabrous, to 3.5 mm. high, 6 mm. broad, the more or less angled tube 
equaled by the narrowly triangular, acute, obscurely and sparsely 
glandular teeth; petals about twice as long as the sepals, obovate- 
spatulate. Perhaps not distinct from E. myrtilloides. Neg. 4168. 

Junin: Tarma, Ruiz & Pavdn. Ancash: Prov. Cajatambo, 4,000 
meters, Weberbauer 2889 (type of E. hypsophila). Catuc, 3,150 
meters, 2506. Cuzco: Raimondi (det. Mattfeld). Monzon, 3,400 
meters, Weberbauer 3720. Ayacucho: 4,000 meters (Pearce?). 
Totorabamba, 3,600 meters, Weberbauer 5468. Huanta, 3,400 
meters, Weberbauer 7487. Huanuco: Ruiz & Pavon, type. Chasqui, 
along stream, 3300; shrub 1.5 meters high; petals white. Tambo de 
Vaca, 3,900 meters, 4425; about 4.5 meters high; the common tree 
of the uplands. Moquehua: Carumas, near brooks on grass steppe, 
3,300 meters, Weberbauer 7267. Without locality: Ruiz & Pavon 
29-93. Bolivia. "Siuba," "chacha," "fassta." 

Weberbauer 5468 was determined by Killip as E. virgata (R. & P.) 
Pers., Chilean, with narrower, petiolate, acute leaves. The type of 
the Rusby species is Mandon 603, Bolivian, referred (in herb.) by 
Engler to E. myrtilloides. Rusby associated with it the Pearce col- 
lection cited above. 

Escallonia cuneifolia (R. & P.) Roem. & Schult. Syst. 5: 328. 
1819. Stereoxylon cuneifolium R. & P. Fl. Peruv. 3: 16. pi. 238. 1802. 



1028 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Apparently similar to E. piurensis, having a high-conic floral 
disk, but the leaves larger, serrate, thick-veined, paler beneath; 
corymbs hirsute; bractlets ovate; calyx teeth minute; capsule tur- 
binate, truncate. Type not seen. The Weberbauer specimen has 
leaves minutely serrulate, about 5 cm. long, 1.5 cm. wide, the close 
panicles puberulent only, but densely sessile-tuberculate-glandular, 
like the calyces. Since the plates of this genus in the Flora Peruviana 
are poor, this collection may possibly represent the Ruiz and Pavon 
species. The disk is distinctly conic. 

Huanuco: Chinchao (Ruiz & Pavon, type). Southwest of Mon- 
zon, 2,500 meters, Weberbauer 3410(1}. 

Escallonia Herrerae Mattf. Notizbl. Bot. Gart. Berlin 10: 
748. 1929. 

Similar to E. pendula, but at once distinguished by the dense 
and long pubescence of reddish, gland-tipped hairs on the younger 
branches; also the lower peduncles are 2-4- rather than 1-flowered. 
As in E. pendula R. & P., the receptacle is free above and cup- 
shaped, with a rim about 1 mm. high, a structure similar to that 
found in Ribes (Mattf eld). Neg. 4171. 

Apurimac: Mollepata, 2,750 meters, Herrera 1237, type. Prov. 
Abancai, Raimondi. Without locality, Gay. "Pauca." 

Escallonia Mandoni Britton, Mem. Torrey Club 3, pt. 3: 
25. 1893. 

Glabrous, the reddish young branches wing-angled; leaves sub- 
sessile, oblanceolate, acute or obtuse, tapering to the base, slightly 
revolute, minutely serrulate, paler and reticulate- veined beneath, 
2.5-4 cm. long, at boul cm. wide; panicles leafy at the base, 2-3 cm. 
long and broad; bracts linear; peduncles and pedicels acutely wing- 
angled, the pedicels 1-2 mm. long; calyx tube 2 mm. long, 5 mm. 
wide, the small teeth obtuse; petals thick, 6 mm. long; filaments and 
style 2 mm. long. Var. microphylla Herzog, Med. Rijks Herb. 27: 
89. 1915, has smaller leaves, more nervose beneath, the inflorescence 
very short. Type referred by Engler (in herb.) to E. resinosa, and 
the differences not apparent, but Mattfeld, Notizbl. Bot. Gart. Ber- 
lin 10: 750. 1929, accepts it as a valid species. 

Cuzco: Without locality, Diehl 2530. Bolivia. 

Escallonia micrantha Mattf. Notizbl. Bot. Gart. Berlin 10: 
747. 1929. 

Branches little angled; leaves oblong-lanceolate, cuneate-atten- 
uate at the base, obtuse, 6-14 cm. long, 1.5-3 cm. wide, entire, puber- 



FLORA OF PERU 1029 

ulent beneath and on the nerves above; panicle terminal, many- 
flowered; pedicels 2-3 mm. long, the filiform bracts about as long; 
petals white, only 3 mm. long; style sometimes deeply divided, the 
2 stigmas broadly reniform; disk plane, pulvinate. Neg. 4173. 
Allied to E. pulverulenta (R. & P.) Pers. and E. millegrana Griseb., 
Chilean, the latter with larger flowers and dentate leaves, the former 
with racemose flowers. By Killip (in herb.) referred toE. paniculata, 
which, however, as to the type at Madrid, is certainly E. floribunda 
var., as determined by Mattfeld. 

Cajamarca: Cascas, 2,000 meters, Raimondi 7739, type. Piura: 
Between Ayavaca and Rio Quiros, 1,800 meters, Weberbauer 6384. 
Without locality: Weberbauer 6365, 6410. "Chachacoma." 

Escallonia myrtilloides L. f. Suppl. 156. 1781. 

A spreading shrub with elongate, straight branches, these bearing 
approximate, short, leafy flowering branchlets; leaves cuneate-obo- 
vate, barely acutish, obscurely lustrous, evenose or slightly veined 
above, mostly 9-12 mm. long, 5 mm. wide, minutely crenate-glan- 
dular or the glands wanting; flowers solitary, more or less nodding; 
calyx glabrous or sparsely sessile-glandular, becoming 8 mm. wide, 
the entire or sparsely glandular, acuminate teeth 3 mm. long, the 
spatulate-obovate petals 6-7 mm. long; style exserted about 5 mm. 
from the fruiting calyx. Compare the closely related E. patens, E. 
tortuosa, and E. corymbosa. The following material perhaps, in part 
at least, belongs elsewhere. The Raimondi specimens were deter- 
mined by Mattfeld. Illustrated, Wedd. Chlor. And. 2: pi. 71. 

Cajamarca: Cutervo, Nancho, Raimondi. Amazonas: Chacha- 
poyas, Weberbauer 2874- Ancash: At 3,600 meters, Raimondi. 
Huanuco: Ambo, Ruiz & Pawn. Arequipa: Raimondi. Junin: 
Raimondi. Huancayo, Raimondi. Apurimac: Raimondi. Without 
locality, Ruiz & Pavon 29-92. Bolivia to Venezuela. "Chacha- 
como," "fassta," "tassta." 

Escallonia paniculata (R. & P.) Roem. & Schult. Syst. 5: 328. 
1819. Stereoxylon paniculatum R. & P. Fl. Peruv. 3: 16. pi. 238. 1802. 

Branchlets reddish, angled, resinous- viscous; blades at first vis- 
cous, especially above, venose, dull, nearly oblong, very minutely 
serrulate, acuminate at each end, about 10 cm. long, 3.5 cm. wide; 
petioles 1-2 cm. long; flowers racemose-paniculate; calyx teeth re- 
duced to calluses; pedicels, calyx, and petals only 2 mm. long; disk 
pulvinate. E. Bridgesii Rusby, Bolivian, has a puberulent inflores- 
cence. Var. floribunda (HBK.) Macbr., comb. nov. (E. floribunda 



1030 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

HBK. Nov. Gen. & Sp. 3: 297. 1819) has smaller, more obtuse leaves 
and denser inflorescence with larger flowers, the petals 3.5 mm. long. 
E. caracasana HBK., to which the type in Madrid has been referred 
by Mattfeld (asE. floribunda var. caracasana Engler), seems, indeed, 
to be the same except that the flowers of the Ruiz and Pavon plant, 
as to the type, appear to be smaller. In any case, the Ruiz and Pavon 
name is earlier. Neg. 29706. 

Huanuco: Iscutunam, Pillao, Ruiz & Pavon, type. Cajamarca: 
Ayavaca, Humboldt, type of the variety. Tambillo, Jelski 312. 
Without locality: Weberbauer 7162, 6391. Colombia. 

Escallonia patens (R. & P.) Killip, Journ. Wash. Acad. Sci. 19: 
192. 1929. Stereoxylon patens R. & P. Fl. Peruv. 3: 13. pi. 234. 1802. 

Shortly and stoutly branched, densely leafy; leaves cuneate- 
obovate, mostly (10) 15-20 mm. long, slightly lustrous to distinctly 
venose above, glandular-serrulate; branches, calyces, and fruit densely 
glandular; calyx 10-15 mm. wide, or wider in fruit, the lobes 5-6 mm. 
long, from a broad base, densely sessile-glandular. Style exserted 
from the fruiting calyx about 1 cm. Description from Geneva and 
Paris specimens; not found at Madrid. "This has been compared with 
E. myrtilloides L. f . from the northern Andes, but is certainly distinct" 
(Killip) ; but he gives no distinctive characters and Engler saw none. 
Negs. 34244, 34669. 

Prov. Huanta: Tambo, 3,200 meters, Weberbauer 5585 (det. 
Killip). Moquehua: Carumas, 3,300 meters, Weberbauer 7267 (det. 
Killip) . Huanuco : Mufia and Pillao, Ruiz & Pavon, type. "Tassta," 
"fassta." 

Escallonia pendula (R. & P.) Pers. Syn. 1: 235. 1805. Stere- 
oxylon pendulum R. & P. Fl. Peruv. 3: 16. pi. 237. 1802. 

A graceful, slender tree with erect-spreading branches, membra- 
naceous, oblong-lanceolate, acute, minutely serrulate leaves, and 
terminal, simple, pendulous racemes sometimes 30 cm. long; leaves 
to 20 cm. long, 5 cm. wide, with some sessile glands along the costa, 
that and the lateral nerves very prominent beneath; racemes gla- 
brous to densely pubescent, the pedicels in fruit nearly 1 cm. long; 
calyx broadly turbinate, the lobes entire; petals linear-spatulate, to 
7 mm. long; stigma 5-lobed; capsule globose. A Gunther and Buch- 
tien specimen with leaves pubescent beneath is probably var. Hum- 
boldtiana Engl. A tree of 4.5-6 meters ( Killip & Smith). Neg. 4147. 
This little tree with pendent flowers is a beautiful sight when in 
blossom; its resistant wood is used for tool handles and other articles; 



FLORA OF PERU 1031 

its leaves are covered with a resin that is applied, crushed as a poul- 
tice, to sprains (Ruiz & Pavon). 

Arequipa: Mollendo, Gunther & Buchtien 709 (det. Bruns as E. 
pulverulenta).Ayacucho: Aina, 750-1,000 meters, Killip & Smith 
23196. Huanuco: Muna, Chaclla, Ruiz & Pavon, type; Raimondi. 
Cajamarca: Cascas, Raimondi (var. Humboldtiana) . Apurimac: 
Hacienda Cotahuadro, at 2,600 meters, Weberbauer 5837. With- 
out locality: Ruiz & Pavdn 50-1, 2900; Weberbauer 6987. North to 
Colombia. "Pumachilca," "pauco." 

Escallonia Pilgeriana Diels, Bot. Jahrb. 37: 531. 1906. 

A shrub with long, pendulous branches, thin, entire leaves, and 
small, white flowers (3-4 mm. long) borne in loose, pyramidal inflores- 
cences about 10 cm. long and 8-9 cm. wide; otherwise nearly E. pani- 
culata HBK., but the very open inflorescence is distinctive, with no 
intermediate forms known. Neg. 4175. 

Cuzco: Urubamba Valley, 1,800 meters, Weberbauer 5061, type. 

Escallonia piurensis Mattf. Notizbl. Bot. Gart. Berlin 10: 
749. 1929. 

A plant with the many-flowered inflorescence of E. paniculata, 
but the puberulent, small leaves, entire and glandular-margined (1-2 
cm. long, 0.6-1 cm. wide), of E. myrtilloides. It is thus equivalent 
to E. resinosa, but the leaf glands are small and obscure and the calyx 
is narrowly turbinate and angulate, only 3 mm. wide at the throat; 
disk narrow, high. Neg. 4176. 

Piura: Huancabamba to Prov. Jae"n, Cajamarca, Raimondi, type. 

Escallonia poliifolia Hook. Icon. 2: pi. 114- 1837. E. macrantha 
Wedd. Chlor. And. 2: 312. 1857. 

A pubescent, glutinous shrub, the densely leafy flowering branch- 
lets 5-7 cm. long; leaves almost glabrous above, densely pubescent 
beneath, rigid, linear-oblong, the entire margins revolute, 2-5 cm. 
long, 3-4 mm. wide; flowers solitary, nodding, on pedicels shorter 
than the leaves; calyx pubescent, and tuberculate with short, hemi- 
spheric tubercles, the teeth broadly subulate; petals 1 cm. long, 4-5 
mm. wide at the apex, spatulate, exceeding the stamens, these equal- 
ing the style. Name written by the author polifolia. 

Amazonas: Chachapoyas, Mathews, type. Between Cajamarca 
and Chachapoyas, 3,350 meters, Raimondi; type of E. macrantha. 



1032 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Escallonia resinosa (R. & P.) Pers. Syn. 1: 235. 1805. Stereoxy- 
lon resinosum R. & P. Fl. Peruv. 3: 14. pi. 235. 1802. E. multiflora 
Presl, Rel. Haenk. 2: 48. pi. 57. 1831. 

Flowering branches virgate, lustrous, reddish, angled, with many 
short branchlets bearing crowded, oblanceolate leaves, rounded at 
the apex; blades 2-3 (-4) cm. long, about 7 mm. wide, glaucous be- 
neath, slightly lustrous and venose above, glandular-margined, the 
glands caducous, stipitate toward the petiolar base; flowers densely 
thyrsoid-paniculate; calyx glabrous, 5 mm. wide in fruit, the teeth 
short; petals and stamens subequal, about 5 mm. long; style with a 
peltate-lobate stigma scarcely 4 mm. long. Wood very durable for 
use in the ground (Raimondi). Neg. 29707. 

Cajamarca: San Miguel, Weberbauer 3916. Huaraz, 3,200 meters, 
Weberbauer 324-4- Huanuco: Ruiz & Pavon, type. Mito, 2,700 
meters, 1556. ^!mbo, Dombey. Without locality, Gay. Lima: Rio 
Blanco, 3,000-3,500 meters, Killip & Smith 21693. Viso, 2,700 me- 
ters, loose soil of steep canyon sides, 562; a small tree; petals white, 
the anthers orange-red. Rio Chillon, above Obrajillo, 3,200 meters, 
rocky canyon, Pennell 14-405. Arequipa: Raimondi. Ayacucho: 
Below Totorabamba, 3,400 meters, Weberbauer 5477. Cuzco: Rai- 
mondi. San Sebastian, 3,400 meters, rocky canyon slope, Pennell 
13630; a tree of 3-6 meters. Near Rio Quencomayo, below Colqui- 
pata, 3,300 meters, rocky, brushy slopes and banks, Pennell 13785. 
Near Cuzco, 3,200-3,500 meters, Herrera 692. Huariaca, 3,000 me- 
ters, 3125 (det. Killip). Bolivia. "Chachas," "tiri encarnado," 
"chachacoma," "chachacomo," "chachacuma." 

Escallonia salicifolia Mattf. Notizbl. Bot. Gart. Berlin 10: 750. 
1929. 

Branches erect, little angled, evanescently puberulent or gla- 
brous, often verruculose-glandular, with many short, densely leafy 
flowering branchlets; blades narrowly lanceolate, 2-3.5 cm. long, 
4-8 mm. wide, acute, cuneate to the petiole, this only 1-2 mm. long, 
the margins, especially toward the apex, minutely serrulate and 
puberulent, the leaf otherwise glabrous, inconspicuously veined; 
racemes 2-4-flowered; flowers pendent, the petals about 12 mm. long, 
inserted with the filaments (7 mm. long) on a disk, slightly longer 
than the peltate style; calyx turbinate-subglobose, the strongly 
dilated, cuplike free portion about 7.5 mm. broad, the lobes 5 mm. 
long, broadly triangular-subulate. Neg. 4181. Suggestive of the 
Chilean species, but with the disk of E. resinosa and relatives, that 
is, broad and plane, cushion-like. 



FLORA OF PERU 1033 

Arequipa: Between Monqui and Chavcona, Raimondi, type. 
Escallonia tortuosa HBK. Nov. Gen. & Sp. 3: 295. 1819. 

Glabrous, with spreading, tortuous, flexuous branchlets; leaves 
coriaceous, dull above or slightly lustrous, the veins faint, mostly 
12 mm. long, 6 mm. wide, obovate-cuneate, acutish-crenate-serru- 
late, with sessile glands in the serrations; calyx lobes glabrous, short- 
glandular, triangular-acute, 3 or 4 times shorter than the linear- 
spatulate petals and the calyx, the tube hemispheric. Seems to be 
referable to E. myrtilloides, or a variety of it. The type at Paris 
is sterile; it is more openly branched than E. corymbosa (as to type), 
with larger leaves, but lacks the numerous small branchlets, the 
leafless branches being very tortuous. 

Peru: Probably. North to Colombia. 

4. HYDRANGEA L. 

Cornidia R. & P.; Sarcostyles Presl. 

Reference: Briquet, Ann. Cons. Jard. Bot. Geneve 20: 393-417. 
1919. 

Only two species of the Hydrangeas described here have, in part, 
petaloid flowers, H . peruviana and H. Weberbaueri, the former dis- 
tinct by the coherence of its petals, the latter by its wine-red, peta- 
loid sepals. H. peruviana, in spite of its name, is apparently known 
only from Ecuador, where collected by Tafalla. Both these species, 
as well as H. Jelskii, are slender-stemmed, scandent shrubs of the 
montana zone, and it is questionable whether the other species 
described as trees are described correctly. With the exception of H. 
tarapotensis and H. Jelskii with 1-3 usually coherent, subulate 
styles, the following species have the styles thickened in age and 
3-4 in number, except those species with petaloid flowers, which 
have only 2 thickened styles. The petals are 4, the stamens 8. Bri- 
quet, Compte Rendu Soc. Phys. Hist. Nat. Geneve 36: 38-43. 1919, 
valued these differences properly as sectional in character, relegating 
Cornidia, still maintained by some students, to synonymy. But his 
species concepts, with more collections, will probably prove to have 
been too finely drawn. Besides the species listed here, Hydrangea 
opuloides Koch, called "hortensia," a native of China and Japan, 
often is planted for ornament, on account of its showy, white, pink, 
or blue flowers. 



1034 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves stellate-pubescent beneath, sometimes very sparsely so. 
Leaves densely stellate- tomentose; styles 1-3, subulate. H. Jelskii. 
Leaves sparsely stellate-pubescent. 
Petaloid flowers none; petals free. 

Inflorescence cymose-pseudo-umbellate; styles short, thick- 
ened. 
Leaves oblong-ovate-elliptic, rarely 7 cm. wide; cymes 

dense. 
Leaves suboblong, acuminate, finally glabrous above. 

H. Sprucei. 

Leaves obovate, obtuse or apiculate, somewhat stellate- 
pubescent above H. Mathewsii. 

Leaves broadly elliptic, 6-10 cm. wide; cymes lax. 

H. Briquetii. 
Inflorescence thyrsoid; styles almost subulate, 2 mm. long. 

H. tarapotensis. 

Petaloid flowers present, showy; petals coherent. . H. peruviana. 
Leaves glabrous beneath. 

Outer flowers radiate; styles 2 H. Weberbaueri. 

None of the flowers radiate; styles 3-4. 

Filaments 1.5-2.5 mm. long; leaves acuminate. 
Leaves to 5 cm. wide, remotely serrulate; cymes dense. 

H. Preslii. 
Leaves to 10 cm. wide, entire or undulate; cymes lax. 

H. Briquetii. 
Filaments 0.5-1 mm. long; leaves obtuse, entire. 

H. ecuadorensis. 

Hydrangea Briquetii Engler, Pflanzenfam. ed. 2. 18a: 207. 
1930. Cornidia umbellata R. & P. Syst. 91. 1798. H. umbellata Briq. 
Ann. Cons. Jard. Bot. Geneve 20: 411. 1919. 

An ample-leaved shrub or small tree with stellate-puberulent 
branchlets and inflorescences, the latter umbellately composed of 
lax cymes; petioles to 3 cm. long; blades to 20 cm. long, half as 
wide, elliptic or oblong-elliptic, caudate-acuminate, obtuse at the 
base, entire, coriaceous, lustrous and glabrous above, paler and 
minutely appressed-stellate-puberulent beneath, the 6-10 lateral 
nerves beautifully anastomosing; pedicels usually 0.5-4 mm. long; 
sepals and petals 4, the sepals 0.4 mm. high, the petals 1.5 mm. 
long; filaments 2-2.5 mm. long, the anthers 1 mm. long; styles 3, 



FLORA OF PERU 1035 

about 1 mm. long. The glabrous, Chilean H. integerrima (H. & A.) 
Engler, to which Peruvian material has sometimes been referred, 
has persistent stamens and 3 free, subulate styles. Neg. 8290. 

Without locality, Ruiz & Pavdn, type. Huanuco: Chinchao, 
Ruiz & Pavon 12-10. Pampayacu, Sawada P44. Muna, 2,100 
meters, 3962. 

Hydrangea ecuadorensis Briq. Ann. Cons. Jard. Bot. Geneve 
20: 410. 1919. 

Branchlets subglabrous; blades elliptic, to 10 cm. long, 1 cm. 
wide, obtuse or very shortly acuminate, entire or obscurely undulate, 
obtuse at the base, coriaceous, glabrous, lustrous above, paler 
beneath, the 7-10 lateral nerves prominent; petioles 1-1.5 cm. 
long; inflorescence ample, the peduncle conspicuously enlarged, 
shortly and densely appressed-stellate-pubescent; cymes equally or 
subequally umbellately disposed, the primary axes to 4 cm. long; 
flowers subsessile, all fertile; sepals 4, ovate-obtuse, 4 mm. long; 
petals ovate-elliptic, to 1.5 mm. long; stamens 8, the filaments 0.5-1 
mm. long; styles 3 or 4, conspicuous, finally extrorse-curved ; ovary 
3-4-celled; capsule 1.5-2 mm. long, inconspicuously nerved. Sepa- 
rated by the author from H. Preslii because of its umbellate 
instead of subspheric inflorescence; the type, Spruce 5058, from an 
unknown locality, presumably Ecuador. H. Bangii Rusby, Bolivian, 
has longer, persistent filaments and obovate, emarginate leaves. 
Neg. 27366. 

Peru: Probably. Ecuador. 

Hydrangea Jelskii Szyszyl. Rospr. Diss. Cl. Math. Phys. Acad. 
Cracov. 29: 218. 1895. 

Younger branchlets and leaves beneath rusty-stellate- tomentose; 
blades elliptic-lanceolate, little narrowed to the obtuse or rounded 
base, acutish, coriaceous, subglabrous above, mostly 10-20 cm. 
long, half as wide; inflorescence cymose, the peduncles 1-5 cm. 
long; flowers greenish, 2 mm. long, the pedicels half as long; calyx 
teeth triangular, acutish; petals concave, sessile equaled by the 8 
glabrous filaments; styles 1-3, short. A more or less climbing 
shrub, similar to H . tarapotensis in the form of the inflorescence, 
but the narrower leaves densely pubescent beneath. 

Cajamarca: Tambillo, Jelski 332, type; Raimondi (det. Mansfeld). 
Amazonas: Molinopampa, 2,400 meters, Weberbauer 4-385. Near 
San Carlos, 2,800 meters, Weberbauer 7157. 



1036 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Hydrangea Mathewsii Briq. Ann. Cons. Jard. Bot. Geneve 20: 
413. 1919. 

Branchlets rusty-stellate- tomentose, the internodes elongate; 
blades obovate, crenate-denticulate toward the obtuse, apiculate 
apex, cuneate-linear to the obtuse or subobtuse base, to 12 cm. long 
and half as wide, coriaceous, rugose, sparsely stellate-pubescent above, 
more or less rusty-tomentose beneath, the lateral nerves (about 10) 
prominent on both sides; petioles 1.5-2 cm. long; cymes densely 
rusty-stellate-pubescent, with primary axes 1-2 cm. long, the pedicels 
to 3 mm. long, often obsolete; flowers all fertile; sepals and petals 4, 
the former 2-4 mm. long, the latter to 1.5 mm. long; stamens 8, the 
filaments 1.5 mm. long, the anthers half as long; styles 3, to 2 mm. 
long. Compare H. Jelskii. Neg. 8291. 

Amazonas: Chachapoyas, Mathews, type. 

Hydrangea peruviana Moric. in DC. Prodr. 4: 14. 1830. Corni- 
dia peruviana Small, N. Amer. Fl. 22: 161. 1905. 

Branchlets tetragonous, stipitately rusty-stellate-tomentose, the 
internodes shorter than the leaves; petioles to 1 cm. long; blades ovate 
or subovate to elliptic, to 7.5 cm. long, 4.5 cm. wide, obtuse at each 
end, more or less densely stipitate-stellate-pubescent, coriaceous, 
regularly and obtusely serrate or crenate-dentate above the middle, 
the lateral nerves about 10, impressed above; inflorescence shortly 
rusty-tomentose, 8-12 cm. broad; fertile pedicels 1-4 mm., the sterile 
3 mm. long; fertile flowers small, the sepals scarcely 0.1 mm. high, 
the petals 1.5 mm. long; stamens and 2 styles 1 mm. long; sterile 
flowers few, the petaloid sepals about 7-9 mm. long. The type 
by Ruiz and Pavon from "Guayaquil," actually was collected by 
Tafalla. Neg. 8036. 

Peru: Probably. Ecuador. 

Hydrangea Preslii Briq. Ann. Cons. Jard. Bot. Geneve 20: 409. 
1919. Sarcostyles peruviana Presl in DC. Prodr. 4: 16. 1830; Rel. 
Haenk. 2: 54. pi. 60. 1831. 

Similar to H. ecuadorensis, but the inflorescence consisting of 
6-9 rays, the shortest only 1.5 cm. long, the primary axes thus 
relatively short; otherwise, from description, apparently the same 
as Briquet's species and quite possibly the earlier name. The 
obscure H. serratifolia (H. & A.) Engler, described as Chilean but 
"thought by Phillipi to be Peruvian" (Engler), has elongate, per- 
sistent stamens and conspicuously dentate leaves, the styles subulate 
as in H. tarapotensis but free. 



FLORA OF PERU 1037 

Huanuco: Haenke, type. Rio Posuso, 3,100 meters, Weberbauer 
6774, 6776 (both det. Engler). Junin: Above San Ramon, 1,300- 
1,700 meters, Schunke A36. 

Hydrangea Sprucei Briq. Ann. Cons. Jard. Bot. Geneve 20: 
412. 1919. 

Branchlets rather sparsely stellate-pubescent; leaves nearly 
oblong, the larger more than 20 cm. long, 8 cm. wide, entire, acumi- 
nate, obtuse or acute at the base, coriaceous, glabrous in age but 
minutely and sparsely stellate-puberulent on both sides at first, the 
lateral nerves about 10, prominent beneath; inflorescence condensed, 
densely many-flowered, the primary axis 1.5-2 cm. long, rusty- 
stellate- tomentose; flowers all fertile; sepals and petals 4, the sepals 
ovate, 2-4 mm. long, the petals 1.5 mm. long, about equaled by the 
filaments and 3 or 4 styles; anthers 1 mm. long. Neg. 8292. 

San Martin: Zepelacio, 1,100 meters, in forest, Klug 3653; 
flowers wine-red. Mt. Campana, near Tarapoto, Spruce 4328, type. 
Colombia. 

Hydrangea tarapotensis Briq. Ann. Cons. Jard. Bot. Geneve 
20: 415. 1919. 

Branchlets shortly and densely stellate-puberulent; blades obo- 
vate, obtuse or shortly acuminate, cuneate to the acute base, to 
10 cm. long and 6 cm. wide, entire, coriaceous, glabrous above, 
beneath minutely appressed-stellate-pubescent, the 6-7 lateral nerves 
prominent; petioles to 2.5 cm. long; inflorescence composed of 
several series of bracteate umbels, ample; pedicels none, or 4 mm. 
long; flowers all fertile; sepals and petals each 4, the petals oblong- 
elliptic, only 2 mm. long; stamens twice as many, 2.5-3 mm. high, 
the anthers 1 mm. long, styles 1-3 and 2 mm. long, often coherent 
in a column. The species is unique in the united styles, when 
these are more than one. Neg. 8293. 

San Martin: Tarapoto, Spruce 4349, type. 

Hydrangea Weberbaueri Engler, Pflanzenfam. ed. 2. 18a: 
206. 1930. 

A vine, the stems 3 cm. in diameter; leaves sharp-pointed, 
roundish-obovate, glabrous beneath; sterile flowers with red, petal- 
like sepals. Allied to H. peruviana in its finally thickened style. 
Neg. 4146. 

Amazonas: Cheto, near Chachapoyas, 2,000 meters, Weberbauer 
4372, type. San Martin: Raimondi. 



1038 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5. PHYLLONOMA Willd. 

Shrubs with alternate leaves which bear, toward their caudate 
apex, from the costa, a short cyme or raceme of small flowers. 
Fruit a berry. Rusby has treated the genus as a distinct family, 
Phyllonomaceae. 

Leaves closely serrulate P. Weberbaueri. 

Leaves entire or very remotely denticulate. 

Leaves entire P. integerrima. 

Leaves remotely denticulate P. ruscifolia. 

Phyllonoma integerrima (Turcz.) Loes. ex Engler, Pflanzen- 
fam. ed. 2. 18a: 224. 1930. Dulongia integerrima Turcz. Bull. Soc. 
Nat. Mosc. 31, pt. 1: 454. 1858. 

Petioles 5-7 mm. long; blades about 8 cm. long, 3 cm. wide, 
caudate-acuminate, the slender pedicels borne at the base of the 
acumen. A shrub 1-4 meters high ( Ule). Neg. 34235. 

Loreto: Cerro de Escalera, Ule 6754 (det. Loesener). Colombia. 

Phyllonoma ruscifolia Willd. ex Roem. & Schult. Syst. 6: 
210. 1820. Dulongia acuminata HBK. Nov. Gen. & Sp. 7: 78. pi. 
623. 1824. 

Similar to P. integerrima, but the blades sometimes smaller and 
the petioles shorter; blades mostly 4-5 cm. long, 1-2 cm. wide; 
petioles 4-5 mm. long; peduncle 2-4 mm. long; petals ovate, persis- 
tent. The dentation is very obscure in the Humboldt Colombian 
specimen. A shrub of 8 meters (Weberbauer). 

Loreto: Moyobamba, 1,100 meters, Weberbauer 4761. Bolivia 
to Colombia. 

Phyllonoma Weberbaueri Engler, Pflanzenfam. ed. 2. 18a: 
225. 1930. 

Differing from the other species in having the short pedicels 
borne on the lower half of the long point of the densely and sharply 
serrate leaves. Neg. 4141. 

Puno: Tambo Ichubamba, Sandia, 2,000 meters, Weberbauer 
1322, type. 

85. CUNONIACEAE 

Reference: Engler, Pflanzenfam. ed. 2. 18a: 229-261. 1930. 

A family of shrubs and trees. About a third of the known species 
are Peruvian, a majority with pinnate leaves, these, typically at 



FLORA OF PERU 1039 

least, opposite or whorled, and in this respect differing usually from 
the closely related Saxifragaceae. Some have regarded Brunellia as 
constituting a distinct family, but it is nearer to Weinmannia than 
to any other group, and therefore, from a practical standpoint, 
the genera should be associated under one family name. 

Leaves opposite. 

Flowers in solitary or corymbose clusters; follicles usually 4-5. 

1. Brunellia. 

Flowers fascicled or solitary, in racemes; follicles 2.3. Weinmannia. 
Leaves alternate 2. Gumillea. 

1. BRUNELLIA R. & P. 

Trees with usually coarsely hairy flowers and capsules. The 
flower parts vary from 4-7, with stamens twice as many as the 
sepals, borne on an 8-11-lobed disk. The original illustration, 
R. & P. Prodr. 71. pi. 12, is of the flower and fruit only, but the 
genus is well illustrated (mostly after Humboldt and Bonpland) in 
Pflanzenfam. op. cit. 227. One species, B. aculeata R. & P. Syst. 
127. 1798, omitted, is, according to Pilger on the type sheet at 
Madrid, an unknown species of Fagara. The genus needs careful 
revision. Some of the Bolivian species may be the same as Peruvian 
ones. Apparently, none of the earlier Colombian names apply, but 
because of the uncertainty they are referred to here. In Peru only 
on the eastern ranges, between 1,800 and 3,000 meters (Weberbauer). 

Leaves simple. 

Leaves obtuse, mostly narrower than 2.5 cm B. Briquetii. 

Leaves acute, 3.5 cm. wide or wider B. dulcis. 

Leaves, at least some of them, pinnate. 

Leaves glabrous beneath. 

Leaves opposite; leaflets 3 pairs, 3 cm. wide or wider. 

B. Weberbaueri. 

Leaves mostly ternate; leaflets 5-7 pairs, narrower. .B. ternata. 
Leaves tomentose or at least puberulent beneath. 
Leaflets more than 3 pairs. 

Sepals 6; petioles and inflorescence glabrous or glabrate. 

B. hexasepala. 

Sepals 4-5; petioles and inflorescence usually densely pubes- 
cent. 



1040 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Inflorescence loosely but densely gray-brown-pilose. 

B. dichapetaloides. 
Inflorescence closely reddish-puberulent-tomentose. 

B. brunnea. 
Leaflets 2 (-3) pairs B. inermis. 

Brunellia Briquetii Baehni, sp. nov. 

Arbor ut videtur ramis baud angulatis, primum ramulis petiolis 
pedunculisque plus minusve brunneo-griseis cum pilis crispe tomen- 
tulosis etiam nonnullis nigris firmiusculis intermixtis lanato-pubes- 
centibus demum glabratis; petiolis superne leviter sulcatis ca. 10 
mm. longis; foliis simplicibus oppositis integris vel obscurissime 
undulato-crenatis oblongis fere basi bene acutis, ad apicem paullo 
vel vix angustatis, apice ipso obtusis vel rotundatis, ad 7 cm. longis, 
2.5 cm. latis, supra nitidulis glabris, nervis venisque obscuris, subtus 
dense ferrugineo-tomentulosis demum glabratis prominente denseque 
reticulato-venosis; paniculis 5-7 cm. longis, strictis, ramulis ca. 1 
cm. longis; floribus ignotis; calycibus 7 mm. latis, segmentis 4-5 
late ovato-acutis utrinque fulvo-puberulis; capsulis sessilibus den- 
sissime fulvo-hirsutis abrupte acutis ca. 7 mm. longis. A species 
remarkable for the small, subentire, opposite leaves. Neg. 27374. 

