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Full text of "Flora of Peru"

THE UNIVERSITY 

OF ILLINOIS 

LIBRARY 




OCT 7 1939 



PUBLICATIONS 



OF 



FIELD MUSEUM OF NATURAL 
HISTORY 



BOTANICAL SERIES 

VOLUME XIII 
PART II 




THE LIBRARY OF THE 

MAR 151939 

UNIVERSITY OF ILLINOIS 
CHICAGO, U.S.A. 

1936-1938 



THE LIBRARY OF THE 

JUL 1-1936 

UNIVERSITY OF ILLINOIS 



BOTANICAL SERIES 



FIELD MUSEUM OF NATURAL HISTORY 

FOUNDED BY MARSHALL FIELD, 1893 

VOLUME XIII 



FLORA OF PERU 

PART II, No. 2 



BY 

J. FRANCIS MACBRIDE 

ASSOCIATE CURATOR, HERBARIUM, DEPARTMENT OF BOTANY 

THE LIBRARY OF THE 

MAR 3 1937 

B. E. DAHLGREtfNIVERSITY OF ILLINOIS 

CHIEF CURATOR, DEPARTMENT OF BOTANY 
EDITOR 



PUBLICATION 379 




CHICAGO, U.S.A. 
MARCH 15, 1937 



PRINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 



58 0. 

FB 

V. ) 2>, 



J. FRANCIS MACBRIDE 



41. CHLORANTHAGEAE. Chloranthus Family 
Reference: Solms in DC. Prodr. 16, pt. 1: 479-485. 1869. 
Not only the articulate branchlets, enlarged at the nodes, but 
also the agreeable fragrance which Ruiz and Pavon describe accu- 
rately as seemingly emanating from the entire shrub, identify at 
once the members of this small family. The resin exuded appears 
as tear-shaped drops, with the color and odor of grains of the alma- 
; ciga; in some places it is collected for the preparation of comforting 
poultices (Ruiz and Pavon). 

1. HEDYOSMUM Sw. 

Tafalla R. & P. Syst. 269. 1798. 

jfc Shrubs or small trees, always readily known by the opposite 

/branchlets jointed at the nodes and by the more or less elongate, 

-.sheath-like, connate stipules. Flowers monoecious or dioecious, 

the staminate in ebracteate aments, the pistillate bracteate and 

"^capitate or cymose, the cymes often branched. Several of the species 

|are doubtfully distinct, but most of them are meagerly known. 

Some forms of H. racemosum are questionably separable from H. 

^arborescens Sw., a species typically West Indian. It is a pity that 

-<|lhe name Tafalla has not been conserved to commemorate the artist 

' \flof the Ruiz and Pavon expedition, who collected so many of the 

*" specimens. 

^ Aments and cymules small, 1-2 sessile in the axils of small leaves; 

leaves conspicuously scabrous beneath H. scabrum. 

Aments and cymules well developed, the latter several, spicate, 

paniculate, or capitate-congested. 
Pistillate flowers in cymules, not capitate-congested. 
Petioles 7-20 mm. long; leaves mostly 10 cm. long. 

Cymules 6-8 mm. long, 3-4 mm. wide H. Dombeyanum. 

Cymules smaller H. racemosum. 

Petioles 3-8 mm. long; leaves mostly smaller. 
Leaves ovate-elliptic or oblong-elliptic, bluntly pointed. 
Branchlets scurfy; drupes 1.5-2 mm. long . . H. Kanehirae. 
257 



258 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branchlets glabrous; drupes 3-3.5 mm. long . H. Huascari. 

Leaves oblong-lanceolate, acuminate H. Lechleri. 

Pistillate flowers capitately congested; leaves oblong-lanceolate, 
coarsely serrate H. glaucum. 

Hedyosmum Dombeyanum Solms in DC. Prodr. 16, pt. 1: 
482. 1869. 

Petioles 1-1.5 cm. long; leaf blades ovate-oblong, 7-10 cm. long, 
2-3 cm. broad, subacuminate, obtusely callous-serrate, glabrous but 
somewhat scabrous and distinctly punctate above; cymules loosely 
paniculate, about 3-flowered, the bracts subequaling the ovate, 
trigonous drupes. 

Huanuco: Cochero (Dombey}. 

Hedyosmum glaucum (R. & P.) Cordem. Adansonia 3: 303. 
1863. Tafalla glauca R. & P. Syst. 271. 1798. 

Branches stout, obtusely angled; leaves acuminate, 10-15 cm. 
long, rarely 3 cm. broad, glabrous; aments terminal, at first ovoid 
and 1.5-2 cm. long, becoming cylindric and 3 cm. long; fruiting in- 
florescence fleshy, to 3 cm. long and 2 cm. broad, the bracts linear, 
shorter than the lustrous, ovate, trigonous drupes. Neg. 8537. 

Huanuco: Cochero and Acomayo, Pavon. "Aitacupi," "alma- 
ciga." 

Hedyosmum Huascari Macbr. Field Mus. Bot. 11: 15. 1931. 

Similar in foliage to H. Kanehirae but the cymules crowded in a 
spike 6-8 mm. long, and the slightly exserted, light brown drupes 
to 3.5 mm. long. Named for one of the last kings of the Incas, 
Huascar. 

Cajamarca: Above Tabaconas, 2,400 meters, Weberbauer 6113, 
type. Amazonas: Chachapoyas, Weberbauer 4353. 

Hedyosmum Kanehirae Macbr. Field Mus. Bot. 11: 14. 1931. 

A tree about 5 meters high with densely scurfy-punctate branch- 
lets having short internodes; leaves crowded, thick, the veins rather 
prominent beneath, the blades closely and minutely callous-crenate, 
dull, glabrous, oblong-elliptic-lanceolate, bluntly narrowed at the 
apex, up to 7 cm. long and 3 cm. wide but mostly smaller; petioles 
3-6 mm. long; cymules approximate in short racemes, many of them 
sessile or nearly so, 4-5 mm. long and 3-4 mm. broad, the bracts 
nearly enclosing the finally black, subtrigonous, acute drupes, these 



FLORA OF PERU 259 

scarcely 2 mm. long. The leaves are employed as a remedy for 
rheumatism (Kanehira). 

Huanuco: Pan de Azucar, Sawada 64, type. Monzon, Weberbauer 
3388. Pampayacu, Kanehira 115, 204- 

Hedyosmum Lechleri Solms in DC. Prodr. 16, pt. 1: 484. 1869. 

Leaves glabrous, thinnish, oblong-lanceolate, acutely acuminate, 
remotely crenate-serrulate, 5-7 cm. long, 1.2 cm. broad, the veins 
obscure, even beneath; cymules in a contracted raceme or spike, 
about 5 mm. long; drupes pale brown, 3 mm. long, little exserted. 

Puno: Talaxara (Lechler 2632, 2660). Sangaban, Lechler 2279. 
Ayacucho: Carrapa, 1,500 meters, Killip & Smith 22403. 

The Killip and Smith specimen, from a tree 7.5 meters high, is 
perhaps a distinct species, the cymules being 7-8 mm. long, the 
drupes included, 4 mm. long. 

Hedyosmum racemosum (R. & P.) G. Don, Gen. Syst. 3: 434. 
1834. Tafalla racemosa R. & P. Syst. 271. 1798. 

A glabrous shrub or small tree with ample, chartaceous, more or 
less serrulate leaves; petioles in the type 7-10 mm. long but often 
2 cm. long or longer; leaf blades lance-elliptic, acuminate, mostly 
10-15 cm. long and 5-7 cm. wide; cymules in simple or nearly simple 
racemes or spikes; drupes in the type trigonous, rather bright (or 
reddish) brown, about 2 mm. long, exserted. Solms recognizes H. 
integrum Cordem. Adansonia 3: 302. 1863, and H. Sprucei Solms in 
DC. Prodr. 16, pt. 1: 483. 1869 (Neg. 7831). The former is distin- 
guished by exactly trigonous, well exserted drupes; the leaves are 
lustrous and the cymules paniculate. H. Sprucei is similar but the 
leaves are nearly dull and the pale drupes are well exserted. Possibly 
these are distinct species but they may be variants, their apparent 
differences being due, at least in part, to the degree of maturity. 
My No. 5798 was a slender tree 7 meters high with a bushy top. 
An infusion of the leaves in alcohol is used as a remedy for rheuma- 
tism (Weberbauer). The odor is that of bergamot (Raimondi) 
Neg. 7830. 

San Martin: Zepelacio, 1,200 meters, King 3273, 3746. Tara- 
poto, Spruce 4436 (H. Sprucei Solms!). Tarapoto, Spruce 4311. 
San Roque, 1,400 meters, Williams 7006, 7500. Junin: Chancha- 
mayo Valley, Schunke 421, 481, 524, 525. La Merced, 1,400 meters, 
5798. Pichis Trail, 1,500-1,900 meters, Killip & Smith 25956. 
Loreto : Pumayacu, 600-1,200 meters, Klug 31 75. Cerro de Escalera, 
Ule 6583. Without locality: Mathews 1994. Huanuco: Haenke 



260 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

(det. Pilger). Monzon, Weberbauer 3549; 256. Rio Posuso, 1,800 
meters, Weberbauer 6738 (det. Mansfeld). Cajamarca: Cutervo, 
Raimondi (det. Krause). Puno: Sandia, Weberbauer 1112 (det. 
Krause); 278. Ayacucho: Near Quillomito, 1,500 meters, Weber- 
bauer 7544- Bolivia. "Anis," "supinum," "carpales," "asar guiru." 

Hedyosmum scabrum (R. & P.) Solms in DC. Prodr. 16, pt. 1: 
480. 1869. Tafalla scabra R. & P. Syst. 270. 1798. H. latifolium 
Cordem. Adansonia 3: 308. 1863. 

Petioles 1-2 cm. long or longer; leaves elliptic-ovate, shortly 
obtuse-pointed, 8-12 cm. long, 4-6 cm. wide, coarsely callous-serrate, 
glabrous above, definitely scabrous and more or less hirsutulous 
beneath ; aments and cymules in pairs, sessile, both in fruit to 10 mm. 
long. Var. Pavonii Solms (Neg. 8535) has glabrous, acuminate 
leaves 3-4 cm. wide and shorter cymules. H. Mandoni Solms of 
Bolivia is to be expected. Its narrower leaves are densely and sharply 
callous-dentate. Neg. 7828. 

Amazonas: Chachapoyas, Raimondi (det. Krause). Cajamarca: 
Cutervo, Jelski 101; Raimondi. Chugur, Weberbauer 4070, 4075 
(det. Krause); 259, 260. Cuzco: Prov. Convention, Weberbauer 
5894, 5893. Huanuco: Chinchao (River o). Muna and Pati, Pavon. 
Mufia, 1,950 meters, 4111. "Aytacupi." Colombia. 

Doubtful Species 

Hedyosmum angustifolium (R. & P.) Solms and H. laciniatum 
(R. & P.) Solms, DC. Prodr. 16, pt. 1: 485. 1869 (under Tafalla 
R. & P. Syst. 272. 1798) are essentially nomina nuda; no specimens 
have been found. Probably both are referable to H. racemosum. 

42. SALICACEAE. Willow Family 

Cultivated members of the family include, according to Herrera, 
Populus tremula L. as "alamo chileno" or "alamo temblon," for 
ornament or for the making of charcoal, and Salix babylonica L., 
the weeping willow, "sauce lloron," which, according to him, "has 
acclimated itself well in canyons of the Department of Cuzco, where 
it is used for decorations at funerals and religious fetes." S. chilensis 
develops into a stately tree in Cuzco and Apurimac (Weberbauer). 

1. SALIX L. Willow 

Apparently only two willows are native in Peru. 
Salix chilensis Molina, Sagg. Nat. Chil. 169. 1782. S. Hum- 
boldtiana Willd. Sp. PI. 4: 657. 1805. 



FLORA OF PERU 261 

Becoming a tree 10 meters tall or higher: leaves linear, acuminate, 
serrulate; stipules deciduous; catkins appearing with the leaves, the 
woolly bracts deciduous; capsules ovate, glabrous, the glabrous 
pedicels longer than the gland. Ascending to more than 3,000 
meters, cultivated only (Weberbauer 85). Illustrated, Mart. Fl. 
Bras. 4, pt. 1: pi. 71. 

Cuzco: Particularly in the provinces of Urubamba, Calca, and 
Quispicanchi (Herrera). Huasao, 3,200 meters, Herrera 3023. 
Arequipa: Region of Mt. Misti (Weberbauer 128, 129). Ancash: 
Puccha Valley (Weberbauer 173). Huaraz (Weberbauer 172). 
Junin: Tarma (Weberbauer 177). Cajamarca: Ocros (Weberbauer 
162). Maranon Valley (Weberbauer 174; 190). Amazonas: Utcu- 
bamba (Weberbauer 192). Lima: Barranco (Weberbauer 148). San 
Lorenzo Island near Callao, in 1852, Andersson. Huara, Ruiz & 
Pawn; Dombey. Piura: (Weberbauer 150). Huanuco, 2,100 meters, 
3232. Argentina and Chile to Texas and the West Indies. "Sauce." 

Salix Martiana Leybold in Mart. Fl. Bras. 4, pt. 1: 227. pi. 72. 
1885. 

Similar to S. chilensis but the catkins looser, the scales of the 
staminate linear-lanceolate and entire instead of ovate, the capsules 
oblong, and the pedicels villous. "Much less common on the Rio 
Ucayali than on the Amazon; found only on the main stream and 
some of the chief tributaries. Nevertheless I found a few isolated 
examples in the great canyon of the Cerro de Canchahuaya" (Huber). 
This is probably the species reported by Ule, Bot. Jahrb. 40: 121-123, 
as S. Humboldtiana. 

Loreto: Quebrada Grande del Cerro de Canchahuaya (Huber 
1 328, 1 564) Florida, Rio Putumayo, 200 meters, in forest, Klug 2075; 
a tree of 5 meters. Fortaleza, 200 meters, Williams 4447. Lower 
Rio Nanay, river banks, Williams 460. Brazil. "Paharbubu." 

43. MYRICACEAE. Bayberry Family 

Reference: Chevallier, Me"m. Soc. Sci. Nat. Math. Cherbourg 
32: 85-340. 1901. 

The pungent aroma that the resinous-glandular leaves or at least 
the berry-like, crowded fruits yield on slight pressure is a well-known 
character of this family, which is represented best in the northern 
hemisphere. The fragrant, whitish wax covering the fruits of certain 
northern species is sometimes largely replaced by or mixed with 
pubescence in the Peruvian species. 



262 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. MYRICA L. 

Nothing about these ament-bearing shrubs serves to distinguish 
them more readily, even at first glance, from other Peruvian amentif- 
erous plants than the extraordinarily crowded and numerous leaves. 
Leaves nearly oblong, about 1 cm. wide; plants dioecious. 

M. Pavonis. 
Leaves lance-obovate, about 2 cm. wide; plants monoecious. 

M. pubescens. 

Myrica Pavonis C. DC. in DC. Prodr. 16, pt. 2: 151. 1864; 287. 
M. Pavonis var. glandulosa Chev. op. cit. 288. 

A slenderly branched shrub or small tree, the youngest parts 
canescent-pilose or puberulent; leaves scarcely narrowed to the obtuse 
tip, 5-7 cm. long, 6-10 mm. wide, short-attenuate to the slender 
petiole, more or less denticulate, resinous-glandular beneath; aments 
1 cm. long; flower bracts ovate, acute, pubescent-margined, shorter 
than the flowers. The type was from Guayaquil, by Ruiz and 
Pavon, who assigned an unpublished specific name referring to the 
Salix-like leaves, which simulate those of the common South Ameri- 
can willow. Neg. 8538. 

Lima: Canta (Simm 88), Cuzco: Pillahuata, 2,400 meters, 
Pennell 14026, Moquehua: Above Moquehua, 3,200 meters, Weber- 
bauer 7391, 7390 (det. Markgraf). "Huacan timbu." 

Myrica pubescens Humb. & Bonpl. ex Willd. Sp. PI. 4: 746. 
1806; 289. 

Robustly branched, the virgate branchlets and leaves more or 
less permanently pilose (rarely glabrous or glabrate), the latter 
resinous beneath; leaves 7-10 cm. long, 1.5-3 cm. wide, coarsely and 
sharply serrate or denticulate, acute or acuminate; aments often 
3-6 cm. long; bracts awl-shaped, acuminate, pilose, exceeding the 
flowers; fruits nearly 5 mm. thick, wax-covered, tomentose (var. 
tomentosa Chev.) or glabrous (var. glabra Chev.). Var. glandulosa 
Chev. is glabrous or nearly so, but the young fruits are tomentose, 
becoming waxy. The young fruits of my No. 3484 were citrous- 
aromatic. "When one takes a bit of the pulverized bark of this shrub 
in place of tobacco, one sneezes 10 or 12 times; the powder does not 
irritate or damage the nose, and after cleaning with the handker- 
chief the sting and the sneezing cease; experiment has shown that 
this cleans out the head and relieves migraine" (Ruiz & Pavon). 
Weberbauer determinations by Markgraf. Illustrated, Denkschr. 
Akad. Wiss. Wien 15: pi. 4 (as "tinctoria"). Negs. 11553, 25097. 



FLORA OF PERU 263 

Cajamarca: Huambos, 3,000 meters, Weberbauer 4160, 4187; 260 
(under an unpublished name). Libertad: Chicama, 1,600 meters, 
Weberbauer 6986 (var. tomentosa}. Ancash: Huaraz, 3,200 meters, 
Weberbauer 3245 (var. glandulosa) . Tallenga, Prov. Cajatambo, 
3,600 meters, Weberbauer 2890 (var. glandulosa); 179 (under an 
unpublished name). Junin: Huacapistana, 2,500 meters, Weber- 
bauer 2192; Killip & Smith 24519 (3-5 meters high). Huanuco: 
Cani, near Mito, dense shrub or tree of grassy slopes, 2,550 meters, 
3484 (det. Killip as a glabrate form). Lima: Canta (Rivero; var. 
tomentosa). Ayacucho: Between Tambo and Rio Apurimac, 2,900 
meters, Weberbauer 5587. Cuzco: Urubamba, 1,800 meters, Weber- 
bauer 5057; 245 (det. Schellenberg). Marcapata, 3,200 meters, Weber- 
bauer 7784- Convencion, Weberbauer 5057 (fruits white- waxy). 
Peru to Venezuela and Costa Rica. "Laurel," "tuppassaire," 
"ssaire." 

44. JUGLANDACEAE. Walnut Family 

Only the black walnut or "nogal" represents this economically 
important family in Peru. 

1. JUGLANS L. Walnut 

Reference: Dode, Bull. Soc. Dendr. France 4: 165-215. 1909. 

Georges H. Barrel, Trop. Woods 10: 51-53, has given an account 
of his personal observation of "nogal" in Peru. He noted it as 
rather common on the upper reaches of the Rio Ucayali, along the 
Pichis Trail and in the Chanchamayo Valley, as the following cita- 
tions bear witness. He found that the native people recognized a 
"nogal bianco" and a "nogal negro." Weberbauer and later Williams 
recorded it from the Department of Amazonas, the former giving its 
altitudinal range in the valleys of the north as 1,600-2,000 meters, 
and on the eastern slopes in the rain forest as between 800 and 1,000 
meters. Herrera has recorded it from Cuzco. Barrel found the 
individual trees rarely clustered, often scattered, which in spite of 
their great timber value (the wood is highly prized for fine construc- 
tion) makes their lumbering expensive. Botanists have reported 
the tree as "plentiful" near Chachapoyas, but probably not from a 
lumberman's standpoint. Apparently the tree usually branches at 
3-5 meters though trees have been reported to 30 meters high that 
branched at 10 meters with a trunk 1 meter in diameter below. 
Walnut is valued locally not only for its wood but also for a dye made 
from a decoction of the bark, leaves, and fruit. According to He- 
rrera, Juglans regia L., the English walnut, is cultivated in Cuzco, 



264 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the nuts being known as "nueces de Chile." The "nogal de la tierra" 
identified by Ruiz and Pa von as J. nigra L., cultivated at Chancai 
for its timber and nuts, has been determined by Markgraf as J. 
neotropica Diels. Williams, Trop. Woods 27: 16-17, has added his 
observations to those of Barrel and Weberbauer, upon which I have 
drawn in part. 

I have not seen Dode's types, and separate the Peruvian material 
studied on the basis of the following key with great misgiving, feeling 
that the differences noted represent only individual variations or 
races. The problem can be solved only by abundant flowering and 
fruiting material from marked trees, since examination of the types 
alone cannot prove much. Practically, there is one acceptable 
name for the walnut of Peru (sens, lat.), Juglans neotropica Diels. 
The other names and descriptions are given for reference conven- 
ience, as the opportunity for further study of the problem may 
present itself. 
Bractlet (staminate) 3-3.5 mm. long, borne under the perianth; 

leaves (at least at flowering time) densely pilose beneath; 

stigmas fleshy, short; young fruits densely pubescent; fruit 

cells 6-8 J. neotropica. 

Bractlet reduced to a woolly tuft on the pedicel ; leaves at flowering 

time densely pubescent, especially beneath; stigmas slender; 

young fruits densely pubescent; lateral fruit cells 4 . . . J. Honorei. 
Bractlet minute, at the base of the receptacle; leaves soon glabrous or 

nearly so, the youngest minutely granular-puberulent; stigmas 

slender; young fruits sparsely pubescent; fruit cells 6-8. 

J. peruviana. 

Juglans Honorei Dode, Bull. Soc. Dendr. France 4: 205. 1909. 

Leaflets 6-15 pairs, ovate-lanceolate, unequally rotund-sub- 
cordate at the base, more or less abruptly acuminate, regularly 
serrate, scabrous-pubescent on both sides, the rough, branched hairs 
more numerous beneath (in youth densely velvety beneath); stami- 
nate aments to 20 cm. long, 1.5 cm. wide; perianth 6-8-lobed, with 
a 2-lobed involucre; anthers 60-70, pubescent at the tip; bractlet 
reduced to a rusty tuft of wool at about the middle of the pedicel; 
pistillate flowers 5-8; stigmatic branches slender; fruit subglobose- 
subconic, 4-5 cm. long, densely villous; nut rather remotely and 
obtusely rugose-costate, depressed at the base, subconic, scarcely 
compressed, to 3.5 cm. high. A tree of 25 meters or taller. The 
native name of "tocte" is recorded by Spruce. In Peru known only 



FLORA OF PERU 265 

from cultivation, but the cultivated specimen by Pavon in Herb. 
Madrid is determined by Markgraf as J. neotropica and, though 
sterile, probably is; the leaves are not truly scabrous but densely 
pubescent, especially on the veins beneath. Fruit illustrated, Dode, 
op. cit. pi. opp. p. 178, from nuts supplied by M. Honore" of Lima; 
leaves in plate on p. 169. 

Lima: Cultivated (Pavon). Ecuador; Colombia. 

Juglans neotropica Diels, Bot. Jahrb. 37: 398. 1906. 

A tree to 30 meters high, the stout young branchlets, especially 
at tip, and the leaf rachis at flowering time conspicuously rusty- 
pubescent; leaflets 7-8 pairs (-14 according to Dode), at first above 
sparsely, beneath densely velvety-pubescent, the largest middle 
leaflets about 12 cm. long and 4 cm. wide, gradually long-acuminate, 
minutely and evenly serrate; staminate aments 20-25 cm. long, 
the flowers partly remote, partly approximate; bractlets 2-3.5 mm. 
long, yellowish-brown-pilose; anthers apically pubescent; pistillate 
flowers 3-10; calyx urceolate, rusty- tomentose, 18 mm. long, 6-7 mm. 
broad, the narrow, reflexed teeth unequal, the larger 5 mm., the 
smaller 2 mm. long; perianth segments 4, reflexed, irregularly den- 
tate, 6 mm. long, 3-4 mm. wide; stigmatic branches broadly lingu- 
late, densely papillose, 8-9 mm. long, 4 mm. broad. Dode associates 
somewhat doubtfully with this species nuts collected by Ruiz and 
Pavon at Huanuco and figures them, op. cit. opposite p. 180. They 
are ovoid, about 4 cm. high, 3.5 cm. thick, moderately rugose, pointed, 
at base rounded, with 8 subelliptic cells. The Raimondi specimen is 
much less pubescent than the type. See also J. peruviana. Illus- 
trated, Weberbauer, pi. 13 opposite p. 199. Neg. 18254. 

Amazonas: Moyabamba, 2,700-3,300 meters, Williams 7606. 
Chachapoyas, 2,700 meters, Williams 7563. Utcubamba, near 
Chachapoyas, 2,000 meters, Weberbauer 4304, type. Lima: Culti- 
vated at Lima and Chancai, Ruiz & Pavon (det. Markgraf); "nueces 
de la Trexna," "nogal." Cajamarca: Chirinos, Raimondi (det. 
Markgraf). Huanuco: Posuso, 900 meters, Weberbauer 6753. 
Ayacucho: Rio Perene", 900 meters, Weberbauer 5632 (det. Nagel). 
Cuzco: Prov. Convention, Santa Ana and Marcapata (Herrera). 
"Nogal." 

Juglans peruviana Dode, Bull. Soc. Dendr. France 4: 208. 1909. 

Separated by the author from J. Honorei by the larger fruits, 
these globose-subconic, at least 5.5 cm. thick, and especially by 
the more numerous fruit cells, these 6-8 and occupying a very large 



266 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

space, particularly the basal ones; other characters given in the 
above key are taken from Weberbauer 6753, with no ripe fruits. 
This collector's 5632 may be the same. In view of the known 
variation in walnut fruits, J. peruviana is probably only a form 
of J. Honor ei, as Dode suggests; in this case the aberrant Weber- 
bauer material is to be taken merely as showing the range of variation 
in the unit J. neotropica, which compare for citation of above col- 
lections. Very similar is J. boliviano, (C. DC.) Dode (J. nigra L. 
var. C. DC.), the strongly ribbed nuts to 6 cm. high and the leaflets 
of the large leaves acuminate and unequal at base. 
Lima: Apparently collected by M. Honor (type). 

45. JULIANIACEAE. Juliania Family 

One of the two genera that constitute this interesting family, 
a group placed by Hemsley between the Juglandales and the Fagales, 
is exclusively Peruvian. In foliage, in the presence of resin, in 
the exalbuminous seed, and anatomically it resembles the Anacardia- 
ceae, but its resemblances with the Juglandaceae may be more 
fundamental, as for example the dissimilarity of the staminate and 
pistillate flowers. Rendle in his Classification of Flowering Plants 
follows Hemsley, but some botanists, as Standley in his Trees and 
Shrubs of Mexico, place it next to the Anacardiaceae. 

1. ORTHOPTERYGIUM Hemsl. 

Reference: Hemsley, Phil. Trans. Roy. Soc. London, Ser. B. 
199: 169-197. 1907. 

A shrub or small tree with dioecious flowers borne rather densely 
at the tips of the stout branchlets before or with the leaves. Leaves 
unequally pinnate. Fruit a samara, consisting of the persistent 
involucre borne on the solitary, wing-like stalk of the inflorescence, 
the wing straight and equal-sided. Asa Gray aptly likens the fruit 
to that of the ash (Fraxinus) inverted. 

Orthopterygium Huaucui (Gray) Hemsl. op. cit. 190. Juliania 
Huaucui Gray, Bot. U. S. Expl. Exped. 1: 371. 1854. 

Leaves at first tomentose, becoming glabrate above, usually 
7-foliolate; petioles several cm. long; leaflets crenulate, oblongish, 
about 1 cm. long; samaras pendulous, 11 cm. long, 8-10 mm. wide. 
The species name records incorrectly the vernacular name. 
Seldom seen without leaves, and always black as if burned or blasted 
(MacLean). The Weberbauer collections determined by Schellen- 
berg. Illustrated, Hemsley, op. cit. pi. 24- 



FLORA OF PERU 267 

Lima: Covering the sides of the base of the Cuesta de Purru- 
chuca, Prov. Canta, in 1831 (Mathews 591, type); (MacLean, 
presumably the type locality). Yanga (Wilkes Exped.). Chosica, 
1,600-1,900 meters, Weberbauer 5362, 5719, 5721, 5681, 5681 a; at 
900 meters, rocky hillside, 2866. Rio de Lomas, 1,000 meters, 
Weberbauer 5741- lea: Above Pisco, Huauyanga-Pampano, Weber- 
bauer 5371, 5372, 5373. Ayacucho: Coracora, 2,800 meters, Weber- 
bauer 5818. "Huancui," "huanarpu." 

46. BETULACEAE. Birch Family 
Reference: Winkler, Pflanzenreich IV. 61. 1904. 

1. ALNUS L. Alder 

The alder is one of the most important trees of the Peruvian 
Andes because it ascends to altitudes (3,500-3,800 meters, according 
to Weberbauer) where it is especially valuable for fuel and for 
small construction. It is sometimes planted. Herrera has recorded 
some form (erroneously as A. acuminata HBK.) as "cultivated 
in great abundance in all the quebradas of the Department of Cuzco, 
its wood greatly valued for all construction purposes." Ruiz and 
Pavon found Betula alba L. cultivated about the convent at Huerta 
de Ocopa near Tarma, one example having nine very tall trunks. 
Weberbauer has reproduced a good photograph of the Peruvian 
alder, showing it in a typical situation Pflanzenw. Peru. Anden, 
opposite p. 295. 

Alnus jorullensis HBK. Nov. Gen. & Sp. 2: 20. 1817; 126. 

Typically Mexican, var. typica Regel having elliptic-oblong 
leaves 5-10 cm. long and 2.5-5 cm. wide, pale rusty-pubescent be- 
neath, this variable species is represented in Peru by several varia- 
tions, two of which are well marked: var. castaneifolia (Mirb.) 
Regel (castaneaefolia) with glabrous (or pubescent in the nerve 
axils), ovate-lanceolate leaves 8-10 cm. long and 1.5-3 cm. wide, 
or larger on the sterile shoots; and var. ferruginea (HBK.) Kuntze, 
with ovate leaves 10 cm. long and 3-6 cm. wide, or twice as large 
on young branches, these and the prominent leaf nerves beneath 
rusty-pilose. Scarcely distinguishable from the last are vars. Mirbelii 
(Spach) Winkl. and acutissima Winkl., the former with coarsely 
serrate leaves, glaucous and pale rusty-pubescent beneath, the latter 
with ovate, acutely acuminate, denticulate leaves having the 13-18 
nerves impressed above but prominent and pilose beneath. 



268 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ruiz and Pavon collected the tree at Pillao and Chacahuasi, 
noting that an infusion of the inner bark tans leather and dyes 
cotton and wool a cinnamon-brown; the leaves crushed with butter 
cicatrize wounds and without fat protect against inflammation; 
applied to recent wounds the leaves stop bleeding. Illustrated, 
Me"m. Mus. Paris 14: 463. pi. 21, 22; vars. castaneifolia and Mirbelii, 
Sargent, Sylva N. Amer. 9: pi. 457. Neg. 11652. 

Cajamarca: Huaraz, 2,200 meters (Weberbauer 172, 179). 
Ancash : Caracha, 1,200 meters, Weberbauer 2650; 162, 168. Samanco 
(Weberbauer 171). Amazonas: Utcubamba (Weberbauer 192). 
Huanuco: Fifteen miles southeast of Huanuco, 2083. Huanuco, 
Kanehira 34- Mito, 2,700 meters, 1527, 1907. Pillao, Ruiz & Pavon 
(var. castaneifolia, det. Mildbraed). Chancai, Huanuco, and Cochero, 
Ruiz & Pavon (var. ferruginea, det. Mildbraed). Huanuco Valley, 
Poeppig (var. acutissima'} . Lima: Matucana, 2,400 meters, 561; 
Weberbauer 182 (var. acutissima). Junin: Tarma, Esposto; at 3,600 
meters (Weberbauer 177, 183); at 2,100 meters, 1021. Huancayo, 
Esposto. Carpapata, edge of forest, 2,700-3,200 meters, Killip & 
Smith 24480; a tree of 4.5-7.5 meters. Ocopa, 3,300 meters, Killip 
& Smith 22008; a tree of 6-12 meters, by roadside. Cuzco: Rio 
Apurimac, 2,800 meters, Weberbauer 5892 (det. Schellenberg). Uru- 
bamba, 2,800 meters (Weberbauer 174, 182, 243). Prov. Quispi- 
canchi, 3,200 meters, Herrera 650 (var. acutissima, det. Mansfeld). 
Calca, Valle del Urubamba, 3,000 meters, Herrera 2092. Libertad : 
Valley of Rio Mixiollo, 2,400 meters, Weberbauer 7040. Puno: 
Sandia, 3,200 meters (Weberbauer 184). Tabina, Lechler 1891 (var. 
Mirbelii, fide Winkler). Without locality, Weberbauer 7040. 
Argentina and Bolivia to Mexico. "Aliso," "ramram," "lambran." 

47. ULMACEAE. Elm Family 
By Charles Baehni 

The Peruvian plants of this family all belong to the Celtideae, 
a group in which the fruit is a drupe containing a curved embryo, a 
character not found in the rest of the family. 
Stamens as many as the perianth segments. 
Leaves alternate. 
Stamens deciduous, included in the minute flowers; leaves 

precocious. 

Staminate flower segments induplicate-valvate, the pistillate 
deciduous.. . .1. Trema. 



FLORA OF PERU 269 

Staminate flower segments imbricate, the pistillate persistent. 

2. Celtis. 
Stamens persistent; leaves appearing after the flowers. 

3. Plagioceltis. 

Leaves opposite 4. Lozanella. 

Stamens twice as many as the perianth segments 5. Ampelocera. 

1. TREMA Lour. 

Sponia Comm. ex Lam. Encycl. 4: 139. 1797. 

Scarcely distinct from Celtis, the flowers sometimes perfect and 
with somewhat imbricate segments. Perianth persisting around 
the drupe. Species poorly understood or very variable in pubes- 
cence and size and discoloration of leaves; cf. Planchon in DC. 
Prodr. 17: 203, under Sponia. 

Trema micrantha (L.) Blume, Mus. Bot. Lugd. Bat. 2: 58. 1853. 
Rhamnus micranthus L. Syst. ed. 10. 937. 1759. Celtis micranthus 
Sw. Prodr. 53. 1788. Sponia micrantha Dene. Nouv. Ann. Mus. 
Paris 3: 498. 1834. C. Lima Sw. loc. cit., non Lam. C. canescens 
HBK. Nov. Gen. & Sp. 2: 28. 1817. C. canescens Dene. op. cit. 
C. macrophylla HBK. op. cit. 30. S. macrophylla Dene. op. cit. 
T. canescens Blume, op. cit. S. peruviana Klotzsch, Linnaea 20: 536. 
1847. S. Chichilea Planch. Ann. Sci. Nat. III. 10: 334. 1849 (not 
1848 as usually cited). T. Chichilea Blume, op. cit. 

An unarmed tree, usually a few meters tall; trunk sometimes 
20-40 cm. in diameter; leaves usually lanceolate, acuminate, minutely 
serrulate, 3-nerved, more or less scabrous above, merely scabrous or 
glabrate to densely and softly pilose beneath, about 10 cm. long; 
flowers greenish white, red to fiery red (Weberbauer). The above 
synonymy is partial, applying to Peruvian specimens cited or found 
in herbaria. Illustrated, Fawc. & Rendle, Fl. Jamaica 1 : 39. Negs. 
25569 (T. Chichilea), 11238 (T. canescens). 

Cuzco: Santa Ana, 1,200 meters, Weberbauer 5038. Rio Pachaca, 
1,100 meters, Weberbauer 5887. Machupicchu, 2,200 meters, Herrera 
3240. Huanuco: Monzon, 900 meters, Weberbauer 3433. Casapi, 
Mathews 2038. Cochero, Poeppig 155, 1247. Pampayacu, Kane- 
hira192. Vilcabamba, 1,800 meters, 5132. Lima(?): Quebrada de 
Pariahuanca (Mathews 829, var. of canescens ace. to Planchon). 
San Martin: Tarapoto, Spruce 4242. Near Moyobamba, 1,110 
meters, Klug 3261 (det. Standley). Lamas, 840 meters, Williams 
6446. Cumbasa, Williams 5765. San Roque, Williams 7376. 



270 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Florida: Rio Putumayo, 180 meters, King 2144 (det. 
Standley). Lower Rio Huallaga, Williams 5006. Mouth of Rio 
Santiago, Tessmann 4431. Yarina-cocha, Tessmann 3227 (glabrate), 
3227a (pilose). Rio Nanay, Williams 368, 366. Rio Santiago, 200 
meters, Mexia 6308, 6238. La Victoria, Williams 3088, 2591 . For- 
taleza, 200 meters, Williams 4225. Rio Masana, Williams 194, 80. 
Puerto Arturo, 135 meters, Killip & Smith 27860; Williams 5152. 
Pebas, Williams 1759. Pinto-cocha, Williams 811. Soledad, 110 
meters, Killip & Smith 29778. Mishuyacu, 100 meters, Klug 1314. 
Iquitos, Killip & Smith 27387. Punchana, Williams 3755. Junin: 
Colonia Perene", 680 meters, Killip & Smith 25021. Chanchamayo 
Valley, 1,500 meters, Schunke 201. Above San Ramon, 1,400-1,700 
meters, Killip & Smith 24621. La Merced, 600 meters, 5229. 
Without locality; Ruiz & Pavon (type of T. Chichilea); Dombey; 
Pavdn. Widely distributed in tropical and subtropical America. 
"Aisegerina" (Huitoto name), "atadijo," "y ana - cas pi-" 

2. CELTIS L. 

References: Planchon, Ann. Sci. Nat. III. 10: 309, 1849; Miquel 
in Mart. Fl. Bras. 4,pt. 1: 173. 1853; Planchon in DC. Prodr. 17: 186. 
1873; Baehni, Candollea 7: 189. 1936. 

Trees, shrubs, or lianas, usually armed with small, inconspicuous 
flowers in axillary clusters or cymes. Perianth 4-5-lobed, deciduous. 
Stigma divided into 2 branches to the base, sometimes each branch 
again 2-cleft. Fruit a drupe. The number of described species is 
rather high. It seems, however, that they all belong to a few well 
defined groups which are considered here as species. Ruiz and 
Pavon record the name "chichillica" for an unindentified species 
from Muna, the bark of which was used for coarse basketry and for 
cords. This reference may well refer to Trema micrantha. 
Adult leaves (except on the nerves) glabrous or practically so. 

Fruit large (6-12 mm. in diam.); young leaves sericeous. 

C. triflora. 

Fruit small (4-6 mm. in diam.) ; young leaves pilose C. iguanea. 

Adult leaves pubescent. 

Inflorescence well developed (3 cm. long or more) C. dichotoma. 

Inflorescence short or glomerulous C. pubescens. 

Celtis dichotoma (Klotzsch) Miq. in Mart. Fl. Bras. 4, pt. 1: 
182. 1853. Momisia dichotoma Klotzsch, Linnaea 20: 539. 1847. C. 
Pavonii Planch. Ann. Sci. Nat. III. 10: 313. 1849. 



FLORA OF PERU 271 

Branchlets, petioles, and young leaves beneath densely covered 
with an indument of golden yellowish (sometimes rusty) hairs; 
spines wanting or very short; leaves ovate or oblong-ovate, acute or 
acuminate, cordate, entire or coarsely dentate toward the apex, 
glabrate in age beneath; staminate inflorescences many-flowered, 
many times longer than the petioles. Fruits large, globose, glabrous, 
yellow and sweet, and (according to Ruiz and Pavon) eaten by the 
Indians. Negs. 11739, 25566 (C. Pavonii). 

Huanuco: Chacahuasi, Posuso, etc., Ruiz & Pavon. 

Celtis iguanea (Jacq.) Sarg. Silva 7: 64. 1895. Rhamnus 
iguaneus Jacq. Enum. PL Carib. 16. 1760. C. aculeatus Sw. Prodr. 
53. 1788. Mertensia laevigata HBK. Nov. Gen. & Sp. 2: 31. 1817. 

A small tree, a shrub, or a liana, glabrous or essentially so except 
for some evanescent pubescence on the young branchlets and leaves 
beneath, usually armed, the spines well developed, geminate; leaves 
ovate or oval-elliptic, entire or remotely serrate toward the tip, the 
young ones pilose, glabrous in age; cymes short, rarely two or three 
times longer than the petioles; fruit small, 4-6 mm. in diameter. 
Illustrated, HBK. Nov. Gen. & Sp. 2: pi. 103. 

San Martin: Tarapoto, Spruce 4236. Alto Rio Huallaga, Williams 
6756. Near Moyobamba, King 3303. Tarapoto, Williams 6666. 
Juan Guerra, 720 meters, Williams 6846. Lamas, 840 meters, 
Williams 6357. Rumizapa, Williams 6784- Loreto: Yarina-cocha, 
Tessmann 3428. Mishuyacu, 100 meters, Klug 1435. Paraiso, 145 
meters, Williams 3365. Junin: La Merced, 700 meters, Killip & 
Smith 24079. Piura: Serran, Weberbauer 6000. Tumbez : Between 
Ricaplaya and Casa Blanqueada, Weberbauer 7741- A species 
widely distributed in South and Central America, West Indies, and 
north to Mexico and Florida. "Palo bianco," "meloncito bianco." 

Geltis pubescens (Humb. & Bonpl.) Spreng. Syst. 1: 931. 1825. 
Mertensia pubescens Humb. & JBonpl. in Schult. Syst. 6: 312. 1820. 
Mertensia brasiliensis Gardn. Lond. Journ. Bot. 2: 339. 1843. 
Momisia brevifolia Klotzsch, Linnaea 20: 538. 1847. C. boliviensis 
Planch. Ann. Sci. Nat. III. 10: 310. 1849. C. velutina Planch, op. 
cit. 313. Momisia flexuosa Wedd. Ann. Sci. Nat. III. 18: 194. 1852. 
M. crenata Wedd. op. cit. 195. 

A tree, a shrub, or a liana, with flexuous branchlets, the young 
twigs puberulous or velvety, armed with straight or slightly curved, 
paired spines; leaves ovate or elliptic-ovate, acute or acuminate, 
cordate or rounded at the base, entire or serrate toward the tip, the 



272 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

young ones often with golden hairs, in age glabrous above, softly 
pubescent beneath; cymes short, not much longer than the petioles; 
fruit small, glabrescent, rough when dry. The synonymy above is 
partial, applying to Peruvian specimens only. Negs. 29617 (crenata), 
29664 (pubescens). 

Junin: Huancayo, Raimondi 9177. Between Sandia and Chun- 
chusmayo, Azalaya, Weberbauer 1126. Chanchamayo, Isern 2322. 
La Merced, 600 meters, 5438, 5280; Killip & Smith 24042, 23406. 
San Martin: Juan Guerra, 720 meters, Williams 6871. Rio Acre: Ule 
9334. Lima(?): Quebrada de Pariahuanca, Mathews 826. Huan- 
cavelica: On the Montaro, upstream from Colcabamba, Weberbauer 
6437. Cajamarca: Prov. Contumaza, Cascas, Raimondi 7973. 
Tropical South America, widely distributed. 

Celtis triflora (Klotzsch) Miq. in Mart. Fl. Bras. 4, pt. 1: 181. 
1853. Momisia triflora Klotzsch, Linnaea 20: 537. 1847. C. glycy- 
carpa Mart, ex Miq. op. cit. 174. 

Branches and branchlets glabrous or evanescently pilose, brown- 
ish, the stout spines mostly solitary, slightly curved ; leaves ovate or 
oblong-elliptic, acuminate, subcordate or cordate, entire or serrate 
toward the tip, asperous above, glabrescent beneath; fruit large, 
globose, smooth. A tree, up to 12 meters high. 

Huanuco: Posuso, Ruiz & Pavon. Loreto: Mouth of Rio 
Santiago, Pongo de Manseriche, Tessmann 4274- Bolivia; Brazil; 
Venezuela. 

3. PLAGIOCELTIS Mildbr., gen. nov. 

A stoutly branched shrub with light yellowish brown bark. 
Leaves appearing with the flowers, or immediately after. Racemes 
terminal or nodal on the leafless branches of the preceding year. 

Plagioceltis dichotoma Mildbr. in Herb. Madrid, sp. nov. 

Frutex racemis atque ramulis novellis exceptis glaber; folia 
juvenilia 5 mm. longe petiolata tenuia subelliptica remotissime 
dentata acuta basi attenuata ad 7 cm. longa et ultra; racemi saepe 
1-ramosi, bracteolis subrotundis; flores circa 2 mm. longi, sepalis 
rotundatis, staminibus persistentibus; ovarium pilosum, stylo 
staminibus longiore. 

Glabrous except for the evanescently puberulent growing branch- 
lets and racemes, the latter often once branched, 1-1.5 cm. long; 
petioles 5 mm. long; leaves (young) thin, very remotely toothed, 
subelliptic, attenuate at the base, acute, to 7 cm. long or longer; 



FLORA OF PERU 273 

bractlets roundish; flowers about 2 mm. long, the pilose ovary soon 
exserted with the conspicuous style from the persistent stamens and 
roundish petals. Description from type in Herb. Madrid. 
Without locality: Ruiz & Pavon. 

4. LOZANELLA Greenm. 

Unarmed trees or shrubs with opposite, serrate leaves and dioe- 
cious inflorescences; fruit a small drupe containing a curved embryo 
with oblong-rotund, scarcely curved cotyledons. The two species 
of this genus are readily distinguished from all other members of the 
Ulmaceae on account of their opposite leaves. 
Adult leaves not densely pubescent beneath; pistillate inflorescence 

conspicuously branched L. enantiophylla. 

Adult leaves densely pubescent beneath ; pistillate inflorescence with 

short lateral branches L. permollis. 

Lozanella enantiophylla (Bonn. Smith) Killip & Morton, 
Journ. Wash. Acad. Sci. 21: 14: 339. 1931. Trema enantiophylla 
Bonn. Smith, Bot. Gaz. 33: 339. 1902. L. trematoides Greenm. Proc. 
Amer. Acad. 41:236. 1905. 

A small tree, 3-7 meters high, or a shrub, with long-petioled, 
ovate to oblong-ovate, leaves, united, deciduous stipules, and open, 
much branched cymes. 

Libertad: Prov. Pataz, Rio Mishiolla Valley, 2,000 meters, 
Weberbauer 7049. Mexico, Guatemala, Colombia. 

Lozanella permollis Killip & Morton, Journ. Wash. Acad. 
Sci. 21:338. 1931. 

A tree up to 8 meters high, the young branchlets covered with 
densely matted hairs; leaves oblique at the base, elliptic, densely 
covered on the veins beneath with long, appressed hairs; fruit a 
small drupe, the styles persistent. 

Cuzco: Lucumayo Valley, Cook & Gilbert 1376. This species has 
been collected several times in Bolivia. 

5. AMPELOCERA Klotzsch 

Reference: Klotzsch, Linnaea 20: 542. 1847. 

An unarmed shrub with remotely serrate, pinnately nerved 
leaves. Flowers articulate, in axillary racemes, mostly geminate, 
corymbose, polygamous, perfect, or staminate by the abortion of 
the ovary. Filaments filiform, exserted. Ovary ovate, the stigma 



274 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

deeply 2-parted, the subulate branches divaricate; ovule pendulous, 
the micropyle superior. Seed arcuate. The following species is the 
type of the genus, to which Grisebach added a Cuban species, with 
some doubt accepted by Urban, Repert. Sp. Nov. 15: 109. Both 
Bentham and Hooker in Genera plantarum and Engler in Pflanzen- 
familien have questioned the disposition of the genus in this family, 
from which Planchon in DC. Prodr. 17: 152 definitely excludes it. 
However, so far as I know, no one has yet placed it elsewhere, and 
Kuhlmann, Arch. Jard. Rio Janeiro, has described two Brazilian 
species which seem to establish it. It may not be Peruvian. The 
genus is placed in Ulmaceae on anatomical grounds by Priemer (?), 
Bot. Jahrb. 17: 466. 1893. The specimen he saw presumably was 
the original from the Lambert Herbarium, but no specimen has 
been found by me in Madrid material, Delessert Herb., or Berlin. 

Ampelocera Ruizii Klotzsch, loc. cit. 

Branches robust, squarrose, the branchlets slender, erect, evanes- 
cently pubescent; leaves short-petioled, oblong or obovate, attenu- 
ate at both ends, glabrous above, glabrate beneath, 5-9 cm. long, 
2.5-3.5 cm. wide; stipules very thin, 2 mm. wide, 6 mm. long; 
racemes 2.5 cm. long, evanescently pubescent; perianth divisions 
obovate, dentate, persistent; style white-tomentose. 

Without locality: (Ruiz & Paron). 

48. MORACEAE. Mulberry Family 

A family of great economic importance, especially in Central 
America, where one member, Castillo,, is a source of rubber. In 
Peru several trees are, at least potentially, of value for their timber, 
notably the tupag or guariuba (Clarisia) with fine-grained, yellowish 
wood and the aita (several species of Brosimum), the wood white, 
on exposure reddish brown. Most of the trees exude a milky latex 
more or less elastic when dried, and discoloring. It has often some 
local use in medicine or otherwise. 

At least two trees of the family are in cultivation: the mulberry 
(Morns') in the coastal valley, and occasionally the breadfruit, arbol 
de pan (Artocarpus incisa), as for example at La Merced. 

The taxonomy of the family is in need of revision. Ducke, 
Archiv. Jard. Bot. Rio Jan. 4: 6-8. 1925, and in previous and sub- 
sequent volumes of the same publication, has given discriminating 
diagnoses and helpful observations on which I have freely drawn. 
In Arch. Inst. Biol. Veg. 2: 29. 1935, he indicates that he considers 



FLORA OF PERU 275 

as excessive the number of genera in the Olmedieae. In the following 
synopsis of Peruvian forms the present day grouping is followed for 
convenience but not in approval, for it results in wide separation of 
similar trees and is absurdly impractical, especially in an economi- 
cally important family. To facilitate determination and especially 
to make clear the dovetailing of characters, the key for genera having 
completely staminate or completely pistillate receptacles is divided, 
one part being exclusively for staminate inflorescences, the other 
for the pistillate, much herbarium material showing only one sort. 
At the end of the general key is one keying all these genera as though 
they pertained to one group. See Field Mus. Bot. 11: 62-63. 1931, 
and remarks under some generic descriptions regarding possible 
relationships. 

Low herbs 1. Dorstenia. 

Trees or shrubs. 
Leaves palmately lobed or divided. 

Leaves somewhat peltate; flowers spicate 2. Cecropia. 

Leaves not peltate; flowers cymose 3. Pourouma. 

Leaves entire, serrate, or lobed but not palmately. 
Flowers on the inner surface of a hollow receptacle open only at 

the top of the receptacle, i.e., a fig. . . . 5. Ficus. 

Flowers not so disposed. 
Flowers spicate or racemose, at least one sort. 
Staminate flowers spicate, the spikes dense; pistillate capi- 
tate or, in Trophis, in short spikes. 
Trees often spiny; leaves never much longer than 10 

cm., often much shorter. 
Filaments exserted; pistillate flowers capitate. 

8. Chlorophora. 
Filaments included or barely exserted; pistillate 

flowers spicate 6. Trophis, 

Trees unarmed, the leaves ample, often 20 cm. long or 

longer; pistillate flowers capitate. 11. Anonocarpus. 

Staminate and pistillate flowers in aments, or the latter 

rarely capitulate but few, or even solitary (Clarisia). 
Staminate perianth present; leaves often somewhat 

toothed. 

Filaments not inflexed; aments lax; pistillate perianth 
tubular or ovoid . . . . 9. Sorocea. 



276 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Filaments in bud inflexed; aments dense, at least 
the pistillate, these fleshy in fruit. 

Pistillate perianth tubular, accrescent; staminate 
segments valvate; leaves usually entire. 

6. Trophis. 

Pistillate perianth segments distinct, the staminate 
imbricate; fruit juicy; leaves serrate. .7. Moras. 

Staminate perianth absent; leaves strictly entire. 

10. Clarisia. 

Flowers not in aments (cf. also Clarisia, sometimes, and 
Chlorophora as to pistillate flowers). 

Petioles mostly under 1 cm. long or if some to about 1.5 
cm., these in the minority; ovules pendulous. 

Receptacles head-like, composed of many stamens and 
1-4 pistils, these deeply inserted. 

Stamens exserted, crowded at throat of the receptacle 
about the exserted style 20. Trymatococcus. 

Stamens separated in all the receptacles by peltate 
bracts 12. Brosimum. 

Receptacles often capitate but completely staminate or 

pistillate. 

Pistillate receptacles (see below for key to staminate). 
Receptacle 1-flowered; styles filiform (cf. Olmedi- 

operebea under Pseudolmedia). 
Ovary inferior, fixed to the perianth. 

17. Pseudolmedia. 

Ovary superior, free in the perianth . . 18. Olmedia. 
Receptacle many-flowered; styles thick to filiform 

(cf. Olmedioperebea) . 
Styles thick, short, the stigma compressed or 

capitate-cushion-like. 

Perianth truncate, compressed, in aggregate 
forming an Anona-like fruit. 

11. Anonocarpus. 

Perianth 4-toothed, the fruit fleshy, but scarcely 
Anona-like (here also Olmedioperebea) . 

14. Perebea. 
Styles or at least stigmas slender or filiform. 



FLORA OF PERU 277 

Perianth lacking; bracteoles peltate (probably 

in Peru) Brosimopsis. 

Perianth present, sometimes pseudobracteolate. 

Perianth segments divided to the base, the 

parts thus bracteole- or scale-like in the 

receptacle 13. Ogcodeia. 

Perianth merely lobed or parted. 
Perianth segments imbricate; receptacles 
and young parts of plant reddish- 

tomentose 19. Helicostylis. 

Perianth segments various; pubescence, if 
present, not characteristically reddish- 
tomentose. 

Perianth 4-lobed, the lobes not perfor- 
ated; bracts all small. .15. Castillo,. 
Perianth segments perforated; inner 

bracts elongate 16. Noyera. 

Staminate receptacles. 

Perianth absent but the bracteoles may form a 

pseudoperianth . 
Bracts seriate, the inner not elongate; leaves 

hirsute above or beneath 15. Castillo,. 

Bracts toward the interior often longer than the 
outer; leaves often smooth or, if hirsutulous, 
only sparsely. 
Bracteoles obvious, spatulate or peltate. 

17. Pseudolmedia. 

Bracteoles (or divided perianths) scalelike; see 

Naucleopsis mentioned under 13. Ogcodeia. 

Perianth present (deeply parted perianths may 

simulate bracteoles). 
Stamens interspersed with peltate bracteoles. 

Brosimopsis. 
Stamens and bracteoles not so intermixed, the 

latter, if not wanting, not peltate. 
Bracts all small, the interior little longer than 

the outer. 
Plant pubescence reddish brown. 

19. Helicostylis. 



278 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plant pubescence, if present, not at all or not 

very reddish. 
Leaves usually scabrous above. 

17. Olmedia. 

Leaves not scabrous above . . 14. Perebea. 
Bracts unequal, the interior longer than the 

outer, closely seriate. 

Leaves glabrous or nearly so; receptacles 

shortly pediceled or sessile . . 13. Ogcodeia. 

Leaves harshly hirsute above; receptacles 

long-pediceled 15. Noyera. 

Petioles mostly 2 cm. long or longer; ovule erect. 
Flowers cymose or the staminate capitulate and cymose; 

stamens 3-4; stigma peltate 3. Pourouma. 

Flowers globose-capitate, the heads single or cymose; 
stamens 1-2; stigma penicillate 4. Coussapoa. 

Aggregate Key to the Species of Genera 13-19, Inclusive 
Leaves scabrous-puberulent, at least on one side. 
Leaves scabrous on both sides. 
Leaves entire; staminate peduncles 5-7 mm. long. 

17. Pseudolmedia scabra. 
Leaves generally undulate-serrate; staminate receptacles sessile 

or subsessile 18. Olmedia aspera. 

Leaves scabrous only beneath. 

Leaves undulate-serrate; staminate receptacle sessile or sub- 
sessile 18. Olmedia Poeppigiana. 

Leaves entire; staminate peduncles to 6 mm. long. 

14. Perebea Chimiqua. 

Leaves glabrous to variously pubescent but not scabrous only. 
Leaves strictly glabrous. 

Leaves mostly or all narrower than 3 cm. 
Leaves caudately acuminate; ovary superior. 

18. Olmedia angustifolia. 
Leaves obtusely acuminate; ovary inferior. 

17. Pseudolmedia laevigata. 
Leaves mostly or all 3.5-6 cm. wide or wider. 

Branchlets spreading-hirsute 17. Pseudolmedia laevis. 



FLORA OF PERU 279 

Branchlets glabrous or puberulent. 
Petioles to 3 mm. long; leaves somewhat acuminate. 

17. Pseudolmedia Mildbraedii. 
Petioles mostly or all much longer; leaves caudate. 
Leaves rarely 6.5 cm. wide, the lateral nerves 15-20. 
Petioles rarely 1 cm. long; nerves beneath rather 

inconspicuous. 
Leaves 3-4 cm. wide, cuneate at the base. 

13. Ogcodeia ternstroemiiflora. 
Leaves 5-6 cm. wide, rounded at the base. 

13. Ogcodeia Tamamuri. 

Petioles often longer; nerves beneath conspicuous. 
Leaves rarely 5 cm. wide, the acumen often 2 cm. 

long 13. Ogcodeia Tessmannii. 

Leaves often 6-7 cm. wide, the acumen rarely 1.5 

cm. long 13. Ogcodeia glabra. 

Leaves mostly 8-12 cm. wide, the lateral nerves 25-30. 

13. Ogcodeia Ulei. 

Leaves more or less pubescent, at least on the nerves beneath. 
Pubescence, at least on the midnerve beneath, setulose or 
hirsute. 

Leaves acute at base or, if subcordate, not at all pseudo- 
peltate. 
Petioles 2-5 mm. long. 

Leaves strongly pubescent on both sides; interior bracts 
of both sorts of receptacles elongate. 

16. Noyera mollis. 

Leaves glabrous (except Castillo) or finally glabrate 
above; interior bracts little or not at all longer than 
the outer. 

Leaves, unless in age, harshly pubescent above; pistil- 
late receptacles several-flowered . . 15. Castilla Ulei. 
Leaves glabrous or glabrate above; pistillate receptacle 

1-flowered. 
Pubescence of plant appressed. 

17. Pseudolmedia macrophylla. 
Pubescence of plant hirtellous, sparse. 

17. Pseudolmedia laevis. 



280 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petioles 8-12 mm. long. 
Leaves gradually acuminate, 3-5 cm. wide; staminate 

perianth lacking 17. Pseudolmedia multinervis. 