Amazonas: Chachapoyas, Mathews; type in herb. Geneva. 

Brunellia brunnea Macbr. Candollea 5: 361. 1934. 

Branches and inflorescence velvety with a short, deep brown 
tomentum, not at all hirsutulous, the branches thick, only slightly 
angled; leaves about 30 cm. long, with mostly 8 pairs of subequal, 
nearly sessile leaflets, these rounded-cordate at the base, acuminate, 
to 15 cm. long, 4.5 cm. wide, undulately crenate-denticulate, scarcely 
lustrous and almost glabrous above, paler beneath and on the veins 
minutely pilose, the midnerve brown-tomentulose, moderately retic- 
ulate-veined on both sides; inflorescence loosely paniculate, 15 cm. 
long, the branchlets curved-spreading; flowers subsessile, 3 mm. 
long, the 5 thick sepals rusty-puberulent on both sides, ovate, acute, 
nearly 2.5 mm. long, equaled by the subulate filaments, these long- 
hirsute at the base, finally exserted; capsules hirsutulous. Near 
the Bolivian B. crenata Engler (B. rhoides Rusby), with 3-4 pairs 
of leaflets and calyx and filaments only 2 mm. long. B. integrifolia 
Szyszyl., Venezuelan, has rounded or barely acute leaflets, strongly 
oblique at the base. 

Cuzco: Between the tambos Tres Cruces and Tambomayo, 2,150 
meters, Weberbauer 6969, type. 



FLORA OF PERU 1041 

Brunellia dichapetaloides Macbr. Candollea 5: 361. 1934. 

Branchlets soon glabrous and lustrous; inflorescence and leaves 
beneath, with the rachis, more or less densely and shortly ashy- 
brown-pilose; petioles 6 cm. long; leaves 5-6-foliolate, 20-25 cm. 
long; leaflets oblong-elliptic, oblique at the base, acute or acuminate, 
subequal, 10-15 cm. long, 3.5-4 cm. wide, obscurely crenate-dentate, 
glabrous and very lustrous above; inflorescence many-flowered, 10 
cm. long and about a third broader, the peduncle 2 cm. long, the 
3 principal branches 3-4 cm. long; flowers crowded on pedicels 
1.5 mm. long, sordid-puberulent; sepals 4, broadly oblong, acutish, 
2 mm. long; stamens exserted, glabrous except at the base; anthers 
red-puncticulate; follicles 4, pilose. B. Brittonii Rusby, Bolivian, 
has leaflets 4.5-6 cm. wide, sepals 6 mm. long, the filaments exserted. 
B. Oliveri (see also under B. Weberbaueri) differs from B. dichapeta- 
loides in its pilose-hirsute branches and obtuse leaves. Nearer the 
latter is B. comocladifolia Humb. & Bonpl., Colombian (not Peru 
as in Index Kew.), with serrulate leaflets, and B. Funckiana Tul., 
similar, but the leaflets acute at the base. B. propinqua HBK., 
reported as Peruvian, is, fide Tulasne, Colombian; it, as to type, 
has smaller leaflets than these other species, 3-4 pairs, obtuse or 
acutish, shorter than 10 cm., and a densely rusty-tomentose inflores- 
cence, the peduncle longer than 5 cm. It is not clear that the 
Peruvian species can be distinguished from B. comocladifolia. 

Junin: Chanchamayo Valley, 1,500 meters, Schunke 83, type; 84. 

Brunellia dulcis Macbr. Candollea 5: 362. 1934. 

A tree of 12 meters, with thick, strongly sulcate-angled, some- 
what appressed-lanate branchlets; leaves simple, oblong-elliptic, 
irregularly denticulate, about 20 cm. long, 5.5 cm. wide, cuneate- 
attenuate at the base, the apex acute, rigid-chartaceous, more or 
less lanate on the nerves above, glabrate, scarcely lustrous, the 
veins obscure, beneath slenderly reticulate-veined and appressed- 
rusty-lanate, the lateral nerves 15-19, prominent beneath; petioles 
1.5-2 cm. long; panicles 15 cm. long, 3-4 cm. wide, the few branches 
densely rusty-lanate and sometimes leafy-bracteate; flowers capi- 
tate-congested, grayish white, nearly 3 mm. long, the sepals ovate, 
acute; stamens 8; filaments 3-4 mm. long, pilose at the base; follicles 
4. The other simple-leaved species (besides B. Briquetii) are: B. 
acutangula Humb. & Bonpl. and B. tomentosa Humb. & Bonpl., 
both Colombian, with whorled leaves, those of the latter 10-20 cm. 
long, 5 cm. wide, merely acute; the former species has flattened 



1042 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

branchlets, puberulent leaves about 10 cm. long, 3.5 cm. wide, 
heavily reticulate-veined beneath, obscurely denticulate. Appar- 
ently too similar to B. tomentosa and from the same country is B. 
Stuebelii Hieron., the branchlets quadrangular, the leaves barely 
acute, opposite, 12-14 cm. long, 3.5-4 cm. wide. Well marked is 
B. ovalifolia Humb. & Bonpl., collected as near as Loja, the leaves 
rotund-elliptic. B. Goudotii Tul. is similar, but the inflorescence is 
said to be densely reddish-tomentose. 

Huanuco: Pan de Azucar, Sawada 74, type. 

Brunellia hexasepala Loes. Bot. Jahrb. 37: 531. 1906. 

Branchlets subglabrous; leaves opposite, on petioles 4-9 cm. 
long, 4-7-foliolate; leaflets rigid, at first, especially beneath, long- 
sericeous, in age glabrescent, prominently and densely reticulate- 
veined beneath, the lateral nerves 18-25, coarsely crenate-serrate, 
rounded at the base, acute at the apex, 4-10 cm. long, 1-3 cm. wide; 
panicles (staminate) 7-11 cm. long, 2.5-5 cm. wide, on peduncles 
3.5-5.5 cm. long, glabrous or sparsely setulose; flowers green, glo- 
merate, subsessile, the 6 free, ovate-lanceolate lobes densely hairy 
within, less so outside, 3.5 mm. long; stamens half as long, the 
filaments very short; disk obscure; follicles spreading, to 13 mm. 
long; seeds 3.5 mm. long, lustrous brown. A shrub 5 meters high. 
Neg. 3276. 

Puno: Sandia, 3,000 meters, Weberbauer 734. 

Brunellia inermis R. & P. Syst. 127. 1798. 

Branchlets only slightly angled, evanescently lanate-puberulent, 
in age glabrate like the leaves beneath; petioles 4 cm. long; leaflets 
2-3 pairs, finely reticulate-veined beneath, glabrous and lustrous 
above, minutely serrulate, unequal at the rounded base, acute at 
the apex, about 15 cm. long, 4.5-8 cm. wide, the lateral nerves 
prominent beneath, only 4-5 mm. apart, numerous (about 25); 
inflorescence densely rusty-pilose, the short branches stoutly long- 
pedunculate, 30 cm. long in fruit; sepals 5, almost 3 mm. long, 
deltoid, acute, appressed-strigose or nearly glabrous within; follicles 
sessile, short-hispid and strigillose, about 7 mm. long. The leaves 
seen consisted of 2 pairs of leaflets and one terminal one, this petio- 
lulate, the lower pair not much smaller than the 3 upper leaflets. 
Ruiz and Pavon described the leaves as ternate and simple. Illus- 
trated, R. & P. Fl. Peruv. 4: pi. 414- 



FLORA OF PERU 1043 

Huanuco: Mufia, on the road to Tambo Nuevo, Ruiz & Pavdn, 
type. Junin: Huacapistana (Weberbauer, 251). Rio Paucartambo, 
Weberbauer 6677. Without locality: Ruiz & Pavdn 23-47. 

Brunellia ternata Loes. Bot. Jahrb. 37: 533. 1906. 

Branchlets terete, densely lenticellate; leaves 8.5-18 cm. long, 
the petiole 1.5-2.5 cm. long; petiolules 1-3 mm. long; leaflets rigid, 
glabrous, dull and reticulate-veined beneath, the lateral nerves 
11-16, crenulate-serrulate, ovate or narrowly ovate-elliptic, the 
more or less unequal base cuneate-obtuse or acute, the apex obtuse 
or scarcely apiculate, mostly 2.5-7.5 cm. long; panicles (pistillate) 
3-4 cm. long, more or less appressed-pilose, the peduncles 4-11 mm. 
long; flowers glomerate, 5-6-parted, the ovate-deltoid segments 
densely hairy within and at the apex outside, 2.5-3 mm. long; 
staminodia(?) about half as long, inserted between 10-12 pilose disk 
lobes; ovaries about 5 and 7 and 2-ovulate. A shrub of 3 meters. 
Neg. 3278. 

Puno: Sandia to Chunchosmayo, 2,000 meters, Weberbauer 1144, 
type. Huanuco: Playapampa, 2,700 meters, wet, rocky forest, 
4882; flowers white. 

Brunellia Weberbaueri Loes. Bot. Jahrb. 37: 532. 1906. 

Similar to B. ternata, but the leaves, at least mostly, opposite, 
with fewer leaflets, these 5.5-11 cm. long, more conspicuously 
reticulate-veined above, and with 13-16 lateral nerves; perianth 
segments puberulent within, glabrous outside; stamens twice as 
long, the elongate filaments filiform. A shrub of 4 meters. B. 
Oliveri Britton (B. boliviana Rusby), Bolivian, has 5-6 pairs of 
crenate-serrate, obtuse leaflets; compare B. dichapetaloides. B. 
racemifera Tul., Colombian, has 4-6 pairs of glabrous, acute leaflets. 
Neg. 3279. 

Huanuco: Monzon, 2,000 meters, Weberbauer 3551, type. Rio 
Posuso, 2,300 meters, Weberbauer 6781. 

2. GUMILLEA R. & P. 

Shrubs with terete, pubescent branches and alternate, pinnate 
leaves, the leaflets ovate-lanceolate. Flowers small, terminal, 
crowded in pubescent spikes. Sepals 5. Petals none. Stamens 5, 
inserted at the base of a conic receptacle, with compressed filaments 
and nearly round anthers. Ovary free, with 2 short, spreading 
styles. Capsule 2-beaked, many-seeded, the small seeds nearly 



1044 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

round. Engler remarks that because of the alternate leaves the 
plant doubtfully belongs to this family. 

Gumillea auriculata R. & P. Syst. 74. 1798. 

Branches becoming glabrous and lustrous, the younger ones and 
the branchlets, like the costa of the leaflets beneath, densely rusty- 
strigose or hispidulous; leaflets entire, the larger 9 cm. long, 3 cm. 
wide, oblong, oblique at the acute base, short-acuminate, about 
7 pairs, alternate, slightly lustrous on both sides and reticulate- 
veined. Illustrated, R. & P. Prodr. pi. 7. Neg. 29330. 

Huanuco: Mufia, Ruiz & Pavon, type. 

3. WEINMANNIA L. 

Reference: Engler in Mart. Fl. Bras. 14, pt. 2: 156-170. 1871; 
Pflanzenfam. ed. 2. 18a: 250-257. 1930. 

These shrubs or small trees, with small flowers, most often 
borne in fascicles on an elongate axis (the inflorescence is here, for 
convenience, called a raceme) exist in a great number of species, 
separated on characters often so slight that they might be described 
as casual. The genus is distinct enough in its flower parts, these 
4 (-5), the anthers 2-lobed, the ovary 2 (-3) -celled, the seeds pubes- 
cent. Kuntze adopted the name Windmannia P. Br., making some 
new combinations under it. I regret that Dr. Jose" Cuatrecasas was 
unable to prepare the account of the Peruvian species, as had been 
planned in connection with his monograph of the genus. My ten- 
tative key and notes, with studies we had made jointly at Madrid, 
have served as the basis for the following treatment. Several 
Andean species not as yet known from Peru have been included, 
especially when they have been misinterpreted or imperfectly 
known. Intentionally omitted areW. intermedia Cham. & Schlecht., 
Mexican, and W. trichosperma Cav., Chilean. The first was credited 
to Peru by Pampanini on the basis of a "Pavon" specimen in herb. 
Boissier, labeled "Peru," but, as is well known, there are a number 
of specimens in that herbarium so labeled whose origin evidently 
is Mexican. Besides, I have not identified (having found no speci- 
mens at Madrid or elsewhere so named) W. corymbosa R. & P. Fl. 
Peruv. 4: pi. 330, 1802, and W. dentata R. & P. op. cit. pi. 334. As 
in the Lauraceae, I have accepted volume four of the Ruiz and 
Pavon work as published, because the species are nearly all identi- 
fiable with the authors' own specimens. Unfortunately, in preparing 
this account in its present form, volume four has not been available. 



FLORA OF PERU 1045 

There is a conflict between the names of D. Don. and Moricand 
(both 1830), and they have been used in accord with the biblio- 
graphic finding of Miss Edith M. Vincent of Field Museum. Ruiz 
and Pavon have noted the names "tiaca" and "machi" for species 
"whose trunks furnish beautiful boards" for furniture and other 
construction. 

Leaves simple, at least some of them, never typically pinnate; if 
trifoliolate, the lateral leaflets reduced, auricle-like. 

Ovary pubescent W. trichocarpa. 

Ovary glabrous. 

Filaments pilose W. latielliptica. 

Filaments glabrous. 
Leaves 4-5 cm. wide. 
Leaves hirsutulous beneath over the whole surface. 

W. latifolia. 
Leaves glabrous or nearly so. 

Leaves coarsely serrate W. Cuatrecasasii. 

Leaves crenate W. laurina. 

Leaves mostly less than 3.5 cm. wide. 

Leaves at least minutely cordate at the base, or rarely 
only broadly rounded. 

Teeth of the leaves incurved, less than 1 mm. deep. 

Capsule 5 mm. long W. crassifolia. 

Capsule to 3.5 mm. long W. sessilifolia. 

Teeth of the leaves coarse, spreading, 1 mm. deep. 

W. heterophylla. 
Leaves acute to cuneate at the base. 

Leaves subentire; flowers minute, the stamens 1 mm. long. 

W. elattantha. 
Leaves more or less crenate-serrate; stamens 2-4 mm. 

long. 

Pubescence coarse on the leaf nerves beneath, shaggy. 

W. nebularum. 
Pubescence fine, pilose, or nearly wanting. 

Leaves simple, usually 4-8 cm. long. 
Leaves broadly rounded at the apex. 



1046 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Veins conspicuously reticulate-veined above; 

stamens 3 mm. long W. ovalis. 

Veins obscure above; stamens 2 mm. long. 

W. elliptica. 
Leaves shortly acute at the apex. 

Petioles rarely as much as 2 mm. long. 

Leaves 25 mm. wide or wider W. ovata. 

Leaves 15-18 mm. wide W. Balbisiana. 

Petioles 5-6 mm. long W. laurina. 

Leaves usually, at least some of them, auricled or 

pinnate, rarely if ever 3 cm. long. 
Pedicels rarely 2 mm. long. 

W. Dzieduszyckii, W. auriculata. 

Pedicels 3-4 mm. long W. Jelskii. 

Leaves typically pinnate or trifoliolate, with 1-many pairs of leaflets 

more or less equaling the terminal one. 

Petioles elongate, usually more or less margined, at least some 
of them 1.5-4 cm. long; leaflets often only 1-2 pairs, ample. 

Ovary pubescent W. ayavacensis. 

Ovary glabrous. 

Leaflets 2-4 pairs, or some leaves trifoliolate. 
Leaflets acute. 

Leaflets elliptic W. Spruceana, W. pentaphylla. 

Leaflets lanceolate W. dictyoneura. 

Leaflets obtuse W. obtusifolia. 

Leaflets 4-6 pairs. 

Leaflets pilose beneath, 17-25 mm. wide W. piurensis. 

Leaflets glabrate or glabrous, 10 mm. wide. . . W. sorbifolia. 
Petioles short, none if any of them much exceeding 1.5 cm.; 

leaflets often several pairs W. ternata, W. crenata. 

Leaflets all small, that is, mostly about 1 cm. long or shorter. 

Leaflets 4-8 mm. long W. microphylla. 

Leaflets about 10 mm. long W. reticulata. 

Leaflets medium in size or, if the lateral ones shorter than 

1 cm., the terminal one usually longer. 
Ovary pilose. 
Leaves trifoliolate. 



FLORA OF PERU 1047 

Petiole densely tomentose-pilose W. ternata. 

Petiole glabrate W. crenata. 

Leaves pinnate. 
Leaves finely reticulate-veined above. 

Ovary sparsely if at all pilose; leaflets typically barely 
1 cm. long W. reticulata. 

Ovary and capsule densely pilose; leaflets about 2 
cm. long W. subsessiliflora. 

Leaves sulcate and prominently nerved, even above. 

Ovary typically glabrous W. glomerata. 

Ovary densely pubescent. 

Leaflets narrower than 1 cm W. Haenkeana. 

Leaflets 1.5-2 cm. wide W. chryseis. 

Ovary glabrous. 
Leaflets 5-17. 

Pedicels obsolete, or finally 1-2 mm. long. 
Leaves more or less pubescent beneath. 
Leaflets rarely 2 cm. long. 
Leaflets sulcate-nerved above, 1.5-3 cm. long. 

W. glomerata. 

Leaflets finely reticulate- veined, typically smaller. 

W. reticulata. 
Leaflets 2.5-3.5 cm. long W. descendens. 

Leaves glabrous, the terminal leaflet elongate. 

Leaflets serrate W. producta. 

Leaflets crenate W. crenata. 

Pedicels obvious, becoming 2.5-6 mm. long. 
Leaflets glabrous or nearly so, cuneate at the base. 
Terminal leaflet elongate; pedicels 3-6 mm. long. 

Leaflets crenate , W. apurimacensis. 

Leaflets deeply serrate W. Weberbaueri. 

Terminal leaflet little if at all elongate; pedicels 2-2.5 
mm. long. 

Leaflets mostly 1-2 cm. long W. glabra. 

Leaflets mostly 2-3 cm. long W. Ulei. 



1048 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaflets pubescent beneath, rounded to acutish at the 

base (cf. W. Weberbaueri). 
Pedicels 3-4 mm. long. 

Pedicels densely hirsute W. lentiscifolia. 

Pedicels glabrous or puberulent. 

Stamens about 2 mm. long W. cinerea. 

Stamens at least 3 mm. long W. hirtella. 

Pedicels shorter than 2.5 mm., even in fruit. 

W. reticulata. 

Leaflets 11-25; at least some of the leaves with 8-12 pairs 
of leaflets W. polyphylla. 

Weinmannia apurimacensis 0. C. Schmidt, Repert. Sp. Nov. 
29: 31. 1931. 

Younger branches densely leafy, angled, short-pilose; leaves 7-11- 
foliolate, to 7 cm. long, the petiole 7-12 mm. long, short-pilose; 
leaflets, except for the lanceolate terminal one (this to 4 cm. long), 
oblong-elliptic, more or less cuneate at the base, obtuse, 2-2.5 cm. 
long, 9-11 mm. wide, glabrous, papyraceous, subcrenate or crenate- 
serrate, obviously venose, especially beneath; rachis wings oblanceo- 
late; racemes to 12 cm. long, puberulent, the sparsely pilose, slender 
pedicels 3-6 mm. long; sepals 1 mm. long, the petals about 2.5 mm. 
long, slightly exceeded by the glabrous filaments; ovary glabrous. 
Compared by the author with W. guanacasana of Colombia, with 
pubescent leaflets of different shape. Neg. 4112. 

Apurimac: Quebrada de Toraya, Prov. Aimarcas, Raimondi, 
type. 

Weinmannia auriculata D. Don, Edinb. New Phil. Journ. 9: 
87. 1830; 164. W. ovata R. & P. Fl. Peruv. 4: pi. 333. 1802, non Cav., 
1801. W. dryadifolia Moric. ex Ser. in DC. Prodr. 4: 9. 1830. 

A densely branched shrub or tree, the younger branches and 
rachis densely hispidulous-tomentose with short, fulvous hairs; leaves 
coriaceous, sparsely pilose or glabrate above, beneath silky-pilose, 
at least on the midrib, short-petiolate (petioles 2-3, rarely 4 mm., 
long), crenulate or crenulate-serrate, oblong-ovate, sometimes deeply 
pinnatifid or exceptionally with 1 pair of lateral leaflets, when pin- 
nate the lateral lobes rounded-ovate, 3-4 times smaller than the 
ovate-spatulate middle lobe; racemes usually 2-3 (-7) cm. long; calyx 
short-pubescent, the lobes ovate-lanceolate; pedicels slightly pubes- 
cent, even in fruit, then only 2 mm. long. Material at Madrid, 



FLORA OF PERU 1049 

Paris, and Geneva is divisible into two forms: in one the leaflets are 
mostly less than 1 cm. wide, 1.5-2 cm. long; in the other to 2.5 cm. 
wide, 3 cm. long. The former form has leaflets a little more acutely 
denticulate; in the latter (in flower) the leaflets are more crenately 
dentate and more pubescent beneath. The first is f . pillavensis Pamp. 
Annali Bot. 2: 64. 1904, and is the plant of Moricand. Better marked 
is var. glabra Macbr., var. nov., ubique glabra vel racemis minute 
puberulentis. Grisar, type, Peru or Ecuador. Neg. 8039. 
Huanuco: Pillao, Ruiz & Pawn. Ecuador. "Machi." 

Weinmannia ayavacensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 772. 1929. 

Young branchlets, leaf nerves beneath, and racemes pilose; peti- 
oles 15-25 mm. long; leaves 5-9-foliolate, dull; leaflets obovate to 
elliptic-lanceolate, 2.5-4 cm. long, 1-1.5 cm. wide (the terminal 
larger), finely veined, remotely serrate; rachis wings 3 mm. wide; 
racemes to 13 cm. long, the flower fascicles dense; pedicels 2-3 mm. 
long, the calyx half as long; sepals somewhat villous; petals 1.5 mm. 
long, the stamens little longer; style 1 mm. long; ovary sub villous. 
Compared by the author with W. hirtella and W. Spruceana, with 
lustrous leaflets, those of the latter species often larger; both species 
with longer stamens. A shrub 3 meters tall (Weberbauer). 

Piura: Among evergreens, 2,700 meters (Weberbauer 6393, type). 

Weinmannia Balbisiana HBK. Nov. Gen. & Sp. 6: 51. pi. 520. 
1823. 

Suggestive of W. ovata but, at least as to the type, the leaflets 
are mostly 15-18 mm. wide, 2-5 cm. long, sparsely pilose on the 
midrib beneath, crenate-serrulate, very shortly acute at the base, 
acute at the apex, subsessile, the petioles barely 1.5 mm. long or 
shorter; stipules 5-10 mm. long, cordate-based; racemes 4-5 cm. long 
in fruit, the rachis pilose, glabrate in age, the pedicels then 2 mm. 
long; sepals 1.5 mm. long; style shorter than the capsule, this about 
3 mm. long. The nodose, verruculose branchlets are compressed 
toward the tips. 

Peru: Reported by Engler. Ecuador; Colombia. 

Weinmannia chryseis Diels, Bot. Jahrb. 37: 414. 1906. 

Young branchlets rusty-hispid-tomentose; leaves coriaceous, gla- 
brous and lustrous above, pilose-tomentulose beneath, 7-15-foliolate, 
the rachis wings obovate, about 5 mm. wide, 1 cm. long or longer; 
leaflets subequal, marginally recurved, somewhat bullate-nervose 



1050 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

above, rotund-elliptic, crenate-serrate, 2-3 (-4) cm. long, 1.5-2 cm. 
wide; racemes 15-25 cm. long, composed of remote, globose glom- 
erules; rachis and pedicels rusty- tomentose, the pedicels 2 mm. 
long; sepals narrowly lanceolate, triangular, 1.5 mm. long; petals 
2.5 mm. long, the stamens twice as long; ovary densely hirsute. 
The leaf pubescence of the dried specimen is orange brown. A shrub 
of 2 meters. W. pubescens HBK., of more northern range, has leaflets 
2-3 times larger. Neg. 27380. 

Amazonas: Chachapoyas, in sparsely shrubby grassland, 2,800 
meters, Weberbauer 4418, type. 

Weinmannia cinerea R. & P. Fl. Peruv. 4: pi. 332. 1802; 167; 
D. Don, Edinb. New Phil. Journ. 9: 90. 1830. W. diversifolia Moric. 
ex Ser. in DC. Prodr. 4: 10. 1830. W. sambucina Ruiz ex Don, loc. 
cit. fide Pamp. Annali Bot. 2: 72. 1904. 

Young branchlets compressed, sparsely puberulent, the older ones 
glabrous, cinereous; leaves coriaceous, at maturity glabrous or the 
midrib beneath sometimes sparsely hispidulous with spreading hairs; 
rachis wings semiobovate, 3-4 mm. wide; leaflets 2-4 pairs, oblong- 
elliptic, scarcely acute, crenate-serrate, 17-20 mm. long, 10-13 mm. 
wide; racemes to 7 cm. long, the rachis yellowish-hispidulous, nar- 
rowly winged between the fascicles of glabrous or glabrate pedicels, 
these 3-4 mm. long; stamens about 2 mm. long; disk cup-shaped, 
thick, like the calyx, its segments subobtuse; ovary glabrous. The 
species, as remarked by Moricand, is not at all cinereous with pubes- 
cence, but the older bark is ashy gray. Neg. 4115. 

Huanuco: Pillao, Ruiz & Pavon, type. 

Weinmannia crassifolia R. & P. Fl. Peruv. 4: pi. 331. 1802; 165. 

Like W. sessili/olia and perhaps the earlier name for that species, 
but known only in fruit; leaves heavy-coriaceous, essentially gla- 
brous, subsessile, rounded at the base, subrotund, or the floral ones 
ovate, those of the branchlets 3.5 cm. wide and 4.5 cm. long or larger; 
pedicels strigillose, 2.5 mm. long; capsule 5 mm. long, 3 mm. thick; 
style nearly 5 mm. long. Neg. 8038. 

Peru: Ruiz & Pavdn, type. Without locality: Lechler 2195. 

Weinmannia crenata Presl, Rel. Haenk. 2: 51. 1831; 166. 

Young branchlets compressed, densely pubescent; leaves coria- 
ceous, lustrous above, glabrous, 3-5-foliolate, the rachis wings narrowly 
obovate, the larger terminal leaflets oblong-lanceolate, obtuse, atten- 
uate at the base, the others suboblong, obtuse, unequally acute at 



FLORA OF PERU 1051 

the base, all beautifully crenate, to 3.5 cm. long, 16 mm. wide; ra- 
cemes spikelike, densely hirsute; pedicels scarcely 1 mm. long, exceed- 
ing the setaceous bracts; calyx lobes ovate, acutish, ciliate, pubescent 
at the base, twice exceeded by the obovate petals; ovary ovate-sub- 
rotund, glabrous. The ovary may be more or less pilose, according 
to Killip and Smith. 

Huanuco: Mountains, Haenke, type. Colombia(?). 

Weinmannia Cuatrecasasii Macbr., nom. nov. W. heterophylla 
HBK. Nov. Gen. & Sp. 6: 53. pi. 522. 1823, non R. & P. 

Branchlets, sometimes the leaf nerves beneath, and racemes 
puberulent, the racemes elongate; leaves sometimes with a pair of 
auricle-like leaflets at their base, decurrent into the petiole, this to 
8 mm. long, the blades ovate, short-acuminate, coarsely and acutely 
serrate, glabrous and lustrous above, the larger ones several centi- 
meters long and about 4 cm. wide; pedicels 3-4 mm. long; flowers 
glabrous except the pubescent sepals, these conspicuously exceeded 
by the subequal petals and stamens. W. macrophylla HBK. op. cit. 
52, Ecuadorean, has much broader (6-8 cm. wide), merely crenulate 
leaves. There seems to be no evidence that either species occurs 
in Peru. 

Peru: (According to Engler). Colombia. 

Weinmannia cymbifolia Diels, Bot. Jahrb. 37: 416. 1906. 

Young branchlets, leaves beneath, and racemes tomentose; leaves 
short-petiolate, fleshy, becoming glabrescent and lustrous above, 9- 
15-foliolate, the rachis wings to 2 mm. long; leaflets subequal, the 
nerves sulcate-impressed above, narrowly elliptic, rounded or obtuse 
at the apex, crenate-serrate but appearing subentire because of the 
recurved margin, 10-18 mm. long, 4-7 mm. wide; racemes 4-6 cm. 
long; pedicels 1-2 mm. long, the narrowly ovate-lanceolate, pilose, 
apically barbate sepals 1.5-2 cm. long, not exceeded by the petals, 
the stamens twice as long; ovary pilose, with the style 3.5 mm. 
long. A shrub 3 meters high, allied to W. tomentosa L. f., but all 
parts much larger and the pubescence different (Diels). 

Amazonas: Above Balsas, 3,300 meters, Weberbauer 4287, type. 

Weinmannia descendens Diels, Bot. Jahrb. 37: 414. 1906. 

Mature branches fibrous, the young ones rusty-sericeous; leaves 
petiolate, chartaceous, lustrous and glabrous above, paler and 
sparsely pilose beneath, 7-11-foliolate, with narrow rachis wings 
10-12 mm. long and 2 mm. wide; leaflets subequal, subelliptic, 



1052 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

appressed-crenate-serrate, 2.5-3.5 cm. long, 1.2-1.4 cm. wide, the 
nerves scarcely impressed above; pedicels 0.5 mm. long; sepals 
triangular, scarcely 1 mm. long; stamens 3.5 mm. long; capsule 
glabrous, ovoid, 2-3 mm. long. A tree 12 meters high, or a shrub. 
Compared by the author with W. cinerea, but distinguished by the 
bark, the pubescence, the nervation, and the short pedicels. 

Loreto: Near Moyobamba, grassy shrub land, 800-900 meters, 
Weberbauer 4572, type. Ecuador. 

Weinmannia dictyoneura Diels, Bot. Jahrb. 37: 414. 1906. 

Glabrous except for the branchlets and minutely pilose racemes 
and rachis; leaves coriaceous, lustrous above, 3-7-foliolate, the 
narrow rachis wings 12-15 mm. long; leaflets subequal, lanceolate, 
coarsely serrate, 3.5-5.5 cm. long, 1-1.5 cm. wide, prominently 
reticulate-veined on both sides by the pale nerves and veins; racemes 
to 10 cm. long; pedicels 1.5-2 mm. long; capsule short-ovoid, to 
4 mm. long, the style much shorter. A shrub 2 meters high, 
separated by the author from W. sorbifolia on foliar differences. 
Neg. 27381. 

Puno: Among shrubs, 2,700 meters, Sandia, Weberbauer 682, type. 

Weinmannia Dzieduszyckii Szyszyl. Diss. Cl. Math. Phys. 
Acad. Cracov. 29: 218. 1894. 

Branchlets shortly and densely sericeous-pilose, in age glabrous 
and black; leaves ternate or rarely simple, coriaceous, glabrous, 
nervose on both sides, the petioles 3-4 mm. long; leaflets sessile, 
obliquely ovate, subobtuse, acute at the base, serrate above the 
middle, 12-18 mm. long, 10-12 mm. wide, the lateral ones much 
smaller, round-ovate, less serrate, 5-6 mm. long, 4-5 mm. wide; 
rachis wings very narrow, pilose; racemes sericeous-pilose; stamens 
and petals equal; capsule oblong. According to the author, related 
to W. dryadifolia. 

Cajamarca: Cutervo (Jelski 345, type). 

Weinmannia elattantha Diels, Bot. Jahrb. 37: 413. 1906. 

A divaricately branched tree, 8 meters high, with simple, 
fleshy-coriaceous, subelliptic leaves on petioles 1-4 mm. long; blades 
shortly or obsoletely crenate-serrate, minutely reticulate-veined, the 
lateral nerves scarcely prominent, 4-5.5 cm. long and half as wide; 
racemes more or less compound, the rachis and pedicels minutely 
hispidulous, the pedicels about 2 mm. long; sepals 1-1.2 mm. long; 
capsule 2-3 mm. long, the style 1-2 mm. long. Growing among 



FLORA OF PERU 1053 

shrubs and small trees. Separated by the author from W. Bal- 
bisiana HBK. on account of the much shorter petioles and smaller 
flowers and capsules. W. boliviensis R. E. Fries is similar, fide 
Engler. Neg. 4117. 

Huanuco: Near Monzon, about 2,500 meters, Weberbauer 3540, 
type. 

Weinmannia elliptica HBK. Nov. Gen. & Sp. 6: 50. 1823. W. 
ovalis R. & P. var. elliptica Pamp. Annali Bot. 2: 66. 1904. 

Perhaps, as suggested by Pampanini, only a variant of W. ovalis 
but, at least as to type, the leaves are smaller and not reticulate- 
veined above; stamens only 2-2.5 mm. long; style pubescent at the 
base, but ovary and filaments glabrous. Neg. 4118. 

Huanuco: Pillao, Ruiz & Pavdn. Ecuador. 

Weinmannia glabra L. f. Suppl. 228. 1781; 167. 

Glabrous, or the young branchlets, petioles, and racemes minutely 
puberulent; leaves membranaceous, glabrous, slightly lustrous, 
venose, short-petioled, with 3-5 pairs of unequal, oblong-obovate, 
obtuse, crenate or serrulate leaflets, with semiobovate or semirhom- 
bic rachis wings; rachis of the raceme terete or scarcely ridged or 
grooved; pedicels little longer than the flowers, minutely hispidu- 
lous; calyx glabrate, with ovate, acute lobes slightly exceeded by 
the obovate-oblong petals; capsule ovoid, glabrous. A relatively 
glabrous plant; lowest leaflets generally subrotund and much smaller 
than those of the middle pairs, these about 15 mm. long, 10-12 mm. 
wide. Illustrated, Usteri, Fl. Sao Paulo 182. 

Peru: According to Engler in Pflanzenfam. Bolivia(?); north 
to Mexico. 

Weinmannia glomerata Presl, Rel. Haenk. 2: 52. 1831. 

Young branchlets and petioles tomentose, the petioles to 
more than 1 cm. long; leaves 5-11-foliolate, glabrous above, the 
rachis wings semiobovate; leaflets (except the terminal) oblong, 
dentate toward the obtuse apex, hispid at least on the costa beneath, 
about 16 mm. long, 8 mm. wide, the slightly larger terminal one 
oblong-lanceolate, serrate, acute; racemes equaling the lower leaves, 
with 5-foliolate basal leaves; flowers in dense glomerules, sessile or 
nearly so, finally 1-1.5 mm. long; bracts ovate, acute, ciliate; calyx 
lobes ovate, acute, sparsely pilose at the apex; ovary, as to the type 
at least, glabrous. Leaves 9-13-foliolate according to Presl, 5-11- 
foliolate according to Engler. Neg. 4119. 