Leaves subcaudate-acuminate, 10-20 cm. wide. 
Leaves not oblique at base; stigmas cushion-like. 

14. Perebea australis. 
Leaves oblique at base; stigmas Ungulate. 

14. Perebea Tessmannii. 
Leaves appearing peltate at the subcordate base. 

14. Perebea pseudopeltata. 

Pubescence merely a puberulence, or a reddish tomentum. 
Pubescence not a reddish tomentum. 
Petioles 2-5 mm. long; staminate receptacle yellowish- 
pubescent, a perianth present. 

17. Pseudolmedia macrophylla. 
Petioles 5-7 mm. long, the receptacles not so pubescent. 

Leaves about 5 cm. wide .14. Perebea elegans. 

Leaves about twice as wide 14. Perebea Standleyi. 

Pubescence a reddish tomentum. . .19. Helicostylis tomentosa. 

1. DORSTENIA L. 

Stemless or subprostrate herbs from thick, tuber-like rootstocks. 
Receptacles large, rounded or lobed, on long or short peduncles. 

Plants stemless; receptacles 4-sided. 

Peduncles and petioles elongate, subequal D. Contrajerva. 

Peduncles shorter than the short petioles D. tubicina. 

Plants caulescent, subprostrate; receptacles orbicular . .D. umbricola. 

Dorstenia Contrajerva L. Sp. PI. 121. 1753. 

Leaves varying from entire to deeply lobed, ovate to suborbi- 
cular, glabrous to pubescent, often 10 cm. long; petioles often 
nearly as long or longer; receptacles green, entire or lobed, borne on 
slender, erect peduncles usually 10-30 cm. long. Known as "contra- 
hierba" in Central America, where in some places the thick rootstocks 
are used to flavor cigarettes (Standley). Illustrated, Bailey, Cycl. 
Hort. 1067. 

San Martin: Tarapoto, Ule 6509; Williams 5905. Rumizapa, 
Williams 6832. Extending to Mexico and the West Indies. 



FLORA OF PERU 281 

Dorstenia tubicina R. & P. Fl. 1: 65. pi. 102. 1798. 

Leaves cordate-ovate or cordate-oblong, doubly serrate, rough 
above, soft-pubescent beneath; receptacles violet in flower, white 
in fruit, recurving on peduncles shorter than the short petioles. 
The fragrant roots are used as "contrahierba." Illustrated, Bot. 
Mag. pi. 2804, from specimens presumably from Trinidad. Neg. 
11614. 

Huanuco: Chinchao and environs, Ruiz. Argentina. "Carpales." 

Dorstenia umbricola A. C. Smith, Bull. Torrey Club 58: 87. 
1931. 

Leaves green, oblong-lanceolate or obovate-lanceolate, cuneate 
or truncate at the base, acute, entire or undulate-crenate, glabrous 
above, sparsely pilose on the nerves beneath, 8-10 cm. long, on 
slender petioles 5-13 mm. long; peduncles at maturity 2.2 cm. 
long; receptacles (mature) 1.5 cm. broad, the staminate and pistillate 
flowers separated. Perhaps a variety of D. argentata Hook, f., 
with variegated leaves and fleshier receptacles, more definitely 
tuberculate marginally. 

Loreto: Soledad on Rio Itaya, Killip & Smith 29639, type. 

2. CECROPIA L. 

Beautiful and common trees of the montana, immediately recog- 
nizable by their smooth, light-colored trunks and branches supporting 
open crowns that bear abundantly large or even huge, long-stalked, 
palmately divided leaves that are usually dark green above and 
silvery or gray-tomentose beneath. Ruiz and Pavon observed that 
the handsome leaves follow the course of the sun, the position of the 
upper surface at night becoming partly reversed. Parts, at least, 
of the hollow trunks and branches appear to be regularly inhabited 
by black ants that rush out almost instantly upon the slightest dis- 
turbance in countless hordes to defend their home. Ruiz and Pavon 
noted, however, that sometimes the hollow, articulated branches or 
trunks are filled with clear, perfectly tasteless sap which the Indians, 
when other water lacked, drank to quench their thirst. The trunks 
are said to be used for tibias or trumpets in the Amazon region, and 
the fiber of the bark has been employed in making mats and coarse 
cloth. The usual common name for the tree in Peru seems to be 
"tacuna," written also "tacuma" and "tacona," or "setico," but 
Spruce found "imbauba" used on the Amazon, and Wallace, who 
observed the Indians using the leaves as a head-dress, recorded the 



282 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

term as "umbooba." Weberbauer, 275, gives a good photograph of 
slender Cecropia trees. It may be helpful to call attention to Pou- 
rouma cecropiaefolia, which in foliage alone simulates Cecropia. 
Leaves with many, usually 12-16, lobes or leaflets. 
Leaflets distinctly petioled or entirely separate, paler but not 

white-pubescent beneath C. sciadophylla. 

Leaflets more or less joined (or the leaves merely lobed), at least 
at the very base, and white- or grayish-pubescent beneath, 
at least between the veins. 
Leaves white-tomentose, pilose, or sericeous-pubescent above, 

not scabrous or not scabrous only. 
Leaves typically 10-11-lobed, greenish above. 
Leaves divided nearly to the base; staminate spikes many. 

C. leucophaea. 
Leaves often only three-fourths divided; staminate spikes 

about 4 C. acutifolia. 

Leaves typically 12-13-lobed, white-tomentose above. 

C. albicans. 

Leaves scabrous above. 
Leaves white- or grayish-tomentose beneath. 

Leaves typically 11-16-lobed C. angustifolia. 

Leaves typically 10-lobed. 

Petioles and leaf nerves hirtellous beneath. . . .C. strigosa. 
Petioles and leaf nerves tomentulose beneath. 

C. montana. 

Leaves reddish-hirsutulous, somewhat white-pubescent be- 
neath C. tubulosa. 

Leaves with 5-11 lobes, rarely parted to the base. 
Leaf lobes entire or merely undulate-repand. 
Leaves more or less pubescent above, usually scabrous. 
Lobes of the leaves typically 10 or 11. 
Leaves softly pubescent above, the scabrosity, if any, 
obscured. 

Leaves parted almost to the base C. leucophaea. 

Leaves merely deeply lobed C. acutifolia. 

Leaves scabrous-pubescent above, other pubescence not 

marked. 
Petioles and leaves beneath more or less hirtellous. 



FLORA OF PERU 283 

Leaves shallowly lobed C. strigosa. 

Leaves very deeply lobed C. tubulosa. 

Petioles and leaves beneath merely white-lanuginose. 

C. montana. 

Lobes of the leaves typically 8 or 9, or fewer. 
Leaf lobes typically 8 or 9. 
Leaves corrugated-rugose above. 
Leaves concolorous beneath, the nerves villous. 

C. nivea. 

Leaves with nerves and veins beneath green or brown. 

C. Klotzschiana. 
Leaves not all or not definitely rugose above. 

Leaf nerves many (20-25) and mostly 1 cm. or less 

apart. 
Staminate spikes many; leaves shallowly lobed. 

C. latiloba. 

Staminate spikes few; leaves deeply lobed. 
Leaves somewhat floccose or more floccose than 

scabrous above C. leucocoma. 

Leaves obviously scabrous above C. bicolor. 

Leaf nerves 12-15 and mostly more than 1 cm. apart. 
Staminate spikes many (-50) ; leaves asperous above. 

C. multi flora. 
Staminate spikes few (-20); leaves often floccose 

above. 

Leaves hirtellous on the nerves beneath . .C. bicolor. 
Leaves tomentose even on the nerves beneath. 

C. obtusa. 

Leaf lobes typically 5, 6, or 7. 

Lobes of the leaves broadly obovate, 10-20 cm. wide. 
Petioles, leaf nerves beneath, and peduncles loosely 

pilose or hirsutulous. 

Leaves asperous above, the lobes obtuse or acute. 

C. ficifolia. 
Leaves hirsutulous above, the lobes acuminate. 

C. Standleyi. 

Petioles, leaf nerves beneath, and, usually, peduncles 
tomentose . . . . C. obtusa. 



284 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Lobes of the leaves oblongish, scarcely at all obovate, 

less than 10 cm. wide C. Engleriana. 

Leaves glabrous above. 
Leaves pubescent beneath. 

Petioles white- tomentose; pistillate spikes 6-10 cm. long. 

C. Francisci. 

Petioles tomentose with soft, white, and setulose, often 
brown hairs mixed ; pistillate spikes twice as long. 

C. flagellifera. 
Leaves glabrous beneath (the nerves mostly minutely puber- 

ulent). 
Leaves very deeply lobed; staminate spikes few, very 

woolly, elongate C. Setico. 

Leaves moderately (to three-fourths) lobed; staminate 

spikes very many C. Tessmannii. 

Leaf lobes, or some of them, dentate, repandly toothed, or pinnat- 
ifid. 

Leaf divisions dentate-serrate C. dentata. 

Leaf divisions more or less pinnately lobed C. polystachya. 

Gecropia acutifolia Tre'cul, Ann. Sci. Nat. III. 8: 81. 1847. 

A tree with 10-11-lobed leaves, the lobes with rotund sinuses, 
elongate and acutely subacuminate, the largest 5-7 cm. wide and 
about 20 cm. long; pubescence of the branchlets and leaves beneath 
early white- tomentose, later pilose on the former and the nerves; 
petioles white- tomentose; staminate peduncles 5-7 cm. long; bracts 
acuminate, 11 cm. long; spikes 4, on pedicels 5-10 mm. long; pistillate 
spikes 6, nearly 20 cm. long, 8 mm. thick, the fruit verruculose, 
oblong-conic, acute. 

Peru: (Pavdn). 

Cecropia albicans Tre'cul, Ann. Sci. Nat. III. 8: 82. 1847. 

Leaves plane, divided halfway to the base, the lobes obtuse or 
obtusely acuminate, white-hirtellous beneath between the very 
slender, reticulate, rufescent veins; petioles 35 cm. long, white with 
a short, dense pubescence; stipules 16 cm. long and 12 cm. broad; 
pistillate peduncle thick, 1.5-2 cm. long; bract 5 cm. long, abruptly 
attenuate; spikes 3-4. Here seems to belong material at Madrid, 
without locality, by Ruiz and Pavon; the staminate spikes are many. 

Peru: Pawn. 



FLORA OF PERU 285 

Cecropia angustifolia Tre"cul, Ann. Sci. Nat. III. 8: 83. 1847. 
C. digitata Klotzsch, Linnaea 20: 534. 1847. 

A tall, handsome tree with pale bark and spreading crown; leaves 
ample, dark green and scabrous above, deeply lobed, the lobes 
oblong-ligulate, the larger about 5 cm. wide and 3-4 times as long; 
petioles 30-40 cm. long; staminate spikes many, the pistillate 2-4, 
about 4 cm. long; fruit oblong, acute. Neg. 11581. 

Junin: La Merced, 1,200 meters, 5741. Above San Ramon, 1,700 
meters, Killip & Smith 24737; a tree of 9-12 meters. Huanuco: 
Ruiz & Pavon. Puno: Sangaban, Lechler(1}. 

Cecropia bicolor Klotzsch, Linnaea 20: 531. 1847. 

Leaves with elongate-oblong, shortly acuminate lobes, green but 
scabrous above, white-tomentose beneath except for the green, 
hirsutulous nerves; pistillate peduncle long-villous, about 7 cm. 
long, bearing 4 spikes 15 to nearly 20 cm. long. The greatly elongate 
spikes are striking. The species may be allied to C. flagellifera. 
Neg. 11582. 

Huanuco: Mufia, Ruiz. 

Cecropia dentata Klotzsch, Linnaea 20: 533. 1847. 

Very imperfectly known; leaf lobes 9, short, obovate, acute, the 
margins dentate-subserrate, green but hirsute-scabrous above, white- 
tomentose beneath except for the muricate-hirsute midrib and nerves; 
petiole 15-20 cm. long, sparsely pilose. 

Huanuco :Muna, (Ruiz & Pavon). 

Cecropia Engleriana Snethl. Notizbl. Bot. Gart. Berlin 8: 365. 
1923. 

Allied to C. ficifolia but the pubescence of the lower leaf surface 
entirely lanuginose, the stipules, petioles, and leaves often smaller, 
and the latter deeply 7-lobed, the lobes oblongish, 5-7 cm. wide and 
with 22-25 lateral nerves; staminate spikes very slender. Neg. 
11586. 

Rio Acre: Ule 9313. 

Cecropia ficifolia Warb. ex Snethl. Notizbl. Bot. Gart. Berlin 8: 
365. 1923. 

A small to tall (30 meters) tree with ample, peltate leaves, greenish 
but densely asperous-pubescent above and white-tomentose beneath, 
not at all deeply 5-6-lobed, the rotund-obovate lobes (to 45 cm. long 



286 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and 20 cm. wide) obtuse or acute, remote and little narrowed below; 
lateral nerves 12-16; petioles 30-45 cm. long; stipules to 15 cm. long, 
pubescent also within; staminate peduncle about 7 cm. long, the 
pistillate twice as long; staminate spikes 12 or more, 5-7 cm. long, 
2-3 mm. thick, the pedicels 1.5-2 cm. long; enveloping hairs 0.5-1 
mm. long; filaments unequal; pistillate spikes 4-6, sessile, 8 cm. long, 
about twice as long in fruit and 8 mm. thick, lightly tomentose; 
stigmas penicillate; fruit 2-3 mm. long, faintly verruculose. Neg. 
11587. 

Rio Acre: UleOSll; Huber 4245. 

Cecropia flagellifera Tre"cul, Ann. Sci. Nat. III. 8: 81. 1847. 
C. Ruiziana Klotzsch, Linnaea 20: 532. 1847. 

Apparently very similar to C. Francisci, but with shorter petioles, 
smaller leaves, and fewer (4) pistillate spikes, these in fruit 20 cm. 
long and 8-9 mm. thick. 

Peru: (Pavori). 

Cecropia Francisci Snethl. Notizbl. Bot. Gart. Berlin 8: 369. 
1923. 

A tree as much as 25 meters high, with punctate-asperous branch- 
lets, long-pubescent stipules to 20 cm. long, petioles nearly 40 cm. 
long, and coriaceous leaves about 9-lobed to the middle, the 
largest of the broadly obovate lobes 40 cm. long and 20 cm. broad; 
peduncles 10 cm. long or longer; pistillate spikes 5-6, sessile, 7-8.5 
cm. long, 4-5 mm. thick, before flowering densely tomentose; stigma 
penicillate. C. palmata Willd. has leaf nerves reddish beneath, 
finely and minutely pubescent or puberulent rather than tomentose- 
hirtellous. Neg. 11588. 

Rio Acre: Ule 9312. 

Cecropia Klotzschiana Miq. in Mart. Fl. Bras. 4, pt. 1: 151. 
1853. C. scabra Ruiz ex Klotzsch, Linnaea 20: 531. 1847, non Mart. 
1831. 

Apparently too near C. nivea, but the pubescence beneath between 
the veins felt-like, the peduncles sparsely hispid, and the pedicels 
hirtellous. Neg. 11591. 

Peru: Ruiz. 

Cecropia latiloba Miq. in Mart. Fl. Bras. 4, pt. 1: 147. 1853. 

Branchlets glabrate; leaves rather similar to those of C. obtusa 

but with many (about 25) lateral nerves 0.5-1 cm. apart; staminate 



FLORA OF PERU 287 

spikes 25-30, about 8 cm. long, the pedicels 5 mm. long, hirsute; 
pistillate spikes sessile, finally 15 cm. long; fruits 3 mm. long, tomen- 
tose at the tip. Neg. 18838. 

Loreto: Yarina-cocha, Tessmann 3467, 3466 (det. Snethlage). 
Rio Masana, Williams 96. Lower Ucayali, Tessmann 3307 (var. ; det. 
Snethlage). Cumaria, Tessmann 3253 (det. Snethlage). Brazil; 
Bolivia. 

Cecropia leucocoma Miq. in Mart. Fl. Bras. 4, pt. 1: 142. 1853. 

Petioles, peduncles, and young leaves above greenish but lightly 
and loosely lanuginose; leaves thin, deeply (even to the base) divided, 
the elongate-obovate lobes apiculate or rounded, white-tomentose 
beneath or merely araneose; lateral nerves about 1 cm. apart or less, 
usually about 20; staminate spikes fewer than 12, about 7 cm. long, 
the pedicels to 5 mm. long; pistillate spikes about 4, to 15 cm. long, 
5-8 mm. thick, subsessile, the slender peduncle 6-8 mm. long. Tess- 
mann 4058 has been given a subspecific herbarium name by Sneth- 
lage, but it has the close, parallel nerves of C. leucocoma and could be 
treated as a variety of the latter with longer, thicker (20 cm. long, 
9 mm. thick), distinctly pediceled pistillate spikes or, more probably, 
as a distinct species. C. adenopus Mart, and C. scabra might be 
sought here. The former may be known by its hispidulous petioles 
and peduncles; the latter by its reduced (asperous and hirtellous) 
leaf pubescence and its merely ternate staminate spikes. Neg. 11592. 
Illustrated, Wawra, Bot. Ergebn. Reise Bras. pi. 85. 

Loreto: Middle Ucayali, Tessmann 3306, 3455 (one a glabrate 
form, det. Snethlage). Yurimaguas, 200 meters, Williams 4173. 
Bolivia; Brazil. "Setico," "siari chal." 

Cecropia leucophaea Poepp. ex Miq. in Mart. Fl. Bras. 4, pt. 
1:151. pi. 50. 1853. 

Probably allied to C. multiflora, but the more numerous acute 
leaf lobes not more scabrous above than otherwise pubescent; nerves 
beneath green but hirsute- villous; staminate spikes very numerous, 
5-7.5 cm. long, on hirsute pedicels 4 mm. long. 

Huanuco: Cochero, Poeppig. 

Cecropia montana Warb. ex Snethl. Notizbl. Bot. Gart. Berlin 
8: 368. 1923. 

A small tree (to 10 meters) with short, white-pilose branchlets, 
rather small stipules (about 5 cm. long) glabrate within, petioles 35 
cm. long, and deeply 10-lobed leaves white-tomentose beneath, the 



288 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lobes oblong, the largest 30 cm. long and a little less than a third as 
wide; lateral nerves 15-25; peduncle 2.5-4 cm. long; pistillate spikes 
4, becoming 3.5-6 cm. long and 6-10 mm. thick; fruit oblong, 2 mm. 
long, minutely tubercled. Neg. 11599. 

Loreto: Cerro de Escalera, 1,300 meters, Ule 6845. 

Cecropia multiflora Snethl. Notizbl. Bot. Gart. Berlin 8: 367. 
1923. 

A small tree (10 meters, so far as known) with minutely pilose 
branchlets and rather deeply lobed leaves, the largest obovate, acutish 
lobes about 30 cm. long and scarcely half as broad, the pubescence 
above merely asperous, beneath white-tomentose, the midrib and 
12-14 nerves minutely pilose; peduncle 5-8 cm. long, finely pilose; 
staminate spikes as many as 50 and 10-15 cm. long, 2-3 mm. thick, 
on pedicels 1-2 cm. long; perianth 1-1.5 mm. long, minutely pubes- 
cent; filaments nearly equal, the anthers before anthesis ecaudate. 
Similar to C. nivea and C. Klotzschiana, but from herbarium material 
distinguishable by the even upper leaf surfaces. Neg. 11600. 

Junin: Chanchamayo Valley, 800 meters, Weberbauer 1837, type. 

Cecropia nivea Poepp. ex Klotzsch, Linnaea 20: 532. 1847. 

Leaves large, deeply 9-parted, the acute, obovate lobes green but 
scabrous-hispid above, loosely and densely white-tomentose beneath ; 
staminate peduncle 3.5-5 cm. long, densely hispid-villous, bearing 
many slender spikes 7 cm. long, their subulate pedicels 6 mm. long. 
Suggests C. multiflora, but the leaves are noticeably rugose. Neg. 
11601. 

Peru: Poeppig. 

Cecropia obtusa Tre"cul, Ann. Sci. Nat. III. 8: 79. 1847. 

Branchlets strigose; leaf lobes obovate-rotund, very obtuse, 
tomentulose above at first, finally asperous, often 20 cm. long and 
half as broad or larger, the lateral nerves 1-2 cm. apart, about 15; 
staminate peduncle about 20 cm. long, bearing 12-15 shortly pedi- 
celed spikes only 4-5 cm. long; perianth tubular, subentire, minutely 
hirtellous; pistillate spikes 8-9 cm. long, 1 cm. thick, on pedicels 2-4 
mm. long; fruit obovate, verruculose, white-tomentose-punctate. 
C. palmata Willd. has leaves glabrate above and pistillate spikes 13-15 
cm. long. Neg. 25536. 

Loreto: La Victoria, Williams 2557. Caballo-cocha, Williams 
2068. Without locality: (Pavdri). Brazil ; British Guiana. "Setico." 



FLORA OF PERU 289 

Cecropia polystachya Tre"cul, Ann. Sci. Nat. III. 8: 80. 1847. 
C. pinnatiloba Klotzsch, Linnaea 20: 533. 1847. 

Well marked (if the character is constant) by the lobed middle 
leaflets; lobes 9-10, ovate, glabrous above, pubescent beneath, the 
larger 7.5 cm. broad, about 20 cm. long; staminate peduncle tomen- 
tose, 7.5 cm. long, the 20 pediceled spikes 5-10 cm. long; pistillate 
spikes about 4 and 6 cm. long, the peduncle a little longer. Neg. 
25538. 

Huanuco: Mufia, Ruiz. 

Cecropia sciadophylla Mart. Flora 24, pt. 2: Beibl. 93. 1841. 

Immediately recognizable by its petioled or entirely disjoined 
leaflets that are glabrous or merely puberulent-tomentulose in the 
areoles, even beneath. Snethlage has distinguished var. decurrens 
Snethl. and var.Juranyiana (A.Richt.) Snethl.,the latter the extreme 
variation with the leaflets actually sessile. This is a medium to tall 
tree with huge leaves (the largest leaflets about 50 cm. long and nearly 
10 cm. wide or wider), glabrate petioles several dm. long, 4-6 pistil- 
late spikes about 10 cm. long, and several staminate spikes, the 
filaments somewhat unequal. Illustrated, Mart. Fl. Bras. 4, pt. 1: 
pi. 46. 

Loreto: Lower Rio Napo, Tessmann 3718 (det. Snethlage). 
Mouth of Rio Santiago, Tessmann 4717, 4278 (det. Snethlage). 
Caballo-cocha, in forest, Williams 2130. Brazil. "Setico." 

Cecropia Setico Snethl. in herb., sp. nov. 

Folia inter maxima, ad 60 cm. lata et ultra, subtus ad nervos 
fere microscopice pulverulenta; pedunculi leviter spinuloso-ciliati 
vel glabrati ad 10 cm. longi; spicae 4 dense lanatae fere sessiles in 
statu fructifero circa 20 cm. longae. 

Leaves very large, at least 60 cm. wide, almost microscopically 
pulverulent on the nerves beneath; peduncles lightly spinulose- 
ciliate or glabrate, to about 10 cm. long; spikes 4, very woolly, 
becoming about 20 cm. long, nearly sessile. Neg. 11608. 

Loreto: Unanana, Lower Rio Napo, 100 meters, Tessmann 3713. 
"Setico." 

Cecropia Standleyi Macbr. Field Mus. Bot. 11: 61. 1931. 

A small, slender tree of 3-6 meters, similar to C. ficifolia but 
easily distinguishable by the soft, scattered trichomes of the upper 
leaf surfaces and the short-acuminate, broadly obovate leaf lobes; 



290 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pubescence beneath not at all lanuginose; staminate aments 8-10, 
9 cm. long, 3 mm. thick, the pedicels 1 cm. long, the filaments 
equal; pistillate spikes 4, subsessile, 18 cm. long, 1 cm. thick, lightly 
tomentose. 

Loreto: Mishuyacu near Iquitos, 100 meters, Klug 399, type. 
Iquitos, 100 meters, Kittip & Smith 27033. "Setico." 

Cecropia strigosa Tre'cul, Ann. Sci. Nat. III. 8: 82. 1847. 

Similar to C. angustifolia, but the leaf lobes fewer and the pubes- 
cence more hirtellous, in this respect approaching C. tubulosa; stami- 
nate spikes about 60; pistillate spikes about 7. 

Peru: (Pawn). 

Cecropia Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 9: 
260. 1925. 

Leaves large, glabrous, deeply (three-fourths their length) 9- 
lobed; spikes similar to those of C. multiflora but not distinctly 
pediceled; filaments very short. Fruit edible. C. laetevirens Huber, 
Bol. Mus. Goeldi 6: 63. 1910, and C. bifurcata Huber, op. cit. 62, 
would be sought here. The leaves of both are broadly and obtusely 
lobed, more or less fulvous-pilose beneath but green and glabrous 
to the eye. Williams 1*313 (part of a leaf) from Yurimaguas is per- 
haps C. Tessmannii. Neg. 11609. 

Loreto: Yarina-cocha, 150 meters, Tessmann 3454- Fortaleza, 
in forest, 200 meters, Williams 4313. Santa Rosa, 200 meters, 
Williams 4949. "Setico," "pungara." 

Cecropia tubulosa Ruiz ex Klotzsch, Linnaea 20: 534. 1847. 

Probably only a form or possibly a variety of C. angustifolia, but 
the leaves beneath more conspicuously reddish-hirsutulous than 
white-tomentulose. C. angustifolia may be rubescent on the nerves. 
Neg. 11610. 

Huanuco: Chinchao, Mufia, Ruiz & Pavon. 

3. POUROUMA Aubl. 

Trees with entire or 3-5-lobed leaves, or one species with 8-12- 
parted leaves simulating those of Cecropia, but the inflorescence 
always cymose and the rather large, often densely velvety-tomentose 
fruits borne distinctly. See Mart. Fl. Bras. 4, pt. 1: 123-132, for 
a number of Amazonian species to be expected, particularly several 
with entire leaves. Other extra-Peruvian species published more 



FLORA OF PERU 291 

recently, and possibly occurring, are mentioned below. Spruce 
found the name "cocura" used for an Amazonian species. 

Leaves not parted or deeply lobed. 

Pubescence fulvous or brown P. phaeotricha. 

Pubescence white or gray P. folleata. 

Leaves palmately parted or lobed, at least typically and some of 
them. 

Leaves 8 (6)-12-parted P. cecropiaefolia. 

Leaves 3-5-lobed. 
Leaves deeply cordate at base, harshly hairy on both sides. 

P. substrigosa. 

Leaves little if at all cordate, or not pubescent. 
Leaves not distinctly, if at all, cordate. 

Branchlets glabrous or puberulent, the leaves white-hairy 
beneath. 

Petioles glabrous or early granulose P. Tessmannii. 

Petioles tomentose, glabrate only in age ... P. Jussieuana. 
Branchlets yellowish-hairy, like the leaves beneath. 

Leaves strigose or scabrous above P. palmata. 

Leaves glabrous above P. triloba. 

Leaves deeply cordate at the base P. Ulei. 

Pourouma cecropiaefolia Mart, ex Miq. in Mart. Fl. Bras. 4, 
pt. 1: 123. pi. 36. 1853. P. multifida Tre"cul, Ann. Sci. Nat. III. 8: 
107. 1847(7). 

The only Peruvian species with Cecropia-like leaves that are 
cordate-rotund and radiately parted; segments obovate-oblong- 
lanceolate, velvety and ashy- or white-tomentose beneath, the 
largest middle ones sometimes 15-30 cm. long and 10-20 cm. broad, 
all more or less connate toward the base; petioles glabrous; peduncles 
puberulent; inflorescence in flower dense, reddish purple with a close 
puberulence, the fasciculate or capitulate staminate flowers sessile 
or nearly so, their minutely setulose segments entirely free. Accord- 
ing to Martius, a tree of 10-15 meters with the habit of a Cecro- 
pia but with juicy fruits that have a pleasant, sweetish-sour taste. 
Sometimes cultivated. The Martius name (Reise 3: 1130. 1831) is 
essentially a nomen nudum until taken up by Miquel. There is no 
doubt, however, as to its application, and Martius gave the common 
names and described the fruit. Lacking positive proof that P. multi- 



292 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

fida Tre"cul is the same, I retain the well-known name. A specimen 
by Killip and Smith (27932) from Puerto Arturo, Loreto, has been 
determined in Herb. Berlin by Mildbraed as P. sapida Karst. It is 
not clear to me that the latter species is distinct. Its leaves have 9 
lobes. In the Field Museum specimen they are separate nearly to 
the base. 

Loreto: Above Pongo de Manseriche, 200 meters, in forest, 
Mexia 6257. Mishuyacu, 100 meters, in clearing, Klug 1185, 1326. 
Mouth of Rio Blanco, Tessmann 3054- Alto Rio Itaya, Williams 
3347. Iquitos, Ducke 7581; Killip & Smith 27381, 29839 (both det. 
by A. C. Smith). Huanuco: Huamalies District, Weberbauer 3705. 
Rio Acre: Ule 9314. Junin: La Merced, 600 meters, 5446. Brazil. 
"Ambauba mansa," "ambauba do vinho," "mapaty," "cucura," 
"uvilla." 

Pourouma folleata Macbr. Field Mus. Bot. 8: 114. 1930. 

Branchlets glabrous; petioles evanescently appressed-setulose, 
2-3 cm. long; leaves entire or repand-undulate, elliptic, acute at the 
base, very shortly acuminate, the larger 12 cm. long, 6 cm. wide, 
glabrous above or the midnerve strigillose, shortly white-tomentulose 
beneath between the 15 nerves, these densely strigillose; stipules 
yellowish-villous, 2.5 cm. long; peduncles 1.5 cm. long; compound 
cymes grayish-brown-hirsutulous, open, the slender-pediceled stami- 
nate flowers not at all capitulate; segments linear-subulate, acumi- 
nate. P. apiculata Spruce and P. tomentosa Mart, have entire 
leaves arachnoid-tomentose beneath, the former distinguished from 
the latter by the apiculate leaf acumination. Its large fruits are 
velvety. P. crassivenosa Mildbr. Notizbl. Bot. Gart. Berlin 10: 419. 
1928, is glabrous beneath to the eye except on the nerves. P. pa- 
raensis Huber is hirtellous on the veins beneath, appressed-lanate in 
the areoles. 

Junin: La Merced, 1,300-1,700 meters, Schunke 416, A95. 

Pourouma Jussieuana Tre"cul, Ann. Sci. Nat. III. 8: 106. 1847. 

A tree with angled, verruculose, finally glabrous branchlets and 
palmately 3-5-lobed, not at all cordate leaves; leaf lobes glabrous 
above, oblong or the middle ovate, repand, acuminate, the larger 12 
cm. wide and 15-25 cm. long, with 17-22 nerves, these conspicuous 
beneath; petioles pubescent, 5-6 cm. long; pistillate inflorescence 
cymose-paniculate, the puberulent peduncle 5 cm. long; fruit ovate, 
densely pale-rusty-puberulent; seed depressed-ovate. P. cuspidata 



FLORA OF PERU 293 

Warb. of Amazonian Brazil is recognizable by the scabrosity of its 
leaves beneath and its small staminate flowers, less than 1 mm. long, 
the branchlets of the inflorescence thick. 

Peru: Buena Vista (Jussieu). "Amandier de Buena- Vista." 

Pourouma palmata Poepp. & Endl. Nov. Gen. & Sp. 2: 29. 
pi 141. 1838. 

Young branchlets densely yellow-scabrous; leaves rounded or 
subcordate, with 3-5 oblong or ovate, acute lobes, harshly scabrous 
above but silky-pubescent beneath, especially on the prominent 
nerves (the lateral about 30) ; petioles subterete, the upper 8 cm. long, 
shortly pubescent; staminate flowers glomerate, the inflorescence 
much branched; perianth segments shortly united, oblong, hirsutu- 
lous at the apex, equaled by the dense filaments; immature fruiting 
inflorescence velvety in appearance, with a dense, reddish brown 
pubescence; pedicels thick, 3-5 cm. long; young fruits ovate, con- 
spicuously capped by the peltate stigma. Fruits edible. Neg. 
11622. 

San Martin: Tocache, Poeppig 2881, Addenda 153. Huanuco: 
Huamalies, Weberbauer 3639. Loreto: Pampa del Sacramento 
(Huber). "Uvilla." 

Pourouma phaeotricha Mildbr. Notizbl. Bot. Gart. Berlin 
10: 193. 1927. 

Young branches, petioles, and inflorescence rusty puberulent- 
tomentose, with paler, rigid hairs intermixed; leaves very regularly 
oblong-elliptic, obtuse at the base, very acutely acuminate, often to 
20 cm. long and 8-9 cm. broad, the mature ones glabrate above, 
beneath on the midrib and 8-10 lateral nerves more or less hirsutulous 
and rusty puberulent-tomentose, the areoles ashy green and under a 
lens white-araneose; fruiting inflorescence with few short branchlets, 
the pedicels 5-8 mm. long; stigma greenish white, pulvinate, densely 
brown-tomentose and white-hirsute; immature fruit 12 mm. long, 
8-9 mm. thick, bright green. A tree of 7 meters, the trunk 8 cm. in 
diameter, with few branches 2 meters above the base. P. aurea 
Mildbr. op. cit. 10: 418, has obovate-oblong leaves broadest at the 
upper third, with 15-17 lateral nerves. Neg. 11624. 

Loreto: Iquitos, Tessmann 5364- 

Pourouma substrigosa Mildbr. Notizbl. Bot. Gart. Berlin 
10: 192. 1927. 

A tree of about 15 meters with rather harshly strigose-hirsute 
foliage and growing parts; petioles densely and shortly hirsute, to 



294 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

25 cm. long; leaves 3-5-lobed, with narrow, deep sinuses, the seg- 
ments short-acuminate, elliptic, the largest middle one 25-35 cm. 
long and 15 cm. wide; staminate inflorescence 2-many times irregu- 
larly branched, reddish brown with dense pubescence of granular 
and club-shaped hairs intermixed, the flowers sessile and glomerulate, 
their linear, more or less dilated segments ciliate at the apex, barely 
1.5 mm. long, exceeding the filaments. Trunk about 17 cm. in 
diameter, with few branches at 6 meters. Fruit edible. Neg. 11625. 

Loreto: Mouth of Rio Santiago, 160 meters, Tessmann 4642, type. 
Streamlet near mouth of Rio Santiago, 400 meters, Mexia 6201; a 
tree of 14 meters, the leaves and twigs inhabited by small ants. 
"Uvilla." 

Pourouma Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 
10: 192. 1927. 

Glabrous or lightly pubescent except for the more or less deeply 
3-lobed leaves, these silvery beneath with an appressed tomentum; 
petioles sulcate, to 15 cm. long; stipules 8 cm. long, acuminate; 
leaves acuminate, 3-nerved from the base, 20 cm. long; staminate 
inflorescence 7 cm. long, the peduncle as long, the branches and 
branchlets chestnut-brown, with a dense covering of clavate hairs; 
flower glomerules dense, the perianth segments subulate-linear, 
closely white-pubescent, about 1.5 mm. long, the filaments half as 
long. A tree of 20 meters, branching at 12 meters, with a trunk 
25 cm. in diameter. Inflorescence early greenish yellow, in anthesis 
bright brown, becoming dark brown. Neg. 11626. 

Loreto: Mouth of Rio Santiago, 160 meters, Tessmann 4236, type. 
Yurimaguas, 200 meters, in forest, Williams 4688. "Setico." 

Pourouma triloba Tre"cul, Ann. Sci. Nat. III. 8: 104. 1847. P. 
triloba Klotzsch, Linnaea 20: 526. Oct., 1847. 

Perhaps only a less pubescent variety of P. palmata; pistillate 
flowers pubescent; staminate glomerules numerous, with the stamens 
about 5 mm. thick; fruit oblong-obovate,. crustaceous, dehiscent by 
two valves. Neg. 11623. 

Huanuco: Macora, Ruiz, type. Pampayacu, Sawada 21. 

Pourouma Ulei Warb. Bot. Jahrb. 40: 132. 1907. 

A tree, the branches and petioles slightly tomentose or glabrate; 
leaves long-petiolate, deeply cordate at the base, with usually over- 
lapping basal lobes, smooth and glabrous above, closely and minutely 



FLORA OF PERU 295 

white-tomentose beneath, the broad lobes abruptly short-acuminate. 
Neg. 11627. 

Loreto: In pasture, El Recreo, 200 meters, Williams 3984- Ama- 
zonian Brazil. "Uvilla." 

4. COUSSAPOA Aubl. 

Rather similar to Pourouma, but both sorts of flowers capitate, 
the pistillate heads solitary or often few. Leaves entire, the petioles 
often about half as long. Stigma capitate-penicillate. Tall trees 
with the name "chichillica," the trunks thick, the foliage abundant, 
the fibers of the bark serving for the manufacture of coarse baskets, 
sacks for the collection of coca and fruits, and various other articles; 
it is used also to fasten and join timbers and beams (Ruiz & Pa von). 
Species very similar and the characters accepted as definitive may 
be variable. 

Some of the following names by Tre"cul are in conflict with those 
of Klotzsch for the same year (1847), but the former were published 
in August while the latter probably appeared toward the last of the 
year. Linnaea for this year consists of 12 numbers, the correction 
page at the end of vol. 20 being signed as written only in August and 
a much earlier article is signed also as written at this date (Nelly 
Dubugnon, Conservatoire Botanique, Geneva). 

Branchlets, especially apically, very villous or setose. 
Leaves broadly rounded and emarginate at the apex. 

C. emarginata. 
Leaves acute or obtuse at the apex, not emarginate. 

Branchlets long-setose; leaves glabrate or lightly tomentose 

beneath C. hirsuta. 

Branchlets hirsute- villous. 

Leaves densely white-tomentose beneath C. villosa. 

Leaves minutely hirsutulous and cobwebby beneath. 

C. vellerea. 
Branchlets glabrate or shortly pubescent. 

Leaves acute, smooth and glabrous above. 

Branchlets puberulent-hirsutulous; leaves 8-17 cm. long. 
Leaves glabrous beneath or the nerves puberulent. 

C. ovalifolia. 

Leaves araneose-tomentulose beneath C. Tessmannii. 

Branchlets glabrous; leaves 20-30 cm. long C. acutifolia. 



296 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves rounded or retuse apically, often more or less asperous. 
Pistillate heads solitary. 

Leaves scabrous on the upper surface C. asperifolia. 

Leaves smooth on the upper surface C. grandiceps. 

Pistillate heads cymose. 

Leaves finely short-hirsutulous beneath C. magnifolia. 

Leaves white-lanuginose beneath, as well as sparsely hirsutu- 
lous C. Standleyi. 

Coussapoa acutifolia Klotzsch, Linnaea 20: 529. 1847. 

With the large leaves of C. magnifolia Tre'cul, but these obviously 
acute and glabrous except for the puberulent nerves beneath; in 
these respects apparently like C. ovalifolia Tre'cul, with much smaller 
and crowded leaves. Neg. 11555. 

Huanuco: Cochero, Ruiz & Pawn. 

Coussapoa asperifolia Trecul, Ann. Sci. Nat. III. 8: 96. 1847. 

Branchlets soon glabrate or glabrous; leaves often almost square- 
sided, about 15 cm. long or smaller, typically very scabrous above 
and rather softly short-pubescent beneath and finely reticulate; 
petioles 1.5-2 cm. long; staminate heads cymose; perianth oblong, 
with short, glabrous teeth; filaments about as long as the flower; 
pistillate flowers in a solitary head, tubular-urceolate; style very 
short, the stigma capitate, villous; fruits very numerous, oblong, 
exteriorly somewhat fleshy. Both the Peruvian specimens are 
essentially glabrous but are typical in the obtuse or retuse leaves. 
C. Martiana Miq. has broadly ovate, somewhat acute leaves, often 
larger and white-arachnoid beneath, in which respect it resembles 
C. subincana Mart, with the leaves somewhat peltate. Neg. 11556. 

Amazonas: Moyobamba, Weberbauer 4472 (det. Mildbraed). 
Junin: La Merced, 55-47. British Guiana. 

Coussapoa emarginata Killip, sp. nov. 

Arbor 12-15-metralis, ramulis crassis densissime pilis longis fulvis 
mollibus pilosis dense foliatis; stipulae lineari-lanceolatae 6.5 cm. 
longae attenuatae extus dense longipilosae; folia mediocria longi- 
petiolata coriacea, petiolo crasso 4.5-5.5 cm. longo dense piloso; 
lamina late obovato-ovalis vel rotundato-ovalis 13-17.5 cm. longa 
9-12 cm. lata apice late rotundata et leviter vel profunde emarginata, 
basi rotundata, supra in sicco fusca ad costam et interdum ad nervos 
pilosa, aliter glabra, nervis non elevatis, subtus pallida ubique sub- 



FLORA OF PERU 297 

sparse adpresse tomentulosa, ad costam nervosque pilis longis ochra- 
ceis sericeo-pilosa, costa crassa elevata, nervis lateralibus utroque 
latere circa 15 rectis elevatis angulo semirecto obliquis in marginem 
desinentibus, nervulis obscuris numerosissimis rectis arete parallelis; 
iniflorescentia (immatura tantum visa) paucicapitata densa, capitulis 
parvis dense paucifloris breviter pedunculatis, pedunculo petiolo 
aequilongo. 

Loreto: Mishuyacu, 100 meters, in forest, Killip & Smith 29955 
(U. S. Nat. Herb., type; duplicate in Herb. Field Mus.). 

Coussapoa grandiceps Killip, sp. nov. 

Arbor, ramulis crassis sparse in statu juvenili hirtello-puberulis 
cito glabratis; stipulae magnae circa 7 cm. longae pallido-tomentosae; 
folia magna coriacea, petiolo crasso 6-10 cm. longo striato glabrato; 
lamina ovali-ovata 24-32 cm. longa 16-19 cm. lata versus apicem 
rotundatum paullo angustata, basi late rotundata brevissime cor- 
data, supra glabra laevis, subtus minute arete pallido-tomentella, 
nervis lateralibus utroque latere circa 18 rectis elevatis, venulis 
ultimis creberrimis prominentibus arete parallelis; capitula feminea 
geminata, pedunculis simplicibus crassis 5-6.5 cm. longis leviter 
tomentulosis glabrescentibus, capitulis densissimis subglobosis multi- 
floris 1.5 cm. diam. 

Loreto: Yanon, wooded banks of lower Rio Huallaga, 135 meters, 
W. J. Dennis (Killip & Smith 29246; type in U. S. Nat. Herb.); 
a tree of 12-15 meters. Puerto Arturo, Yurimaguas, 200 meters, in 
forest, Williams 5349. Apparently conspecific is Williams 4179 
from Yurimaguas. "Renaco caspi." 

Coussapoa hirsuta Tre"cul, Ann. Sci. Nat. III. 8: 97. 1847. 
C. setosa Klotzsch, Linnaea 20: 528. 1847. 

Branchlets conspicuously long-setose toward the tips; petioles 
to 3 cm. long, evanescently pilose; leaves obovate, rounded at the 
base, acute, about 15 cm. long, 10 cm. wide or narrower, slightly 
floccose on the nerves beneath; pistillate heads cymose-corymbose, 
the urceolate flowers thin and including the subdrupaceous fruit; 
seed irregular. Mildbraed (in herb. Madrid) maintains the Klotzsch 
name. Neg. 11560. 

Huanuco: Macora, Ruiz & Pawn. 

Coussapoa magnifolia Trecul, Ann. Sci. Nat. III. 8: 98. 1847. 
C. Ruizii Klotzsch, Linnaea 20: 529. 1847. 

Suggesting C. asperifolia Tre"cul, but the leaves often 30 cm. 
long and nearly as wide; lateral nerves 6-7; petioles glabrate, to 10 



298 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cm. long; pistillate heads several; ovary subobovate. Mildbraed 
employs the Klotzsch name. Neg. 11557. 

Huanuco: Cochero, Poeppig 1371; Ruiz & Pavon, type. 
Junin: La Merced, 600 meters, 5447. 

Coussapoa ovalifolia Tre"cul, Ann. Sci. Nat. III. 8: 95. 1847. 
C. puberula Klotzsch, Linnaea 20: 529. 1847. 

Petioles 4-6 cm. long; leaves ovate or ovate-oblong, rounded 
at the base, mostly 8-12, rarely 15, cm. long, half as wide; lateral 
nerves 13-15, the veins obvious; pistillate heads umbellate-cymose, 
the flowers puberulent only at the tip; stigma penicillate; ovary 
oblong. Smoothish forms of C. asperifolia may be distinguished by 
their obtuse leaves and solitary pistillate heads. C. trinervia Mildbr. 
is glabrous, with strongly 3-nerved leaves. C. nitida Miq. is sparsely 
hirtellous and the pistillate flowers glabrous. Neg. 11559. 

Junin: Rio Paucartambo Valley, 700 meters, Killip & Smith 
25341; a tree of 9-12 meters. La Merced, 600 meters, 5594. 
Huanuco: Chinchao, Pillao, Posuso, etc., /fou'z. Loreto: Puerto 
Metendez, Tessmann 3922 (det. Mildbraed). Rio Acre: Vie 9316? 
(det. Mildbraed). Without locality: Ruiz. 

Coussapoa Standleyi Macbr., sp. nov. 

C. magnifoliae affinis; petiolis (12 cm. longis), nervis venisque 
minutissime parceque hispidulis; foliis apice rotundato-obtusis ad 
40 cm. longis supra glabris subtus dense arachnoideis, margine valde 
undulato-repandis; nervis lateralibus circa 16 prominentibus; pedun- 
culis dense ferrugineo-tomentuloso-hirsutulis; capitulis fern, circa 
15 plerumque geminato-connatis, dense hirsutulis. Seemingly well 
distinct in character of pubescence, and the numerous pistillate heads 
(these are undeveloped). Mildbraed referred the specimen to C. 
magnifolia with a query. The pubescence suggests that of the 
different C. vellerea. Neg. 29511. 

Huanuco: Huamalies, 700-800 meters, Weberbauer 3702, type 
(Hb. Berlin). 

Coussapoa Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 
10:413. 1928. 

Youngest branchlets rusty-hirsutulous; leaves ovate, nearly 
truncate at the base, triangular-acute at the apex, 13-17 cm. long, 
9-12 cm. wide, on stout petioles a third as long; lateral nerves about 
11, the veins and veinlets beneath densely reticulate and prominent, 
the areoles thereby sunken; pistillate inflorescence many-headed, 



FLORA OF PERU 299 

on peduncles 1.5-2 cm. long, the short, thick branchlets somewhat 
hirsute and granular; heads 5 to nearly 10 mm. thick, sometimes 
geminate-connate; flowers 3 mm. long, glabrous, the shorter bracts 
filiform or slightly widened at the apex. Differs from C. puberula 
in the pubescence. C. intermedia Mart, has very fine, scarcely at all 
reticulate or obvious veins. Neg. 11562. 

Loreto: Mouth of Rio Santiago, Tessmann 4673, type. "Renaco." 

Coussapoa vellerea Klotzsch, Linnaea 20: 527. 1847. 

Similar to C. villosa, but the leaf pubescence not at all tomentose, 
the arachnoid hairs evanescent; leaves to 40 cm. long, 30 cm. wide; 
peduncles 5 cm. long, to 11 cm. in fruit, geminate, the fruiting peri- 
anth reddish-brown-hirsute, 3 mm. long. 

Huanuco: Macora, Ruiz & Pavon, type. 

Coussapoa villosa Poepp. & Endl. Nov. Gen. & Sp. 2: 33. pi. 147. 
1838. 

Branches, petioles, bracts, and peduncles villous-hirsute, often 
even shaggy; leaves ample, sometimes 30 cm. long and a third as 
broad or larger, broadly ovate and more or less cordate, acute or 
obtuse; staminate heads few to rather many, corymbose-paniculate; 
bractlets spatulate, often emarginate; perianth turbinate, trifid, 
pubescent; pistillate heads solitary or geminate; perianth contracted 
below the apex, oblong; style slender; fruit ovate-oblong. C. Sprucei 
Mildbr. has oval-oblong leaves, subvelvety-pubescent beneath. Neg. 
11564. 

Huanuco: Casapi, Poeppig, type. Loreto: Mouth of Rio 
Santiago, Tessmann 4696 (det. Mildbraed). Santa Rosa, Lower Rio 
Huallaga, 135 meters, Kittip & Smith 28800 (det. A. C. Smith). 
Ecuador. 

5. FICUS L. Fig 

By Paul C. Standley 

Small or large trees with milky sap; leaves entire in the native 
American species; easy of recognition among all American trees 
by the form of the fruit, similar to that of the cultivated fig, but in the 
wild trees much smaller, consisting of a hollow, more or less fleshy 
receptacle, with a small apical opening (ostiole), this closed by 
small, overlapping scales; flowers minute, densely inserted all over 
the interior wall of the receptacle. A large genus in tropical America, 
doubtless with more Peruvian representatives than are listed here. 
A few other Peruvian species, in fact, are represented in the herba- 



300 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rium by fragmentary material, insufficient for their diagnosis. The 
plants often or usually begin life as epiphytes, the seeds germinating 
upon a branch of some host tree. The seedlings send down aerial 
roots that develop into long, cord-like stems which finally envelop 
and strangle the host. The wood is soft and of no economic 
importance. The larger fruits are fleshy, sweet, and edible (espe- 
cially in the subgenus Pharmacosycea} . They are seldom eaten by 
people but are much sought by birds and other animals. The latex 
yields a kind of rubber. From the bark the ancient Mexicans pre- 
pared paper upon which some of their manuscripts were written. 

Receptacles solitary in the leaf axils; leaves often scabrous; involucre 
3-lobate. Subgenus Pharmacosycea. 

Leaf blades deeply emarginate at the base F. Macbridei. 

Leaf blades not emarginate at the base. 

Leaves gradually acute or acuminate, or cuspidate-acuminate. 

F. glabrata. 

Leaves obtuse or acutish, sometimes abruptly short-acute. 
Leaves with very numerous, close lateral nerves. 

F. anthelmintica. 

Leaves with few distant lateral nerves F. radula. 

Receptacles geminate; leaves never scabrous; involucre 2-lobate. 

Subgenus Urostigma. 

Branches hirsute with long, spreading, brown hairs. Fruit sessile. 
Fruits large, about 12 mm. in diameter, densely hirsute. 

F. juruensis. 

Fruits about 6 mm. in diameter, glabrate F. Llewelyni. 

Branches not hirsute, usually glabrous or nearly so. 
Receptacles small, mostly 4-6 mm. in diameter. 

Leaves large, about 35 cm. long, attenuate to the base, nar- 
rowly oblanceolate-oblong F. caballina. 

Leaves much smaller, chiefly 5-14 cm. long. 

Young branches and petioles glabrous. Fruit glabrous. 
Leaves 3-nerved at the base, commonly about 6 cm. long. 

F. nitida. 

Leaves not 3-nerved at the base, mostly 7-12 cm. long. 

F. Matthewsii. 

Young branches and petioles, at least at first, strigose or 
appressed-pilose. 



FLORA OF PERU 301 

Receptacles strigose at first, becoming glabrate. 

F. Killipii. 

Receptacles glabrous from the first. 
Nerves of the leaves very oblique, salient on the lower 

surface F. Weberbaueri. 

Nerves divergent at an angle of more than 60 degrees, 
not elevated on the lower surface. .F. casapiensis. 
Receptacles large, 8-20 mm. in diameter or even larger. 
Fruits sessile. 
Leaves rounded at the apex; involucre about half as long 

as the fruit F. Urbaniana. 

Leaves abruptly cuspidate-acuminate; involucre small and 

inconspicuous F. paraensis. 

Fruits pedunculate. 

Leaves narrowed to the acute or obtuse base. . .F. gemina. 
Leaves broad toward the truncate or shallowly cordate base. 

F. Ruiziana. 

Ficus anthelmintica Mart. Syst. Mat. Med. Bras. 88. 1843. 
Pharmacosycea anthelmintica Miq. Lond. Journ. Bot. 7: 66. 1848. 

A large tree, glabrous throughout; stipules narrow, long-attenuate, 
as much as 11 cm. long; leaves long-petiolate, the blades elliptic- 
oblong or elliptic, 13-23 cm. long, 5.5-10 cm. wide, acute or acutish, 
subacute to rounded at the base, the lateral nerves close together, 
prominent beneath, slender, 15-25 or more on each side, diverging 
at a wide angle; receptacles pedunculate or almost sessile, globose, 
commonly 2-2.5 cm. in diameter. Illustrated, Mart. Fl. Bras. 4, 
pt. 1 : pi. 25, f. 2. 

Junin: La Merced, 600 meters, 5295; a large tree with widely 
spreading, dense crown. Loreto: Iquitos, in 1932, W. G. Scherer: 
at 100 meters, Killip & Smith. Pebas, in forest, Williams 1867. 
Widely distributed in Brazil. "Oje," "huito." 

The sap (leche de oje) and bark (corteza de oje") are used medicin- 
ally in the region of Iquitos, particularly as a local remedy for 
tertian fevers. 

Ficus caballina Stand!., sp. nov. 

Omnino glabra, ramulis crassis ut videtur dense foliatis ochraceis; 
stipulae non visae; folia inter maxima subcoriacea breviter petiolata, 
petiolo crasso 1.5 cm. tantum longo; lamina anguste oblanceolato- 



302 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

oblonga circa 36 cm. longa et 11 cm. lata obtusa basin versus longe 
sensim attenuata, basi ipsa attenuato-cuneata, supra in sicco brun- 
nescens, costa nervisque prominentibus, costa gracili subtus elevata, 
nervis lateralibus utroque latere circa 15 gracilibus prominentibus 
angulo lato divergentibus fere rectis versus marginem inaequaliter 
conjunctis; receptacula numerosa ut videtur ex axillis defoliatis 
nascentia globosa 5-6 mm. diam. glabra, ostiolo plus minusve 
elevato, involucre bilobo, lobis brevibus late rotundatis vix 2.5 mm. 
longis; pedunculi graciles 6-8 mm. longi. 

Loreto: Caballo-cocha, in forest, Williams 2075 (type in Herb. 
Field Mus. No. 608,842). "Renaquillo." 

A species easily recognized by the very large, long and narrow 
leaves and remarkably small receptacles. 

Ficus Carica L. Sp. PI. 1059. 1753. 

The common fig ("higo"), native of the Mediterranean region, 
is cultivated frequently in various parts of Peru. It differs from all 
American species in its deeply lobate leaves. 

Ficus casapiensis Miq. Ann. Mus. Bot. Lugd. Bat. 3: 298. 
1867. Urostigma casapiense Miq. Lond. Journ. Bot. 6: 548. 1847. 

Branchlets thick, sparsely pilose or glabrate; stipules 2 cm. long 
or less, acuminate, dorsally sericeous-pilose with long, pale hairs; 
leaves petiolate, coriaceous, the blades oblong to oblong-obovate, 
very abruptly short-acuminate, very obtuse to subacute at the base, 
sparsely pilose beneath along the costa, elsewhere glabrous, the 
lateral nerves about 17 pairs, divergent at a wide angle, very slender; 
receptacles sessile, glabrous, globose, "slightly larger than a pea. 
The specimens from Loreto are referred here with some doubt. 

Huanuco: Casapi, Mathews, type. Loreto: Santa Rosa, 135 
meters, in forest, Killip & Smith 27846. Puerto Arturo, 135 meters, 
in forest, Killip & Smith 27846. 

Ficus gemina Ruiz ex Miq. in Mart. Fl. Bras. 4, pt. 1: 98, in 
syn. 1853. Urostigma geminum Miq. loc. cit. 

A tree of 5-8 meters or more, glabrous throughout, the branches 
relatively slender; stipules small and narrow, about 1 cm. long; 
leaves slender-petiolate, subcoriaceous, the blades ovate-elliptic to 
oblong-elliptic, commonly 6-12 cm. long and 2-5 cm. wide, acuminate 
or abruptly acute, obtuse or subacute at the base and rather con- 
spicuously triplinerved, the lateral nerves 8-15 on each side, very 
slender, divergent at a rather wide angle; receptacles short-peduncu- 



FLORA OF PERU 303 

late, globose, generally 7-8 mm. in diameter, often spotted with red, 
the ostiole plane or conspicuously elevated. Neg. 11713. Illustrated, 
Mart. Fl. Bras. 4, pt. 1: pi. 27, f. 3. 

Huanuco: Pampayacu, Sawada 13, 17. Posuso, Ruiz, probably 
type material. Junin: La Merced, 600 meters, along stream, 5484, 
a large tree with spreading crown, the bark white; Killip & Smith 
23463, fruits yellow to orange with darker spots. Loreto: Mishu- 
yacu, 100 meters, Killip & Smith 29899. Moyobamba, Weberbauer 
4523. San Martin: Tarapoto, Ule 6510; Williams 5502; Spruce 
4274. Chazuta, 260 meters, in forest, Klug 3972. Also in Colombia, 
and reported from Brazil and Surinam. "Renaco," "higo silvestre." 

Ficus glabrata HBK. Nov. Gen. & Sp. 2: 47. 1817. 

A large tree, glabrous throughout; stipules narrow, usually not 
more than 2 cm. long, often much smaller; leaves petiolate, the 
blades elliptic-oblong to elliptic-oval, 12-25 cm. long, 5-10 cm. wide, 
abruptly acute or acuminate, sometimes caudate-acuminate, acute 
to obtuse at the base and 3-5-nerved, pale green when dried, the 
lateral nerves remote, usually 8-15 on each side, rather stout, 
divergent at a wide angle; receptacles pedunculate, globose, 1.5-4 
cm. in diameter, glabrous or obscurely scaberulous, the ostiole 
prominent, the involucre very small. 

Junin: Chanchamayo, Isern 2329. Loreto: Lower Rio Huallaga, 
135 meters, Killip & Smith 29009, fruit rich purple. San Antonio, 
145 meters, Williams 3548; Killip & Smith 29458, fruit deep and 
pale green mottled. Iquitos, 120 meters, edge of forest, Williams 
3747. Rio Masana, Williams 100. Balsapuerto, 150-350 meters, 
Killip & Smith 28381. San Martin: Tarapoto, Williams 6311, 
5463. San Roque, Williams 7628. Extending to Brazil and north- 
ward through Central America. "Oje"." Called "higueron" in Ecuador. 

Ficus juruensis Warb. Bot. Jahrb. 40: 140. 1907, nomen. 

A small or medium-sized tree, the branchlets densely brownish- 
hirsute with spreading hairs; stipules brown-hirsute, often long- 
persistent; leaves short-petiolate, the blades obovate-oblong to 
elliptic or almost oval, mostly 7-15 cm. long and up to 6.5 cm. wide, 
acuminate or sometimes obtuse and abruptly short-pointed, coria- 
ceous, rounded or obtuse at the base, scabrous and somewhat pilose 
on the upper surface or finally glabrate, beneath densely short-pilose 
with soft, brownish hairs, the lateral nerves about 10 pairs, very 
prominent beneath, almost straight, ascending at a rather wide 
angle; receptacles sessile, globose, about 12 mm. in diameter, densely 



304 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

brown-hirsute. Negs. 11706, 11710 (determined by Warburg as a 
new species; Ule 5271). 

Junin: Huacapistana, 1,800 meters, 5819; Weberbauer 2321. 
Loreto: Lower Rio Huallaga, 200 meters, Williams 4702. Without 
locality, Ule 5271. Amazonian Brazil. 

Ficus Killipii Standl., sp. nov. 

Arbor 6-30-metralis, ramulis crassiusculis griseis vel fuscis 
rimosis primo sparse strigosis cito glabratis dense foliatis; stipulae 
circa 1 cm. longae anguste triangulares longiattenuatae puberulae vel 
glabratae; folia petiolata coriacea, petiolo crassiusculo 1-1.5 cm. longo 
glabrato; lamina oblonga vel obovato-oblonga 6-12.5 cm. longa 2.5-5 
cm. lata obtusa vel breviter abrupte obtuso-acuminata, basi obtusa 
vel anguste rotundata atque 5-nervia, costa utrinque prominente, 
nervis lateralibus utrinque prominentibus utroque latere 9-11 
tenuibus angulo latiusculo divergentibus prope marginem arcuato- 
conjunctis fere rectis; receptacula geminata sessilia parva globosa 
5 mm. diam. primo sparse strigosa cito glabrata, ostiolo paullo 
prominente, involucre brevi, lobis late rotundatis. 

Junin : Rio Paucartambo Valley, near Perene" Bridge, 700 meters, 
in forest, Kittip & Smith 25252 (Herb. Field Mus. No. 616,750, 
type). Ayacucho: Choimacota Valley, 700 meters, Weberbauer 7538. 

Weberbauer reports the local name of the latex as "aceite Maria," 
and states that it is taken by women as a remedy for sterility. 

Ficus Llewelyni Standl., sp. nov. 

Arbor, ramulis crassis ochraceis vel brunnescentibus primo 
pilis longis brunneis dense paten ti-hirsutis; stipulae circa 1 cm. 
longae dense brunneo-hirsutae; folia mediocria petiolata subcoriacea, 
petiolo crasso vel subgracili 8-15 mm. longo hirsuto vel glabrato; 
lamina obovato-elliptica vel oblongo-elliptica 7.5-12 cm. longa 4-6.5 
cm. lata apice obtusa vel rotundata et abrupte breviter acutata, basi 
obtusa vel anguste rotundata, supra in sicco fuscescens glabra, 
costa nervisque impressis, subtus praesertim ad nervos brunneo- 
pilosa vel subhirsuta, costa gracili elevata, nervis lateralibus utroque 
latere 8-12 gracilibus prominentibus fere rectis obliquis marginem 
fere attingentibus; receptacula geminata sessilia globosa circa 6 mm. 
diam. primo strigosa cito glabrata, ostiolo prominente, involucro 
parvo bilobo extus strigoso. 

Loreto: Caballo-cocha on the Amazon, in forest, Williams 2308 
(Herb. Field Mus. No. 608,283, type). San Martin: Chazuta, 260 



FLORA OF PERU 305 

meters, in forest, epiphytic, Klug 4049. Huanuco: Vitoc, Ruiz 
& Pawn. Junin : Chanchamayo, Isern 2231 . 

Ruiz and Pavon assigned to the plant an unpublished specific 
name now untenable. 

Ficus Macbridei Standl., sp. nov. 

Arbor 10-metralis, ramulis crassissimis primo albido-pilosis cito 
glabratis, internodiis brevibus vel elongatis; stipulae angustae attenu- 
atae circa 5.5 cm. longae glabrae; folia inter maxima petiolata sub- 
coriacea, petiolo crasso circa 7 cm. longo glabrato; lamina obovato- 
elliptica circa 30 cm. longa et 17 cm. lata apice obtusa et abrupte 
breviter acutata, basin versus paullo angustata, basi ipsa anguste 
rotundata et profunde (6 mm.) emarginata, supra glabra costa 
venisque vix elevatis, subtus praesertim ad nervos scaberula vel ad 
costam breviter hirtella, costa crassa elevata, nervis lateralibus 
utroque latere circa 12 crassiusculis remotis angulo lato divergentibus 
prope marginem arcuato-conjunctis; receptacula solitaria sessilia (?) 
globosa 2 cm. diam. dense pilis albidis hirtello, involucre parvo 
breviter trilobo. 

Huanuco: Pampayacu, mouth of Rio Chinchao, 1,050 meters, 
Macbride 5090 (Herb. Field Mus. No. 536,141, type). 

Ficus Matthewsii Miq. Ann. Mus. Bot. Lugd. Bat. 3: 298. 
1867. Urostigma Matthewsii Miq. Lond. Journ. Bot. 6: 549. 1847. 
F. oblanceolata Rusby, Bull. N. Y. Bot. Gard. 4: 446. 1907. 

A small or medium-sized tree, glabrous throughout; petioles 
short or elongate, the blades obovate-oblong to narrowly oblan- 
ceolate-oblong, mostly 5.5-12 cm. long and 2.5-4 cm. wide, acute or 
obtuse, the extreme tip obtuse, acute or obtuse at the base, coria- 
ceous, the costa very prominent beneath, the lateral nerves fine, very 
numerous, closely parallel, divergent often at almost a right angle; 
receptacles numerous, deep red or pinkish or yellowish dotted with 
red, short-pedunculate, globose, 5-6 mm. in diameter, the involucre 
minute. 

Type collected in Peru by Mathews. Huanuco: Huacachi, 1,950 
meters, the trunk attached to a rock, 3866. Junin: La Merced, 
Weberbauer 1881. Loreto: Iquitos, 120 meters, Williams 8012, 3737, 
8079, 3694, 3784; Killip & Smith 27077. Timbuchi, Rio Nanay, 
Williams 927. Mishuyacu, 100 meters, in forest, Killip & Smith 
29897. Bolivia to Venezuela. "Renaco." 



306 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ficus nitida Thunb. Ficus Diss. 10. 1786. 

Glabrous throughout; a small or often a very large tree with 
broad crown and numerous aerial roots; stipules narrow, attenuate, 
about 1 cm. long; leaves small, petiolate, coriaceous, the blades 
obovate, acute at the base, the apex acute or acutish but the tip 
obtuse, the lateral nerves fine and inconspicuous; receptacles sessile, 
globose, about 6 mm. in diameter. 

An Asiatic species, said to be the most frequent street tree of 
Lima. It is planted commonly for shade and ornament in many 
parts of tropical America. 

Ficus paraensis Miq. Ann. Mus. Bot. Lugd. Bat. 3: 298. 1867. 
Urostigma paraense Miq. Lond. Journ. Bot. 6: 534. 1847. 

A small or medium-sized tree, glabrous throughout; stipules 3 cm. 
long or less, narrow, attenuate, caducous; leaves medium-sized, sub- 
coriaceous, petiolate, the blades oblong to oval-oblong, mostly 10-18 
cm. long and 6-8 cm. wide, abruptly acuminate or sometimes cau- 
date-acuminate, subacute to broadly rounded and often emarginate 
at the base, the lateral nerves numerous, divergent at a wide angle; 
receptacles sessile or nearly so, globose, about 1.5 cm. in diameter, 
the ostiole elevated, the involucre small and inconspicuous. Illus- 
trated, Mart. Fl. Bras. 4, pt. 1: pi. 28, f. 1. Neg. 18826. 

Huanuco: Pampayacu, Sawada 4- Junin: La Merced, 600 meters, 
river bank, 5571; Killip & Smith 23711, the fruit green striped with 
purple. Pichis Trail, 500 meters, Killip & Smith 26213. Rio Perene, 
600 meters, Killip & Smith 25237. Loreto: Yurimaguas, 200 meters, 
edge of forest, Williams 3803. Puerto Arturo, 200 meters, Williams 
503 If. Caballo-cocha, Williams 2171. Manfinfa, Williams 1141. 
Iquitos, 100 meters, in forest, Killip & Smith 27484- Amazonian 
Brazil; reported from British Guiana. "Renaco," "sacha-oje." 

Ficus radula Willd. Sp. PI. 4: 1144. 1806. Pharmacosycea 
radula Liebm. Dansk. Vid. Selsk. Skrift. V. 2: 330. 1851. 

A large or medium-sized tree, glabrous or the branches and leaves 
sometimes minutely puberulent; stipules narrow, attenuate, mostly 
1-1.5 cm. long, caducous; leaves petiolate, coriaceous, the blades 
oblong to obovate or oval, 8-16 cm. long, 4-8 cm. wide, rounded or 
obtuse at the apex and abruptly short-pointed, obtuse or acutish at 
the 5-nerved base, often somewhat emarginate, the lateral nerves 
prominent beneath, 7-12 on each side, divergent at a wide angle; 
receptacles short-pedunculate, globose, 1.5-3 cm. in diameter, sca- 
brous, the involucre very small. 



FLORA OF PERU 307 

Amazonas: Chachapoyas, Weberbauer 4309. Junin: La Merced, 
700 meters, Kittip & Smith 23534. Rio Perene", 600 meters, in forest, 
Killip & Smith 25166. Loreto: Florida, 200 meters, in forest, Klug 
2099. Without locality: Ruiz & Pavon. Ranging northward to 
southern Mexico. "Zauchama caspi" (Klug). 

Klug reports that the Indians prepare from the bark a kind of 
cloth that they use for their clothing. In various parts of the earth 
numerous Moraceae are or have been employed in much the same 
manner. 

Ficus Ruiziana Standl., sp. nov. 

Arbor alta fere omnino glabra, ramulis crassiusculis cinnamomeis 
vel ochraceis; stipulae anguste triangulares attenuatae caducae 
extus puberulae; folia mediocria vel majuscula subcoriacea longi- 
petiolata, petiolo gracili glabro 3.5-10 cm. longo; lamina oblongo- 
ovalis vel ovali-elliptica 9-25 cm. longa 5.5-12 cm. lata apice rotun- 
data vel obtusa et abrupte in acumen angustum acuminatum brevem 
contracta, basi late rotundata vel breviter cordata, glabra, in sicco 
viridescens vel brunnescens, costa nervisque supra prominentibus, 
costa subtus elevata gracili, nervis lateralibus utroque latere 8-16 
remotis gracilibus prominentibus fere rectis angulo lato divergentibus 
prope marginem arcuato-conjunctis; receptacula globosa 12-15 mm. 
diam. pedunculata glabra geminata, ostiolo prominente, involucro 
bilobo, lobis patentibus late rotundatis vix ultra 2.5 mm. longis; 
pedunculi crassi 3-5 mm. longi. 

Junin: La Merced, 1,200 meters, Macbride 5795 (Herb. Field 
Mus. No. 536,824, type). Huanuco: Cochero and Posuso, Ruiz (a 
tracing of a leaf in Herb. Berlin, apparently referable here). Loreto: 
Santa Rosa, 200 meters, in forest, Williams 4877. Iquitos, 120 
meters, Williams 7998. Caballo-cocha, in forest, Williams 2114. 
Cuzco: Santa Rosa, Urubamba Valley, 1,200 meters, Cook & Gilbert 
1722. Probably also in Amazonian Brazil. "Renaco." 

Ficus Urbaniana Warb. in Urban, Symb. Ant. 3: 459. 1903. 

An almost glabrous tree, the branchlets very thick; stipules 
broadly lanceolate, glabrous; leaves large, petiolate, the blades 
rounded-obovate to obovate or oblong-obovate, 14-22 cm. long, 7-14 
cm. wide, broadly rounded at the apex, obtuse or rounded at the base, 
the lateral nerves 7-9 pairs, remote, ascending at a wide angle; 
receptacles sessile or nearly so, globose, 1.5-2.5 cm. in diameter or 
even larger, densely velutinous-pubescent, the large ostiole promi- 



308 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nent; involucre at first enclosing the young fruit, in age sometimes 
2 cm. long, sericeous-puberulent. 

Libertad: Below Hacienda Membrillal, 1,400 meters, Weberbaucr 
6984- Ecuador, Venezuela, and the West Indies. 

Ficus Weberbaueri Standl., sp. nov. 

Ramuli crassi ochracei sparse strigosi densiuscule foliati, inter- 
nodiis brevibus; stipulae triangulares attenuato-acuminatae 5-6 mm. 
longae extus dorso strigosae; folia inter minora coriacea petiolata, 
petiolo 10-17 mm. longo gracili sparse strigoso; lamina elliptico- 
oblonga 6-10 cm. longa 2.5-4.5 cm. lata acuta vel subacuta, basi 
paullo angustata obtusa, glabra vel glabrata, in sicco brunnescens, 
costa supra elevata, nervis non elevatis, costa subtus elevata gracili, 
nervis lateralibus utroque latere 8-10 arete elevatis fere rectis valde 
obliquis angulo fere semirecto adscendentibus juxta marginem 
arcuato-conjunctis; receptacula sessilia geminata parva globosa 
glabra 5 mm. tan turn diam., ostiolo depresso, involucro bilobo extus 
sparse strigoso, lobis late rotundatis vix 2 mm. longis. 

Cuzco: Casnipata, Prov. Paucartambo, 700-800 meters, Weber- 
bauer 6954 (Herb. Field Mus. No. 628,234, type). Loreto: Iquitos, 
125 meters, Williams 8138. "Renaco." 

6. TROPHIS L. 

Similar to Chlorophora but the pistillate flowers borne in short, 
dense racemes, tubular and 4-dentate. Fruit globose, scantily fleshy, 
edible. 

Trophis racemosa (L.) Urban, Symb. Antill. 4: 195. 1905. 
Bucephalon racemosum L. Sp. PI. 1190. 1753. 

Leaves oblong-elliptic, 8-10 cm. long, entire; staminate aments 
5-10 cm. long, the pistillate about 1 cm. long. The Peruvian tree, 
sometimes 10-15 meters high, may have serrulate leaves, var. meri- 
dionalis (Bur.) Macbr., comb. nov. (T. americana L. var. meri- 
dionalis Bur. in DC. Prodr. 17: 253. 1873). The leaves and branch- 
lets of the species are said to serve as a nutritious cattle food 
(Standley). Illustrated, Fawc. & Rendle, Fl. Jam. 3, pt. 1: 41. 

San Martin: Juanjui, 400 meters, in forest, Klug 3794. Tarapoto, 
Spruce 4521 (the var.); Williams 6573, 6542. Loreto: Mouth of 
Rio Santiago, Tessmann 4083. Yurimaguas, 135 meters, in forest, 
Killip & Smith 27685. Puerto Arturo, 200 meters, in forest, Williams 
5243 (narrow-leaved form). Florida, 200 meters, Klug 2079. Near 
mouth of Rio Tigre, 115 meters, Killip & Smith 27531. Caballo- 



FLORA OF PERU 309 

cocha, in forest, Williams 2086. Rio Masana, Williams 201. Sole- 
dad, 110 meters, Killip & Smith 29576. Huallaga, Yurimaguas, 200 
meters, Williams 4658. Above Pongo de Manseriche, 210 meters, 
Mexia 6303, 6262. Ranging to Mexico and the West Indies. "Cu- 
chara-caspi, ' ' ' 'uspai-manchinga. ' ' 

7. MORUS L. Mulberry 

Reference: Bureau in DC. Prodr. 17: 237-249. 1873. 
Flowers green, monoecious, each kind in separate aments. Seeds 
included in the succulent fruiting perianths which, borne densely 
in spikes, form the well known mulberry or "mora," M. nigra L. 
or M. alba L., both of which, Asiatic in origin, are sometimes culti- 
vated in Peru. In addition, according to Bureau, there are the two 
following American species. Perhaps Herrera's reference in Contr. 
Fl. Dep. Cuzco, ed. 2. 65. 1921, to M. nigra as "growing wild in 
abundance in the Province of Convencion, from Huiro to Echarati," 
should apply rather to one of the following. 

Morus celtidifolia HBK. Nov. Gen. & Sp. 2: 33. 1817; 246. 

Branches and branchlets slender, glabrous or somewhat pubes- 
cent; leaves ovate, rounded or cordate at the base, acuminate, sca- 
brous above, hirtellous on the nerves beneath; stipules 1-10 mm. 
long; spikes laxly flowered, both the staminate and pistillate peduncu- 
late, the peduncle at least 5 mm. long. A tree to 9 meters tall, the 
wood valued in Ecuador for building purposes. Illustrated, Sarg. 
Man. Trees N. Amer. 305. 

Peru: (Mathews). Extending north to Mexico. 

Morus insignis Bureau in DC. Prodr. 17: 247. 1873. 

Branchlets at first whitish-tomentose, the branches glabrous, 
flexuous; leaves broadly ovate, unequal at the subcordate base, 
acuminate, very scabrous above, more or less pubescent beneath, 
densely so in youth; stipules 2 cm. long; staminate spikes 4-8 cm. 
long, the peduncles 3-4 mm. long; fruiting spikes often greatly elon- 
gate, subsessile. 

Peru: Ruiz & Pavon. Colombia; Costa Rica. 

8. CHLOROPHORA Gaud. Fustic 

Small trees or shrubs, variable vegetatively, spiny to unarmed, 
with entire to coarsely serrate or 3-lobed leaves, glabrate or hirtellous, 
but distinctive by its unisexual inflorescences, the staminate ament- 
like spikes or sometimes globose heads, the pistillate globose heads; 



310 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

fruit many-seeded and semifleshy. Perianth 4-parted, that of the 
pistillate flower cupulate, thickened at the apex. 

Branches armed with stout, recurved spines; staminate flowers 
capitate C. brasiliensis. 

Branches unarmed or with straight spines; staminate flowers in 
slender aments C. tinctoria. 

Chlorophora brasiliensis (Mart.) Standl., comb. nov. Brous- 
sonetia brasiliensis Mart. Flora 24: Beibl. 10. 1841. Madura bra- 
siliensis Endl. Gen. Suppl. 4: 34. 1847. 

A shrub or small tree, almost glabrous, the branches armed with 
numerous long, abruptly recurved spines; leaves slender-petiolate, 
elliptic or broadly ovate, abruptly short-acuminate, obtuse at the 
base, thin, glabrous, entire or nearly so; staminate heads 5-6 mm. 
in diameter. Illustrated, Mart. Fl. Bras. 4, pt. 1: pi. 54. 

San Martin: Juan Guerra, 720 meters, Williams 6855. Brazil. 

Described by the collector as a vine. The generic position of the 
plant is somewhat doubtful but for the present, at least, it may be 
placed in Chlorophora. 

Chlorophora tinctoria (L.) Gaud. Bot. Freyc. Voy. 508. 1826. 
Morus tinctoria L. Sp. PI. 986. 1753. Madura affinis Miq. in Mart. 
Fl. Bras. 4, pt. 1: 155. 1853. Madura tinctoria (L.) D. Don, var. 
affinis (Miq.) Bureau in DC. Prodr. 17: 230. 1873; var. chlorocarpa 
Bureau, op. cit. 229. 

Leaves varying from lanceolate to elliptic, usually about 10 cm. 
long and several cm. wide, entire or serrate, on petioles 1 cm. long; 
staminate aments 3 to several cm. long, pedunculate as also the 
globose pistillate heads.- This tree is well known as the source of 
fustic dye-wood, furnishing the olive-drab of khaki and other tones 
in greens, browns, and yellows. Its light yellow wood, close-grained, 
durable and taking a high polish, is used in the manufacture of 
furniture, wheels, etc., in some parts of its range. Variable in leaf 
form. Illustrated, Mart. Fl. Bras. 4, pt. 1: pi. 51, 52. 

Amazonas (?): (Mathews 1981}. Yurimaguas, Poeppig 2384. 
Without locality, Ruiz & Pavon; (Mathews). Loreto: Rio Ucayali, 
Tessmann 3366. Iquitos, 120 meters, Williams 8040, 8050, 7217. 
Rio Nanay, Williams 501, 672. Caballo-cocha, Williams 2441. 
Parana-pura, 200 meters, Williams 2441. San Martin: Pongo de 
Cainarachi, 230 meters, Klug 2607. Tarapoto, 750 meters, Williams 
5435. Rio Mayo, in forest, Williams 6254. Brazil to Mexico and 
the West Indies. "Insira," "insira caspi," "limulana." 



FLORA OF PERU 311 

9. SOROCEA St. Hil. 

Small trees with subentire or spinulose-toothed leaves and laxly 
flowered, unisexual racemes or spikes. Staminate perianth present, 
4-parted. Otherwise very similar to Clarisia. 

Leaves spinulose-dentate S. ilicifolia. 

Leaves, if toothed, repandly and bluntly. 
Branchlets puberulent or glabrous. 
Petioles 8-10 mm. long, or longer. 
Leaves bluntly toothed; pedicels 5 mm. long, or longer. 

S. Briquetii. 

Leaves entire or repandly toothed ; pedicels very short. 
Leaves elliptic-obovate, to nearly 10 cm. wide or smaller. 

S. Ulei. 

Leaves elliptic-oblong, to 6 cm. wide or smaller. .S. opima. 
Petioles 2-5 mm. long. 

Leaves glabrous S. muriculata. 

Leaves finely pubescent beneath S. Sprucei. 

Branchlets and especially the leaves beneath hirtellous . . S. hirtella. 

Sorocea Briquetii Macbr. Candollea 4: 311. 1931. 

Glabrous except for a minute puberulence on the pistillate 
peduncle and pedicels; leaves firm-membranaceous, ovate-oblong- 
lanceolate, acute at the base, obtusely acuminate, mostly 10 cm. long 
and 3.5 cm. wide, irregularly serrate, especially above the middle, 
lustrous above and rather prominently net-veined, paler beneath, 
the about 10 principal lateral nerves arcuately joined before the 
margin; peduncle 5-7 mm. long; pedicels to 10 mm. long; fruit sub- 
globose, 5 mm. thick; stigmas apparently acute. Neg. 11637. 

San Martin: Tarapoto, Spruce 4220, type. Rumizapa, 800 
meters, Williams 6803. 

Sorocea hirtella Mildbr. Notizbl. Bot. Gart. Berlin 10: 183. 
1927. 

Young branchlets and leaf nerves beneath short-hirsute with 
spreading hairs; petioles 1 cm. long; leaves often slightly unequal 
at the rounded base, long-acuminate, mostly 10-13 cm. long and 
6 cm. wide, glabrous above, the lateral nerves 10-12; rachis and 
peltate bracts hirtellous; perianth segments glabrate, to 2.5 mm. 
long and nearly 2 mm. broad. A tree of 13 meters growing in upland 
woods, the trunk 21 cm. in diameter. Probably here belongs a Ruiz 



312 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and Pavon fruiting collection in Herb. Boissier with original label 
"Clarisia gen. nov.," on the basis of which the name Clarisia nitida 
(Allem.) Macbr. was proposed (see Clarisia racemosa). Its largest 
elliptic-acuminate leaves are over 20 cm. long, and 10-12 cm. wide. 
The axillary fruiting spikes are minutely pilose, 2-4 cm. long, the 
sessile, crowded, puberulent, globose fruits 5-6 mm. in diameter. 
Neg. 11640. 

Loreto: Mouth of Rio Santiago, 160 meters, Tessmann 4016, 
type. Mishuyacu, 100 meters, in forest, King 739. Caballo-cocha, 
Williams 2092. Without locality: Ruiz & Pavon (?). 

Sorocea ilicifolia Miq. in Mart. Fl. Bras. 4, pt. 1: 114. 1853. 

Distinctive because of the spinulose leaf serration; a glabrous 
shrub or small tree with oblong-elliptic or broadly elliptic, sharply 
acuminate leaves on petioles 1 cm. long or often shorter and with 
racemes 4-6 cm. long. S. Klotzschiana Baill. (S. castaneifolia Huber, 
Bol. Mus. Goeldi 5: 333. 1909) of the Amazon is not obviously 
different. Neg. 18810. 

Junin: La Merced, Weberbauer 1858. Brazil. 

Sorocea muriculata Miq. in Mart. Fl. Bras. 4, pt. 1: 113. pi. 
34. 1853. 

A densely leafy, slender, often flexuously branched shrub or 
small tree, the young branchlets, petioles, and inflorescences puberu- 
lent; leaves thin, oblong-elliptic, bluntly caudate-acuminate, acute 
at the base, mostly 8-10 cm. long and 3 cm. wide, paler and reticulate 
beneath; staminate spikes to 4 cm. long, the shorter pistillate ones 
often fasciculate; stigmas obtuse; fruit muricate-scabrous, thick- 
pediceled. S. Steinbachii Mildbr., with mostly broader leaves, some 
with a tooth or two, might be sought here. Its fruits are mostly 
sessile. S. amazonica (Mart.) Miq. is scarcely distinguishable from 
S. muriculata except by its acute stigmas and smooth fruits. 

San Martin: Near Moyobamba, Klug 3566 (probably, in flower). 
Loreto: Iquitos, 120 meters, Williams 8010. Mishuyacu, 100 
meters, Klug 153. Mouth of Rio Santiago, Tessmann 4059, 4111 
(det. Mildbraed), 5251. 

Sorocea opima Macbr. Field Mus. Bot. 11: 64. 1931. 

A shrub with sparsely puberulent branchlets, densely puberulent 
pistillate racemes, and glabrous leaves; petioles 8 mm. long; leaves 
chartaceous, gray-green, subopaque, rather obscurely veined above, 
conspicuously reticulate beneath, the primary lateral nerves 10-12, 



FLORA OF PERU 313 

subrotund to the acutish or obtuse base, caudate-acuminate, the 
apex very broad (about 5 mm. at the middle) and up to 2 cm. long, 
entire or undulate-margined, the largest to 20, the smallest 10 cm. 
long; racemes extra-axillary, in fruit to 5 cm. long; fruit sessile or 
subsessile on the strongly thickened rachis, finally glabrous, 1.5 cm. 
long and 1 cm. thick, distinctly beaked. 

Loreto: Caballo-cocha, in forest, Williams 2357, type. San 
Martin: Zepelacio 1,200-1,600 meters, mountain forest, Klug 3566. 

Sorocea Sprucei (Baill.) Macbr. Field Mus. Bot. 11: 16. 1931. 
Pseudosorocea Sprucei Baill. Adansonia 11: 296. 1875. 

Apparently glabrous but under a lens the branchlet tips, petioles, 
and leaf nerves beneath more or less puberulent; branchlets short 
and stiff; petioles scarcely 5 mm. long; leaves subcoriaceous, lustrous 
above, paler beneath, mostly about 8 cm. long and 3 cm. wide, short- 
acuminate, entire; staminate spikes subsessile, 1-2 cm. long, the 
flowers about 2 cm. wide, minutely pubescent; pistillate inflorescence 
unknown. Described as entirely glabrous. 

San Martin: Near Tarapoto, Spruce 4483, type. 

Sorocea Ulei Warb. ex Ule, Bot. Jahrb. 40: 141. 1907, nomen. 

Glabrous, with thin, green or grayish leaves, all entire or more 
usually somewhat repand-toothed, elliptic-obovate, to 20 cm. long 
and nearly half as broad but mostly smaller, rather abruptly caudate, 
acuminate, the obtuse point about 1 cm. long, subobliquely and little 
narrowed, or even subrotund, to the acutish base; petioles to 1.5 cm. 
long, somewhat hirsutulous; petals glabrous, about 1 cm. long. 
Described from Ule 5871, from Brazil. S. Steinbachii Mildbr. is appar- 
ently similar but its leaves are more nearly entire, much more 
cuneate below, and on usually shorter petioles. Neg. 25563. 

Loreto: Rio Ucayali, Ule. Cultivated, from the Ucayali, Ducke 
17044; Huber 6977. Brazil. 

10. CLARISIA R. & P. 

Reference: Lanjouw, Recueil Trav. Bot. Ne"erl. 33: 254-276. 1936. 

Trees or shrubs with alternate, bistipulate, petiolate, entire or 
dentate leaves. Flowers dioecious. Staminate spikes peduncled, 
axillary or in short racemes. Perianth none, the 1-staminate flowers 
intermixed with usually peltate bractlets. Pistillate flowers racemose 
or 1-several in small heads, the ovoid perianth adnate to the inferior 
or semi-inferior ovary. Style short, the exserted stigmas short or 



314 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

long. Tall, thick- trunked trees, the wood valuable. On incision, 
the trunk yields abundantly a white resin that in the air is brownish 
and somewhat elastic; it is excellent for water-proofing, and the 
Indians cover their blow guns, etc., with it (Ruiz & Pavon). 

Leaves 2-4 cm. wide, abruptly short-acuminate C. racemosa. 

Leaves 5-7 cm. wide, long caudate-acuminate C. biflora. 

Clarisia biflora R. & P. Syst. 255. 1798. 

A tree with numerous short, very leafy branchlets; petioles sul- 
cate above, minutely appressed-pubescent; leaves oblong-obovate 
to obovate, rounded at the base, abruptly long-cuspidate, the acumen 
nearly 1.5 cm. long, 5-7 cm. wide, the blades 10-15 cm. long or longer, 
chartaceous, conspicuously reticulate and laterally nerved beneath, 
somewhat lustrous above, paler or often reddish brown beneath; 
stipitate pistillate flowers apparently binate at the base of the 
petiole; pedicels 2-3 mm. long, in fruit thick, 7 mm. long; perianth 
4-bracted, 1 bract smaller than the others, suborbicular, subpeltate, 
fimbriate; perianth subglobose, about 5 mm. in diameter, the apical 
opening with a short, cylindric, irregularly lobed collar; styles 7 
mm. long; fruit globose, 2 cm. in diameter. Neg. 29482. 

Huanuco: Cochero, Chinchao, Posuso, Pillao, Chacahuasi, Ruiz 
& Pavdn. San Martin: Zepelacio, 1,100 meters, in forest, Klug 
3698; a tree of 15 meters. Without locality: Weberbauer 7057. 
"Yasmich," "piamich." 

Clarisia racemosa R. & P. Syst. 255. 1798. Soaresia nitida 
Allem. Arch. Palestr. Scien. Rio Jan. 1 : 142. 1858. C. nitida Macbr. 
Field Mus. Bot. 11:15. 1931. 

A tree to 30 meters, the ultimate branches numerous, slender; 
petioles glabrous or sparsely puberulous, 4-15 mm. long; leaves 
oblong-elliptic, rounded at the base, abruptly and obtusely acumi- 
nate, the acumen about 5 mm. long, lustrous above, olivaceous and 
paler beneath, the about 16 lateral nerves scarcely more prominent 
than the reticulate veins; staminate aments usually several in 1-2 
axillary, racemosely arranged inflorescences, the common peduncle 
pubescent, 2-4 cm. long; spike peduncle 3 mm. long, the spikes 2-6 
cm. long, linear-cylindric; stamens intermixed with variously shaped 
bracts, these sometimes coherent basally as a pseudoperianth, the 
rows of stamens bordered by subpeltate bractlets; pistillate flowers 
in racemiform panicles, often 2-3 in the leaf axils, the rachis pilose, 
5-10 cm. long; pedicels 5-10 mm. long, densely pubescent; perianth 



FLORA OF PERU 315 

subtended by 4 broad, membranous-edged bracts, dentate and 
pubescent; perianth cupulate, glabrous except at the thin, obscurely 
lobed opening; stigmas 4-5 mm. long; fruit to 22 mm. long, 16 mm. 
broad, woody, the bracts persisting. C. ilicifolia (Spreng.) Lanj. 
& Rossb., with acute-based, remotely dentate leaves, and C. Spruceana 
Lanj., with congested staminate spikes, the pubescence of the pedun- 
cles reflexed and uncinate, may be expected in Amazonian Peru. Ruiz 
and Pavon found the very durable and highly desirable wood already 
known and appreciated, describing the cortex as blood-red, the roots 
so red that they appeared to have bled. Ducke has observed that 
the leaves vary in size and acumination. He has described the bark 
as brown, verrucose, intensely red within as also the branchlets, the 
wood as yellow, finally becoming brown-yellow, the grain fine. Ruiz 
and Pavon recorded the vernacular name as "tulpay," Tessmann as 
"mashonasti," but the usual Brazilian term is "guariuba." Illus- 
trated, Allem. op. cit. Neg. 29483. 

Huanuco: Chinchao, Posuso, Ruiz & Pavon. Pampayacu, Poep- 
pig. Cochero, Poeppig 1087. Junin: La Merced, Weberbauer 1885, 
1912. Loreto: Yarina-cocha, Tessmann 5433 (det. Mildbraed). 
La Victoria, Williams 3097, 2684. Rio Acre: Ule 9317. Cultivated 
from the Ucayali (Ducke; Huber 6977). Brazil. "Capinuri." 

11. ANONOCARPUS Ducke 

A dioecious tree, the staminate flowers in long spikes, the pistillate 
in elliptic heads forming at maturity a fleshy, subglobose, anona-like 
fruit. Staminate perianth 2-4-parted. Pistillate perianth fleshy, 
strongly compressed, obliquely tubular, the truncate apex with an 
elongate cleft from which are exserted the 2 subsessile stigmas. 

Anonocarpus amazonicus Ducke, Archiv. Jard. Bot. Rio Jan. 
3: 39. 1922. 

A tree, the youngest branchlets and petioles somewhat puberu- 
lent, as also the leaf nerves beneath; petioles to 1 cm. long; leaves 
obovate-oblong, commonly 10-15 cm. long, 4-7 cm. wide, charta- 
ceous; staminate spikes about 1 cm. long or shorter, the white-puberu- 
lent peduncles 1 cm. long; perianth and bractlets ciliate, about 1.5 
mm. long; pistillate peduncle stout, about 1 cm. long, pubescent like 
the head, the latter to 5 cm. long and 4 cm. thick. Neg. 11628. 

Loreto: Puerto Arturo, Yurimaguas, 200 meters, in forest, 
Williams 5334. Amazonian Brazil. "Mashunaste." 



316 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

12. BROSIMUM Sw. 

Reference: Ducke, Archiv. Jard. Bot. Rio Jan. 3: 29-30. 1922. 

Shrubs or trees, marked among the Peruvian genera by the 
globose receptacles that are composed of numerous crowded, round- 
ish, thin-edged bractlets intermixed with the sometimes exserted 
stamens, and contain one or more deeply imbedded pistillate flowers. 
Brosimopsis S. Moore is similar in aspect but the receptacles are 
completely staminate or pistillate, the latter containing several 
flowers; it is allied to Olmedia. There are a number of Amazonian 
species, mostly very well marked in contrast to the Peruvian one, 
which seem, except B. echinocarpum, to center around B. guianense 
(Aubl.) Huber, Bol. Mus. Goeldi 5: 337. 1909, cf. Ducke. Apparently 
nearest ours is B. Lecointei (B. Le Cointei) Ducke, Archiv. Jard. Bot. 
Rio Jan. 3: 28. 1922, "aita." It has the small leaves ofB.Tess- 
mannii, but they are bluntly short-acuminate. Its heartwood is 
white, finally brownish rose instead of reddish purple as that of 
typical B. guianense ("bois de lettre" or "nuirapinima"), to which 
B. Aubletii has been referred. The former, however, has thicker, 
scarcely (and obtusely) pointed leaves. B. discolor Schott has, appar- 
ently, no clear difference, but the wood appears to be unknown. 
From herbarium material and descriptions it seems probable that our 
Brosimums, as here named, are distinct from B. guianense, to which 
Poeppig, Miquel, and Huber referred them, unless, in fact, there is 
but one variable specific entity, as Ducke suggests, for this particular 
group. He has transferred all the Brazilian species to the earlier 
name Piratinera Aubl., but Brosimum Sw. was already conserved. 
Pittier, however (Contr. U. S. Nat. Herb. 20: 99. 1918), retains the 
Aublet name for those species with 2 or more pistillate flowers, the 
staminate being provided with a perianth, a logical segregation in 
view of the acceptance of the presence or absence of a perianth as 
indicating generic demarcation elsewhere in the family. Ducke 
with reason rejects the character in this instance, although accepting 
it elsewhere. 

Leaves faintly and remotely serrate, the largest 7 cm. wide. 

B. echinocarpum. 
Leaves entire. 

Leaves pubescent beneath. 

Leaves obtusely acuminate, about 3 cm. wide. . . .B. Aubletii. 
Leaves acutely acuminate, usually 1.5-2.5 cm. wide. 

B. Tessmannii. 



FLORA OF PERU 317 

Leaves glabrous beneath. 
Leaves mostly 5-6.5 cm. wide, pale when dried, oval-elliptic, 

gradually acuminate B. Uleanum. 

Leaves mostly 3-4 cm. wide, brownish when dried, oblong-oval, 

abruptly caudate-acuminate B. paraense. 

Brosimum Aubletii P. & E. Nov. Gen. & Sp. 2: 34. 1838. B. 
guianense (Aubl.) Huber, as to synonymy only. 

A small tree (as known) with many oblong-obovate, obtusely and 
shortly acuminate leaves mostly about 7 cm. long, lustrous above, 
grayish-puberulent beneath; staminate flowers unknown; pistillate 
peduncle nodding, only equaling the short petiole; bractlets thin at 
the margins; flowers 2 (always); fruit red, succulent, edible, the 2 
seeds globose, smooth. Neg. 11654. 

Loreto: Yurimaguas, Poeppig 24-13. Guianas. 

Brosimum echinocarpum P. & E. Nov. Gen. & Sp. 2: 34. pi. 
148. 1838. 

A shrub or tree with oblong-ovate leaves, subcuneate at the base 
and rather gradually long caudate-acuminate, distinctly but remotely 
serrate, thin, glabrous, to 15 cm. long and 7 cm. wide; heads axillary, 
peduncled, the pistillate rigidly setose, the pubescent setae 4-5 mm. 
long; fruit tuberculate-asperous, orange. Neg. 11656. 

Loreto: Yurimaguas, Poeppig 2255, type. 

Brosimum paraense Huber, Bol. Mus. Goeldi 6: 67. 1910. 

A large tree, glabrous throughout; stipules very narrow, elongate, 
as much as 1.5 cm. long, brown; petioles short, the blades mostly 
oblong-elliptic and 4.5-10 cm. long, coriaceous, usually lustrous, 
the numerous lateral nerves diverging at a wide angle; fruiting heads 
about 12 mm. in diameter, short-stalked. Neg. 11662. 

Loreto: Caballo-cocha, in forest, Williams 2252. Forest near 
Kokamas, Rio Nanay, Williams 725. Forest between Rio Nanay 
and Rio Napo, Williams 683. Encanto, Fox 93. Amazonian Brazil 
and the Guianas. "Palo de sangre." 

There is some doubt regarding the proper determination of the 
Peruvian tree, but the specimens agree rather well with authentic 
material of B. paraense, and are probably conspecific. The tree 
furnishes a valuable timber, used for furniture and other purposes. 

Brosimum Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 
10: 190. 1927. 



318 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Much \ikeB. discolor, according to the author, but the papery 
leaves (mostly 6 cm. long) gradually long-acuminate, the lateral 
nerves about 10; staminate receptacle borne on a slender peduncle 
1 cm. long, subglobose, the yellow, peltate bracts numerous above, 
few below. A tree 25 meters high, the trunk diameter 55 cm., the 
branches much branched. A "magic tree," eaten by the magicians 
(Tessmann). Neg. 11667. 

Loreto: Middle Rio Ucayali, 155 meters, Tessmann 3268, type. 
Puerto Mele"ndez, Tessmann 4860. "Waira caspi." 

Brosimum Uleanum Mildbr. Notizbl. Bot. Gart. Berlin 10: 
191. 1927. 

Glabrous except for the very minutely puberulent young branch- 
lets, stipules, petioles (5 mm. long), peduncles (scarcely 1 cm.), 
and tips of the 2 large bracts; leaves usually oval-elliptic, obtuse 
at the base, short-acuminate, about 8 cm. long and 4-6.5 cm. wide, 
the 16 lateral nerves prominent beneath; receptacles green, sub- 
globose, mostly staminate, 1 cm. broad, a few of the peltate bracts 
1 mm. wide; stamens numerous, exserted, to over 2 mm. long; 
style branches puberulent, nearly 2 cm. long. A tree 40 meters 
high with a trunk diameter of 1.25 meters. In upland forest. Neg. 
11688. 

Loreto: Mouth of Rio Santiago, Tessmann 4523, type. Rio 
Acre: Ule 9324. 

13. OGCODEIA Bureau 

Acanthosphaera Warb. Verh. Bot. Ver. Brandenb. 48: 150. pi. 2. 
1907. 

Pistillate receptacles more or less "spiny" in appearance by the 
accrescence of the subulate bracteoles; staminate receptacles with 
broad, seriately arranged bracts. Branchlets usually conspicuously 
provided with large, stiff stipules. See Mildbraed, Notizbl. Bot. 
Gart. Berlin 10: 186. 1927; 11: 418. 1932, for a redefinition of 
the genus and its relationship with Naucleopsis macrophylla Miq., 
which apparently is unknown from Peru unless 0. Ulei is a synonym 
(cf. Ducke, Arch. Inst. Biol. Veg. 2: 30. 1935). It is not clear to 
me that Ogcodeia is soundly separated; there is too much stress 
placed perhaps on presence or absence of perianth in one sort of 
flower and in bract development, especially when the bracteoles 
here as elsewhere, or the perianth segments when completely divided, 
may separate the flowers indiscriminately, thus morphologically 
serving the same purpose. Besides the following there is 0. amara 



FLORA OF PERU 319 

Ducke with leaves 5-10 cm. broad, the lateral nerves 15-18, and the 
bractlets of the mature receptacle aculeate-linear and 3-5 cm. long. 
It has a very little latex, and is used as a remedy for fevers under the 
names "quina" or "balsamo," terms applied also to Quassia amara 
and Myroxylon peruiferum respectively (Ducke). Another Amazo- 
nian species has been indicated, 0. caloneura (Huber) Macbr. It 
suggests 0. Tessmannii, but the young parts and the bracts are 
puberulent-tomentulose. 

Leaves mostly 3-6.5 cm. wide, with 15-20 lateral nerves. 
Petioles rarely 1 cm. long; nerves rather inconspicuous beneath. 
Leaves cuneate at the base, 3-4 cm. wide. .O.ternstroemiiflora. 

Leaves rounded at the base, 5-6 cm. wide 0. Tamamuri. 

Petioles often longer; nerves conspicuous beneath. 
Leaves rarely 5 cm. wide, the acumen often 2 cm. long. 

0. Tessmannii. 
Leaves often 6-7 cm. wide, the acumen rarely 1.5 cm. long. 

0. glabra. 
Leaves mostly 8-12 cm. wide, the lateral nerves 25-30. . . .0. Ulei. 

Ogcodeia glabra (Spruce) Mildbr. Notizbl. Bot. Gart. Berlin 
11: 418. 1932. Naucleopsis glabra Spruce in herb. 

Branchlets and leaves glabrous, the latter coriaceous, with a very 
prominent costa beneath; lateral nerves 12-14, the veins reticulate; 
petioles 2-2.5 cm. long; leaf blades 4-6 cm. wide, about 15 cm. long, 
the acumen 1 cm. long; pistillate inflorescence 2 cm. wide; bracts 
recurving, oblong-lanceolate, acutish, very minutely puberulent, 
nearly 1.5 cm. long, or the innermost longer; perianth segments 
puberulent, linear-clavate, 5 mm. long; style 2 mm. long, the 
branches twice as long. Description of pistillate inflorescence from 
the following collection, from a tree of 6 meters with brownish 
yellow flowers. Neg. 11649. 

Loreto: Mishuyacu near Iquitos, 100 meters, in forest, Klug 257. 
Amazonian Brazil. 

Ogcodeia Tamamuri Macbr. Field Mus. Bot. 11: 64. 1931. 

Similar to 0. ternstroemiiflora, but the leaves often 16-17 cm. 
long and 5-6 cm. wide, and the staminate receptacles geminate. 
0. caloneura (Huber) Macbr. Field Mus. Bot. 11: 17. 1931, if sought 
here, is distinguishable by its rusty-tomentulose branchlets. 

Loreto: Timbuchi on the Rio Nanay, in forest, Williams 992, 
type. "Tamamuri." 



320 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ogcodeia ternstroemiiflora Mildbr. Notizbl. Bot. Gart. Berlin 
10: 188. 1927. 

Youngest branchlets minutely puberulent; leaves oblongish, cune- 
ate at the base, with an acumen 1-1.5 cm. long, with this mostly 
12-13 cm. long and 3-4 cm. wide, drying pale brown beneath, 
densely but not conspicuously reticulate- veined ; staminate receptacles 
axillary, to 4-fasciculate, on peduncles 5 mm. long, the spiraled 
bracts about 9, puberulent, oval to orbicular, 5 mm. long; bractlets 
1-1.5 mm. long, 0.75 mm. broad, the blade plicate; filaments 1.5 mm. 
long, the anthers scarcely 1 mm. long. A small tree, to 5 meters 
high, or larger, the trunk 20 cm. thick; inflorescences greenish or 
white or dirty white. Fruit eaten. In upland areas (Tessmann}. 
Neg. 11650. 

Loreto: Yarina-cocha, Tessmann 5467, 3228. Iquitos, Tessmann 
5358. Rio Maranon near mouth of Rio Tigre, 115 meters, Killip 
& Smith 27525. San Antonio, 110 meters, in forest, Killip & Smith 
29305. Mishuyacu, 100 meters, King 1131. Soledad, Killip & 
Smith 29810. "Lana." Amazonian Brazil. 

Ogcodeia Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 10: 
189. 1927; 11: 420. 1932. 

Similar to 0. ternstroemiiflora but quite glabrous; leaves thinner, 
with large, conspicuous nerves and veins beneath and glabrous bracts 
about 7 mm. long; pistillate heads nearly 2.5 cm. wide; bracts and 
bractlets very minutely puberulent, the larger inner bracts triangular- 
ovate, acuminate, about 1 cm. long, the perianth segments 4 mm. 
long, rigid-aculeate; style base 2 mm. long, the branches 6 mm. long. 
A tree about 8 meters high, the trunk diameter 8 cm.; bracts 
yellowish. Neg. 11651. 

Loreto: Mouth of Rio Santiago, Tessmann 4109, type. Alto 
Rio Itaya, Williams 3511. "Naccho-huasu." 

Ogcodeia Ulei (Warb.) Macbr. Field Mus. Bot. 11: 16. 1931. 
Acanthosphaera Ulei Warb. Verh. Bot.Ver. Brandenb. 48: 150. pi. 2. 
1907. Naucleopsis Ulei Ducke, Archiv. Jard. Bot. Rio Jan. 3: 38. 
1922. 

A small to tall tree with conspicuous, persistent stipules 1.5-2.5 
cm. long, acuminate from a broad base, and heavy, oblong-elliptic, 
glabrous leaves on short, thick petioles 0.5-1.5 cm. long, the blades 
with many (25-30) parallel lateral nerves and reticulate- veined, 
especially beneath, several dm. long and 8-15 cm. wide; inflores- 
cences axillary, sessile, the staminate 7 mm. long, 4 mm. broad; 



FLORA OF PERU 321 

stamens with their anthers and acute bractlets prominent, the 
former 3 mm., the latter 2 mm. long; pistillate inflorescences 5 mm. 
broad and long, the bractlets rounded at the tip, subasperous; style 
2 mm. long, the branches longer; fruit globose, the aculeate bractlets 
3.5 cm. broad, 3 cm. long, the bractlets themselves 8 mm. long and 
2 wide. Mature fruit pale yellow, sweet, edible (Ducke). The 
Killip and Smith numbers were determined by Smith as N. macro- 
phylla, "probably identical with N. Ulei, the specific status of which 
is doubtful." 

Loreto: Iquitos, Ule 6257; (Ducke}. Yurimaguas, 200 meters, 
Williams 5194; Killip & Smith 28179, 28244. San Antonio, 110 
meters, Killip & Smith 29300. Amazonian Brazil. 

14. PEREBEA Aubl. 

Trees with usually thick leaves and unisexual, flattish to sub- 
globose, many-flowered receptacles, the pistillate conspicuously 
pedunculate. Staminate perianth 4-parted, the pistillate 4-toothed. 
Ovary semi-inferior or completely free. Style very short, bidentate. 
The rubber trees, Castillo,, to which one of the following species has 
been referred, are very similar but the style is typically longer and 
the stamens are mixed with scales. Filiform stigmas and elongate 
bracts, the inner exceeding the perianth, are characters that serve 
to distinguish Noyera. Olmedioperebea Ducke, to be expected, 
differs in having only 1-4 pistillate flowers, these coalescent; the 
staminate perianths, too, are very short. In all these characters, 
other related plants considered, one may see degrees of development 
rather than fundamental distinctions. Olmedia calophylla P. & E., 
which probably will be found, may well belong to Perebea (P. calo- 
phylla Benth. & Hook.). It has heavy, callous-margined leaves, 
obviously scabrous beneath. P. guianensis Aubl. has already been 
found on the upper Amazon; its nearly glabrous, very large, and 
thin leaves are somewhat serrate and its ovary is completely superior 
(Ducke). 

Leaves distinctly pseudopeltate P. pseudopeltata. 

Leaves acute at base or merely subcordate. 
Leaves 20 cm. long or longer. 

Pubescence neither scabrous nor stellate; leaves subcordate or 
emarginate at the base. 

Leaves oblique at the base; stigmas Ungulate. P. Tessmannii. 

Leaves not oblique; stigmas cushion-shaped P. australis. 



322 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence of the leaves beneath scabrous or stellate-puberu- 
lent; leaves rounded to an acute base. 

Leaves scabrous beneath P. Chimiqua. 

Leaves stellate-puberulent beneath P. Standleyi. 

Leaves 10 cm. long or shorter P. elegans. 

Perebea australis (Hemsl.) Macbr. Field Mus. Bot. 11: 17. 
1931. Castillo, australis Hemsl. in Hook. Icon. IV. 7: pi. 2676. 1901. 

Young branchlets and leaves silky-tomentose or the latter gla- 
brous above and hirsute beneath mostly on the costa and lateral 
nerves; petioles 1 cm. long; blades coriaceous, oblong-lanceolate, 
30-40 cm. long and 10-20 cm. wide, slightly emarginate at the base, 
abruptly acuminate; primary nerves 17-19, joined before the undu- 
late margin, prominent only beneath; pistillate receptacles axillary, 
solitary, on stipes 2-2.5 cm. long; bracts in several series, acuminate, 
free or nearly so; perianth free(?), lobed, pubescent; style short, 
thick, hairy, the stigmas cushion-like. Pittier, Contr. U. S. Nat. 
Herb. 13: 258-260. 1910, cites this species as the only Castilla with 
cushion-shaped stigmas. The species, as apparently well illustrated, 
seems to be a typical Perebea. Probably the well known name Cas- 
tilla, later than Perebea, should be conserved to include the latter 
group and several other segregates, as remarked elsewhere. "Com- 
mon in woods," according to the collector, at 1,300-1,700 meters, 
in January, 1866, but apparently never collected since and the 
staminate flowers unknown. 

Cuzco(?): Moro Zungo (Pearce). Junin: Near La Merced, 800- 
1,300 meters, Killip & Smith 23877; a tree of 15-20 meters, the trunk 
45 cm. in diameter. 

Perebea Chimiqua Macbr. Field Mus. Bot. 11: 63. 1931. 

Leaves distinctly although finely scabrous-puberulent beneath, 
shortly caudate-acuminate, not at all cordate at the base, to 25 cm. 
long and 12 cm. wide, often less than half as large, on petioles 1 cm. 
long, thinnish; staminate receptacles 2-4 in the axils, short-peduncu- 
late, 5 mm. wide; bracts broadly ovate, 1.5 mm. wide, scarcely acute, 
strigose; perianth segments 4, little thickened at the apex; fila- 
ments very short; anthers subrotund. Near P. guianensis Aubl., 
with leaves slightly cordate and nerves more or less hirsute. 

Loreto: San Antonio, Alto Rio Itaya, 145 meters, Williams 3412, 
type. "Chimiqua." 



FLORA OF PERU 323 

Perebea elegans Macbr. Field Mus. Bot. 11: 63. 1931. 

A tree of 20 meters; young branchlets rusty- tomentulose; petioles 
5-7 mm. long; leaves oblong-elliptic, slightly oblique at the acute 
base, short-acuminate, to 5 cm. wide, slightly puberulent on the reticu- 
late veins and nerves beneath, the latter about 12; staminate recep- 
tacles in 3's (mostly), globose, 7 mm. thick, the peduncles nearly 5 
mm. long; bracts scale-like; perianth segments 4 mm. long, strongly 
fleshy-cucullate at the apex, puberulent; filaments 4, thick, 2 exserted. 
Perhaps nearest P. laurifolia. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 603, type. 

Perebea pseudopeltata Mildbr. Notizbl. Bot. Gart. Berlin 
10: 184. 1927. 

A tree, readily known by its foliage, the lobes, of its deeply cordate 
leaves so closely investing the petiole apex that is sunken in the leaf 
base that the leaves appear to be peltate; they are oval-oblong, 
abruptly acuminate, about 50 cm. long and nearly 25 cm. wide, 
glabrous above, but appressed setose-pilose and puberulent on the 
obvious (20-23) lateral nerves and the finely reticulate venation 
beneath; stipules densely long-silky-pubescent; staminate recep- 
tacles 6-12-fasciculate on short branchlets, 1.5-2.5 cm. wide (young), 
the many marginal bracts in 2-3 irregular series, about 2.5 mm. long, 
the outer broader, fewer; perianth segments 4, widened and thickened 
below the tip. Greenish yellow heads unpleasantly scented. A 
tree about 15 meters high, branching at 8 meters, the trunk 22 cm. 
in diameter. In upland woods. Neg. 11680. 

Loreto: Mouth of Rio Santiago, in upland forest, Tessmann 4552, 
type. 

Perebea Standleyi Macbr., sp. nov. 

Arbor; ramulis junioribus pedunculisque fulvo-stellato-tomen- 
tulosis; petiolis circa 6 mm. longis; foliis circa 30 cm. longis 12 cm. 
latis, membranaceis, basi rotundato-acutis, apice ignoto, ubique 
costa subtus excepta minute parceque stellato-puberulenta glabris, 
nervis lateralibus circa 18 solum subtus cum venis reticulatis promi- 
nulis; pedunculis circa 1 cm. longis; receptaculis 2 cm. crassis; fructi- 
bus angulatis obovoideis 1 cm. longis, ad apicem exceptis glabris, 
apice dense fulvo-stellato-tomentosis 7 mm. crassis. 

A tree, the branchlet tips and peduncles rusty stellate-tomentu- 
lose; petioles 6 mm. long; leaves nearly 30 cm. long, 12 cm. wide, 
rounded to the acute base, the apex unknown, membranous, glabrous 



324 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

except the minutely and sparsely stellate-puberulent costa beneath; 
lateral nerves about 18, with the reticulate veins prominent only 
beneath; peduncles 1 cm. long; fruiting receptacles 2 cm. thick, the 
angled fruits obovoid, 1 cm. long, glabrous except at the 7 mm. 
broad apex which is velvety rusty-tomentose. 

Loreto: Yurimaguas, Killip & Smith 28340 (Herb. Field Mus. 
No. 615,971, type). 

Perebea Tessmannii Mildbr. Notizbl. Bot. Gart. Berlin 10: 
185. 1927. 

A tree 25 meters tall, with few branches, the young ones chestnut- 
colored and appressed-setulose; petioles to 12 mm. long; leaf blades 
ovate or elliptic-ovate, 25-30 cm. long and usually 12-15 cm. wide, 
more or less markedly oblique at the base, with an acute acumen as 
much as 2.5 cm. long, at first somewhat appressed-setulose above, 
beneath laxly sericeous-setulose on the costa and conspicuous (about 
20) lateral nerves and veins; pistillate receptacle 1.5 cm. in diameter, 
the squamate bracts passing from rotund-ovate to narrowly triangu- 
lar; perianth urceolate, 5 mm. long, pubescent, the Ungulate stigma 
1.5 mm. long, glabrous; staminate receptacles 3-5 in the axils, on 
peduncles 1.5 cm. long, somewhat hispid-pilose, 8-12 mm. broad, the 
bracts similar but fewer than in the pistillate; perianth segments 
rounded at the tip, tomentose, 2 of them broader; filaments rigid, 
connate at the base. Var. Ulei Mildbr. (Rio Acre) has smaller 
leaves with 15-16 nerves, the midrib pubescence beneath spreading. 
Trunk diameter to 31 cm.; branching at 12 meters. In upland 
forests. The Klug specimen has leaves to 40 cm. long, distinctly 
cordate at the base, the basal lobes overlapping, and suggests P. 
pseudopeltata. Neg. 11681. 

San Martin: Pongo de Cainarachi, Klug 2719 (det. Standley). 
Loreto: Mouth of Rio Santiago, Tessmann 4174, 4601. Rio Acre: 
Ule 9325. 

15. GASTILLA Cervantes 

Reference: Pittier, Contr. U. S. Nat. Herb. 13: 247-279. 1910. 

Very similar to Perebea, but the single Peruvian species distin- 
guished by the short, harsh pubescence on the upper surface of at 
least all but the oldest leaves. Ducke has remarked that the foliage 
of P. guianensis, as yet not known from Peru, is, however, similar 
to that of C. Ulei, and this similarity probably substantiates the 
suggestion that the genera are not natural (see remarks under Pere- 



FLORA OF PERU 325 

bea}. In Central America the genus is a source of rubber. The 
name has been written Castilloa. 

Castilla Ulei Warb. Bot. Jahrb. 35: 674. 1905. 

Branchlets more or less rusty-hispid, glabrous in age; petioles 
5 mm. long; leaves oblong-elliptic, short-acuminate or apiculate, 
10-25 cm. long, 4-6 cm. wide, appressed-tomentose beneath; fruit 
solitary, axillary, subglobose, 2 cm. long, 2-2.5 cm. wide, the 3-5 
seeds 12 mm. long, 6 mm. wide. Illustrated, loc. cit.; Contr. U. S. 
Nat. Herb. 13: pi. 22. 

Loreto: Pebas, Williams 1802. Rio Acre: Ule 9318. Brazil. 
"Caucho negro," "caucho." 

16. NOYERA Trecul 

A tree with the foliage of the Peruvian species of Castilla, but the 
interior bracts of both sorts of receptacles much longer than the 
outer and the staminate receptacles long-peduncled. Style branches 
(stigmas) long and slender. 

Noyera mollis (Poepp.) Ducke, Archiv. Jard. Bot. Rio Jan. 3: 
37. 1922. Olmedia mollis Poepp. Nov. Gen. & Sp. 2: 31. pi 144- 1838. 