Huanuco: Mountains, Haenke. Chinchao, Rivero. 



1054 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Weinmannia Haenkeana Engler, Linnaea 36: 632. 1870; 170. 

Younger branchlets densely cinereous-tomentose, blackening with 
age; leaves extremely coriaceous, short-petiolate, glabrous and 
lustrous above, long-pubescent beneath, with 4-6 pairs of subequal, 
oblong leaflets, 10-15 mm. long, 4-6 mm. wide, the rachis wings 
semielliptic, sulcate medially; leaflets slightly attenuate toward the 
base, entire or indistinctly reflexed-crenate, deeply sulcate along 
the nerves; racemes equaling the leaves; pedicels short, densely con- 
gested; calyx lobes ovate- triangular, acute, ciliate; capsule 2 mm. 
long, ovate-globose, densely and shortly cinereous-pilose, the style half 
as long. Engler (in Pflanzenfam.) supplies the locality. Neg. 19320. 

Puno: Between Sandia and Chunchosmayo, Haenke, type. 

Weinmannia heterophylla R. & P. Fl. Peruv. 4: pi. 331. 1802. 
W. cordata D. Don, Edinb. New Phil. Journ. 9: 87. 1830. W. alnifolia 
Domb. in herb. W. heterophylla HBK. as interpreted by Engler, non 
HBK. W. subcordata Moric. ex Ser. in DC. Prodr. 4: 8. 1830. 

Leaves simple, sessile, subrotund, coarsely crenate, mostly 4 cm. 
long and 3 cm. wide, lustrous, reticulate-veined on both sides, 
glabrous except on the costa beneath; angled rachis and pedicels 
slightly pilose; racemes on peduncles 8-12 cm. long, dense in flower; 
pedicels mostly 2-3 mm. long, whorled; flowers glabrous; stamens 
at least 2 mm. long; young fruits and styles together 4 mm. long, 
subequal in length. Leaves crowded; well marked by their coarse, 
spreading crenations. Neg. 29331. 

Huanuco: Huasa-huasi, Dombey; Ruiz & Pawn. 

Weinmannia hirtella HBK. Nov. Gen. & Sp. 6: 56. 1823; 168. 

A dense shrub with small, 9-15-foliolate leaves, the petioles 6-12 
mm. long, the leaflets thick-coriaceous, oblong-elliptic, glabrous and 
lustrous above, more or less pilose on the midnerve or nerves beneath, 
slightly crenate-serrate, 12-15 mm. long, at least half as wide; pedi- 
cels solitary or 2-3 together, puberulent; calyx glabrous, the stamens 
3-4 times longer, at least 3 mm. long; ovary glabrous; style 2 mm. 
long. Leaflets to 3 cm. long (Engler). W. sulcata Engler, Ecuador- 
ean, has as many as 8 or 9 pairs of somewhat larger leaflets with few 
or no rachis wings. 

Peru: Probably. Ecuador; Colombia. 

Weinmannia Jelskii Szyszyl. Diss. Cl. Math. Phys. Acad. 
Cracov. 29: 219. 1894. 

Younger branchlets pilose, compressed-trigonous, the older ones 
glabrous, terete; leaves simple, subsessile, trapezoid or oval, obtuse, 



FLORA OF PERU 1055 

cuneate at the base, serrate above the middle, coriaceous, glabrous 
except on the midrib beneath, 12-25 mm. long, 11-16 mm. wide; 
racemes pilose; pedicels 3-4 mm. long, the glabrous capsules about 
as long, the styles half as long; seeds winged. The author compared 
his species with W. guyanensis. 

Cajamarca: Cutervo (Jelski 309, type). 

Weinmannia latielliptica 0. C. Schmidt, Repert. Sp. Nov. 29: 
31. 1931. 

Branches subverruculose, lenticellate, the somewhat angled 
younger ones slightly pubescent; leaves simple, broadly elliptic or 
rarely ovate, 5-5.5 cm. long, 3.5-4.5 cm. wide, shortly narrowed to 
the petiole (this about 2 mm. long), slightly mucronate-serrate, 
coriaceous, prominently veined and rather lustrous on both sides; 
racemes to 10 cm. long, the rachis and pedicels pilose, the pedicels 
3-5 mm. long; sepals 1 mm. long, the petals twice as long, exceeded 
by the short-pilose filaments; ovary glabrous. Not clearly distinct 
from W. ovalis unless by the pilose filaments, but these are not quite 
glabrous in the type of Ruiz and Pavon's species. Neg. 4126. 

Amazonas: Entrada al Valle de Huayabamba, Raimondi, type. 

Weinmannia latifolia Presl, Rel. Haenk. 2: 51. 1831; 164. 

Youngest branchlets, leaves beneath, and racemes hirsute; leaves 
coriaceous, rounded-ovate, to 7.5 cm. long, 5 cm. wide, strongly 
net-veined, especially beneath, equally crenate-serrate, glabrate 
above; petioles 2 mm. long; stipules 6 mm. long; racemes many- 
flowered, with a few serrulate, linear-lanceolate basal leaves; pedi- 
cels 2.5-4 mm. long; calyx short-pilose, with ovate-triangular, 
subobtuse, sometimes glabrous lobes; petals 5 mm. long; stamens 
glabrous, 5 mm. long; young fruit sparsely hispidulous toward the 
apex, equaled by the glabrous styles, these 2 mm. long. The floral 
measurements are from Hartweg 743, of Ecuador, type of W. rugosa 
Benth., referred here by Engler; the young capsules are described by 
Presl as glabrous. Compare W. Cuatrecasasii and the similar W. 
Bangii Rusby, Pflanzenfam. 251, Bolivian. Neg. 4127. 

Huanuco: Mountains, Haenke. Ecuador. 

Weinmannia laurina HBK. Nov. Gen. & Sp. 6: 51. 1823. 

Apparently scarcely distinct from W. ovata but, as to type, the 
leaves elliptic, not at all obovate, tapering to the petiole, this 5-6 
mm. long, acute, 4 cm. wide or wider, 8 cm. long, sometimes larger, 
crenate, coriaceous, reticulate-veined on both sides, glabrous except 



1056 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

for a few hairs on the midnerve beneath; peduncles and rachis pilose, 
the rachis terete; racemes about 5 cm. long; pedicels in fruit puber- 
ulent-pilose, 2-3 mm. long; capsules nearly oblong, 4 mm. long, the 
styles scarcely half as long. Type locality unknown, but probably 
Colombian; Linden 717 is a good match for the type, better than 
925 cited by Engler, which has much longer pedicels. 
Peru: Probably. Venezuela(?); Colombia(?). 

Weinmannia Lechleriana Engler, Linnaea 36: 630. 1870; 170. 

A shrub with straggling or ascending branches, the branchlets 
densely and shortly hispidulous-tomentose; leaves with 3-5 pairs of 
lightly sulcate, oblong-elliptic, obtuse, serrulate leaflets, these gla- 
brous above, more or less long-hirsute-pilose beneath, especially on 
the costa and rachis, mostly 12 (10-20) mm. long, 8 mm. wide, 
reticulate-veined, dull, even above; rachis wings obovate, 4 mm. 
wide; racemes 4-7 cm. long in fruit, the rachis and pedicels hispid- 
ulous, the pedicels slender, flexuous, nearly 5 mm. long in fruit; 
flowers minute; stamens glabrous, 1.2 mm. long; capsule pilose, 
nearly 2 mm. long, the glabrous style about as long. Neg. 8040. 

Puno: Sachapata, Lechler 2661, type. 

Weinmannia lentiscifolia Presl, Rel. Haenk. 2: 52. 1831; 167. 

Compressed young branchlets hirsute with spreading, yellowish 
hairs; petioles 8 mm. long; leaves 7-11-foliolate, coriaceous, reticu- 
late-veined on both sides, glabrous above, hirsute beneath, the 
rachis wings obovate; leaflets oblong, except the terminal one (this 
oblong-lanceolate, acute), acutish at the base, obtuse at the apex, 
callous-dentate, to 18 mm. long, 12 mm. wide; racemes geminate, 
terminal, equaling the leaves, spikelike; pedicels to 4 mm. long, 
densely hirsute; bracts setaceous; calyx lobes ovate, acute, ciliate; 
ovary ovate, glabrous. 

Huanuco: Mountains, Haenke, type. 

Weinmannia microphylla R. & P. Fl. Peruv. 4: pi. 334- 1802. 
W. fagaroides HBK. Nov. Gen. & Sp. 6: 54. pi. 524. 1823; 168. W. 
parvifolia Ruiz ex D. Don, Edinb. New Phil. Journ. 9: 89. 1830. 
W. microphylla R. & P. var. parvifolia Pamp. Annali Bot. 2: 79. 
1904. W. Baccariniana Pamp. op. cit. 81. 

A densely branched, densely leafy shrub or tree, the younger 
branchlets nearly tomentose with a fine, short pubescence; leaves 
1.5-2.5 cm. long, with mostly 2-3 pairs of leaflets below the terminal 
3; rachis often hispid, wing-margined; leaflets obscurely veined, lus- 



FLORA OF PERU 1057 

trous, obscurely 1-4-crenate-serrulate at or toward the apex, obovate, 
4-5 mm. wide, 5-6 mm. long; racemes densely flowered, only in fruit 
exceeding the leaves; filaments glabrous, 2 mm. long; rachis and pedi- 
cels minutely pilose, the pedicels 1.5-2 mm. long, in fruit shorter than 
the ovoid capsules, these longer than the glabrous style. W. Bac- 
cariniana is a form with 4 pairs of leaflets, these 7 mm. long, with 7 
crenations. W. microphylla R. & P. var. tenuior (Diels) Macbr., 
comb. nov. (W. parvifolia Don, var. tenuior Diels, Bot. Jahrb. 37: 416. 
1906), has more slender leaflets than the typical form, the margins 
incurved. A more distinct variation is var. Weddellii Macbr., var. 
nov., foliis 5-jugis opacis; foliolis 7 mm. longis; racemis ad 8 cm. 
longis. This passes to W. hirta Sw.; cf. Engler, Pflanzenfam. ed. 2. 
18a: 254. W. microphylla HBK. Nov. Gen. & Sp. 6: 54. pi 523. 1823, 
from Loja, Ecuador, has lustrous leaves consisting of only 3 leaflets, 
at least in the type; cf. Engler, Pflanzenfam. ed. 2. 18a: 252. A Will- 
denow name, published in synonymy, is available for it; cf. Pamp. 
Annali Bot. 2: 70. 1904. Negs. 8043, 34636. There is a photograph 
of this tree in Bull. Torrey Club 56: 369. 1929. 

Huanuco: Pillao, Ruiz & Pavdn. Junin: Tarma, 2,800 meters, 
Weberbauer 2282, 2479 (var. tenuior). Amazonas: Chachapoyas, 
Mathews (type of W. Baccariniana). Puno: Carabaya, Weddell 
4637 (type of var. Weddellii). Without locality: Gay. Bolivia to 
Colombia. "Machi." 

Weinmannia nebularum Diels, Bot. Jahrb. 37: 413. 1906. 

An arborescent shrub, the young branches cinereous-pubescent; 
petioles 5-8 mm. long; leaves simple, chartaceous, glabrate above, 
at first sericeous-tomentose, later sparsely pilose beneath, elliptic, 
5-6 cm. long, 2-3 cm. wide, conspicuously serrate, the lateral nerves 
bullate-impressed above, prominent beneath; peduncles 12-15 mm. 
long; racemes geminate, much exceeding the leaves, 12-15 cm. long, 
minutely pilose; pedicels 3-4 mm. long; sepals 1.5-2 mm. long, the 
petals slightly longer. To 8 meters high, in thickets and open woods. 
Distinguished by the author from W. cordata Don by the petiolate, 
acute-based leaves. Neg. 27385. 

Cajamarca: Chugur, 3,000 meters, Weberbauer 4093, type. 
Junin: Huacapistana, 2,700 meters, Weberbauer 2274, 2320. "Shalle." 

Weinmannia obtusifolia Rusby, Bull. N. Y. Bot. Gard. 6: 
508. 1910. 

Younger parts puberulent; petioles 12-25 mm. long; leaflets 3-5, 
the lateral ones 3^4 cm. long, 1-2 cm. wide, oblong-ovate, obtuse, 



1058 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

broadly cuneate at the base, obtuse-serrate, somewhat lustrous, 
reticulate-veined; racemes 7-10 cm. long, the fascicles dense; pedicels 
1-1.5 mm. long; sepals 1 mm. long, the stamens 3 times as long. 
"Near W. diversifolia" (Rusby), i.e. W. cinerea. 

Peru: Without locality (Mathews, fide Rusby). Cuzco: Rio 
Chaupimayo, Soukup 794- Bolivia. 

Weinmannia ovalis R. & P. Fl. Peruv. 4: pi. 333. 1802; 160; D. 
Don. Edinb. New Phil. Journ. 9: 86. 1830. W. elliptica HBK. Nov. 
Gen. & Sp. 6: 50. 1823. 

Glabrous, even to the youngest leaves, except for a sparse, crisped 
strigillosity on the short peduncles, rachis, and pedicels; petioles 1-3 
mm. long; leaves coriaceous, conspicuously and finely reticulate- 
veined on both sides, ovate-elliptic or somewhat obovate, shortly 
acute at the base, rounded and obtuse or slightly narrowed and acut- 
ish at the apex, mostly 5 cm. long, 3 cm. wide, or the larger ones 8 
cm. long, 5 cm. wide, crenate-serrulate, the costa strongly developed 
beneath; racemes about 7 cm. long; pedicels in fruit 2-4 mm. long, 
mostly in fascicles of 3-5; stamens nearly 3 mm. long; styles glabrous, 
2 mm. long, about equaling the narrowly ovoid capsule. A slender 
tree, 7 meters high (Killip & Smith). Neg. 8044. 

Huanuco: Pillao, 2,000 meters, Ruiz & Pawn, type. Without 
locality, Mathews 1203. Junin: Huacapistana, Killip & Smith 24257. 
North to Ecuador and the Guianas. 

Weinmannia ovata Cav. Icon. 6: 45. pi. 566. 1801; 165. 

Branchlets sparsely pilose; leaves rigid, reticulate- veined and 
glabrous on both sides, petioled (petioles 2 mm. long), oblong-elliptic, 
7.5 cm. long, 2.5 cm. wide or wider, acute at each end, crenate-serrate; 
racemes with lanceolate basal leaves, elongate; bracts ovate, obtuse; 
pedicels minutely puberulent, 1-2 mm. long; calyx short-pilose, the 
lobes oblong-ovate, obtuse; capsule oblong. A tree 6 meters tall. 
Neg. 4129. 

Lima: San Buenaventura, growing with Cantua flexuosa, etc., 
Nee, type. Cuzco: (Pennell 14099 [?]; cf. Killip & Smith, Bull. 
Torrey Club 56: 366. 1929). 

Weinmannia pentaphylla R. & P. Fl. Peruv. 4: pi. 330. 1802. 
W. marginata Moric. ex Ser. in DC. Prodr. 4: 10. 1830. W. bifida 
Poepp. ex Engler, Linnaea 36: 608. 1870. 

Branchlets soon glabrous, strongly compressed and nodose; leaves 
typically, even the uppermost, with 1-2 pairs of leaflets below the 



FLORA OF PERU 1059 

terminal 3, but some branchlets bearing only ternate but long- 
petiolate leaves; petioles more or less margined or winged; leaflets 
subequal, oblique and narrowed at the base (the terminal cuneate 
at the base and somewhat larger), more or less acuminate or acute, 
crenate-serrate or in age serrate, lustrous and reticulate-veined on 
both sides, glabrous or slightly pilose on the costa beneath, oblong 
or ovate-lanceolate, finally becoming 9 cm. long, 3.5 cm. wide, mostly 
somewhat shorter and about 3 cm. wide; stipules glabrous, fugacious; 
peduncles, rachis, and pedicels more or less densely strigillose-pilose; 
pedicels in fruit 4 mm. long, the fascicles whorled ; rachis ridged but 
sub terete; racemes terminal, geminate, the peduncles 1.5 cm. long, 
the racemes about 10 cm. long; flowers glabrous; sepals minute; sta- 
mens 2.5 mm. long; capsule narrowly ovoid, glabrous, with the style 
4 mm. long. The original label of Poeppig reads "Peruvia suban- 
dina," but there are mislabeled specimens in herbaria. Negs. 8041, 
4113, 29332. 

Huanuco: Cochero, Dombey; Ruiz & Pavon. Without locality, 
River o; Poeppig 1273 (type of W. bifida). 

Weinmannia piurensis 0. C. Schmidt, Repert. Sp. Nov. 29: 
30. 1931. 

Petioles of the 9-13-foliolate leaves 1.5-2.5 cm. long, short-pilose; 
leaflets broadly lanceolate, 3-6 cm. long, 17-25 mm. wide, narrowed 
to the rounded base and the acute apex, soon glabrous above, pilose 
on the prominent nerves beneath, mucronate-serrate; rachis wings 
6-9 mm. wide; racemes densely flowered, to 13 cm. long, the rachis 
villous; pedicels 1.5-3 mm. long, densely pilose; petals 2 mm. long, 
twice as long as the sepals, subequaling the glabrous filaments; ovary- 
glabrous. Related to W. Spruceana, with longer stamens. A small 
tree (Raimondi). Neg. 4130. 

Piura: Near Anchalai, Raimondi, type. 

Weinmannia polyphylla Moric. ex DC. Prodr. 4: 11. 1830. 

Branches little compressed, soon glabrate, at first, like the short 
petioles, coarsely and densely hispid; leaves 17-25-foliolate, about 10 
cm. long; leaflets subequal (except the middle one of the terminal 3, 
it more or less elongate), about 15 mm. long, half as wide, nearly ob- 
long, minutely serrate toward the rounded apex, lustrous but sparsely 
hispidulous on the obvious nerves above, or glabrate, glabrous 
beneath except for the hispid costa and sometimes also the reticulate 
veins; rachis wings rarely 4 mm. wide; peduncles several centimeters 
long, hispidulous like the rachis, this 6-10 cm. long; flowers scattered 



1060 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or slightly clustered, the divaricate pedicels minutely pilose, soon 5-7 
mm. long; sepals glabrous except for a tuft of hairs at the apex, 1 mm. 
long; styles and glabrous ovary subequal. Type from "Guayaquil." 
Neg. 8046. 

Peru: Probably. Ecuador. 

Weinmannia producta Moric. ex DC. Prodr. 4: 11. 1830. 

Branchlets crisp-puberulent, soon glabrate; leaves with 3-5 pairs 
of leaflets below the terminal 3, these very unequal, the middle one 
as much as 3 cm. long; lateral leaflets progressively reduced toward 
the short petiole, the lowest ones subrotund, only 5 mm. long, the 
middle ones 10-15 mm. long, about 6 mm. wide, oblong-elliptic, 
serrate toward the rounded apex, lustrous, glabrous and venose on 
both sides; rachis wings 2-3 mm. wide; racemes dense, the glomerules 
contiguous, short-peduncled, 7-10 cm. long, exceeding the leaves; 
angled rachis and pedicels sparsely pilose or glabrate, the pedicels 
in flower 1.5 mm. long; sepals glabrous except at the apex; stamens 
and style 2 mm. long, exceeding the conspicuous, elliptic petals; 
ovary glabrous. With "W. sessiliflora" in Herb. Moric., now Herb. 
Delessert; W. subsessiflora is meant, and presumably the locality is 
the same. Negs. 34054, 34637. 

Peru: Ruiz & Pavon, type; probably from Pillao. 

Weinmannia reticulata R. & P. Fl. Peruv. 4: pi. 332. 1802; 
Pav. ex Ser. in DC. Prodr. 4: 10. 1830; 169. 

A compact shrub with erect or ascending branches, the branch- 
lets shortly rusty-pilose toward the tips; leaves about 4 cm. long, 
with 3-5 pairs of crenate, oblong-elliptic, obtuse leaflets, these mostly 
10 mm. long, 6 mm. wide, slightly pilose on both sides or soon gla- 
brate, usually hispid beneath on the costa, lustrous above; rachis 
wings obovate, 3 mm. wide; racemes dense, 4-5 cm. long, rusty-pilose; 
pedicels only 2 mm. long in fruit; stamens scarcely 2 mm. long; sepals 
ciliate; capsule glabrous, equaled by the styles. Neg. 4132. 

Huanuco: Pillao and Acomayo, Ruiz & Pavdn, type. Without 
locality: Ruiz & Pavdn 12-5. 

Weinmannia sessilifolia Rusby, Bull. N. Y. Bot. Gard. 6: 
509. 1910. 

Similar to W. ovalis, but the uppermost stipules more appressed- 
strigose; leaves subrotund or the upper oblong-elliptic, rounded at 
both ends or barely acutish at the rounded apex, sparsely long-pilose 
on the costa beneath, mostly 2.2 cm. wide and 4 cm. long, net-veined 



FLORA OF PERU 1061 

on both sides or at first obscurely so above; pedicels 1-3 or nearly 4 
mm. long in fruit, approximate; racemes 4-6 cm. long; calyx lobes 
acute, ciliate; capsule broadly ovoid, 3 mm. long, nearly as broad, 
the style scarcely 2.5 mm. long, sparsely pubescent at the base. 
According to Pampanini, this is the same as W. crassifolia. 

Puno : Tatanara, Lechler 21 95. Bolivia. 

Weinmannia sorbifolia HBK. Nov. Gen. & Sp. 6: 57. 1823. 

A tree with leaves, as to type, having 3-4 pairs of leaflets below the 
terminal 3, these about 3 cm. long, 1 cm. wide, suboblong, glabrous, 
acute, acutely serrate; petioles 1.5-3 cm. long; racemes to 10 cm. 
long; pedicels 2.5 mm. long; flowers minute, the stamens scarcely 2 
mm. long; style and capsule glabrous, subequal. In Peru, according 
to Engler, but unless the species is more variable than seems probable, 
his specimens belong elsewhere. Killip & Smith, Bull. Torrey Club, 
56: 366. 1929, suggest that the species is W. crenata; the type locality 
is unknown. Neg. 4134. 

Peru: Perhaps. Colombia(?). 

Weinmannia Spruceana Engler in Mart. Fl. Bras. 14, pt. 2. 
166. 1871. W. bifida Poepp. f. alata Pamp. Annali Bot. 2: 71. 1904. 

Branchlets glabrate, somewhat quadrangular; leaves glabrous 
and lustrous above, with narrowly winged petioles, the rachis wings 
semi-oblong-cuneate; leaflets 2-3 pairs, the terminal one ovate, acute, 
sinuate, attenuate to the base, the others oblong, subobtuse at both 
ends, to 7.5 cm. long, crenate-serrate; racemes slightly longer than 
the leaves, pilose with spreading hairs, the narrow basal leaves 5-f olio- 
late; pedicels longer than the flowers; bracts obovate, glabrous like 
the calyx, the ovate calyx segments acute; petals oblong; ovary gla- 
brous; stamens 3 mm. long; style 2 mm. long. Neg. 4136. 

Amazonas: Chachapoyas, Mathews 3035. Ecuador. 

Weinmannia subsessiliflora R. & P. Fl. Peruv. 4: pi. 33^ 1802; 
D. Don, Edinb. New Phil. Journ. 9: 90. 1830; 166. 

Branches marked with white lenticels, the compressed youngest 
ones as well as the petioles, peduncles, and angled raceme rachis 
puberulent-pilose; leaves mostly with 4 pairs of oblong-elliptic, crenu- 
late-serrulate leaflets, these 12-22 mm. long, about 8 mm. wide, the 
terminal one somewhat larger, glabrous on both sides except on the 
costa, finely reticulate-veined; rachis wings obovate-rhombic, 4-6 
mm. wide; racemes 6-10 cm. long; pedicels in fruit only 1 mm. long; 



1062 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sepals puberulent, minute; stamens glabrous, scarcely 2 mm. 
long; capsule broadly ovoid, puberulent, 2 mm. long, the style less 
than half as long. Neg. 27387. 

Huanuco: Pillao, Ruiz & Pavdn, type. Lima: Cheuchin, Dombey. 

Weinmannia ternata Engler in Mart. Fl. Bras. 14, pt. 2: 
165. 1871. 

Branchlets, petioles, and rachis of the racemes shortly tomentose- 
pilose; leaves ternate, single, or geminate, subequal, lustrous above, 
paler beneath, with a few long hairs on the costa beneath or slightly 
puberulent on the costa above, elegantly reticulate-veined on both 
sides, about 2.5 cm. long, 1 cm. wide, oblong-ovate, crenate at the 
base, acutish, crenulate; racemes 4 cm. long, densely flowered, 
appearing spicate; sepals minute, hispidulous; stamens glabrous, 1.5 
mm. long; ovary densely pubescent, longer than the glabrous style. 
W. microphylla R. & P., with small, sessile leaves, if sought here may 
be found under W. fagarioides. Neg. 4138. 

San Martin: Tarapoto, Spruce 4238, type. 

Weinmannia trichocarpa Pamp. Annali Bot. 2: 67. 1904. 

Branches and branchlets slender, the younger ones brownish- 
puberulent; leaves simple, sessile, oblong-elliptic, short-cuneate at 
the base, remotely crenate, coriaceous, glabrous and prominently 
reticulate-veined on both sides, 2.5-4.5 cm. long, 1.5-2 cm. wide, the 
upper ones smaller; racemes 6-8 cm. long; pedicels appressed-puberu- 
lent, about as long as the flowers; calyx pilose only at the apex; sta- 
mens twice as long as the petals; ovary densely pilose. Neg. 8049. 

Cajamarca: Chota and montafia de Nancho, Raimondi (det. 
Schmidt). Amazonas: Chachapoyas, Mathews 3034, type. 

Weinmannia Ulei Diels, Bot. Jahrb. 37: 415. 1906. 

Younger branches minutely hirsutulous; leaves subsessile, char- 
taceous, somewhat lustrous above, dull and paler beneath, glabrous 
except for a few long hairs on the nerves, 3-15-foliolate, the rachis 
wings 7 mm. wide; lower leaflets progressively smaller, the ter- 
minal one slightly larger, all elliptic or obovate-elliptic, crenate- 
serrate, 2-3 cm. long, 8-14 mm. wide; racemes slender, elongate, 
10-25 cm. long, the minutely pilose pedicels glabrate, 2.5 mm. long; 
sepals glabrous, subovate, 1 mm. long, the stamens about 3 times as 
long; capsule glabrous. A shrub or tree 2-8 meters high, separated 
by the author from W. hirta Swartz. Neg. 4139. 

Loreto: Cerro de Escalera, 1,300 meters, Ule 6752, type. 



FLORA OF PERU 1063 

Weinmannia Weberbaueri Diels, Bot. Jahrb. 37: 415. 1906. 

Blackish purple branches glabrate, or the younger ones sparsely 
puberulent; leaves short-petiolate, thin-chartaceous, minutely squam- 
ulose beneath, otherwise glabrous, 11-17-foliolate, the rachis wings 
1-1.5 mm. wide; lateral leaflets shorter than the terminal one, obo- 
vate-elliptic, acute at the base, deeply serrate, 13-18 mm. long, 5-8 
mm. wide; racemes axillary, with the peduncle 5-15 cm. long, 
sericeous-pilose, the puberulent pedicels 3-5 mm. long, the sepals 
half as long, slightly exceeded by the rose-suffused petals; capsule 
glabrous, 3-4 mm. long. A shrub 2 meters high, distinct from 
W. microphylla R. & P. in the larger, coarsely serrate leaflets. 

Ancash: In thickets, 3,200 meters, Huaraz, Weberbauer 3240, 
type. Prov. Cajatambo, Weberbauer 2879. "Perejil." 

86. ROSACEAE. Rose Family 

Although roses are not native in Peru, the family is represented 
by a number of trees and shrubs familiar to North Americans and 
Europeans, such as choke cherries (Prunus) and blackberries (Rubus). 
Several almost cosmopolitan genera are meagerly, if at all, repre- 
sented, like the strawberry (Fragaria), cinquefoil (Potentilla), and 
hawthorn (Crataegus) . There are a number of conspicuous and inter- 
esting plants of the family, notably the beautiful quinuares (Poly- 
lepis), several of which are of economic value, and the curious and 
useful lloque (Kageneckia). In cultivation to a greater or less 
extent are such widely known fruits as the goma de pais or apricot, 
Armeniaca vulgaris L.; the durazno, abidor, melocotonero, or peach, 
Persica vulgaris L.; the almendro or almond, Amygdalus communis 
L.; plums, ciruelas, Prunus domestica L.; peral or pear, Pyrus com- 
munis L.; manzano or apple, Mains communis DC.; and membrillo 
or quince, Cydonia vulgaris Pers. The cultivated roses include R. 
centifolia L. and R. indica L. According to Herrera, all these species 
are found in the Department of Cuzco and most of them, at least, 
in the more central and northern inter-Andean valleys also. 

Fruit an apple-like drupe; calyx tube and ovary adnate; leaves 

simple; flowers corymbose 1. Hesperomeles. 

Fruit not apple-like, sometimes plumlike, berry-like, or dry and 

carpellate (achenes). 
Trees or treelike shrubs. 

Carpel 1; inflorescence usually many-flowered or the flowers 
large. 



1064 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers not in pendulous spikes; leaves simple. 
Style lateral from near the base of the ovary. 
Ovary 1-celled, at the base of the calyx tube; flowers 

small 2. Licania. 

Ovary laterally inserted in the calyx throat; flowers 

often rather large, or the stamens conspicuous. 
Ovary 2-celled (septum not always complete); flowers 

medium-sized 3. Parinarium. 

Ovary 1-celled; flowers conspicuous, at least by the 

long stamens. 
Stamens 3-7, the filaments conspicuous. 

4. Hirtella. 

Stamens 15 to many 5. Couepia. 

Style terminal 6. Prunus. 

Flowers in pendulous spikes; leaves compound ... 7. Polylepis- 
Carpels 5, stellately spreading; flowers few; leaves subentire or 
minutely serrulate. 

Leaves entire 8. Quillaja. 

Leaves serrulate 9. Kageneckia. 

Ligneous or entirely herbaceous, but not treelike shrubs (see 

Margyricarpus) ; carpels often many. 

Ligneous, usually prickly shrubs or vines; fruit edible . 10. Rubus. 
Herbs, usually perennial, suffrutescent, if at all, only at the base, 

except in Margyricarpus. 
Carpels many. 

Style elongate 11. Geum. 

Style not elongate. 
Receptacle more or less fleshy, strawberry-like. 

12. Fragaria. 

Receptacle dry 13. Potentilla. 

Carpels 1-3. 

Calyx accompanied by bractlets; petals none; depressed or 

low herbs 14. Alchemilla. 

Calyx without bractlets. 
Shrubs, rigid, often somewhat spinose. 

15. Margyricarpus. 
Herbs. 
Petals none; calyx prickles wanting or barbed. 



FLORA OF PERU 1065 

Calyx valvate; fruit aculeate 16. Acaena. 

Calyx imbricate; fruit not aculeate . . 17. Sanguisorba. 

1. HESPEROMELES Lindl. 
Reference: C. K. Schneider, Bot. Jahrb. 42: 85-88. 1908. 

Shrubs, often tall, sometimes depressed and gnarled, often much 
branched, always with simple, entire or more or less dentate leaves. 
Flowers cymose-corymbose; bractlets narrow. Calyx teeth 5, per- 
sistent. Disk glabrous or tomentose. Ovary cells 5, 1-ovuled. 
Fruit a small, edible drupe. An American genus, conveniently 
maintained as distinct from the habitally different and pinnate- 
leaved Osteomeles of China and Hawaii. 

Leaves cuneate-obovate, glabrous or glabrate, rarely exceeding 
3 cm. in length or 1 cm. in width. 

Leaves 10 mm. long, 3-4 (-6) mm. wide H. escalloniaefolia. 

Leaves, at least some of them, a third larger. 

Leaves mostly 10-12 mm. long, 5-7 mm. wide, definitely den- 
ticulate at the subtruncate apex H. cuneata. 

Leaves mostly 15 mm. long, 8-9 mm. wide, subentire or crenu- 

late-denticulate H. pernettyoides. 

Leaves somewhat or much larger, or at least wider, often pubescent 

beneath, often more or less rounded at the base. 
Leaves coriaceous and soon reticulate-veined, especially beneath, 

typically about 3.5 cm. wide H. lanuginosa. 

Leaves chartaceous or subcoriaceous, often little if at all net- 
veined, or smaller. 

Leaves mostly 5-6 cm. wide (see H. lanuginosa) . . . H. latifolia. 
Leaves mostly 1.5 cm. wide. 
Leaves mostly 4-6 cm. long, 2.5-3 cm. wide, more or less 

pubescent beneath H. Weberbaueri. 

Leaves mostly 2-2.5 cm. long and about 1.5 cm. wide, some- 
what pubescent beneath. 
Pubescence ferruginous; leaves irregularly dentate, shorter 

than the corymbs H. palcensis. 

Pubescence pale; leaves evenly dentate, about as long as 

the corymbs H. Gayana. 

Leaves mostly 3^1 cm. long, 1.5-2.5 cm. wide, glabrous or 
glabrate beneath H. heterophylla. 



1066 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Hesperomeles cuneata Lindl. Bot. Reg. 23: sub pi. 1956. 1837. 
Osteomeles cuneata Dene. Nouv. Arch. Mus. Paris 10: 184. 1874. 

Spinescent or with short, stiff branchlets; leaves cimeate, above 
the middle finely dentate and rarely subtrilobate, that is, with 2-3 
large, irregularly placed teeth, lustrous and soon glabrous above, 
paler, dull, and finally glabrous beneath, 1-2 cm. long, 0.5-0.7 cm. 
wide, or on older branchlets smaller; corymbs at the tips of short 
branchlets, shorter than the leaves, slightly villous; calyx lobes nar- 
rowly subulate, recurved; petals entire or the edges slightly undulate; 
disk glabrous or somewhat pubescent at the center. Neg. 27482. 

Junin: Huacapistana, 3,500 meters, Weberbauer 2242 (det. 
Schneider). Lima: Above Lima, Mathews 577, type. Huanuco: 
Pampayacu, Kanehira /0. Ayacucho: Quinua, Weberbauer 5541 
(det. Mansfeld). Chancobango, Isern 2041- San Mateo, Isern 2003. 

Hesperomeles escalloniaefolia (Schlecht.) Schneid. Bot. Jahrb. 
42: 88. 1908. Crataegus escalloniaefolia Schlecht. Linnaea 27: 468. 
1856. Osteomeles escalloniaefolia Dene. Nouv. Arch. Mus. Paris 10: 
184. 1874. H. pernettyoides Wedd. var. microphylla Wedd. Chlor. 
And. 2: 231. 1857, saltern pro parte. 