Branchlets hirsute; leaves subsessile, oblong, acute, about 20 cm. 
long, 7 cm. wide, pustulate-setulose above, hirsute beneath; staminate 
receptacles several, fasciculate, with linear interior bracts; perianth 
3-4-parted; stamens 2-4; pistillate receptacles sessile, 8-12-flowered; 
ovary semi-inferior. A tree as much as 20 meters high, with an 
abundant, grayish yellow latex, the pale yellow wood worthless; 
mature fruits greenish yellow, the drupes separating easily (Ducke). 
At Obidos called "muiratinga da terra firma" (Ducke). Neg. 11687. 

Loreto: Tessmann 5465 (det. Mildbraed). Amazonian Brazil. 
"Caucho-rana." 

17. PSEUDOLMEDIA Trecul 

Small to large trees with smooth, puberulent, or sparsely hir- 
tellous leaves and small, unisexual, bracteate receptacles, the pistil- 
late 1 (-3) -flowered, and the bracts of these all tiny, those of the 
staminate often elongate, at least the interior. Ovary adherent to 
the perianth. Olmedioperebea Ducke is similar to this and Olmedia 
but the pistillate flowers if more than 1 (1-4) are concrescent, the 
stigmas are short and thick, and bracteoles are lacking in both sorts 
of receptacles. It is nearly Perebea except for its few-flowered pistil- 
late receptacles, and its existence suggests that Perebea must naturally 



326 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

include all the Olmedieae, including Castilla, which well-known name 
ought to be conserved for the genus, sens. lat. The latter, as well 
denned by Pittier, illustrates the great variation in characters else- 
where accepted as of generic import. There is no sound distinction 
between these small groups. The recently redefined Ogcodeia may 
have some merit as a segregate because of its exaggerated bracteole 
development and divided perianths, but these are relative develop- 
ments, as is illustrated in the case of the former by Perebea acantho- 
gyne Ducke. 

Petioles 2-5 mm. long; leaf areoles shallow. 
Branchlets and leaves glabrous, the latter mostly 2-4 cm. wide. 

Leaves thin, 2-3 cm. wide P. laevigata. 

Leaves thick, mostly 4-4.5 cm. wide P. Mildbraedii. 

Branchlets and leaves (5-10 cm. wide) somewhat pubescent, not 
distinctly scabrous. 

Pubescence of appressed puberulence P. macrophylla. 

Pubescence sparse, hirtellous, evanescent P. laevis. 

Branchlets and leaves distinctly scabrous P. scabra. 

Petioles 8-10 mm. long; leaf areoles very deep P. multinervis. 

Pseudolmedia laevigata Tre"cul, Ann. Sci. Nat. III. 8: 131. 
1847. 

Similar to P. laevis but glabrous and with thinnish, much smaller, 
oblong-lanceolate, obtusely acuminate leaves with an acute, slightly 
oblique base, 5-9 cm. long; lateral nerves 15-18, not prominent; 
staminate receptacles sessile, 2-several, the roundish bracts appressed- 
puberulent, with scarious, ciliate-subfimbriate margins. P. Huberi 
Macbr. Field Mus. Bot. 11: 16. 1931 (P. obliqua [Huber] Ducke, not 
[Karst.] Benth. & Hook.) has broader leaves, very oblique at the 
rounded or truncate base. 

Peru: (Pawn?}, Herb. Webb (type). Loreto: Iquitos, Ducke 
751 3 (?). Without locality: Weberbauer 6965. 

Pseudolmedia laevis (R. & P.) Macbr. Field Mus. Bot. 11: 
16. 1931. Olmedia laevis R. & P. Syst. 258. 1798. 

Young branchlets subcompressed, hirsute with spreading hairs, 
the leaves beneath on the conspicuous lateral nerves (15-18) sparsely 
so but subappressed or finally glabrous; leaves subcordate or rotund 
at the base, gradually acuminate, rather thick, about 15 cm. long and 
half as wide; pistillate receptacles sessile in the leaf axils, the bract- 



FLORA OF PERU 327 

lets strigose, the perianth densely soft-pubescent. Perhaps only a 
single variable species is concerned with this, together with P. 
laevigata and P. macrophylla. P. ferruginea (P. & E.) Tre'cul, to be 
expected, is similar to P. laevis, but the branchlets are hirsute and 
the leaves are very oblique at the base. The trunk of P. laevis on 
incision yields abundantly a white latex which on exposure to the 
air condenses into an elastic resin, reddish chestnut in color (Ruiz & 
Pavon). Neg. 11670. 

Junin: La Merced, Weberbauer 1899. Huanuco: Posuso region, 
Ruiz & Pavon. San Martin: Juanjui, 400 meters, in forest, King 
3819; a tree of 15 meters. 

Pseudolmedia macrophylla Tre'cul, Ann. Sci. Nat. III. 8: 132. 
1847. 

A tree or shrub with densely appressed-pubescent branchlets 
(eventually glabrate) prominently marked by the stipular scars; 
leaves oblong-elliptic, subrotund at the base, rather abruptly acumi- 
nate, with a linear, obtuse tip, appressed-pubescent beneath on the 
nerves, 15-20 cm. long, 5-10 cm. wide, the lateral nerves 17-18; 
staminate receptacles geminate, 7-8 mm. broad, the roundish bracts 
densely yellowish appressed-pubescent. Neg. 18815. 

Peru: (Pavdn). 

Pseudolmedia Mildbraedii Macbr. Field Mus. Bot. 11: 61. 
1931. 

A tree about 15 meters high, the staminate receptacles star-like, 
the light yellow stamens and several narrow bracts spreading as 
points; leaves 8-9 cm. long, the 15 lateral nerves conspicuous, as 
also the reticulate venation above, not at all oblique at the acute 
base, obtusely and broadly caudate at the apex or shortly obtuse- 
acuminate; bracts thin but not scarious and obscurely ciliate on the 
margins. 

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

Pseudolmedia multinervis Mildbr. Notizbl. Bot. Gart. Berlin 
10: 189. 1927. 

A tree sometimes 60 meters high, with a trunk diameter of 1 
meter, the densely leafy branchlets closely and persistently hirsute; 
leaves ovate-oblong-lanceolate, mostly obliquely rounded at the 
base, more or less gradually acuminate, usually 9-14 cm. long, 3-5 
cm. wide, above glabrous except on the sunken costa, which is strongly 



328 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

hirsute beneath, the 17-22 conspicuous nerves less so, the areoles 
between the numerous veins deeply granulate-rugulose; staminate 
receptacles 1-1.5 cm. wide, the principal bracts broadly triangular, 
sericeous-pubescent, as also the tips of the (2-3 mm. long) spatulate 
bractlets; anthers 1.5-2 mm. long, apiculate, glabrous. In upland 
forest. Fruit eaten. Neg. 11671. 

Loreto: Mouth of Rio Santiago, Tessmann 4697. Soledad, 110 
meters, Killip & Smith 29809. Fortaleza, Lower Rio Huallaga, 
Williams 4379. Rio Acre: Ule 9322. "Chimicua," "itauba amarilla." 

Pseudolmedia scabra Macbr. Field Mus. Bot. 11: 62. 1931. 

A tree of 4 meters, readily recognizable by the dense and fine but 
"sticky," scabrous puberulence that covers the leaves on both sides; 
leaves oblong-elliptic, equal and acute at the base, obtuse or nearly 
so at the apex, finally 11 cm. long, 4-5 cm. wide, paler beneath, the 
9 lateral nerves and veins prominent; staminate receptacles axillary, 
2-3, on peduncles 5-7 mm. long, globose, 5 mm. thick; bractlets 
little thickened at the apex, 1 mm. long; filaments slender, nearly 
1.5 mm. long, the anthers nearly 1 mm. long. 

Loreto: Mishuyacu near Iquitos, 100 meters, in forest, King 384, 
type. 

18. OLMEDIA R. & P. 

Very much like Pseudolmedia in aspect, but the staminate flowers 
ebracteolate and the ovary free. 0. calophylla P. & E. (possibly a 
Perebea) is well marked by its large, thick leaves, lustrous above, 
opaque and scabrous beneath, and with a definite cartilaginous 
margin. 

Leaves glabrous, 1-2.5 cm. wide '. 0. angustifolia. 

Leaves scabrous, at least beneath, mostly wider. 

Leaves somewhat scabrous-pubescent, even above 0. aspera. 

Leaves smooth above 0. Poeppigiana. 

Olmedia angustifolia P. & E. Nov. Gen. & Sp. 2: 30. pi. 143. 

1838. 

A small tree with slender, flexuous branchlets, the youngest 
strigose-puberulent, and with numerous thin, oblong, entire (or 
with 1 or 2 remote teeth toward the tip) leaves mostly 8-10 cm. long, 
acute at the base, caudately acuminate, the tip often 1.5 cm. long; 
receptacles with ovate-deltoid, sparsely strigillose bracts and many 
silky-pubescent flowers. 

Loreto: Yurimaguas, Poeppig 2270, type. 



FLORA OF PERU 329 

Olmedia aspera R. & P. Syst. 257. 1798. 

Very well marked by the extremely scabrous leaves and, to a 
less extent, the younger branchlets; leaves thinnish, obscurely 
and remotely toothed, often 15 cm. long and 4-6 cm. wide or larger; 
staminate receptacles sessile or nearly so. A shrub or tree of 3-10 
meters (Ule) . Incised, the trunk emits copiously a white latex similar 
to that of Pseudolmedia laevis (Ruiz & Pavon). A large tree; the 
inner bark, beaten and washed, used for making mats in which the 
Indians carry their baggage on journeys, to serve them in place of 
beds (Huber). Neg. 11674. 

Huanuco: Posuso and Chinchao, Ruiz & Pavon. Cochero, 
Poeppig 1267. Junin: La Merced, 600 meters, 5269; Weberbauer 
1916, 1871; Killip & Smith 23464, 23816. Chanchamayo Valley, 
1,500 meters, Schunke 273, 272. Colonia Perene*, 680 meters, Killip & 
Smith 25057. Rio Paucartambo Valley, 700 meters, in forest, Killip 
& Smith 25369. Colonia Perene, Weberbauer 91. San Martin: 
Rio Chipurana, Huber 1548. San Roque, 1,400 meters, Williams 
7195, 7082. Tarapoto, 750 meters, Williams 6022, 6019, 6544. 
Juanjui, 400 meters, in forest, Klug 3787. Without locality: Weber- 
bauer 7059. Ecuador to Bolivia. "Llanchama," "minshi-pata." 

Olmedia Poeppigiana Klotzsch, Linnaea 20: 525. 1847. 

Probably only a variety or form of the above. Of greater interest 
is the related gigantic tree, 0. maxima Ducke, Archiv. Jard. Bot. Rio 
Jan. 3: 32. pi. 3. 1922, "muiratinga" of the Amazon, with a yellowish 
white wood, said to be valueless. Its rather abruptly acuminate 
leaves are minutely pilose or glabrate beneath and the staminate 
receptacles are pedunculate, with a few small, tomentulose bracts 
at the base. Neg. 11677. 

Huanuco: Cochero, Poeppig 1267. Loreto: Puerto Mele"ndez, 
Tessmann 3914- 

19. HELICOSTYLIS Trecul 

A tree similar to Perebea, but the stigmas long and slender and 
somewhat contorted. The typical and Peruvian species is dis- 
tinguishable, however, by the characteristic reddish brown tomentum 
of the receptacles and growing parts, which is at least less markedly 
present in other Moraceous genera. 

Helicostylis tomentosa (P. & E.) Macbr. Field Mus. Bot. 
11: 16. 1931. Olmedia tomentosa P. & E. Nov. Gen. & Sp. 2: 32. 
pi. 145. 1838. H. Poeppigiana Trecul, Ann. Sci. Nat. III. 8: 134. 1847. 



330 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Typically a large tree, the petioles, leaves beneath (on the 
prominent nerves and veins), peduncles, and receptacles reddish 
velvety-pubescent; leaf blades oblong, acute at the base, acuminate, 
glabrous above except on the 11-16 nerves, 8-18 cm. long, 4-6 cm. 
broad; petioles 6-10 mm. long; staminate receptacles 3-10-fascicu- 
late, the peduncles 5-13 mm. long; pistillate receptacles solitary, 
globose. The Rivero plant is a variety with puberulent branchlets, 
elliptic-oblong leaves subrotund at the base, abruptly acuminate, 
glabrous above and shortly hirtellous or glabrate beneath, 12-25 
cm. long, 5.5-9 cm. broad, on petioles 1.5 cm. long. The specimen 
may be rather a Perebea. 

Loreto: Florida, Rio Putumayo, 180 meters, in forest, a tree of 
5 meters, King 2295(1}. Without locality: (Rivero]. Brazil; 
British Guiana. 

20. TRYMATOCOCCUS P. & E. 

Reference: Lanjouw, Recueil Trav. Bot. Neerl. 32: 262-272. 1935. 

Generally a small tree with oblongish, acuminate, undulate, 
very shortly petioled leaves and solitary, short-pediceled, subglobose, 
nodding receptacles. Pistillate flowers deeply immersed, the perianth 
wanting, the staminate 3-lobed, with 3 stamens. Ducke has 
recorded an abundant latex, which Poeppig did not find, perhaps 
testing only young, fertile branches. 

Trymatococcus amazonicus P. & E. Nov. Gen. & Sp. 2: 30. 
pi 142. 1838. 

Leaves rigid, glabrous, or slightly lepidote above, to 15 cm. long 
and 7 cm. broad, or often smaller, conspicuously yellowish net- 
veined beneath; stamens included, the filaments coalescent below; 
style exserted, bifid; fruit hard, 1-seeded, about 2 cm. thick. T. 
turbinatus (Baill.) Ducke, Archiv. Jard. Bot. Rio Jan. 3: 73. 1922, has 
tomentose branchlets and petioles and much larger (2.5-3 cm. 
across in fruit) receptacles that are velvety-pubescent. The thickly 
veined, obovate-elliptic leaves of T. paraensis Ducke, Archiv. Jard. 
Bot. Rio Jan. 3: 22. 1922, are mostly smaller, being only 4-5 cm. 
long and 3 cm. wide, and the latex is greenish. 

San Martin: Tarapoto, Spruce 3895. Pongo de Cainarachi, 230 
meters, Klug 2718. Loreto: Iquitos, Tessmann 5350, 5354 (det. 
Mildbraed); (Ducke 7537); Killip & Smith 26997. Yurimaguas, 
Poeppig 2630, type; at 135 meters, Killip & Smith 29052, 29114; 
in forest, Williams 4525, 3875. Punchana, 120 meters, in forest, 



FLORA OF PERU 331 

Williams 3767. Pebas, Williams 1882. San Juan, Williams 3735. 
La Victoria, edge of forest, Williams 2965. Brazil. 

49. URTICACEAE. Nettle Family 
By E. P. Killip 

References: Weddell, Ann. Sci. Nat. III. 18: 197-232. 1852; IV. 
1: 173-212. 1854; Arch. Mus. Paris 9. 1856-1857; DC. Prodr. 16, 
pt. 1: 32-235. 1869; Blume, Mus. Bot. Lugd. Bat. 2: 43-170. 1852- 
1856; Killip, "Notes on Peruvian Urticaceae of the Marshall Field 
Exploration," Journ. Wash. Acad. Sci. 15: 48-56. 1925. 

Monoecious or dioecious herbs, shrubs, or trees, sometimes armed 
with stinging hairs and often bearing cystoliths; leaves simple, 
alternate, with those of the adjacent nodes often unequal, or opposite, 
with those of a pair often unequal, entire, toothed, or rarely lobed, 
stipulate; flowers small, greenish or greenish white, in unisexual or 
androgynous clusters, the perianth 2-5-lobed or parted, or sometimes 
wanting; fruit an achene. 

Leaves opposite, those of a pair often unequal; plants mostly herba- 
ceous. 
Plants armed with stinging hairs; perianth of pistillate flowers 4- 

parted 1. Urtica. 

Plants without stinging hairs; perianth of pistillate flowers 3-parted, 

or tubular and 2-4-dentate. 
Stigma short-penicillate; inflorescence paniculate, cymose, or 

capituliform; plants usually herbaceous 2. Pilea. 

Stigma filiform; flowers in sessile clusters in the leaf axils or in 

spikes; plants woody 3. Boehmeria. 

Leaves alternate. 

Perianth of pistillate flowers none. 

Flowers in axillary glomerules 4. Phenax. 

Flowers in long, slender spikes 5. Myriocarpa. 

Perianth of pistillate flowers present. 
Plants woody. 

Stigma sessile, penicillate; plants usually armed with stinging 

hairs 6. Urera. 

Stigma filiform; plants without stinging hairs. 

Pistillate perianth strongly nerved ; achenes crustaceous, light 
brown to white; leaves entire (toothed in one Peruvian 
species), those of the adjacent nodes subequal. 

7. Pouzolzia. 



332 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pistillate perianth nerveless; leaves toothed, those of the 

adjacent nodes often very unequal 3. Boehmeria. 

Plants herbaceous. 
Leaves toothed ; plants armed with stinging hairs; inflorescence 

paniculate 8. Fleurya. 

Leaves entire; plants unarmed; inflorescence glomerulate. 

9. Parietaria. 
1. URTICA L. Nettle 

Annual or perennial herbs armed with stinging hairs; leaves 
petiolate, toothed or incised; plants monoecious or dioecious, the 
flowers in androgynous or unisexual clusters, axillary and subsessile, 
or forming elongate spikes or panicles, 4-parted. 

Leaves flabellate-incised, the segments lobed U. flabellata. 

Leaves ovate to ovate-lanceolate, or cordate. 

Inflorescences unisexual; leaves rufo-hirsutulous beneath. 

U. Macbridei. 
Inflorescences androgynous; leaves glabrous, or pilose or pilosulous 

with pale hairs. 
Achenes not more than 1 mm. long, very slightly flattened; 

leaves thick, usually rugose-bullate U. echinata. 

Achenes 1.2-2 mm. long, strongly flattened; leaves thin, flat. 
Leaves incised-dentate; flowers in glomerules; plants annual. 

U. urens. 

Leaves coarsely dentate or serrate; flowers in glomerules or 
elongate spikes; plants perennial U. magellanica. 

Urtica echinata Benth. PI. Hartw. 252. 1846. U. andicola 
Wedd. Ann. Sci. Nat. III. 18: 198. 1852. 

An annual herb up to 40 cm. high, usually densely covered 
throughout with stinging hairs; stem pilosulous; leaves ovate or 
ovate-orbicular, 0.5-3 cm. long, coarsely dentate, acute, usually 
rugose-bullate and pubescent beneath; flowers in sessile, androgyn- 
ous clusters shorter than the petioles; achenes 0.8-1 mm. long, 
apiculate, very slightly compressed. 

Huanuco: Bafios, Wilkes Expedition. Junin: At 4,000 meters, 
Mackenzie in 1924. Arequipa: Nevado de Chachani, 4,200 meters, 
Pennell 13289. Tacna: Volcan Tacora, 4,000-4,500 meters, Weddell 
4440 (type of U. andicola); Werdermann 1147. Cuzco: Cordilleras 
del Pachatusan, 4,400 meters, Herrera 2578. Also in Ecuador and 
Bolivia. 



FLORA OF PERU 333 

Urtica flabellata HBK. Nov. Gen. & Sp. 2: 40. 1817. 

An annual, branched from the base, sparingly bristly on the 
stem, densely so on the foliage; leaves thick, flabellate-incised, with 
the segments sub trilobate, rugose-bullate; panicles androgynous, 
densely flowered, shorter than the petioles; achenes 1.5-2 mm. long, 
acute, strongly flattened. 

Huanuco: Pampas de Bombon, Ruiz & Pavon. Junin: Raimondi 
1186. Huancayo, 3,400 meters, Killip & Smith 22118. Cuzco: 
Paucartambo Valley, 3,600 meters, Herrera 2321. Puno: San 
Antonio, Lechler 1802. Sandia, Weberbauer 969. Without locality: 
Ruiz & Pavon; Gay 979. Also in southern Colombia, Ecuador, and 
Bolivia. 

Urtica Macbridei Killip, Journ. Wash. Acad. Sci. 15: 49. 1925. 

An erect herb, about 1.5 meters high, armed with a few weak 
bristles; leaves sparse, cordate-ovate, 6-12 cm. long, 4-8 cm. wide, 
acuminate, crenate, densely hirsutulous on the nerves and veins 
beneath; plants monoecious, the flower spikes unisexual, the stami- 
nate spikes 12-15 cm. long, in the upper axils, the pistillate 3-3.5 
cm. long, in the lower axils; achenes 1 mm. long. 

Huanuco: Mufia, 2,400 meters, 4271 (type). 

Urtica magellanica Poir. in Lam. Encycl. Suppl. 4: 223. 1816. 

A perennial herb, up to 80 cm. high, the stem glabrous or pubes- 
cent, sparingly or sometimes densely armed with bristles; leaves 
ovate or ovate-lanceolate, 2-7 cm. long, 1.5-5 cm. wide, acuminate, 
cordate or rounded at the base, coarsely serrate or dentate; flowers 
in androgynous glomerules or slender spikes, the staminate flowers 
about 2 mm. wide; achenes 1.2-1.5 mm. long, strongly compressed. 

Cajamarca: Cutervo, Raimondi 4101. Tambillo, Raimondi 
4340, 4392, 7053. Lima: Rio Blanco, 3,300 meters, Killip & Smith 
21629. Huanuco: Llata, 2,100 meters, 2268. Junin: La Oroya, 
Rose 1 8699. Cuzco: Ollantaitambo, 3,000 meters, Cook & Gilbert 
636, 798, 1204. Cuzco, Herrera 533, 679. Puno: Puno, Soukup 
12. Dept. uncertain: San Damian, 2,400-3,000 meters, Hrdlicka 
in 1913. Without locality: Gay 589, 2159. Also in Bolivia, Chile, 
and Argentina. "Apiquisa," "eelula quisa," "quisa," "mula-quisa," 
"yana-quisa." 

These specimens show much variation in the length of the 
inflorescences, and perhaps more than a single species is represented. 
U. ballotaefolia, a common plant of Colombia, is similar to U. 



334 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

magellanica, and some of the Peruvian material listed above has been 
referred to it. Other specimens have been cited in literature as U. 
bracteata Steud. 

Urtica urens L. Sp. PI. 984. 1753. 

An erect or ascending annual, 15-50 cm. high, with a simple or 
few-branched stem; leaves crowded, oval, 2-5 cm. long, 0.6-3 cm. 
wide, rarely larger, rounded or acute at the ends, incised-dentate; 
panicles androgynous, subsessile, usually shorter than the petioles; 
achenes 1.5-2 mm. long. 

Lima: Matucana, 2,400 meters, 11 '4. Surco, 1,500 meters, Killip 
& Smith 21534. Rio Blanco, 3,000-3,500 meters, Killip & Smith 
21627, 21709. Junin: La Oroya, 3,650 meters, 974. Tarma, 3,100 
meters, Killip & Smith 21781. Cuzco: Ollantaitambo, 3,000 meters, 
Cook & Gilbert 778, 788, 1202. Cuzco, Hen era 305. Without 
locality: Humboldt & Bonpland; Lechler 1803; Savatier; Gay 2285. 
Introduced from Europe and widely distributed in the New World. 
"Quisa," "cohuiquisa," "ortiga menor." 

2. PILEA Lindl. 

Reference: Killip, Contr. U. S. Nat. Herb. 26: 367-394. 1936. 
Annual or perennial, monoecious or dioecious, usually succulent 
herbs, sometimes suffrutescent; leaves opposite, entire or toothed, 
trinerved or triplinerved, occasionally penninerved, those of a pair 
sometimes unequal and dissimilar; flowers in clusters, rarely solitary, 
forming dense heads, cymes, panicles, or umbels, which are andro- 
gynous or unisexual; staminate flowers 4 (rarely 2-3)-parted, the 
pistillate normally 3-parted, the segments unequal, the stigma short- 
penicillate, sessile; achenes compressed. 
Leaves all entire, usually less than 1 cm. long. 
Leaves faintly 1-nerved. 
Cymes sessile or subsessile; leaves, at least the larger one of a 

pair, obovate or oblong, cuneate. 

Cystoliths not elevated, very faint on under surface of the 

leaves; plant suberect, up to 30 cm. high . . P. microphylla. 

Cystoliths strongly elevated on both surfaces of the leaves; 

plant lax, the branches 50 cm. or more long . . P. foliosa. 

Cymes pedunculate; leaves nearly globular, appearing orbicular 

when dry P. serpyllacea. 

Leaves trinerved. 

Staminate flowers borne singly or in pairs; leaves cordiform; 
plant prostrate, much branched P. nerteroides. 



FLORA OF PERU 335 

Staminate flowers in sessile, androgynous heads; plant erect, 
unbranched P. nitida. 

Leaves (at least the larger one of a pair) toothed, sometimes sub- 
entire in P. marginata. 

A. Leaves of a pair dissimilar or very unequal, the larger more 
than twice as long as the smaller. 

Plants with densely pubescent branches and petioles; stipules 
more than 2 mm. long, subpersistent. 

Leaves less than 7 mm. long, the cystoliths conspicuous. 

P. pulegifolia. 

Leaves 1-3 cm. long, the cystoliths faint beneath. 

P. ramosissima. 
Plants glabrous throughout. 

Foliar cystoliths none P. Weberbaueri. 

Foliar cystoliths present. 
Larger leaves at a node not more than 4 mm. wide. 

P. diversifolia. 

Larger leaves at a node more than 4 mm. wide. 

Leaves appearing to be in fours at the nodes, owing to 
the presence of a short branchlet bearing a pair of 
leaves P. nutans. 

Leaves otherwise. 

Smaller leaf of a pair oblique, subentire; pistillate cymes 

sessile. 

Larger leaves very oblique, more than 7 cm. long, 
fully 7 times longer than the smaller ones. 

P. Bassleriana. 

Larger leaves symmetrical or nearly so, less than 
7 cm. long, and proportionately shorter. 

P. imparifolia. 

Smaller leaf symmetrical, toothed; pistillate cymes 

peduncled. 
Cystoliths all punctiform P. pichisana. 

Cystoliths linear or fusiform, occasionally puncti- 
form ones intermingled. 

Achenes longitudinally costate; larger leaves not 
more than 3.5 cm. long P. costata. 



336 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Achenes, as far as known, ecostate; larger leaves 

more than 3.5 cm. long. 

Larger leaves more than 3.5 cm. wide, more 
than 5 times longer than the smaller ones. 

P. Haenkei. 

Larger leaves not more than 3.5 cm. wide, less 

than 5 times longer than the smaller ones. 

P. macrocystolithica. 

A. Leaves of a node similar and equal or, if unequal, the larger 

one not more than twice as long as the smaller. 
B. Leaves averaging not more than 2 cm. long, usually massed 
at the end of the stem or branches; plants slender, usually 
prostrate or repent, the branches scarcely 10 cm. long. 
Cymes androgynous; leaves coarsely crenate-dentate; stem 

unbranched P. lamioides. 

Cymes usually unisexual; leaves serrulate or crenate-ser- 
rulate; stem usually branched. 

Pistillate cymes sessile P. Dombeyana. 

Pistillate cymes peduncled. 

Stem long-repent and rooting at most of the nodes, the 
leaves rather evenly spaced out . . P. nummularifolia. 
Stem repent only in the lower part, the leaves massed at 
the end of the stem and branches. 

Stem densely pubescent P. strigosa. 

Stem glabrous. 

Cystoliths on lower surface of leaves linear or fusi- 
form. 

Leaves broadly ovate or suborbicular, toothed 
nearly to the base, the cystoliths inconspicu- 
ous P. dauciodora. 

Leaves spatulate, rhombic, or suborbicular, 
toothed only in the upper half, the 

cystoliths conspicuous P. Fendleri. 

Cystoliths on lower surface of leaves punctiform. 
Achenes less than 1 mm. wide; leaves suborbicular 
or subreniform ; plants monoecious . . P. pusilla. 
Achenes more than 1 mm. wide; leaves ovate- 
lanceolate; plants apparently dioecious. 

P. delicatula. 



FLORA OF PERU 337 

B. Leaves averaging more than 2 cm. long; plants erect, more 

than 10 cm. high, usually leafy throughout. 
C. Plants glabrous throughout. 

Leaves coarsely and sharply serrate-dentate; inflorescence 
androgynous, sessile, much shorter than the petioles; 
plants annual, with a simple stem P. hyalina. 

Leaves not coarsely and sharply serrate-dentate; inflores- 
cence unisexual, pedunculate. 

Staminate flowers in dense, globose heads, these solitary 
and long-peduncled or forming a once or twice 
branched cyme. 
Plants dioecious; pistillate inflorescences diffuse, longer 

than the petioles P. minutiflora. 

Plants monoecious; pistillate inflorescences compact, 

short-peduncled P. Macbridei. 

Staminate flowers in cymes or panicles. 
Cystoliths stellate, 3-rayed; leaves subentire or denti- 
culate only toward the apex P. marginata. 

Cystoliths simple; leaves definitely toothed. 
Stem verrucose-roughened, woody; leaves averaging 

less than 3 cm. long P. verrucosa. 

Stem smooth, usually herbaceous; leaves longer. 
Leaves sessile or subsessile, sometimes clasping. 

P. subamplexicaulis. 

Leaves distinctly petioled, never clasping. 
Leaves ovate, triplinerved well above the base; 
inflorescences usually shorter than the 

petioles P. punctata. 

Leaves trinerved or subtriplinerved near the 

base; inflorescences longer than the petioles. 

Cystoliths of upper surface of leaves more 

than 0.5 mm. long, coarse, all fusiform; 

leaves thick-carnose P. suffruticosa. 

Cystoliths of upper surface of leaves shorter 

and finer, linear, fusiform, or punctiform; 

leaves usually of a thinner texture. 

Achenes minute, about 0.5 mm. long; 

leaves cordulate . . .P. citriodora. 



338 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Achenes 1-1.5 mm. long; leaves narrowed 

at the base. 
Stipules 3-10 mm. long, persistent. 

P. Poeppigiana. 
Stipules less than 3 mm. long, soon 

deciduous P. multiflora. 

C. Plants pubescent throughout, or at least with some 

indument on either the stem, leaves, or inflorescence. 
Leaves of nearly equal size at all the nodes, the internodes 
subequal, the inflorescence not confined to the upper 
axils. 

Plant annual, slender, with a pellucid stem; cymes 
androgynous; leaves sharply serrate-dentate. 

P. hyalina. 

Plant perennial, with a coarse stem. 
Stem succulent, essentially glabrous ... P. cushiensis. 
Stem ligneous or subligneous, densely rufo-hirsute. 
Leaves short-acuminate; plants monoecious, the 
staminate flowers in a dense, globose head. 

P. Pavonii. 
Leaves caudate-acuminate; plants dioecious, the 

staminate flowers in panicles P. hirsuta. 

Leaves massed at the end of the stem and branches, or a 
few, much smaller, at the lower nodes; inflorescence 
mainly in the upper axils. 

Staminate inflorescence borne at the rooting, leafless 
nodes, the perianth lobes about 2 mm. long. 

P. submissa. 

Staminate inflorescence borne with the pistillate at the 
upper nodes, or the flowers mixed with the pistillate 
ones. 

Cystoliths on upper surface of leaves all linear or fusi- 
form, not elevated P. pubescens. 

Cystoliths on upper surface of leaves punctiform or a 
few fusiform, elevated. 

Leaves acuminate, densely hirsute above. 

P. ceratocalyx. 

Leaves obtuse or subacute, glabrous above. 

P. Spruceana. 



FLORA OF PERU 339 

Pilea Bassleriana Killip, Contr. U. S. Nat. Herb. 26: 381. 1936. 

A coarse, succulent, glabrous herb, up to 50 cm. high, terrestrial 
or on tree trunks; leaves of a node very unequal and dissimilar, the 
larger elliptic, ovate-elliptic, or elliptic-lanceolate, 8-15 cm. long, 
2.5-7 cm. wide, caudate-acuminate, subsessile or short-petioled, 
coarsely crenate-serrate, the smaller ones orbicular-reniform, 5-15 
mm. long, strongly oblique, sessile; plants monoecious or dioecious; 
staminate cymes borne along the naked rooting portion of the stem, 
pedunculate; pistillate cymes sessile in the axils of the leaves. 

Loreto: Balsapuerto, 150-300 meters, Killip & Smith 28429, 
28467, 28471 (type); King 2870. Yurimaguas, Killip & Smith 
28993. Pongo de Manseriche, Killip, Smith & Dennis, 29145; 
Mexia 6355, 6360; Tessmann 4603. Mouth of Rio Pastaza (Killip, 
Smith & Dennis, 29196). 

Pilea ceratocalyx Wedd. in DC. Prodr. 16, pt. 1: 148. 1869. 

A low herb, densely grayish-hirsute throughout; leaves oblong 
or elliptic-ovate, 3-7 cm. long, 1-3 cm. wide, acuminate, narrowed 
at the base, sharply serrate, the cystoliths fusiform and punctiform; 
plants monoecious, the cymes apparently unisexual, the staminate 
compact, subsessile, the pistillate pedunculate, much longer than the 
adjacent petiole. 

Huanuco: Cochero, Poeppig 1032. Without locality: Poeppig 
3046 (type). These specimens are variously labeled "Peru" and 
"Brazil" in herbaria. The general relationship is with the Peruvian 
species. 

Pilea citriodora Wedd. Ann. Sci. Nat. III. 18: 216. 1852. 
Urtica limoniodora Pavon ex Wedd. loc. cit., as synonym. P. tar- 
mensis Killip, Journ. Wash. Acad. Sci. 15: 51. 1925. 

An erect, glabrous herb up to 60 cm. high, the stem un- 
branched; leaves ovate-lanceolate, 6-14 cm. long, 2.5-6 cm. wide, 
obtuse or short-acuminate, cordulate, crenate-serrate, the cystoliths 
fusiform, linear, and punctiform; plants dioecious, both the staminate 
and pistillate flowers borne in diffuse, long-peduncled panicles. 

Huanuco: Casapi, Poeppig 1260. Junin: Huacapistana, Province 
of Tarma, 2,800 meters, 5822 (type of P. tarmensis); Weberbauer 
1776a. Chanchamayo, Raimondi 2398. Without locality: Ruiz 
& Pavon (type). 

Pilea costata Killip, Contr. U. S. Nat. Herb. 26: 382. 1936. 
A succulent, glabrous herb about 50 cm. high, branched toward 
the apex; leaves of a node unequal and dissimilar, crenate-serrate 



340 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nearly to the base, the cystoliths linear, the larger leaf lance-elliptic, 
2.5-3.5 cm. long, 1.2-1.5 cm. wide, acuminate, the smaller one ovate, 
1.5-2 cm. long, obtuse or subacute; plants dioecious, the pistillate 
cymes 3-4 mm. wide, sessile or nearly so; achenes longitudinally 
costate. 

Junin: Enenas, Pichis Trail, 1,700 meters, Killip & Smith 
25638 (type). 

Pilea cushiensis Killip, sp. nov. 

Herba dioica, caule foliato, infra glabro, ad apicem sparse rufo- 
hirsutulo; folia elliptico-ovata, crenato-serrata, triplinervia, supra 
glabra, subtus in costis sparse rufo-hirsutula, cystolithis linearibus, 
creberrimis; inflorescentiae paniculiformes, in axillis fere omnibus. 

A succulent, perennial herb 1-1.5 meters high; stem sparsely 
rufo-hirsutulous toward the apex, otherwise glabrous; stipules ovate, 
about 3 mm. long, obtuse, soon deciduous; leaves of a node similar 
and subequal, elliptic-ovate, 7-15 cm. long, 3.5-6 cm. wide, acumi- 
nate, nan-owed to a cordulate base, coarsely crenate-serrate, tri- 
plinerved (lateral nerves extending to the upper quarter of the blade), 
the cystoliths linear, dense, faint beneath, the petioles 2-12 cm. long, 
those at a node often very unequal; plants dioecious, the staminate 
inflorescences borne in most of the axils, paniculiform, about 3 cm. 
long (not fully developed), rufo-hirsutulous, the perianth segments 
suborbicular. 

Huanuco: Cuschi, about 1,500 meters, June 19-23, 1923, on forest 
floor, Macbride 4826 (type, Field Mus. No. 535,863). 

In the key to the Andean species of Pilea (Contr. U. S. Nat. Herb. 
26: 368-376. 1936) this species would be placed at the very end, being 
differentiated from the Colombian P. Castronis by the shape of the 
leaves and the cystolithic marking, and by having a more diffuse 
inflorescence. 

Pilea dauciodora (R. & P.) Wedd. Ann. Sci. Nat. III. 18: 223. 
1852. Urtica dauciodora R. & P. ex Wedd. loc. cit., as synonym. P. 
uncidens Wedd. op. cit. 224. P. dauciodora var. uncidens Wedd. in 
DC. Prodr. 16, pt. 1: 138. 1869. 

A slender plant, often forming mats, glabrous throughout; leaves 
broadly ovate or orbicular-ovate, averaging 1.5 cm. long and 1 cm. 
wide, crenate-serrate, densely covered with linear and fusiform 
cystoliths; plants monoecious or dioecious, the clusters unisexual, 
though often both kinds borne at the same axils, rarely androgynous, 
slender-peduncled . 



FLORA OF PERU 341 

Huanuco: Muna, 2,000 meters, 4117. Junin: Huacapistana, 
1,800-2,700 meters, Kittip & Smith 24160, 24408. Acobamba, Rai- 
mondi 2800. Ayacucho: Carrapa, 1,200 meters, Killip & Smith 
22439. Puno: Sandia, Weberbauer 753. Without locality : "Andes," 
Ruiz & Pavdn (type) ; Spruce. Also in Central America, Colombia, 
Venezuela, and Bolivia. 

Pilea delicatula Killip, Journ. Wash. Acad. Sci. 15: 51. 1925. 

A slender, glabrous herb, up to 10 cm. high; leaves ovate-lanceo- 
late, 1-1.5 cm. long, acute, sharply mucronate-serrate, the cystoliths 
of the upper surface linear, faint, those of the lower surface puncti- 
form; plants apparently dioecious; pistillate flowers in small, globose, 
short-peduncled cymes. 

Huanuco: Tambo de Vaca, 4,000 meters, 4400 (type). 

Pilea diversifolia Wedd. Ann. Sci. Nat. III. 18: 212. 1852. 

A lax, much branched, glabrous herb, the stem up to 1.5 meters 
long; larger leaves of a node narrowly cuneate-oblong, 5-10 mm. 
long, 1-4 mm. wide, coarsely 3-7-toothed, 1-nerved, the smaller 
leaves suborbicular, 3-5 mm. wide, entire or few-toothed, faintly 
triplinerved; cystoliths linear; plants dioecious; pistillate flowers in 
short-peduncled cymes. 

Huanuco: Muna, 2,100 meters, 3984- Junin: Dos de Mayo, 
Pichis Trail, 1,800 meters, Killip & Smith 25885. Ayacucho: 
Carrapa, 1,500 meters, Kittip & Smith 22357. Without locality: 
Ruiz & Pavon (type). 

Pilea Dombeyana Wedd. Ann. Sci. Nat. III. 18: 221. 1852. P. 
orbiculata Killip, Journ. Wash. Acad. Sci. 15: 53. 1925. 

A low, succulent herb; stipules broadly ovate, persistent; leaves 
borne mostly near the ends of the branches, nearly orbicular, 7-25 
mm. wide, short-petiolate, the cystoliths faint, fusiform; plants 
monoecious or dioecious, the inflorescences unisexual, the staminate 
flowers sessile in globose clusters in a few-branched panicle, the 
pistillate in small, densely flowered, sessile cymes. 

Huanuco: Chasqui, 3289 (type of P. orbiculata). Dept. uncer- 
tain: Between Callcan and Pelechuco, 3,600 meters, Pearce in 1864; 
"Andes," Dombey (type). Without locality: MacLean; "Herb. 
Hooker 2031." 

Pilea Fendleri Killip, nom. nov. Urtica dichroa Poepp. ex Wedd. 
in DC. Prodr. 16, pt. 1: 61. 1869, as synonym. P. dauciodora var. 



342 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

crenata Wedd. op. cit. 139, not P. crenata Britt. & Wils. P. daucio- 
dora var. pilosula Wedd. loc. cit. P. leptophylla Killip, Contr. U. S. 
Nat. Herb. 26: 387. 1936, not Urban. 

A very slender plant, with the stem repent toward the base, at 
length ascending and few-several-branched ; leaves rotund-spatulate, 
rhombic-orbicular or nearly orbicular, 5-20 mm. long, subequal at a 
node, cuneate, crenate or crenate-serrate above the middle, thin, 
glabrous or sparsely pilosulous with hyaline hairs, the cystoliths 
linear or fusiform, conspicuous; plants monoecious or dioecious, the 
heads unisexual or androgynous, peduncled in the upper axils. 

Huanuco: Cochero, Poeppig 1552. Without locality: Poeppig 
1383. Also in Venezuela and Colombia. 

In a list in the Prodromus of species excluded from Urtica Weddell 
gives U. dichroa Poepp., an herbarium name apparently, as equaling 
Pilea nummularifolia. He does not, however, cite the name in the 
synonymy of the latter. Poeppig's No. 1552, in the Vienna herbarium, 
bears this name in Poeppig's handwriting, though the specimen is 
clearly not P. nummularifolia. 

Pilea foliosa Killip, Contr. U. S. Nat. Herb. 26: 377. 1936. 

A glabrous, succulent herb with lax branches 50-150 cm. long; 
leaves of a node unequal and dissimilar, the longer obovate, 5-10 
mm. long, entire or obscurely undulate, the smaller subreniform, 
1.5-3 mm. wide, entire; cystoliths linear, strongly elevated; plants 
monoecious, the staminate flowers usually solitary, the pistillate 
solitary or in small, subsessile clusters. 

Junin: Carpapata, 3,000 meters, Killip & Smith 24400 (type). 
Huacapistana, 1,900 meters, Killip & Smith 24281. Chanchamayo 
Valley, Schunke 498, 678, 992. 

Pilea Haenkei Killip, Contr. U. S. Nat. Herb. 26: 384. 1936. 

A glabrous herb 25 cm. high or more; leaves of a node similar but 
very unequal, ovate-lanceolate, acuminate, cordulate, crenate- 
serrate to the base, triplinerved, densely covered with fusiform and 
punctiform cystoliths above and fusiform ones beneath, the larger 
leaves 9-13 cm. long, 3.5-4 cm. wide, petiolate, the smaller ones 
1.5-2 cm. long, subsessile; plants dioecious, the staminate cymes 
sessile or short-peduncled, densely flowered. 

Peru, collected in the montana, but the precise locality unknown, 
Haenke 1870 (type). 



FLORA OF PERU 343 

Pilea hirsuta (Pavon) Wedd. Ann. Sci. Nat. III. 18: 220. 1852. 
Urtica hirsuta Pavon ex Wedd. loc. cit., as synonym. 

An erect, robust herb, the stem ferruginous- villous; leaves ovate- 
elliptic, 7-15 cm. long, 3-7 cm. wide, coarsely serrate, rugulose, fer- 
ruginous-hirsute on the nerves and veins, the cystoliths fusiform; 
plants dioecious, the staminate and pistillate inflorescences similar, 
paniculate, slightly longer than the adjacent petiole. 

Peru, without locality, Ruiz (type). 

Pilea hyalina Fenzl, Denkschr. Akad. Wiss. Math. Naturw. 
(Wien) 1: 256. 1850. Urtica arvensis Poepp. ex Fenzl, loc. cit., as 
synonym. 

A slender, erect annual, the stem unbranched, pellucid, glabrous; 
leaves rhombic-elliptic or ovate, 1-6 cm. long, 0.8-4.5 cm. wide, 
acute, cuneate, coarsely serrate-dentate, thin-membranous, glabrous, 
or sparsely hyaline-strigillose above, the cystoliths linear, faint; 
plants monoecious, the inflorescences androgynous, cymose-pani- 
culate, much shorter than the petioles. 

Amazonas: Moyobamba, Mathews 1555, San Martin: San 
Roque, 1,400 meters, Williams 7149. Huanuco: Cochero, Poeppig 
1539. Jum'n: La Merced, 800 meters, Killip & Smith 23593. San 
Ramon, 1,600 meters, Killip & Smith 24676. Yapas, Pichis Trail, 
1,400 meters, Killip & Smith 25580. Ayacucho: Aina, 750-1,000 
meters, Killip & Smith 22746. Without locality: Mathews 2032; 
Gay; Ruiz & Pavon. Widely distributed in the American tropics, 
from Mexico to Chile, Brazil, and Argentina, at elevations up to 
1,500 meters. 

Pilea imparifolia W T edd. Ann. Sci. Nat. III. 18: 212. 1852. P. 
dendrophila Miq. in Mart. Fl. Bras. 4, pt. 1 : 202. 1853. 

A decumbent, glabrous herb with numerous suberect branches; 
larger leaf rhombic-ovate to elliptic-oblong, rarely obovate, 2-6 
cm. long, 0.8-2 cm. wide, crenate-serrate above the middle, oblique, 
sessile or short-^etioled, the smaller leaf obovate-orbicular or 
orbicular-reniform, 0.8-1.5 cm. long, very asymmetrical, subentire; 
plants dioecious, the cymes sessile or subsessile, few-flowered. 

San Martin: Tarapoto, Spruce 4434- Loreto: Rio Itaya, Killip 
& Smith 29308, 29515, 29572. Mouth of Rio Napo, Tessmann 3721. 
Mouth of Rio Santiago, Mexia 6365. Maucallacta, Klug 3950. 
Between Rio Ucayali and Rio Huallaga, Huber 1519. Junin: San 
Nicolas, Pichis Trail, Killip & Smith 26038. Colombia to northern 



344 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Peru, eastward to the Guianas and Amazonian Brazil; at low 
elevations. 

Pilea lamioides Wedd. Ann. Sci. Nat. III. 18: 213. 1852. 

An unbranched herb up to 12 cm. high; leaves ovate, 1-2 cm. 
long, 6-12 mm. wide, coarsely crenate-dentate, obtuse; plants 
monoecious, the cymes androgynous, borne in the upper axils, the 
staminate flowers short-pediceled in sessile clusters, the pistillate 
sessile in short-peduncled clusters. 

Lima: San Geronimo, 150 meters, 5910. Atocongo, Pennell 
14751. Amancais, Weberbauer 1594- Without locality: Dombey 
(type) ; Pavon. 

Pilea Macbridei Killip, Journ. Wash. Acad. Sci. 15: 52. 1925. 

A glabrous plant about 1 meter high, the stem slightly woody, 
much branched; leaves ovate or orbicular-ovate, 3-5 cm. long, 1.5- 
2.5 cm. wide, acuminate, cordulate, serrate, the cystoliths linear 
above, punctiform beneath; plants monoecious, the staminate flowers 
in a subglobose, slender-peduncled head or in glomerules forming a 
once branched panicle, the pistillate in short-peduncled cymes. 

Huanuco: Rio Chinchao, 2,800 meters, 5179 (type). Junin: 
Carpapata, 3,000 meters, Killip & Smith 24457. 

Pilea macrocystolithica Killip, Contr. U. S. Nat. Herb. 26: 
384. 1936. 

A glabrous, unbranched herb about 30 cm. high; leaves of a node 
dissimilar and unequal, the larger one ovate or ovate-lanceolate, 
4-6 cm. long, 2-3 cm. wide, obtuse or obtusely acuminate, short- 
petiolate, serrulate, the smaller one suborbicular, 1.5-2 cm. wide, 
subsessile, the cystoliths of the upper surface very numerous, linear, 
elevated, 0.7-0.8 mm. long, those of the lower surface shorter and 
obscure; plants apparently dioecious, the staminate cymes sub- 
globose, on very slender peduncles about 2 cm. long. 

Peru, collected in the montana, the precise locality unknown, 
Haenke 1860 (type). 

In the original account of this species a line in the description of 
the cystoliths was accidentally omitted, so that the reason for the 
selection of the specific name was not at all apparent. 

Pilea marginata (Poepp.) Wedd. Arch. Mus. Paris 9: 238. 
1856-57. Urtica marginata Poepp. ex Wedd. loc. cit., as synonym. 

An erect, glabrous herb 80 cm. or more high; leaves oblong or 
elliptic-oblong, 10-20 cm. long, 4-8 cm. wide, attenuate-acuminate, 



FLORA OF PERU 345 

undulate, subentire or denticulate toward the apex, coriaceous, the 
cystoliths stellate, 3-rayed; plants apparently dioecious, the stami- 
nate flowers in small glomerules in a short, narrow panicle. 

San Martin: Tarapoto, Ule 6508. Huanuco: Cochero, Poeppig 
3045. Pampayacu, 1,100 meters, 5086. Junin: San Nicolas, Pichis 
Trail, 1,100 meters, Killip & Smith 26032. Dept. uncertain: Rio 
Amazonas, Poeppig 1088 (type), 2088. 

Pilea microphylla (L.) Liebm. Dansk. Vid. Selsk. Skrift. V. 2: 
296. 1851. Parietaria microphylla L. Syst. ed. 10. 1308. 1759. Pilea 
muscosa Lindl. Coll. Bot. pi. 4- 1821. 

A glabrous, succulent herb, 4-30 cm. high, variable in size and 
habit; leaves usually crowded, thick, short-petioled, obtuse or sub- 
acute, entire, those of a pair unequal, the larger usually obovate, up 
to 1 cm. long, the smaller orbicular or obovate-orbicular, up to 3 
mm. long, the cystoliths linear, transverse across the blade; plants 
monoecious, rarely dioecious, the flower clusters androgynous or 
unisexual, sessile or subsessile. 

Amazonas: Chachapoyas, Mathews. San Martin: Tarapoto, 
Williams 6071; Spruce 4028; Ule 6657. San Roque, Williams 7233. 
Rio Huallaga, Williams 4082, 6662. Zepelacio, King 3719,Loreto: 
Iquitos, Killip & Smith 27500. Puerto Mele*ndez, Tessmann 4742. 
Huanuco: Piedra Grande, 1,500 meters, 3690. Cochero, Poeppig in 
1830. Junin: La Merced, 700 meters, Killip & Smith 24073. Rio 
Paucartambo, Killip & Smith 25329. Without locality: Ruiz & 
Pawn. Throughout the American tropics. Often cultivated as a 
border plant or in pots. 

Pilea minutiflora Krause, Bot. Jahrb. 37: 529. 1906. 

A glabrous herb 30-40 cm. high; leaves ovate-lanceolate or ovate- 
elliptic, 3-8 cm. long, 1.5-3 cm. wide, acuminate, rounded or cordu- 
late at the base, crenate-serrate, the cystoliths few and faint; plants 
usually dioecious, the flowers in small clusters, the staminate ones 
forming a once branched panicle, the pistillate forming subdichoto- 
mcus cymes up to 7 cm. long. 

Junin: Huacapistana, 1,900-2,000 meters, Weberbauer 2027 (type). 
Huariaca, 2,900 meters, 3126. Puno: Sandia, Weberbauer 575. 

The Indians are said to use this as a remedy for unrequited love. 

Pilea multiflora (Poir.) Wedd. Ann. Sci. Nat. III. 18: 218. 
1852, in part. Urtica multiflora Poir. in Lam. Encycl. Suppl. 4: 
223. 1816. Pilea anomala Wedd. Ann. Sci. Nat. III. 18: 217. 1852. 



346 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

An erect, glabrous herb up to 1 meter high, the stem simple or 
few-branched; leaves oblong-lanceolate to ovate-lanceolate, 4-15 
cm. long, 1.5-4 cm. wide, suboblique, long-acuminate, sharply ser- 
rulate, the cystoliths linear and punctiform, faint; plants monoecious 
or dioecious, the inflorescences unisexual, both kinds cymose-pani- 
culate, diffuse. 

Huanuco: Pampayacu, 2,100 meters, Weberbauer 6812. Junin: 
Pichis Trail, about 1,700 meters, Kittip & Smith, 25775, 25801, 
25913, 25929. Puno: Sandia, Weberbauer 6521. Cuzco: Cerro de 
Cusilluyoc, Pennell 14013. Without locality: Jussieu (type). Also 
in Bolivia. 

Pilea nerteroides Killip, Contr. U. S. Nat. Herb. 26: 377. 1936. 
P. cordifolia Killip, Journ. Wash. Acad. Sci. 15: 50. 1925, not Benth. 

A slender, prostrate herb, leafy throughout; stem villosulous; 
stipules ovate, persistent; leaves cordate, up to 4 mm. long and 5 mm. 
wide, villosulous, covered with punctiform cystoliths; staminate 
flowers borne singly or in pairs, the peduncles 3-6 mm. long; pistil- 
late flowers in 4-6-flowered, pedunculate umbels. 

Huanuco: Tambo de Vaca, 4,000 meters, 4395 (type). 

Pilea nitida Wedd. Ann. Sci. Nat. III. 18: 211. 1852. 

A very slender, glabrous plant, up to 10 cm. high; leaves elliptic- 
ovate, obtuse at both ends, 4-15 mm. long, 2.5-8 mm. wide, entire 
or rarely subcrenulate, obscurely 3-nerved, bearing linear cystoliths 
on the upper surface; plants monoecious, the flowers in compact, 
androgynous heads shorter than the petioles. 

Cajamarca: San Pablo, 2,300 meters, Weberbauer 3872. Lima: 
San Agustin, Weberbauer 5244- Without locality: Dombey (or Ruiz 
& Pawn[1}; type). 

Pilea nummularifolia (Sw.) Wedd. Ann. Sci. Nat. III. 18: 
225. 1852. Urtica nummularifolia Sw. Act. Holm. 1787: 63. pi. 1, 
f. 2. 1787. 

A repent or trailing plant, with a slender, villosulous or glabres- 
cent stem; leaves orbicular, 5-12 mm. wide, crenate, strigillose with 
stiff, hyaline hairs; plants monoecious or dioecious, the inflorescences 
apparently unisexual, sessile, the staminate subumbellate, the pistil- 
late cymose. 

Loreto: Pebas, on the Rio Amazonas, Williams 1917. Also in 
Venezuela and the West Indies. 



FLORA OF PERU 347 

Pilea nutans (Poepp.) Wedd. Arch. Mus. Paris 9: 196. pi. 7, 
f. 11-13. 1856-57. Urtica nutans Poepp. ex Wedd. loc. cit., as 
synonym. Pilea Herrerae Mildbr. ex Herrera, Anal. Univ. Cuzco 1 : 
147. 1926, name only. 

A diffuse, much branched, glabrous herb; leaves appearing in 
4's (rarely in 3's) due to the presence at each node of 1 or 2 borne on 
very short branchlets, serrate or crenate-serrate, triplinerved, the 
cystoliths linear, the larger leaves of a node rhombic-ovate or 
rhombic-lanceolate, 8-25 mm. long, 4-15 mm. wide, the smaller ones 
rhombic or broadly ovate, up to 5 mm. long; plants dioecious, the 
staminate flowers in small, 8-12-flowered, globose heads. 

Huanuco: Cochero, Poeppig 1565 (type). Junin: Huacapistana, 
1,800 meters, Kittip & Smith 24274- Dos de Mayo, Pichis Trail, 
Killip & Smith 25789. Porvenir, Pichis Trail, Kittip & Smith 
25895. Pasla, Raimondi 9290. Cuzco: Rio Yanamayo, Pennell 
14049. Urubamba Valley, Cook & Gilbert 1095. Cosnipata Valley, 
Herrera 6. Puno: Ollachca, Raimondi 9649. Without locality: 
Dombey. Also in Bolivia. "Chia-chia," "quisa," "yuncaquisa." 

Pilea Pavonii Wedd. Ann. Sci. Nat. III. 18: 219. 1852. Urtica 
cymosa Pavon ex Wedd. loc. cit., as synonym. 

An erect herb, 40-60 cm. high, with a stout, quadrangular, rufo- 
villosulous stem; leaves broadly ovate, 6-10 cm. long, 4-6 cm. wide, 
short-acuminate, crenate-serrate, subcoriaceous, appressed-pilosu- 
lous above, densely ferruginous-hirsute beneath, the cystoliths fusi- 
form and punctiform beneath; plants monoecious, the staminate 
inflorescences borne in the upper axils on stout peduncles, globose, 
compact, the pistillate sessile or subsessile in the middle axils, cymose- 
paniculate, much shorter than the adjacent petiole. 

Huanuco: Muna, 2,000 meters, -4075. Without locality: Pavon 
(type). 

Pilea pichisana Killip, Contr. U. S. Nat. Herb. 26: 383. 1936. 

A slender, glabrous, subrepent herb, with lax branches up to 20 
cm. long; leaves of a node unequal but similar, ovate-lanceolate, 
acute or acuminate, petiolate, crenate-serrate nearly to the base, the 
cystoliths all punctiform, the larger leaf 2-2.8 cm. long, the smaller 
one 0.5-1.2 cm. long; plants dioecious, the pistillate flowers in sub- 
sessile cymes. 

Junin: Dos de Mayo, Pichis Trail, 1,800 meters, Kittip & Smith 
25876 (type). 



348 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pilea Poeppigiana Wedd. Ann. Sci. Nat. III. 18: 225. 1852. 

A glabrous plant, with the stem repent, at length erect, 30-50 
cm. high; stipules broadly ovate, persistent; leaves ovate-lanceolate, 
rarely oblanceolate, up to 15 cm. long and 7 cm. wide, acuminate, 
cuneate at the base, coarsely crenate-serrate, the petioles up to 7 
cm. long, the cystoliths fusiform and punctiform; plants dioecious, 
the pistillate inflorescence cymose-paniculate, borne in the upper 
axils, long-peduncled. 

San Martin: Tarapoto, Ule 6844. Huanuco: Yanano, 1,800 
meters, 3770. Cochero, Poeppig 1032, 1539B.Pichis Trail, 1,800 
meters, Killip & Smith 25826, 25902. Department uncertain: 
"Subandean region," Poeppig (type). Also in the Amazon basin of 
Colombia and Ecuador. 

Pilea pubescens Liebm. Dansk. Vid. Selsk. Skrift. V. 2: 302. 
1851. 

Stem repent, at length erect and usually with several erect or 
ascending branches, strigillose; leaves massed at the end of the stem 
or branches, wanting or much reduced below, broadly ovate or 
elliptic-ovate, up to 7 cm. long and 6 cm. wide, crenate-serrate, 
sparingly strigillose above with hyaline hairs, or nearly glabrous, 
hirsutulous beneath on the nerves and veins, the cystoliths linear 
and fusiform above; plants monoecious or dioecious, the staminate 
flowers in a sessile cluster at the base of the cymose-paniculate, 
peduncled pistillate inflorescences, or scattered among the pistillate 
flowers. 