Stoutly branched, the ferruginous-puberulent young branchlets 
terminating in a spinose tip; leaves obovate or oblanceolate, with 
2-4 minute teeth above the middle on each side, rounded at the apex, 
acute at the base, crenate, 9-10 mm. long, 3-6 mm. wide, glabrous, 
lustrous above, paler, dull, and venose beneath; corymbs about 5- 
flowered, the pedicels sparsely pilose; bracts linear-lanceolate, 4-7 
mm. long; calyx glabrous, the lobes ovate, with long, acuminate, 
recurved tips, 1.5 mm. long; petals slightly crenulate; disk and 
styles densely long-pilose; stamens 3 mm. long. Description from 
Lechler material. Doubtfully distinct from H. cuneata. Neg. 27483. 

Ancash: Tallenga, 3,600 meters, Weberbauer 2881 (det. Schneider). 
Puno: Tabina, Lechler 2060, type. Cuyocuyo, 3,100 meters, 
Weberbauer 846 (det. Schneider) . Cuzco : Gay. "Toen." 

Hesperomeles Gayana (Dene.) Macbr., comb. nov. Osteomeles 
Gayana Dene. Nouv. Arch. Mus. Paris 10: 183. 1874. 

Some of the shorter branches spinose-tipped ; leaf blades 2-2.5 
cm. long, crenate or denticulate to the subtruncate apex, broadly 
elliptic-oblong or slightly obovate, acute at the base, chartaceous, 
lustrous above, pale and slightly pilose beneath; petioles 2-4 mm. 
long; inflorescence at first densely sericeous-puberulent, dense; 



FLORA OF PERU 1067 

bractlets filiform; calyx teeth subulate-acuminate; petals subentire, 
glabrous; disk villous. Perhaps only a pubescent variety of H. 
heterophylla. Neg. 34729. 

CuzcoiGay 1553, type; also 367, 372 (fruit). 

Hesperomeles heterophylla (R. & P.) Hook. Icon. 9: pi. 846. 
1852. Crataegus obtusifolia Pers. Syn. 2: 37. 1806. Mespilus Per- 
soonia Spreng. Syst. 2: 506. 1825. M. heterophylla R. & P. Fl. Peruv. 
4: pi. 4%5b. 1802. H. obtusifolia Lindl. Bot. Reg. 23: sub pi. 1956. 
1837. H. Fieldii Macbr. Field Mus. Bot. 4: 81. 1925. 

Gray-barked, the reddish, lenticellate branchlets and the corymbs, 
including the calyces, evanescently and slightly pilose, otherwise 
glabrous; petioles 5-8 mm. long; blades chartaceous, lustrous above, 
pale beneath, finely reticulate-veined on both sides, elliptic or 
oblong-elliptic, acute at the base, mostly rounded at the apex or 
slightly narrowed and subacute, the larger ones usually 3-4 cm. 
long, 1.5-2.5 cm. wide, the short flowering branchlets often with 2 or 
3 much smaller leaves at the base, all more or less obscurely crenate- 
serrate from below the middle; corymbs dense, shorter than the 
leaves; calyx tube ovate-acuminate, about 1 cm. long; petals some- 
what crenulate; disk pilose. Fruit edible (Dombey). Description 
from type material. Osteomeles glabrata HBK. seems to be a large- 
leaved variety. A depressed form is var. vacciniifolia Hook. loc. cit. 

Huanuco: Chaclla, Weberbauer 6795 (det. Mansfeld). Pillao, Ruiz. 
Junin: Huasa-huasi, Ruiz & Pavon; Dombey, type. San Carlos 
(Mathews 1445, fide Hook.). Ayacucho: Prov. Huanta, Weberbauer 
(det. Mansfeld). Cuzco: Herreral37 (det. Markgraf). Cajamarca: 
Cutervo, Jelski 278. Colombia; Ecuador. "Milucapa." 

Hesperomeles lanuginosa R. & P. ex Hook. Icon. 9: pi. 846. 
1852. Mespilus lanuginosa R. & P. Fl. Peruv. 4: pi. 425a. 1802. 
Crataegus ferruginea Pers. Syn. 2: 37. 1806. Eriobotrya cordata 
Lindl. Trans. Linn. Soc. 13: 102. 1822. Osteomeles ferruginea HBK. 
Nov. Gen. & Sp. 6: 211. 1824. H. oblonga Lindl. Bot. Reg. 23: sub 
pi. 1956. 1837. Mespilus ferruginea Poir. Encycl. Suppl. 4: 73. 1816. 
H. ferruginea Benth. PI. Hartw. 129. 1844. 

Branches spreading, stout, rather tardily glabrate, with pale 
lenticels; branchlets, young leaves beneath, and corymbs densely 
ferruginous-tomentose, the pubescence persistent or on the leaves 
tardily disappearing in part or becoming cinereous; petioles about 
1 cm. long; blades coriaceous, evanescently puberulent above, later 
lustrous, slightly rugulose by the impressed nerves and veins, these 



1068 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

soon strongly reticulate beneath; blades typically ovate or subelliptic- 
ovate, subacute or often subcaudate at the base, rounded at the apex, 
about 5 cm. long and 3.5 cm. wide; corymbs dense, shorter than the 
leaves; bractlets filiform; calyx tube ovate, acute at the base, scarcely 
1 mm. long; petals oblong-cuneate, pilose, white; stamens short. 
Description from the Dombey material in herb. Paris. Species 
variable. Mathews 888, not seen, the type of H. oblonga, is described 
as having "leaves oblong, rugose, above glabrous, beneath ferrugi- 
nous-villous; cymes lanate, shorter than the leaves." Petals in some 
forms are glabrous, as in the doubtfully distinct 0. rubescens Dene., 
of Colombia, with larger flowers. A form from Colombia with more 
oblong leaves and more grayish pubescence may be distinct. Some- 
times a tree of 5 meters. Petals sometimes eciliate (Weberbauer 4023) . 
Junin: Palca, 3,200 meters, Weberbauer 2495 (det. Schneider). 
Carpapata, Killip & Smith 24454- Huasa-huasi, Dombey, type. 
Ayacucho: Pampalca, Killip & Smith 23257. Quinua, 3,300 meters, 
Weberbauer 5546 (det. Mansfeld). Pillao, Ruiz. Huanuco: Monzon, 
3,300 meters, Weberbauer 3374 (det. Schneider). Pampayacu, 
Kanehira 27; Sawada P22. Cani, 3409. Mito, 1470. Cajamarca: 
Hualgayoc, Weberbauer 4023 (det. Schneider). Cuzco: Gay 349. 
Ollantaitambo, Herrera 3419, 1471. Bolivia to Colombia. "Mi- 
llucassa," "llinlli." 

Hesperomeles latifolia (HBK.) M. Roem. Syn. Rosifl. 225. 
1847. Osteomeles latifolia HBK. Nov. Gen. & Sp. 6: 212. pi. 554. 1824. 

A shrub 4 meters high; branchlets, leaves beneath, and corymbs 
densely ferruginous-pilose or tomentulose; petioles 7-15 mm. long; 
blades broadly elliptic, truncate or slightly cordate at the suboblique 
base, rounded at the apex, to 10 cm. long and 7 cm. wide, or the lower 
probably larger, dull and glabrous above except the more or less 
tomentulose costa and the obscure nerves, these 7 mm. distant, in 
age slightly reticulate-veined beneath; corymbs compact, shorter than 
the leaves; bractlets filiform; calyx tube subulate, short, much shorter 
than the woody receptacle. Doubtfully distinct from H. lanuginosa, 
sens. lat. Neg. 3376. 

Cuzco: Yanamanchi, 3,300 meters, Weberbauer 4970 (det. Schnei- 
der). Junin: Huacapistana, Killip & Smith 24236. Piura: Aya- 
vaca, Humboldt, type. "Lengli." 

Hesperomeles palcensis Schneid. Bot. Jahrb. 42: 86. 1908. 
Prostrate, unarmed, the young parts ferruginous- tomentose; 
leaves elliptic-obovate, more or less rounded at the apex, rounded or 



FLORA OF PERU 1069 

acute at the base, irregularly dentate, 1.5-2.5 cm. wide; inflorescence 
distinctly exceeding the leaves, glabrate or pubescent; flowers 4-5 
mm. broad; calyx lobes triangular, short-acuminate, little shorter 
than the receptacle. Compared by the author with H. ferruginea, 
but not surely distinct from that or from H. Gayana. Neg. 1285. 

Junin: Palca, Weberbauer 1782, type. 

Hesperomeles pernettyoides Wedd. Chlor. And. 2: 230. 1857. 
Osteomeles pernettyoides Dene. Nouv. Arch. Mus. Paris 10: 184. 
1874. H, pernettyoides var. glaucophylla Wedd. op. cit. 231. 

Shorter branchlets sometimes spinose, at first puberulent; petioles 
2-3 cm. long; blades more elliptic-oblong than obovate, acute at the 
base, rounded at the apex, slightly lustrous above, sparsely and 
evanescently pilose beneath, mostly about 15 cm. long, 8-10 cm. wide, 
often much smaller on the same branch, minutely and evenly crenu- 
late-denticulate to below the middle; corymbs villous, dense, few- 
flowered; calyx sparsely villous, the filiform-subulate tube 1.5 mm. 
long; petals glabrous; disk sparsely pubescent. Very doubtfully dis- 
tinct from H. cuneata. The Gay specimen from Cuzco cited by 
Weddell I have referred to H. escalloniaefolia. Neg. 34709. 

Junin: Yanahuanca, 1286. Lima: Matucana, 2,300 meters, 
Weberbauer 116 (det. Schneider). Cajamarca: Ocros, 2,400 meters, 
Weberbauer 2735 (det. Schneider) . Caya and Chachapoyas, Raimondi, 
type of var. glaucophylla. Puno: Sandia, 2,100 meters, Weberbauer 
511 (det. Schneider). Huanuco: Chinchapalca, 1590. Mito, 1507. 
Tomaiquichua, 2418. Bolivia; Colombia. 

Hesperomeles Weberbaueri Schneid. Bot. Jahrb. 42: 85. 1908. 

A shrub about 2 meters high; young branchlets (sometimes api- 
cally spinose), leaves beneath, and inflorescence more or less ferrugi- 
nous-pilose; leaves mostly oblong to broadly elliptic, subrotund or 
somewhat cuneate at the base, crehate-dentate; sepals narrowly tri- 
angular, acuminate, longer than the receptacle; petals ovate-oblong, 
sometimes denticulate, greenish white; ovules solitary. Neg. 1286. 

Ayacucho: Above Osno, 2,500 meters, Weberbauer 5602 (det. 
Mansfeld). Lima: San Mateo, Isern 2254. Huanuco: Monzon, 
2,400 meters, Weberbauer 3424, type. Llata, 2267. Pampayacu, 
Kanehira 103. Tambo de Vaca, 4415. Puno: Sandia, 2,200 meters, 
Weberbauer 528 (det. Schneider). Bolivia. 



1070 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

2. LICANIAAubl. 

Moquilea Aubl. 

Reference: J. D. Hooker in Mart. Fl. Bras. 14, pt. 2: 8-19. 1867. 

Shrubs or trees with alternate, simple, persistent leaves, these 
most often glabrous above and tomentose beneath, the petioles some- 
times biglandular at the apex. Flowers small, in divaricately 
branched panicles, short-pedicellate or sessile, 3-bracteolate. Calyx 
tube villous or naked within, the 5 lobes small. Stamens 3-10, included 
or exserted, minute, sometimes connate in a ring. Ovary inserted at 
the base of the calyx tube, 1-celled. Fruit 1-seeded, ligneous or 
sometimes fleshy. The similar genus Parinarium has the style aris- 
ing at the apex of the ovary. 

Leaves glabrous, or the nerves of the younger ones slightly pubescent. 
Inflorescence cinereous. 

Leaves ample, obscurely reticulate- veined L. elata. 

Leaves small, finely reticulate- veined on both sides . .L. octandra. 

Inflorescence fulvous L. intrapetiolaris. 

Leaves pubescent beneath, the pubescence sometimes araneose. 
Leaves broadly elliptic or subrotund, rarely half longer than broad. 
Petioles 10-15 mm. long. 
Leaves conspicuously reticulate- veined beneath. 

L. parinarioides. 

Leaves only slightly reticulate-veined L. retusa. 

Petioles 2-4 mm. long L. octandra. 

Leaves narrower or, if elliptic, about twice as long as wide. 
Stamens conspicuously exserted. 

Leaves rarely 3 (-4.5) cm. wide, the venation dense. 

L. octandra. 

Leaves usually 4 (-7) cm. wide, the venation lax L. lata. 

Stamens very short. 

Leaves ferruginous- villous beneath L. lucida. 

Leaves pale or white-tomentulose. 

Branchlets ferruginous-pilose L. trigonioides. 

Branchlets glabrate L. microcarpa. 

Licania elata Pilger, comb. nov. Moquilea elata Pilger, Notizbl. 
Bot. Gart. Berlin 6: 136. 1914. 

A tall tree, glabrous except for the villous inflorescence; branchlets 
conspicuously lenticellate; blades dull, oval to oblong, short-acumi- 



FLORA OF PERU 1071 

nate, 10-13 cm. long, 4.5-6 cm. wide; petioles 11-13 mm. long; pani- 
cles terminal, to 18 cm. long, the spreading branches to 8 cm. long; 
flowers densely fasciculate, urceolate-campanulate, 2.5-3 mm. long, 
densely villous at the base of the stamens; petals villous-ciliate; fila- 
ments glabrous. Reported by Tessmann as a tree 15 meters high, 
the trunk 50 cm. in diameter. In appearance similar to L. pallida 
Benth. Neg. 18072. 

Rio Acre: Ule 9446. Loreto: Rio Ucayali, Tessmann 3215, 3250. 
Puerto Arturo, Williams 5064- Yarina-cocha, Tessmann 5468. 

Licania intrapetiolaris Spruce ex Hook. f. in Mart. Fl. Bras. 
14, pt. 2: 11. 1867. 

A nearly glabrous tree, as much as 10 meters high, the ample 
panicle conspicuously fulvous-pubescent; leaves oblong-elliptic, some- 
times broadly so, 15-25 cm. long, 7-10 cm. wide, glabrous or the 
prominent costa and 7-9 lateral nerves beneath evanescently ap- 
pressed-pilose, thick-coriaceous; stipules as long as the stout petioles, 
often somewhat connate, borne between the petiole and the branchlet, 
the glands often obscure; flowers 3 mm. long; calyx sericeous-strigil- 
lose, not araneose within; petals and stamens 5, the short filaments 
connate; style and ovary pubescent. Illustrated, Mart. Fl. Bras. 
14, pt. 2: pi. 2. The form with pubescent nerves and somewhat 
smaller flowers (2 mm. long) is var. brevis Macbr. Candollea 5: 
369. 1934. 

Loreto: Mishuyacu, near Iquitos, Klug 612. Amazonian Brazil. 

Licania lata Macbr. Candollea 5: 369. 1934. 

A tree of 18 meters, the branchlets soon glabrous; petioles 7-10 
mm. long; blades chartaceous-coriaceous, lustrous above, white- 
tomentulose beneath, slender-veined, oblong-elliptic or sometimes 
elliptic, rounded or rarely somewhat acute at the base, short-acumi- 
nate, to 16 cm. long, about 4.5 cm. or even 6-7 cm. wide; glands 
more or less prominent; panicles ferruginous-pubescent, 10-15 cm. 
long, the branches 2.5-8 cm. long, densely flowered, the white or 
yellowish flowers sessile; calyx campanulate, cinereous- tomentulose, 
3 mm. long, densely lanuginous within, the lobes broadly ovate, 
acute; petals none; stamens about 12; filaments glabrous, twice 
exceeding the calyx lobes; ovary and style long-pilose. Perhaps a 
form of L. octandra. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 560, type; 
587. Santa Rosa, Kittip & Smith 28942. 



1072 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Licania lucida Macbr. Candollea 5: 369. 1934. 

Branchlets, leaves beneath, and panicles softly and shortly fer- 
ruginous- villous; petioles 2-3 mm. long; stipules linear-lanceolate, 
5-7 mm. long; blades ovate-elliptic, slightly cordate at the base, 
subabruptly acuminate, mostly 10-12 cm. long, 5-7 cm. wide, some 
of the leaves much smaller, glabrous and very lustrous above, the 
veins reticulate beneath; panicle 7-8 cm. long, the branches 2.5-3.5 
cm. long; flowers crowded, sessile; calyx urceolate, 3 mm. long, 
strigillose outside, obscurely costate, more or less lanate-villous 
within; stamens about 10, the filaments short; style appressed-white- 
pilose, 2 mm. long, the similarly pilose ovary scarcely 1 mm. long. 
More pubescent than L. mollis, and the style longer. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 462, type; 
also 499(1). 

Licania microcarpa Hook. f. in Mart. Fl. Bras. 14, pt. 2: 
15. 1867. 

Branchlets slender, glabrous; blades elliptic-ovate, acuminate, 
cordate at the base, glabrous above, scarcely lustrous, cinereous and 
appressed-tomentulose beneath, 6-10 cm. long, 3-4 cm. wide, the 
lateral nerves 10-12, obsolete above, well marked beneath; petioles 
3-4 mm. long, eglandular; stipules subulate, persistent; panicle 
tomentulose, cinereous or pale ferruginous, open, in fruit to several 
centimeters long; calyx tomentose within, araneose; fruit irregularly 
globose, about 7 mm. thick, short-stipitate. Typically a large tree, 
to 25 meters high, with much branched crown, the wood very hard 
(Spruce). Neg. 3370. 

Loreto: Mishuyacu, 100 meters, Klug 1525. San Martin: 
Zepelacio, Klug 3740. Brazil. "Caraipe," "canida." 

Licania octandra (Hoffmansegg) Pilger, comb. nov. Hirtella 
octandra Hoffmansegg ex Roem. & Schult. Syst. 5: 274. 1819. 
L. Turiuva Cham. & Schlecht. Linnaea 2: 550. 1827. 

A small tree, with slender, gray-barked branches, the youngest 
ones sparsely soft-pilose, as much as 7 meters high, sometimes gla- 
brous; petioles 2-4 mm. long; blades oblong-elliptic, acuminate, 
acute at the base, mostly 8 cm. long and 3.5 cm. wide, densely retic- 
ulate-veined, lustrous above, beneath more or less araneose-tomen- 
tulose; panicle 10-15 cm. long, the longer branches 4-5 cm. long, 
rather laxly flowered; flowers greenish yellow, sessile in clusters of 
2-several, cinereous-puberulent, about 3 mm. long; stamens long- 
exserted, united and villous at the base. 



FLORA OF PERU 1073 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 729. Pun- 
chana, near Iquitos, Williams 1331. Rio Ucayali, Tessmann 3446. 
Brazil. 

Licania parinarioides Huber, Bol. Mus. Goeldi 5: 368. 1909. 

A tree of 15 meters, the branchlets rusty-pilose, finally gla- 
brescent; petioles 10-15 mm. long, rusty- tomentose above, distinctly 
biglandular below the apex; blades broadly elliptic to nearly rotund, 
rounded at the base or apex, subcoriaceous, glabrescent and lustrous 
above, cinereous-araneose and minutely pilose beneath, 10-20 cm. 
long, 7-10 cm. wide, with many prominent lateral nerves, these 
puberulent and sparsely long-pilose, the veins beautifully reticulate 
beneath; inflorescence ample, the few divaricate to ascending, strict 
branches velvety with a yellowish tomentum, the rather crowded 
flowers subsessile; calyx in fruit 6 mm. long, white-hispid beneath the 
20 stamens, the lobes 3 mm. long, acute; style villous at the base, the 
slightly excentric ovary 1-celled. 

Loreto: Soledad, Tessmann 5219 (det. Pilger). Mishuyacu, 100 
meters, Klug 1526. Amazonian Brazil. 

Licania retusa Pilger, Notizbl. Bot. Gart. Berlin 6: 137. 1914. 

A tree, as much as 35 meters high, the rigid, more or less roundish 
leaves retuse at the apex, 7-11 cm. long, 4.5-8 cm. wide, very 
minutely but densely tomentulose beneath; petioles 10-13 mm. long; 
panicle to 14 cm. long, densely fulvous- tomentulose; flowers sessile, 
glomerulate, 2.5-3.5 mm. long, the spatulate, villous petals 1 mm. 
long; stamens in a complete ring, short; filaments hirsute- villous 
and united below. Neg. 18080. 

Rio Acre: Vie 9568, type. 

Licania trigonioides Macbr. Candollea 5: 368. 1934. 

A low shrub with elongate, slender, densely ferruginous-hirsutulous 
branchlets; stipules linear-subulate, 7 mm. long, densely appressed- 
ferruginous-pilose; leaves subsessile, the petioles 1-4 mm. long, the 
blades oblong-ovate-elliptic, subcordate at the base, more or less acu- 
minate, mostly about 10 cm. long and 4.5 cm. wide, chartaceous, 
somewhat lustrous above, glabrous except on the costa, beneath 
densely cinereous-tomentulose, the nerves and especially the costa 
sparsely ferruginous-villous, the veins obscure, slender-reticulate on 
both sides; panicles remotely branched, ferruginous-hirsute, about 
20 cm. long, the spreading, remotely flowered branches to 7 cm. long; 
flowers sessile in fascicles of 2-3, sordid-tomentulose and pilose; 



1074 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

calyx tube urceolate, hirsute within, scarcely 15 mm. long, the lobes 
1 mm. long, ovate, acute; petals none; stamens 5-7, the filaments 
densely villous; style villous, 2 mm. long; ovary velvety-pilose. 
Suggestive of L. incana Aubl., of Brazil and the Guianas. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 449, type. 

3. PARINARIUM Juss. 

Trees with alternate, persistent, often coriaceous, entire leaves, 
sometimes 2-glandular at the base, the stipules subulate or lanceolate. 
Flowers racemose, corymbose, or paniculate, 2-bracteolate, perfect. 
Stamens 10 to many, short-connate, often unilateral. Ovary uni- 
laterally adnate, exserted, 2-celled, the septum sometimes incomplete, 
the cells 1-ovuled, the basal style often hirsute. Drupe sometimes 
edible. 

Parinarium parile Macbr. Candollea 5: 367. 1934. 

Branchlets somewhat tuberculate, glabrous; petioles thick, 8-10 
mm. long; blades oblong-elliptic or elliptic-lanceolate, subrotund at 
the base, acuminate at the apex, to 16 cm. long and 5 cm. wide, or 
often 9 cm. long and 3 cm. wide, coriaceous or coriaceous-charta- 
ceous, glabrous and somewhat lustrous above, cinereous-tomen- 
tulose between the veins beneath, finally glabrate, the nerves about 
25, prominent, especially beneath; panicle branches pubescent, with 
ferruginous, somewhat spreading hairs, the branchlets and flowers 
appressed-cinereous-pilose; pedicels obsolete or 1 mm. long; bracts 
2.5 mm. long, caducous; calyx tube turbinate, 2 mm. long, densely 
pilose within only at the throat, the acuminate, narrowly ovate lobes 
little longer; petals caducous, oblong-elliptic, scarcely 1.5 mm. long; 
stamens to 6, unilateral; filaments glabrous, free, scarcely as long or 
little longer than the sepals; style pilose at the base. 

Loreto: Manfinfa, upper Rio Nanay, Williams 1140, type; 1113. 
San Martin: Juanjui, Klug 3870. "Uchpa-umari." 

4. HIRTELLA L. 

Cosmibuena R. & P. Prodr. 10. pi. 2. 1794. 

Reference: Hooker in Mart. Fl. Bras. 14, pt. 2: 27-40. 1867. 

Shrubs or trees with short-petioled, alternate, entire leaves, the 
narrow stipules deciduous. Flowers in axillary and terminal racemes 
or panicles, the panicles usually racemiform. Sepals reflexed, the 
petals early deciduous, much exceeded by the 3 to several stamens, 



FLORA OF PERU 1075 

these partly staminodial and confined to one side of the tubular 
receptacle. Fruit a drupe, sometimes juicy, 1-seeded. 

Leaves bulbous-inflated at the base. 

Flowers in long racemes H. guainiae. 

Flowers in short racemes, almost fasciculate H. physophora. 

Leaves not bulbous-inflated at the base. 

Bracts glandular, or many of them with at least 1 large gland. 

Leaves to 4.5 cm. wide H. subglanduligera. 

Leaves 5-8.5 cm. wide H. aureohirsuta. 

Bracts eglandular, or the glands at least small and promptly 

caducous. 
Leaves conspicuously ciliate with spreading hairs. 

H. pilosissima. 

Leaves not ciliate, or the marginal hairs appressed. 
Stamens 3 (-4) ; inflorescence rarely a simple raceme. 

Inflorescence usually a racemiform panicle, glabrate or 

more or less pubescent H . triandra. 

Inflorescence a setulose raceme H. plumbea. 

Stamens 5-7. 

Inflorescence an open, cymose panicle H. rasa. 

Inflorescence a simple raceme, or racemiform. 

Inflorescence a racemiform panicle H. elongata. 

Inflorescence a simple raceme. 
Leaves oblong-lanceolate, more or less petiolate. 

H. racemosa. 
Leaves elliptic, subcordate, subsessile . . . . H . Standleyi. 

Hirtella aureohirsuta Pilger, Bot. Jahrb. 37: 538. 1906. 

A woody vine, all the growing parts conspicuously yellow-hirsute; 
leaves oblong, subsessile, short-acuminate, glabrate and rugose above, 
hirsute beneath, 10-20 cm. long, 5-8.5 cm. wide; racemes to 20 cm. 
long; bracts lance-subulate, mostly biglandular; pedicels to 8 mm. 
long, stipitate-glandular; flowers 5 to many; petals elliptic, 5 mm. 
long, the similar sepals 4-4.5 mm. long. My specimens are from 
a shrub with open, sprawling branches, the flowers greenish red. 
Species apparently very near several older ones. Neg. 3331. 

Amazonas: Moyobamba, 800 meters, Weberbauer 4487, type. 
Junin: La Merced, 5532 (det. Killip). Chanchamayo Valley, Schunke 



1076 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

328, 329. San Ramon, Kittip & Smith 24794- Rio Paucartambo 
Valley, Kittip & Smith 25276. 

Hirtella elongata Mart. & Zucc. Abh. Math. Nat. Cl. Akad. 
Munch. 1: 384. 1832; 39; Flora 15, pt. 2: Beibl. 1: 85. 1832. 

Branchlets and panicles cinereous-puberulent, like the short 
petioles and the leaf nerves beneath; blades oblong-elliptic, acumi- 
nate, rounded or cordate at the base, glabrous and lustrous above 
except on the costa, often bullate, the 12 lateral nerves prominent; 
stipules linear, subulate, much longer than the petiole, soon decid- 
uous; panicle elongate, narrowly racemiform, the stout rachis and 
the branchlets lenticellate, the branchlets numerous, about 1.5 cm. 
long, apically 1-2-flowered; stamens 6; ovary very hirsute, but the 
style glabrous, like the reddish fruit at maturity. An irregularly 
branched tree, 6 meters high, the branches hollow, housing wasps 
and ants (Spruce). A liana (Klug). Illustrated, Mart. Fl. Bras. 14, 
pt. 2: pi. 11. Neg. 3337. 

Loreto: Mishuyacu, 100 meters, Klug 916, 726, 2530, 2575. 
Brazil. 

Hirtella guainiae Spruce ex Hook, in Mart. Fl. Bras. 14, 
pt. 2: 31. 1867. 

Branchlets and leaves beneath hispid with long, spreading hairs; 
cordate leaf base saccately enlarged on each side of the short petiole, 
the vesicles hispid like the prominent leaf nerves beneath; stipules 
filiform, about 12 mm. long; blades coriaceous, glabrous above, 
lustrous on both sides; racemes tomentose, simple, the pedicels 
4 mm. long, nearly equaled by the filiform bracts; stamens 3; style 
pilose toward the base. A shrub 2-3 meters high, with violet 
flowers (Klug). Illustrated, Mart. Fl. Bras. 14, pt. 2: pi. 10. 

Loreto: Mishuyacu, 100 meters, Klug 27, 512; Kittip & Smith 
29947. Amazonian Brazil. 

Hirtella physophora Mart. & Zucc. Abh. Math. Nat. Cl. Akad. 
Munch. 1: 374. 1832; 29; Flora 15, pt. 2: Beibl. 1: 79. 1832. 

In general similar to H. guainiae, but spreading-hirsute through- 
out, even on the upper leaf surface; racemes scarcely 2.5 cm. long, 
fasciculate; flowers 2-2.5 cm. broad, the pedicels 8 mm. long; sta- 
mens 7; style glabrous; ovary strigose, almost included in the calyx 
tube. Flowers greenish or bright brown (Klug). H. myrmecophila 
Pilger is similar, but the inflorescence is very sparsely hispid and 
the ovary is near the base of the tube. Neg. 3348. 

Loreto: Mishuyacu, 100 meters, Klug 989. Brazil. 



FLORA OF PERU 1077 

Hirtella pilosissima Mart. & Zucc. Abh. Math. Nat. Cl. Akad. 
Munch. 1: 373. 1832; 30; Flora 15, pt. 2: Beibl. 1: 78. 1832. 

Well marked by the broadly elliptic, oblong, or somewhat obo- 
vate, setulose-margined leaves, these chartaceous, subacute at the 
base, cuspidate, hirsute above with slender hairs; petioles very 
short; pedicels 3-4 mm. long, equaled by the subulate bracts; 
stamens 3-4; ovary strigose, the style pilose. Neg. 3349. 

Loreto: Rio Nanay, Williams 804. Marquisapa, Williams 1183. 
Brazil. 

Hirtella plumbea Pilger, Notizbl. Bot. Gart. Berlin 6: 139. 1914. 

A shrub or perhaps a tree, with lustrous, lead-colored foliage; 
stipules to 8 mm. long; leaves sparsely setose beneath, long-acumi- 
nate, to 18 cm. long and 6.5 cm. wide; inflorescence 15-30 cm. long, 
fuscous-hirsute-setose; pedicels 3-4 mm. long; sepals 3-3.5 mm. long, 
the lilac petals little longer; stamens 4, about 12 mm. long, the style 
not much longer. Neg. 3350. 

Rio Acre: Ule 79br, type. 

Hirtella rasa Standl. Field Mus. Bot. 17: 252. 1937. 

A tree 6-27 meters tall; leaves elliptic-oblong or oblong, acute 
or short-acuminate, 11-13 cm. long, 4-6.5 cm. wide, lustrous above, 
sparsely and minutely hispidulous or almost glabrous, somewhat 
paler beneath and sparsely and minutely strigillose-scabrous or 
almost glabrous, subcoriaceous; petioles rather thick, 5-7 mm. long; 
inflorescence terminal, 8-19 cm. long, minutely and densely ap- 
pressed-pilose with cinereous or fulvous hairs; bracts broadly ovate; 
flowers subsessile; sepals 3-4 mm. long, densely fulvous- or cinereous- 
sericeous, about equaling the white, glabrous petals; stamens 5; 
fruit densely spreading-pilose. 

San Martin: Juanjui, King 4249. Amazonian Brazil. 

Hirtella racemosa Lam. Encycl. 3: 133. 1789; 33. H. ameri- 
cana Auct., non L. 

A shrub or small tree, the leaves oblong or oblong-lanceolate, 
acute to long-acuminate, usually 3-4.5 cm. wide, about 10 cm. long, 
sometimes to 6.5 cm. wide and 16 cm. long, glabrate or somewhat 
strigose on the costa and sometimes on the nerves beneath, mem- 
branaceous or becoming subcoriaceous; racemes axillary and termi- 
nal, greatly elongate in fruit, slightly puberulent to hirsutulous; bracts 
subulate, usually shorter than the pedicels, these slender, 5-15 mm. 
long; stamens 5-7; fruit glabrous. Variable, especially in the leaves, 



1078 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

a number of varieties under the name H. americana having been 
designated by Hooker. 

San Martin: Tarapoto, Ule 6541, 6453; Williams 6154, 5887. 
Chazuta, King 4025. Pongo de Cainarachi, King 2704, 2714 
Loreto: Yurimaguas, Ule 6264; Williams 3884, 7832, 4041, 3829; 
Killip & Smith 27549, 29651. Mishuyacu, King 203, 263, 1333. 
Florida, Klug 1 989. Masisea, Tessmann 3115. La Victoria, Williams 
2765, 2990. Caballo-cocha, Williams 2358(1}. Amazonas: Chico- 
playa, Ruiz & Pavdn (det. Pilger). Moyobamba, Weberbauer 4573. 
Rio Acre: Ule 9412, 9415. Widely distributed in South America, to 
Central America and the West Indies. 

Hirtella Standleyi Baehni & Macbr., sp. nov. 

Arbor parva 3-5 m. alta, ramulis robustis, racemis, petiolisque 
dense pilis brevibus fulvis patentibus pubescentibus; stipulis 5 mm. 
longis; petiolis 1-15 mm. longis haud incrassatis; foliis oblongo-ellip- 
ticis vel interdum late oblongo-ovatis basi manifeste sed minute 
cordatis apice abrupte acuminatis (acumen ca. 4 mm. longus), plerum- 
que 8-10 cm. longis, 5-6 cm. latis, rigido-coriaceis, supra costa excepta 
nitidulis glabratis, subtus ad nervos plus minusve hirsutulis, sub- 
nitidis, nervis lateralibus cum venis laxe reticulatis subtus prominen- 
tibus, supra obscuris; racemis ad 15 cm. longis densifloris; pedicellis 
vix 2.5 mm. longis; bracteis setaceis 3 mm. longis; bracteolis minutis 
glanduliformibus; petalis late obovatis 4 mm. longis, 3 mm. latis; 
drupa oblongo-ovoidea breviter stipitata sparse pilosa 3.5 mm. 
longa, 2.5 mm. crassa. Species otherwise similar to H. scaberula 
Spruce in Mart. Fl. Bras. 14, pt. 2: 33. 1867, of the upper Amazon, 
with thinner leaves that are not at all cordate, borne on glabrate, 
much thickened petioles, the racemes sparsely pilose, and with smaller, 
spatulate-oblong petals. We acknowledge our thanks to Sir Arthur 
W. Hill for the loan of Spruce's type. 

Loreto: A tree with lilac flowers, Balsapuerto, Klug 3022, type; 
also 2869. 

Hirtella subglanduligera Pilger, Notizbl. Bot. Gart. Berlin 
6: 140. 1914. 

A tree of 10-25 meters, the younger parts and the many inflores- 
cences more or less villous-hirsute; leaves coriaceous, oval or elliptic, 
broadly cuneate at the base, narrowed to the obtuse apex, 8-9 cm. 
long, to 4.5 cm. wide, subsessile; panicle branches short, 1-few- 
flowered; glands of the bractlets large, sessile or subsessile; sepals 



FLORA OF PERU 1079 

3.5-4 mm. long; petals 5 mm. long; stamens 4, about 13 mm. long, 
the longer style hirsute below. Neg. 3357. 

Rio Acre: Ule 9414, type. 