San Martin: San Roque, 1,350-1,500 meters, Williams 6957, 
7695. Tarapoto, Spruce 4455. Widely distributed in the American 
tropics. 

Pilea pulegifolia (Poir.) Wedd. Ann. Sci. Nat. III. 18: 218. 
1852. Urtica pulegifolia Poir. in Lam. Encycl. Suppl. 4: 224. 1816. 

An herb with an elongate stem and numerous densely tomen- 
tellous, leafy branches; leaves appearing in 4's due to the presence 
at each node of a pair borne on a very short branchlet, ovate, 3-6 
mm. long, 2-4 mm. wide (those of a node similar but slightly un- 
equal), obtuse, crenate-serrulate, the petioles tomentellous, the 
cystoliths linear, conspicuous; plants dioecious, the staminate flow- 
ers in small, pedunculate cymes in the upper axils. 

Peru: Churugallana, Dombey (type). 



FLORA OF PERU 349 

Pilea punctata (HBK.) Wedd. Ann. Sci. Nat. III. 18: 222. 
1852. Urtica punctata HBK. Nov. Gen. & Sp. 2: 38. 1817. 

An erect or ascending, glabrous herb, 20-40 cm. high; leaves 
ovate, 3-8 cm. long, 1.5-3 cm. wide, acuminate, rounded at the base, 
coarsely crenate-serrate, triplinerved well above the base, black- 
punctate beneath, the cystoliths fusiform; plants monoecious or 
dioecious, the inflorescences androgynous or unisexual, cymose, 
longer than the adjacent petioles. 

Cajamarca: Zaulaca, Humboldt & Bonpland (type). Amazonas: 
Chachapoyas, 1,600 meters, Weberbauer 1+307. Without locality: 
Ruiz & Pawn. 

Pilea pusilla Krause, Bot. Jahrb. 37: 530. 1906. 

A slender herb up to 10 cm. high, the stem filiform; leaves sub- 
orbicular or subreniform, up to 1 cm. long and 1.5 cm. wide, obtuse, 
subtruncate at the base, sharply serrulate, the cystoliths all puncti- 
form; plants monoecious, the flower clusters forming slender- 
peduncled panicles. 

Junin: Palca, 1,900-2,000 meters, Weberbauer 2023 (type). 

Pilea ramosissima Killip, Contr. U. S. Nat. Herb. 26: 380. 1936. 

A diffuse herb, the stem slightly woody, with numerous lateral, 
pubescent branches; leaves appearing in 4's, unequal and slightly 
dissimilar, crenate-serrate, essentially glabrous except for the pubes- 
cent petioles, the larger leaves rhombic-ovate, 1-3 cm. long, 7-15 
mm. wide, obtuse, the smaller ones orbicular or ovate-orbicular, 
4-5 mm. long, the cystoliths faint on the under surface; plants 
dioecious, the pistillate flowers in small, 10-flowered, compact cymes, 
their peduncles 1-1.3 cm. long. 

Huanuco: Chaglla, 2,800 meters, 3650 (type). 

Pilea serpyllacea (HBK.) Wedd. Ann. Sci. Nat. III. 18: 205. 
1852. Urtica serpyllacea HBK. Nov. Gen. & Sp. 2: 37. 1817. U. 
thymifolia HBK. loc. cit. Pilea globosa Wedd. Ann. Sci. Nat. III. 
18 : 208. 1852. P. thymifolia Blume, Mus. Bot. Lugd. Bat. 2 : 44. 1855. 

A glabrous, very succulent herb, red-tinged throughout; leaves 
nearly globular, 1-5 mm. in diameter, entire or shallowly crenate, 
transversely striate with linear cystoliths; plants monoecious, or the 
staminate flowers sometimes wholly wanting, the pistillate flowers 
in peduncled cymes. 

Cajamarca: Hualgayoc, Weberbauer 4054; Raimondi 2209. 
Callacate, Raimondi 5639. Huanuco: Huacachi, 2,000 meters, 



350 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3868, 4087. Huanuco, 2,200 meters, 3512; Pearce 118. Casapi, 
Poeppig 1381. Lima: Matucana, 2,500 meters, 447. Rio Blanco, 
3,200 meters, Killip & Smith 21601. Junin: Carpapata, 2,400 
meters, Killip & Smith 24334- Tambo de Viso, Weberbauer 158. 
Cuzco: Cuzco, Herrera 51; Weberbauer 4892. Urubamba Valley, 
1,800 meters, Cook & Gilbert 262, 1035; Herrera 2288. Apurimac 
Valley, Herrera 3065. Ollantaitambo, Pennell 1 3657. Without 
locality: Ruiz & Pavon (type) ; Jussieu. Western Venezuela to Peru 
and Bolivia, 2,000-3,500 meters altitude. "Accoicarpa," "quisa," 
"kkuru-quisa." 

Pilea Spruceana Wedd. in DC. Prodr. 16, pt. 1: 161. 1869. 

A low herb, the stem repent, at length erect, sparsely villous; 
leaves oblong or ovate-oblong, 2-8 cm. long, 1.5-4 cm. wide, obtuse 
or subacute, crenate-serrate, sparingly ciliate, strigillose above, 
villous beneath, the cystoliths punctiform and fusiform, the latter 
more numerous at the margin; plants monoecious or dioecious, the 
pistillate flowers in short-peduncled cymes, the staminate flowers 
subsessile at the base of the pistillate inflorescence. 

San Martin: Tarapoto, Spruce 4376 (type). Loreto: Cumbasa, 
Ule 6843. Huanuco: Posuso, Pearce 284- Also in Bolivia. 

Pilea strigosa Wedd. Ann. Sci. Nat. III. 18: 225. 1852. P. 
repens var. strigosa Wedd. in DC. Prodr. 16, pt. 1: 156. 1869. 

A low herb, the stem ferruginous-hirsute, repent at the base, with 
lax branches; leaves suborbicular to broadly ovate, 8-25 mm. long, 
rounded or subacute at the apex, crenate-serrate, strigose above, 
ferruginous-hirsutulous on the nerves beneath; plants usually 
monoecious, the cymes unisexual, pedunculate, the staminate sub- 
globose, the pistillate paniculiform. 

Loreto: Santa Rosa, 135 meters, Killip & Smith 28835, 28843. 
Huanuco: Cochero, Poeppig 1552 in part. Junin: Dos de Mayo, 
Pichis Trail, Killip & Smith 25795. Without locality: Mathews 
2031 (type). Also in Bolivia. 

Pilea subamplexicaulis Killip, Contr. U. S. Nat. Herb. 26: 
390. 1936. 

A glabrous plant 35 cm. high or more; leaves lanceolate or oblong- 
lanceolate, 7-20 cm. long, 2.5-6 cm. wide, acuminate or attenuate- 
acuminate, cordate or slightly clasping at the base, with numerous 
fusiform and fewer punctiform cystoliths; plants dioecious; stami- 



FLORA OF PERU 391 

nate flowers in loose cymes up to 2 cm. long, the pistillate in short, 
sessile cymes. 

San Martin: Tarapoto, 1,100 meters, Ule 6588 (type). San 
Roque, 1,350-1,500 meters, Williams 7425. 

Pilea submissa Wedd. in DC. Prodr. 16, pt. 1: 151. 1869. 

Plant terrestrial, or repent on tree trunks, the erect or ascending 
portion of the stem less than 10 cm. long, pubescent; leaves rhombic- 
elliptic, 4-9 cm. long, 1.5-3.5 cm. wide (extremes up to 15 cm. long 
and 5 cm. wide), narrowed at both ends, short-petioled, crenate- 
serrulate, glabrous above, hispidulous on the nerves and veins 
beneath, the cystoliths fusiform, faint; plants monoecious or dioe- 
cious, the cymes unisexual, the staminate borne at the rooting, 
leafless nodes, their peduncles 4-8 cm. long, the pistillate at the upper 
axils, their peduncles 2.5-3 cm. long. 

San Martin: Tarapoto, Spruce 4155 (type). Loreto: Pongo de 
Manseriche, 250 meters, Mexia 6359. Pumuyacu, King 3186. 
Junin: San Nicolas, Pichis Trail, 1,100 meters, Kittip & Smith 26023. 

Pilea suffruticosa Krause, Bot. Jahrb. 37: 529. 1906. 

An erect, glabrous, suffrutescent plant, about 2 meters high, the 
stem unbranched; leaves narrowly elliptic to ovate-elliptic, 5-8 
cm. long, 2-3 cm. wide, acute, serrulate, the cystoliths fusiform, 
prominent; plants apparently dioecious, the staminate inflorescences 
cymose-paniculate, much longer than the adjacent petiole. 

Amazonas: Chachapoyas, 2,500 meters, Weberbauer 4387 (type). 

Pilea verrucosa Killip, Journ. Wash. Acad. Sci. 15: 53. 1925. 

A few-branched, glabrous shrub, about 1 meter high, the stem 
verrucose-roughened; leaves ovate or subrhombic, 2-3.5 cm. long, 
1-2.5 cm. wide, acute, crenate-serrate, the cystoliths punctiform; 
plants monoecious, the inflorescences unisexual, the staminate 
flowers in much branched panicles in the upper axils, the pistillate in 
subsessile cymes in the lower axils. 

Huanuco: Rio Chinchao, 2,800 meters, 5201 (type). 

Pilea Weberbaueri Killip, Contr. U. S. Nat. Herb. 26: 380. 1936. 

A branched, glabrous herb, up to 30 cm. high; leaves opposite or 
sometimes appearing in 3's or 4's due to the presence at a node of a 
pair of leaves on very short secondary branches, without cystoliths, 
the larger leaves oblong-oblanceolate, 7-12 mm. long, 3-6 mm. 
wide, obtusely acuminate, petiolate, 3-5-toothed near the apex, the 



352 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

smaller ones orbicular-reniform, 3-5 mm. long, 4-6 mm. wide, 
sessile or subsessile; plants apparently dioecious, the pistillate 
flowers in dense clusters in a compact cyme, the peduncles filiform. 

Junin: Between Palca and Huacapistana, 1,900-2,000 meters, 
Weberbauer 2022 (type). 

Species doubtfully occurring in Peru 

Pilea angustata Killip, Contr. U. S. Nat. Herb. 26: 378. 1936. 
This is based upon a Grisar collection in the Paris Herbarium, said 
to have come from Ecuador or Peru. In view of other specimens in 
this collection, it is probable that most, if not all, came from Ecuador. 

3. BOEHMERIA Jacq. 

Trees, shrubs, or perennial herbs, unarmed; leaves opposite or 
alternate, toothed, 3-nerved, those of the adjacent nodes sometimes 
unequal and dissimilar; plants monoecious or dioecious, the flowers 
in globose, usually unisexual clusters in the leaf axils or forming a 
spike; staminate flowers 4 (rarely 3 or 5) -parted; pistillate flowers 
tubular, contracted at the throat, 2-4-toothed or entire, the stigma 
filiform; achene enclosed in the persistent perianth. 

Flower clusters forming long spikes. Leaves opposite . . .B. caudata. 
Flower clusters in the leaf axils. 

Leaves opposite B. Mathewsii. 

Leaves alternate. 

Under surface of leaves appressed-silvery-sericeous. .B. Pavonii. 
Under surface of leaves not sericeous. 

Leaves coriaceous, strongly bullate, the larger usually nar- 
rowly lanceolate B. aspera. 

Leaves membranous, flat or slightly rugulose, the larger 

ovate-lanceolate or oblong-lanceolate. 
Styles about 1 mm. long; leaves of the adjacent nodes 
similar and subequal J5. brevirostris. 

Styles 1.5-2 mm. long; leaves of the adjacent nodes very 
dissimilar and unequal. 

Leaves coarsely crenate-serrate, pale and pilosulous 
beneath B. fallax. 

Leaves closely serrulate, concolorous, appressed-hispid- 
ulous beneath . . . B. anomala. 



FLORA OF PERU 353 

Boehmeria anomala (Wedd.) Killip, sp. nov. B. Pavonii 
var. anomala Wedd. in DC. Prodr. 16, pt. 1: 199. 1869. 

A slender, open shrub, or a tree, about 4 meters high, the branches 
flexuose, appressed-hirsutulous; leaves alternate, those of the adja- 
cent nodes very unequal and dissimilar, the larger ovate-lanceolate 
or oblong-lanceolate, 6-15 cm. long, 1.5-6 cm. wide, attenuate- 
acuminate, often strongly oblique, closely serrulate, short-petioled, 
membranous, appressed-hispidulous on both surfaces, concolorous, 
the smaller leaves (rarely present in herbarium specimens) ovate or 
suborbicular, 3-5 mm. long, 3 mm. wide, sessile; plants apparently 
dioecious, the flower clusters about 5 mm. wide. 

San Martin: Tarapoto, Spruce 3962 (type). Junin: La Merced, 
600 meters, 5268. Ayacucho: Choimacota Valley, 1,400 meters, 
Weberbauer 7529. 

This is more closely related to B. fallax than to B. Pavonii. 

Boehmeria aspera Wedd. Arch. Mus. Paris 9: 349. pi. 11, f. 
24-28. 1856-57. B. diversifolia Wedd. Ann. Sci. Nat. IV. 1: 202. 
1854, not Miquel, 1851. B. Pavonii var. diversifolia Wedd. in DC. 
Prodr. 16, pt. 1: 199. 1869, in part. 

A shrub 1-3 meters high, densely leafy throughout, the branch- 
lets grayish- or blackish-hirsute; leaves alternate, those of the adja- 
cent nodes very unequal and somewhat dissimilar, the larger lanceo- 
late, usually narrowly so, 4-13 cm. long, 1-4 cm. wide, long-acuminate, 
crenate-serrate, short-petioled, coriaceous, strongly bullate, smooth 
or hispid above, densely cano-hirsute beneath, the smaller leaves 
ovate, 1-3 cm. long, 0.8-1.5 cm. wide, acute, subsessile; plants 
monoecious, the flower clusters up to 8 mm. wide, predominately 
pistillate, sometimes with a few staminate flowers intermingled. 

Huanuco: Piedra Grande, 1,500 meters, 3698. Muiia, Pearce 
120. Junin: Palca, Dombey (type of B. diversifolia Wedd.). Hua- 
capistana, 1,800-2,400 meters, Killip & Smith 24109, 24352; Weber- 
bauer 1994. Ayacucho: Carrapa, 1,500 meters, Killip & Smith 
22324- Without locality: Ruiz & Pavon. Also in Colombia. 

Boehmeria brevirostris Wedd. Ann. Sci. Nat. IV. 1: 201. 1854. 

A shrub about 3 meters high, the branchlets cano-hirsute; leaves 
of the adjacent nodes similar and subequal, ovate or broadly ovate- 
lanceolate, 3-15 cm. long, 2-7 cm. wide, attenuate-acuminate, 
usually cordate, coarsely crenate-serrate, membranous, flat, ap- 
pressed-hispid above, softly pubescent beneath; plants monoecious 



354 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

or dioecious, the flower clusters 4-6 mm. wide, unisexual or 
with a few staminate flowers in the pistillate heads; styles about 
1 mm. long. 

Junin: Rio Perene", near Colonia Perene", 600 meters, Killip & 
Smith 25168. Without locality: Mathews 2039 in part (type). 

Boehmeria caudata Sw. Prodr. 34. 1788. B. peruviana Blume, 
Mus. Bot. Lugd. Bat. 2: 224. 1856. 

A shrub or a small tree, up to 8 meters high; leaves opposite, 
large, broadly ovate to elliptic, up to 25 cm. long and 10 cm. wide, 
acute or acuminate, finely serrate to crenate-serrate, membranous, 
flat or rugulose, hispid above, sparingly to densely pubescent beneath ; 
plants dioecious, the flowers in dense clusters forming elongate 
spikes; fruiting perianth at length broadly obovate, compressed. 

Junin: Alcotanga, Raimondi 9199. Ayacucho: Carrapa, Killip 
& Smith 22338. Cuzco: Urubamba Valley, Cook & Gilbert 886. 
Santa Ana, Cook & Gilbert 1644- Cedrobamba, Herrera 2008. 
Machupicchu, Herrera 3214, 3216, 3244, 3245. Puno: Sandia, 
Weberbauer 649. Without locality: Gay 636. Widely distributed in 
tropical America. "Quisa-quisa" (Cuzco). 

Boehmeria fallax Wedd. Arch. Mus. Paris 9: 346. 1856-57. B. 
fallax var. cordata Wedd. in DC. Prodr. 16, pt. 1: 198. 1869. 

A much branched shrub 2-5 meters high, with slender, hir- 
sutulous or puberulent branches; leaves alternate, those of the 
adjacent nodes very unequal and dissimilar, the larger ovate- 
lanceolate or oblong-lanceolate, 2-15 cm. long 1-6 cm. wide, long- 
acuminate, short-petioled, coarsely crenate-serrate, membranous, 
flat or slightly rugulose, scaberulous and more or less hispidulous 
above, pilosulous and pale beneath, the smaller leaves suborbicular- 
reniform, about 5 mm. wide, few- toothed, sessile, soon deciduous; 
plants monoecious or rarely dioecious, the flower clusters androgy- 
nous or unisexual, up to 5 mm. wide. 

Huanuco: Casapi, Mathews 2036 (type). Junin: La Merced, 700 
meters, Killip & Smith 23662, 23712. Between San Nicolas and 
Azupizu, Pichis Trail, Killip & Smith 26113. Ayacucho: Carrapa, 
1,500 meters, Killip & Smith 22954- Aina, 1,000 meters, Killip & 
Smith 22765. Estrella, 500 meters, Killip & Smith 23087. Cuzco: 
Urubamba Valley, 1,800 meters, Cook & Gilbert 1118. Without 
locality: Pavon (type of B. fallax var. cordata); Poeppig 1623; 
Mathews 2037. Also in Bolivia. 



FLORA OF PERU 355 

Boehmeria Mathewsii Killip, nom. nov. Boehmeria hirta Wedd. 
Ann. Sci. Nat. IV. 1: 202. 1854, not Swartz, 1788. B. Weddelliana 
Killip, Journ. Wash. Acad. Sci. 15: 55. 1925, not Vidal, 1886. 

A compact shrub 1.5-2 meters high, the branches densely hir- 
sute; leaves opposite, broadly ovate, 6-10 cm. long, 3-6 cm. wide 
(extremes up to 21 cm. long and 13 cm. wide), acuminate, rounded 
at the base, crenate-serrate, membranous, rugulose, hispid above, 
cano-tomentose beneath, becoming glabrescent; plants apparently 
dioecious, the flower clusters axillary, 8-10 mm. wide, the perianth 
of the pistillate flowers about 2 mm. long. 

Huanuco: Mufia, 2,200 meters, 4012. Without locality: Matheivs 
2039 in part (type of B. hirta Wedd.). 

Boehmeria Pavonii Wedd. Ann. Sci. Nat. IV. 1: 202. 1854. 
Procris longifolia Pavon ex Wedd. loc. cit., as synonym. Boehmeria 
Pavonii var. diversifolia Wedd. in DC. Prodr. 16, pt. 1: 199. 1869, 
in part. Phenax pallidus Rusby, Mem. Torrey Club 4: 259. 1895. 
B. pallida Killip, Journ. Wash. Acad. Sci. 21: 347. 1931, excluding 
synonym "B. diversifolia Wedd." 

A shrub or a tree, 2-6 meters high, the branchlets nearly straight, 
finely appressed-pubescent; leaves alternate, those of the adjacent 
nodes similar but usually very unequal, oblong-lanceolate or elliptic- 
lanceolate, serrulate or crenate-serrulate, strongly trinerved, dis- 
tinctly petioled, subcoriaceous, flat or slightly rugulose, appressed- 
hispidulous above, pale and closely appressed-short-silvery-sericeous 
beneath, the hairs pointing toward the middle of the areoles, the 
larger leaves 4-16 cm. long, 1.5-4.5 cm. wide, attenuate-acuminate, 
the smaller ones 1-3 cm. long, 0.5-1.5 cm. wide, sessile or short- 
petioled; plants dioecious, rarely monoecious, the flower clusters 
6-8 mm. wide, unisexual, rarely androgynous. 

San Martin: Tarapoto, Spruce 4029, 4951; Williams 5772. 
Zepelacio, 1,200-1,600 meters, King 3572. Alto Rio Huallaga, 
Williams 6314, 6781. Loreto: Balsapuerto, 220 meters, King 
2915. Junin: La Merced, 700 meters, 'Killip & Smith 23442. 
Ayacucho: Aina, 800 meters, Killip & Smith 23125. Without 
locality: Ruiz & Pav6n (type). Also in Bolivia. "Ishanga." 

There has been much confusion regarding this species and B. 
diversifolia Wedd., due to Weddell's varying interpretations in his 
three surveys of the genus, and to the misidentification of historic 
collections of Boehmeria in several European herbaria. Many of the 
specimens cited above were distributed as B. pallida. 



356 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

4. PHENAX Wedd. 

Unarmed shrubs or suffrutescent herbs; leaves alternate, toothed, 
rarely entire, 3-5-nerved; plants monoecious or dioecious, the flow- 
ers in dense, sessile, axillary clusters, the staminate usually 4-lobed, 
the pistillate without a perianth, subtended by several bractlets, the 
stigma elongate-filiform, persistent. 

Plants dioecious; leaves usually narrowly lanceolate, attenuate- 
acuminate P. angustifolius. 

Plants monoecious; leaves ovate or ovate-elliptic (lanceolate in one 

species), acute, acuminate, or obtuse. 
Leaves entire or with 1 or 2 coarse teeth. 

Inflorescence densely flowered; leaves lanceolate, 1-2 cm. long, 

quintuplinerved P. integrifolius. 

Inflorescence loosely flowered; leaves ovate or ovate-lanceolate, 

less than 1 cm. long, trinerved P. Weddellianus. 

Leaves with several teeth. 
Flower clusters unisexual, loosely flowered; leaves coarsely and 

unequally crenate P. laxiflorus. 

Flowers clusters androgynous; leaves serrate or crenate-serrate. 
Leaves essentially glabrous, usually drying light green. 

P. laevigatus. 

Leaves pubescent, at least beneath, drying darker. 
Stem slender, deeply angled, pilosulous; leaves membra- 
nous; staminate and pistillate flowers subequal in num- 
ber to a cluster P. hirtus. 

Stem stout, subangular, hirsute; leaves coriaceous or sub- 
coriaceous; staminate and pistillate flowers often very 
unequal in number to a cluster P. rugosus. 

Phenax angustifolius (HBK.) Wedd. Ann. Sci. Nat. IV. 1: 
193. 1854. Boehmeria angustifolia HBK. Nov. Gen. & Sp. 2: 34. 
1817. Procris longifolia Poepp. ex Wedd. loc. cit., as synonym. 
Phenax Ulei Krause, Verh. Bot. Ver. Brandenb. 48: 151. 1906. 

A slender shrub or suffrutescent herb, up to 3 meters high, with 
several branches; leaves linear-lanceolate or lanceolate, 5-15 cm. 
long, 0.5-5 cm. wide, gradually attenuate-acuminate, minutely 
serrulate, glabrous above, sparingly strigillose on the nerves and 
veins beneath, the nerves impressed above; plants dioecious; achenes 
minute, granular, the style 4-5 mm. long. 



FLORA OF PERU 357 

Loreto: Raimondi 2203. Iquitos, Killip & Smith 27446. Near 
Pongo de Manseriche, 100-200 meters, Killip, Smith & Dennis 
29146; Mexia 6323. Cerro de Escolar (Escalera), 1,200 meters, Ule 
6842 (type of P. Ulei). Junin: Colonia Perene", 700 meters, Killip 
& Smith 24911, 25336. La Merced, Killip & Smith 23568. Aya- 
cucho: Aina, Killip & Smith 22513. Without locality: Poeppig 
1315. Costa Rica to Colombia, Bolivia, and Brazil. 

Phenax Ulei appears to be merely a variant with proportion- 
ately broader leaves. 

Phenax hirtus (Sw.) Wedd. in DC. Prodr. 16, pt. 1: 235 38 . 
1869. Boehmeria hirta Sw. Prodr. 34. 1788. P. urticaefolius Wedd. 
Ann. Sci. Nat. IV. 1: 192. 1854, in part. P. petiolaris Wedd. loc. cit. 

A shrub or suffrutescent herb, 0.5-2.5 meters high, the stem and 
branches slender, deeply angled, pilosulous; leaves ovate or rarely 
ovate-lanceolate, 3-12 cm. long, 1.5-7 cm. wide, acute or acuminate, 
dentate-serrate, thin, flat or slightly rugulose, sparingly hispid above, 
hirtellous on the nerves beneath; plants monoecious, the glomerules 
androgynous with the staminate and pistillate flowers nearly equal 
in number; achenes verruculose. 

San Martin: San Roque, 1,400 meters, Williams 7161. Caja- 
marca: Cajamarca, Raimondi 1765. Hualgayoc, Raimondi 4176. 
Chorrillos, Raimondi 7204. Tambillo, Raimondi 4630, 6039. Ca- 
llacate, Raimondi 4227. Junin: Alcotanga, Raimondi 9198. Aya- 
cucho: Carrapa, 1,200 meters, Killip & Smith 22436. Aina, Killip 
& Smith 23176. Without locality: Ruiz & Pawn. This species is 
widely distributed in tropical America. Some of the specimens 
cited above have proportionately longer leaves than in typical 
Jamaican material, and may represent a variety. 

Phenax hirtus var. minor Wedd. in DC. Prodr. 16, pt. 1: 
235 39 . 1869. P. urticaefolius var. minor Wedd. Arch. Mus. Paris 
9: 496. 1856-57. 

Leaves 0.8-3 cm. long, 0.5-2 cm. wide; flower clusters usually 
smaller than in the typical form. 

Loreto: Near Yurimaguas, 135 meters, Killip & Smith 27863, 
28849. Cajamarca: Chorrillos, Raimondi 7760. Lima: Matucana, 
233, 2883. Obrajillo, Wilkes Expedition. Lima, Raimondi 12894- 

Phenax integrifolius Wedd. Ann. Sci. Nat. IV. Bot. 1: 193. 1854. 

Plant suffrutescent, the stem repent, at length procumbent or 

ascending, diffusely branched, villous; leaves crowded, lanceolate, 



358 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1-2 cm. long, 3-8 mm. wide, entire, quintuplinerved, pilose; plants 
monoecious, the glomerules densely flowered, androgynous; achenes 
minutely granular. 

Type a Peruvian plant collected by Pavon. 

Phenax laevigatus Wedd. Ann. Sci. Nat. IV. 1: 192. 1854; 
Arch. Mus. Paris 9: 497. pi 16 A. 1856-57. 

A shrub 1-2.5 meters high, with spreading branches, the younger 
ones pilosulous; leaves ovate, 3-9 cm. long, 1.5-4 cm. wide, acute or 
acuminate, coarsely crenate-serrate or serrate, coriaceous, essentially 
glabrous, sublustrous; plants monoecious, the glomerules androgy- 
nous, 5-8 mm. in diameter. 

Amazonas: Chachapoyas, Mathews 1556. Huanuco: Mito, 
2,700 meters, 1503. Huacachi, 2,000 meters, 4153. Pillao, Ruiz & 
Pavon. Junin: Huasa-huasi, Dombey (type). Huacapistana, Killip 
& Smith 24099. Palca, Weberbauer 1775. Lima: Above Lima, 
Raimondi 12548. Ayacucho: Carrapa, 2,200 meters, Killip & 
Smith 22312. Cuzco: Valle del Apurimac, Herrera. Also in Ecua- 
dor and Bolivia. 

Phenax laxiflorus Wedd. Arch. Mus. Paris 9: 499. 1856-57. 

Plant suffrutescent, the branches slender, puberulent; leaves 
ovate or elliptic, 1.5-4 cm. long, 1-2 cm. wide, obtuse, coarsely and 
unequally crenate with 2-4 crenations to a side, pilosulous above, 
finely pubescent beneath; plants monoecious, the glomerules loosely 
flowered, unisexual, the staminate in the lower axils, the pistillate 
in the upper. 

Huanuco: Pillao, Ruiz & Pavon (type). Muna, 2,500 meters, 
4282, referred here doubtfully. 

Phenax rugosus (Poir.) Wedd. in DC. Prodr. 16, pt. 1: 235 38 . 
1869. Procris rugosa Poir. in Lam. Encycl. 5: 628. 1804. Boehmeria 
ballotaefolia HBK. Nov. Gen. & Sp. 2: 35. 1817. Phenax ballotae- 
folius Wedd. Ann. Sci. Nat. IV. 1: 192. 1854. 

A shrub 2-3.5 meters high, the stem stout, subangular, densely 
hirsute; leaves ovate, 4-12 cm. long, 2.5-7 cm. wide, crenate-ser- 
rate, coriaceous or subcoriaceous, rugose, usually villous-tomentose 
beneath ; plants monoecious, the glomerules very dense, the staminate 
and pistillate flowers often very unequal in number in a glomerule; 
achenes verruculose. 

Huanuco: Muna, Pearce 158. Junin: Carpapata, 3,000 meters, 
Killip & Smith 24420. Cuzco: Ollantaitambo, 3,000 meters, Cook 



FLORA OF PERU 359 

& Gilbert 588. Urubamba Vallay, Herrera 2248. Vilcanota Valley, 
H err era 1077. Dept. uncertain: Panahuanca, Mathews 931. 
Without locality: Jussieu (type); Mathews 2033; Gay 360, 1661. 
Mexico to Venezuela and Bolivia. "Monte-pespeta" (Cuzco). 

Phenax rugosus var. minor Wedd. in DC. Prodr. 16, pt. 1: 
235 38 . 1869. 

Leaves elliptic-ovate, 1.5-2 cm. long, 0.8-1 cm. wide, subsessile, 
serrate, thick-coriaceous, bullate, lustrous, glabrous above, tomentel- 
lous beneath. 

Junin: Tarma, Mathews 669 (type). 

Phenax Weddellianus Killip, Journ. Wash. Acad. Sci. 24: 43. 
1934. 

Plant suffrutescent, with a slender, glabrescent stem, diffusely 
branched; leaves ovate or ovate-lanceolate, 5-18 mm. long, 3-8 
mm. wide, entire or with 1-2 coarse teeth on either side, trinerved, 
sparingly strigillose above, pilosulous beneath; plants monoecious, 
the clusters loosely flowered, androgynous or rarely unisexual; 
achenes narrowly ovoid. 

Cuzco: Gay (type). 

5. MYRIOCARPA Benth. 

Unarmed trees or shrubs; leaves alternate, petiolate, toothed or 
rarely subentire, trinerved, bearing cystoliths; plants dioecious, 
rarely monoecious; staminate flowers in glomerules forming slender, 
dichotomous spikes, the perianth 4-parted; pistillate flowers in 
slender, elongate, dichotomous, sub-unilateral spikes, rarely in pani- 
cles, without a perianth, the achenes compressed, stipitate or sessile, 
often setose at the margin, the stigma lateral, semilunate. 

Leaves ovate to ovate-oblong, conspicuously toothed nearly to 
the base, membranous, the cystoliths of the upper surface 
conspicuous M. stipitata. 

Leaves prevailingly obovate, undulate-serrulate in the upper half, 
coriaceous, the cystoliths obscure M. laevigata. 

Myriocarpa laevigata Killip, Journ. Wash. Acad. Sci. 15: 55. 
1925. 

A compact tree or shrub, up to 5 meters high, essentially glabrous 
throughout; leaves obovate, rarely ovate, 7-14 cm. long, 4-8 cm. wide, 
abruptly acuminate or acute, shallowly undulate-serrulate above the 
middle, coriaceous, the cystoliths obscure; plants dioecious or some- 



360 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

times monoecious, the spikes unisexual, once or twice forked near the 
base, 6-15 cm. long; staminate flowers sessile in contiguous clusters; 
achenes ovate-elliptic, setose at the margin, stipitate. 

Huanuco: Muna, 2,300 meters, 3925 (type). Yanano, 1,800 
meters, 3783. Rio Posuso, 1,900-2,000 meters, Weberbauer 6735, 
6743. Junin: Huacapistana, 1,800 meters, Killip & Smith 24186. 

Myriocarpa stipitata Benth. Bot. Voy. Sulph. 168. pi. 55. 1844. 
M. densiflora Benth. op. cit. 169. M. Dombeyana Wedd. Ann. Sci. 
Nat. III. 18: 232. 1852. M. densiflora var. Dombeyana Wedd. 
Arch. Mus. Paris 9: 487. 1856-57. 

A tree 3-9 meters high, with a slender trunk, the branchlets 
usually hirsute- tomentose; leaves broadly ovate to ovate-oblong, 
10-25 cm. long, 6-12 cm. wide, acuminate, rounded or cordulate at 
the base, crenate to crenate-serrulate, rugulose with age, bearing on 
the upper surface conspicuous cystoliths which radiate from the 
center of the areoles, membranous, essentially glabrous above, 
sparingly pilosulous to densely tomentose beneath; spikes 1-3 times 
forked near the base, the staminate shorter than the leaves, the 
pistillate often much longer, up to 30 cm. long; achenes elliptic, 
1-1.5 mm. long, subsessile or usually stipitate, generally setose at 
the margin. 

Piura: Prov. Huancabamba, 1,300 meters, Weberbauer 601 4- 
San Martin: Tarapoto, 750 meters, Spruce 4286; Ule 6507; Williams 
6145. Pongo de Cainarachi, 230 meters, Klug 2748. Zepelacio, Klug 
3536. Loreto: Balsapuerto, 220 meters, Klug 2898. Pongo de 
Manseriche, Mexia 6252. Rio Nanay, Williams 414' Cuschi, 
1,600 meters, 4838. Pampayacu, Sawada 16. Junin: Huasa-huasi, 
Dombey (type of M. Dombeyana). Carpapata, 2,700 meters, Killip 
& Smith 24374- Huacapistana, 2,000 meters, Killip & Smith 
24139. La Merced, 600 meters, 5454; Killip & Smith 23475, 24059, 
24064, 24075, 24077, 25386. Dos de Mayo, Pichis Trail, Killip & 
Smith 25837, 25856. Ayacucho: Carrapa, 1,000 meters, Killip & 
Smith 22467, 22487. Aina, 900 meters, Killip & Smith 22687. 
Estrella, 500 meters, Killip & Smith 23073. Kimpitiriki, 400 meters, 
Killip & Smith 22952. Cuzco: Urubamba Valley, 1,800 meters, Cook 
& Gilbert 937, 1047, 1147. Without locality: Mathews 2040 (type of 
M. densiflora). Colombia to Venezuela and Ecuador. "Ishanga" 
(Tarapoto), "tigre-tigre" (Cuzco). 

M. densiflora was differentiated from M. stipitata on the basis of 
the leaves being softly tomentose beneath and the achenes sub- 



FLORA OF PERU 361 

sessile rather than stipitate, and the two species have always been 
considered distinct. The large number of specimens now available 
fail to show a correlation of these or any other characters; the degree 
of indument is highly variable and, even in a single spike, there is 
variation in the relative length of the stipe to the body of the achene. 

6. URERA Gaud. 

Trees or shrubs, usually with stinging hairs; leaves alternate, 
stipulate, petiolate, the cystoliths punctiform, linear, or wanting; 
plants usually dioecious, the flowers small, in axillary, dichotomous 
or irregularly branched cymes, the staminate with a 4-5-parted 
perianth and 4-5 stamens, the pistillate with 4 equal or unequal 
segments; stigma penicillate persistent; achene straight or oblique, 
at least partially surrounded by the fleshy, enlarged perianth. 

Leaves incised-lobed usually more than halfway to the midnerve. 

U. laciniata. 

Leaves not lobed, dentate, crenate, or subentire. 
Achenes more than 2 mm. long; leaves coarsely dentate or sinuate- 
dentate U. baccifera. 

Achenes less than 2 mm. long; leaves crenate-dentate. 

U. caracasana. 

Urera baccifera (L.) Gaud, in Freyc. Voy. Uran. Bot. 497. 
1826. Urtica baccifera L. Sp. PI. ed. 2. 1398. 1763. 

A coarse, erect, subligneous herb, or a shrub, 1-4 meters high, the 
stem densely covered with short, stout, stinging prickles; leaves 
broadly ovate or round-ovate to oblong-ovate, up to 35 cm. long and 
15 cm. wide, rounded or cordate at the base, coarsely dentate or 
irregularly sinuate-dentate, glabrescent, rarely densely pubescent, 
or with hairs or prickles, especially on the nerves; plants dioecious, 
the flowers in much branched cymes; fruit succulent, white or rose- 
colored, 3-5 mm. long. 

San Martin: Chazuta, 260 meters, Klug 4029. Loreto: Rio 
Maranon Valley, 150 meters, Killip, Smith & Dennis 29169. Rio 
Itaya, 110 meters, Killip & Smith 29376, 29465, 29579; Williams 
3423. Mishuyacu, 100 meters, Klug 1085. Lower Rio Huallaga, 
Williams 4350. Huanuco: Muna, 2,200 meters, 3910. Monzon, 
Weberbauer 3497, 3498. Junm: Pichis Trail, Killip & Smith 25904, 
26105, 26356. Ayacucho: Aina, 900 meters, Killip & Smith 22794. 
Madre de Dios: Seringal, Ule 9330. Widely distributed in 
tropical America. 



362 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Urera caracasana (Jacq.) Gaud, ex Griseb. Fl. Brit. W. Ind. 
154. 1859. Urtica caracasana Jacq. Hort. Schoenbr. 3: 71. pi. 396. 
1798. Urera Jacquini Wedd. Ann. Sci. Nat. III. 18: 200. 1852. U. 
subpeltata Miq. in Mart. Fl. Bras. 4, pt. 1: 189. pi. 66. 1853. U. 
acuminata Miq. op. cit. 190, not Gaud. U. Jacquini var. subpeltata 
Wedd. Arch. Mus. Paris 9: 145. 1856-57. U. Jacquini var. Miqueli 
Wedd. loc. cit. U. caracasana var. subpeltata Wedd. in DC. Prodr. 
16, pt. 1: 90. 1869. U. caracasana var. Miqueli Wedd. loc. cit. U. 
capitata var. Pavonii Wedd. op. cit. 92. 

A shrub 1-6 meters high, with elongate branches, the young 
twigs, petioles, inflorescence, and leaf veins usually armed with 
stinging hairs; leaves broadly ovate or sometimes ovate-lanceolate, 
variable in size, up to 30 cm. long and 25 cm. wide, acuminate, cor- 
date at the base, usually with an open sinus but sometimes with a 
closed sinus and overlapping basal lobes, crenate-dentate, scabrous 
above, more or less pubescent beneath; plants dioecious, the cymes 
regularly dichotomous, shorter than the petioles or often much 
longer, the staminate flowers sessile in distinct, compact glomerules, 
the pistillate flowers usually ternate, sometimes single or in dense 
clusters, pedicellate or subsessile. 

San Martin: Zepelacio, Klug 3554- Tarapoto, Spruce 4305. 
Loreto: Lower Rio Huallaga, Williams 4288, 4308, 4796, 4945. 
Iquitos, Tessmann 3584; Williams 8041- Rio Maranon, Killip & 
Smith 27518, 29230; Tessmann 3857, 4118. Pumayacu, Klug 3174. 
Mishuyacu, Klug 441, 1071. Rio Putumayo, Klug 1639. Rio 
Napo, Mexia 6469. Rio Amazonas, Williams 1861, 2005, 2673, 
2789. Rio Nanay, Williams 490, 505, 802, 1256. Cerro de Escalera, 
Ule 45 in part. Yarina-cocha, Tessmann 3198. Huanuco: Casapi, 
Mathews 2030. Monzon, Weberbauer 3704. Junin: La Merced, 
Killip & Smith 23929. Rio Perene", Killip & Smith 25151. Ayacu- 
cho: Kimpitiriki, Killip & Smith 22935. Madre de Dios: Seringal, 
Ule 9332. Dept. uncertain: Pampa del Sacramento, Castelnau. 
Without locality: Ruiz & Pavdn (type of U. capitata var. Pavonii); 
Gay; Poeppig; Fox 32. Widely distributed in tropical America. 
"Ishanga," "ishangu del agua." 

Urera caracasana is here interpreted in its widest sense. Many 
of the specimens from northern Peru cited above have proportion- 
ately narrower leaves than in the typical form and represent the 
variety subpeltata. If further study proves that U. capitata, with 
the pistillate flowers sessile in dense heads, is specifically distinct 
from U. caracasana, some of this material, including the type of U. 



FLORA OF PERU 363 

capitata var. Pavonii, should be placed in that species. At present 
it seems best to treat these as representing a young stage of U. 
caracasana. 

Urera laciniata (Goudot) Wedd. Ann. Sci. Nat. III. 18: 203. 
1852. Urtica laciniata Goudot ex Wedd. loc. cit., as synonym. 

An erect, tree-like herb or often truly arborescent, 1-4 meters 
high, the branches densely covered with stout bristles or spines; 
leaves 15-35 cm. long and wide, deeply incised-lobed (lobes acumi- 
nate, entire or few-toothed), membranous, spiny on the nerves 
beneath, glabrescent; plants dioecious, the cymes up to 20 cm. long, 
the staminate flowers in glomerules, the pistillate in glomerules or 
distinct; achenes suborbicular, nearly 2 mm. long. 

San Martin: Tarapoto, Spruce 4140. Alto Rio Huallaga, Wil- 
liams 6799. San Roque, 1,350-1,500 meters, Williams 7785. 
Loreto: Mishuyacu, 100 meters, Klug 1079. Rio Nanay, Williams 
451. Yurimaguas, 200 meters, Williams 4937. Rio Ucayali, Tess- 
mann 3177. Huanuco: Posuso, 600 meters, 4703. Junin: La 
Merced, 600 meters, 5315. Puerto Yessup, 400 meters, Killip & 
Smith 26319. Ayacucho: Estrella, 500 meters, Killip & Smith 
22645. Without locality: Haenke 1730; Ruiz & Pawn. Costa Rica 
to Venezuela, Peru, and Bolivia. "Ishanga." 

7. POUZOLZIA Gaud. 

Low shrubs, rarely woody vines; leaves alternate, entire, toothed 
in only a few American species, petiolate, 3-nerved, the upper sur- 
face densely covered with punctiform cystoliths; plants monoecious, 
rarely dioecious, the flowers in small, axillary clusters or in leafless 
spikes; staminate perianth 4-5-parted or lobed, the stamens 3-5; 
pistillate perianth tubular, 2-4-toothed, usually costate, the ovary 
included, the stigma filiform, at length deciduous, the achenes 
crustaceous, shiny. 

Leaves entire. 

Flower clusters in leafless spikes; plants scandent, dioecious, 

glabrescent P. formicaria. 

Flower clusters in the leaf axils; plants monoecious, densely 

pubescent. 
Under surface of the leaves hirtellous on the nerves, compactly 

white-lanate between the nerves P. Poeppigiana. 

Under surface of leaves densely brown-hirsute P. obliqua. 

Leaves coarsely crenate-serrate P. longipes. 



364 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pouzolzia formicaria (Poepp.) Wedd. Arch. Mus. Paris 9: 
407. 1856-57. Boehmeria formicaria Poepp. ex Wedd. Ann. Sci. Nat. 
IV. 1:201. 1854. 

A woody vine or a scandent shrub; leaves oblong-lanceolate, 4- 
12 cm. long, 2-6 cm. wide, acuminate, rounded or cordulate at the 
base, entire, subcoriaceous, concolorous, scabrid above, essentially 
glabrous; plants dioecious, the flower clusters borne in spikes 8-15 
cm. long; pistillate perianth about 2 mm. long, hispidulous; achenes 
conical, about 1.5 mm. long, brownish white, the style about 3 times 
longer, densely pubescent. 

Loreto: Mainas, Poeppig 2287 (type). Rio Paranapura, 200 
meters, Klug 3944- Rancho Indiana, 110 meters, Mexia 6411. 
Mishuyacu, 100 meters, Klug 930. Iquitos, 120 meters, Williams 
8117; Tessmann 3585, 3897, 3898. Rio Ucayali, Tessmann 3336. 
Lower Rio Nanay, Williams 557. Florida, Rio Putumayo, 200 
meters, Klug 1998, 2030, 2354. 

Pouzolzia longipes Killip, sp. nov. P. procridioides var. hirsuta 
Wedd. in DC. Prodr. 16, pt. 1: 231. 1869(?). 

Herba monoica; folia late ovata, acuminata, grosse crenato-ser- 
rata, supra setulosa, subtus in nervis puberula; glomeruli axillares, 
androgyni vel unisexuales, perianthio masc. 4-lobato; achaenia 
conica. 

An herb 60-75 cm. high, woody below, the stem sparingly pilo- 
sulous, at length glabrous, angulate; leaves broadly ovate, 5-10 
cm. long, 3-6 cm. wide, acuminate, rounded and abruptly narrowed 
at the base, coarsely crenate-serrate, membranous, sparingly setulose 
above, puberulent on the nerves beneath, otherwise glabrous, the 
petioles 2-5 cm. long; plants monoecious, the flower clusters axillary, 
5-8 mm. wide, androgynous or unisexual; staminate perianth about 
2.5 mm. long, deeply 4-lobed; pistillate perianth 1-1.5 mm. long, 
scaberulous; achenes conical, nearly 1 mm. long, light brown, the 
styles 3 mm. long, pubescent. 

Junin: Enefias, Pichis Trail, 1,700-1,900 meters, dense forest, 
July 2, 1929, Killip & Smith 25753 (type, U. S. Nat. Herb. No. 
1,359,857; Field Mus. No. 632,801). 

This species, one of the few American ones with toothed leaves, is 
nearest the African P. procridioides (E. Mey.) Wedd., and agrees 
well with the Bonpland specimen at Paris, labeled merely "Amer. 
Trop.," which Weddell referred to P. procridioides. In addition to 
the improbability of the African species occurring in Peru, there are 



FLORA OF PERU 365 

slight differences in the toothing of the leaves, the indument, and the 
size of the achenes which suggest that another species is represented. 
The type of P. procridioides var. hirsuta, a Pavon specimen in the 
Boissier Herbarium, not examined in connection with the present 
studies, may be P. longipes, though the description of the indument 
does not well apply to the specimen cited above. 

Pouzolzia obliqua (Poepp.) Wedd. Arch. Mus. Paris 9: 405. 
1856-57. Margarocarpus obliquus Wedd. Ann. Sci. Nat. IV. 1: 
204. 1854. Boehmeria obliqua Poepp. ex Wedd. loc. cit., as synonym. 

A shrub or a small tree, 2-5 meters high, densely hirsute nearly 
throughout; leaves oblong or oblong-lanceolate, 2-14 cm. long, 1-5 
cm. wide, attenuate-acuminate, rounded or cordulate at the base, 
oblique, short-petioled, hispid above; plants monoecious or dioecious, 
the flower clusters axillary, androgynous or unisexual; achenes light 
brown or white, the styles 8-10 mm. long. 

San Martin: Juanjui, Klug 1+169. Loreto: Mainas, Poeppig 2032 
(type). Guatemala to Venezuela and Peru, evidently uncommon 
in Peru. 

Pouzolzia Poeppigiana (Wedd.) Killip, Journ. Wash. Acad. 
Sci. 21: 347. 1931. Margarocarpus Poeppigianus Wedd. Ann. Sci. 
Nat. IV. 1 : 204. 1854. M. asper Wedd. loc. cit. Boehmeria discolor 
Poepp. ex Blume, Mus. Bot. Lugd. Bat. 2: 206. 1856. B. aspera 
Blume, loc. cit., footnote, not Wedd. Pouzolzia discolor Wedd. 
Arch. Mus. Paris 9: 408. pi. 13B, f. 18-24- 1856-57. P. aspera 
Wedd. in DC. Prodr. 16, pt. 1: 233. 1869, not Wight. 

A shrub 1-5 meters high, with elongate, villous branches; leaves 
lanceolate or oblong-lanceolate, 2-15 cm. long, 1-4.5 cm. wide, 
acuminate, rounded at the base, symmetrical or nearly so, entire, 
above scabrous and hirsutulous, beneath rufo-hirtellous on the 
nerves and compactly white-lanate between them; plants monoe- 
cious, the flower clusters axillary, androgynous or unisexual, the 
styles up to 1.5 cm. long, the achenes light brown or white, about 
1 rnm. long. 

San Martin: Lamas, 840 meters, Williams 6442; Spruce. Alto 
Rio Huallaga, Williams 6683. Zepelacio, 1,400 meters, Klug 3437. 
-Loreto: Rio Amazonas, Poeppig 3044- Huanuco: Rio Huallaga 
Canyon, 1,200 meters, 4328. Junin: La Merced, 700 meters, Killip 
& Smith 23386, 23528, 23586, 23938. Colonia Perene", Killip 
& Smith 25024- Ayacucho: Carrapa, 1,000 meters, Killip & Smith 
22471. Cuzco: Bues in 1930. Without locality: "Peru subandina," 



366 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Poeppig 1284 (type; also type of P. discolor); Ruiz & Pawn; Gay; 
Mathews 2036, 2037. Also in Bolivia. 

8. FLEURYA Gaud. 

Annual herbs, usually with stinging hairs; leaves alternate, 
petiolate, toothed, trinerved; plants monoecious or dioecious, the 
flowers in clusters in large panicles, the staminate 4-5-parted, the 
pistillate with 4 imbricate perianth segments, the stigma papillose, 
at length hooked; achenes oblique, compressed. 

Fleurya aestuans (L.) Gaud, in Freyc. Voy. Uran. Bot. 497. 
1826. Urtica aestuans L. Sp. PI. ed. 2. 1397. 1762. F. glandulosa 
Wedd. Ann. Sci. Nat. III. 18: 205. 1852. F. aestuans var. glandu- 
losa Wedd. Arch. Mus. Paris 9: 112. 1856-57. F. aestuans var. race- 
mosa Wedd. in DC. Prodr. 16, pt. 1: 72. 1869. 

An erect, simple or few-branched herb, up to 1.2 meters high, the 
stem sometimes glandular; leaves broadly ovate, 7-17 cm. long, 2.5- 
12 cm. wide, acute or acuminate, coarsely and sharply dentate, 
slender-petioled, membranous, usually with a few stinging hairs 
on the upper surface; panicles usually androgynous; achenes 
1-1.5 mm. long. 

Loreto: Fortaleza, Williams 4462. Puerto Arturo, 135 meters, 
Killip & Smith 27755; Williams 5130, 5195. Iquitos, 100 meters, 
Kittip & Smith 27074; Williams 1340, 1348, 8099, 8193. Lower 
Rio Nanay, Williams 353. La Victoria, Williams 2603. Nauta, 
Raimondi 2173. Without locality: Poeppig 2086; Ruiz & Pawn. 
Throughout tropical America. 

9. PARIETARIA L. 

Diffuse or tufted, slender herbs, without stinging hairs; leaves 
alternate, entire, trinerved; flowers in small, axillary glomerules, the 
staminate, pistillate, and perfect intermixed; perianth tubular, 
4-parted; achenes straight, ovoid, lustrous. 

Parietaria debilis Forst. Fl. Ins. Austr. Prodr. 73. 1786. P. 
debilis var. ceratosantha Wedd. Arch. Mus. Paris 9: 515. 1856-57. 

An erect or diffuse, pilosulous or glabrescent, annual herb; leaves 
ovate, rhombic-ovate, or orbicular-ovate, 5-20 mm. long, obtuse or 
rounded at the apex; clusters few-flowered. 

Cajamarca: Cascas, Raimondi 8174. Tambillo, Raimondi 7995. 
Lima: Matucana, 2,500 meters, 262. Lima, 5870; Rose 18581. 
Lurin, 5968. Atocongo, Pennell 14795. Arequipa: Mollendo, 



FLORA OF PERU 367 

Hitchcock 22391, 22410; Johnston 3552. Tiabaya, 2,000 meters, 
Pennell 13072. Arequipa, 2,600 meters Pennell 1 3194. Locality 
uncertain: Dombey; Weddell. Widely distributed in the warmer 
parts of the world. 

50. PROTEACEAE. Protea Family 

Besides the following genera of this family, which is best repre- 
sented in Australia, the silk-oak of Australia, Grevillea robusta Cunn., 
is doubtless cultivated, as in all warm regions, for shade. It may be 
known by its fern-like leaves, silky-pubescent beneath. 

Flowers showy, conspicuous above the leaves; hypogynous glands 

large, or the disk entire or obscurely 3-lobed ... 1. Embothrium. 

Flowers rather small, often inconspicuous; hypogynous glands or 

scales 3-4 or the disk 4-lobed. 
Inflorescence axillary, at least in part, 1-5 cm. long; hypogynous 

glands 3; style laterally expanded 2. Lomatia. 

Inflorescence usually terminal, often longer than the leaves; 

hypogynous scales or glands 4. 
Leaves conspicuously net-veined; fruit tardily dehiscent. 

Leaves entire 3. Panopsis. 

Leaves pinnate 4. Euplassa. 

Leaves inconspicuously net- veined; fruit early dehiscent. 

5. Roupala. 
1. EMBOTHRIUM Forst. 

Oreocallis R. Br. Trans. Linn. Soc. 10: 196. 1811. 

Tall shrubs with long, willow-like branches, the ultimate ter- 
minated by dense racemes of showy, strap-shaped, 1-petaled flowers. 
Style oblong-fusiform or oblique at the tip. The herbarium name 
Catas Dombey is mentioned in synonymy by Lamarck, Encycl. 
2: 355. 1786. 

Inflorescence more or less reddish-pubescent E. grandiflorum. 

Inflorescence glabrous, often pruinose-glaucous. 
Leaves mostly 3-3.5 cm. wide, rarely 10 cm. long. 

E. mucronatum. 
Leaves mostly 4-5 cm. wide, usually 10-15 cm. long. 

E. Weberbaueri. 

Embothrium grandiflorum Lam. Encycl. 2: 354. 1786. E. 
emarginatum R. & P. Fl. 1 : 62. pi. 95. 1798. Oreocallis grandiflora 
R. Br. Trans. Linn. Soc. 10: 197. 1811. 



368 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Branchlets dark-barked, the youngest reddish-tomentose as the 
new leaves beneath; petioles 1-2 cm. long; leaves coriaceous, gla- 
brous and lustrous above, elliptic, acutish at the base, rounded and 
mucronate or somewhat emarginate at the apex, about 10 cm. long 
and half as wide; racemes often finally 10 cm. long; pedicels 1 cm. 
long in flower, 3 cm. in fruit; flowers reddish, curved, 4-5 cm. long, 
the perianth deeply 4-parted; anthers ovate, sessile; styles as long 
as the perianth, persisting with the obliquely directed, peltate 
stigma on the oblong-cylindric capsule; seeds winged, orbicular, 
compressed. A shrub or tree of 1-6 meters. Meisner in DC. Prodr. 
14: 445. 1856, distinguished, under the name Oreocallis grandiflora: 
var. emarginata. (R. & P.) Meisn., the leaves beneath and the branch- 
lets pubescent; var. obtusifolia Meisn., the leaves glabrate; and 
var. acutifolia Meisn., similar to the last but the leaves acute. Web- 
erbauer (86) notes that the species is a widely spread east-Andean 
type which, however, occurs in the inter- Andean valleys of the Hua- 
llaga, etc., even to the western slopes in Cajamarca. Ruiz and Pavon 
found the crushed leaves applied to bruises and to aching teeth, and 
the flowering branches used to adorn altars and the arches carried 
in processions. Illustrated, Weberbauer, 160. 

Cajamarca: San Pablo, 2,400 meters, Weberbauer 3826; 256. 
Huambos, 2,600-3,000 meters (Weberbauer 260). Toward Hual- 
gayoc, 2,200 meters (Weberbauer 189). Ancash: Huaraz (Weber- 
bauer 179). Pichiu, 3,500 meters, Weberbauer 2922. Huanuco: 
Mito, 1380. Fifteen miles northeast of Huanuco, 3,000 meters, 
2149. Yanano, 1,800 meters, 4936. Palca and Huasa-huasi, Ruiz 
& Pavdn. Between Huanuco and Pampayacu, Kanehira 66. 
Junin: Huacapistana, 1,700 meters, Weberbauer 1742; 246. Chan- 
chamayo, Isern 2179, 2302,Cuzco: Valle de Lares (Hen era 788). 
Santa Ana, 1,900 meters, Weberbauer 5000; 281. Without locality, 
Diehl 2523. Pillahuata, 2,800 meters, Pennell 14084- Ecuador. 
"Salta-perico," "tsacpa," "cucharilla," "zacpa," "llamas," "chappa," 
"cocaniro," "picahua," "catas," "machinparrani," "mastimpanrani." 

Embothrium mucronatum Willd. ex R. & S. Syst. 3: 33. 1818. 
Oreocallis Ruizii Klotzsch, Linnaea 20: 474. 1847. 

Apparently very similar to E. grandiflorum but glabrous or nearly 
so and the leaves often much smaller, oblongish, mucronate, some- 
what lustrous above; terminal thyrse lax, the bracts sparsely pilose. 
A small shrub, perhaps only a variety of E. grandiflorum. Neg. 
11772. 



FLORA OF PERU 369 

Huanuco: Ruiz (type). Pampayacu, Sawada P41- Amazonas: 
Chachapoyas, Raimondi (det. Diels); at 2,700 meters, Williams 
7551. Ecuador. "Picahuay." 

Embothrium Weberbaueri Perk. Bot. Jahrb. 45: 434. 1911. 

A shrub of 3-5 meters, resembling E. mucronatum in lack of 
pubescence but perhaps distinguishable by the larger, mostly 
emarginate, obscurely mucronulate leaves and the terminal, racemose 
inflorescence; racemes 20 cm. long or longer, the slender pedicels 
2 cm. long; flowers rose-colored, 3.5-4 cm. long; fruits 4 cm. long, 
the beak 2.5-3 cm. long, on stipes to nearly 4 cm. long. Neg. 11773. 

Huanuco: Monzon, Prov. Huamalies, 1,000-1,500 meters, 
Weberbauer 3464. (type). San Martin: East of Moyobamba, 1,100 
meters, Weberbauer 4762 (det. Perkins). Near Moyobamba, Klug 
3417. San Roque, 1,400 meters, Williams 7820, 7668. 

2. LOMATIA R. Br. 

A shrub allied to Embothrium, but the axillary racemes rela- 
tively short and few-flowered. Style expanded laterally at the 
tip. The generic name has been conserved. 

Lomatia hirsuta (Lam.) Diels, comb. nov. Embothrium hirsu- 
tum Lam. Encycl. 2: 355. 1786. E. obliquum R. & P. Fl. 1: 63. pi. 
97. 1798. L. obliqua R. Br. Trans. Linn. Soc. 10: 196. 1811. 