Hirtella triandra Swartz, Prodr. 51. 1788. H. Cosmibuena 
Lam. Illustr. 2: 114. 1797. H. racemosa R. & P. Fl. Peruv. 3: 5. 
1802, non Lam. 1789. 

A shrub or slender tree, sometimes 10 meters high, with virgate, 
glabrate or somewhat pubescent branchlets; leaves usually oblong 
or oblong-lanceolate, acute, acuminate, or caudate-acuminate, usually 
rounded at the base, often about 10 cm. long and 4.5 cm. wide; 
flowers racemose-paniculate, the pedicels slender, 5-10 mm. long, 
sometimes shorter or longer; bracts small, subulate, deciduous, 
eglandular; stamens 3, rarely 4; petals white, roundish, about 
equaling the calyx; fruit juicy, plumlike, purplish, not edible, 
1 cm. long. 

Loreto: Yurimaguas, Williams 4653. Lower Rio Huallaga, 
Killip & Smith 2927. Florida, King 2250. San Martin: Zepe- 
lacio, King 3724. Rio Acre: Ule 9413. Widely distributed in 
tropical America. "Quinulla," "isiguiro-ey" (Huitoto names). 

5. COUEPIA Aubl. 

Reference: J. D. Hooker in Mart. Fl. Bras. 14, pt. 2: 40-49. 1867. 

Glabrous or tomentose trees or shrubs with alternate, coriaceous, 
entire leaves, the short petioles sometimes biglandular toward the 
apex. Stipules often setaceous and deciduous. Flowers in axillary 
or terminal panicles or racemes, usually white-tomentose and often 
large, like the more or less caducous bracts. Calyx tube elongate. 
Stamens 15 to many, somewhat monadelphous at the base, some- 
times laterally disposed. Ovary 1-celled, laterally inserted in the 
calyx throat. Style elongate, usually villous. Fruit often edible. 

Leaves small, all or most of them narrower than 4 cm. 

Calyx glabrous C. Williamsii. 

Calyx pubescent C. Ulei. 

Leaves medium-sized to large, mostly 5 cm. wide or wider. 

Branchlets villous C. canomensis. 

Branchlets glabrous or nearly so. 

Calyx more or less cinereous-puberulent C. paraensis. 

Calyx densely fulvous-pubescent, pale in C. subcordata. 



1080 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Calyx tube about 1 cm. long; style pubescent below the 
middle. 

Leaves obscurely reticulate- veined C. subcordata. 

Leaves coarsely and prominently reticulate-veined. 

C. racemosa. 
Calyx tube about 2 cm. long, or the style pubescent. 

Calyx subsessile, about 5 mm. thick C. macrophylla. 

Calyx pedicellate, about 3 mm. thick C. chrysocalyx. 

Couepia canomensis (Mart.) Benth. in Mart. Fl. Bras. 14, 
pt. 2: 42. 1867. Moquilea canomensis Mart. Nov. Gen. & Sp. 2: 
80. pi. 166. 1826. 

A small tree, to 7 meters high, very distinct among Peruvian 
species by its softly reddish- villous branchlets; leaves oblong- 
lanceolate, in age coriaceous and glabrous above, to 15 cm. long, 
sordidly white-tomentulose beneath, the nerves villous, the reticu- 
lation becoming prominent; racemes solitary, sometimes nodding, 
to 10 cm. long, rufous-pubescent throughout; bracts lanceolate, 
exceeding the cylindric calyx tube; flowers almost 12 mm. long, the 
petals much narrower but not longer than the oblong (typically 
6 mm. long) calyx lobes; fruit globose, pubescent, lanate within, 
3.5 cm. thick. 

Loreto: Iquitos, Williams 3780; Klug 142 (sepals longer); Tess- 
mann 3690, 5109 (det. Pilger). Brazil. 

Couepia chrysocalyx (Poepp. & Endl.) Benth. ex Hook. f. in 
Mart. Fl. Bras. 14, pt. 2: 42. 1867. Moquilea chrysocalyx Poepp. 
& Endl. Nov. Gen. & Sp. 3: 75. pi. 286. 1845. 

A rather large tree, up to 16 meters high, much branched from 
the base (Spruce), or 10 meters high, with a trunk 30-40 cm. in 
diameter (Tessmann), the youngest branchlets and the leaves 
beneath white-araneose-tomentose; leaves oblong, usually more or 
less acute, finally coriaceous, glabrous above, 15-25 cm. long; racemes 
short, axillary, corymbose; bracts small, deciduous before an thesis; 
flowers about 2.5 cm. long; calyx tube 12-15 mm. long; petals white, 
oblong, exceeding the roundish calyx lobes; ovary and style densely 
pubescent. 

Loreto: Puerto Metendez, Tessmann 3918 (det. Pilger). Yuri- 
maguas, Kittip & Smith 27933; Williams 5012. San Martin: Tara- 
poto, Spruce 4616 (calyx shorter). Pongo de Cainarachi, Klug 2656. 
Chazuta, Klug 3971. "Parinari," "sacha umapu." 



FLORA OF PERU 1081 

Couepia macrophylla Spruce ex Hook. f. in Mart. Fl. Bras. 14, 
pt. 2: 41. 1867. Couepia speciosa Pilger, Bot. Jahrb. 37: 539. 1906. 

A small tree, 6 meters high, with suboblong or oblong-obovate, 
coriaceous leaves, those of the flowering branchlets about 20 cm. 
long and 6-8 cm. wide, the lower to 35 cm. long and nearly 15 cm. 
wide, shortly obtuse-acuminate, sometimes cuneate at the base, 
pale above, dark and minutely tomentose beneath; petioles about 
1 cm. long, stout; racemes dense, 6-8 cm. long; calyx densely seri- 
ceous- tomentose; sepals 5, orbicular-ovate, 1 cm. long; petals white, 
villous-margined; staminal circle complete, the filaments 2.5 cm. 
long. The fleshy, ovate bracts are caducous. Williams 888, sterile, 
from Timbuchi, called "chibo-caspi" and "capucarnia," is perhaps 
this species. Neg. 17976. 

Amazonas: Moyobamba, 1,400 meters, Weberbauer 4649. 
Loreto: Florida, Klug 2089. Amazonian Brazil. 

Couepia paraensis (Mart. & Zucc.) Benth. in Mart. Fl. Bras. 
14, pt. 2: 48. 1867. Moquilea paraensis Mart. & Zucc. Abh. Math. 
Phys. Cl. Akad. Munch. 1: 390. 1832. 

A tree of 10-13 meters; branchlets glabrous, white-punctate; 
petioles 8 mm. long, tomentose, biglandular; blades oblong or obo- 
vate-oblong, abruptly acute or acuminate, rounded at the base or 
rarely cordate, 7.5-15 cm. long, glabrous and lustrous above, the 
slender veins impressed, glabrate beneath, with 12-20 slender nerves, 
the veins obscure; stipules subulate, deciduous; panicles solitary, 
terminal, longer than the leaves, erect or nodding, many-flowered, 
even the flowers cinereous- tomentose; bractlets early caducous; 
pedicels slender, 12-16 mm. long; calyx tube 8 mm. long, attenuate 
at the base, the lobes rounded, little exceeded by the marginally 
pubescent, white or salmon-pink petals; stamens 50 or fewer, 8 mm. 
long, more or less unilateral; ovary, like the style base, villous; 
fruit oblong, 2.5 cm. long, glabrous within. 

Loreto: Iquitos, 100 meters, Williams 1490; Killip & Smith 
27184, 29963. Mishuyacu, 100 meters, Klug 1444, 2503, 2520. 
Amazonian Brazil. 

Couepia racemosa Benth. in Mart. Fl. Bras. 14, pt. 2: 43. 1867. 

Branchlets not white-punctate, glabrous; petioles 8-12 mm. long; 
blades thick-coriaceous, glabrous but not lustrous above, the nerves 
deeply impressed, densely reticulate-veined beneath, the 8-12 lateral 
nerves prominent, 10-20 cm. long, oblong or obovate-lanceolate, 
rounded to the acute base, acuminate; racemes axillary, somewhat 



1082 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

shorter than the leaves, ferruginous- tomentose, 5-10 cm. long; 
pedicels slender; calyx tube 6 mm. long, conic-cylindric, gibbous 
at the base, the oblong lobes obtuse; petals oblong, glabrous, white, 
exserted; stamens 40 or fewer, 12 mm. long; ovary hirsute, the 
style villous at the base; fruit large, ovoid, woody, obscurely 6-costate 
pubescent within. Stipules none, according to Spruce. Williams 
1161 seems to be var. reticulata Pilger, the leaves closely reticulate- 
veined. 

Loreto: Timbuchi, Williams 1161. Brazil. "Mashu sacha." 

Couepia subcordata Benth. in Mart. Fl. Bras. 14, pt. 2: 46. 
pi. 16. 1867. C. amazonica Fritsch, Ann. Hofmus. Wien 5: 12. 1890. 

A tree of 12 meters, with dense crown; branchlets thick, at first 
cinereous-puberulent; petioles pubescent, eglandular, 6-12 mm. long; 
blades oblong-lanceolate, acute or acuminate, more or less cordate 
at the base, 7.5-15 cm. long, submembranaceous, only slightly 
lustrous above, the nerves slender, beneath ferruginous (rarely 
cinereous), the 12-15 prominent nerves glabrous; stipules deciduous, 
subulate; panicles dense, axillary and terminal, equaling the leaves 
or longer, white-tomentose even to the flowers; pedicels 8-12 mm. 
long, 3-5 together on each ascending branchlet; calyx tube cylindric, 
the acute, oblong lobes 6 mm. long; petals narrow, ciliate; stamens 
to 40, the filaments connate at the base; ovary villous, the style 
densely hirsute below the middle. 

Loreto: La Victoria, Williams 3173. Amazonian Brazil. 

Couepia Ulei Pilger, Verh. Bot. Ver. Brandenb. 47: 149. 1905. 

A shrub, characterized by its small leaves, the larger ones 
7-10 cm. long; petioles rarely 5 mm. long; blades smooth and lustrous 
above, ferruginous-tomentulose beneath, only the lateral nerves 
obvious; inflorescence terminal, or 1-2 racemes from the upper leaf 
axils, olive-gray-pubescent; petals 5-7 mm. long, somewhat pubes- 
cent, at least on the margins; calyx lobes 4.5-5 mm. long; style 
1-1.5 cm. long, villous toward the base. The fruit of Williams 8157 
and 2346 is of the size and shape of a small pear; it is brown, with 
numerous pale flecks. Neg. 17981. 

Loreto: Rio Itaya, Williams 7, 29, 63, 88. Rio Mazan, near 
Iquitos, Williams 8157. Caballo-cocha, Williams 23^6. Amazonian 
Brazil. "Parinari," "suichi-parinari." 

Couepia Williamsii Macbr. Candollea 5: 366. 1934. 
A small tree with slender, somewhat tortuous branchlets; petioles 
5 mm. long; blades narrowly oblong or elliptic, short-acute or sub- 



FLORA OF PERU 1083 

rotund at the base, subabruptly acute-acuminate, mostly 11 cm. 
long and 3.5 cm. wide, but sometimes 16 cm. long and nearly 6 cm. 
wide, lustrous and glabrous above, densely appressed-ferruginous- 
tomentulose beneath, chartaceous-membranaceous, the nerves con- 
spicuous beneath; racemes slender, axillary and terminal, 6-10- 
flowered, glabrous; pedicels 5 mm. long; flowers about 2 cm. long; 
calyx slightly curved, the cylindric tube somewhat gibbous at the 
base, about 1.5 cm. long, the broadly ovate lobes 6 mm. long; 
stamens almost 1 cm. long; style 9-11 mm. long, densely villous, 
even to the tip. Close to C. leptostachys Benth., but in that Brazilian 
species the style is pubescent only at the base. 

Loreto : La Victoria, Williams 2975, type. 

6. PRUNUS L. 

Reference: Koehne, Bot. Jahrb. 52: 279-333. 1915. 

This well known genus is characterized chiefly by its free calyx 
tube, which does not crown the drupaceous fruit. The stamens in 
the Peruvian species are often more than 20. The cherry, commonly 
grown for food knd for ornament near Chosica, Cuzco, and many 
other places in the mountain valleys is P. Capuli Cav., originally 
of Mexico. When the genus is sufficiently known, the total number 
of Peruvian species probably will be materially reduced. 

Petals pink; flowers subsessile, occurring in clusters before the leaves 

appear; ovary velvety- tomentose P. Persica. 

Petals white; flowers pediceled, in racemes with or after the leaves; 
ovary glabrous. 

Cupule under the ovary persistent P. Capuli. 

Cupule under the ovary deciduous. 
Leaves entire. 

Leaves about 3 cm. long, 2 cm. wide P. ovalis. 

Leaves much larger. 

Inflorescence usually with 1-4 slightly pubescent branches. 

P. subcorymbosa. 

Inflorescence always a simple, glabrous raceme. 
Leaves broadly oval or subrotund, mostly 7-11 cm. wide. 
Leaves broadly oval; petioles very stout. 

P. amplifolia. 

Leaves subrotund or broadly elliptic; petioles relatively 
slender, sometimes short. 



1084 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels 2-2.5 mm. long; petals 1.5 mm. long. 

P. detrita. 
Pedicels 3-4 mm. long; petals to 3.5 mm. long. 

P. rotunda. 
Leaves suboblong-elliptic, mostly narrower. 

Petioles 9-12 mm. long, or the pedicels short and stout. 
Pedicels stout and short, to 4 mm. long in fruit. 

P. integrifolia. 
Pedicels slender, about 6 mm. long in flower. 

P. Williamsii. 

Petioles 3-8 mm. long; pedicels slender, at least 2.5 
mm. long, except sometimes in P. Ruiziana. 

Leaves 6 cm. long, 3 cm. wide. 
Leaves obscurely veined, short-pointed. 

P. Ruiziana. 

Leaves conspicuously veined P. stipulata. 

Leaves mostly twice as large or larger. 

Petioles very short and stout P. pleiantha. 

Petioles 5 mm. long or more. 
Petioles 5-10 mm. long; pedicels stout and 

short P. integrifolia. 

Petioles 5-8 mm. long; pedicels slender or 

filiform. 
Pedicels 1.5-3 mm. long. 

Glands basal on the leaf blade .... P. sana. 
Glands borne above the base of the leaf 

blade. 
Stamens to 2 mm. long; leaves obscurely 

venose P. vana. 

Stamens to 3 mm. long; leaves reticulate- 
veined beneath P. debilis. 

Pedicels to 6 mm. long in flower. 

P. Williamsii. 
Leaves serrulate, or at least some of them with a few teeth 

near the base. 
Leaves mostly about 8 cm. long and 3.5 cm. wide, or smaller. 

Leaves acuminate P. rigida. 

Leaves rounded at the apex. 



FLORA OF PERU 1085 

Leaves distinctly narrower than wide P. oblonga. 

Leaves subrotund P. ovalis. 

Leaves mostly twice as large, 13-17 cm. long, 5-7 cm. wide. 

P. huantensis. 

Primus amplifolia Pilger, Bot. Jahrb. 37: 538. 1906. 

Lenticels of the branches many; petioles 5-7 mm. long, 2-2.5 
mm. thick; blades coriaceous, broadly elliptic, often somewhat 
one-sided, 15-20 cm. long, 8-11 cm. wide, apically (apparently) 
shortly obtuse-acuminate, scarcely lustrous; nerves rather con- 
spicuous beneath, the glands almost basal; racemes 4-5 cm. long, 
the rachis 1 mm. thick; pedicels slender, 2-2.5 mm. long; petals 
suborbicular, 1.5 mm. long, undulate. Neg. 3311. 

Puno: Chunchosmayo, 900 meters, Weberbauer 1265, type. 
Huanuco: Cochero, Poeppig 1774. 

Primus Capuli Cav. Anal. Hist. Nat. Madrid 2: 110. 1800. 

A shrub or tree, to 15 meters high; petioles 1-2.5 cm. long; 
leaf blades lanceolate to ovate, narrowed and acute at the base, 
caudate-acuminate at the apex, lustrous above, smooth beneath, 
6.5-14 cm. long, 2.5-4.5 cm. wide; racemes terminating leafy 
branches, the peduncles leafy at the base, 3-10 cm. long; flowers 
small, white; ovary glabrous; fruit red or black, 1 cm. in diameter 
or smaller. 

Junin: Tarma, Killip & Smith 21897; Kanehira 61. Huanuco: 
Mito, 1390. "Cerezo," "cereza." 

This tree, native of Mexico, is planted and sometimes is locally 
established along stream banks or in clearings in Peru. It is highly 
esteemed for its large, juicy fruit. 

Prunus debilis Koehne, Bot. Jahrb. 52: 317. 1915. 

Glabrous, the very slender branches with minute lenticels; 
petioles 5-8 mm. long; leaf glands almost basal, 1 mm. wide; blades 
broadly cuneate at the base, gradually and obtusely long-acuminate, 
11-16 cm. long, 4-6 cm. wide, papyraceous or membranaceous, the 
slender nerves prominent-reticulate beneath; racemes 3-5.5 cm. 
long, lax, the nearly filiform pedicels 1.5-3 mm. long; calyx cup 
1.8 mm. long, 2.2 mm. wide; pedicels 1.5 mm. long and broad; 
anthers 0.25 mm. long; stigma 0.3 mm. broad. Neg. 31347. 

Loreto: Mainas, Yurimaguas, Poeppig, type. 



1086 FIELD MUSEUM OF NATURAL HISTORY BOTANY, *VOL. XIII 

Primus detrita Macbr. Candollea 5: 365. 1934. 

A glabrous tree about 8 meters high, with slender, obscurely 
lenticellate branches; petioles 12 mm. long; glands subbasal; blades 
elliptic-oval or somewhat ovate-elliptic, rounded to the acute base, 
abruptly and obtusely acuminate at the apex, 12-15 cm. long, 
6.5-8 cm. wide, entire, coriaceous-chartaceous, lustrous above; 
racemes mostly 5 cm. long; pedicels slender, 2.5-3 mm. long; petals 
1.5 mm. long; stamens 2-2.5 mm. long; anthers minute. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 1094, type. 

Primus huantensis Pilger, Bot. Jahrb. 50: Beibl. Ill: 8. 1913. 

A shrub of 3 meters, the branches brown, with many lenticels; 
leaves rigid, coriaceous, broadly rounded at the base, ovate, gradually 
narrowed to the acuminate apex, rarely subelliptic, shortly and 
rigidly setose-denticulate, more or less bullate, 13-17 cm. long, 
5.5-7 cm. wide, the thick petiole to 1 cm. long, the nerves impressed 
above, prominent beneath; racemes to 17 cm. long; pedicels 4-6 
mm. long; sepals 1 mm. long; petals 2.5-3 mm. long; fruit about 
1 cm. thick. 

Ayacucho: Prov. Huanta, 2,900 meters, Weberbauer 5583, type. 
Huanuco: Mito, 1701. Cani, 8399. "Inca-inca." 

Prunus integrifolia (Presl) Walp. Ann. Bot. 3: 854. 1852-53. 
Cerasus integrifolia Presl, Epim. Bot. 194. 1849. 

A tree of 10-15 meters, the branchlets virgate, smooth; petioles 
5-10 mm. long; blades entire, oblong-elliptic, mostly about 12 cm. 
long and 4 cm. wide, or those on young shoots nearly 20 cm. long and 
8 cm. wide, slightly narrowed to the acute apex, equally rounded at 
the subcordate base, subcoriaceous, dull, scarcely venose; glands 
basal, obscure; racemes about 10 cm. long, the rachis 2 mm. thick; 
pedicels 2 mm. long (in fruit 4 mm.), scarcely 1 mm. thick; calyx 
3 mm. deep, 5 mm. wide; anthers 0.5 mm. long; fruit 10 mm. long. 
The Bolivian P. Brittoniana Rusby is very near this species (see 
under P. rigida). Neg. 3314. 

Huanuco: Cochero, by wooded streams, Ruiz & Pawn; Dombey. 
Mountains of Huanuco (Haenke, type). Panao, 3629. Posuso, 
Weberbauer 6747. Pampayacu, Sawada Pi 5. Junin: Carpapata, 
Killip & Smith 24453. "Imainca." 

Prunus oblonga Macbr. Candollea 5: 364. 1934. 
A glabrous, densely leafy shrub, the rigid, reticulate-veined, 
oblong-elliptic leaves subsessile (petioles scarcely 1 mm. long), 



FLORA OF PERU 1087 

mostly 4 cm. long and half as wide; racemes about 3 cm. long, dense, 
the rachis 2 mm. thick, the pedicels 3-4 mm. long; petals 3 mm. long; 
stamens 2.5 mm. long, the anthers 0.9 mm. long. 
Huanuco : Posuso, Weberbauer 6732, type. 

Primus ovalis Ruiz ex Koehne, Bot. Jahrb. 52: 315. 1915. 

Densely and rigidly branched, with numerous rigid, oval leaves, 
mostly 2-3 cm. long; petioles 2-3 mm. long; blades rounded or sub- 
cordate at the base, densely mucronate-serrulate to entire, incon- 
spicuously veined; glands basal; racemes axillary, 3-4 cm. long, 
dense, the pedicels 1 mm. long; calyx 2.5 mm. long, the subrotund 
petals of equal length. Plant entirely glabrous, even the calyx 
within; stigma about 0.8 mm. broad; anthers 0.7 mm. long. P. num- 
mularia Koehne, loc. cit., has punctate glands and somewhat larger, 
suborbicular leaves, lustrous above. Neg. 3317. 

Huanuco: Pillao, Ruiz, type. Amazonas: Chachapoyas 
(Mathews). 

Primus Persica (L.) Stokes, Bot. Mat. Med. 3: 190. 1812. 
Amygdalus Persica L. Sp. PI. 472. 1753. 

A small tree with broadly lanceolate or lance-oblong, serrate, 
long-acuminate leaves; flowers from a scaly bud; fruit subglobose. 

Junin: Matucana, 11$. 

The peach, native of Asia, is sometimes found escaped from 
cultivation in thickets and along banks of streams in Peru. 

Prunus pleiantha Pilger, Bot. Jahrb. 37: 538. 1906; 317. 

Branchlets drying blackish, the lenticels numerous; leaves sub- 
sessile (petioles about 3 mm. long), coriaceous, oblong or ovate- 
elliptic, rounded or little narrowed at the base, narrowed to an 
elongate, obtuse-acuminate apex, 13-20 cm. long, 5-8 cm. wide, 
the nerves slightly impressed above, scarcely conspicuous, only the 
costa prominent beneath; basal gland 1; racemes many, 8-12 mm. 
long, the stout, angled rachis glabrous; pedicels finally 3-5 mm. 
long; petals suborbicular. Negs. 3319, 31353. 

Huanuco: Monzon, 1,600 meters, Weberbauer 3507, type. 

Prunus rigida Koehne, Bot. Jahrb. 52: 316. 1915. 

A glabrous shrub to 3 meters high, with obscurely lenticellate 
branchlets and rigid, ovate-lanceolate, sparsely denticulate leaves 
on petioles 5-7 mm. long; glands usually 3-5 mm. above the leaf 
base, 0.5-1 mm. wide; teeth mostly or all below the middle of the 



1088 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

blades; racemes numerous but solitary, 4-6 cm. long, the rachis 1-1.2 
mm. thick; calyx cup 3 mm. long and broad; stigma 0.8 mm. broad; 
anthers 1.2 mm. long. Var. subintegra Koehne has longer petioles, 
larger leaves, 10-15 cm. long, more sparsely denticulate, and remote 
racemes. P. Brittoniana Rusby has larger, thinner leaves, obscurely 
denticulate above the middle. Neg. 3321. 

Cajamarca: Above San Miguel, 2,800 meters, Weberbauer 3945, 
type. Cuzco: Gay (the variety). 

Primus rotunda Macbr. Candollea 5: 365. 1934. 

A glabrous tree, well marked by the oval or subrotund, abruptly 
acuminate leaves, lustrous above, dull beneath, about 10 cm. long, 
6-8 cm. wide; branchlets slender, with few obscure lenticels; petioles 
7 mm. long; glands suprabasal, 1 mm. long; racemes about 4 cm. 
long, the rachis 1 mm. thick; pedicels 3^1 mm. long; petals 3.5 mm. 
long; stamens 3 mm. long, the anthers minute. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 797, 823. 
Pomayaco, Klug 3136. 

Prunus Ruiziana Koehne, Bot. Jahrb. 52: 316. 1915. 

Glabrous, the branches obscurely lenticellate; leaf glands basal, 
0.5-1 mm. broad; leaves rounded at the base, narrowly decurrent to 
the (4-7 mm. long) petiole, oblong, to 7 cm. long and nearly 4 cm. 
wide, very obtuse or barely acute, entire, rather conspicuously 
reticulate- veined beneath, rigid-coriaceous; racemes solitary, 4-7 cm. 
long, the rachis 1.5-1.7 mm. thick, the pedicels 2-3 mm. long; calyx 
cup 3 mm. long, 4 mm. broad; petals 4 mm. long; stigma 1.3 mm. 
broad; anthers 0.9 mm. long. Doubtfully distinct from P. integri- 
folia. Neg. 3323. 

Huanuco : Pillao, Ruiz 4, type. 

Prunus sana Macbr. Candollea 5: 363. 1934. 

Glabrous, the robust, leafy branches with numerous conspicuous 
lenticels; glands basal, 2 mm. long; leaves dull, chartaceous, acute at 
each end, elliptic-lanceolate, as much as 13 cm. long and 5 cm. wide, 
the veins, especially beneath, slightly reticulate; racemes crowded 
but solitary, about 4 cm. long, dense, the rachis 1 mm. thick; pedicels 
slender, to 2 mm. long, the petals and stamens scarcely as long; 
anthers minute. 

Cuzco: Paucartambo, Weberbauer 6958, type. 



FLORA OF PERU 1089 

Primus stipulata Macbr. Candollea 5: 363. 1934. 

Branchlets densely and conspicuously tuberculate-lenticellate; 
stipules tardily deciduous, roundish, more than 5 mm. wide; petioles 
2.5-3 mm. long; leaf glands basal; blades ovate-elliptic, rounded at 
the base, acuminate at the apex, 5-7 cm. long, 2.5-3 cm. wide, rigid- 
chartaceous, lustrous, conspicuously reticulate-veined on both sides, 
the slender veins impressed above; racemes 4-6 cm. long, the rachis 
scarcely 1.5 mm. thick; pedicels 4-5 mm. long, densely tuberculate; 
petals and stamens 2.5 mm. long; anthers 0.6 mm. long. 

Peru: Without locality, Weberbauer 6930, type. 

Primus subcorymbosa Ruiz exKoehne, Bot. Jahrb. 52:315. 1915. 

Branches slender, at first rather densely hirtellous, like the some- 
what branched racemes; petioles 6-13 mm. long, glabrous; leaf 
glands basal, 1 mm. broad; blades rounded-elliptic or broadly elliptic, 
or rarely ovate or oblong, 9.5-16 cm. long, 5-8 (-10) cm. wide, entire, 
glabrous, slightly reticulate-veined beneath, papyraceous, nearly 
dull above; racemes clustered and with 1 to several branches, 2-6 cm. 
long, the rachis 0.8-1.5 mm. thick at the base, the pedicels 2-3.5 mm. 
long; sepals 1 mm. long, hirsute within, like the cup outside; petals 
rotund, 2 mm. long, hirsute above the base; ovary glabrous; fruit 
oval, 2 cm. long; stigma 1 mm. broad; anthers 0.8 mm. long. 

Peru: Ruiz, without locality, type. 

Prunus vana Macbr. Candollea 5: 362. 1934. 

A glabrous tree, 5-7 meters high, the branchlets with few minute 
lenticels; petioles 4-7 mm. long; blades broadly ovate-elliptic, mostly 
12-14 cm. long and 5-6 cm. wide, acute at the base, broadly obtuse- 
acuminate at the apex, rigid-chartaceous, very lustrous above, dull 
beneath, almost veinless; racemes to 4.5 cm. long; flowers violet and 
orange (Klug); rachis nearly 1 mm. thick; pedicels to 2 mm. long, 
stout; petals and stamens scarcely as long, the anthers minute. 

Junin: La Merced, Killip & Smith 23772, type. Yapas, Killip & 
Smith 25467. San Martin: Near Moyobamba, King 3613. 

Prunus Williamsii Macbr. Candollea 5: 364. 1934. 

Glabrous, the slender branchlets with numerous lenticels; petioles 
mostly 4-10 mm. long; blades lustrous above, chartaceous, incon- 
spicuously veined, broadly ovate-elliptic, to 14 cm. long and 6 cm. 
wide, rotund-acute at the base; glands 2 mm. long; racemes 6-9 cm.' 



1090 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long, lax, the rachis 1 mm. thick, the slender pedicels 6-7 mm. 
long; petals 2 mm. long; stamens 2.5 mm. long, the anthers 0.3- 
0.4 mm. long. 

San Martin: San Roque, Williams 7396, type. Chazuta, Klug 
4122, 4124- 

1. POLYLEPIS R. & P. 
Reference: Bitter, Bot. Jahrb. 45: 564-655. 1911. 

Trees, notable for their leafless, tortuous branchlets, the alternate, 
3-foliolate or unequally pinnate leaves crowded at their tips. Petioles 
imbricate at the broad, membranous, vaginate base. Racemes 
slender, loosely flowered, pendulous, the flowers bracteate. Stamens 
5 to many, with short, glabrous filaments, the anthers often pilose. 
Carpel 1, included in the persistent, angled, spinose or winged, 
turbinate calyx tube. The chestnut or red-brown, thin bark of the 
flowering branches, which soon exfoliates, is a conspicuous feature 
of these trees. 

Leaflets 7-8 pairs, about 3 cm. long P. multijuga. 

Leaflets 3-5 (1-6) pairs, often smaller. 

Leaflets densely appressed-pilose beneath and little or not at all 

tomentose. 

Leaflets 5 pairs, serrate P. serrata. 

Leaflets 3-4 pairs, entire, or the leaves 3-foliolate. 

Leaflets lustrous above, white-pubescent beneath . P. albicans. 

Leaflets dull above, sordid-pubescent beneath . P. subsericans. 

Leaflets more or less densely tomentose or also rarely pilose or 
glabrate but, if pilose, the hairs not appressed. 

Leaflets barely 10 mm. long, usually shorter .... P. tarapacana. 
Leaflets 11 mm. long or more, at least some of them. 
Leaflets entire, often retuse. 
Leaflets pilose and tomentose beneath, dull above. 

P. subsericans. 
Leaflets densely tomentose beneath, lustrous above. 

P. Weberbaueri. 

Leaflets distinctly, but sometimes minutely, crenate-serrulate, 
at least toward the apex. 

Pubescence of the leaflets beneath dense or sparse but 
loosely tomentose or villous; leaflets often 2-3 cm. long. 



FLORA OF PERU 1091 

Leaflets 3 pairs, slightly villous beneath .... P. racemosa. 
Leaflets mostly 3. 
Leaflets less than 1.5 cm. long, densely villous- tomen- 

tose beneath P. rugulosa. 

Leaflets 2-3 cm. long, slightly villous P. villosa. 

Pubescence of the leaflets beneath compact-tomentose, 
sometimes fine, or, if partly pilose, appressed; leaflets 
often 1.5 cm. long. 

Plants erect shrubs or trees, 1 to several meters high. 

Pubescence tomentose P. incana. 

Pubescence sericeous P. tomentella. 

Plants low, 50 cm. high, with spreading branches. 

P. subquinquefolia. 

Polylepis albicans Pilger, Bot. Jahrb. 37: 535. 1906; 605. 

A shrub, up to 3 meters high; leaves unequally pinnate; leaflets 
3 pairs, linear-elliptic, entire or slightly crenulate toward the rounded, 
incised-emarginate apex, somewhat unequal at the base, to 2 cm. 
long, finally glabrous above, beneath and marginally densely pubes- 
cent with long, white, sericeous hairs; inflorescence densely yellowish- 
sericeous- villous, the ovate, acuminate bracts 4 mm. long; sepals 
3 or 4, rotund-ovate, 3 mm. long; stamens 10-12. Neg. 3383. 

Ancash: Above Huaraz, Weberbauer 3229, type. "Quinar," 
"quinual." 

Polylepis incana HBK. Nov. Gen. & Sp. 6: 227. 1824; 639. 
P. incana subsp. icosandra Bitter, Bot. Jahrb. 45: 641. 1911. P. in- 
cana subsp. micranthera Bitter, op. cit. 642. P. incana subsp. incarum 
Bitter, op. cit. 643. P. pallidistigma Bitter, op. cit. 645. 

A densely branched shrub or small tree with trifoliolate leaves, 
the subcoriaceous leaflets oblanceolate or oblong, crenate, obtuse or 
somewhat emarginate, glabrous and almost dull above, or sometimes 
lustrous, more or less densely tomentose with undulate hairs beneath, 
to 3 cm. long, 9-11 mm. wide, usually about half as large; stamens 
about 20, rarely as few as 5, the anthers pilose, especially toward 
the apex; fruit 4 mm. long, gibbous or short-aculeate. A variable 
species. Bitter's subsp. micranthera is P. tomentella Wedd., fide 
Fritsch. P. pallidistigma may be distinct, at least varietally, from 
both P. incana and P. tomentella by its obovate leaflets, these about 
2 cm. long, 8-10 mm. wide. Neg. 3386. Used to tan hides because 
of the considerable amount of tannin it contains (Herrera). 



1092 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Junin: Near Cerro dePasco, KanehiraS. Tarma, Killip & Smith 
21899. Ancash: Near Tallenga, Weberbauer 2886 (subsp. icosandra). 
Huanuco: Caxamarquilla, Haenke (subsp. micranthera) . Llata, a 
common tree of stream banks, 2,000 meters, 2249 (det. Killip as 
subsp. incarum). Chiclin, Sawada P83. Puno: Lake Titicaca, 
Jardin del Inca, Salar 148 (subsp. incarum). Munani, 3,600 meters, 
Weberbauer 1369 (type of P. pallidistigma) . Puno, Soukup 365. 
Ayacucho: Chavina, Weberbauer 5776. Totorabamba, Weberbauer 
5487. Lima: Rio Blanco, Killip & Smith 21745. Cuzco: Paucar- 
tambo Valley, H err era 1129. Pisac, Pennell 13731. Bolivia to 
Colombia. "Manzanita," "cceuiia," "queuna." 

Polylepis multijuga Pilger, Bot. Jahrb. 37: 536. 1906; 588. 

A shrub or tree, to 8 meters in height, with yellowish, long- 
pubescent sheaths and leaves; leaflets oblong, rounded at the base, 
the obtuse apex not at all emarginate, remotely serrate, 2-5 cm. 
long, finally glabrous above; racemes to 25 cm. long, pendent, the 
rachis yellowish-villous; bracts 5-8 mm. long; sepals rounded-ovate, 
3 mm. long, the pubescence at the subacute apex hirsute; stamens 
10-12, the anthers pilose. Neg. 27496. 

Cajamarca: Toward Hualgayoc, 2,700 meters, Weberbauer 4098, 
type. Amazonas: Chachapoyas, 3,000 meters, Weberbauer 4429. 

Polylepis racemosa R. & P. Syst. 139. 1798; 617. 

Branchlets thick; lower sheaths about 1 cm. broad, densely long- 
pilose, especially at the margins; petioles 1.5-2 cm. long; leaflets 3 
pairs, scarcely coriaceous, broadly lanceolate or obovate, unequally 
crenate, revolute, glabrous above, prominently reticulate-veined and 
densely pilose beneath, to nearly 3 cm. long and 1.5 cm. wide; ra- 
cemes becoming 10 cm. long or longer, few-flowered, the rachis short- 
pilose, the bracts lanceolate, long-acuminate, to 7 mm. long; sepals 4, 
greenish, unequal; stamens 18-20, the filaments 3 mm. long; fruit 
obconic, about 5 mm. long, 4 mm. broad, more or less pubescent, 
each of the 4 angles terminating in a short spine. The wood is much 
used for fuel, the ashes for cleaning and bleaching cloth, and the 
wood was used in mines. According to Weberbauer, the tree is 
planted for the same purposes, but Herrera says it is used only for 
fuel, the wood being inferior. Neg. 3392. 

Lima: San Buenaventura, Nee. Junin: Cacray, 3,360 meters, 
Weberbauer 220 (det. Bitter). Tarma, Isern 2094. Cuzco: Gay 588, 
pro parte (det. Bitter). In all Cuzco (Herrera). Without locality, 
Humboldt3141. "Cceufia," "quinuar," "queuna." 



FLORA OF PERU 1093 

Polylepis rugulosa Bitter, Bot. Jahrb. 45: 638. 1911. 

Sheaths densely pilose, the leaves 3-foliolate, the leaflets oblong or 
obovate, obtuse, crenate, rugulose above, finally glabrescent, densely 
cinereous- tomentose, even in age, 10-13 mm. long, 5.5-6 mm. wide; 
racemes finally 5-6 cm. long, the slender rachis densely pubescent, 
the pilose, lanceolate, acuminate bracts 5-7 mm. long; sepals in fruit 
4 mm. long, densely pilose below, densely papillose on the thin mar- 
gins and apex; stamens 10-15, the filaments to 5 mm. long, the 
purple anthers densely pubescent; fruit obconic, 5-6 mm. long, 
gradually attenuate to the base, densely pubescent, the angles 
aculeate and usually gibbous only between the angles. A shrub 
about 3 meters high, separated by the author from P. tomentella by 
the densely pilose sepals and anthers. 

Arequipa: Pampa de Arrieros, 3,700 meters, Weberbauer 4881, 
type. Nevado de Chachani, Pennell 13295. Moquehua: Above 
Torata, Weberbauer 7470. 

Polylepis serrata Pilger, Bot. Jahrb. 37: 536. 1906; 591. 

Similar to P. multijuga, but the leaves shorter, to 8 cm. long, 
the leaflets fewer, in age often glabrescent, even beneath, the racemes 
to 20 cm. long, and the stamens only 5-7, the larger anthers glabrous. 
-Neg. 3394. 

Huanuco: Near Monzon, 3,400 meters, Weberbauer 3354, type. 
Cuzco: Above Yanamanche, 3,800 meters, Weberbauer 4954- 
Marcapata, 2,900 meters, Weberbauer 7803. "Qqueuna," "queuria." 

Polylepis subquinquefolia Bitter, Bot. Jahrb. 45: 636. 1911. 

A low shrub with procumbent-ascending branches, to about 50 
cm. high; petioles 5-8 mm. long; leaves usually 5-foliolate, the leaflets 
broadly oblong or obovate, obtuse, often glabrescent and more or 
less lustrous above, densely cinereous-tomentulose beneath, crenate 
above the middle, the oblique, sessile lateral leaflets to 17 mm. long 
and 8 mm. wide, the petiolule of the slightly larger terminal leaflets 
3^4 mm. long; stamens 8-9, the anthers densely pilose, like the style; 
fruit densely pubescent. Neg. 3396. 

Puno: Cuyoenyo, 3,600 meters, Weberbauer 931, type. 

Polylepis subsericans Macbr. Candollea 5: 367. 1934. P. incana 
HBK. var. flavipila Bitter, Bot. Jahrb. 45: 640. 1911. 

Basal internodes naked, as much as 3.5 cm. long; leaflets at first 
appressed-pilose on both sides, later glabrate above, permanently 



1094 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

yellowish-strigose-pilose beneath; leaflets 16-17 mm. long, 4-6 mm. 
wide, typically entire or 2-4-crenate toward the apex, or the cre- 
nations sometimes more numerous. 

Ayacucho: Hacienda Totorabamba, Weberbauer 5487, type. 
Above Pisco, on stony slopes, Weberbauer 51+33 (type of the variety). 

Polylepis tarapacana Philippi, Viaje Tarapaca 21. 1891; 651. 

Stoutly branched, with many crowded, appressed branchlets, 
these densely clothed with the petiole bases; petioles 6 mm. long, 
within at the base almost shaggy-villous, on the outer edges (like the 
trifoliolate leaves beneath) yellow- tomentulose; leaflets barely 10 mm. 
long, 4 mm. wide, obscurely crenulate toward the apex, puberulent 
above; sepals 3, broadly lanceolate, unequal, pilose; stamens 8-13, 
the anthers glabrous at the base, densely pilose toward the tip. 
Illustrated, Bitter, Bot. Jahrb. 45: pi. 10. 

Tacna: Tacora, Weddell; Philippi; Pentland; Werdermann 1143. 
Bolivia. "Quinual," "quinoa." 

Polylepis tomentella Wedd. Chlor. And. 2: 237. pi. 78. 1857. 

A sprawling shrub, the subprostrate branches clothed with 
lustrous, exfoliating, reddish bark; branchlets short, flexuous, ascend- 
ing to erect, at first puberulent-tomentose, like the petioles, leaves, 
perianth, and racemes; petioles short- villous on one side and at the 
enlarged base; leaflets 3, mostly 9 mm. long and 4 mm. wide, or even 
15 mm. long and 6 mm. wide, minutely denticulate toward the apex, 
soon glabrous and dull above, yellowish-puberulent, tomentulose 
beneath; racemes 1-2 cm. long, glabrous or glabrate in age, even to 
the rotund, cuspidate calyx lobes. Neg. 34733. 

Cuzco: Gay. Ayacucho: Chavina, Weberbauer 5776 (var.; det. 
Bitter). 

Polylepis villosa HBK. Nov. Gen. & Sp. 6: 228. 1824; 637. 

A shrub, the branches with long internodes; petioles 1-2.5 cm. 
long, densely brown- villous like the persistent sheaths; leaves mostly 
trifoliolate, the leaflets as much as 5.5 cm. long and 1.7 cm. wide, in 
the type 3 cm. long, 1 cm. wide, oblong, rounded-obtuse, in the type 
retuse, the veins impressed above, prominently reticulate beneath; 
sepals broadly ovate or lanceolate, 5 mm. long, pubescent outside; 
stamens about 20, the anthers villous; fruiting calyx villous, the turbi- 
nate tube 6-8-angled and dentate, spinose above. The Ruiz and 
Pa von and the Haenke specimens are P. incana HBK. var. pri- 



FLORA OF PERU 1095 

movestita Bitter and var. connectens Bitter, Bot. Jahrb. 45: 645, 
respectively, both of which seem better referable to P. villosa. Neg. 
3398. 

Cajamarca: Bonpland 3685, type. Huancayo, Esposto (det. 
Pilger as P. racemosa). Huanuco: Haenke. Caxamarquilla, Ruiz & 
Pav6n. Cuzco: Gay (det. Bitter). 

Polylepis Weberbaueri Pilger, Bot. Jahrb. 37: 535. 1906. 

Branchlets short, only the young sheaths villous; leaflets 3, 
rarely 2-4 pairs, distinctly emarginate, elliptic-obovate, 12-17 mm. 
long, lustrous and finally glabrous above, densely short-tomentose 
beneath; racemes longer than the leaves, the bracts clasping, to 6.5 
mm. long; sepals broadly ovate, in fruit 5 mm. long. Neg. 3399. 

Ancash: Yungai, 3,700 meters, Weberbauer 3287, type. "Quinar," 
"quhmal." 

8. QUILLAJA Molina 
Smegmadermos R. & P. Prodr. 144. pi. 31. 1794. 

Similar to Kageneckia, but the leaves subentire. Flowers 3-5, on 
axillary and terminal peduncles, polygamo-dioecious, the calyx lobes 
valvate. Disk fleshy; stamens 10. Carpels coherent. 

Quillaja brasiliensis (St. Hil. & Tul.) Mart. Syst. Mat. Med. 
127. 1843. Fontenella brasiliensis St. Hil. & Tul. Ann. Sci. Nat. II. 
17 : 142. pZ. 7. 1842. 

Glabrous; leaves lustrous above, oblong-lanceolate, acute at each 
end, obscurely and remotely sinuate-dentate or entire, 5-10 cm. long, 
12-22 mm. wide; petals scarcely longer than the spreading calyx 
lobes; disk deeply 5-lobed. Used like soap, and has the property of 
removing grease from dyed cloth without affecting the color (Herrera) . 
In all the valleys of the Department of Cuzco ( Herrera) ; but surely 
as an introduction. The generic name has been written Quillaia. 

Cuzco: Prov. Convencion and Calca (Herrera). Uruguay; Brazil. 
"Quillai." 

9. KAGENECKIA R. & P. 

Evergreen shrubs or trees, the more or less glutinous leaves 
crowded toward the tips of the branches, minutely serrate. Stipules 
caducous. Flowers unisexual, the pistillate solitary, the staminate 
in short racemes or corymbs. Calyx persistent, the lobes imbricate. 
Stamens 10-20, in 1 series. Disk slender, free. Carpels 5, tomentose, 
stellately spreading, many-seeded. 



1096 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Kageneckia lanceolata R. & P. Syst. 290. 1798. K. glutinosa 
HBK. Nov. Gen. & Sp. 6: 237. 1824. K. amygdalifolia Presl, Epim. 
Bot. 195. 1849. Osteomeles Pentlandiana Dene. Nouv. Arch. Mus. 
Paris 10: 183. 1874. 

Branches elongate, bearing many short, densely leafy flowering 
branchlets, these somewhat resinous toward the tips; leaves lanceo- 
late or oblanceolate, long-cuneate to the base, rounded or short- 
acute at the apex, usually about 3.5 cm. long and 1 cm. wide, some- 
times twice as broad or even broader, finely glandular-serrulate from 
the base to the apex; flowers about 7 mm. wide, corymbose, few, on 
pedicels to 7 mm. long; calyx lobes acute, slightly pilose, especially 
marginally, rarely throughout; fruits solitary, densely brown- villous- 
tomentose, star-shaped at maturity, each of the 5 spreading carpels 
suggesting a somewhat one-sided, Napoleonic hat, 10 mm. long, 8 
mm. wide above; seeds 11 mm. long, the wing oblique at the base, 
7 mm. long. Perhaps more than one species, or at least variety, 
can be distinguished. K. oblonga R. & P., of Chile, may be the same, 
but typically it has larger leaves, less finely serrulate and more 
abruptly cuneate to a well defined petiole. An infusion of the bitter 
bark and leaves of the tree was used in the treatment of fevers 
(tercianas), according to Ruiz and Pa von. The wood, because of 
its toughness, according to Herrera, is a favorite one for walking 
sticks and for supports of the hanging bridges. The leaves furnish 
a black dye. 

Cajamarca: Jae"n de Bracamoros, Humboldt. Ramblan, 2,100 
meters, Weberbauer 4804; 190. Ayacucho: Chavina, Weberbauer 
5780. Above Pisco, Weberbauer 5425. Rio de Lomas, Weberbauer 
5769. Lima: Chosica, Weberbauer 5722. Above Lima, Castelnau. 
Canta, Ruiz & Pavdn. Huanuco: Maria del Valle, 3549 (det. 
Killip). Llata, 2281, 2295,Cuzco: Gay. San Sebastian, 3,300 
meters (Herrera 1515). Vilcanota, Pennell 14190. Urubamba 
Valley (Weberbauer, 175). Bolivia. "Uritumicuna," "lloque," 
"loque." 

10. RUBUS L. 

More or less suffrutescent and usually prickly shrubs, some of 
the Peruvian species prostrate. Calyx lobes persistent, ebracteolate. 
Stamens and carpels many, the drupes not included in the calyx. 

Creeping or spreading plants with conspicuous, usually broad stipules. 
Leaves all simple. 

Stipules narrowly lanceolate; petioles very short. . .R. extensus. 



FLORA OF PERU 1097 

Stipules broadly ovate; petioles long. 
Leaves not at all or only slightly bullate-reticulate. 
Leaflets glabrous beneath or sparsely pubescent on the 

nerves R. coriaceus. 

Leaflets more or less densely pubescent beneath. 

R. acanthophyllus. 

Leaves conspicuously bullate-reticulate R. betonicifolius. 

Leaves all or nearly all 3-foliolate. 

Leaves white-pubescent beneath, the leaflets rarely over 2.5 cm. 
wide R. Weberbaueri. 

Leaves glabrate beneath, at least not white-pubescent, the 
leaflets usually more than 2.5 cm. wide. 

Leaves softly pubescent beneath R. nubigenus. 

Leaves usually glabrate beneath. 

Petiole longer than the terminal petiolule R. roseus. 

Petiole shorter than the terminal petiolule. . . . .R. andicola. 
Tall, erect or clambering plants with linear-subulate stipules. 
Branchlets, petioles, and inflorescences abundantly stipitate- 

glandular. 

Leaves glabrous on the nerves beneath or slightly pilose, lustrous. 

R. erythrocladus. 
Leaves finely pilose, at least beneath, dull. 

Leaves glabrous above R. adenothallus. 

Leaves pubescent above R. bogotensis. 

Branchlets, petioles, and inflorescence glabrous to setose, little if 
at all glandular (except sometimes in R. robustus var.). 

Plants more or less densely reddish-setose R. urticaefolius. 

Plants glabrous, puberulent, or villous. 
Plants puberulent or villous. 

Inflorescences terminal, usually many-flowered; pubescence 
villous-tomentose R. robustus. 

Inflorescences lateral, loosely few-flowered; pubescence 
spreading-villous R. sparsiflorus. 

Plants glabrous and glaucous. 

Leaves closely white-tomentose beneath R . glaucus. 

Leaves glabrate or slightly crisp-pubescent beneath. 

R. megalococcus. 



1098 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rubus acanthophyllus Focke, Abh. Nat. Ver. Bremen 4: 161. 
1874. R. Jelskii Fritsch in Szyszy. Diss. Cl. Math. Phys. Acad. 
Cracov. 29: 220. 1894. 

Prostrate or matted, the flowering branches more or less pilose- 
hirsute and subulate-aculeate; petioles 1-3 cm. long; stipules broad, 
mostly dentate; leaves suboblong or oblong-ovate-lanceolate, some- 
times 3-lobed, unequally crenate-serrate, softly pilose-tomentose 
above and more densely white-tomentose beneath, there often 
aculeate on the nerves; flowers almost 2 cm. broad, borne toward 
the ends of the branches on tomentose, stipitate-glandular peduncles; 
sepals 8 mm. long, the purple petals longer; stamens and style gla- 
brous, the numerous carpels villous. Possibly a pubescent state of 
R. coriaceus. 

Amazonas: Chachapoyas, 3,300 meters, Weberbauer 4424' 
Cuzco: Yanamanche, 4,400 meters, Weberbauer 4943. Huanuco: 
Monzon, 3,500 meters, Weberbauer 3308. Mito, 1827. Huanuco, 
2176. Cajamarca: Cutervo, Jelski 6. 

Rubus adenothallus Focke, Medd. Rijks Herb. 19: 56. 1913. 

Branches, branchlets, petioles, leaves beneath, peduncles, and 
calyces densely purple-glandular; lower leaves with 5 leaflets; petioles 
10 cm. long, moderately aculeate; terminal petiolule 4 cm. long, the 
lower 1 cm.; leaflets subcoriaceous, lanceolate, about 10 cm. long, 
3-3.5 cm. wide, minutely mucronate-serrate, truncate at the base, 
long-acuminate, beneath glandular-punctate and puberulent; flower- 
ing branches angled; stipules filiform; principal terminal inflorescence 
rather dense; sepals almost 1 cm. long, the suborbicular petals little 
longer. Illustrated, loc. cit. Neg. 27501. 

Ayacucho: Osno, 3,500 meters, Weberbauer 5559 (det. Focke). 
Huanuco: Vitoc, Isern 2358 (smaller-flowered; toward R. bogotensis). 
Mito, 1841. Yanano, 3735. Cuzco :Gay. Bolivia. 

Rubus andicola Focke, Bibl. Bot. 72: 36. 1910. 

A creeping plant with long, pilose branches, armed with short, 
recurved prickles; terminal petiolule 2-4 cm. long, or usually 3-4 
times longer than the petiole; leaflets subcoriaceous, elliptic, acute, 
about 10 cm. long and 6 cm. wide, serrate, pilose above, appressed- 
pubescent beneath; stipules dentate, equaling or longer than the 
petioles; inflorescence lax, the peduncles villous, aculeate; sepals lance- 
olate, often caudate, villous at the base; petals longer, purple; stamens 
and styleglabrous; carpels villous. Illustrated, op. cit. 35. Neg. 18026. 

Amazonas: Moyobamba, Weberbauer 4748. Ecuador. 



FLORA OF PERU 1099 

Rubus betonicifolius Focke, Bibl. Bot. 72: 33. 1910. 

A vine, readily known by its cordate-ovate, rarely 3-lobed, bul- 
late-rugose leaves, as much as 7 cm. wide, coarsely and unequally 
dentate, more or less pilose on the nerves beneath; stipules broadly 
ovate-serrate, much shorter than the petioles, which are 3-4 cm. long; 
flowers axillary or terminal and corymbose, the hirsute peduncles 
densely aculeate, sometimes glandular; calyx setose-aculeate or 
unarmed ; filaments and style filiform. Neg. 18027. 

Puno: Sandia, Weberbauer 670. Bolivia. "Nuto-karikari." 

Rubus bogotensis HBK. Nov. Gen. & Sp. 6: 220. 1824. R. 
porphyromallos Focke, Repert. Sp. Nov. 9: 235. 1911(?). 

A robust plant, commonly stipitate-glandular throughout; 
prickles on fertile branches small and few, or lacking in the inflores- 
cence, numerous on the long petioles and petiolules; leaflets mem- 
branaceous, densely serrate, densely pubescent above in age, sparsely 
so, glandular, and pilose beneath, especially on the nerves, broadly 
ovate-elliptic, subcordate, abruptly short-acuminate, 4-7 cm. wide, 
or in the Peruvian plants often narrower and gradually acuminate; 
inflorescence elongate; flowers sometimes almost 2 cm. wide; petals 
nearly equaled by the sepals or exceeded by them, the sepals then 
caudate, to 12 mm. long; carpels glabrous, 6 mm. long. The Peru- 
vian plants may constitute a variety. Neg. 34745. 

Huanuco : Tambillo, 3569. Yanano, 4942. Junin : Huacapistana, 
Weberbauer 2190. Carpapata, 2,700 meters, Killip & Smith 24384. 
Huasa-huasi, Ruiz & Pavon; Dombey (det. Bolle). Ayacucho: 
Carrapa, 2,800 meters, Killip & Smith 22285; Weberbauer 7561. 
Cuzco: Urubamba (Herrera 1552). Valle de San Miguel, Machu- 
picchu, Herrera 2010. Puno: Puno, Soukup 323. 

Rubus coriaceus Poir. Encycl. 6: 237. 1804. 

Similar to R. acanthophyllus, but the coriaceous leaves glabrous 
beneath, or the nerves hirsutulous; sepals and petals subequal. 
Illustrated, Bibl. Bot. 72: 32. Negs. 18025, 34744. 

Junin: Palca, Ruiz; Dombey; Weberbauer 2477. Colombia; 
Ecuador. 

Rubus erythrocladus Mart. Fl. Bras. 14, pt. 2: 62. 1867. 

Allied to R. adenothallus ; branchlets densely red-setose; leaves 
lustrous, mostly with 5 oblong-lanceolate, acuminate leaflets only 2-3 



1100 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cm. wide, typically glabrous, but sometimes puberulent on the nerves 
beneath; flowers often smaller. 

Junin : Palca, Weberbauer 24-98 (det. Focke) . Brazil. 

Rubus extensus Fritsch in Szyszy. Diss. Cl. Math. Phys. Acad. 
Cracov.29:221. 1894. 

A slender-stemmed plant, well characterized by its mostly lanceo- 
late stipules; stems, petioles, and leaves beneath more or less tomen- 
tose; leaflets ovate-elliptic, evenly serrulate, 1.5-4.5 cm. wide, on 
petiolules 7-18 mm. long. Perhaps referable to R. loxensis Benth. 
Neg. 18028. 

Cajamarca : Chonta Cruz, Jelski 3. Cutervo (Jelski) . 

Rubus glaucus Benth. PI. Hartweg. 173. 1845. 

Flowering branches elongate, glabrous and glaucous, pruinose, 
sparsely aculeate; leaves all 3-foliolate, the leaflets ovate-lanceolate, 
long-acuminate, irregularly mucronate-serrulate, glabrous above, 
white-tomentose beneath; inflorescence terminal, racemose-panicu- 
late, short, some of the flowers axillary; pedicels long, divaricate, 
more or less aculeolate; sepals ovate-cordate, mucronate, glabrate; 
petals ovate, white; carpels tomentulose. 

Huanuco: Vilcabamba, 5199. Ranging to Costa Rica. 

Rubus megalococcus Focke, Abh. Nat. Ver. Bremen 4 : 175. 1874. 

A nearly glabrous, glaucescent shrub, similar to R. glaucus, but 
the inflorescence leafy and interrupted, or the flowers axillary and 
racemose, few; pedicels puberulent, somewhat glandular and more or 
less aculeolate; flowers large, nodding, the roseate petals longer than 
the triangular-lanceolate sepals; carpels many, glabrous, the fruit 
almost 8 mm. long. 

Junin: Palca, 3,400 meters, Weberbauer 2503. Bolivia. 

Rubus nubigenus HBK. Nov. Gen. & Sp. 6: 220. 1824. 

More or less villous, stipitate-glandular, prickly; petioles 3-4 cm. 
long; leaflets petiolulate, oblong-elliptic, 3-6 cm. long, 3-4 cm. wide, 
merely acute, crenate-serrate, dull and appressed-pilose or tomentose 
above, more or less canescent-velutinous beneath; stipules and bracts 
ovate-reniform; inflorescence a narrow, interrupted panicle; flowers 
2-3 cm. wide, the roundish, purplish petals little longer than the 
sepals; carpels villous. The type has leaves spreading-pilose only 
on the prominent nerves and veins beneath. The following speci- 
mens appear to be too near R. roseus, but the Ruiz and Pavon speci- 



FLORA OF PERU 1101 

men at Madrid was determined by Bolle as R. glabratus HBK. This 
species, as to type, has the leaves all 3-foliolate, glabrous, dull and 
rugose above, scarcely acute, to 4 cm. long and 2 cm. wide, or much 
smaller; petioles 2-3 cm. long; petiolules a few millimeters long; 
prickles few; sepals broadly ovate, abruptly acute, 8-10 mm. long; 
petals as long as the sepals; flowers solitary; pedicels 1.5-2 cm. 
long. Neg. 34748. 

Huanuco: Mito, 1995. Junin: Vitoc, Isern 2356, 2206 (leaflets 
5 cm. wide; cf. R. macrocarpa) ; Martinet (leaflets 2.5 cm. wide). 
Without locality, Ruiz & Pavdn (or R. glabratus; see note above). 
Bolivia; Ecuador. 

Rubus robustus Presl, Epim. Bot. 196. 1849. R. floribundus 
HBK. Nov. Gen. & Sp. 6: 219. pi. 557. 1824, non Weihe. R. helio- 
scopus Focke, Bot. Jahrb. 54: Beibl. 117: 41. 1916. R. boliviensis 
Focke, Abh. Nat. Ver. Bremen 4: 158. 1874. R. peruvianus Fritsch 
in Szyszy. Diss. Cl. Math. Phys. Acad. Cracov. 29: 220. 1894. 

Robust, arcuately and abundantly branched; branches and leaf- 
lets beneath commonly conspicuously villous-tomentose; petioles and 
petiolules more or less villous-tomentose, recurved-aculeate; leaflets 
elliptic-oblong, acuminate, appressed-pubescent above, the nerves 9- 
12 (-15); inflorescence terminal, sometimes narrow, often much 
branched and leafy below; sepals and petals subequal, the sepals 
often a little longer; carpels usually many, glabrous. Variable in 
pubescence, size of flowers, and number of carpels (see Field Mus. 
Bot. 8: 118. 1930). The HBK. type is much less pubescent. Possibly 
varieties are distinguishable. The plant is typically esetose. My 
numbers 3894, 1218, and 5200, with glandular or nonglandular 
setae, suggestive of hybridization with R. bogotensis or R. urticaefolius, 
were described as R. floribundus var. nimbatus Macbr. loc. cit., 
but should become R. robustus Presl, var. nimbatus Macbr., 
comb. nov. 

Junin: Palca, 700 meters, Weberbauer 1801, 643 (det. Focke); 
Isern 2199. Puno: Sandia, Weberbauer 833 (det. Focke). Hua- 
nuco: Near Mufia, 3894- Yanahuanca, 1218. Chanchamayo, Isern 
2309. Vitoc, Isern 2359. Cani, 3410. Mito, 1405, 1939. Vilca- 
bamba, hacienda on Rio Chinchao, 5200, 5177. Ayacucho: Prov. 
Huanta, Weberbauer 5580 (type of R. helioscopus) . Piura: Ayavaca, 
Weberbauer 6388 (det. Focke). Cuzco: Gay. Cerro de Cusilluyoc, 
Pennell 14088. Without locality: Haenke, type. Cajamarca: 
Celendin, Woytkowski. 



1102 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rubus roseus Poir. Encycl. 6: 245. 1804. 

Typically glabrous, or in the varieties more or less hirsute- 
villous and the peduncles often glandular; upper leaves simple; 
petioles 2-2.5 cm. long; leaflets petiolulate, elliptic, serrate; flowers 
1 to several and panicled, on short or long pedicels; stipules elongate, 
dentate; calyx often setose; flowers 2-3 cm. broad; fruit large, the 
carpels glabrous to villous. A polymorphic species of which the 
following recorded varieties occur in Peru: var. rosaeflorus Hook. 
Icon. 1: pi. 46. 1837, more or less hirsute; var. santarosensis (Kuntze) 
Macbr. Field Mus. Bot. 8: 118. 1930, also hirsute, with smaller flowers 
and more dentate stipules; var. Lechleri (Focke) Macbr. Field Mus. 
Bot. 8: 118. 1930, likewise hirsute but with short-pedicellate, more 
or less paniculate flowers. The last might be reduced to R. nubigenus 
or considered a variety of that species. The acid-sweet fruits are 
used for making refreshing drinks (Herrera). Neg. 34750. 

Huanuco: Pillao, Ruiz & Pavon (det. Bolle). Mufia, 4288 (var. 
santarosensis), 4286 (var. rosaeflorus). Churupallana, Dombey. 
Huanuco, 2143. Vitoc, Isern 2357 (var. Lechleri), 2363, 2355 
Amazonas: Molinopampa, 2,300 meters, Weberbauer 4375. Cuzco: 
Paucartambo, Vargas 324; Soukup 399. Yanamanche, Weberbauer 
4967. Cerro de Cusilluyoc, Pennell 14104, 14123, 13831 (var. 
Lechleri). Cuzco, Herrera 1. Puno: Sandia, Weberbauer 674- 
Agapata, Lechler (type of var. Lechleri). "Cjari-cjari," "zarzamora," 
"chilifruta." 

Rubus sparsiflorus Macbr. Field Mus. Bot. 8: 117. 1930. 

Moderately robust, the more or less prickly branches and petioles 
soft- villous; petioles slender, mostly 6-10 cm. long; stipules almost 
1 cm. long, persistent; leaflets abruptly acuminate, evenly serrulate, 
mostly 6 cm. long and 3.5 cm. wide, glabrate above, subappressed- 
villous beneath, especially on the nerves; racemes axillary and 
terminal, 2-4 cm. long, 3-6-flowered; flowers 1-1.5 cm. wide, deep 
rose; pedicels scarcely 5 mm. long, densely rusty- villous, eglandular; 
sepals pilose on both sides; carpels glabrous or sparsely pilose at 
the apex. 

Ayacucho: Pampalca, 3,200 meters, Killip & Smith 23238. 
Junin: Vitoc, Martinet 1321. Huanuco: Mito, 1674, type. Caja- 
marca: San Miguel, Weberbauer 3940. 

Rubus urticaefolius Poir. Encycl. 6: 246. 1804. 
Well characterized by the widely spreading, reddish setae that 
more or less abundantly clothe the remotely aculeate stems; stipules 



FLORA OF PERU 1103 

filiform; leaflets sometimes 5, pubescent above, white- tomentose 
beneath; inflorescence ample, becoming open, to 30 cm. long and 10 
cm. wide; flowers small, short-pediceled, the fruiting pedicels 5-10 
mm. long; sepals white-pubescent, scarcely setigerous, little shorter 
than the petals; carpels glabrous, the edible fruit globose, small. 
The leaves on young shoots may be simple and broadly elliptic. 
Neg. 34751. 

San Martin: San Roque, Williams 7157, 6971. Near Moyo- 
bamba, Klug 3585. Junin: Huacapistana, Killip & Smith 24.104. 
Pichis Trail, 350 meters, Killip & Smith 25449. Chanchamayo, Diehl 
2459; Schunke 41 3. Huanuco: Yanano, 3757. Monzon, Weberbauer 
3362,Ayacucho: Aina, Killip & Smith 23105, 22561. Carrapa, 
Killip & Smith 22333. Ancash: Supa-acros, Weberbauer 2647. 
Lima: Barranco, Weberbauer 17. Lima, Ruiz & Pavdn; Dombey. 
Apurimac : Abancai, Vargas 467. Cuzco : Urubamba, 3,000 meters 
(Herrera 1524). South America. "Cjari-cjari." 

Rubus Weberbaueri Focke, Bibl. Bot. 72: 38. /. 11. 1910. 
R. nubigenus HBK. subsp. Ruizii Focke, Abh. Nat. Ver. Bremen 4: 
162. 1874. 

Branchlets and petioles closely cinereous-tomentulose and some- 
what glandular and aculeate; leaves 3-foliolate, the firm leaflets 
5-9 cm. long, 2-2.5 (-4) cm. wide, subequally serrulate, dull and 
green above, compactly white-tomentose beneath, the terminal one 
distinctly petiolulate; sepals lance-ovate, sometimes dentate; corymbs 
interrupted, few-flowered; pedicels hirsute- tomentose, aculeate; 
flowers about 2 cm. wide; petals longer than the unarmed calyx; 
fruit elongate, villous. Neg. 18037. 

Huanuco : Monzon, 3,200 meters, Weberbauer 3362, type. Pillao, 
Ruiz & Pawn. Tambo de Vaca, 4393. 

11. GEUM L. 
Reference: Bolle, Repert. Sp. Nov. Beih. 72: 1-119. 1933. 

Low, rhizomatous, herbaceous perennials with unequally pinnate, 
crowded, subsessile leaves, those of the stems few and often 3- 
foliolate or reduced. Flowers often corymbose, rather large, with 5 
broad petals usually exceeding the bracteolate calyx lobes, the calyx 
tube persistent. Stamens many, the filaments filiform. Achenes 
many, the receptacle short or clavate, the styles straight or geniculate, 
slender, elongate. 



1104 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Geum peruvianum Focke, Bot. Jahrb. 37: 540. 1906. 

Stems spreading- villous below, hirsute and branched above, tall; 
leaves hirsute throughout, the lower ones interrupted-pinnate, the 
terminal leaflet much larger, obtusely crenate-lobed, the lateral 
ones very unequal, subelliptic, incised-crenate; cauline leaves sub- 
sessile, 3-foliolate or deeply trifid, the narrower divisions incised- 
serrate, the terminal one often 3-lobed; flowers 1-1.5 cm. broad or 
smaller; petals suborbicular, yellow; carpels densely sericeous- villous, 
about as long as the upper segment of the style, nearly or quite gla- 
brous. This is G. magellanicum of authors, in part. Neg. 3616. 

Cajamarca: Hualgayoc, 3,500 meters, Weberbauer 4007, type; 
Raimondi. Ayacucho: Prov. Huanta, Weberbauer 5571. Huanuco: 
Tambo de Vaca, 4454- Mito, 1401. Ecuador; Colombia. 

Geum Quellyon Sweet, Brit. Fl. Gard. 1: 3. pi. 292. 1829. 
G. chiloense Balb. ex Ser. in DC. Prodr. 2: 551. 1825, in syn. 

Similar to G. peruvianum, but the lower leaves elongate; upper 
articulation of the style hirsute; petals deep orange to red. The 
root has the odor of cinnamon (Ruiz & Pawn}. In Peru introduced. 
Illustrated, Feuill. PI. Med. Perou Chil. 2: pi. 27. 

Cajamarca: In garden at Hacienda Tahoma, Hualgayoc, Weber- 
bauer 4089 (det. Focke). Junin: Huasa-huasi, Ruiz & Pavdn (det. 
Bolle). Chile. "Canelillo." 

12. FRAGARIA L. 

Technically the genus is distinguished from Potentilla only by 
the fleshy receptacle and by the many carpels. The genus Duchesnea 
Smith, with yellow flowers and less succulent receptacle, is included 
here for convenience. 

Petals yellow; receptacle not juicy F. indica. 

Petals white; receptacle juicy F. vesca. 

Fragaria indica Andr. Bot. Rep. 7: pi. 479. 1807. Duchesnea 
indica Focke in Engl. & Prantl, Pflanzenfam. 3, pt. 3: 33. 1894. 

A silky-pubescent, strawberry-like plant with leafy runners and 
3-foliolate leaves; leaflets obovate or broadly oval, crenate-dentate; 
calyx 5-parted, the lobes alternating with larger, spreading, leaflike, 
3-dentate appendages; petals yellow, 1-1.5 cm. wide; fruit red, spongy, 
inedible. Plant intermediate between Fragaria and Potentilla and, 
except for convenience, better placed in the latter genus; cf. Wolf, 



FLORA OF PERU 1105 

Monogr. 664, where it has the name Potentilla indica (Andr.) Wolf. 
Illustrated, Bois, Diet. Hort. 553. 

Lima: Huariaca, 8104- Junin: Yanahuanca, 1170. Naturalized 
from Asia. 

Fragaria vesca L. Sp. PI. 494. 1753. 