A glabrous-leaved shrub 2-3 meters high with reddish-villous 
racemes about as long as the leaves, these coriaceous, lustrous above, 
often colored beneath, ovate, very unequally serrate, 5-12 cm. 
long, 2-5 cm. broad; perianth 1 cm. long, white; fruits pedicellate, 
2.5-3.5 cm. long. Some part of the plant is said to supply a dye. 

Cajamarca: San Miguel, 2,600 meters, Weberbauer 3890. Cu- 
tervo, Jelski. Piura: At 3,000 meters, Weberbauer 6053. Rio Piura 
and Nancho, Raimondi (det. Diels). Libertad: East of the Mara- 
fion, Raimondi (det. Diels). Chile. "Raral," "andaga," "garo," 
"shiapash." 

3. PANOPSIS Salisb. 

Andriapetalum Pohl, PI. Bras. Icon. 1: 113. 1827. 

Shrubs or small trees with entire, often verticillate leaves. 
Racemes axillary and terminal. Filaments obvious, borne below the 
middle of the perianth lobes. Disk cupulate, 4-lobed. Fruit a hard, 
1-seeded drupe. The later name of Pohl was accredited to Schott 
by Endlicher as Andripetalum. 



370 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels rigid, 10-12 mm. long, glabrous P. acuminata. 

Pedicels lax, 5-6 mm. long, pubescent P. rubescens. 

Panopsis acuminata (Meisn.) Macbr. Field Mus. Bot. 11: 66. 
1931. Andriapetalum rubescens Pohl, var. acuminatum Meisn. in 
DC. Prodr. 14: 346. 1856. 

Leaves subsessile, oblong-lanceolate, acute to long-acuminate, 
2.5-4 cm. wide, finely reticulate-veined on both sides, glabrous; 
racemes glabrous, the stiff pedicels divaricate; flowers cream-colored. 
Perhaps better regarded as a variety of the next. A tree of 
8 meters (Klug). Neg. 7445. 

Loreto: Mishuyacu, near Iquitos, 100 meters, Klug 1540. Brazil. 

Panopsis rubescens (Pohl) Pittier, Contr. Fl. Venez. 21. 1923. 
Andriapetalum rubescens Pohl, op. cit. 114. pi. 91. 

Similar to the above, but the young branchlets, leaves, and 
racemes rusty-tomentulose; leaves narrowed to both ends, obtusish, 
glabrous in age, beautifully reticulate-veined and suffused with red 
from the midnerve. Flowers pure white, with the odor of vanilla; 
fruit 4-6 cm. long, 1-1.5 cm. thick, densely tomentose, indehiscent 
(Ducke). The Peruvian form, var. simulans Macbr. Field Mus. 
Bot. 11: 67. 1931, has acute or acuminate leaves. Neg. 19067. 

Loreto: Timbuchi, Rio Nanay, Williams 1044 (type of var. 
simulans}. Manfmfa, upper Rio Nanay, Williams 1121. Brazil; 
British Guiana. 

4. EUPLASSA Salisb. 

Pinnately leaved shrubs or trees with racemes or narrow, raceme- 
like panicles of rather small flowers. Otherwise like Panopsis but 
perianth oblique in bud, anthers subsessile, and disk entire. 

Euplassa Isernii Cuatrecasas, sp. nov. 

Ramis ignotis; foliis petiolatis, petiolo 8-9 cm. longo tereti 
inferne applanato minute puberulento; foliolis 8-jugis breve petio- 
lulatis paullo alternantibus, petiolulo 3-4 mm. longo tereti striato 
puberulento, oblongo-ellipticis, acuminatis basi late acutis margine 
repando-serratis vel subintegris 8-12 cm. longis, circa 4 cm. latis, 
chartaceo-coriaceis conspicue reticulatis utrinque subnitidulis supra 
viridibus glabris subtus castaneis ad nervum medianum minute 
sparseque puberulentis; floribus paniculam angustam spurie race- 
mosam referentibus; paniculis cum pedunculo 6 cm. longo 30 cm. 
longis cum pilis adpressis minutis paullo fulvis parce pubescentibus; 



FLORA OF PERU 371 

pedicellis ad mediam connatis circa 7 mm. longis; petalis 10 mm. 
longis; ovario glabro. Apparently, by reason of its many leaflets 
and glabrous ovary, very well marked, and the second species west 
of the Andes, the other being E. occidentalis I. M. Johnston of Ecua- 
dor, with 4 pairs of obovate leaflets. Included here with Dr. Cua- 
trecasas' permission, the description supplied by me. 

Junin: Chanchamayo, Isern 2283, type, Herb. Madrid. 

5. ROUPALA Aubl. 

Stoutly branched shrubs with alternate leaves. Racemes or 
spikes slender, often more or less tomentose, the rather small flowers 
often divaricate on short pedicels. Anthers subsessile on the upper 
part of the perianth lobes. A fruiting specimen (Williams 7439, 
San Roque) has entire, ovate, long-acuminate, long-petioled leaves 
exactly simulating those of R. macropoda Karst. of Colombia, but 
the fruits are cylindric, 1 cm. long, 2 mm. thick; in the absence of 
flowers the generic identity is uncertain. The Aublet name has been 
written Rhopala, Ropala, and Rupala and, at least by typographical 
error, still otherwise, using the same letters. 

Flowers sessile, the spikes much exceeding the leaves . . . R. spicata. 
Flowers always more or less pedicellate. 

Mature leaves glabrous except the midnerve beneath, the trichomes 
not distinct, usually forming a gray indument. 

Leaves rotund-cordate; pubescence beneath grayish. 

R. cordifolia. 

Leaves not cordate, or at least the pubescence not a gray 
indument. 

Leaves simple, at least those of the flowering branchlets. 
Leaves rounded-truncate at the base . . . R. dolichopoda. 
Leaves cuneate at the base or at least acute. 

Racemes distinctly pubescent; leaves 3.5-4 cm. wide. 
Leaves finally glabrous beneath . . . . R. complicata. 
Leaves pubescent beneath on the costa . . R. ferruginea. 
Racemes glabrous; leaves mostly wider.... R. Dielsii, 

Leaves pinnate, but only the sterile branchlets known. 

R. Raimondii. 

Mature leaves more or less pubescent beneath, the trichomes dis- 
tinct, often red or fulvous. 



372 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence red-brown, very dense on the leaf nerves beneath; 

pedicels often only 1-2 mm. long R. monosperma. 

Pubescence grayish-fulvous, moderate; pedicels 3-5 mm. long. 

R. pinnata. 

Roupala complicata HBK. Nov. Gen. & Sp. 2: 153. pi 119. 1817. 

A slender shrub or tree with thick, subglabrous, glaucescent 
leaves and pallidly rusty-tomentose racemes that about equal them; 
leaf blades entire or remotely and obtusely few-dentate, decurrent 
into the petiole, this 2-3 cm. long; leaf nerves more or less elevated 
or subimmersed, the veins obscurely and laxly reticulate; racemes 
about 10 cm. long; pedicels 2-3 mm. long; ovary rusty-tomentose; 
stigma scarcely clavate. Variable; the specimens by Killip and 
Smith were determined by the latter as R. Gardneri Meisn., a form 
typically of interior Brazil with more elevated leaf nervation. See 
Field Mus. Bot. 11: 66. 1931. Neg. 11749. 

San Martin: Morales, Alto Rio Huallaga, 900 meters, Williams 
5703. Zepelacio, 1,100 meters, Klug 3742; a tree of 4 meters, the 
flowers light yellow. San Roque, 1,400 meters, Williams 7439. 
Moyobamba, 800 meters, Weberbauer 4480; 290. Junin: Chan- 
chamayo Valley, 1,200 meters, Schunke 426; Isern 2327. San 
Ramon, 900-1,300 meters, Killip & Smith 24782, 24891. Rio 
Perene", 600 meters, in forest, Killip & Smith 25114; a slender shrub 
of 1.5-2 meters; flowers creamy white. Colonia Perene", 680 meters, 
Killip & Smith 25035. Loreto: Along the Rio Ucayali, Tessmann 
3431. Cuzco: Santa Ana, 800 meters, Weberbauer 5024; also Uru- 
bamba (280). North to the Guianas and Costa Rica. "Ingaina," 
"arellan." 

Roupala cordifolia HBK. Nov. Gen. & Sp. 2: 152. pi. 118. 1817. 

Much branched, the branchlets glabrous; petioles to 2.5 cm. 
long; leaves to about 10 cm. long, remotely and coarsely dentate 
or subentire, fleshy-coriaceous, soon glabrous, the laxly reticulate 
veins subimmersed; racemes reddish-pubescent, very much longer 
than the leaves; flowers 8-10 mm. long, white, the slender, rusty- 
tomentose pedicels 2-3 times longer; stigma clavate. 

Cajamarca: Jae"n de Bracamoros (Humboldt, type). Huambos, 
Weberbauer 4192. Ecuador; Colombia. 

Roupala Dielsii Macbr. Field Mus. Bot. 11: 65. 1931. 
A glabrous shrub with slender, few-branched, densely leafy 
branchlets; leaves elliptic-oblong or somewhat obovate, basally 



FLORA OF PERU 373 

attenuate to the (3-5 mm. long) petiole, shortly and broadly acu- 
minate, 10 cm. long, 4 cm. broad, somewhat lustrous above but 
scarcely veiny, opaque beneath, the lateral veins elevated but 
obscurely reticulate, finally subcoriaceous, entire below but more 
or less denticulate above the middle; racemes with flowers about 
2 cm. wide, as long as the leaves, glabrous or glabrescent, only the 
ovary reddish-tomentose; flowers 3.5 mm. long, a little longer than 
the pedicels; style 6 mm. long, the stigma distinctly clavate. 
Loreto: Rio Itaya, Williams 114 (type). 

Roupala dolichopoda Diels, Bot. Jahrb. 54: Beibl. 117: 35. 
1916. 

A shrub of 2 meters; petioles 4-9.5 cm. long, slightly pilose 
basally; leaves coriaceous, finally glabrous, ovate, short-acuminate, 
nearly truncate at the base, appressed callous-serrate, 10-15 cm. 
long, 6.5-11 cm. wide; racemes sessile, rusty-pilose, becoming gla- 
brate, about 10 cm. long; pedicels 1.5-2.5 cm. long; perianth yellow- 
ish, rusty-pilose, 7-8 mm. long; ovary rusty-tomentose, the glabrous 
style obovate apically. Allied by the author with R. complicate/,. 
Neg. 11752. 

Piura: Above Ayavaca, 1,700 meters, Weberbauer 6364 (type). 

Roupala ferruginea HBK. Nov. Gen. & Sp. 2: 152. 1817. 

Apparently too near R. complicate, HBK., but the leaves obovate- 
oblong, entire, the costa beneath pubescent, even in age, otherwise 
glabrous, reticulate-veined, about 7 cm. long; stigma clavate. 
A shrub of 2 meters. 

Huanuco: Mito, 2,550 meters, 3447 (det. Johnston). Caja- 
marca(?): Chamay, Bracamoros (Humboldt, type). 

Roupala monosperma (R. & P.) I. M. Johnston, Contr. Gray 
Herb. 73: 42. 1924. Embothrium monospermum R. & P. Fl. 1: 63. 
pi. 98. 1798. R. peruviana R. Br. Trans. Linn. Soc. 10: 192. 1811. 

Branchlet tips, leaves, especially beneath, and racemes densely 
pubescent with a more or less evanescent, nearly bright red or red- 
brown tomentum; petioles 2-2.5 cm. long; leaf blades glabrous above 
in age, obscurely toothed, somewhat obliquely elliptic, shortly acute 
at both ends, finally about 10 cm. long and 6 cm. wide; racemes 
10 cm. long; pedicels 1-2 mm. long; flowers glabrate, 6 mm. long, 
the tube about three times longer than the limb; fruit 3 cm. long, 
long-necked at the base, acute. Flowers greenish yellow or yellow 
(Weberbauer). R. Varelana Diels (named for the Director of the 



374 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Madrid Bot. Garden), from Ecuador by Ruiz and Pavon, differs 
in its smaller, oblong-oval leaves and congested racemes, 3-5 cm. 
long. Negs. 11759, 27824. 

Junin: Huacapistana, 2,000 meters, Weberbauer 2168; 246. 
Mantaro Valley, 2,500 meters, Weberbauer 6587. Huanuco: Pam- 
payacu, Sawada P. 28. Near Panao, Ruiz & Pavon. "Paco-paco 
de la sierra." 

Roupala pinnata (R. & P.) Diels in herb., comb. nov. Embo- 
thrium pinnatum R. & P. Fl. 1 : 163. pi. 97. 1798. R. diversifolia R. 
Br. Trans. Linn. Soc. 10: 193. 1810. 

Branchlets, racemes, and younger leaves beneath more or less 
rusty-tomentose; petioles 1.5 cm. long, rarely 2.5 cm.; leaves of 
young shoots more or less coarsely serrate or lobed or even pinnate, 
these with 5 pairs of leaflets, softly pubescent on both sides, strongly 
oblique, the twice larger terminal one to 8 cm. long and 4 cm. wide; 
normal leaves coarsely serrate, oval or roundish, almost equally 
attenuate at the base, sharply acute or cuspidately acuminate, about 
6 cm. long and 4 cm. wide, lustrous above; racemes 10 cm. long; 
pedicels 3-5 mm. long; fruits short-necked at the base, acute, 2.5 cm. 
long. A compact tree of 5 meters (my specimen) with creamy 
white flowers. Bark fetid (Ruiz & Pavon). Leaves of the Jelski 
specimen very heavy, nearly round. Negs. 11765, 27822. 

Huanuco: Muna, Ruiz & Pavdn; at 2,100 meters, 3948. Posuso, 
1,200 meters, Weberbauer 6777. Cajamarca : Cutervo, Jelski 97(1}. 
"Paco-paco." 

Roupala Raimondii Macbr. Field Mus. Bot. 11: 65. 1931. 

Glabrous except the slender, densely leafy branchlet tips, these 
evanescently reddish-pubescent; petioles to 4 cm. long; leaves finally 
20-25 cm. long, the 4-7 pairs of ovate-lanceolate leaflets strongly 
unequal at the entire, cuneate base, medially spinescent-serrate, 
apically entire, long-acuminate, to 14 cm. long and 4 cm. wide, 
often smaller or the terminal a little larger; veins and veinlets 
obscurely and laxly reticulate on both sides; upper leaf surface 
slightly lustrous, the lower opaque, reddish; teeth mostly 4 mm. 
long, 2.5 mm. broad at the base, a few larger, all straight or nearly 
so, apiculate, the sinus acute. Type sterile and possibly, but not 
at all certainly, the juvenile leaves of R. complicata or some other 
species, but those of R. complicata (rarely pinnate) are not at all 
spinescent-serrate, the teeth short. Neg. 11786. 



FLORA OF PERU 375 

San Martin: Crown of sierras, Tarapoto, 750 meters, Williams 
5997 (type). Between Moyobamba and Tarapoto, Raimondi. 

Roupala spicata Baehni, sp. nov. 

Frutex vel arbor; ramuli petiolique leviter puberulo-tomentosi. 
Laminae (in sicco conduplicatae) ovatae, basi et apice acutae, margine 
indistincte dentatae, supra glabrescentes, subtus puberulae. Spicae 
axillares, ferrugineo-pilosae. Flores sessiles; sepala semper ad basin 
libera et medio coalescentia; antherae filamenta brevia; stylus glaber 
apice clavatus, ovarium uniloculare (semina 2) lanatum, glandulae 
hypogynae 4, liberae. Petioli 1.5-2.5 cm. longi; laminae 4-5 cm. 
latae, 5-6 cm. longae. Perianthium 8 mm. longum. Neg. 29570. 

Branches, petioles, and leaves beneath lightly puberulent- 
tomentose; petiole 1.5-2.5 cm. long; leaf blades broadly ovate, shortly 
acute at base and apex, obscurely dentate, 4-5 cm. broad, 5-6 cm. 
long; spikes rusty-pilose, 2-3 times longer than the leaves; perianth 
segments coalescent at the middle, 8 mm. long; anthers subsessile; 
style clavate. Dr. Charles Baehni of the Conservatoire Botanique, 
Geneva, has studied this plant, which had been determined, obvi- 
ously in error, as R. cordifolia HBK. with pedicellate flowers (as all 
other species!). 

Cajamarca: Huambos, 2,300-2,500 meters, Weberbauer 1+192, 
type in Cons. Bot. Geneva. 

51. LORANTHACEAE. Mistletoe Family 

Reference: Engler & Krause, Pflanzenfam. ed. 2. 16b: 98-203. 
1935. 

Traditionally known for the romantically and economically 
important mistletoe, this family in recent times has acquired addi- 
tional interest in the discovery that it contains a drug that affects 
blood pressure (see Gautier, Semaine Me"dicale 513. 1907) and rubber 
in important quantities, this from several members including 
Peruvian species (see Warburg, Tropenpflanzer 9: 633. 1905 and 
Engler & Krause, op. cit. 131 for additional references on both these 
subjects). Many of the Peruvian species, notably in the genera 
Phrygilanthus, Psittacanthus, and Gaiadendron, supply in flower 
brilliant patches of color in the green mass of vegetation along 
forest trails or river banks open to the sun. 

The family is very closely knit. The forms with a calyx or 
calyculus have been sorted into two groups, one comprising those 
with a 1-celled ovary, the other those with the ovary 2-several- 
celled. The segregation of these main divisions is based principally 



376 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

on the presence or absence of endosperm in the seed, supplemented 
by stamen characters. These characters occur in both Old and New 
World forms and, significantly, are not associated concomitantly or 
with other diagnostic features, not repetitive in some other com- 
bination. It may be questioned, therefore, whether they show a 
common origin and indicate a natural relationship; the genera 
resulting do not suggest it. Perhaps there have been similar parallel 
developments affording different combinations of the same characters 
in this very homogeneous family. However this may be, it seems 
possible to recognize the recently accepted genera so far as Peruvian 
species are concerned, but if one were to include the genus Loranthus 
L., only since Eichler restricted to the Old World, it would be appar- 
ent that only the presence of endosperm distinguishes the Peruvian 
group Aetanthus; if this establishes correctly a precedent, taxonomy 
of a family may be chiefly chemical. As a practical matter and as a 
matter of fact(!), there is a single highly natural genus, Loranthus 
L., world-wide in distribution, divisible into sections that may indi- 
cate the theoretical relationship of the species and groups of species. 
Those who believe that the presence or absence of endosperm is 
fundamental may thus express their judgment without depriving 
the phytogeographer, ecologist, pharmacist, traveler, to mention a 
few other than the professional taxonomist, of a group name for a 
group obviously and reasonably self-contained in nature. The 
extremes to which segregation must be carried logically have been 
shown recently by Danser. It is noteworthy that the segregate 
genera have been maintained mostly on characters, such as endo- 
sperm and ovary cells, completely different from those on which 
their authors founded them. 

In the following treatment the receptacle and its more or less 
developed edge are called the calyx, for which there is good prece- 
dent, inasmuch as it is unknown whether the edge of this structure is 
a reduced calyx or the edge of the blossom axil (receptacle); cf. 
Engler & Krause, op. cit. 119. It may be remarked that those who 
use the word calyculus for this edge, in practice nearly always apply 
the name to the entire structure, as evidenced in giving the length ; 
the term calyx, therefore, will be understood by everyone. The 
bracteal development that results in a cupula, so called, more or less 
enclosing the calyx and ovary, is mentioned here simply as cup. 

Flowers minute, more or less immersed in a fleshy rachis or borne in 
axillary clusters and strobiles, then sometimes tiny; calyx 
obsolete, except more or less evident in Oryctanthus. 



FLORA OF PERU 377 

Flowers clustered and strobilate. 
Leaves alternate. 

Filaments elongate, free 1. Antidaphne. 

Filaments short, mostly united to the perianth segments. 

2. Eremolepis. 

Leaves opposite 3. Lepidoceras. 

Flowers spicate. 

Flower parts 3; calyx wanting; spikes articulate. 
Anther cells confluent, 1-pored ; leaves often small or wanting. 

4. Dendrophthora. 

Anthers 2-celled, dehiscing longitudinally; leaves well de- 
veloped 5. Phorandendron. 

Flower parts 6; calyx present; spikes not articulate. 

6. Oryctanthus. 

Flowers small to very large, or rarely tiny but never inserted in the 
rachis; calyx present. 

Anthers basifixed; ovary 1-celled; endosperm lacking. 

7. Aetanthus. 
Anthers versatile; ovary 1-several-celled. 

Ovary 2-several-celled ; endosperm ruminate; leaves more or less 
punctate beneath; flowers in 3's, each calyx subtended by 
a persistent, spreading bractlet 8. Gaiadendron. 

Ovary 1-celled; characters other than above, at least in part. 
Flowers showy, usually 1 cm. long to much longer. 
Endosperm present; calyx not enclosed in a cup but if 
obviously bracteolate (bractlets often promptly cadu- 
cous or minute), the bractlet sometimes foliaceous 
but the flowers then not ternate. . .9. Phrygilanthus. 

Endosperm lacking; each calyx more or less enclosed in a 
cup, this sometimes shallow, or the calyces, borne in 
3's, often subtended by one concave bractlet, this 
sometimes foliaceous 10. Psittacanthus. 

Flowers small, rarely 6 mm. long. 
Crowded inflorescences basally white-bracteolate. 

11. Peristethium. 

Open or simple inflorescences not white-bracteolate. 

12. Struthanthus. 



378 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. ANTIDAPHNE P. & E. 

A small shrub with alternate, roundish-obovate leaves and small, 
clustered, strobile-like spikes, their imbricate bracts caducous at 
anthesis. Staminate flowers apetalous; perianth segments of the 
pistillate flower 3. The Peruvian species grows especially on 
Lauraceae. 

Antidaphne viscoidea P. & E. Nov. Gen. & Sp. 2: 70. pi. 199. 
1838. 

Leaves on flowering branchlets obovate, about 5 cm. long, 3 
cm. wide, the terminal and those on older branches rotund-obovate, 
to 6 cm. wide and 9 cm. long, the reticulate venation prominent; 
flowers cream-colored. Weberbauer found it between 2,700 and 
3,000 meters. Neg. 11822. 

Cajamarca: San Miguel, Weberbauer 3929; 258. Chugur, Weber- 
bauer 4099; 259. San Martin: Zepelacio, 1,200-1,600 meters, Klug 
3594.. Huanuco: Cochero and Pampayacu, Poeppig 1308. Bolivia 
to Colombia. 

2. EREMOLEPIS Griseb. 

Shrubs allied to Antidaphne, but the perianth segments often 
4 and present also in the staminate flowers. Leaves 3-5-nerved. 
No Peruvian specimens found cited, but in the Pflanzenfamilien 
it is stated that there are 6 species in "Chile, Peru, Brazil, and 
Cuba." Perhaps most to be expected is 

Eremolepis punctulata (Clos) Griseb. emend. Benth. in Benth. 
& Hook. f. Gen. PI. 3: 215. 1880. Lepidoceras punctulata Clos in 
Gay, Fl. Chile 3: 165. 1847. 

Pistillate and staminate flowers on separate plants; perianth 
segments deciduous from the fruit. E. Schottii (Eichl.) Engler has 
the two sorts of flowers on different branches of the same shrub and 
perianth segments persistent; E. Glazioui (van Tiegh.) Engler has 
both sorts of flowers together, the perianth segments concrete with 
the pistil, and fruits borne at the base of short, leafy branchlets. 

Peru: (Possibly). Chile. 

3. LEPIDOCERAS Hook. f. 

Much branched shrubs with opposite leaves and axillary racemes, 
the tiny flowers dioecious. Unlike the two preceding related genera, 
the seeds are without endosperm. Grows on Myrtaceae. Author- 
itatively (see Pflanzenfam.) but surely not unquestionably Peruvian. 



FLORA OF PERU 379 

Lepidoceras Dombeyi Hook. f. Fl. Ant. 2: 293. 1846. 

Branches very slender; leaves elliptic-obovate, typically sharply 
apiculate; staminate flowers closely bracteate, the pistillate bracts 
soon foliaceous. Isern 2136, "Chanchamayo," but without original 
label, the leaves obtuse or mucronulate, is L. Kingii Hook. f. As 
Chanchamayo, Peru, is a region totally different phytogeographically 
from the known range of the genus, the Isern locality is almost 
surely an error. 

4. DENDROPHTHORA Eichl. 

Leafy plants, similar in appearance to Phoradendron, from which 
they must be distinguished as indicated in the key, except that in 
general they are less robust or smaller, with the flowers in single rows 
on each side of the rachis. 

Leaves developed. 

Leaves mostly or all of them 2 cm. long or longer. 
Petioles obvious. 
Leaves 4-6 cm. long. 
Petioles 1-2 cm. long; pistillate spikes 4-5 cm. long. 

D. leucocarpa. 

Petioles 1 cm. long; pistillate spikes 2 cm. long. .D. nodosa. 
Leaves 1.5-4 cm. long. 
Internodes terete; staminate flowers 50-100. 

D. hexasticha. 

Internodes (upper) compressed; staminate flowers rarely 60. 
Dioecious; petioles 5-12 mm. long. . . .D. chrysostachya. 

Monoecious; petioles 1.5-2.5 mm. long D. clavata. 

Petioles lacking; leaves linear D. linearifolia. 

Leaves, or at least most of them, 1.5 cm. long or shorter. 
Petioles obvious. 

Leaves acuminate; flowers monoecious D. Negeriana. 

Leaves obtuse; flowers dioecious. 

Leaves linear-spatulate D. Urbaniana. 

Leaves obovate-orbicular D. crassuloides. 

Petioles obsolete. 
Leaves 6-14 mm. long, nerveless. 

Leaves 6-10 (15) mm. long; spikes pistillate above, stam- 
inate below . . D. mesembryanthemifolia. 



380 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves 8-14 mm. long; spikes entirely staminate. 

D. ferruginea. 

Leaves 5 mm. long, the lateral nerves obvious on both sides. 

D. ramosa. 

Leaves wanting or reduced to scales. 

Branches terete or quadrate; flowers 4-seriate. .D. fasciculata. 
Branches complanate; flowers 2-seriate D. Pavoni. 

Dendrophthora chrysostachya (Presl) Urban, Ber. Deutsch. 
Bot. Ges. 14: 285. 1896; 68. Viscum chrysostachyum Presl, Epim. 
Bot. 254. 1849. V. globuliflorum Presl, op. cit. 134. Phoradendron 
chrysostachyum Eichl. in Mart. Fl. Bras. 5, pt. 2: 107. 1868. 

Dioecious; internodes 2-5 cm. long; petioles 1.5-2.5 mm. long; 
leaves elliptic-oblong, obtuse, narrowed at the base, 2-4 cm. long, 
7-20 mm. broad, basally 3-5-nerved; staminate spikes pedunculate, 
4-7 mm. long; fruit globose, nearly smooth. 

Huanuco: (Haenke). Muna, 2,100 meters, 3950. Vilcabamba, 
1,800 meters, 5135. 

Dendrophthora clavata (Benth.) Urban, Ber. Deutsch. Bot. 
Ges. 14: 285. 1896. Viscum clavatum Benth. PI. Hartw. 189. 1845. 
Phoradendron clavatum Eichl. in Mart. Fl. Bras. 5, pt. 2: 107. 1868. 

Allied to D. crassuloides but monoecious; leaves 1.5-4 cm. long, 
7-20 mm. wide; peduncles 5-15 mm. long; flowers 6-7-seriate in 
1-3- jointed spikes 0.5-2 cm. long, the lower staminate joint 24-66- 
flowered, the upper pistillate 18-42-flowered. Apparently not col- 
lected in Peru, but certainly occurring there. Neg. 29151. 

Peru: (Probably). Bolivia to Ecuador and Venezuela. 

Dendrophthora crassuloides (Presl) Urban, Ber. Deutsch. 
Bot. Ges. 14: 285. 1896; 62. Viscum crassuloides Presl, Epim. Bot. 
252. 1849. Phoradendron crassuloides Eichl. in Mart. Fl. Bras. 5, 
pt. 2: 107. 1868. 

Internodes soon terete, very shortly pilose, 2.5-5 cm. long; 
petioles 3-5 mm. long; leaves rounded at the apex, gradually nar- 
rowed to the petiole, 1-1.5 cm. long, nearly as broad, basally incon- 
spicuously 3-5-nerved; pistillate spikes to 1 cm. long, 1-2-articulate, 
the 4-8 flowers 4-5-seriate; peduncles 4-10 mm. long. Closely 
allied is D. portulacoides (Presl) Urban, with subcordate leaves and 
staminate spikes with 50-120 flowers in each joint, about 10-seriate. 

Huanuco: (Haenke). Puno: Sandia, Weberbauer 737, 6124; 240- 



FLORA OF PERU 381 

Dendrophthora fasciculata Patsch. Bot. Jahrb. 45: 441. 1911. 

Densely and fasciculately branched; spikes fasciculate, 4-5 cm. 
long, 5-7-articulate, the superior flowers often staminate, the lower 
pistillate, 8-12 in each joint; fruit white, to 1 cm. long, 3 mm. 
thick. On Euphorbia. 

Ancash: Prov. Huari, 2,700-2,800 meters, Weberbauer 3300 (type). 

Dendrophthora ferruginea Patsch. Bot. Jahrb. 45: 441. 1911. 

A slender, branching shrub, the internodes 2-2.5 cm. long; 
leaves linear, acuminate, 2 mm. wide; spikes solitary, 3-9 mm. 
long, 1-2-articulate, the peduncle 5 mm. long; flowers dioecious, 
ferruginous, 4-seriate, about 20 in each joint. 

Huanuco: Monz6n, 3,300-3,500 meters, Weberbauer 3382 (type). 

Dendrophthora hexasticha van Tiegh. Bull. Soc. Bot. France 
43: 182. 1896; 67. 

Similar to D. chrysostachya; young branchlets papillose-scabrous; 
petioles 3-10 mm. long; staminate spikes with 3-4 joints, the lower 
6-seriate, sometimes 10-seriate, with 50-100 flowers, the pistillate 
2-3-articulate, 6-seriate, the flowers 20-36. On Vaccinium. Neg. 
29152. 

Puno: Tabma,Lechlerl927. Without locality: Dombey. 

Dendrophthora leucocarpa (Patsch.) Trel. Gen. Phor. 218. 
1916. Phoradendron leucocarpum Patsch. Bot. Jahrb. 45: 438. 1911. 

A papillose-scabrous shrub, similar to D. nodosa; internodes 
3-5 cm. long; leaves 2-2.5 cm. broad; pistillate spikes 5-7-jointed, 
each joint 30-34-flowered. Neg. 18189. 

Junin: Huacapistana, 2,500-3,100 meters, Weberbauer 2193, 2439. 

Dendrophthora linearifolia Patsch. Bot. Jahrb. 45: 440. 1911. 

Glabrous, the branches terete or somewhat angled, the internodes 
3 cm. long; leaves obtuse, 3-5.5 cm. long, 4 mm. broad; spikes soli- 
tary, sessile, not articulate, 5 mm. long, the upper flowers staminate, 
the lower pistillate, 4-seriate ; fruit white, 5 mm. long, 3 mm. thick. 
On Croton. 

Puno: Sandia, at 2,100-2,300 meters, Weberbauer 539 (type); 238. 

Dendrophthora mesembryanthemifolia Urban, Ber; Deutsch. 
Bot. Ges. 14: 285. 1896; 70. 

Monoecious, the younger branches definitely striate-pilose and, 
like the leaf margins, minutely papillose-pilose; leaves oblong- or 



382 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

linear-spatulate, the tip acute, recurved or obtuse and mucronulate, 
gradually narrowed to the base, 1-3 mm. broad; spikes mostly 2- 
jointed, the joints 3-12 mm. long, 6-seriate, 8-50-flowered. On Salvia. 
Cuzco: Valle del Apurimac, Herrera. Cuyocuyo, Weberbauer 868 
(det. Patschovsky). Pillahuata, 2,800 meters, Pennell 14098. Mo- 
llepata, Herrera 1228 (det. Patschovsky). Puno: Agapata, Lechler 
1893. Ayacucho: Totorabamba, 3,500 meters, Weberbauer 5465. 
Huanuco: Tambo de Vaca, 3,900 meters, 4383. Mito, 2,700 meters, 
1606. Cani, 2,550 meters, 3477. 

Dendrophthora Negeriana Patsch. Bot. Jahrb. 45: 440. 1911. 

A slender shrub, the younger branches compressed, somewhat 
scabrous, the internodes 3 cm. long; leaves lance-ovate, narrowed 
to the (2 mm. long) petiole, 8-15 mm. long, 3-5 mm. broad; spikes 
long-peduncled, solitary, 2-5 cm. long, 3-articulate, the flowers in 
each spike ferruginous, 2-seriate, 2-8 in each joint. 

Junin: Huacapistana, 2,500 meters, Weberbauer 2138 (type). 

Dendrophthora nodosa Patsch. Bot. Jahrb. 45: 439. 1911. 

Branches slender, papillose-scabrous, the internodes 2-2.5 cm. 
long; leaves lanceolate, obtuse, narrowed to the petiole, prominently 
nerved, 1-2 cm. broad; spikes solitary, the very slender staminate 
ones 4 cm. long, 3-articulate, on a peduncle 6 mm. long; flowers 
dioecious, 6-seriate, about 42 to each joint; pistillate spikes 2-articu- 
late, the 2-5-seriate flowers 4-15 in each joint. 

Puno: Toward Chunchosmayo, 1,800-2,600 meters, Weberbauer 
1078 (type). 

Dendrophthora Pavoni van Tiegh. Bull. Soc. Bot. France 43: 
182. 1896; 71. 

Leafless; flowers 1-2-seriate. Allied by the author to D. Man- 
cinellae Eichl. of Cuba. D. Poeppigii van Tiegh., of the upper 
Amazon, is monoecious, the 2-5-jointed spikes staminate above, 
pistillate below, the lower joint 4-6-flowered. 

Peru, the locality unknown. 

Dendrophthora ramosa Patsch. Bot. Jahrb. 45: 440. 1911. 

A strongly branched shrub, the internodes 2 cm. long; leaves 
spatulate, narrowed toward the tip, 2 mm. wide; spikes solitary, 
the peduncle 10-13 mm. long, with 1 or rarely 2 joints 10-27 mm. 
long; staminate flowers ferruginous, 4-seriate, 40-82 on each joint. 

Huanuco: Near Monzon, 3,400 meters, Weberbauer 3369 (type). 



FLORA OF PERU 383 

Dendrophthora Urbaniana Patsch. Bot. Jahrb. 45: 440. 1911. 

A slender, much branched, often pilose shrub, drying yellow; 
branches terete, the internodes 2-3.5 cm. long; leaves 8-14 mm. 
long, 3 mm. broad, rarely pilose, finely nerved, narrowed to the 
(3 mm. long) petiole; spikes mostly solitary, the staminate 1-2 
cm. long, the 2-seriate flowers 12-18 in the 1-2 joints; peduncle 
3-5 mm. long; pistillate spikes 7 mm. long, each joint 4-flowered; 
fruit globose, drying yellow, plicate-rugose, 4 mm. long, 3 mm. thick. 
Neg. 27825. 

Amazonas: Near Chachapoyas, 2,400 meters, Weberbauer 4404, 
4388; 263. 

5. PHORADENDRON Nutt. 

Reference: Trelease, Gen. Phorad. 1916. 

The key characters may serve to identify this well known genus. 
It contains species that are harmful on account of their parasitic 
habit on useful plants, notably in the Amazonian region on planta- 
tions of young Hevea rubber trees (Warburg). The following com- 
pilation is almost wholly drawn from the monograph by Trelease. 
Three new species by him, kindly communicated by Mr. Ellsworth 
P. Killip, are included, for which I have supplied Latin diagnoses. 

Scales present only on the basal joint of each branch. 
Leaves 2-5 mm. wide, 1-3 cm. long. 

Leaves 3 cm. long, 3-nerved P. Ernestianum. 

Leaves 1 cm. long, enervose P. virgatum. 

Leaves about 1 cm. wide, 12-13 cm. long P. angustifolium. 

Leaves much broader. 
Leaves basinerved. 

Nerves fine; leaves rather thin P. semiteres. 

Nerves coarse; leaves fleshy P. obliqua. 

Leaves pinnately nerved. 

Spikes scarcely 2 cm. long, very slender P. Mathewsii. 

Spikes mostly longer, in any case rather stout. 

Spikes clustered, at least at some nodes . . P. Englerianum. 

Spikes solitary P. peruvianum. 

Scales present on all nodes. 

Stems continuous or prevailingly not branching at all nodes; 
flowers in 4-10 series. 



384 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves obscurely pinnate- veined beneath; flowers 4 (-6). 

P. piperoides. 

Leaves basinerved; flowers in (4-) 6 or more series. 
Flowers in 6 or fewer series, the spikes 2-5 cm. long. 
Spikes often clustered; leaves rather obscurely nerved. 
Leaves broadly elliptic, 3-10 cm. wide; scales several. 

P. crassifolium. 

Leaves nearly orbicular, 7-12 cm. wide; scales a single pair. 

P. Albert-Smithii. 

Spikes mostly or all solitary; leaves heavily nerved beneath. 
Scales a solitary pair; leaves 5-7 cm. wide. 

P. Urbanianum. 

Scales several; leaves 3-5.5 cm. wide. . . P. ayacuchanum. 
Flowers in 8(6)-10 series, the spikes about 6 cm. long. 
Leaf nerves obvious at the base; scales about 1 cm. above 

the node P. Lindavianum. 

Leaf nerves completely lacking; scales nodal. P. Macbridei. 

Stems normally forking at each node; flowers in 2-4 series; leaves 

basally nerved P. huallagense. 

Phoradendron Albert-Smithii Trelease, sp. nov. 

Internodiis gracilibus, 3-4 mm. crassis, 15 cm. longis, teretibus; 
vaginis cataphyllaribus 2 ad omnia internodia, paullo supra basin 
abeuntibus; foliis 7-12 cm. latis fere rotundatis, ad basin longe 
(1 cm.) angustatis, vix coriaceis basinerviis, nervis tenuibus; 
spicis plerumque 3-5.3 cm. longis, 5-articulatis; floribus circa 22 in 
quovis articulo in seriebus 4 (2)-6 dispositis. Aequatoriales- 
Percurrentes. 

Glabrous, the long, slender internodes 3-4 mm. thick, 15 cm. 
long, terete, with an annular pair of cataphylls shortly above the 
base of each; leaves 7-12 cm. broad, drying rather thin, dull green 
and finely basinerved on both sides, cuneately tapered at the base 
for about 1 cm.; spikes yellow, before flowering 3 cm. long, with 
about 5 joints, these about 22-flowered, the flowers in 4 (2) or 6 series. 

Ayacucho: Aina, 750-1,000 meters (Killip & Smith 11598; 
type in U. S. Nat. Herb.). 

Phoradendron angustifolium (HBK.) Eichl. in Mart. Fl. 
Bras. 5, pt. 2: 115. 1868; 65. Loranthus angustifolius HBK. Nov. 
Gen. & Sp. 3: 442. 1820. 



FLORA OF PERU 385 

Not forked, the rather thin, basally finely nerved leaves about 
10 cm. long; scales a single pair 5-10 mm. above the base of the 
joint, spreading; spikes somewhat clustered, 2 cm. long, slender, 
with 3-6 oblong joints, these 30-40-flowered, the pistillate flowers 
in about 4 series; peduncle 4-6 mm. long, the scales narrowly white- 
margined; fruit 4 cm. thick, smooth, the sepals inflexed. Allied is 
P. parietioides Trel., to which Tessmann 4880 from the Maranon 
may possibly belong; it has somewhat falcate, finely nerved, veiny 
leaves 1.5-2 cm. wide; fruit ellipsoid, 3 mm. thick, 4.5 mm. long, the 
sepals spreading. Illustrated, Trelease pi. 79. 

Cajamarca(?) : Olleras to Mt. Aipate (Bonpland 3508). With- 
out locality: Pavon (scales ciliate; pistillate spikes 30 mm. long, 
about 12-flowered at the end of the joint). Bolivia. 

Phoradendron ayacuchanum Trelease, sp. nov. 

Internodiis 3-4 mm. latis 13 cm. longis teretibus sed versus nodos 
paullo dilatatis; vaginis cataphyllaribus ad omnia internodia, 2 ad 
infimum internodium tantum obviis, 3 vel 4 alteris supra basin 
insertis; foliis falcato-lanceolatis, basi apiceque cuneato-angustatis, 
obtusis, 3-5.5 cm. latis, 15-17 cm. longis, nervis e basi conspicuis; 
spicis plerumque solitariis 2 cm. longis 4-5-articulatis, floribus 
4-seriatis circa 10 in quovis articulo; perianthio clause. Aequa- 
toriales-Crassifoliae. 

Glabrous, dark green, the leaves drying coriaceous and dull 
brown; internodes 3-4 mm. thick, 13 cm. long or longer, terete 
except at the slightly flattened nodes; cataphylls a sterile basal pair 
and 3 or 4 subequally spaced, fertile pairs between each 2 leaf nodes, 
broad and pointed; leaves falcately lanceolate, gradually blunt- 
acuminate, 15-17 cm. long, cuneately subsessile, heavily basinerved; 
spikes subsolitary, 2 cm. long, with 4 or 5 round-turbinate joints, 
these about 10-flowered, the flowers in 4 (2) series; sepals closed. 

Ayacucho: Aina, 750-1,000 meters (Killip & Smith 22713; type 
in U. S. Nat. Herb.). 

Phoradendron crassifolium Eichl. in Mart. Fl. Bras. 5, pt. 
2: 125. pi 48. 1868; 144. 

Occasionally pseudodichotomous; scales several pairs, only the 
upper (often soon deciduous) fertile; leaves very thick and dull, 
more or less lanceolately or elliptically ovate, obtuse or bluntly 
pointed, 3-10 cm. wide, 8-16 cm. long, rounded or attenuate at the 
base; spikes occasionally forming a compound, terminal inflorescence; 



386 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

joints about 5, scarcely 10-flowered; peduncle 5 mm. long, often 
with several pairs of crowded scales; fruit yellowish, slightly granular, 
4 mm. thick, the sepals closely inflexed. The Mathews specimen is 
ascribed by Trelease to "Martens," presumably an error. Illus- 
trated, Trelease pi. 213-214. 

San Martin: Tarapoto, Williams 6126, 6588. Moyobamba 
(Mathews 1622; leaves 9 cm. wide, 16 cm. long, acuminate). 
Junin : Chanchamayo Valley, 1,500 meters, Schunke 215. La Merced, 
600 meters, 5531; Weberbauer 1860; 283. Cuzco: Valle de Santa 
Ana, Herrera 992. Loreto: Mishuyacu, 100 meters, Klug 420, 42. 
Bolivia to Central America and the West Indies. 

Phoradendron Englerianum Patsch. Bot. Jahrb. 45: 439. 
1911; 133. 

Similar to P. peruvianum, the internodes sometimes somewhat 
quadrangular; scales often 2 or even 3 pairs, within 10 mm. of the 
base; leaves 2.5-6 cm. wide, 9-15 cm. long, cuneate to the petiole, 
this 5 cm. long; spikes more or less clustered, subsessile. Illustrated, 
Trelease pi. 196. 

Junin: Tarma, Weberbauer 1903, 1904; 283. 

Phoradendron Ernestianum Patsch. Bot. Jahrb. 45: 439. 
1911; 121. 

Somewhat pseudodichotomous, the upper internodes rhombically 
4-angled; scales a single pair, nearly basal; leaves lanceolate, acute, 
3-nerved, cuneately subsessile; spikes mostly solitary, subsessile, 
10-25 mm. long, the 3-5 rounded joints about 6-flowered, the 
flowers in about 4 series; scales scarcely ciliate; fruit granular, 
nearly 3 mm. thick, the erect sepals separated. Illustrated, Tre- 
lease pi. 177. 

Cajamarca: Balsas to Celedin, Weberbauer 4251. 

Phoradendron huallagense Ule, Verh. Bot. Ver. Brandenb. 48: 
158. 1906; 155. 

Cymosely dichotomous; scales a solitary, nearly basal pair; 
leaves ovate, lustrous, mucronate, acuminate to very obtuse, 2-2.5 
cm. wide, 5-6 cm. long, cuneately subsessile; spikes solitary, 1 to 
finally 3 cm. long, with about 4 short, 4-flowered joints; peduncle 
scarcely 2 cm. long; fruit white, granular, roughened, 2 mm. thick, 
4 mm. long, the slightly parted sepals erect. Illustrated, Trelease 
pi. 267. 



FLORA OF PERU 387 

Loreto: Huallaga, Ule 6664, type. Mishuyacu, 100 meters, 
King 1123. Florida, 180 meters, King 21 75. "Beguefide" (Huitoto 

name). 

Phoradendron Lindavianum Patsch. Bot. Jahrb. 45: 438. 
1911; 155. 

Not dichotomous or scarcely so, the long, thick branches some- 
what flattened below the nodes, the elliptic-ovate leaves drying 
golden yellow; scales one pair, sharply deltoid, keeled, borne about 
1 cm. above the base of the joint; leaves more or less mucronate, 
obtuse to emarginate, 4-9 cm. wide, 10-15 cm. long, decurrent on 
the petiole, this 10 mm. long; joints about 4, often over 100-flowered, 
the granular peduncle 5-10 mm. long; fruit (immature) 3 mm. 
thick, 5 mm. long, the sepals closely inflexed. Illustrated, Trelease 
pi 235. 

Puno: On Aralia, Weberbauer 1288 (type). Huanuco: Cochero, 
Poeppig. 

Phoradendron Macbridei Standl., sp. nov. 

Kami crassissimi subcompressi, internodiis valde elongatis infra 
nodos dilatatis, cataphyllis ad basin internodii infimi tantum insertis; 
folia magna crassissima, petiolis crassissimis vix ultra 6 mm. longis; 
lamina oblonga vel oblanceolato-oblonga 8-15 cm. longa 3.5-7 cm. 
lata, apice late rotundata vel breviter emarginata, basi acuta vel 
basin versus sensim angustata, utrinque profunde dense rugulosa; 
spicae solitariae sessiles circa 4 cm. longae crassae, nodis circa 6, 
floribus numerosis circa hexastichis. Berries white. 

Huanuco: Muna, 2,100 meters, Macbride 3954 (type in Herb. 
Field Mus.). 

Phoradendron Mathewsi Trelease, Gen. Phorad. 133. 1916. 

Allied to P. peruvianum, and similar; leaves somewhat obliquely 
elliptic-lanceolate, 2-4 cm. wide, 16 cm. long, cuneately decurrent on 
the petiole, this less than 1 cm. long; spikes clustered, 1.5-2 cm. long, 
the 3-4 very Blender joints sometimes 8-flowered, the flowers in 
about 4 series; peduncle 2 mm. long; fruit (immature) 2 mm. thick. 
Illustrated, Trelease pi. 197. 

Amazonas: Chachapoyas, Mathews, type. San Martin: San 
Roque, 1,400 meters, Williams 6979. "Pishcuisman." 

Phoradendron obliquum (Presl) Eichl. in Mart. Fl. Bras. 5, 
pt. 2: 134. 1868; 86. Viscum obliquum Presl, Epim. Bot. 225. 1849. 



388 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Thick-stemmed, the nearly basal scales sometimes 2 pairs and 
ordinarily another pair 2-3 cm. higher, tubular; leaves lanceolate, 
often falcate, very obtuse, 4-8 cm. wide, 10-23 cm. long, basally 
attenuate to the thick petiole; spikes 5-6 cm. long, the short, swollen 
joints sometimes 30-40-flowered, the flowers in 4-6 series; peduncle 
stout, scarcely 5 mm. long, with about 3 pairs of scales; fruit smooth, 
3 mm. thick (immature), the sepals closed. Illustrated, Trelease 
pi. 117. 

Huanuco: Huanuco Mountains (Haenke, type). Yanano, 1,800 
meters, 3811. San Martin: San Roque, 1,400 meters, Williams 
7008. Ecuador. 

Phoradendron peruvianum Eichl. in Mart. Fl. Bras. 5, pt. 
2: 123. 1868; 131. 

Scarcely dichotomous, the internodes ancipitally compressed; 
scales a single pair, nearly basal, white-margined; leaves broadly 
lanceolate, sometimes falcate, acute to obtuse, 3-5 cm. wide, 6-12 
cm. long, basally acute, the petiole 5-7 mm. long; spikes solitary, 
finally 4-5 cm. long, the 3-5 stout, oblong joints about 50-flowered, 
the flowers in 6 series; peduncle stout, 3-4 mm. long; fruit wide, 
smooth, 4 mm. thick, the sepals closed. Illustrated, Trelease pi. 193. 

Without locality, Ruiz & Pavdn. Huanuco: Muna, 2,100 
meters, 3970. 

Phoradendron piperoides (HBK.) Trelease, Gen. Phorad. 
145. 1916. Loranthus piperoides HBK. Nov. Gen. & Sp. 3: 443. 1820. 

Rather frequently pseudodichotomous or dichotomous; scales 
1-5 pairs toward the base of the lowest joint or 10-15 mm. above 
the base of the other joints, white-margined; leaves lanceolate to 
round-ovate, mucronately subacute to bluntly acuminate, 2.5-5 
cm. wide, 6-10 cm. long; spikes slender, mostly clustered, usually 
reddish, 3-6 cm. long, with about 6 joints, these 10-15-flowered; 
peduncle 2-3 cm. long; fruit yellow or orange, warty to smooth, 
about 4 mm. thick, 5 mm. long, the ascending sepals slightly parted. 
Illustrated, Trelease pi. 217, 222. 

Loreto: Along Rio Ucayali, Tessmann 3345, 3052. El Recreo, 
200 meters, Williams 3943. Lower Rio Nanay, Williams 615. 
Near mouth of Rio Tigre, 115 meters, Killip & Smith 27523. 
Without locality: Ruiz & Pavdn. Huanuco: Rio Huallaga Canyon, 
1,200 meters, 4246. Yanano, 1,800 meters, 3720. San Martin: 
Rio Mayo, Williams 6284- Juan Guerra, 720 meters, Williams 



FLORA OF PERU 389 

6915. San Roque, 1,400 meters, Williams 6980. Juanjui, 400 
meters, Klug 3868. Argentina to Mexico. "Suelda con suelda." 

Phoradendron platycaulon Eichl. in Mart. Fl. Bras. 5, pt. 2: 
108. pi. 33. 1868. 

Branches with scales on all the joints, strongly compressed and 
much dilated, striate, bright green; leaves oblong or linear-oblong, 
3-5 cm. long, 3-6 mm. wide, obtuse; spikes clustered, 25 mm. long 
or less, the joints about 3, 4-6-flowered. Easily recognized by the 
rather slender and strongly compressed branches. 

Loreto: Caballo-cocha, Williams 2342. Amazonian Brazil and 
French Guiana. 

Phoradendron quadrangulare (HBK.) Krug & Urban, Bot. 
Jahrb. 24: 35. 1898. Loranthus quadrangularis HBK. Nov. Gen. 
&Sp. 3:444. 1820. 

Plants rather slender, the branches conspicuously 4-angulate, 
only the basal joints with cataphylls; leaves narrowly oblong or 
elliptic-oblong, conspicuously petiolate, 4-6 cm. long, obtuse, cune- 
ately narrowed at the base; spikes clustered, 3-4 cm. long, the 3-5 
slender joints 12-26-flowered; berries white, subglobose, 3 mm. in 
diameter, the sepals closely inflexed. Illustrated, Trelease pi. 154, 
155. 

Loreto: Iquitos, 120 meters, Williams 8026. San Martin: 
Tarapoto, 800 meters, Williams 5496, 6307. Ecuador and Colombia. 
"Pishco isman." 

Phoradendron semiteres Trelease, Gen. Phorad. 69. pi. 85. 
1916. 

Somewhat pseudodichotomous; scales a single pair, tubular- 
bifid, scarious-margined ; leaves obliquely or subfalcately lanceolate, 
obtuse, about 3 cm. wide, 10 cm. long, 5-7-nerved, cuneately decur- 
rent for 10-15 mm.; spikes more or less clustered, to 3 cm. long in 
fruit, the usually 3 ellipsoid joints 10-23-flowered, the flowers in 
about 4 series; peduncle 2 mm. long, the scarious-margined scales 
eciliate; fruit red, nearly smooth, 3 mm. thick, the sepals closed. 

Peru: Without locality (Ruiz & Pavori). Bolivia. 

Phoradendron Urbanianum Ule, Verh. Bot. Ver. Brandenb. 
48: 157. 1906; 155. 

Very sparingly forked, drying olive; scales a single pair, about 
5 mm. above the base of the joint, sharply deltoid, keeled; leaves 



390 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

elliptic-lanceolate, mucronately acuminate to obtuse, 5-7 cm. wide, 
8-13 cm. long, cuneately wing-petioled for about 10 cm.; joints 
about 4 and 30-flowered, the slightly granular peduncle 3 cm. thick, 
4 cm. long. Illustrated, Trelease pi. 234. 

Loreto(?): Cerro de Escalera, Vie 6681 (type). 

Phoradendron virgatum Trelease, sp. nov. 

Ramis elongatis, internodiis 2-4 cm. longis teretibus; vaginis 
cataphyllaribus ad basin ramorum, ad internodia caetera nullis; 
foliis oblongis 2 mm. latis 10 mm. longis obtusis sessilibus, nervis 
obsoletis; spicis 1-5 ad 3 cm. longis circa 3-articulatis; floribus in 
articulo 9 vel 11, 4-seriatis; baccis in statu juvenili globosis, sepalis 
inflexis. Aequitoriales-Virgatae. 

Dark green, glabrous, the long, virgate branches with short 
(2-4 cm.), terete internodes; cataphylls (not seen) evidently limited 
to the basal joints; leaves oblong, 2 mm. wide, reflexed, obtuse, sessile, 
drying concave, coriaceous, not evidently nerved; spikes commonly 
clustered, about 3 cm. long, with about 3 swollen joints, these 
9-11-flowered, the flowers in 4 (2) ranks; peduncle 3 mm. long, the 
scales blunt, connate; fruit (immature) globose, with inflexed or 
meeting sepals. 

Junin: Carpapata, 2,400 meters (Killip & Smith 24350; type in 
Herb. Field Mus.). Huanuco: Mito, 2,700 meters, 3343. 

6. ORYCTANTHUS (Griseb.) Eichl. 
Reference: Eichler in Mart. Fl. Bras. 5, pt. 2: 87-92. 1868. 

Plants parasitic on shrubs and trees, usually with aerial roots 
and short, axillary racemes of small flowers, the flowers of the 
Peruvian species perfect. Closely allied to Struthanthus, but the 
calyx throat barely developed and the bractlets greatly reduced or 
obsolete. The genus is well illustrated in Pflanzenfam. ed. 2. 16b: 
174 (0. ruficaulis Eichl.). The species or the forms so recognized 
are defined very indefinitely, even the presence or absence of the 
very minute bractlets, often promptly caducous, being of question- 
able worth as a specific character. 

Leaves broad and somewhat clasping 0. amplexicaulis. 

Leaves broadly ovate to lanceolate, not clasping. 
Spikes pedunculate; leaves often 3 cm. wide or wider. 

Spikes mostly in a terminal raceme; bractlets obvious. 

0. amazonicus. 



FLORA OF PERU 391 

Spikes, at least many of them, axillary. 

Leaves opposite 0. botryostachys. 

Leaves alternate 0. ovalifolius. 

Spikes sessile or subsessile, all axillary; leaves 2-2.5 (-3) cm. wide. 

0. florulentus. 

Oryctanthus aniazonicus Ule, Verb. Bot. Ver. Brandenb. 48: 
152. 1907. 

Similar to 0. botryostachys Eichl., but the spikes in the upper leaf 
axils passing into a terminal raceme 10-25 cm. long, and with 
tooth-like but caducous bractlets present; branchlets, petioles, and 
spikes rusty-red-furfuraceous; leaves ovate, 7-10 cm. long, 3-5 cm. 
broad, nerved and reticulately veined ; flowers crowded, yellowish. 
Doubtfully distinct. Neg. 11818. 

Loreto: Iquitos, Ule 6252 (type). Pisco, Rio Nanay, Williams 
1284. 

Oryctanthus amplexicaulis (HBK.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 88. 1868. Loranthus amplexifolius HBK. Nov. Gen. & Sp. 
3: 445. 1820. 

Nearly glabrous, with subrotund or oval, obsoletely nerved 
leaves several cm. wide that are subamplexicaul ; spikes mostly in 
a terminal raceme, the peduncles 5-12 mm. long, the spikes little 
longer. Well marked by its bluntly rounded, sessile leaves. 

Loreto: San Isidro, Tessmann ^951 (det. Krause). Colombia 
to the Guianas. "Suelda con suelda." 

Oryctanthus botryostachys Eichl. in Mart. Fl. Bras. 5, pt. 
2: 89. pi. 29. 1868. 

Terete branchlets, petioles (2-4 mm. long), and peduncles more 
or less rusty-furfuraceous, becoming glabrate; leaves broadly ovate 
or oblong-ovate, rounded at the apex or rather acuminate, 5-8 cm. 
long, 2.5-5 cm. broad, rounded to the acute juncture with the 
petiole; nervation obvious but faint; spikes in several leaf axils and 
in a terminal raceme several cm. to 10 cm. long or longer; peduncles 
about 1 cm. long, the spikes often twice as long; flowers 2-3 mm. 
distant; bractlets rudimentary, caducous. The Weberbauer plant 
is more than usually reddish and scurfy, and the determination 
perhaps questionable. The name is sometimes written by recent 
authors bothryostachys. 0. spicatus (Jacq.) Eichl. has no (or 
poorly developed) terminal inflorescence, the flowers 4-5 mm. distant. 
Neg. 19045. 



392 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

San Martin: Moyobamba, 1,500 meters, Weberbauer 4766; 
290 (det. Patschovsky). San Roque, Williams 7290. Junin: 
Hacienda Schunke, 1,600 meters, Killip & Smith 24829 (toward 0. 
florulentus). Chanchamayo Valley, Schunke 248.- Loreto: Iquitos, 
on Siparuna, Killip & Smith 27218 (det. Smith). Yurimaguas, 
Williams 3819 (form with subsessile spikes). Contamana, 150 
meters, on Anona, Killip & Smith 26875. Brazil to the Guianas. 
"Pishcu isman," "suelda con suelda," "pishcumicuna." 

Oryctanthus florulentus (Rich.) Urban, Bot. Jahrb. 24: 31. 
1897. Loranthus florulentus Rich. Act. Soc. Hist. Nat. Paris 107. 
1792. L. ruficaulis P. & E. Nov. Gen. & Sp. 2: 61. pi. 185. 1838. 0. 
ruflcaulis Eichl. in Mart. Fl. Bras. 5, pt. 2: 90. pi. 30. 1868. 