Plant slightly villous; leaflets ovate or broadly oval, coarsely 
dentate, rather thin, pale and glabrate on the lower surface; scape 
usually exceeding the leaves, few-flowered, becoming irregular and 
raceme-like, with the primary branches unequal; sepals usually 
reflexed in fruit, much shorter than the fruit; achenes superficial, not 
imbedded in pits. 

Cuzco: Calca, Weberbauer 7894- Native of Europe and Asia; 
cultivated in many regions. "Fresa," "frutilla." 

13. POTENTILLA L. 

Reference: Wolf, Bibl. Bot. 71: 1-714. 1908. 

In general similar to Geum, but the leaves most often digitately 
3-7-foliolate. Calyx 4-5-bracteolate. Receptacle dry. Style most 
often ventral, not elongating. 

Leaflets subrotund or broader than long, sparsely pilose; bractlets 
and sepals subequal P. Dombeyi. 

Leaflets broadly ovate, longer than broad, densely pilose; bractlets 
longer than the sepals P. Weddellii. 

Potentilla Dombeyi Nestler, Monogr. Potent. 38. pi. 5. 1816; 397. 

Plants cespitose, greenish brown, sparsely long-pilose throughout, 
the stems 10-15 cm. long, prostrate-ascending; petioles 2-3 cm. long; 
stipules to 1 cm. long, the sheath 6-7 mm. long, the free tips narrowly 
ovate-lanceolate, acuminate, subobtuse; leaves all similar, subrotund, 
to 15 mm. wide and 12 mm. long, trifoliolate or deeply 3-lobed or 
often with only 1 lobe separated to the base, the lobes dentate half- 
way to the midrib, the teeth oblong, obtuse, mostly 6 on each lobe; 
uppermost leaves sessile, reduced; flowers few; pedicels in fruit 15 
mm. long; petals broadly obovate, 5 mm. long; sepals narrowly 
ovate, acute, scarcely 4 mm. long, equaled by the elliptic, obtuse 
bractlets; achenes smooth. Neg. 34755. 

Huanuco: Prostrate in open places in bunch-grass slopes, 3,500 
meters, Mito, 1882. Without locality, Dombey, type. 



1106 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Potentilla Weddellii Macbr., sp. nov. 

Habitu ad P. Dombeyi accedit sed differt caulibus minus dense 
pilosis, foliis late ovato-rotundatis, foliolis ad 15 mm. longis, 12 mm. 
latis subadpresse cinereo-strigosis; inflorescentia densiore; petalis ca. 
4 mm. longis; sepalis bracteolis minoribus. Cespitose, the slightly 
pilose, prostrate-ascending stems 5-15 cm. long; lowest petioles 2.5 
cm. long, appressed-pilose; leaves trifoliolate or the reduced upper 
ones 3-lobed, broadly ovate-rotund, the leaflets 15 mm. long and 12 
mm. wide or smaller, dentate about one-third the distance to the 
costa, the subobtuse teeth suboblong, grayish-pubescent on both 
sides with long, appressed but lax, lustrous hairs; flowers few, 
crowded ; petals 4 mm. long; calyx 3.5 mm. long, the bractlets elliptic, 
obtuse, the smaller sepals acute. There is in the Gay collection 
from Peru a specimen of Potentilla with trifoliolate leaves, probably 
new, supposed to have been collected in the temperate region of the 
Andes at Cuzco (Wedd. Chlor. And. 2: 234. 1857). Neg. 34756. 

Cuzco(?):Gcw/ (type, herb. Paris). 

14. ALCHEMILLA L. 

Reference: Perry, Contr. Gray Herb. 84: 1-57. 1929. 

Perennials, sometimes shrubby, often prostrate, or rarely annual. 
Leaves usually palmately or pinnately lobed or parted, rarely all 
radical. Stipules mostly foliaceous, lobed or cleft and connate to 
form a sheath or, if membranous, all or mostly entire. Flowers axil- 
lary or terminal, often cymose, glomerulate. Hypanthium persistent, 
often bracteate. The Peruvian species are chiefly of the section 
Lachemilla, with 2 (3-4) stamens, opposite the sepals, the anthers 
extrorse, in contrast to Alchemilla sensu strict., with 4 (5) stamens, 
alternate with the sepals, the anthers introrse. The section Aphanes, 
annual, the single stamen with an extrorse anther, is represented by 
one species. Rothmaler, Repert. Sp. Nov. 42: 164-173. 1937, re- 
establishes these sections as genera, a procedure scarcely to be 
followed outside monographic work. This author himself calls 
attention to the interesting analogy between the sections and their 
ranges. I acknowledge my indebtedness to Rothmaler's studies in 
the section Lachemilla, particularly his Alchemillae Colombianae, 
Trab. Mus. Cienc. Nat. Madrid, Bot. 31. 1935. 

Leaves appendaged with 2 leafy blades along the costa .A.diplophylla. 
Leaves more or less lobed or pinnate, not appendaged. 



FLORA OF PERU 1107 

A. Leaves palmately cleft or parted, sometimes shallowly or 
obscurely so. 

Basal leaves orbicular, obscurely 5-13-lobed. 

Leaves deeply cordate, 9-11-lobed; petioles 3-10 cm. long or 
longer. 

Styles 2; flowering branches short, robust, erect, leafless. 

A. orbiculata. 

Styles 4-8; flowering branches elongate, slender, leafy. 

A. pectinata. 

Leaves slightly cordate, 5-7-lobed; petioles to 3.5 cm. long. 

A. Lechleriana. 
Basal leaves 3-5-cleft or parted (in A. Williamsii orbicular). 

Cauline leaves developed, 3 (-5) -parted. 

Flowers, at least some of them, distinctly pedicellate or, 
if glomerate and subsessile, the hypanthium pubescent 
within. 

Hypanthium pubescent outside; flowers pedicellate. 
Leaves appearing 5-lobed by the laterally cleft lateral 
lobes. 

Leaves 3-parted A. andina. 

Leaves 5-7-lobed A. pseudovenusta. 

Leaves appearing 3-lobed, the lateral lobes not deeply 
cleft. 

Plants slightly pilose A. vulcanica. 

Plants densely pubescent A. Jamesonii. 

Hypanthium glabrous outside; plants spreading- villous- 

hirsute A. frigida. 

Flowers glomerate and subsessile; hypanthium glabrous 

within; plants, if pubescent, spreading-villous. 
Basal leaves palmately cleft; plants, if small, silvery- 
pubescent. 

Leaves more or less villous and orbicular, the lobes 
several-dentate. 

Flowers hirsute- villous A. Williamsii. 

Flowers glabrous or nearly so A. frigida. 

Leaves pilose or sericeous, the leaf lobes entire or bifid ; 
flowers often glabrous or but slightly pubescent. 



1108 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves more or less pilose A. aphanoides. 

Leaves silvery-villous; plants very small. 

A. holosericea. 

Basal leaves subpinnately cleft; plants small, not silvery- 
pubescent. 

Plants 1 cm. high, the leaves minute. . .A. sandiensis. 
Plants at least 3 cm. high, the leaves to 7 mm. long. 

A. repens. 
Cauline leaves reduced, forming, with the stipules, verticillate 

sheaths. 

Stipule lobes definitely spreading or reflexed, loosely hirsute. 
Stipule lobes 6 mm. long, little revolute; stigma sub- 

clavate A. galioides. 

Stipule lobes 4-5 mm. long, revolute; stigma globose. 

A. rivulorum. 
Stipule lobes ascending-appressed, appressed-hirsute. 

A. nivalis. 
A. Leaves pinnately cleft or parted. 

Basal leaves, at least, bipinnate, soft (see some species, perhaps 
Peruvian, under A. pinnata). 

Stipules all golden brown, membranous A. achilleaefolia. 

Stipules of the cauline leaves herbaceous. 

Calyx lobes subequal A. pinnata. 

Calyx lobes unequal, the outer 4 smaller, linear. 

A. erodiifolia. 
Basal leaves pinnate, the upper ones 3-parted, all rigid. 

A. barbata. 

Alchemilla achilleaefolia Re"my, Ann. Sci. Nat. III. 8: 224. 
1847. A. pinnata Re"my, op. cit. 6: 354. 1846, non Ruiz & Pavon. 
Lachemilla achilleifolia Rothm. Repert. Sp. Nov. 42: 169. 1937. 

Plants matted, from a fusiform root; radical leaves pinnate, the 
leaflets biparted, the segments oval, entire, obtuse, ciliate, with 
lanceolate, acute, brown, membranous, ciliate stipules; cauline leaves 
many-parted, villous; flowers involucrate, in small, axillary heads; 
perianth campanulate, hirsute, pedicellate, with 8 equal, oval, sub- 
obtuse, sericeous teeth; stamens 2. Allied to A. barbata, and sharing 
with it the conspicuous, brown stipules. See also A. erodiifolia, to 
which Perry referred my collection. 

Junin: San Jose', 4,000 meters, 1104 (det. Rothmaler). Bolivia. 



FLORA OF PERU 1109 

Aphanes andicola Rothmaler, sp. nov. 

Planta annua parva, 0.5-1 cm. alta, paullo ramosa, caulibus 
glabrescentibus, internodiis brevissimis; folia sessilia dense imbricata 
basi angustata cum stipulis connata, 3-loba, lobo mediano trifido, vel 
5-loba, margine longe ciliata, lobis lanceolatis obtusis, lateralibus 
integris, longe piloso-hirsutis; stipulae herbaceae inter se et cum 
lamina connatae, 2-fidae, lobis lanceolatis obtusis; flores in axillis 
stipularum 4-6-glomerulati, stipulas multo superantes, breviter pedi- 
cellati, 1.5-1.75 mm. longi, 0.75 mm. lati, subclausi, hypanthio ovato, 
brunneo, 4-costato, costis sat longe pilosis, intervallibus puberulis, 
sepalis 4 triangularibus, nervo dorsali piloso, apice pilo longiore 
obsitis praediti; episepala nulla. 

Ayacucho: Near Santa Ine*s, between Pisco and Ayacucho, in 
puna, 4,300-4,400 meters, Weberbauer 5454 (type in herb. Berol.). 

Alchemilla andina (Perry) Macbr. Candollea 5: 366. 1934. 
A. procumbens Rose, var. andina Perry, Contr. Gray Herb. 84: 23. 
1929. Lachemilla andina Rothm. Repert. Sp. Nov. 42: 169. 1937. 

Plants creeping, more or less appressed-pubescent or glabrous; 
leaves 0.5-1.5 cm. long, 3-parted, the obovate-cuneate divisions 
serrate, with 2-4 teeth on each side toward the apex; stipules leaflike; 
inflorescence loosely cymose, the filiform pedicels 3-10 mm. (1-5 mm. 
fide Rothmaler) long; hypanthium broadly campanulate, 1.5-2 mm. 
long, strongly constricted by the disk, the teeth subequal; styles 
3-5; stigmas globose. A. hirta (Perry) Rothm. Trab. Mus. Nac. 
Madrid Bot. 31 : 12. 1935, of Ecuador and Colombia, is similar but 
the stem pubescence is widely spreading and the bractlets are longer 
than the calyx teeth. A. fulvescens (Perry) Rothm. op. cit. 35, 
known as near as Ecuador, might be sought here because its reni- 
form basal leaves are deeply incised-5-lobed. The former species 
is illustrated, Rothmaler, op. cit. 13. 

Ancash: Above Huaraz, 3,200 meters, Weberbauer 3236. 
Ayacucho: Totorabamba, 3,600 meters, Weberbauer 5475 (det. 
Perry). Cuzco: Gay. Ollantaitambo, 3,000 meters, Cook & Gilbert 
696. Puno: Agapata, Lechler 1923. Bolivia to Colombia. 

Alchemilla aphanoides Mutis ex L. f. Suppl. 129. 1781. Lache- 
milla aphanoides Rothm. Repert. Sp. Nov. 42: 170. 1937. 

Plants forming mats, or the stems erect and crowded, densely or 
sparsely pilose; leaves 3-parted, the sessile upper ones usually gla- 
brous, the segments divided or 2-3-dentate; stipules with the sessile 
upper leaves forming many-lobed, perfoliate leaves; flowers crowded 



1110 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

in axillary and terminal cymes, at maturity typically glabrous, sub- 
sessile, the hypanthium 1 mm. long, the sepals and bractlets sub- 
equal; styles 1-3, usually 2, the stigma capitate. Var. tripartite, 
(R. & P.) Perry, 40 (A. tripartite, R. & P. Fl. Peruv. 1: 68. 1798), 
the usual Peruvian form, is more villous, the lateral leaf lobes often 
bifid, the flowers somewhat pilose, even at maturity. A. Sprucei 
Perry, 40, Ecuadorean, has all the stem leaves sessile and 3-5- 
lobed, the lobes deeply dentate; see also A. frigida below and A. ran- 
unculoides, this under A. Williamsii. The variety, according to 
Ruiz and Pavon, grew at Tarma, Pillao, and Huasa-huasi. 

Amazonas: Chachapoyas, Williams 7539. Huanuco: Panao, 
2208. Lima: Rio Blanco, 686. 

Alchemilla barbata Presl, Epim. Bot. 199. 1849; 55. A. 
Weberbaueri Pilger, Bot. Jahrb. 37: 537. 1906. Lachemilla barbata 
Rothm. Repert. Sp. Nov. 42: 169. 1937. 

Plants cespitose, well marked by the large, rigid leaves, fulvous 
stipules, and the entire leaflets 3-4 mm. long, barbate at the apex; 
stipules entire or lacerate, scarious; flowers terminal, densely glomer- 
ulate, the pedicels hirsute; hypanthium glabrous. 

Ancash: Between Ocros and Chonta, Weberbauer 2778. Huanta, 
4,000 meters (Pearce). Without locality: (Haenke, type). 

Alchemilla diplophylla Diels, Bot. Jahrb. 40: 277. 1908; 56. 
A. appendiculata Wedd. ex Murbeck, Lunds Univ. Arsskr. N. F. 2. 
11, pt. 8: 12. /. 3-4. 1915. Lachemilla diplophylla Rothm. Repert. 
Sp. Nov. 42: 169. 1937. 

Plants rhizomatous, 1-2 cm. high, the margins of the young leaves 
sparsely pilose; blades cuneate-obovate, 3-8 mm. long, 3-5 mm. wide, 
deeply tridentate, the middle tooth smallest, the appendages sub- 
equaling the leaf blade; petioles to 2 cm. long; stipules scarious, 
entire; flowers almost glabrous; peduncles bracteolate below the 
middle; bractlets none. This curious plant suggests in aspect some 
umbellifer. Moreover, its leaves closely simulate Peruvian species 
of Caltha, with which it has been confused in herbaria. 

Junin: Yauli, 4,400 meters, Weberbauer 288, type. Cuzco: In 
Distichia cushions, La Ray a, 4,400 meters, Pennell 13498, in part; 
Weddell 4412. Bolivia. 

Alchemilla erodiifolia Wedd. Chlor. And. 2: 247. 1857; 53. 
Lachemilla erodiifolia Rothm. Repert. Sp. Nov. 42: 172. 1937. 



FLORA OF PERU 1111 

Plants pilose nearly throughout, rooting at the nodes, the stems 
several decimeters long; radical leaves more or less bipinnatifid, with 
conspicuous, brownish, cleft stipules; cauline leaves pinnatifid, the 
segments 3-5-dissected ; flowers glomerulate, the perianth 8-dentate, 
the tube glabrous, the teeth sparsely pilose; carpels 2. The type 
was not designated but the description cited the following collection 
by Dombey and one by Weddell from Bolivia. Confused, according 
to Rothmaler, with A. achilleaefolia by Perry, who reduced the 
latter to A. pinnata. In the Dombey specimen the stipules are sub- 
herbaceous and cleft. Neg. 34281. 

Huanuco: Cheuchin, Dombey. Cuzco: Lucumayo Valley, Cook 
& Gilbert 1253 (fide Perry). Ecuador; Bolivia. 

Alchemilla frigida Wedd. Chlor. And. 2: 245. 1857; 27. 

Hirsute, with elongate, prostrate branches; basal leaves 1.5-2 cm. 
wide, subreniform, 3-5-lobed or cleft, the lobes incised-dentate with 
subobtuse teeth, sparsely hirsute above or glabrate; stipules mem- 
branous, lanceolate, acute, those of the subsessile cauline leaves 2-4- 
lobed; inflorescence terminal on short, lateral branchlets; flowers 
about 2 mm. long, short-pediceled, glabrous or with a few hairs on the 
lobes, the lobes 0.5 mm. long, the lance-ovate bractlets slightly 
shorter and narrower; styles 1-2; stigma clavate. Weberbauer 2691 
was referred by Markgraf to A. hirsuta HBK., a disposition followed 
by me in Candollea 5: 366. 1934. Rothmaler, however, has identified 
the HBK. plant as A. aphanoides; see Notizbl. Bot. Gart. Berlin 12: 
488. 1935. Neg. 34694. 

Cajamarca: Ocros, Weberbauer 2691. Bolivia; Argentina. 

Alchemilla galioides Benth. PI. Hartw. 134. 1844; 45. 

Stems loosely and densely hirsute; leaf sheath 8-10-lobed, at 
first loosely ascending; inflorescence glomerulate, axillary and ter- 
minal; flowers pedicellate; hypanthium 1.5 mm. long, only the lobes 
hirsute. Negs. 18016, 34285. 

Peru: Andes of Puitac, 3,000 meters (Pearce). Huanuco: Mon- 
zon, 3,400 meters, Weberbauer 3348 (det. Pilger). Ecuador. 

Alchemilla hirsuta HBK. Nov. Gen. & Sp. 6: 224. 1824; 57. 

Closely allied to A. aphanoides, but rather abundantly spreading- 
villous-hirsute; hypanthium pubescent within. Type locality un- 
known : Humboldt 31 48 in Herb. Willd. Neg. 34696. 

Ancash: Ocros, Weberbauer 2691 (det. Markgraf). 



1112 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Alchemilla holosericea Perry, Contr. Gray Herb. 84: 42. 1929. 
Lachemilla holosericea Rothm. Repert. Sp. Nov. 42: 170. 1937. 

A minute plant, rooting at the nodes, the ascending branches 1-2 
cm. high; leaves about 3 mm. long, sessile, the divisions of the lower 
ones entire; stipules much divided, appearing to form a sheath with 
the large segments revolute-margined; hypanthium 2 mm. long, 
appressed-villous; flowers axillary and solitary or terminally glomeru- 
late and pedicellate. 

Junin: Tambo de Vaca, 3,900 meters, 4398, type. Colombia; 
Ecuador. 

Alchemilla Jamesonii Perry, Contr. Gray Herb. 84: 25. 1929. 
Lachemilla Jamesonii Rothm. Repert. Sp. Nov. 42: 169. 1937. 

Similar to A. vukanica, but much more densely appressed-pubes- 
cent, the stipules strict, and the hypanthium conspicuously villous. 
According to the author, the achenes are larger than those of any 
other species. Neg. 27432. 

Cajamarca: Above Hualgayoc, 4,000 meters, Weberbauer 3966 (ex 
descr. ; determined at Berlin as A. rupestris) . Ecuador. 

Alchemilla Lechleriana Griseb. Goett. Abh. 24: 124. 1879; 16. 
Lachemilla Lechleriana Rothm. Repert. Sp. Nov. 42: 171. 1937. 

Like A. orbiculata, but the stolons leafy, the leaves often larger, 
usually densely appressed-villous beneath, and the inflorescence 
loosely racemose. 

Puno: Sachapata, Lechler 2606, type. 

Alchemilla nivalis HBK. Nov. Gen. & Sp. 6: 223. pi. 560. 1824; 
48. Lachemilla nivalis Rothm. Repert. Sp. Nov. 42: 170. 1937. 

Similar in aspect to A. galioides, but the 10-15 stipule lobes 
remaining erect, or only slightly spreading. 

Peru: Portachuelo (Mathews 1149). Amazonas: Balsas, Weber- 
bauer 4299 (det. Pilger). Without locality: River o. Colombia; 
Ecuador. 

Alchemilla orbiculata R. & P. Fl. Peruv. 1: 68. 1798; 13. 
Aphanes orbiculata Pers. Syn. PI. 1: 150. 1805. Lachemilla orbiculata 
Rydb. N. Amer. Fl. 22: 381. 1908. 

A decumbent herb with orbicular-reniform leaves and with con- 
spicuous, leafless stolons, the leaves scalelike or greatly reduced, 
1.5-2.5 cm. wide, the sinus open or closed, glabrate above, villous 
beneath, at least on the nerves; stipules membranous, light brown; 



FLORA OF PERU 1113 

inflorescence loosely corymbose, nearly leafless; flowers 3-4 mm. long, 
sericeous-villous. The pubescence of forma villifera (Perry) Rothm. 
is widely spreading, especially on the petioles. Neg. 18017. 

Cajamarca: Hualgayoc, 4,000 meters, Weberbauer 397 (det. 
Pilger). Junin: Rio Masamerich, 2,500 meters, Weberbauer 6671. 
Huacapistana, Weberbauer 2261 (det. Pilger). Chanchamayo, Isern 
2298. Huanuco: (Pearce 66). Southeast of Huanuco, wet bank at 
edge of thicket, 3,150 meters, 2088 (det. Perry). Mito, 2,700 meters, 
1439 (det. Perry). Panao, Chaclla, Pillao, Ruiz & Pavdn, type. 
Huasa-huasi, Dombey. Colombia to Peru. 

Alchemilla pectinata HBK. Nov. Gen. & Sp. 6: 226. 1824; 14. 
Lachemilla pectinata Rothm. Repert. Sp. Nov. 42: 171. 1937. 

In general characters similar to A. orbiculata, but the stolons 
leafy; leaves subcoriaceous, shallowly 9-11-lobed, pectinate-serrate; 
inflorescence axillary or terminal on leafy shoots; floral bracts am- 
plexicaul but not sheathing; achenes 4-6. 

Peru: (fide Rothmaler). Bolivia to Mexico. 

Alchemilla pinnata R. & P. Fl. Peruv. 1 : 69. 1798; 52. Aphanes 
pinnatus Pers. Syn. PI. 1: 150. 1805. Zygalchemilla pinnata Rydb. 
N.Amer.F1.22:385. 1908. 

Usually softly gray-villous, but sometimes greenish ; leaves bipin- 
natifid, 1.5-6 cm. long, the numerous pinnae usually imbricate and 
unequally divided; flowers solitary and axillary or terminal and 
glomerulate, distinctly pedicellate, villous to glabrate. Variable 
in pubescence. Forma argentea Rothm. is smaller and densely seri- 
ceous-pilose; forma minima Wedd. ex Rothm. (var. minima Wedd. ex 
Griseb. Goett. Abh. 19: 137. 1874; PI. Lorentz. 89. 1874) is glaucous, 
sparsely hirsute or glabrate. The very distinct A. paludicola Rothm. 
op. cit. 41 and A. Mandoniana Wedd. have the bractlets reduced or 
wanting; the basal leaves of the former have 2-3 pairs of leaflets, 
those of the latter are nearly pinnately parted. Since these two 
closely related forms have been found in Bolivia as well as in Ecuador, 
their occurrence in Peru is probable. Illustrated, Trab. Mus. Nac. 
Madrid Bot. 31: 32 and 42. Neg. 18018. Ruiz and Pavon listed the 
localities as Tarma, Huasa-huasi, Huariaca, Payanchacra, Rondos, 
Chaulan, and Pillao. 

Cajamarca: Ocros, Weberbauer 2687 (det. Pilger). Cuzco: Cuzco, 
Herrera 2385. Lima: La Raya, Pennell 13527, 13525. Matucana, 
Weberbauer 190 (det. Pilger). Rio Blanco, 807. Cheuchin, Ruiz & 



1114 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pawn. Junin: Palca, Isern 2292. Yauli, 4,400 meters, Weberbauer 
318. Puno: Chuquibambilla, Pennell 13411. 

Alchemilla pseudovenusta Rothm. Notizbl. Bot. Gart. Berlin 
12:489.1935. 

A ligneous, creeping plant, the branched, procumbent stems root- 
ing at the tips, to 15 cm. long, densely pilose with erect-spreading 
hairs; basal leaves subrosulate, reniform, acute, 2 cm. broad, 1.5 cm. 
long, 5- to partially 7-lobed or incised to the middle, the divisions 
with 8-10 oval teeth on each side, more or less densely pubescent, 
especially on the nerves; cauline leaves similar, petioled; lower 
stipules membranous, the upper herbaceous, many of them multi- 
parted; flowers in dense cymes, aggregate toward the tips of the 
branchlets, solitary in the axils of the bracts, densely hirsute or sub- 
sericeous, short-pedicellate; hypanthium turbinate, 1.75 mm. long, 
the subequal lobes and bractlets 1.25 mm. long; styles 4-5, exserted, 
the stigma long-clavate; stamens 2. The type specimen was referred 
originally by Markgraf to A. orbiculata R. & P., a disposition which 
I had accepted; the author compares it with A. pectinata HBK. The 
petioles are as much as 5 cm. in length. 

Junin: In sphagnum, near Huacapistana, 3,200 meters, Weber- 
bauer 2216, type. 

Alchemilla repens Presl, Epim. Bot. 199. 1849; 43. Lachemilla 
repens Rothm. Repert. Sp. Nov. 42: 172. 1937. 

A little plant, resembling A. sandiensis, but at least 3 cm. high, 
the bractlets much shorter than the sepals. See also A. Mandoniana 
(under A. pinnata), which might be sought here because of its 
subpinnate basal leaves. 

Lima: Open, rocky slope, Huaros, 3,500 meters, Pennell 14716. 
Without locality: Haenke, type. 

Alchemilla rivulorum Rothm. Notizbl. Bot. Gart. Berlin 12: 
490. 1935. Lachemilla rivulorum Rothm. Repert. Sp. Nov. 42 : 170. 1937. 

With the habit of A. galioides, but the stems much more branched ; 
basal leaves deeply 3-lobed, the lateral lobes bifid, the margins revo- 
lute; cauline leaves sessile, the lower 3-parted and slightly longer 
than the sepals; sheath divisions 9, lanceolate; hypanthium globose, 
1.25 mm. long, more or less hirsute toward the tip; styles 2-3. 
The type was referred originally by Pilger to A. galioides, a disposi- 
tion that had been accepted by the present writer. Neg. 34287. 

Ancash: In stones at brook margin, mountains above Huaraz, 
4,200 meters, Weberbauer 2973, type. Ecuador. 



FLORA OF PERU 1115 

Alchemilla sandiensis Pilger, Bot. Jahrb. 37: 537. 1906; 43. 

Stoloniferous, the stems 2-4 cm. long, slightly villous; basal 
leaves orbicular-ovate, 3-parted, the terminal segment 6-8-lobed, 
the lateral segments 4-5-lobed, the lobes oblong, acute to subobtuse; 
stipules subovate, membranaceous; cauline leaves 3-parted, with 
leaflike stipules; flowers 1.5-2 mm. long; hypanthium turbinate, 
appressed-pubescent; sepals ovate; pistils 1 or 2. Neg. 18019. 

Puno: Between Poto and Ananea, Sandia, Weberbauer 945, type. 

Alchemilla vulcanica Schlecht. & Cham. Linnaea 5: 573. 
1830; 24. Lachemilla vulcanica Rydb. N. Amer. Fl. 22: 382. 1908. 

Often dense, the creeping stems suffrutescent, the ascending stems 
and branches appressed-pilose; lateral leaf lobes very rarely bifid, 
sometimes glabrous above, the narrowly cuneate lobes 3-7; petioles 
3-5 mm. long, shorter than the linear-lobed, spreading stipules; 
inflorescence cymose, the pedicels 1-5 mm. long; hypanthium 1.5 mm. 
long, sparsely pilose within. 

Ancash: Above Huaraz, 4,200 meters, Weberbauer 3110. 
Amazonas: Chachapoyas (Mathews). Junin: Yauli, 4,400 meters, 
Weberbauer 348. Puno: Cuyocuyo, 3,600 meters, Weberbauer 926. 
Colombia; Bolivia; Guatemala; Mexico. 

Alchemilla Williamsii Perry, Contr. Gray Herb. 84: 26. 1929. 
Lachemilla Williamsii Rothm. Repert. Sp. Nov. 42: 172. 1937. 

Appressed-villous, at least on the leaf nerves beneath, low or 
depressed; leaves orbicular-reniform, glabrate above, the lobes more 
or less marked, often obscurely so, the teeth 2-3 mm. long, linear- 
acute; stipules lance-ovate; cauline leaves 3-parted; flowers glomeru- 
late; hypanthium villous outside, 2 mm. long. Neg. 34706. 

Junin: Palca, Isern 2205(1). CuzcoiGay. Bolivia. 

15. MARGYRICARPUS R. & P. 

Somewhat tortuously branched shrubs, often spinescent from the 

rigid leaf rachises, the leaves imbricate, unequally pinnate, with 

broad, vaginate petioles, or the leaflets apparently fasciculate. 

Flowers sessile, inconspicuous, solitary in the leaf axils; petals none. 

Calyx ebracteolate, persistent, winged or tuberculate, constricted at 

the throat. Stamens 2 or 3. 

Leaves at least in part appearing simple, fasciculate; fruits wing- 
angled M. strictus. 

Leaves all pinnate, the leaflets evanescently setose at the apex; fruits 
nerve-angled M . pinnatus. 



1116 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Margyricarpus pinnatus (Lam.) Kuntze, Rev. Gen. 3, pt. 2: 
77. 1898. Empetrum pinnatum Lam. Encycl. 1 : 567. 1785. Ancistrum 
barbatum Lam. Illustr. 1: 77. 1791. M. setosus R. & P. Fl. Peruv. 
1: 28. pi. 8. 1798. 

A decumbent, sprawling shrub; leaves about 5 mm. long, the 
linear leaflets lustrous, glabrous except for caducous setae at the tip ; 
petioles vaginate; drupe white, slightly fleshy, suggesting a pearl 
(whence the common name), sweet and edible (Ruiz & Pavdn); drupe, 
when dry, rib-angled, the ribs entire. 

Cajamarca: Cutervo, Jelski 326. Huanuco: Mito, 1515. Junin: 
Tarma, Huasa-huasi, Xaurxa, Huarocheri, Conchucos, Ruiz & Pawn. 
Palca, 3,000 meters, Weberbauer 24-50; 248. Cuzco: Pampa de Anta, 
3,400 meters, Herrera 699. Valle de Cuzco (Weberbauer 181), 4881, 
4899 (det. Pilger). Without locality: Jussieu, type. Argentina and 
Chile. "Yerba de perlilla," "perlillas," "canlli," "china-canlli." 

Margyricarpus strictus (Poepp.) Macbr., comb. nov. Tetra- 
glochin strictum Poepp. Fragm. Syn. 26. 1833. M. alatus Gill, in 
Hook. Bot. Misc. 3: 305. 1833. M. imberbis Presl, Epim. Bot. 
200. 1849(7). 

Leaves often spiniform, or sometimes with 1 or 2 leaflets, other 
leaves much smaller, fasciculate, trifoliolate; fruit 5-8 mm. long, 
3-5-winged, the wings often somewhat denticulate. This could be 
regarded as a distinct genus, but habitally the shrubs are similar, 
the fruit more or less wing-angled. The Peruvian plant seems to be 
M. cristatus Britton, Mem. Torrey Club 3, pt. 3: 25. 1893, this 
possibly a distinct variety. Illustrated, Wedd. Chlor. And. 2: pi. 77. 

Lima: Banos (WilkesExped.). Rio Blanco, 3,000 meters, in firm, 
stony soil, 732, 3003. Puno: Puno, Soukup 212, 213. Sanborg, 
Weberbauer 1389 (det. Pilger). Chuquibambilla, Pennell 13356 
Cuzco: Gay. Pucara, Weberbauer 407 (det. Pilger). Sacsahuaman, 
3,500 meters (Herrera 102). Tacna: Werdermann 1122. Lake Titi- 
caca, Meyen. Arequipa: Pampa de Arrieros, Pennell 13332. Boli- 
via; Chile. "Canlli," "orcco-canlli." 

16. ACAENA L. 

Reference: Bitter, Bibl. Bot. 74: 1-336. 1910-1911. 

Decumbent or ascending herbs, often suffrutescent at the base, 
with erect, usually scapelike flowering branches. Leaves unequally 
pinnate, the leaflets incised-serrate or divided. Stipules vaginate, 
adnate to the petiole. Stamens 1-10. Fruit usually tuberculate or 
aculeate, included in the persistent calyx. Carpels 1-2. 



FLORA OF PERU 1117 

Inflorescence capitate. 

Leaflets oval, about 1 cm. long A. ovalifolia. 

Leaflets narrowly ovate-lanceolate, about 2 cm. long. .A. argentea. 
Inflorescence spicate. 

Leaflets white-pubescent beneath A. cylindristachya. 

Leaflets green beneath, often somewhat pubescent. 

Fruit 4-5 mm. long; leaflets 5-7 pairs, subequal. .A. torilicarpa. 

Fruit 7-8.5 mm. long; leaflets 4 pairs, with 2-3 much smaller 

ones A. elongata. 

Acaena argentea R. & P. Fl. Peruv. 1: 67. pi. 103. 1798; 215. 

Branches suberect, with old leaves below, 30 cm. high, the creep- 
ing, branching, ligneous stems becoming many decimeters long, the 
internodes 5-6 cm. long, more or less densely long-pubescent; leaflets 
mostly 9, rarely 11-13, oblong-lanceolate, acutish, glabrous above, 
long-appressed-gray-pilose beneath, at least on the nerves, mostly 
2.5-3.5 cm. long, nearly 1 (-1.5) cm. wide, the lower ones greatly 
reduced; flowers capitate, perfect; stamens 3-4; fruit with 4 spines, 
the longer ones 6-7 mm. long, apically 4-5-glochidiate, the body of 
the fruit 5 mm. long, attenuate. Type from Chile. A. subincisa 
Wedd. Chlor. And. 2: 242. 1857; 218, Ecuadorean, is similar but the 
leaves are 5-6-pinnate, the leaflets thinner, deeply serrate-dentate, 
yellowish-sericeous beneath. Neg. 29606. 

Piura: Between Huancabamba and Ayavaca, Weberbauer 6325. 
Patagonia to Ecuador. 

Acaena cylindristachya R. & P. Fl. Peruv. 1: 68. pi. 104. 
1798; 46. 

A species well marked by the numerous leaflets of the (6-8 cm. 
long) leaves, these densely appressed-pilose, greenish above, silvery- 
sericeous beneath, 1 cm. long, 4 mm. wide, suboblong; fruit turbinate, 
with 3-5 very unequal spines, the longest 4 mm. long. A tufted 
plant from a stout caudex, the nearly naked, appressed-pilose pedun- 
cle 10-20 cm. high, the cylindric spikes 2 cm. long. Illustrated, 
Wedd. Chlor. And. 2: pi. 76. Neg. 17997. 