Branches more or less compressed-angulate and, with the young 
leaves, reddish-scurfy, becoming glabrous; petioles 3-5 mm. long; 
leaves ovate to oblong-elliptic, usually 3-4.5 cm. long, 2-3 cm. 
broad, most often rounded at each end, or subacute; spikes 1-3 cm. 
long; flowers red, bracteolate. 0. spicatus (Jacq.) Eichl. has been 
mistaken for this species; it resembles more 0. botryostachys, which 
compare. Neg. 21396. 

San Martin: Moyobamba, Weberbauer 4490; 290 (det. Patschov- 
sky). San Roque, Williams 7288. Huanuco: Monzon, Weberbauer 
3545; 256 (det. Patschovsky as 0. spicatus; cf. also 0. botryostachys). 
Loreto: Iquitos, Williams 3596, 1407. Pebas, on Citrus, Williams 
1764- Brazil; Guianas. 

Oryctanthus ovalifolius (R. & P.) Macbr. Field Mus. Bot. 
11: 17. 1931. Loranthus ovalifolius R. & P. Fl. 3: 50. pi. 177. 1802. 
Struthanthus ovalifolius G. Don, Gen. Syst. 3: 411. 1834. 

Simple-stemmed, glabrous, very fleshy; leaves short-petioled, 
oval, about 2.5 cm. long; spikes solitary or geminate, 1 cm. long; 
flowers sessile in the fleshy rachis, crowded; calyx none or minute; 
fruit oval, purplish. Referred by Eichler to 0. botryostachys with a 
query; perhaps a Struthanthus. 

Huanuco: Pillao; Chacahuasi (Ruiz & Pavdn). 

7. AETANTHUS (Eichl.) Engler 

Nodosely jointed, dichotomously branched plants with heavy 
leaves and short, axillary racemes of showy flowers. The group could 
be retained, as by Eichler, in Psittacanthus, but the anthers are very 
slender and basifixed as in Loranthus of the Old World, which it 
resembles except for the lack of endosperm ; see remark at beginning 



FLORA OF PERU 393 

of family. The nearly related Psathyranthus amazonicus Ule, Verb. 
Bot. Ver. Brandenb. 48: 156. pi. 2. 1907, to be expected, has fleshy 
anthers. The following names represent, probably, in reality only 
about three variable species. 

Anthers cross-celled. 

Petioles slender, 1-2.5 cm. long. 
Leaves obovate, long-attenuate at the base, 4-8 cm. wide. 

A. cauliflorus. 

Leaves elliptic-lanceolate, 4-5 cm. wide A. coriaceus. 

Petioles stout, rarely as long as 12 mm. 

Flowers 7.5-10 cm. long; leaves about 6 cm. wide .... A. nodosus. 
Flowers 5-6 cm. long; leaves to 4.5 cm. wide. . .A. subandinus. 
Anthers not cross-celled. 

Leaves 3-4.5 cm. wide, obscurely veined A. ornatus. 

Leaves 5-6 cm. wide, obviously veined A. Paxianus. 

Aetanthus cauliflorus Ule, Verh. Bot. Ver. Brandenb. 48: 153. 

1907. 

Leaves 3-5-verticillate, 12-20 cm. long, long-attenuate to the 
petiole, this 1.5-2 cm. long, often attenuate also to the obtusish 
apex; flowers 6-7 cm. long; filaments sharply papillose. Otherwise 
apparently very similar to A. subandinus. Neg. 3293. 

Loreto: Cerro de Escalera, 1,400 meters, Ule 6908 (type). 

Aetanthus coriaceus Patsch. Bot. Jahrb. 45: 437. 1911. 

Petioles 1.5-2.5 cm. long; leaves obtuse, 7-8.5 cm. long, the lateral 
nerves slender; calyculus 5 mm. long; flowers to 8 cm. long, red below, 
orange above; fruit 2 cm. long, 7 mm. thick. 

Cajamarca: Above San Miguel, Weberbauer 3938 (type); 258. 

Aetanthus nodosus (Desr.) Engler, Pflanzenfam. Nachtr. 3, 
pt. 1: 136. 1897. Loranthus nodosus Desr. in Lam. Encycl. 3: 601. 
1792. L. dichotomus R. & P. Fl. 3: 45. pi. 274. 1802. Psittacanthus 
dichotomus Eichl. in Mart. Fl. Bras. 5, pt. 2: 27. 1868. 

Branches often ternate; petioles short or 10-12 mm. long; leaves 
verticillate, 6-10 cm. long, 4-7 cm. wide, lightly nerved, the lateral 
nerves about 7 ; peduncles about equaling the petioles, 4-6-flowered ; 
flowers tubular, pendulous, red below, yellow above; fruit greenish 
yellow, oval, 2 cm. long, 12 mm. thick. In Herb. Madrid the Ruiz and 
Pavon specimen is identified by Krause as Psittacanthus dichotomus! 



394 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Muna, Ruiz & Pavdn; (Jussieu, type). Pampayacu, 
Sawada 34. Tambo de Vaca, 3,900 meters, 4900. "Mocma," 
"mucma." 

Ae tan thus ornatus Krause, Notizbl. Bot. Gart. Berlin 8: 206. 
1922. 

Petioles 10-13 mm. long; leaves narrowly elliptic or obovate, 
narrowed at the base, 5-8 cm. long; pedicels 5-8 mm. long; flowers 
scarlet, with orange limb, the tube 14-18 mm. long, 3 mm. thick 
at the base to 6 mm. at the apex, the lobes 3.5-4.5 cm. long, 1-1.5 
mm. broad; filaments 14-16 mm. long, the anthers 12-15 mm. long. 
Nearly A. Mutisii (HBK.) Engler, of Ecuador, but the leaves and 
flowers smaller and the petioles a little longer. Neg. 3295. 

Piura: Above Palambla, Weberbauer 6057 (type). 

Aetanthus Paxianus Patsch. Bot. Jahrb. 45: 437. 1911. 

Not apparently distinguishable from A. ornatus except by the 
broader, ovate leaves, 8-10 cm. long, the flowers 6-8.5 cm. long, 
and the filaments 2-2.5 cm. long; fruit ovate, 2 cm. long, 13 mm. 
thick. Neg. 3296. 

Huanuco: Near Monzon, 2,500 meters, Weberbauer 3390 (type); 
255. 

Aetanthus subandinus Ule, Verh. Bot. Ver. Brandenb. 48: 
154. 1907. 

Branches terete, rarely trichotomous; leaves ternate, obovate 
or broadly oblong, 7-9 cm. long, 3.5-4.5 cm. wide, narrowed to the 
(3-5 mm. long) petiole, coriaceous, indistinctly nerved; pedicels 
2-3 mm. long; calyculus 2-3 mm. long; flowers scarlet; filaments 
4 mm. long, epapillose. Neg. 3297. 

San Martin: Cerro de Cumbasa, 1,000 meters, Vie 6318, type. 
Loreto: Mouth of Rio Santiago, Tessmann 4367. Pumayacu, 600- 
1,200 meters, King 3164 (det. Standley). Florida, 200 meters, Klug 
2106; flowers brick-red and yellow. Caballo-cocha, Williams 2072. 
Florida, Klug 2254- "Ucuguifise" (Huitoto Indian name). 

8. GAIADENDRON G. Don 

Typically well marked as indicated in the key; also the original 
species becomes a tree and subsists then independently. However, 
two of the Peruvian species are, so far as known, parasitic shrubs, 
and have been determined in Herb. Madrid by Krause as Phrygi- 
lanthus sp., following Eichler who, however, reduced the genus Gaia- 



FLORA OF PERU 395 

dendron to Phrygilanthus, although Krause himself accepts it; cL 
Pflanzenfam. loc. cit. Dr. Charles Baehni has kindly verified my 
assumption from appearance that these parasitic shrubs must be 
associated with the original Gaiadendron, if the genus is maintained, 
and the transfer of the species is here made for him, with his permis- 
sion. G. Don included Phrygilanthus eugenioides (HBK.) Eichl. 
here on the basis of its punctate leaves and sometimes terrestrial 
habit, but it lacks the persistent bractlets and, presumably, has a 
1-celled ovary. 

Leaves lanceolate, acuminate; flowers white or yellowish white. 

G. Tagua. 

Leaves rounded at the apex or merely acute, often obovate; flowers 
yellow. 

Leaves more or less obovate, rounded or blunt, densely punctate 
beneath; terrestrial trees. 

Bracts not foliaceous. 

Flowers about 2 cm. long; ovary 1-1.75 mm. wide. 

G. puracense. 

Flowers about 3 cm. long; ovary 2 mm. wide. .G. punctatum. 
Bracts foliaceous G. macranthum. 

Leaves not obovate, acutish, with few or obscure punctations; 
parasitic shrubs. 

Leaves lance-oblong, to 7 cm. long, less than half as broad. 

G. lanceolatum. 

Leaves elliptic, mostly 2.5-4 cm. long G. ellipticum. 

Gaiadendron ellipticum (R. & P.) Baehni, comb. nov. 
Loranthus ellipticus R. & P. Fl. 3: 47. pi. 276. 1802. Phrygilanthus 
ellipticus Eichl. in Mart. Fl. Bras. 5, pt. 2: 48. 1868. 

Apparently too similar to P. lanceolatus, but the leaf blades 
somewhat shorter, the longest about 5 cm. long, on petioles 4-6 
mm. long; racemes densely flowered, 3 cm. long. The leaves of both 
species are sparsely punctate beneath. Neg. 29469. 

Huanuco: Posuso, Ruiz & Pavon. Cuzco: Ollantaitambo, 2,800 
meters, Herrera 683. Calca, 3,000 meters, Herrera 2091. Ollan- 
taitambo, 3,000 meters, Pennell 13666. "Matapalo." 

Gaiadendron lanceolatum (R. & P.) Baehni, comb. nov. 
Loranthus lanceolatus R. & P. Fl. 3: 47. pi. 278. 1802. L. lancifolius 



396 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Poir. ex R. & S. Syst. 7: 142. 1829. Phrygilanthus lanceolatus Eichl. 
in Mart. Fl. Bras. 5, pt. 2: 48. 1868. 

A brown-barked shrub with yellow flowers borne ternately in 
axillary and terminal racemes 4-7 cm. long; leaves oblong-lanceolate, 
2-3 cm. broad, slightly lustrous above, punctate and drying brown 
beneath, obsoletely veined; petioles 7-10 mm. long; pedicels 2-3 
mm. long; bracts ovate, 4 mm. long, reflexed, persistent; flowers 
about 12 mm. long, the 7-8 divisions nearly filiform; fruit oblong. 
Neg. 29460. 

Huanuco: Pillao, Ruiz & Pavon (type). 

Gaiadendron macranthum Killip, Journ. Wash. Acad. Sci. 
24: 44. 1934. 

Leaves obovate, 3.5-5 cm. long, 1.5-2.3 cm. wide; bracts similar 
to the leaves, 10-25 mm. long, 4-10 mm. wide, persistent; bractlets 
ovate, 5-7 mm. long, 4-5 mm. wide; flowers in 3's, the lateral short- 
pediceled; calyx cylindric, 4 mm. long, the margin flaring, entire; 
petals 7, orange, 3-3.5 cm. long. Not clearly distinct from G. 
punctatum, but the author separates it on "the large leaf -like bracts, 
larger bractlets, and longer petals." A shrub about 2 meters high 
(Weberbauer). 

Ayacucho: Putis, Choimacota valley, 3,400 meters, Weberbauer 
7523 (type). 

Gaiadendron punctatum (R. & P.) G. Don, Gen. Syst. 3: 
432. 1834. Loranthus punctatus R. & P. Fl. 3: 47. pi. 277. 1802. 
Taguaria punctata Raf. Sylva Tell. 125. 1838. Phrygilanthus punc- 
tatus Eichl. in Mart. Fl. Bras. 5, pt. 2: 48. 1868. 

A tall shrub or small tree with very erect, terete branches; 
petioles 4-5 mm. long; leaf blades oval or somewhat obovate, acutish 
at the base, rounded at the blunt apex, to 7 cm. long and 4 cm. wide, 
slightly lustrous above, dull and densely punctate beneath ; flowers 
in dense, terminal and upper-axillary racemes, these 10 cm. long or 
longer; peduncles 3-4 mm. long, 2-3-flowered, the elliptic bractlets 
7 mm. long; petals 3 cm. long. Often a tall tree that at flowering 
time is brilliant because of the myriads of bright yellow flowers, 
borne so profusely that they seem to clothe all the upper crowded 
branchlets. Petals 6 in the original specimen, but this scarcely con- 
stant. My collection was from a tree more than 20 meters tall. 

Cajamarca: Huambos, 2,600 meters (Weberbauer 260). 
Huanuco: Pampayacu, Sawada 59, 40; Kanehira 35. Playapampa, 



FLORA OF PERU 397 

2,700 meters, 4894. Vilcabamba, 1,800 meters, 4971. Pan de 
Azucar, Sawada 61. Rio Monzon (Weberbauer 255). Tambo de 
Vaca, 4458. Junin: Huasa-huasi, Ruiz & Pavon. Above Huacapis- 
tana, 3,000 meters, Killip & Smith 24456; (Weberbauer 246). 
Ayacucho: Carrapa, 1,200 meters, Killip & Smith 23234, 22296. 
Cuzco: Pillahuata, 2,800 meters, Pennell 14086 (det. Krause). 
Urubamba (Herrera). Bolivia; Colombia. "Matapalo." 

Gaiadendron puracense (HBK.) G. Don, Gen. Syst. 3: 432. 
1834. Loranthus puracensis HBK. Nov. Gen. & Sp. 3: 437. 1820. 

Perhaps only a variety of G. punctatum, but slenderer and smaller 
in all its parts; leaves lance-oblong, slightly obovate, 4-6 cm. long, 
2 cm. wide; flowers in terminal, rather long racemes; petals 6-8. 
The shorter, slenderer flowers, slender ovary, and smaller bracts are 
at once apparent in the following Weberbauer material, in contrast 
to cotype specimens of G. punctatum. 

Cajamarca: Huambos, 3,000 meters, Weberbauer 4164 (det. 
Baehni, ex char.). Junin: Huacapistana, 2,000 meters, Weberbauer 
2119 (det. Baehni, ex char.). Colombia. 

Gaiadendron Tagua (HBK.) G. Don, Gen. Syst. 3: 431. 1834. 
Loranthus Tagua HBK. Nov. Gen. & Sp. 3: 436. 1820. 

Leaves oblongish, fleshy-coriaceous, minutely punctate beneath, 
to nearly 10 cm. long, 3-4 cm. wide; racemes 2-3 cm. long, axillary, 
solitary; bractlets about as long as the calyx; petals 7, 2 cm. long; 
flowers yellowish white, fragrant. Peru to Costa Rica, British 
Guiana, and Venezuela, according to Krause. 

Peru: Apparently. Colombia. 

9. PHRYGILANTHUS Eichl. 

Half parasitic shrubs or rarely terrestrial trees, with mostly 
opposite leaves and conspicuous, red, yellow, or white flowers. 
Calyx rim obvious. Filaments definitely narrowed beneath the 
versatile anthers in contrast to those of Aetanthus. Besides the 
following, P. heterophyllus (R. & P.) Eichl., P. tetrandrus (R. & P.) 
Eichl., and P. aphyllus (Miers) Eichl. have been reported in Peru 
in literature as recently as in the new edition of the PflanzenJ "ami- 
lien but I have seen no specimens or definite citations that are 
not highly questionable, as for example Dombey material given as 
Peruvian. A "P. Lehmanianus" mentioned by Weberbauer, 162, "to- 
ward Ocros," Cajamarca, seems to be an unpublished name for a 
specimen probably referred to one of the following. 



398 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers 6-8-parted; inflorescences mostly axillary. 
Pedicels of the flowers subequal. 

Leaves about 3 cm. long or shorter P. ligustrinus. 

Leaves 5-10 cm. long. 

Parasitic shrubs; bracts promptly deciduous. 
Leaves 5-7 cm. long; pedicels 1-3-flowered . P. acutifolius. 

Leaves 8-10 cm. long; pedicels 3-flowered P. repens. 

Trees; bracts rather tardily deciduous P. eugenioides. 

Pedicels unequal, the middle much shorter, or the flowers sessile. 

P. suaveolens. 

Flowers 5-parted; inflorescences mostly terminal. 
Flowers about 10 cm. long or longer. 

Bracts conspicuously exceeding the calyculus. 

Leaves attenuate at the base, narrowly lanceolate. 

P. monzoniensis. 
Leaves rounded at the base, broadly ovate or elliptic. 

P. grandiflorus. 
Bracts small, little if at all longer than the calyculus. 

Leaves short-petioled P. Chodatianus. 

Leaves sessile P. peruvianus. 

Flowers 2-5 cm. long. 

Leaves opposite; bracts conspicuous P. longebracteatus. 

Leaves often ternate; bracts small P. verticillatus. 

Phrygilanthus acutifolius (R. & P.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 49. 1868. Loranthus acutifolius R. & P. Fl. 3: 48. pi. 274. 
1802. Psittacanthus acutifolius G. Don, Gen. Syst. 3: 417. 1834. 

Very similar to P. eugenioides, but perhaps always parasitic; 
leaves lanceolate, acutely long-acuminate, obscurely veined, dull 
on both sides, 5-10 cm. long, about 2 (-2.5) cm. broad; racemes 3 
cm. long; ternations and flowers pedicellate, the pedicels of each 
about 3 mm. long; bracts promptly deciduous; flowers white, 12 mm. 
long; fruit oval, black, the size of a pea. The bracts are not "tardily 
deciduous" on the type specimen. Loranthus albiflorus Hook. Icon. 
pi. 683 is the same, according to Eichler. Neg. 29457. 

Huanuco: Ruiz & Pavdn. Ayacucho: Rocky cliffs, Weberbauer 
5518. Ecuador. 



FLORA OF PERU 399 

Phrygilanthus Chodatianus Patsch. Bot. Jahrb. 45: 435. 1911. 

Internodes 1-2 cm. long; leaves obovate, strongly cuneate to 
the base, 3-4.5 cm. long, about 0.7 cm. broad; racemes secund, 4.5 
cm. long; bracts 4-5 mm. long; calyx 2-3 mm. long; flowers 4 cm. 
long, scarlet; fruit 1 cm. long, scarlet. On Polylepis. Negs. 3286. 
27827. 

Ancash: Pichiu, 4,000-4,100 meters, Weberbauer 2934 (type); 
217, 225. lea: Above Pisco, Weberbauer 5434 (det. Krause). 

Phrygilanthus eugenioides (HBK.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 50. 1868. Loranthus eugenioides HBK. Nov. Gen. & Sp. 
3: 435. 1820. Gaiadendron eugenioides G. Don, Gen. Syst. 3: 431. 
1834. 

Branches rugose; leaves ovate-lanceolate, sometimes narrowly 
so, basally attenuate to the slender petiole, long-acuminate, usually 
about 10 cm. long, lustrous above, more or less obviously veiny 
and punctate beneath; racemes axillary and terminal, often several 
cm. long; bracts promptly caducous; flowers pedicellate, 12-16 mm. 
long, greenish white; calyx scarcely 1 mm. high; fruit black, 8 mm. 
long. Variable in shape and size of leaves; very doubtfully distinct 
from P. acutifolius. Half parasitic on trees or growing independently. 
Illustrated, Mart. Fl. Bras. 5, pt. 2: pi. 12. 

Huanuco: Huariaca, 2,850 meters, 2402. San Rafael, Sawada 
115. Ambo, 2,100 meters, 3198. Amazonas: Chachapoyas, Weber- 
bauer 4386; 263 (det. Patschovsky). Piura: Huancabamba (Hum- 
boldt, type). Negritos, Haught. Argentina to Brazil. 

Phrygilanthus grandiflorus (R. & P.) Eichl. in Mart. Fl. 
Bras. 5, pt. 2: 46. 1868. Loranthus grandiflorus R. & P. Fl. 3: 45. 
pi. 273. 1802. 

Branches more or less pendent, bearing terminally several 
scarlet flowers 10-15 cm. long; leaves usually opposite, oval or elliptic, 
blunt, slightly lustrous above, somewhat veiny; bracts persistent, 
foliaceous, well exceeding the calyx; fruit oval, bluish, the size of 
an olive. 

Huanuco: Posuso, Muna, Pillao (Ruiz & Pavdn). Monzon, 
Weberbauer 3366 (det. Patschovsky). Cajamarca: Nancho, Rai- 
mondi. "Hatun-mocma," "hatun-mucma," "moma."' 

Phrygilanthus ligustrinus (Willd.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 48. 1868. Loranthus ligustrinus Willd. ex R. & S. Syst. 
7: 136. 1829. L. ligustrifolius Willd. op. cit. 135. 



400 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Similar in general to P. acutifolius, but the leaves only about 
half as large and the flowers corymbose or racemosely congested. 
In Peru according to Engler & Krause, Pflanzenfam. Neg. 11820. 

Peru(?): (Haenke, type of L. ligustrifolius) . "Middle America" 
(Humboldt). 

Phrygilanthus longebracteatus (Desr.) Macbr. Field Mus. 
Bot. 11: 17. 1931. Loranthus longebracteatus Desr. in Lam. Encycl. 
3: 599. 1792. L. glaucus R. & P. Fl. 3: 45. pi. 275. 1802. L. corym- 
bosus Dietr. Gaert. Lex. Nachtr. 4: 468. 1815-21. P. corymbosus 
Eichl. in Mart. Fl. Bras. 5, pt. 2: 46. 1868. 

Allied to P. verticillatus, but v/ith showy, reddish flowers, medially 
yellow, 3.5-5 cm. long, and conspicuous bracts 1 cm. long; leaves 
glaucous, opposite, ovate-lanceolate, obtusely subacuminate; inflo- 
rescences terminal, few-flowered; fruit deep purple, ovate-oval. 
Flowers scarlet, according to Weberbauer. Neg. 29458 (P. glaucus}. 

Huanuco: Yanano, 1,800 meters, 3737. Mito, 2,700 meters, 
3344. Huasa-huasi, Pillao, Muna,. Chinchao, Ruiz & Pavdn. 
Cajamarca: Cutervo, Raimondi (det. Krause). Ancash: Chiquian, 
Weberbauer 2849 (det. Krause). Junin: Huancayo, Raimondi. 
Chile to Colombia. "Pupa." 

Phrygilanthus monzoniensis Patsch. Bot. Jahrb. 45: 436. 1911. 

Branches papillose-verrucose, the internodes 2 cm. long; leaves 
lanceolate, acuminate, about 4 cm. long, 4-8 mm. broad, the nar- 
rowed petiolar base 3-5 mm. long; flowers 8 cm. long, the foliaceous 
bracts 2 cm. long; calyx dentate, 6 mm. long; fruit ellipsoid, 12 mm. 
long, black. Flowers bright red, orange-zoned. Negs. 3288, 27828. 

Huanuco: Near Monzon, 3,400 meters, Weberbauer 3342 (type). 

Phrygilanthus peruvianus Patsch. Bot. Jahrb. 45: 435. 1911. 

A shrub of 2 meters, the internodes 6-9 cm. long; leaves ovate- 
lanceolate, acute, 4-5.5 cm. long, 3-4 cm. broad; flowers to 9 cm. 
long, red, in terminal, secund racemes; bracts acute, 4-5 mm. long, 
2 mm. broad; calyx twice as long; fruit red, 1 cm. long. Neg. 3287. 

Ancash: Ocros, 2,400-3,500 meters, Weberbauer 2117, 3147. 
Lima: San Buenaventura, 2,800 meters, Pennell 14545; flowers 
scarlet-red. 

Phrygilanthus repens Patsch. Bot. Jahrb. 45: 436. 1911. 
A parasitic shrub with long adventitious roots; internodes 3-4 
cm. long; leaves lanceolate, 1-1.5 cm. broad, acutely acuminate, 



FLORA OF PERU 401 

narrowed into a petiole 2 cm. long; racemes about 8 cm. long, the 
pedicels 3-flowered; petals 6, white; calyx 3 mm. long, dentate; 
fruit 7 mm. long. Perhaps not a Phrygilanthus; parasitic on Schinus 
Molle, the white flowers very fragrant. Neg. 3289. 

Ancash: Valley of the Puccha, Weberbauer 3733 (type). 

Phrygilanthus suaveolens (HBK.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 49. 1868. Loranthus suaveolens HBK. Nov. Gen. & Sp. 3: 
435. 1820. 

Branches terete, smooth; leaves ovate-oblong, rounded at the 
base, acuminate, lustrous above, about 5 cm. long, half as wide; 
petioles 4-5 mm. long; racemes 2-4 in the axils; flowers white, short- 
pediceled, the middle subsessile, 6-8 mm. long, ebracteate; calyx 
obsoletely dentate. Neg. 3290. 

Cajamarca(?) : Gonzanama (Humboldt). 

Phrygilanthus verticillatus (R. & P.) Eichl. in Mart. Fl. 
Bras. 5, pt. 2: 47. 1868. Loranthus verticillatus R. & P. Fl. 3: 47. 
1802. L. Poeppigii DC. Prodr. 4: 308. 1830; P. & E. Nov. Gen. & 
Sp. 2: 61. pi. 184. 1838. 

A half parasitic shrub with verticillate, erect branchlets and 
subverticillate, lanceolate, enervose leaves; flowers crowded, grena- 
dine red, mostly borne ternately, 3-5 cm. long; bracts shorter than 
the calyx; fruit oval, purplish black. 

Cuzco: Urubamba (Weberbauer 4901); (Herrera 2271; det. 
Krause). Chile; Bolivia. "Ictriho," "itiu," "ictrigo," "bela-bela." 

10. PSITTACANTHUS Mart. 

Reference: Eichler in Mart. Fl. Bras. 5, pt. 2: 23-45. 1868. 

Except for the negative character, "endosperm lacking," the 
genus can not be distinguished, all species considered, from Phrygi- 
lanthus. The majority of species, however, may be recognized by 
the presence of a cup-like bracteal development in which the ovary 
is partly concealed and which may represent the complete union of 
the bractlets; it is hereafter referred to as a "cup." 

Flowers solitary or few, axillary or nodose; nodes not or little 
enlarged. 

Leaves about 1 cm. long P. cuneifolius. 

Leaves about 10 cm. long or longer. 

Petioles often 1 cm. long; leaves cuneate or acute at the base; 
flowers red. 



402 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pedicels verruculose; flowers acute P. crassifolius. 

Pedicels smooth; flowers truncate P. peculiaris. 

Petioles to 7 mm. long; leaves not at all acute at the base; 

flowers red or yellow P. Krausei. 

Flowers racemose, the racemes often short, or in branched inflores- 
cences, the nodes enlarged in many cases. 
Leaves opposite or ternate (see also P. calcaratus). 
Leaves cordate-amplexicaul. 

Leaves auricled at the base P. amplexifolius. 

Leaves merely slightly clasping the stems P. cardiphyllus. 

Leaves little if at all cordate. 
Leaves narrowly oblong or, if ovate, round-obtuse at the apex. 

Cup cyathiform; leaves 3-6 cm. wide P. Cordiae. 

Cup patelliform; leaves rarely as wide as 3 cm. 
Leaves subsessile; peduncles 6-8 mm. long. . . P. obovatus. 

Leaves sessile; peduncles 15 mm. long P. tumbecensis. 

Leaves ovate-lanceolate, acuminate or at least narrowed to 

the apex. 

Calyx much exceeded by a foliaceous bract .... P. cucullaris. 
Calyx not so subtended. 
Leaves 10 cm. long or much longer. 

Flowers pedicellate P. subalatus. 

Flowers sessile P. cupulifer. 

Leaves 3 cm. long or shorter P. coccineus. 

Leaves alternate or not strictly opposite. 

Flowers usually 8-10 cm. long P. calcaratus. 

Flowers 1-4 cm. long. 
Leaves roundish, not at all narrowed at the base. 

P. Weberbaueri. 

Leaves cuneate at the base, narrow. 
Leaves petioled; calyx twice longer than the cup. 

P. divaricatus. 
Leaves sessile; calyx slightly longer than the cup. 

P. linearis. 

Psittacanthus amplexifolius (van Tiegh.) Engler, Pflanzen- 
fam. Nachtr. 1: 136. 1897. Arthraxon amplexifolius van Tiegh. 
Bull. Soc. Bot. France 42: 353. 1895. 



FLORA OF PERU 403 

Allied to P. cardiphyllus, but the leaves auricled at the pro- 
nouncedly amplexicaul base, as in P. cordatus, but rounded at the 
apex; perianth in bud not enlarged apically. 

Cajamarca: Jae"n (Vidal Senege). 

Psittacanthus calcaratus A. C. Smith, Bull. Torrey Club 58: 
89. 1931. 

Branches stout, subquadrate; leaves veiny, rather thin, blunt 
or rounded at the apex, cuneate to the base, subelliptic, mostly 11-12 
cm. long, 5 cm. broad; petioles 1-1.5 cm. long; inflorescence terminal, 
crowded; peduncles to 4 cm. long, the stout pedicels about half as 
long; cup 2 mm. long, 4-5 mm. wide at the rim, subpatellif orm ; 
calyx 4 mm. long, truncate, minutely denticulate or entire; perianth 
blunt, 7-9.5 cm. long, each interior margin of the 6 linear lobes 
triangularly spurred near the tip; filaments attached 11 mm. below 
the tip of the perianth, 5-6 mm. long, equaling the pubescent an- 
thers. Flowers red and yellow. The related P. clusiifolius (Willd.) 
Eichl., with leathery leaves and stamens attached near the tip of the 
perianth segments, may occur. 

Loreto: Mishuyacu, 100 meters, Klug 694 (type). 

Psittacanthus cardiphyllus (Willd.) Eichl. in Mart. Fl. Bras. 
5, pt. 2: 27. 1868. Loranthus cardiphyllus Willd. ex R. & S. Syst. 7, 
pt. 1:162.1829. 

Like P. Cordiae, but the leaves distinctly cordate at the base; 
terminal racemes many-flowered, with about 10 nodes, the pedicels 
3-4 mm. long. Of this affinity are P. bicalyculatus (Mart.) G. Don 
and L. cordatus (Hoffm.) G. Don, the former with longer pedicels, 
2-4 inflorescence nodes, and oblongish leaves, the latter with lan- 
ceolate, acuminate leaves, hastately cordate at the base. 

Peru(?) : Probably. Brazil. 

Psittacanthus coccineus Patsch. Bot. Jahrb. 45: 436. 1911. 

Glabrous; leaves opposite, sessile, lanceolate, 3 cm. long, 4-8 mm. 
broad, subcoriaceous, the nervation evident; flowers in terminal 
racemes, on pedicels 4 mm. long; cup 2 mm. long, the usually dentate 
calyx 3 mm. long; perianth segments free, red, nearly 3 cm. long, 
the filiform filaments attached below the middle; fruit elliptic- 
oblong, black, 4 mm. long, 2 mm. thick. On Salix and Schinus 
Molle. Neg. 3299. 

Ancash: Near Ocros, Weberbauer 2643 (type). 



404 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Psittacanthus Cordiae Krause, Notizbl. Bot. Gart. Berlin 8: 
206. 1922. 

Branchlets terete; leaves sessile or nearly so, rigid-coriaceous, 
ovate or roundish, rarely slightly emarginate at the base, 5-8 cm. 
long, 3-6 cm. wide; nerves many, obvious; flowers short-pediceled, 
in terminal racemes; bracts broadly ovate, 2-2.5 mm. long, about as 
long as the cyathiform cup, this scarcely equaling the ovary; perianth 
scarlet, the linear-lanceolate lobes usually free to the base, 3-3.4 
cm. long, sparsely and very shortly pilose without; filaments about 
medially inserted; anthers 2-2.5 mm. long. P. chanduyensis Eichl. 
and P. obovatus Benth., both Ecuadorian, may be expected; the former 
has smaller leaves, rarely 3.5 cm. long, and very short racemes; 
the latter obovate leaves, more or less attenuate basally, distinctly 
nervose, 2.5-5 cm. long. Neg. 3298. P. cinctus (Mart.) G. Don 
with scurfy inflorescence and orange-tomentose flowers would also 
be sought here. 

Piura: On Cordia rotundifolia, near Serran, 200-300 meters, 
Weberbauer 5996 (type). La Brea, 150 meters, Weberbauer 7766. 
Without locality, Raimondi. "Piiia." 

Psittacanthus crassifolius (Mart.) G. Don, Gen. Syst. 3: 
417. 1834. Loranthus crassifolius Mart, ex R. & S. Syst. 7, pt. 1: 
123. 1829. 

Young parts obsoletely reddish-scurfy; branches terete; leaves 
broadly ovate or elliptic, fleshy-coriaceous, opaque, 8-12 cm. long, 
5-7 cm. broad or exceptionally half again as large; flowers mostly 
ternately arranged in the axils, the common peduncle 4-6 mm. long; 
perianth to 9 cm. long, the filaments attached to about 1 cm. below 
the apex, the anthers 5-6 mm. long. A. C. Smith, from whom this 
description, in part, is drawn, observed 1 or 2 spurs on each perianth 
lobe. Neg. 19056. 

Loreto: Mishuyacu, 100 meters, Klug 119 (det. Smith). Brazil. 

Psittacanthus cucullaris (Lam.) G. Don, Gen. Syst. 3 :416. 1834. 
Loranthus cucullaris Lam. Journ. Hist. Nat. Paris 1: 444. pi. 23. 1793. 

Completely glabrous, drying black; petioles 2-6 mm. long; leaves 
elliptic-oblong or linear-lanceolate, acute, to 10 cm. long, 1.5-4 
cm. broad, subcoriaceous, with 5-7 nerves; flowers terminally 
paniculate, the peduncle of the ternations to 12 mm. long; bracts 
cucullate-concave, foliaceous, 12-20 mm. long; perianth segments 
yellow, orange, or reddish, the filaments medially inserted; fruit 
ellipsoid. Illustrated, Mart. Fl. Bras. 5, pt. 2: pi. 7. 



FLORA OF PERU 405 

Loreto: Mishuyacu, Klug 1317 (det. Smith). Brazil to the 
Guianas. 

Psittacanthus cupulifer (HBK.) G. Don, Gen. Syst. 3: 417. 
1834. Loranthus cupulifer HBK. Nov. Gen. & Sp. 3: 438. 1820. 

Allied toP.cucullaris; branches terete; leaves subsessile, obliquely 
ovate-oblong, narrowed to the obtuse apex, reticulate, 2.5-5 cm. 
broad; inflorescences densely flowered, 3-6 cm. long, the flowers 
sessile or subsessile, 4 cm. long, subtended by a concave bract 5 mm. 
long; calyx 6-8 mm. long; filaments medially adnate; fruit ellipsoid. 
Weberbauer noted the flowers as scarlet below, golden yellow 
medially, orange toward the tips; Klug as yellowish green and red, 
yellow and red, and yellow. Here as elsewhere G. Don has been 
accepted as second authority for the name instead of Blume, who in 
R. & S. Syst. 7, pt. 2: 1730. 1830, indicated the transfer without 
actually making it. 

San Martin: Moyobamba, Weberbauer 4718 (det. Patschovsky). 
Loreto: Maucallacta, 200 meters, Klug 3955. Pebas, Williams 
1871. Florida, 180 meters, Klug 2189, 2325. Fortaleza, 140 meters, 
Williams 4315; Klug 2804- Lower Rio Nanay, Williams 614, 531. 
Yurimaguas, 180 meters, Williams 7875, 7874- Caballo-cocha, 
Williams 244%- Rio Nanay, Raimondi. Middle Rio Apaga, Tess- 
mann 3838 (det. Krause). Lower Rio Blanco, Tessmann 3076 (det. 
Krause). Ecuador to Bolivia. "Pishcu-isman," "jucu-gui-o" 
(Huitoto name), "suelda con suelda." 

Psittacanthus cuneifolius (R. & P.) G. Don, Gen. Syst. 3: 
416. 1834. Loranthus cuneifolius R. & P. Fl. 3: 46. pi. 276. 1802. 
Phrygilanthus cuneifolius Eichl. in Mart. Fl. Bras. 5, pt. 2: 49. 1868. 

Densely branched, the slender branches spreading; leaves oblong- 
cuneate or narrowly obovate, subsessile, 1.5-2 cm. long, about 5 
mm. broad, apically rounded, mucronate, faintly 3-nerved; flowers 
solitary or 3-5 in axillary fascicles or racemes scarcely 1 cm. long, the 
pedicels 5-7 mm. long; cup 2.5 mm. deep, the teeth marked, the 
calyx nearly twice as long; flowers about 3 cm. long, yellowish red; 
petals coalescent; filaments filiform, subulate; fruit 8-10 mm. long. 
Subsp. peruanus Engler is apparently the typical, i.e. the Peruvian, 
form. A shrub of 5 meters on Rhamnaceae (Killip & Smith). The 
fruits are used for the making of bird lime, which boys smear on the 
branches of trees to catch small birds, particularly the "chchainas" 
(jilgueros)', the gum is employed also as a plaster for fractures, etc. 
(Herrera). Neg. 29462. 



406 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Cheuchin, Ruiz & Pawn. Chulki, Sawada 57. Mito, 
2,700 meters, 1781. Huanuco, 2,100 meters, 2046. Junin: Between 
Viques and Ingahuasi, 3,150 meters, Killip & Smith 22173. Cuzco: 
Capana, Prov. Paucartambo (Herrera 635). Parcocella, 3,600 
meters, Herrera 1144- Arequipa: Yura, 2,700 meters, Weberbauer 
1415. Above Pampa Redonda, Weberbauer 5767. Ayacucho: 
Ayacucho, 3,100 meters, Weberbauer 5530. Chile to Brazil. "Pupa," 
"liga," "suelda con suelda." 

Psittacanthus divaricatus (HBK.) G. Don, Gen. Syst. 3: 
417. 1834. Loranthus divaricatus HBK. Nov. Gen. & Sp. 3: 433. 1820. 

Branches spreading; leaves subsessile, elliptic-oblong, rounded 
at the apex, coriaceous, 2-3 cm. long, 10-18 mm. wide; peduncles 
3-flowered, paniculately disposed; flowers 2.5 cm. long or longer, 
on pedicels to 4 mm. long; calyx obsoletely dentate. The original 
locality is given as perhaps the Rio Orinoco, but the species is 
accredited in literature to Chile. 

Cajamarca: Nancho, Raimondi. Brazil (?); Chile(?). 

Psittacanthus Krausei Macbr. Candollea 5: 349. 1934. 

Branches terete, markedly nodose; petioles 6-7 mm. long; leaves 
ovate, attenuately and acutely acuminate, to 10.5 cm. long, about 
3.5 cm. broad; flowers 2-3 in corymbs and umbels, the common 
peduncle 12-15 mm. long; pedicels glabrous, 10-12 mm. long; flowers 
lustrous, 6-6.5 cm. long; anthers 3 mm. long, with a mucro 2 
mm. long. P. peronopetalus Eichl., with larger leaves, puberulent 
inflorescence, shorter pedicels, and short-mucronulate anthers, may 
be expected. P. biternatus (Hoffm.) G. Don, similar, has elliptic 
leaves rounded at both ends. 

Loreto: Mishuyacu, 100 meters, King 1323 (type). 

Psittacanthus linearis (Killip) Macbr., comb. nov. Phrygi- 
lanthus linearis Killip, Journ. Wash. Acad. Sci. 24: 45. 1934. 

Glabrous, with terete branches and linear, obtuse leaves, slightly 
narrowed at the base, sometimes subopposite, 2-3.5 cm. long, 1.5-5 
mm. wide, obscurely 1-nerved; inflorescence compactly subcorym- 
bose, the spreading pedicels of the ternately arranged flowers 3-5 
mm. long; bractlets triangular-ovate, 2-3 mm. long; cup 2-2.5 
mm. deep, denticulate, slightly exceeded by the subentire calyx; 
petals 6, bright red or scarlet, 2-2.5 cm. long; filaments about 2 
cm. long, thus attached toward the base. This plant, from descrip- 
tion, must be a Psittacanthus, apparently similar to P. coccineus. 



FLORA OF PERU 407 

According to Haught, it is slender, semi-trailing, almost always on 
"algarroba," the branches 2 meters long or longer, flowering all the 
year and broken off with long poles by goat herders for their animals 
that seem to prefer this to any other plant. On Acacia and Prosopis 
limensis. 

Piura: Prov. Paita (Haught 9, type). Negritos, Haught F21. 
La Brea, Weberbauer 7765. Prov. Paita (Townsend 823?; "but 
leaves to 6 cm. long, 15 mm. wide"). "Pina." 

Psittacanthus obovatus Benth. ex Eichl. in Mart. Fl. Bras. 
5, pt. 2: 27. 1868. Loranthus obovatus Benth. Bot. Voy. Sulph. 103. 
1844. 

Slightly scabrous; branches dichotomous or verticillate; leaves 
subsessile, obovate, obtuse, rounded or cuneate at the base, faintly 
reticulate- veiny, 2.5-5 cm. long, 12-25 mm. wide; racemes with 
5-6 short nodes; peduncles 6-8 mm. long, finely puberulent; flowers 
in 3's, the pedicels 4-6 mm. long; cup a little shorter than the calyx; 
petals 6, 3.5-4 cm. long. Allied by Eichler with P. chanduyensis 
Eichl. with sessile, not at all obovate leaves. 

Piura: Negritos, Haught F 22. Ecuador. "Pina." 

Psittacanthus peculiaris A. C. Smith, Bull. Torrey Club 
58: 90. 1931. 

Leaves ovate-lanceolate, falcate to broadly ovate, cuneate at the 
base, narrowly acuminate or sometimes only acute, 4-8 cm. wide, 
mostly 10-12 cm. long; petioles about 1 cm. long; flowers red, usually 
2 or 3, the common peduncle and pedicels 2-3 mm. long; cup sub- 
patelliform, 0.8 mm. long, 2 mm. wide at the rim; corolla finally to 
5 cm. long, very slender toward the base, the throat narrowed, the 
linear lobes each partly obscured by a fleshy, oblong, horizontal spur 
nearly 2 mm. long; filaments scarcely 1 mm. long. 

Loreto: Mishuyacu, 100 meters, Killip & Smith 29967 (type); 
King 118, 1206. Pefia Blanca, Rio Itaya, 110 meters, Killip & 
Smith 29653. Leticia, Williams 3147. Lower Rio Nanay, Wittiams 
593, 646. 

Psittacanthus subalatus Krause, Notizbl. Bot. Gart. Berlin 
8: 207. 1922. 

Branchlets stout, 4-angled and wing-margined; petioles 1-3 mm. 
long; leaves coriaceous, rarely cordate-emarginate at the base, 
mostly subfalcate, to 20 cm. long, 4-9 cm. wide; lateral nerves dis- 
tinct on both sides; flowers pedicellate; bracts broadly ovate, con- 



408 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cave, 2-3 mm. long, the subpatelliform cup 2-3 mm. long, the calyx 
2 cm. long; ovary 5-6 mm. long; perianth lobes roseate, free to the 
base; filaments medially attached, 3-4 cm. long; style 7-8.5 cm. 
long. Differs from P. cupulifer (HBK.) Eichl. in its larger and 
broader leaves and much smaller cup. P. fakifrons (Mart.) G. Don, 
with golden yellow flowers and oblong-lanceolate leaves 1-4 cm. 
wide, may be expected. Neg. 3300. 

Piura: Above Palambla, 1,400 meters, Weberbauer 6030 (type). 

Psittacanthus tumbecensis (Killip) Macbr., comb. nov. 
Phrygilanthus tumbecensis Killip, Journ. Wash. Acad. Sci. 24: 45. 
1936. 

Leaves narrowly oblong, subfalcate, 5-10 cm. long, 1.5-3 cm. 
wide, obtuse, narrowed at the base, with many obscure nerves; 
inflorescence loosely corymbose; bractlets 2-3 mm. long; cup 2-2.5 
mm. long, 4-5 mm. wide, the calyx slightly longer; petals 6, 3-4 
cm. long, "fiery red with yellow point." Apparently allied to 
P. obovatus. 

Tumbez: Mountains east of Hacienda Chicama, 500-700 meters, 
Weberbauer 7668, type. 

Psittacanthus Weberbaueri Patsch. Bot. Jahrb. 45: 437. 1911. 

Glabrous, erect; leaves sessile, oblong-ovate, obtuse or rounded 
at the apex, 3-3.5 cm. long, 2-2.5 cm. broad, the lateral nerves 
reticulate; flowers in terminal racemes on pedicels 3-4 mm. long; cup 
cucullate, the calyx 3 mm. long; perianth 2.3-3.5 cm. long; filaments 
filiform, medially attached; fruit oblong, 5 mm. long, 3 mm. thick. 
Neg. 3301. 

Cajamarca: Llallan, 600-700 meters, Weberbauer 3796 (type). 

11. PERISTETHIUM van Tiegh. 
Reference: van Tiegh. Bull. Soc. Bot. France 42: 175. 1895. 

A segregate of Phrygilanthus, but as well marked as any genus 
by virtue of its small flowers crowded in axillary inflorescences and 
basally subtended by white bracts that on falling leave a little collar, 
to which peculiarity the generic name refers. No Peruvian specimen 
appears to exist, but the plant is included here because definitely 
stated to occur in Peru by all authorities! 

Peristethium leptostachyum (HBK.) van Tiegh. loc. cit. 
Loranthus leptostachyus HBK. Nov. Gen. & Sp. 3: 440. 1820. 



FLORA OF PERU 409 

Petioles 10-12 mm. long; leaf blades ovate-elliptic, acuminate, 
net-veined, about 10 cm. long and half as wide; axillary spikes 
2-several, much shorter than the leaves; flowers scarcely 4 mm. long. 

Peru: (Occurrence doubtful). Colombia. 

12. STRUTHANTHUS Mart. 
Phthirusa Mart. Flora 13, pt. 1: 110. 1830. 
Reference: Eichler in Mart. Fl. Bras. 5, pt. 2: 52-87. 1868. 

Erect or not infrequently scandent shrubs, often developing 
adventitious roots. Flowers tiny, in simple or branched, axillary or 
terminal spikes, racemes, or panicles, or rarely solitary or subsoli- 
tary, usually dioecious or perfect. Stamens often unequal, the fila- 
ments of the longer then sometimes scalloped at each side, appar- 
ently by pressure of the anthers of the shorter ones. Filaments in 
varying degree thin and slender, short and thick, or even lacking, 
even in staminate flowers. Except in treating Lor anthus L. sensu lat., 
this group must be maintained as distinct. Even though its char- 
acter of small flowers is intangible, this is obviously a natural genus 
or, in the former case, subgenus. Here, as elsewhere in the family, 
G. Don is given as second authority in place of Blume in R. & S. 
Syst. 7, pt. 2: 1731. 1830, where the combinations are only indicated. 
Rubber has been produced commercially from two of the species 
found in Peru, S. aduncus and S. pyrifolius, the fruits having yielded 
as high as 7 per cent rubber on an average. The practicability of 
cultivation is highly questionable (Krause), so that no economic 
significance can be envisaged, the exploitation of wild plants for this 
purpose surely being very limited. 

Plants glabrous; filaments (except in 2 species) slender. 

Leaves 5-7.5 cm. wide, at least most of them, or the filaments 
thick, often scalloped. 

Filaments thin, straight; leaves all or most of them wider than 
5 cm. 

Verticels bracteolate, pedicellate, more or less paniculate. 

S. retroflexus. 

Verticels spicate, ebracteolate or the bractlets promptly 
caducous S. polystachyus. 

Filaments thick, often laterally scalloped; leaves mostly 4-5 
cm. wide. 

All the flowers panicled; anthers acuminate. . . .S. aduncus. 



410 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

All or nearly all the flowers spicate or racemose; anthers 

mucronate S. Eichleri. 

Leaves regularly narrower, rarely if ever some of them about 5 

cm. wide. 

Verticels ebracteolate or the bractlets very promptly caducous, 
free; leaves little longer than broad. 

Racemes several cm. long S, orbicularis. 

Racemes less than 1 cm. long S. lucarquensis. 

Verticels bracteolate, the bractlets more or less united; leaves 

usually distinctly longer than broad. 

Flower clusters all in corymbs or pseudocymes, the inflores- 
cence rarely longer than 2 cm., usually shorter. 

Leaves obovate-suborbicular S. retusus. 

Leaves oblong-lanceolate S. dichotrianthus. 

Flower clusters in spikes or racemes often longer than 2 cm., 

the terminal inflorescence rarely paniculate. 
Flower clusters borne in spikes (typically). 
Spikes 2-5 cm. long; flowers to about 6 mm. long; leaves 

nearly nerveless S. concinnus. 

Spikes 1 cm. long; flowers to 3 mm. long; leaves nerved. 

S. tennis. 
Flower clusters pedicellate, that is, racemose (cf. S. tennis). 

Leaves fleshy, veinless S. acuminatus. 

Leaves thin, veiny on both sides S. sarmentosus. 

Plants more or less reddish-furfuraceous; filaments thick. 

Inflorescence branchlet not conspicuously compressed; fruit 

reflexed S. pyrifolius. 

Inflorescence branchlet strikingly compressed; fruit spreading. 

S. platycladus. 

Struthanthus acuminatus (R. & P.) Blume ex R. & S. Syst. 
7, pt. 2: 1731. 1830. Loranthus acuminatus R. & P. Fl. 3: 49. 1802. 

Slender, with long, virgate, slightly angled branches, the narrowly 
ovate-lanceolate, long-acuminate leaves remote; petioles 3-5 mm. 
long; leaf blades 6-7 cm. long, 1.5-2.5 cm. broad, rounded or scarcely 
acute at the base, dull; racemes solitary, about 4 cm. long, the 
slender rachis sharply quadrate; ternation pedicels 2 mm. long, the 
flowers sessile; bracteoles tardily caducous, nearly free, subequal, 
shorter than the calyx; flowers pale yellowish, 4 mm. long, the 



FLORA OF PERU 411 

medially attached filaments very short; fruit oval. In aspect similar 
to some forms of S. thyrsiflorus (C. & S.) Blume (S. syringifolius 
Mart.), with short-acuminate and somewhat veiny leaves and one 
bractlet longer than the other. Neg. 29464. 

Huanuco: Pillao, Chinchao, Panao, Ruiz & Pavon. 

Struthanthus aduncus (Meyer) G. Don, Gen. Syst. 3: 414. 
1834. Loranthus aduncus Meyer, Prim. Fl. Esseq. 149. 1818. L. pani- 
culatus HBK. Nov. Gen. & Sp. 3: 442. 1820. L. conduplicatus HBK. 
op. cit. 441. L. Theobromae Willd. ex R. & S. Syst. 7: 132. 1829. 
Phthirusa Theobromae Eichl. in Mart. Fl. Bras. 5, pt. 2: 56. pi. 14. 
1868. P. paniculata Macbr. Field Mus. Bot. 11: 17. 1931. 

Glabrous, erect or scandent, the bark grayish; petioles 5-10 mm. 
long; leaf blades ovate or ovate-lanceolate, somewhat acute, 3-10 
cm. long, 3-7 cm. wide, coriaceous; flowers usually white, 4-6 mm. 
long, the perianth segments spreading, paniculate or the lower 
axillary spikes simple, the ternations sessile or subsessile; calyx mem- 
branous, truncate; fruit red, oblong. Not at all clearly distinct and 
more or less typically represented by some of the following col- 
lections is var. orinocensis (Spreng.) Baehni & Macbr., comb. nov. 
(Phthirusa orinocensis [Spreng.] Eichl. op. cit. 60), with the terna- 
tions more or less pediceled and the racemes not at all panicled. 

San Martin: Morales, Williams 5693. Zepelacio, 1,200-1,600 
meters, Klug 3605. San Roque, 1,400 meters, Williams 7064. 
Loreto: Iquitos, Tessmann 3530, 5068; Williams 7976, 8095. Mishu- 
yacu, 100 meters, Klug 3937. Florida, 200 meters, Klug 2134, 
2047. Mainas, Poeppig 2383,Cuzco: Urubamba, Weberbauer 5063 
(flowers yellow). Widely distributed in tropical America. "Suelda 
con suelda," "pishco isman," "rochido-ey" (Huitoto name). 

Struthanthus concinnus Mart, ex G. Don, Gen. Syst. 3: 411. 
1834. 

Branches long, slender, terete; petioles 3-6 mm. long; leaf blades 
lanceolate, acute at the base, gradually and acutely acuminate, 
4-7 cm. long, 1.5-2.5 cm. broad, nearly nerveless, thin, dull beneath; 
ternations sessile or subsessile; flowers greenish yellow, the stami- 
nate to 6 mm., the pistillate to 4 mm. long; filaments affixed below 
the middle of the linear perianth segments; calyx denticulate; fruit 
ellipsoid. S. rubens Mart, ex G. Don has coriaceous leaves lustrous 
on both sides, the lateral nerves obvious. The Peruvian specimens 
may be rather S. sarmentosus; at any rate, they are aberrant here by 



412 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

reason of the pediceled flowers. Illustrated, Mart. Fl. Bras. 5, pt. 
2: pi. 24- 

Junin: Huacapistana, Weberbauer 2157 (det. Patschovsky). 
Colonia Perene", 680 meters, Killip & Smith 25058. La Merced, 
700 meters, Killip & Smith 23507. Chanchamayo Valley, 1,200 
meters, Schunke 331. Above San Ramon, 1,300-1,700 meters, 
Schunke A93. Carpapata, 2,400 meters, Killip & Smith 24366. 
Huacapistana, 1,800-2,400 meters, Killip & Smith 24314- Huanuco: 
Yanano, 1,800 meters, 3724- Ayacucho: Carrapa, 2,200 meters, 
Killip & Smith 22310. Brazil; Surinam. 

Struthanthus dichotrianthus Eichl. in Mart. Fl. Bras. 5, 
pt. 2: 75. 1868. 

Densely branched, the branches more or less scandent; petioles 
3-10 mm. long; leaf blades oblong-lanceolate, obviously nervose 
beneath, attenuate at both ends, 2.5-7 cm. long, 1-3 cm. wide; 
ternations pedicellate, the pedicels 2-5 mm. long, in 2-3-chotomous, 
axillary umbels; bracts minute; flowers white, the larger staminate 
ones 5-6 mm. long; calyculus entire; fruit oblong, 5-6 mm. long, 
3 mm. thick. The Peruvian collection was determined apparently 
by Patschovsky. 

Loreto: Puerto Limon, Tessmann 4886. Surinam and British 
Guiana; Venezuela. 

Struthanthus Eichleri Baehni & Macbr., nom. nov. Phthirusa 
polystachya Eichl. in Mart. Fl. Bras. 5, pt. 2: 57. pi. 19. 1868, not S. 
polystachyus (R. & P.) G. Don. 

Nearly S. aduncus, but the leaves rather less coriaceous, about 10 
cm. long, the flowers all in simple, axillary spikes and, especially, the 
filaments slightly glandular, the anthers shortly produced. Flowers 
purple (Ule). S. nitens (Mart.) Baehni & Macbr. would be sought 
here; its lustrous, narrowly ovate-lanceolate leaves are attenuately 
acuminate, its spikes solitary, the filaments eglandular, the anthers 
apiculate. 

Rio Acre: Ule 9337 (det. Krause). Brazil. 

Struthanthus lucarquensis (HBK.) Macbr., comb. nov. Lor- 
anthus lucarquensis HBK. Nov. Gen. & Sp. 3: 440. 1820. Phthirusa 
lucarquensis G. Don, Gen. Syst. 3: 421. 1834. 

Much branched, the branchlets terete; leaves somewhat mem- 
branous and veiny, 2.5 cm. long, 18-20 mm. broad; flowers sub- 
sessile, 3 mm. long, whitish yellow; racemes only 6 mm. long; petals 



FLORA OF PERU 413 

4, oblong, acute, slightly concave; filaments membranous, nearly 
free; berry ovoid. Because of the slender filament, this plant must 
belong to the group Struthanthus. 

Peru(?) : Lucarque ( Humboldt}. Ecuador. 

Struthanthus orbicularis (HBK.) Blume ex R. & S. Syst. 7, 
pt. 2: 1731. 1850. Loranthus orbicularis HBK. Nov. Gen. & Sp. 3: 
434. 1820. 

A shrub with long, flexuous branches, these often pendent and 
twining, the young leaves prehensile; petioles 8-12 mm. long; blades 
more or less approaching orbicular, 2.5-5 cm. long, attenuate or 
contracted to the petiole, coriaceous; ternations subsessile in axillary 
spikes 2.5-10 cm. long, the rachis sharply quadrate; flowers yellowish 
green, to nearly 6 mm. long, more or less pediceled; bracts free, cadu- 
cous; fruit oval, red, 1 cm. long. Here would key Tessmann 3248 
with more elliptic leaves, from the Ucayali; the ternations are 
pedicellate, the racemes only 1-2 cm. long. 

San Martin: Rioja, west of Moyobamba, Weberbauer 4702; 
290 (det. Patschovsky). Brazil, north to Central America. 

Struthanthus platycladus (Ule) Baehni & Macbr., comb. nov. 
Phthirusa platyclada Ule, Verh. Bot. Ver. Brandenb. 48: 153. 1907. 

Similar to S. pyrifolius, to which it could be joined as a variety; 
leaves coriaceous, rounded-obtuse, mucronate; flowers 2-3 mm. 
long, yellowish green; fruit ascending-spreading. Perhaps all the 
Peruvian material is rather referable to S. pyrifolius. 

San Martin: Tarapoto, Wittiams 5711. Loreto: Yurimaguas, 
Williams 4984- Middle Rio Blanco, Tessmann 3141- Brazil. 
"Pishco isman." 

Struthanthus polystachyus (R. & P.) G. Don, Gen. Syst. 
3: 410. 1834. Loranthus polystachyus R. & P. Fl. 3: 50. 1802. 

Branches long, terete, the branchlets more or less compressed 
at the nodes; petioles 1-1.5 cm. long; blades broadly ovate, acumi- 
nate, acute at the base, to 15 cm. long, 7 cm. wide, perfectly opaque, 
coriaceous, cartilaginous-margined, obviously finely reticulate- veined 
on both sides; spikes 2 or more together, to 6 cm. long, the rachis 
angled; ternations 5-7 mm. apart; bractlets inconspicuous; flowers 
tiny, purple; fruit oval, brownish yellow, 6 mm. thick. Flowers 
lacking in the Madrid material. Neg. 29465. 

Huanuco: Muna, Pillao, Panao, Pati, Chinchao, Ruiz & Pawn. 