Junin: Tarma, Weberbauer 2509 (det. Bitter); Ruiz & Pawn. 
Cuzco: Paucartambo, Vargas 317. Cerro de Cusilluyoc, Pennell 
13833. Huanuco: Chaclla, Ruiz & Pavon. Mito, 1816. Puno: 
Sandia, Soukup 152. Bolivia to Costa Rica. 

Acaena elongata L. Mant. 2: 200. 1771; 28. A. lappacea R. & 
P. Fl. Peruv. 1 : 66. pi. 103. 1798. 



1118 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

More or less tortuously much branched, fruticose at the base, 
the branches clothed with old petiole bases; flowering stems erect- 
ascending; leaves 2-6 cm. long or longer, 4-7- (rarely 9-) pinnate; 
sheaths slightly connate at the base, minutely stipitate-glandular 
or glabrous, the stipules foliaceous; lower leaflets minute, entire, the 
upper ones oblong-obovate, 10-15 mm. long, 4-5 mm. wide or larger, 
acute, crenate-serrate, the teeth usually pilose at the apex, pilose 
beneath on the costa, glabrous above; raceme more or less peduncled, 
the lower bracts often 3-lobed; flowers erect, subsessile; body of the 
nodding, mature fruit 7-8.5 mm. long, pilose, with 10-12 spines. 
The Madrid specimen of the Ruiz and Pavon plant is referred by 
Pilger to this species, but unfortunately it is without locality; the 
specimen in herb. Berlin is labeled as from Tarma. 

Junin: Tarma, Ruiz & Pavdn. Acobamba, Picoy, Tapu, Ruiz & 
Pavon. Lima: Matucana, 463. North to Mexico. 

Acaena ovalifolia R. & P. Fl. Peruv. 1: 67. pi. 103. 1798; 236. 
A. ovalifolia var. serrata Bitter, Bibl. Bot. 74: 238. 1910. 

A sprawling, ligneous, densely leafy plant; leaves several-11- 
foliolate (fide Bitter), in the typical form with about 3 pairs of leaflets, 
the lower ones much reduced; leaflets oval-elliptic or slightly obovate, 
obtuse, sharply dentate-crenate, green and glabrous above, more or 
less sericeous beneath; scapes suberect, leafless above, little elongate 
in fruit, the capitate inflorescence then 1.5 cm. broad; fruit 2.5-4 mm. 
long, abruptly attenuate at the base, appressed-pilose, the spines 
usually 2, rarely 4, as much as 12 mm. long, glochidiate only at the 
apex. A highly variable species, forming carpet-like mats. 

Cajamarca: Hualgayoc, Weberbauer 4006 (det. Pilger). Ayacu- 
cho: Pampalca, Killip & Smith 22211. Junin: Tarma, Ruiz & 
Pavon; Dombey. Huanuco: Open meadow land, Huacapistana, 
Weberbauer 2097 (det. Pilger) ; Killip & Smith 24407. Chinchupalca, 
1595. Near Huanuco, 2076 (det. Killip). Pillao, Acobamba, Huasa- 
huasi, Ruiz & Pavon. South American Andes. "Canaharamjo," 
"broquin." 

Acaena torilicarpa Bitter, Bibl. Bot. 74: 33. 1910. 

Similar to A. elongata, but the leaflets often more numerous and 
subequal, the fruit smaller, the body 3-5 mm. long, the 1-3 spines 
retrorse-glochidiate even to the base. Pilger refers this species to 
A. lappacea (here treated as a synonym of A. elongata). Bitter, as 
usual, gives a separate taxonomic status to each herbarium sheet: 
var. 1, robusta Bitter, loc. cit.; var. 2, brevidentata Bitter, subvar. 



FLORA OF PERU 1119 

parvifolia Bitter, op. cit. 34; var. 3, gracilis Bitter, op. cit. 34. Accord- 
ing to Weberbauer, this species is a shrub 0.5-2 meters high. 
Neg. 18005. 

Cuzco: Urubamba, Weberbauer 4921. Lima: Matucana, Weber- 
bauer 177. Ancash: Huailas, 3,000 meters, Weberbauer 3158. 

17. SANGUISORBA L. 

Annuals or perennials with alternate, pinnatifid, stipulate leaves 
and small, perfect flowers in dense, peduncled, cylindric-oblong 
spikes. Calyx lobes 4, deciduous, the angled tube dry and winged in 
fruit; carpels included, usually 1. Stamens 2-12. 

Sanguisorba officinalis L. Sp. PI. 116. 1753. 

A glabrous perennial with a rootstock, sometimes more than 
1 meter high; leaflets 9-13, ovate to lance-ovate, serrate, 2-8 cm. 
long; upper stipules foliaceous, often lunate, serrate; spikes 1-3 cm. 
long, 1 cm. thick; filaments usually included ; sepals dark purple, 2-2.5 
mm. long. 

Cuzco: Quebrada de Miscahuara (Hen era 1528). Adventive 
from Europe. "Pimpinela." 

87. CONNARACEAE 

By Julian A. Steyermark 

Reference: Schellenberg, Pflanzenreich IV. 127. 1938. 
Trees, shrubs, or woody vines. Leaves alternate, exstipulate, 
1-3-foliolate or odd-pinnate, the leaflets coriaceous, entire. Flowers 
usually paniculate in the Peruvian species, rarely racemose, mostly 
perfect. Calyx 5- or rarely 2-parted, usually persistent and surround- 
ing the base of the fruit, the lobes imbricate in the Peruvian species. 
Petals 5, free or slightly connate, imbricate. Stamens 10 in the 
Peruvian species, opposite the petals, 5 of them longer than the 
others, perigynous or hypogynous, the filaments filiform, monadel- 
phous at the base, often united at the base into an annular disk. 
Carpels 5 or 1, 1-celled, free, superior. Style slender. Stigma capi- 
tellate and simple or bilobed. Ovules 2. Capsule follicular, 1-2- 
seeded. Seeds often arillate. 

Carpel 1, solitary 4. Connarus. 

Carpels 5. 

Inflorescence terminal or, if axillary, the epidermis of lower leaf 
surface scarcely papillose; flower maturing 1 solitary follicle. 

3. Rourea. 



1120 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Inflorescence axillary; flower maturing several follicles. 

Epidermis of lower leaf surface papillose. . .1. Pseudoconnarus. 
Epidermis of lower leaf surface not papillose .... 2. Bernardinia. 

1. PSEUDOCONNARUS Radlk. 

Leaves 3-foliolate. Panicles axillary, many-flowered. Petals 5, 
obovate-cuneate. 

Pseudoconnarus reticulatus Schellenb. Pflanzenreich 103. IV. 
127: 94. 1938. 

Branches densely tomentose; petioles velutinous; leaflets 3, the 
terminal one broadly oval, shortly and acutely acuminate at the 
apex, rounded at the base, the lateral ones oblique, 3.5-10 cm. long, 
2.5-6.7 cm. broad, coriaceous, subtriplinerved, the lower surface 
finely reticulate-veined, papillose and tomentulose; inflorescence 
paniculate, axillary, with a velutinous rachis; flowers unknown; 
fruit 9 mm. long, clavate, glabrous. 

Loreto: Iquitos, 100 meters, Ducke, type. Specimen not seen. 

2. BERNARDINIA Planch. 

Small, woody vines in the Peruvian species. Leaves 3-foliolate 
in the Peruvian species. Panicles axillary. Calyx deeply parted into 
5 lobes, imbricate, scarcely increasing in size after anthesis. Petals 5. 
Stamens 10, the filaments glabrous. Carpels 5. 

Bernardinia agelaeoides Schellenb. Pflanzenreich IV. 127: 100. 
1938. 

A woody vine with glabrous branches; leaves long-petioled, the 
rachis 4-7 cm. long, glabrous; leaflets 3, with thickened petiolules 
4-7 mm. long, the blades of the leaflets ovate-elliptic, 8-14 cm. 
long, 4-7.5 cm. broad, long-acuminate at the apex, cuneate at the 
base, coriaceous, glabrous, the nerves and veins on the upper surface 
conspicuous, the nerves and costa on the lower surface very promi- 
nent, strongly subtriplinerved, the 2 lowest nerves joined to the 
costa 0.5-1 cm. from the base of the blade, subglaucescent and 
densely areolate-reticulate on the lower surface; inflorescence appear- 
ing after the leaves, fasciculate-paniculate, subsessile, 5-7.5 cm. 
long, the branches sparsely pilosulous to nearly glabrous, congested 
and densely many-flowered; pedicels slender, 4-4.5 mm. long, mostly 
glabrous; bracts minute; calyx 2.5 mm. long, glabrous outside, with 
linear-oblong, obtuse, ascending lobes 2 mm. long, much longer than 
the tube; petals white, linear, glabrous, 3-4 mm. long, 1 mm. broad; 
fruit not seen. 



FLORA OF PERU 1121 

Loreto: Florida, Rio Putumayo, 200 meters, Klug 1986, type. 

This species has not as yet been found in fruit and, therefore, is 
placed only provisionally, out of respect to Schellenberg, in Bernar- 
dinia, although on the basis of the imbricate calyx lobes, inflorescence 
appearing after the leaves, and on flower characters, it could just as 
well be included in Rourea. 

3. ROUREA Aubl. 

Leaves evergreen, coriaceous, odd-pinnate. Panicles axillary in 
the Peruvian species, many-flowered, erect or pendulous. Flowers 
small. Calyx increasing in size after anthesis. Petals 5, white or 
yellowish, longer than the calyx, linear-oblong to suborbicular. 
Capsule curved. 

Lower leaf surface, petioles, and youngest branchlets brown-velu- 

tinous R. rhynchosioides. 

Lower leaf surface, petioles, and youngest branchlets glabrous. 

Sepals spreading in fruit; pedicels 4-10 mm. long in anthesis. 

Panicles loosely flowered; pedicels 6-10 mm. long. R. cuspidata. 

Panicles densely flowered; pedicels 4-5 mm. long.fi. densiflora. 

Sepals erect-ascending or appressed in fruit; pedicels stout, 2-3 mm. 

long in anthesis. 

Calyx lobes suborbicular; petals large, oblong-spatulate, 7-8 
mm. long; leaflets usually 5-7, rarely 3 R. camptoneura. 
Calyx lobes oval or oblong-lanceolate; petals smaller, linear- 
oblong, 4-5 mm. long; leaflets usually 3. 

Calyx lobes oval R. puberula. 

Calyx lobes oblong-lanceolate R. amazonica. 

Rourea amazonica Huber, Bol. Mus. Goeldi 5: 373. 1909. 

A woody vine with usually 3-foliolate, rarely 5-foliolate leaves; 
leaflets broadly ovate or broadly elliptic, 8-22 cm. long, 3-11 cm. 
broad, abruptly acuminate, rounded or gradually curved at the base, 
coriaceous; panicles 8-12 cm. long, the branches minutely puberulent; 
flowers white, relatively large, loosely scattered, on pedicels 2-2.5 mm. 
long, these becoming in fruit 3.5-5 mm. long; calyx lobes ovate- 
lanceolate, obtuse, in fruit longer than the tube; petals 4-5 mm. 
long, white, subacute; fruit obovoid, arcuate, conspicuously striate, 
2-2.5 times longer than the calyx. Neg. 30172. 

Loreto: Rio Itaya, Williams 221. Rio Huallaga, 150-210 meters, 
Williams 3885, 4091, 4107, 4142, 4180, 4619, 4871. Yurimaguas, 
Poeppig D2152. Amazonian Brazil. 



1122 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Rourea camptoneura Radlk. Sitzungsb. Akad. Wissensch. 
Munchen 16: 375. 1886. 

A woody vine with 3-7-foliolate leaves; leaflets ovate, 8-14 cm. 
long, 3-7.5 cm. broad, acuminate, rounded at the base, coriaceous; 
panicles 10-15 cm. long, the branches minutely puberulent; flowers 
white, relatively large, loosely scattered, on pedicels 2-3 mm. long; 
calyx lobes ovate-rotund, rounded at the apex, in fruit about equaling 
the tube; petals narrowly oblong-spatulate, rounded at the apex, 
7-8 mm. long; fruit elliptic-obovate, 1.5 cm. long, striate, 2.5-3 
times longer than the calyx. Neg. 19293. 

Loreto: Rio Nanay, Williams 1009. Mishuyacu, near Iquitos, 
100 meters, Klug 458, 905. Iquitos, 110 meters, Mexia 641 la. 
Brazil. 

Rourea cuspidata Benth. ex Baker in Mart. Fl. Bras. 14, pt. 2: 
181. 1871. 

A woody vine; leaves usually 3-foliolate or the uppermost ones 
simple; leaflets elliptic-oblong, 8-18 cm. long, 3.5-9 cm. broad, long- 
acuminate, rounded or tapering at the base, coriaceous, glabrous, 
somewhat paler beneath ; panicles 7-12 cm. long, the branches slender, 
glabrate; flowers white, loosely scattered on slender, elongate pedicels 
6-10 mm. long; calyx lobes oblong, obtuse, in flower and fruit exceed- 
ing the length of the tube; petals oblong, obtuse, 4-6 mm. long; 
fruit slightly curved, 1-1.3 cm. long. Neg. 5824. 

Loreto: La Victoria, Williams 2963. Iquitos, 120 meters, Wil- 
liams 1336, 1513, 8109. Mishuyacu, near Iquitos, 100 meters, Klug 
231,801,805. Brazil. 

Rourea densiflora Steyermark, sp. nov. 

Frutex scandens, ramulis glabris; foliis longipetiolatis; foliolis 3 
crasse 3-6 mm. longe petiolulatis ovato-ellipticis 10-16 cm. longis, 
6-8 cm. latis, apice longe acuminatis, basi cuneatis, coriaceis, glabris, 
nervis venulisque supra non elevatis, costa nervisque subtus valde 
elevatis, plus minusve concoloribus; inflorescentia paniculata termi- 
nali pedunculata 14 cm. longa conferte ramosa, ramis patenti- 
adscendentibus glabris dense multifloris, pedicellis tenuibus 4-5 mm. 
longis, plerumque glabris, bracteis minutis sed conspicuis, parce 
pubescentibus; calyce ca. 2 mm. longo extus parce pilosulo, lobis 
oblongis obtusis patenti-adscendentibus tubo multo longioribus; 
petalis albidis oblongis 1.5-1.75 mm. longis, plerumque glabratis; 
fructu non viso. 



FLORA OF PERU 1123 

Loreto: Mishuyacu, near Iquitos, 100 meters, King 796 (type in 
Herb. Field Mus.). 

This new species of Rourea is closely related to R. cuspidata 
Benth., from which it differs in its more closely and numerously 
flowered panicles and shorter pedicels. 

Rourea puberula Baker in Mart. Fl. Bras. 14, pt. 2: 179. 1871. 

A woody vine with usually 3-, rarely 1-foliolate leaves; leaflets 
elliptic, 6-12 cm. long, 3-6 cm. broad, long-acuminate at the apex, 
rounded or more or less narrowed at the base, chartaceous; panicles 
7-9 cm. long, the branches puberulent; flowers white, relatively small, 
on pedicels 2 mm. long; calyx lobes elliptic or oblong, obtuse, puberu- 
lous outside; petals 5 mm. long, slightly longer than the calyx; 
fruit 1.2 cm. long. 

Loreto: Florida, Rio Putumayo, 180 meters, Klug 2138. Yuri- 
maguas, lower Rio Huallaga, 135 meters, Killip & Smith 29037. 
Mishuyacu, near Iquitos, 100 meters, Klug 31*3. Brazil; Bolivia. 

Rourea rhynchosioides Standl. Field Mus. Bot. 11: 158. 1936. 

A woody vine with 3-foliolate leaves; leaflets broadly oval or 
obovate-rotund, 8-12 cm. long, 6-8.5 cm. broad, the terminal one 
somewhat larger and broader, rounded or shortly subacute at the 
apex, rounded or subemarginate at the base, above mostly glabrous 
except for the fulvous-pilosulous nerves, beneath densely fulvous- 
pilose, especially on the nerves, veins, and costa; panicles 12-17 cm. 
long, the branches densely fulvous-tomentose, remotely many- 
flowered; flowers yellowish, very small, on thickened pedicels scarcely 
1 mm. long; calyx lobes triangular-ovate, obtuse, erect; petals 4 mm. 
long, obtuse; fruit not seen. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 578, type. 
Amazonian Brazil. 

4. GONNARUSL. 

Leaves odd-pinnate. Panicles axillary or terminal, many- 
flowered, racemose or spicate. Flowers small. Calyx not increasing 
in size after anthesis. Petals 5, longer than the calyx, ligulate or 
oval, white, cream-colored, yellow, or greenish brown. 

Nerves on upper surface of leaflets very prominent. 

Leaflets usually 3; inflorescence elongate, racemose-paniculate. 

C. Sprucei. 
Leaflets usually 7-11; inflorescence short, spicate. .C. pachyneurus. 



1124 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Nerves on upper surface of leaflets inconspicuous, or at least not 

prominent. 
Leaflets 5-11; rachis of inflorescence ferruginous-tomentose. 

C. Patrisii. 

Leaflets 3; rachis of inflorescence smooth or slightly grayish-hairy. 

C. Martii. 

Connarus Martii Schellenb. Pflanzenreich IV. 127: 243. 1938. 

A shrub; leaves long-petiolate, the rachis glabrous; leaflets 3-5, 
oblong, 3.5-9 cm. long, 1.5-3.8 cm. broad, shortly and obtusely 
acuminate at the apex, rounded at the base, glabrous, concolorous, 
the lateral nerves 6-8 pairs, inconspicuous; calyx moderately pilosu- 
lous outside, with oblong-lanceolate lobes 2.5-3 mm. long; petals 
4 mm. long, obtuse, glabrous; filaments all glandular; fruit 1.8 
cm. long. 

San Martin: Alto Rio Huallaga, Williams 6641- Bolivia; Brazil. 

Connarus pachyneurus Radlk. Sitzungsb. Akad. Wissensch. 
Munchen 16: 365. 1886. Connarus Klugii Standl. Field Mus. Bot. 
11:157.1936. 

A shrub, 1-4 meters tall; leaves large, long-petiolate, the rachis 
glabrous; leaflets 7-11, oblong or ovate-oblong, 15-26 cm. long, 5-7.5 
cm. broad, abruptly cuspidate-acuminate at the apex, cuneate- 
obtuse or rounded at the base, glabrous, concolorous, the veins and 
nerves on the lower surface elevated and prominent; panicles axillary, 
short, densely many-flowered, spiciform, with simple, subsessile 
branches 1-3 cm. long, shortly ferruginous-pilose; calyx densely 
stellate-ferruginous-tomentose outside, 2.5 mm. long, with 2 broadly 
ovate calyx lobes; petals yellow or brownish, oval or broadly ovate, 
3 mm. long, glabrous; filaments glabrous, shortly connate at the 
base; fruit 2-2.5 cm. long. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 500, 1494- 
Brazil. 

This species, remarkable because of its 2-lobed calyx, may well 
constitute a distinct genus. It is placed provisionally in Connarus. 

Connarus Patrisii (DC.) Planch. Linnaea 23: 432. 1850. Om- 
phalobium Patrisii DC. Prodr. 2: 86. 1825. 

A small tree, 5-8 meters tall, with glabrous branches; leaves long- 
petiolate, the rachis glabrous; leaflets of the lower leaves usually 5-7, 
of the upper mostly 3, obovate-oblong or oblong-elliptic, abruptly 



FLORA OF PERU 1125 

cuspidate-acuminate, rounded at the base, glabrous, concolorous, 
the veins and nerves on the lower surface slightly prominent; panicles 
closely many-flowered, the ultimate branches elongate, racemose, 
4.5-12 cm. long, shortly ferruginous-pilose; calyx sparsely ferrugi- 
nous-pilose outside; petals white, greenish, or cream-colored, ligulate, 
obtuse, 4.5-6 mm. long, much longer than the calyx; fruit scarlet, 
about 2.5 cm. long. 

San Martin: Juanjui, 400 meters, Klug 3855. Rio Huallaga, 360- 
900 meters, Williams 5670, 5733, 6209, 6214, 6641, 6808Loreto: 
Florida, Rio Putumayo, 180 meters, Klug 2366. French Guiana; 
Brazil; Ecuador. 

Connarus Sprucei Baker in Mart. Fl. Bras. 14, pt. 2: 187. 1871. 

A shrub or small tree with glabrous branches; leaves long-petio- 
late, the rachis glabrous; leaflets usually 3, elliptic-oblong, abruptly 
cuspidate-acuminate, rounded at the base, glabrous, concolorous, 
the nerves on both surfaces very prominent; panicles large, loosely 
many-flowered, the ultimate branches elongate and racemose, 4-16 
cm. long, shortly ferruginous-pilosulous to glabrate; calyx sparsely 
puberulous outside; petals brownish yellow, ligulate, obtuse, 2.5-4 
mm. long, longer than the calyx. Neg. 19291. 

Loreto: Timbuchi, Rio Nanay, Williams 1010, 1149. Iquitos, 
120 meters, Williams 3664. Lower Rio Huallaga, 155-210 meters, 
Williams 3806. Balsapuerto, 220 meters, Klug 2937. Yurimaguas, 
lower Rio Huallaga, 135 meters, Killip & Smith 27563. Between 
Yurimaguas and Balsapuerto, 135-150 meters, Killip & Smith 28296. 
Amazonian Brazil. 



ADDITIONS AND CORRECTIONS 

Page 96. Add the following new species: 

Peperomia Soukupii Trelease, sp. nov. 

Herba ut videtur modica caespitosa glabra, caule in sicco valde 
angulato; folia ad nodum 4-6-lanceolata, basi acuta, ca. 3.5 cm. longa 
atque 1.5 cm. lata, in sicco dura aurea, revoluta, supra profunde sul- 
cata, subtus trinervia; inflorescentia non visa. 

Cuzco: Growing on Theobroma, Quillabamba, J. Soukup 137 
(type in Herb. Field Mus.). 

Page 113. Line "B" should be changed to read: Midrib without 
strong, ascending branches upwards. 



Page 253. PLEIOSTACH YOPIPER Trelease, Proc. Amer. Phil. 
Soc. 73: 328. 1934 should replace the generic heading on this page. 
The citation for the single species should read: Pleiostachyopiper 
nudilimbum (C. DC.) Trelease, Proc. Amer. Phil. Soc. 73: 329. 
pi a. 1934. 

Page 306. After Ficus paraensis add the following species: 

Ficus peruviana (Miq.) Rossberg, Repert. Sp. Nov. 42: 61. 
1937. Pharmacosycea peruviana Miq. Lond. Journ. Bot. 7: 72. 1848. 

Branchlets sparsely pilose with slender, appressed, fugacious 
hairs; petioles 1-1.5 cm. long; blades broadly elliptic, 11-13 cm. long, 
6 cm. wide, abruptly and shortly obtuse-acuminate, rounded at the 
base and trinerved, coriaceous, glabrous, the lateral nerves 6-10 on 
each side; stipules almost as long as the petioles, lanceolate; recep- 
tacles globose, glabrous. The description is based upon the original 
one by Miquel. Rossberg states that the species does not belong to 
the subgenus Pharmacosycea. Because of lack of data regarding the 
receptacles in the original description, I am unable to decide whether 
the species is the same as one of those that I have described from 
Peru. Paul C. Standley. 

Without definite locality: Mathews 2061, type. 



Page 307. After Ficus Ruiziana add the following species: 

Ficus Ulei Rossberg, Repert. Sp. Nov. 42: 60. 1937. 

A tree; stipules 6 mm. long, subglabrous; petioles 2.5 cm. long 
or less, the blades chartaceous, oblong or elongate-elliptic, rarely 
obovate, 13-24 cm. long, 8.5 cm. wide or less, cuspidate, obtuse or 

1126 



FLORA OF PERU 1127 

somewhat cuneate at the base, glabrous above or sparsely puberulent 
on the costa, glabrous beneath, the lateral nerves 7-10 on each side, 
divergent at an angle of 70-85 degrees; receptacles globose, on pedun- 
cles 3 mm. long or less, 14-17 mm. in diameter, glabrous. Referable to 
subgenus Pharmacosycea; apparently closely related toF. radula Willd. 
Loreto : Mainas Alto (Poeppig) . Amazonian Brazil. 



Page 421 . In the treatment of the Olacaceae there were several 
unfortunate complications that necessitate some extensive additions 
and corrections to the family. The manuscript for the group was 
prepared several years before it was printed, although an attempt 
was made to check literature just before printing. Some of the 
recently described species had not yet been entered or at least filed 
in the Gray Herbarium card catalogue of new species, and somehow 
or other the appearance of the Olacaceae in the second edition of 
the Pflanzenfamilien escaped notice. It is necessary to present here 
a new key to the genera of the family, and a complete new treatment 
of the genus Heisteria. The writer is greatly obliged to Dr. H. 
Sleumer, who has kindly forwarded useful notes regarding the dis- 
position of the Peruvian species of Heisteria. Paul C. Standley. 

Stamens twice as many as the petals. 

Flowers fasciculate in the leaf axils; calyx usually greatly enlarged 

and colored in fruit Heisteria. 

Flowers spicate or racemose; calyx not or scarcely accrescent in 

fruit Minquartia. 

Stamens as many as the petals or fewer. Calyx not accrescent in 

fruit, or but little enlarged. 
Stamens free, as many as the petals or fewer. 

Petals 5-6, the stamens fewer Liriosma. 

Petals 4, the stamens of the same number Tetrastylidium. 

Stamens united to form a tube, of the same number as the petals. 

Stamens united to form a tube about the style Schoepfia. 

Stamens united with the petals Aptandra. 



Page 422. Below is a new key to the Peruvian species of Heisteria, 
with an enumeration of them. The number of species here recognized 
is almost double that of the original treatment. The available material 
is much less than would be desirable for study, especially since some 
of the species seem to be rather vaguely limited. I have some doubt 



1128 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

that all species listed here are valid ones, but it may be that ample 
material will show that the number of Peruvian forms is even greater. 

Fruiting calyx deeply lobate, much longer than the drupe, often 

enclosing it. 

Lateral nerves of the leaves about 15 on each side . . . H. Sleumeri. 
Lateral nerves of the leaves about 10 on each side. 

Leaves mostly 5.5-7 cm. long H. iquitensis. 

Leaves mostly 10-15 cm. long or larger H. Spruceana. 

Fruiting calyx subentire or only shallowly lobate, usually shorter 
than the drupe, often reflexed, or at least not enclosing the fruit. 
Fruit oval or ellipsoid, conspicuously longer than broad. 
Calyx broad, strongly accrescent, colored, shallowly lobate. 

H. cyanocarpa. 
Calyx cupular, not or scarcely accrescent, green, truncate. 

Fruit globose, about as broad as long. 

Plants scandent; lateral nerves of the leaves 3-4 on each side. 

H. scandens. 

Plants erect shrubs or trees; lateral nerves of the leaves more 
numerous. 

Leaves dull, not lustrous H. pallida. 

Leaves very lustrous. 

Leaf blades very obtuse at the base, the lateral nerves 
strongly arcuate H. caloneura. 

Leaf blades acute to broadly cuneate at the base, the 
lateral nerves almost straight H. nitida. 

Heisteria caloneura Sleumer, Notizbl. Bot. Gart. Berlin 12: 
66. 1934. 

A tree about 20 meters high; leaves short-petiolate, ovate or 
ovate-oblong, 8-12 cm. long, 3.5-5.5 cm. wide, rather abruptly 
acuminate, rounded-cuneate and asymmetric at the base, coriaceous, 
very lustrous, the lateral nerves 5-6 on each side; pedicels in flower 
about 5 mm. long, numerous; petals ovate, acuminate, 2 mm. long; 
immature fruit globose, the calyx accrescent, almost entire, spreading. 

Loreto: Mouth of Rio Santiago, 160 meters, in upland forest, 
Tessmann 4419, type. Amazonian Brazil. 

Heisteria cyanocarpa P. & E. 

The treatment for this species remains the same as on page 423. 



FLORA OF PERU 1129 

Heisteria iquitensis Sleumer, Repert. Sp. Nov. 38: 207. 1935. 

A tree of 35 meters; petioles about 1 cm. long; blades obovate- 
elliptic or oblong-elliptic, 5.5-9.5 cm. long, 3-4.5 cm. wide, shortly 
obtuse-acuminate, broadly cuneate at the base, subcoriaceous, some- 
what lustrous above, dull beneath, the lateral nerves about 12 on 
each side; pedicels 2.5 mm. long; petals elliptic, 2.5 mm. long, gla- 
brous outside, pilosulous within; fruiting calyx red, 5-lobate, the lobes 
ovate, 2.5-3 cm. long; drupe ellipsoid, white, 13 mm. long. 

Loreto: Iquitos, upland forest, Ducke 19561. Amazonian Brazil. 

Heisteria nitida Engler in Mart. Fl. Bras. 12, pt. 2: 18. 1872. 

A shrub or tree, 10 meters high or less; leaves lance-elliptic or 
lance-oblong, mostly 7-10 cm. long but sometimes larger, acute or 
acuminate, acute to broadly cuneate at the base, very lustrous, the 
lateral nerves numerous and very oblique, almost straight, the nerves 
and veins conspicuous on both surfaces; pedicels numerous, 3-5 mm. 
long; fruiting calyx purple-red, about 1 cm. long, spreading, sub- 
entire; fruit subglobose, 1 cm. long. 

Loreto: Fortaleza, in forest, Williams 4248. Iquitos, Tessmann 
5280 (fide Sleumer). San Martin: Juanjui, 400-800 meters, King 
4287. Tarapoto, Spruce 4148. Junin: Chanchamayo, Isern 2171. 
La Merced, Weberbauer 1865 (fide Sleumer). Huanuco: Posuso, 
Ruiz & Pawn (fide Sleumer); Ruiz & Pavon 14-52. Amazonian 
Brazil. 

Heisteria pallida Engler. 

The treatment of this species remains as on page 424. 

Heisteria parvicalyx A. C. Smith, Brittonia 2: 146. 1936. 

A shrub or small tree, sometimes 7.5 meters high, with a trunk 
7 cm. in diameter; petioles 1.5 cm. long or less; blades subcoriaceous, 
elliptic-oblong, 13-22 cm. long, 4-9 cm. wide, short-acuminate, 
attenuate at the base, often very lustrous, the lateral nerves 6-9 on 
each side; flowers usually few in a fascicle, the stout pedicels 1-2 mm. 
long; petals lance-ovate, 4-5 mm. long; fruit ellipsoid, about 10 mm. 
long and 7 mm. broad. 

Loreto: Paraiso, Alto Rio Itaya, 145 meters, Williams 3379. 
La Victoria, Williams 2957. Amazonian Brazil. 

Williams 3379 is referred (in litt.) by Sleumer doubtfully to H. 
iquitensis, but it seems to be undoubtedly referable here, as also 
Krukoff 8278, referred doubtfully by A. C. Smith to H. Spruceana. 



1130 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

On page 424 of this Flora, Williams 3379 was placed incorrectly 
under H. densifrons Engler, a species not known to occur in Peru. 

Heisteria scandens Ducke, Archiv. Jard. Bot. Rio Jan. 4: 9. 
1925. H. eurycarpa Standl. Field Mus. Bot. 11: 148. 1936. 

A shrub or usually a large, woody vine; see the description on 
page 424. Known in Peru only from the single collection cited as the 
type of H. eurycarpa. Also in Amazonian Brazil. 

Heisteria Sleumeri Standl., sp. nov. 

Frutex omnino glaber, ramulis crassiusculis solemniter angulatis, 
internodiis elongatis; folia breviter petiolata subcoriacea magna, 
petiolo crasso 1-1.5 cm. longo; lamina anguste elongato-lanceolata 
ca. 26 cm. longa, 5.5-6.5 cm. lata, longe sensim attenuata, basi 
breviter cuneato-acuta, supra sublucida, nervis venisque plus minusve 
impressis, subtus paullo pallidior, brunnescens, costa crassiuscula 
elevata, nervis lateralibus utroque latere ca. 15 elevatis, angulo lato 
divergentibus, prope marginem pulchre junctis et nervum intramar- 
ginalem efformantibus, nervis aliis intermediis bene evolutis atque 
prominentibus inter nervos primaries interpositis; flores in axillis 
subsessiles; calyx fructifer rubro-purpureus fere 2.5 cm. longus, 
profunde lobatus, lobis rotundato-ovatis apice anguste rotundatis. 

Loreto: Forest between Rio Nanay and Rio Napo, Williams 698 
(type in Herb. Field Mus.). 

Apparently a quite distinct species, noteworthy for the unusually 
numerous nerves of the exceptionally long and narrow leaves. Con- 
spicuous is the anastomosing of the lateral nerves to form an almost 
regular submarginal nerve. 

Heisteria Spruceana Engler in Mart. Fl. Bras. 12, pt. 2: 15. 1872. 

The description and citation of specimens (with very few excep- 
tions) for Heisteria cauliflora Smith on page 422 are referable to H. 
Spruceana, which occurs also in Amazonian Brazil. H. cauliflora, 
as limited by Sleumer, does not occur in Peru. A synonym of 
this species is H. Tessmanniana Sleumer, Notizbl. Bot. Gart. 
Berlin 12: 67. 1934, based on Tessmann 5060, from Iquitos. Dr. 
Sleumer writes that new Heisteria material from eastern Peru shows 
that the characters upon which it was separated from H. Spruceana 
do not hold. 



Page 426. Sleumer (Pflanzenfam. ed. 2. 16b: 31. 1935) reports 
Schoepfia Schreberi Gmel. from Peru, without indication of the 



FLORA OF PERU 1131 

region. This species may occur in Peru ; I have seen no Peruvian speci- 
mens or records. 

Page 427. After Aptandra tubicina add the two following genera: 
MINQUARTIA Aubl. 

Large trees; leaves short-petiolate, elliptic or oblong-elliptic, 
more or less coriaceous, acuminate, glabrous and lustrous above, 
minutely pubescent beneath; inflorescences solitary in the leaf axils, 
racemose, many-flowered, pubescent; calyx small, 5-dentate, per- 
sistent; petals usually 5, united below; stamens 10, 5 of them opposite 
the petals, the others opposite the sepals; ovary 5-celled; fruit 
drupaceous, 1-seeded, with a somewhat fleshy exocarp. 

Minquartia punctata (Radlk.) Sleumer, Repert. Sp. Nov. 
39: 282. 1936. Endusa punctata Radlk. Sitzungsb. Akad. Wiss. 
Miinchen 16: 311. 1886. M. macrophylla Ducke, Archiv. Inst. 
Biol. Rio de Janeiro 2: 33. 1935. 

A tree of 12-15 meters, stellate-puberulent with a brownish 
indument; petioles mostly 3-4 cm. long; blades oblong to elliptic- 
oblong, 20-35 cm. long, cuspidate-acuminate, narrowly rounded to 
broadly cuneate at the base, the lateral nerves conspicuous beneath ; 
inflorescences 9 cm. long or less, simple, many-flowered, the flowers 
aggregate, short-ped