414 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Struthanthus pyrifolius (HBK.) G. Don, Gen. Syst. 3: 413. 
1834. Loranthus pyrifolius HBK. Nov. Gen. & Sp. 3: 441. 1820. 
Phthirusa pyrifolia Eichl. in Mart. Fl. Bras. 5, pt. 2: 63. pi. 17. 1868. 

Branchlets compressed, more or less reddish-scurfy but soon 
glabrous; petioles rarely 10 mm. long; blades ovate-lanceolate, 
often acuminate, scarcely coriaceous, distinctly nervose, about 10 
cm. long and half as wide or smaller; ternations of red-brown perfect 
flowers sessile or subsessile in axillary spikes or rarely in a terminal 
panicle (var. terminalis Macbr., comb, nov.; P. pyrifolia var. 
terminalis Macbr. Candollea 5: 349. 1934); calyx obsoletely dentic- 
ulate; perianth segments 6, narrowly lanceolate, 1.5-2 mm. long; 
filaments short; fruit oblong, red. The name is sometimes modern- 
ized or corrected to read "pirifolius." S. rufus (Mart.) Macbr. 
(comb, nov.; Loranthus rufus Mart.) has dioecious flowers 5-6 mm. 
long, borne in a terminal panicle. Also to be expected are S. micran- 
thus (Eichl.) Baehni & Macbr. (comb, nov.; Phthirusa micrantha 
Eichl.) and S. phaeoclados (Eichl.) Baehni & Macbr. (comb, nov.; 
Phthirusa phaeoclados Eichl.), both with usually 4-merous flowers 
and sessile anthers, the former with them in axillary glomerules, the 
latter in axillary spikes, the leaves emarginate and 5-7-plinerved. S. 
santaremensis (Eichl.) Baehni & Macbr. (comb, nov.; Phthirusa 
santaremensis Eichl.), similar, has pinnately nerved leaves. 

San Martin: San Roque, 1,400 meters, Williams 7482 (type of var. 
terminalis Macbr.). Morales, Williams 5711. Rumizapa, Williams 
6790 (toward S. platydadus}. Amazonas: Chachapoyas, Weber- 
bauer 4458. Junin: Colonia Perene", 680 meters, Killip & Smith 
24996,25071. Rio Peren<, 600 meters, Killip & Smith 25716. La 
Merced, 600 meters, 5337. Loreto: Iquitos, Ule 6249 (flowers 
purple); Williams 3584; Killip & Smith 27310. Caballo-cocha, 
Williams 2257. La Victoria, Williams 2695. Puerto Arturo, 
Williams 4984. Tropical South America to Central America. 
"Suelda con suelda," "pishco isman." 

Struthanthus retroflexus (R. & P.) G. Don, Gen. Syst. 3: 
415. 1834. Loranthus retroflexus R. & P. Fl. 3: 49. pi 279. 1802. 

Stoutly branched, the branches terete; petioles nearly 1.5 cm. 
long; leaves broadly elliptic or oval, about 10 cm. long, dull or nearly 
so, coriaceous, cartilaginous-margined, very faintly veiny on both 
sides, acute at both ends; panicles axillary and terminal, to 15 cm. 
long, the ternation pedicels scarcely 2 mm. long; rachis minutely 
papillose, angled; bractlets persistent, broadly ovate; flowers sessile, 



FLORA OF PERU 415 

tiny; fruit oval, nearly 10 mm. long, 6 mm. thick. The type at 
Madrid has been referred by Krause, following Eichler's suggestion, 
to S. marginatus (Desr.) Blume, var. paniculatus Eichl. That Brazil- 
ian species, however, seems to be uniformly a much more slenderly 
branched plant, with typically narrower, thinner leaves. Neg. 29466. 
Huanuco: Chinchao, Macora, Ruiz & Pavon. 

Struthanthus retusus (Gham. & Schlecht.) Blume ex R. & S. 
Syst. 7, pt. 2: 1731. 1830. Loranthus retusus Cham. & Schlecht. 
Linnaea 3: 215. 1828. L. polyrrhizos Mart, ex R. & S. Syst. 7, pt. 
1: 139. 1829. S. polyrrhizos Mart, ex Don, Gen. Syst. 3: 412. 1834. 

Scandent, flexuous, branched; petioles 8-12 mm. long; leaves 
obovate-suborbicular, 2.5-3 (-5) cm. long, 0.5-3 cm. broad, usually 
marginate, sometimes deeply so, coriaceous; ternations in axillary, 
peduncled corymbs, more or less pedicellate; flowers greenish yellow, 
4-6 mm. long, the pedicels of the pistillate ones as long, those of the 
staminate shorter; fruit ovoid-oblong, under 6 mm. long. The 
Peruvian collection is apparently determined by Patschovsky, as 
"aff. polyrrhizus" ; but see remark under S. orbicularis. 

Loreto: Middle Rio Ucayali, Tessmann 3298. Brazil. 

Struthanthus sarmentosus (R. & P.) Blume ex R. & S. Syst. 
7, pt. 2: 1731. 1830. Loranthus sarmentosus R. & P. Fl. 3: 49. pi. 
278. 1802. 

Similar to S. acuminatus, but the shorter branchlets nearly terete, 
the leaves more shortly acuminate, thin and distinctly veiny, dull 
beneath, lustrous above, clearly acute at the base or slightly decur- 
rent on the slender petiole, this about 1 cm. long; terminal racemes 
panicled. The Killip and Smith material has been referred to S. 
concinnus Mart., but as its flower clusters are pedicellate, it rather 
belongs here or to S. tennis Patsch., but the petioles are only 4-5 
mm. long. Neg. 27839. 

Huanuco: Muna, Ruiz & Pavon; at 2,100 meters, 401 9. Junin: 
Huacapistana region, Killip & Smith 24314, 24366. Loreto: 
Caballo-cocha, Williams 2144- Ayacucho: Carrapa, Killip & 
Smith 22310. "Liga." 

Struthanthus tenuis Patsch. Bot. Jahrb. 45: 438. 1911. 

Slender-stemmed, the internodes to 10 cm. long; petioles 1 cm. 
long; leaves broadly ovate-lanceolate, acuminate, 5-6 cm. long, 
3.5 cm. wide; bractlets lanceolate, 2 mm. long; filaments medially 



416 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

adnate to the greenish perianth segments. This is probably to be 
included in S. sarmentosus (R. & P.) Blume. 

Junin: La Merced, 1,000 meters, Weberbauer 1948, type; 283. 

52. SANTALACEAE. Sandalwood Family 
Reference: Pilger, Pflanzenfam. ed. 2. 16b: 52-91. 1935. 
The source of sandalwood and oil, the family is outstanding in 
Peru in no respect unless by the contrast between its representatives 
there: a genus of small trees strikingly reddish-pubescent, and two 
genera of inconspicuous herbs. 

Woody plants with oval, more or less pubescent leaves. 

1. Cervantesia. 
Herbs. 
Bractlets not united; plants diminutive, creeping, the hidden 

stems yellow 2. Arjona. 

Bractlets united to form a cup; erect or ascending plants. 

3. Quinchamalium. 

1. CERVANTESIA R. & P. 

Casimiroa Dombey ex Baill. Adansonia 3: 126. 1862-3. 

Small trees with elliptic leaves that are pubescent, especially 
beneath, as well as the branchlets and the terminal or subterminal 
flower clusters. Tube of the perianth shortly campanulate above 
the ovary. Named for Vicente Cervantes, Mexican botanist. 

Cervantesia bicolor Cav. Icon. 5: 49. pi. 475. 1799. 

Similar to C. tomentosa, but the branchlets and leaf pubescence 
beneath a dense tomentum of reddish, crisped or crinkly hairs; leaves 
5-8 cm. long, mostly 5-6 cm., 2-2.5 cm. broad; perianth segments 
rounded or subacute. 

Lima: Obrajillo and San Buenaventura, Nee. Ecuador. 

Cervantesia tomentosa R. & P. Prodr. 39. pi 7. 1794; Fl. 3: 
19. pi 24. 1802. 

Branchlets and leaves beneath yellowish or somewhat rusty, 
pilose or lanuginose with long, more or less flexuous hairs, especially 
those of the leaves; leaves soon lustrous and glabrous above, mostly 
9-10 cm. long, 2-2.5 cm. broad; young leaves and inflorescences 
densely red-pilose, the small flower parts hidden in the abundant 
pubescence; perianth segments acute, persistent and nearly enclos- 
ing the acorn-like fruits, these about 1 cm. long, 8 mm. thick. A 



FLORA OF PERU 417 

shrub of 8 meters, with fiery red fruits (Weberbauer). C. glabrata 
Stapf, Kew Bull. 1906: 76. 1906, of Ecuador, has glabrate, elliptic 
leaves only 3-4 cm. long. Neg. 27874. 

Junin: (Ruiz & Pavori). Huasa-huasi, Ruiz & Pavdn, type; also, 
according to Ruiz & Pavon, at Huarocheri and Canta. Chancha- 
mayo, Isern 2395. Lima: Chorrillos, Raimondi. Libertad: Otuza, 
Weberbauer 6989. 

2. ARJONA Cav. 

Reference: Skottsberg, Svensk Bot. Tidskr. 10: 520-528. 1916. 

Herbs with underground stems or rhizomes, alternate, linear or 
linear-lanceolate leaves, and one to several terminal or subterminal, 
white, lilac, or purplish red flowers. Tube of the perianth elongate 
above the ovary. Placenta thick and short. The name has been 
written variously since its publication as above, in honor of Francisco 
Arjona, a teacher of botany at Cadiz. 

Arjona glaberrima Pilger, Bot. Jahrb. 37: 398. 1906. 

Rhizomes branching, remaining attached to the original plant 
from which rise the 1-2-flowered branches, these scarcely more than 
2 cm. long or much shorter; leaves 2-3 mm. long; flowers glabrous, 
purplish, the tube 6 mm. long, the lanceolate lobes 1.5-2 mm. long. 
In wet cushions of Distichia, etc. Neg. 14343. 

Lima: At 4,500 meters, Weberbauer 5157. 

3. QUINCHAMALIUM J uss . 

Low herbs, often branched at the caudex, from thick roots. 
Flowers mostly terminally aggregate, yellowish green to brown or 
reddish brown. Regarding the name see Pilger, op. cit. 90, in foot- 
note. As this author remarks, "The species are nearly related, difficult 
to separate"; perhaps, it may be added, the species are in reality 
rather forms or races of fewer variable or unstabilized specific entities. 

Anthers barely exserted from the tube Q. brevistaminatum. 

Anthers well exserted to at least a third the length of the lobes. 

Leaves 3-4 (-5) mm. long; perianth tube 5-6 mm. long. 

Q. Stuebelii. 
Leaves mostly longer; flowers often 7-8 mm. long. 

Perianth segments cucullate at the tip. 

Leaves 5-9 mm. long; flowers apparently reddish from 
anthesis . . Q. Raimondii. 



418 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves mostly 1-2 cm. long; flowers at anthesis yellowish 
green .Q. procumbens. 

Perianth segments plane at the tip or merely curved-apiculate. 

Leaves 1.5-2.5 mm. broad Q. lomae. 

Leaves scarcely 1 mm. broad. 

Stems several dm. long Q. elongatum. 

Stems rarely 20 cm. long Q. Hoppii. 

Quinchamalium brevistaminatum Pilger, Notizbl. Bot. Gart. 
Berlin 10: 1030. 1930. 

Stems ascending, about 10 cm. long, sparsely leafy; leaves linear 
or narrowly lanceolate, 1-1.5 cm. long, 1-3 mm. broad; calyculus 
subglobose, 2 mm. across; perianth tube narrow, 4-6 mm. long, the 
divisions scarcely longer than 2 mm.; style slightly, the stamens 
scarcely exserted from the tube; anthers nearly 1 mm. long, subsessile. 
Flowers at first greenish yellow, later reddish brown. 

Lima: Loma formation, railway to San Agustin, 300-400 meters, 
Weberbauer 5718, type. 

Quinchamalium elongatum Pilger, Notizbl. Bot. Gart. 
Berlin 10: 1031. 1930. 

Similar to a robust plant of Q. procumbens, but the leaves nearly 
filiform and the fruiting racemes to 3 cm. long; perianth tube 8.5-10 
mm. long, the divisions 3-4 mm. long; anthers 1.5-2 mm. long. Neg. 
14352. 

Ancash: Ocros, 2,400 meters, Weberbauer 2661, type. Between 
Samanco and Huaras, 3,000-3,400 meters, Weberbauer 3122; 171. 

Quinchamalium Hoppii Pilger, Notizbl. Bot. Gart. Berlin 10: 
1030. 1930. 

Not clearly distinct from Q. elongatum unless somewhat smaller 
in all its parts; stamens attaining three-fourths the length of the 
perianth divisions. Neg. 14354. 

Arequipa: At 3,000-3,500 meters, Hopp 56. 

Quinchamalium lomae Pilger, Notizbl. Bot. Gart. Berlin 10: 
1030. 1930. 

Branches to 30 cm. long, the leaves finally attaining 2 cm.; 
perianth tube 7-10 mm. long, the divisions at least 3 mm. long; 
anthers 1.5 mm. long. Allied by the author with Q. majus Brongn. 



FLORA OF PERU 419 

(Q. chilense var. majus Speg.) with shorter perianth divisions in 
proportion to the tube. Neg. 14356. 

Arequipa: Loma formation, between Atiquipa and Chala, 280 
meters, Weberbauer 7187, type. 

Quinchamalium procumbens R. & P. Fl. 2: 1. pi. 107b. 1799. 
Q. linifolium Meyen ex Walp. Nov. Act. Nat. Cur. 19: Suppl. 1: 
1412. 1843, fide Pilger. 

Stems few to many, a few cm. to 20-30 cm. long, procumbent- 
ascending to suberect from a woody caudex surmounting the stout 
taproot; leaves 1-2 or sometimes 3 cm. long, about 1 cm. wide, 
glabrous; calyculus scarcely 2 mm. thick; flowers soon reddish 
brown, the tube 5-6 mm. long, the anthers reaching to about one- 
half the length of the (2 mm. long) perianth divisions, these apically 
cucullate; anthers 1.5 mm. long. An earlier name may be Q. chilense 
Lam. Encycl. 111. 2: 142. pi. 125. 1797, or a more suitable, var. 
procumbens (R. & P.) Kuntze, Rev. Gen. 3: 284. 1893, but to judge 
from the Lamarck illustration of the type by Jussieu, the name might 
as well apply to some other species, as for instance to Q. brevista- 
minatum. Other names which may apply to the Peruvian forms 
include Q. ericoides Brongn., Q. Dombeyi Brongn., and Q. gracile 
Brongn. in Duperr. Voy. Coq. Bot. 231. pi. 52. 1826-1829, the 
last Latin name used repeatedly by Weberbauer for the Peruvian 
plant. It has been treated also as Q. chilense var. gracile (Brongn.) 
Hook, (fide Kuntze). The Ruiz and Pa von localities were given 
as Huarocheri, Canta, Tarma, and Conception. The above descrip- 
tion is drawn from the Madrid type, without locality. 

Ayacucho: Below Coracora, 2,600 meters, Weberbauer 5793. 
Chavina, 3,300 meters, Weberbauer 5783. Ayacucho, Weberbauer 
5524- Lima: Chilca, Casapalta (Ball, as Q. ericoides). Obrajillo, 
Nee. Canta, 3,100 meters, open bank, Pennell 14599. San Buena- 
ventura, 2,700 meters, open slopes, Pennell 14501. Quebrada de 
San Mateo, Isern 2077. Piura: Huaitara, 2,700 meters, Weberbauer 
5412. Cuzco: Herrera. Urubamba, 3,500-3,700 meters (Weber- 
bauer 181). Puno: Pisacoma (Meyen, type of Q. linifolium). Bolivia. 
"Quinchamoli," "chinchimali," "chinchinami." 

Quinchamalium Raimondii Pilger, Notizbl. Bot. Gart. Berlin 
10: 1031. 1930. 

Branches divaricately spreading; leaves less than 1 mm. broad; 
calyculus 2 mm. thick; perianth divisions 3 mm. long, apically 
short-cucullate, the stamens reaching to about three-fourths their 



420 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

length, the anthers 1.75 mm. long. This seems to be the same as 
Q. procumbens or possibly Q. gracile Brongn. Neg. 14360. 

Lima: Raimondi. 

Quinchamalium Stuebelii Hieron. Bot. Jahrb. 21: 306. 1896. 

Annual, the branches to 2.5 cm. long; leaves papillose, puberulent, 
mucronate; calyculus 2 mm. long, the larger teeth nearly 0.5 mm. 
long; perianth divisions 2 mm. long, scarcely half as wide, the stamens 
shorter, the style equaling the latter. Neg. 14361. 

Puno(?): Tomarape", 4,200 meters, Stuebel 112a. Bolivia. 

53. OPILIACEAE. Opilia Family 
By Paul C. Standley 

Trees or shrubs; leaves alternate, without stipules; flowers very 
small, racemose or paniculate, perfect or unisexual; calyx none or 
poorly developed, not accrescent in fruit; perianth segments 4-5, 
free or more or less united; stamens as many as the petals and usually 
opposite them, inserted upon the disk or at the base of the petals; 
disk 4-5-lobate; ovary free or sunk in the disk, 1-celled, 1-ovulate; 
fruit drupaceous. Only one other genus of the family is known in 
America, in Venezuela. 

1. AGONANDRA Miers 

Slender shrubs or trees, usually glabrous or nearly so; flowers 
minute, dioecious, greenish, racemose, the racemes axillary or borne 
at defoliate nodes; calyx minute, cupular, 4-lobate; petals 4 in the 
staminate flower, the stamens 4, the filaments filiform, exserted; 
petals none in the pistillate flower; ovary glabrous, the stigma sessile, 
discoid. 

Agonandra brasiliensis Miers, Ann. Nat. Hist. II. 8: 172. 
1851, nomen; Benth. & Hook. Gen. PI. 1: 349. 1862. 

A slender tree of 4-8 meters or more, the branches and leaves 
glabrous, the bark pale; mature leaves subcoriaceous, on short 
or elongate petioles, oblong-elliptic or ovate, 4-9 cm. long, 2-4 cm. 
wide, gradually or abruptly acuminate, often long-acuminate, acute 
or obtuse at the base or often contracted and decurrent, the nerves 
obscure; racemes 1-5 cm. long, few- or many-flowered, slender, 
solitary or geminate opposite the leaves, usually densely and minutely 
puberulent, the pedicels 1-4 mm. long; petals 1.5-2.5 mm. long, green; 
drupes oval, red, about 1 cm. long. Neg. 19070. 



FLORA OF PERU 421 

Loreto: Mishuyacu, 100 meters, in forest, Klug 397. Bolivia 
and Brazil to Colombia. 

The South American material referred to this species is decidedly 
variable in its more obvious characters, and it is quite probable that 
a careful revision, with ample material, will show that it is divisible 
into several species. 

54. OLACACEAE. Olax Family 
By Paul C. Standley 

Trees or shrubs; leaves usually alternate and entire, penninerved, 
without stipules; inflorescence commonly axillary and few-flowered, 
the flowers solitary, fasciculate, cymose, or racemose, small, greenish 
or white, regular, perfect or unisexual; calyx small, with 4-6 teeth 
or lobes, sometimes accrescent in fruit; petals 4-6, free or more or 
less united, valvate or slightly imbricate; stamens 4-12, inserted 
with the petals and more or less adnate to them, all fertile or partly 
sterile, the filaments free or rarely monadelphous, the anthers 2- 
celled; disk various; ovary free, 1-celled or imperfectly 3-5-celled; 
ovules usually 2 or 3; fruit drupaceous, commonly 1-celled and 
1-seeded. 

Calyx in fruit becoming greatly enlarged and brightly colored, often 
flat and rotate. Stamens twice as many as the petals; flowers 
fasciculate in the leaf axils 1. Heisteria. 

Calyx little or not at all enlarged in fruit, not brightly colored. 

Stamens free, fewer than the petals. Petals 5-6 2. Liriosma. 

Stamens united to form a tube, of the same number as the petals. 

Stamens united to form a tube about the style. . . .3. Schoepfia. 

Stamens united with the petals 4. Aptandra. 

1. HEISTERIA Jacq. 

Glabrous trees or shrubs with entire, membranaceous or coria- 
ceous, short-petiolate leaves; flowers minute, short-pedicellate or 
sessile, fascicled in the leaf axils; calyx minute, 5-6-dentate or lobate, 
in fruit greatly enlarged and persistent, erect and enclosing the fruit 
or often reflexed and exposing it, usually red or purple, subentire 
to deeply lobate; petals small, more or less villous within; stamens 
usually 10-12, rarely 5-6, hypogynous or adnate at the base to the 
petals; ovary depressed-globose, 3-celled; fruit a drupe, globose or 
oblong, the flesh thin, the endocarp crustaceous. 



422 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Fruiting calyx lobate almost to the base, much longer than the 

drupe H. cauliflora. 

Fruiting calyx subentire or only very shallowly lobate, usually 
shorter than the drupe, often reflexed or at least not enclosing 
the fruit. 

Fruit globose or depressed-globose, as broad as long or broader. 
Calyx spreading in fruit; leaves acute at the base. . . H. pallida. 
Calyx abruptly reflexed in fruit; leaves obtuse or rounded at 

the base H. eurycarpa. 

Fruit ellipsoid or oval, conspicuously longer than broad. 

Fruiting calyx conspicuous, spreading, more than half as long 

as the fruit H. cyanocarpa. 

Fruiting calyx inconspicuous, erect, less than one-third as long 
as the fruit H. densifrons. 

Heisteria cauliflora Smith in Rees, Cycl. 17: No. 2. 1819. 

A shrub or tree 3-10 meters high ; leaves short-petiolate, coria- 
ceous, blades variable, chiefly oblong or oblong-lanceolate, 8-25 cm. 
long, acute to obtuse or narrowly rounded at the base, long-attenuate 
to the apex, lustrous; flowers clustered in the leaf axils or on naked 
branches below the leaves, short-pedicellate or subsessile; pedicels 
in fruit usually short but sometimes much elongate; calyx in flower 
minute, dentate; petals whitish, 2.5 mm. long; fruiting calyx pink 
or red, usually 2 cm. long or more, spreading or rather closely en- 
closing the fruit, lobed almost to the base, the lobes obtuse; drupe 
oval or rounded-oval, about 1 cm. long, longer than broad. Neg. 
27860. 

Junin: Chanchamayo, Isern 2171. Loreto: Rio Itaya, Mexia 
6477. Timbuchi, Alto Rio Nanay, in forest, Williams 996, 1034. 
Rio Masana, Williams 18, 98, 25, 26, 27, 8160. Mishuyacu, near 
Iquitos, 100 meters, King 385, 326, 1363, 906. Pinto-cocha on the 
Rio Nanay, Williams 834, 798, 836, 834- Iquitos, 100 meters, in 
forest, Williams 8058; Killip & Smith 27188, 27190. La Victoria, 
on the Amazon River, Williams 2957. Forest between Rio Nanay 
and Rio Napo, Williams 698. Puerto Arturo, Yurimaguas, edge 
of forest, Williams 5191. Fortaleza, Yurimaguas, in forest, Williams 
4248. Recreo, Yurimaguas, in forest, Williams 4161. Yurimaguas, 
135 meters, in forest, Killip & Smith 27696. Without locality: Ruiz 
& Pavdn 14-52. San Martin: Zepelacio, King 3305. Also in the 
Guianas and doubtless in Amazonian Brazil. "Platina-caspi," 
"huangana caspi." 



FLORA OF PERU 423 

The numerous specimens exhibit remarkable variation in the 
form of their leaves, but they appear to be uniform in fruit charac- 
ters, and these alone seem to be dependable for limitation of species. 
One or two of the numbers listed perhaps do represent distinct 
species, so far as one may judge from leaf form, but unfortunately 
they do not bear fruits. 

Heisteria cyanocarpa P. & E. Nov. Gen. & Sp. 3: 35. pi. 241. 
1845 ; Engl. in Mart. Fl. Bras. 12, pt. 2 : 17. pi. 5, f. 2. 1872. H. nitida 
Engl. op. cit. 18. 1872. 

A shrub or tree 3-9 meters high; leaves short-petiolate, the 
blades somewhat coriaceous, lance-oblong to narrowly elliptic, 
mostly 10-20 cm. long but often larger, obtuse to acute at the 
base, long-attenuate to the apex, usually lustrous; flowers clustered 
in the leaf axils, the pedicels about 2 mm. long, in fruit equaling or 
shorter than the calyx; calyx in flower 1 mm. long, 5-dentate; 
petals whitish, lanceolate, 3 mm. long; fruiting calyx red, orbicular, 
1 cm. long or less, subentire, usually reflexed; drupe blue, about 1 cm. 
long. Negs. 27861, 27865. 

Loreto: Balsapuerto, Klug 2899. Rio Putumayo, Klug 1607. 
Florida, Klug 2161. Rio Santiago, Mexia 6183. Type from Yuri- 
maguas, Mainas, Poeppig. La Victoria, on the Amazon, Williams 
2949, 2745, 2805, 2808. Caballo-cocha, on the Amazon, Williams 
2038. Leticia, on the Amazon, Williams 3169. Pebas, on the Ama- 
zon, in forest, Williams 1572, 1741. Lower Rio Nanay, in jungle, 
Williams 477. Rio Masana, in forest, Williams 86. Rio Maranon, 
near mouth of Rio Tigre, 115 meters, wooded bank, Killip & Smith 
27513, 27536. Huallaga, Yurimaguas, Williams 4661. Mishuyacu, 
near Iquitos, 100 meters, Klug 1138, 414, 1038; Killip & Smith 
29942. Yurimaguas, Killip & Smith 27691. Wooded banks of Rio 
Itaya, above Iquitos, 110 meters, Killip & Smith 29396. Iquitos, 
Kittip & Smith 27399. Without locality: Ruiz & Pavon 14-54- Also 
in the Amazon Valley of Brazil. "Cotoma masacey" (Huitoto name). 

The type of H. nitida is Spruce 4148 from Tarapoto. I have seen 
no material of that number, but the characters upon which the species 
was separated from H. cyanocarpa do not appear to be of taxonomic 
importance. 

Heisteria densifrons Engl. in Mart. Fl. Bras. 12, pt. 2: 17. 

pl.5,f. 1. 1872. 

A glabrous tree; leaves short-petiolate, ovate to elliptic-oblong, 
7-17 cm. long, 3-6 cm. wide, rather abruptly long-acuminate, acute 



424 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

at the base; flowers few, the pedicels very short and thick; calyx 
in anthesis minute, urceolate, very shallowly denticulate, in fruit 
accrescent and cyathiform, about 3 mm. long; fruit obovoid, broadly 
rounded at the apex, almost 1 cm. long. Neg. 19085. 

Loreto: Paraiso, Alto Rio Itaya, 145 meters, Williams 3379. 
Amazonian Brazil. "Parinari" (perhaps an erroneous name). 

Heisteria eurycarpa Standl. Field Mus. Bot. 11: 148. 1936. 

A shrub; leaves short-petiolate, the blades subcoriaceous, elliptic 
or oblong-elliptic, 5-9 cm. long, rounded or very obtuse at the base, 
abruptly narrow-acuminate at the apex, lustrous; flowers densely 
clustered in the leaf axils, the pedicels very short; fruit depressed- 
globose, much broader than high, about 1 cm. wide; fruiting calyx 
apparently green, abruptly reflexed, shallowly lobate, the lobes obtuse. 

Loreto: La Victoria, on the Amazon River, in forest, Williams 
2719, type. 

Heisteria pallida Engl. in Mart. Fl. Bras. 12, pt. 2: 16. 1872. 

A medium-sized tree; leaves short-petiolate, subcoriaceous, the 
blades oblong or lance-oblong, 9-13 cm. long, obtuse or acutish at the 
base, narrowed to the bluntly short-acuminate apex, lustrous; pedicels 
in flower mostly 5-9 mm. long, in fruit longer than the petiole; 
fruiting calyx 1 cm. long, spreading, membranaceous, indistinctly 
lobate; drupe globose, 1 cm. in diameter, pruinose. Neg. 27865. 

San Martin: Type collected near Tarapoto, Spruce 4531. Tara- 
poto, in forest, Williams 6719, 5966, 5813. "Chuchuhuasha," 
"huaami chuchu huasha." 

2. LIRIOSMA P. & E. 

Trees or shrubs with slender branches, glabrous or somewhat 
pubescent; leaves at first very thin, becoming membranaceous or 
subcoriaceous; flowers very small, in short or somewhat elongate, 
simple or branched, axillary racemes; calyx minute, cupuliform, 
truncate, the limb somewhat accrescent in fruit; petals 6, connate to 
the middle; fertile stamens 3, the filaments lanate, becoming gla- 
brate; sterile stamens 6, spatulate, bifid at the apex; ovary 3-celled 
at the base, 3-ovulate; fruit drupaceous, 1-seeded. 

Leaves finely puberulent beneath, soft to the touch . . . .L. adhaerens. 
Leaves glabrous. 

Petals glabrous outside L. gracilis. 

Petals puberulent or pilosulous outside L. Spruceana. 



FLORA OF PERU 425 

Liriosma adhaerens Spruce ex Engl. in Mart. Fl. Bras. 12, 
pt. 2:26. pi 6,f.3. 1872. 

A slender shrub of 2-3.5 meters, the branches sometimes elongate 
and somewhat trailing, minutely puberulent or glabrate, green; 
leaves short-petiolate, thin, oblong-ovate to broadly ovate, 5-9 cm. 
long, 2.5-4 cm. wide, acute or acuminate, acutish to rounded at 
the base, glabrous above or nearly so, minutely soft-puberulent 
beneath; racemes short, longer than the petioles, few-flowered, 
flexuous, puberulent throughout, the pedicels 2 mm. long; petals 
6 mm. long; fruit apparently oval, red, 2.5 cm. long. Negs. 21938, 
19072. 

Junin: Puerto Bermudez, 375 meters, in forest, Killip & Smith 
26475, 26664; flowers green or white. Loreto : Manfinfa, Alto 
Nanay, Williams 1109. Yurimaguas, in forest, Williams 4748. 
San Martin: Tarapoto, Spruce 4207, type collection; Williams 6632, 
5520, 6315. Amazonian Brazil, and reported from Rio de Janeiro. 

Liriosma gracilis A. C. Smith, Bull. Torrey Club 58: 91. 1931. 

A slender shrub or tree 1-5 meters high, sometimes with elongate, 
clambering branches, the branchlets glabrous or sparsely puberulent; 
leaves short-petiolate, ovate or oblong-ovate, mostly 6-8.5 cm. long 
and 2.5^4.5 cm. wide, acute or acuminate, acute to rounded at the 
base, rather thick, glabrous or practically so; racemes 3-6-flowered, 
little exceeding the petioles, the rachis minutely pilose; petals 6-7 
mm. long, glabrous; fruit ellipsoid, about 2 cm. long. The flowers 
are described as white, yellow, or orange-yellow, the fruit as orange- 
yellow. 

Loreto: Mishuyacu, in clearing, 100 meters, Klug 44, type 
collection; 848, 851, 1557. Lower Rio Nanay, Williams 659. Iqui- 
tos, 100 meters, in forest, Killip & Smith 27320. Morona-cocha, 115 
meters, in second-growth, Mexia 6513. Florida, 200 meters, in 
forest, Klug 2014, 2242. "Senimoro-ey" (Huitoto name). 

Liriosma Spruceana Engl. in Mart. Fl. Bras. 12, pt. 2: 24. 1872. 

A slender shrub or tree, 5.5 meters high or less, the green branches 
glabrous or nearly so, often much elongate; leaves thin or rather 
thick, short-petiolate, ovate or oblong-ovate, 7-15 cm. long, 3.5-8 
cm. wide, acuminate, acute or obtuse at the base; racemes short or 
elongate, flexuous, glabrous, mostly 6-8-flowered, the pedicels 4-5 
mm. long; flowers 5 mm. long; fruit ellipsoid, 2 cm. long. Neg. 
27871. 



426 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Puerto Arturo, 200 meters, in forest, Williams 5055, 
5008, 5304. La Victoria, in forest, Williams 2532. Florida, 180 
meters, in forest, King 2293; flowers white. Iquitos, 100 meters, 
Killip & Smith 26990; fruit pale orange. San Martin: Mount 
Guayrapurina, near Tarapoto, Spruce 4884, type. 

3. SCHOEPFIA Schreb. 

Shrubs or small trees; leaves glabrous, membranaceous or coria- 
ceous; flowers few, in short, axillary racemes; calyx small, cupular, 
obscurely 5-denticulate, unchanged in fruit; petals 4-6, inserted on 
the margin of the disk, united for most of their length, with a fascicle 
of hairs within above the middle; stamens as many as the petals and 
opposite them, the filiform filaments adnate to the corolla for almost 
their whole length; ovary partially 3-celled, 3-ovulate; fruit dru- 
paceous, enclosed except at the apex in the accrescent disk, 1-seeded. 

Schoepfia flexuosa (R. & P.) R. & S. Syst. Veg. 5: 160. 1819. 
Haenkea flexuosa R. & P. Fl. 3: 8. pi. 231. 1802. 

A glabrous shrub or small tree, as much as 5 meters high, with 
more or less flexuous branches; leaves somewhat coriaceous when 
dry, short-petiolate, narrowly or broadly ovate, about 6 cm. long 
and 4 cm. wide, acute, almost rounded to subacute at the base, the 
lateral nerves few and distant, the veins obsolete; racemes solitary 
or geminate, less than half as long as the leaves, few-flowered, the 
flowers almost sessile, each subtended by a small, ovate bract, 4-5- 
parted; perianth yellowish, about 6 mm. long, the segments united 
for somewhat more than half their length, the lobes spreading or 
reflexed, glabrous within; drupe said to be "as large as a chick-pea 
(deer)." 

Huanuco: Type collected in the mountains of Pillao and near 
San Antonio de Playa Grande, Ruiz & Pavon. Without locality, 
Ruiz & Pavon. San Antonio de Playa Grande, Ruiz & Pavon 29- 
59. Libertad: Hacienda Membrillal, 1,200-1,400 meters, Weber- 
bauer 6983. 

4. APTANDRA Miers 

Small or large trees, glabrous throughout, with slender branches; 
leaves rather thin, acuminate; flowers in small, axillary panicles; 
calyx very small, patelliform, 4-dentate, fleshy, in fruit slightly 
accrescent and cupuliform, partly enclosing the fruit; petals 4, fleshy, 
almost linear, inflexed at the apex, finally reflexed and spirally con- 
volute; disk composed of squamiform glands; stamens 4, the fila- 



FLORA OF PERU 427 

ments united to form a fleshy tube; ovary 1-celled, 1-ovulate. One 
other species is known from Brazil, and another occurs in western 
Africa. 

Flowers on long, slender pedicels A. Spruceana. 

Flowers short-pedicellate or almost sessile A. tubicina. 

Aptandra Spruceana Miers, Ann. Nat. Hist. II. 7: 202. 1851. 

A small or medium-sized tree, reported to reach a height of 16 
meters, with long, slender branches; leaves short-petiolate, oblong 
or elliptic-oblong, mostly 9-12 cm. long and 4-5 cm. wide, abruptly 
and narrowly obtuse-acuminate, acutish at the base; panicles broad, 
shorter than the leaves, the very numerous white flowers umbellate 
at the ends of the branches, the pedicels sometimes 2 cm. long but 
mostly much shorter; petals 3 mm. long; drupe globose, as much as 
2 cm. in diameter. Illustrated, Miers, Contr. Bot. 1: pi. 1; Mart. 
Fl. Bras. 12, pt. 2:pl.l,f. 2. Neg. 19075. 

Loreto: Santa Rosa, 200 meters, edge of forest, Williams 4798. 
San Ramon, Yurimaguas, 200 meters, in forest, Williams 4550. 
Timbuchi, edge of forest, Williams 964- San Martin: Pongo de 
Cainarachi, 230 meters, in forest, Klug 2698. Amazonian Brazil. 
"Pamashto," "trompo huayo." 

Aptandra tubicina (Poepp.) Benth. ex Miers, Ann. Nat. Hist. 
III. 4: 360. 1859. Heisteria tubicina Poepp. Nov. Gen. & Sp. 3: 35. 
pi. 251. 1845. 

A small tree with slender, green branches; leaves short-petiolate, 
elliptic, 10-13 cm. long, 5-6 cm. wide, linear-attenuate at the apex; 
panicles one-fourth or one-third as long as the leaves, the flowers 
on pedicels only 1 mm. long, subracemose, the pedicels in fruit 
elongate and thickened; fruit blackish, ovoid-globose, 2.5 cm. long. 
Illustrated (flower), Mart. Fl. Bras. 12, pt. 2: pi. 1 J.I. 

Loreto: Type collected in forest near Yurimaguas, Poeppig. 
Amazonian Brazil. 

55. BALANOPHORACEAE. Balanophora Family 

Reference: Harms, Pflanzenfam. ed. 2. 16b: 296-339. 1935. 

Fleshy, yellowish or red root parasites that at least superficially 

suggest some strange fungus rather than a flowering plant. A 

Brazilian species, Lophophytum mirabile Schott & Endl. (illustrated, 

Pflanzenfam. ed. 2. 16b: 325), that reminds one of a sea-anemone 

as much as of a plant, and is represented in Peru by a variety, has 



428 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the local reputation of endowing with magical power (Harms). The 
rhizomes often bear or are more or less enveloped in gall-like struc- 
tures of symbiotic character; cf. Strigl, Sitzungsber. Akad. Wiss. 
Wien 1127-1172. 1908. 

Rhizomes scaly, with deciduous bracts, the elongate flower body 

basally bracteate; perianth lacking 1. Lophophytum. 

Rhizomes ebracteate. 
Flower body without imbricate bracts. 
Flowers borne externally; perianths lacking. 
Staminate flowers on the upper part of the flower body. 

2. Ombrophytum. 
Staminate flowers on the lower part of the flower body or 

absent 3. Juelia. 

Flowers sunk in hair-like paleae, the staminate with a perianth. 

Perianth 3-parted; flower body often oval 5. Helosis. 

Perianth tubular; flower body often elongate. . .6. Corynaea. 
Flower body bracteate, the bracts imbricate 4. Langsdorffia. 

1. LOPHOPHYTUM Schott & Endl. 

Flower body devoid of a cup-like base, whitish to reddish violet. 
Some medicinal use of the Brazilian species has been made, par- 
ticularly in the treatment of rhachitis (Peckolt). 

Lophophytum bolivianum Wedd. Ann. Sci. Nat. III. 14: 185. 
pi. 10. 1850. 

Rhizomes lobed, the cone-like inflorescences 10-30 cm. long, 
sometimes half buried; staminate flowers red, hidden by the sharp, 
erect, brown bracts, the pistillate ebracteolate, 4-5 mm. long; styles 
exserted but deciduous in age. Doubtfully distinct from L. mirabile 
Schott & Endl.; cf. Hook. f. Trans. Linn. Soc. 22: 48. 1856. Tess- 
mann noted the color of the plants as reddish below, shading to light 
brown or white toward the tip. 

Loreto: Mouth of Rio Santiago, Tessmann 4202 (det. Harms). 
Bolivia. 

2. OMBROPHYTUM Poepp. 

Allied to Lophophytum, but the stamens and pistils separately 
borne around a disk or knob-like structure, the flower body basally 
enclosed in a deep cup or volva; lacking the cone-like scales of the 
related genera, the plants are totally different in appearance, these 
suggesting an ear of maize. Styles short, falling promptly. The 



FLORA OF PERU 429 

plants develop rapidly after a shower, enduring only a short time 
(the author therefore called them "rain plants"), or they are soon 
killed, like fleshy fungi, by insect larvae (Poeppig). The authority 
for the generic name is as given! 

Anthers many. 

Volva cup-like, 3-lobed 0. peruvianum. 

Volva entire, tube-like 0. zamioides. 

Anthers few, about 7 0. Ulei. 

Ombrophytum peruvianum P. & E. Nov. Gen. & Sp. 2: 40. 
pi. 155. 1838. 

Yellowish, 10 cm. tall or taller, the broad volva more or less 
3-lobed and as much as 10 cm. deep. Illustrated, Pflanzenfam. 327. 

Huanuco: Cochero (Poeppig). Loreto: Upper Maranon (Tess- 
mann 3994, det. Harms). Rio Acre: ( Ule, fide Harms). Brazil. 
"Maiz del monte." 

Ombrophytum Ulei Harms, Pflanzenfam. ed. 2. 16b: 328. 
/. 165. 1935. 

Distinguished especially by the low staminate disk and the few 
stamens; volva narrow, entire. Illustrated, Pflanzenfam. ed. 2. 16b: 
327. 

Rio Acre: (Ule). Loreto: Rio Maranon above Pongo de Man- 
seriche, 200 meters, Mexia 6337a, on floor in dense forest. 

Ombrophytum zamioides Wedd. Ann. Sci. Nat. III. 14: 184. 
pi. 10. 1850. 

Apparently like 0. peruvianum, but smaller, the tubular volva 
entire. Perhaps here belongs a similar violet-colored plant from 
Tocache which Poeppig mentions. Hooker f. suggests, Trans. Linn. 
Soc. 22: 50, that the plate of 0. peruvianum by Poeppig and End- 
licher is partly from memory, the differences of the Weddell plant, 
therefore, more apparent than real, and reduces the latter name. 
Only further collections can prove this assumption. 

Peru(?): Prov. de las Cordilleras (Weddell). 

3. JUELIA Asplund 

Fleshy, the tuberous rhizome without scales. Flowers ebracteo- 
late, chiefly pistillate, with staminate intermixed toward the base of 
the flower body. Stamens 2. Ovary 1-celled, destitute of free pla- 
centae as in Ombrophytum at first. 



430 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Juelia subterranea Asplund, Svensk. Bot. Tidskr. 22: 274. 1928. 

Rhizome about 5 cm. thick; peduncular portion of the plant equal- 
ing or exceeding the volva; flowers rosaceous or pale rose; filaments 
1-1.5 mm. long, the linear anthers twice as long; styles 1 mm. long. 
The bracts are stiped, with a one-sided, pointed, peltate upper end. 
In the type only one per cent of 100-150 flowers were staminate. As 
the original locality is near the boundary of the Department of 
Puno, the plant in all probability grows also in Peru, with Lepi- 
dophyllum quadrangulare (Meyen) Benth. & Hook., the host plant. 

Peru: (Probably). Bolivia. 

4. LANGSDORFFIA Mart. 

Rhizomes at first tomentulose, more or less contorted, bearing 
1-3 egg-shaped inflorescences subtended by several imbricate rows 
of pointed bracts. Pistillate flowers imbedded in tissue, the free 
staminate ones with a 2-3-parted perianth. Candles prepared from 
the rhizomes, which are rich in wax, are sold in the markets of Colom- 
bia under the name "siejos" (Harms). 

Langsdorffia hypogaea Mart, in Eschwege, Journ. Bras. 2: 178. 
pi. 5. 1818. 

Plants to several cm. high, the flower bodies 2-4 cm. in diameter; 
staminate flowers many, 8-12 mm. long. On roots of palms and fig 
trees (Ficus). Illustrated, Pflanzenfam. ed. 2. 16b: 336. The cita- 
tion has not been verified. 

Huanuco: Cochero (Poeppig). Mexico to southern Brazil. 

5. HELOSIS L. C. Rich. 

Stipes of the roundish flower bodies raised from a short, cup-like 
base of the branching rhizomes. Staminate perianth 3-parted. The 
name has been conserved. 

Helosis cayennensis (Sw.) Spreng. Syst. 3: 765. 1826. Cyno- 
morium cayenense Sw. Fl. Ind. Occ. 1: 13. 1797. H. guyanensis 
Rich. Me"m. Mus. Hist. Nat. Paris 8: 416. pi. 20. 1822. 

Flowering stipes 5-30 cm. long, the basal cup very short; flower 
body ovoid, 2-4 cm. long, the flowers blood-red and more or less 
exserted. Plants bright yellowish or reddish yellow, or the stems 
purplish red. On Myrtaceae. Illustrated, Mart. Nov. Gen. & Sp. 
3: pi. 298, 300. 

Loreto: Mouth of Rio Santiago, Tessmann 4402. Lower Rio 
Napo, Tessmann 3714. Cumaria, Tessmann 5077. (All det. Harms.) 



FLORA OF PERU 431 

6. CORYNAEA Hook. f. 

Similar to Helosis except as indicated. Besides the following, 
the Colombian and Ecuadorian C. Sprucei Eichl. may occur in Peru; 
it may be known by its unisexual flower bodies. The genus is illus- 
trated, Trans. Linn. Soc. 22: pi. 13, 14. 

Corynaea Purdiei Hook. f. Trans. Linn. Soc. 22: 55. 1856. 

Rhizome broad, flattened, lobed, sometimes several dm. long; 
stipes white, 2-2.5 cm. high, 12 mm. thick, bearing a subglobose or 
ovoid, bisexual flower body about 3.5 cm. thick, this at first white 
but soon covered with brown-red, hexagonal, peltate scales. On 
Cinchona roots. The related species being Colombian, the locality 
as Peruvian may be questioned. 

Peru(?): San Juan del Oro (Weddell). Colombia. 

56. ARISTOLOCHIACEAE. Birthwort Family 

Reference: 0. C. Schmidt, Pflanzenfam. ed. 2. 16b: 204-242. 
1935. 

The penguin flower, A. brasiliensis Mart. & Zucc., with blossoms 
15 cm. long, the lower lip as broad Pflanzenfam. /. 122c is 
cultivated at the Lima Botanic Garden (Killip) and may possibly be 
found in Peru. 

1. ARISTOLOCHIA L. 

References: Duchartre in DC. Prodr. 15, pt. 1: 432-498. 1864; 
Schmidt, Repert. Sp. Nov. 23: 282-299. 1927; 292-293. 1930; 30: 
65-75. 1932. 

The determinations of practically all recent collections are by 
Schmidt or have been verified by him. The reference numbers after 
the citations are to the treatment by Duchartre. 

Leaves truncate at the base or very shallowly cordate. 

Leaves subrotund, mostly more than 11 cm. wide (cf. A. bicolor). 

Pubescence of the 5-nerved leaves beneath cobwebby. 

A. maranonensis. 
Pubescence of the 7-nerved leaves beneath merely crisped. 

Flowers purple and blotched about the edge, yellow within, 

about 2 cm. wide at the tip A. Pilgeriana. 

Flowers twice or thrice larger, purple-blotched within. 

A. Ruiziana. 



432 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves deltoid-ovate, mostly narrower than 10 cm. (cf. A. bicolor). 
Leaves mostly very lightly cordate at the base, glabrous or the 

pubescence minute. 

Stipules present (that is, here as elsewhere, pseudo-stipules). 
Leaves glabrous, not sculptured beneath. . . .A. physodes. 
Leaves pubescent, subcircularly sculptured beneath. 

A. deltoidea. 
Stipules none. 

Leaf nerves 7; leaves 11 cm. wide A. bicolor. 

Leaf nerves 5; leaves 6-7 cm. wide A. putumayensis. 

Leaves rounded- truncate at the base, crisp-pubescent beneath. 

A. Guentheri. 

Leaves distinctly cordate, usually cordately lobed, at the base. 
Leaves glabrous beneath or the pubescence microscopic or con- 
fined, as in A. Weberbaueri, to the nerves. 
Leaves ovate, pandurate or hastate. 

Leaves pandurate; flowers large A. pandurata. 

Leaves not pandurate, often ovate or subovate; flowers 

medium-sized. 
Leaves deltoid-cordate; petioles to 4 cm. long. 

A. Weberbaueri. 
Leaves broadly hastate; petioles 5.5-7 cm. long. 

A. mishuyacensis. 
Leaves subrotund. 

Leaf blades 10-20 cm. long. 
Leaves concolorous; flowers about 5 cm. long. 

Perianth lip wide and flaring, purple A. floribunda. 

Perianth lip narrow, yellowish A. cauliflora. 

Leaves glaucous beneath; flowers about twice as long. 

A. didyma. 
Leaf blades 4-8 cm. long. 

Perianth lip rounded, flaring A. elegans. 

Perianth lip split, narrow A. lingulata. 

Leaves distinctly but often finely pubescent all over beneath. 
Stems glabrous. 

Width of leaves usually less than 8 cm. (4.5-8 cm.); flowers 
solitary. 



FLORA OF PERU 433 

Leaves oblongish, 15-18 cm. long; perianth limb bifid. 

A. iquitensis. 

Leaves ovate-cordate, 7-10 cm. long; perianth limb entire. 

A. Rimbachii. 
Width of leaves usually more than 8 cm. (6-18 cm.); flowers 

several. 
Leaves gradually acuminate ; flowers roseate. A . peruviana. 

Leaves short-acuminate; flowers yellowish. 

A. maranonensis. 
Stems pubescent. 

Trichomes of the stems 1.2-2 mm. long, spreading. 
Leaf pubescence beneath short, dense; perianth lip 2-3 cm. 

wide. 
Leaves to 5 cm. long or longer, broadly ovate; perianth 

lip about 2 cm. wide A. amazonica. 

Leaves often larger, ovate-deltoid; perianth lip wider. 

A. truncata. 

Leaf pubescence beneath hirsutulous; perianth lip 8 mm. 
wide A. pilosa. 

Trichomes appressed or short. 
Trichomes subappressed; leaves sometimes pandurate. 

Leaves pandurate A. Burelae. 

Leaves not pandurate A. Macbrideana. 

Trichomes spreading; leaves not pandurate. 

Leaves velvety beneath, the sinus under 1 cm. wide; 
flowers several A. asperifolia. 

Leaves more or less pubescent beneath, the sinus often 
broader; flowers solitary or binate. 

Leaves 5 (-T)-nerved, scabrous or pubescent above; 
perianth limb short-acuminate. 

Leaves deltoid, subobtuse A. Killipiana. 

Leaves ovate, acuminate A. fragrantissima. 

Leaves 3-5-nerved, glabrous above; perianth limb 
I typically long-acuminate A. Mathewsii. 

Aristolochia amazonica Ule, Verh. Bot. Ver. Brandenb. 47: 
122. 1905. 



434 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Stems herbaceous, setose-pilose; petioles 3-5 cm. long; leaves 
sometimes 7 cm. long and nearly as wide, acute, glabrous above, 
shortly and densely tomentose beneath; flowers axillary, solitary, 
the pedicel with ovary 5-6 cm. long; perianth densely pilose, about 
7 cm. long; tube slightly curved above the middle, cylindric, finally 
broadened and expanded to the one-lobed limb, this 2-2.5 cm. long; 
lip broadly elliptic, rounded-obtuse, to nearly 3 cm. long, pubescent 
within, with subulate processes. Remarkable for the long (to 3 cm.) 
utricle. Flowers purple or olive with dark markings, the lip in the 
middle bright brown-yellow, at the tip thickly flecked with purple. 
Neg. 4896. 

Loreto: Cainarachi, Ule 6406, type. Yurimaguas, 135 meters, 
in forest, Kittip & Smith 27583, 27996. El Recreo, 200 meters, edge 
of forest, Williams 3957. Balsapuerto, 220 meters, in forest, Klug 
3042. 

Aristolochia asperifolia Ule, Verh. Bot. Ver. Brandenb. 47: 
121. 1905. 

A high-climbing vine with round and forking stems; petioles 3-4 
cm. long, that is, little longer than the depth of the leaf sinus; 
leaves to 16.5 cm. long and 9 cm. broad, ovate-oblong, acuminate, 
above rough-pubescent, indistinctly 5-nerved; flowers 6-7 cm. long, 
green, in short racemes; tube 2.5 cm. long, pubescent without, ventri- 
cose-ovoid at the base, refracted, recurved, expanding above the 
middle to the cordate-ovate, acutely cuspidate limb, above densely 
glandular; capsule to 6.5 cm. long and 3.2 cm. thick; seeds tri- 
angular, broadly margined, verruculose. Near A. Sprucei Mast., 
but distinguished by its rough leaves, narrowly cordate at the base. 
Neg. 4899. 

San Martin: Tarapoto, Ule 6502 (type). Alto Rio Huallaga, 
Williams 5653. Junin: La Merced, 600 meters, in thicket, 5459. 
Rio Acre: Ule 9340. "Canastilla." 

Aristolochia bicolor Ule, Verh. Bot. Ver. Brandenb. 47: 124. 
1905. 

Herbaceous, glabrous except for the grayish white, finely pubes- 
cent leaves beneath; petiole 5-7 cm. long; leaves triangular-acute, 
thin, dark green above, nearly silvery beneath (but the 7 nerves 
glabrous), 12-13.5 cm. long, 11-12.5 cm. wide; capsule rugose, 
ellipsoid, 5-6 cm. long; seeds 6 mm. long and 3 mm. broad, minutely 
pubescent, acute at the base, incised at the apex. Neg. 4900. 

Loreto: Yurimaguas, Ule. Brazil. 



FLORA OF PERU 435 

Aristolochia Burelae Herzog, Medd. Rijks Herb. Leiden 40: 
2. 1921. 

Slender, the youngest stems, leaves on both sides, and flowers 
tomentose; petioles 1.5-2 cm. long or somewhat longer; leaves 
oblong-lanceolate or narrowly deltoid, acute, 5.5-12 cm. long, 3.5- 
8.5 cm. wide, 5-7-nerved; flowers axillary, solitary; tube 2 cm. long, 
the lip 3 cm. long, 8 mm. wide at the lightly cordate base, narrowly 
oblong-lanceolate, acute, black-hirsute and glandular-punctate with 
yellow-bordered, black spots about the opening. Neg. 4903. 

Rio Acre: Ule 9342. Bolivia. 

Aristolochia cauliflora Ule, Verh. Bot. Ver. Brandenb. 47: 
120. 1905, 

A thick-stemmed, high-climbing liana with cordate-acuminate, 
thick leaves; petioles about 15 cm. long; leaves blunt, with a broad, 
deep sinus, 13.5-22 cm. long, 11-20 cm. broad; flowers few, in 
fascicles, yellowish with lilac-red veins, to 10 cm. long; tube 2-2.5 
cm. long; limb 1-lobed, lanceolate, sparsely pubescent within; lip 
4-6 cm. long but only 2 cm. broad; capsule long-beaked, cylindric. 
The tube throat is yellow at the edge, toward the mouth lilac or 
brown-purple, from the middle to the tip yellow, the tip itself 
flecked with dark purple and black-purple-papillate. Neg. 4904. 

San Martin: Tarapoto, Ule 6634 (type). Chazuta, 260 meters, 
King 4125. Loreto: Almeria, Middle Rio Ucayali, Tessmann 3516. 

Aristolochia deltoidea HBK. Nov. Gen. & Sp. 2: 146. pi. 112. 
1817. Howardia deltoidea Klotzsch, Monatsb. Akad. Berlin 1859: 
611. 1859. 

Stems sulcate, puberulent above and at the nodes; leaves deltoid, 
5-nerved, somewhat pubescent on the nerves beneath but not silvery, 
6-9 cm. long and 5-6 cm. broad; pseudo-stipules orbicular-reniform, 
sessile; flowers axillary, small, greenish without, with brown veins, 
within yellowish, the tip of the lip pale green with dark brown veins; 
utricle ovoid, puberulent, 3 mm. long; tube to 10 cm. long, strongly 
infundibuliform, with one rounded lobe only 3 mm. long, the other 
1.5 cm. long. 

Cajamarca: Maranon Valley, Humboldt 3645 (type). Tabaconas 
Valley in half -xerophy tic shrub, 1,200 meters, Weberbauer 6166. 
Bellavista, in evergreen shrub, 500 meters, Weberbauer 6210. Jaen, 
Raimondi. Amazonas: Bagua, Raimondi. Brazil. 



436 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Aristolochia didyma S. Moore, Journ. Bot. 53: 7. pi. 535. 1915. 

Stems angled, glabrous; petioles 5-10 (-25) cm. long; leaves 
cordate-suborbicular, 5-nerved, glabrous, papyraceous, 10 cm. wide 
and 12 cm. long or sometimes twice as large; flowers 8-10 cm. long, 
purplish, red-lineate, axillary, solitary, the peduncles 6-10 cm. 
long; utricle 3-3.5 (-5) cm. long, the tube 2.5 cm. long, the limb nearly 
twice as long; lower lip densely long-ciliate within, the upper, erect 
lip 2-lobed, the ovate, obtuse segments glabrous without, ciliate 
within; capsule 3.5 cm. long, nearly 3 cm. thick. Emits, as so many 
species, the odor of carrion. Flowers pale yellow, with purple stripes 
and margins (Ule). 

Rio Acre: Uk 9338. Brazil. "Zapato difunto." 

Aristolochia elegans Mast. Card. Chron. n. ser. 24: 301. /. 64. 
1885. 

Flowers solitary from the herbaceous shoots; petioles to 5.5 cm. 
long; blades ovate-reniform, openly but evidently cordate at the base, 
obtuse, glabrous, beneath glaucescent, to 8.5 cm. long and 9.5 cm. 
broad; flowers long-stalked, the slightly distended (2 cm. long) tube 
abruptly bent upward, cream-colored with many purplish brown 
markings within, the golden-yellow throat (6.3 cm. long) surrounded 
by a deep purple blotch, velvety in texture; limb obliquely cordate- 
ovate, 7.5-8 cm. long, 5 cm. wide; utricle 3.4 cm. long, 12 mm. 
thick, within at the base arachnoid-pubescent; gynostemium 6 mm. 
long, the anthers 4 mm. long. Widely cultivated and naturalized 
in tropical lands. 

Junin: In hedge at La Merced, 600 meters, 5322. Santa Rosa, 
Killip & Smith 28923. 

Aristolochia floribunda Lem. 111. Hort. 15: pi. 568. 1868. A. 
juruana Ule, Verh. Bot. Ver. Brandenb. 7: 118. 1905. 

Allied to A. cauliflora, but the thinner, cordate-reniform, acumi- 
nate leaves with a shallow sinus; flowers large, greenish yellow, 
the throat within dark carmine, with white veins, the tube mouth 
greenish yellow, the tube itself 1 cm. long, the throat 4 cm. long 
and nearly as wide; limb cordate-ovate, peltate; peduncles 4 cm. long, 
fasciculate-racemose, the flowers few or as many as 12. Neg. 4919. 

Loreto: Mouth of Rio Santiago, Tessmann 4193. Brazil. 

Aristolochia fragrantissima Ruiz, Mem. Virt. Bejuco Estrella 
46. 1805. Howar dia fragrantissima Klotzsch, Monatsb. Akad. Berlin 
1859: 615. 1859. 



FLORA OF PERU 437 

High climbing or procumbent and diffuse, the very long, sulcate- 
angled branchlets rusty-pubescent; leaves ovate-cordate, with a deep 
sinus, acuminate, above scabrous, beneath densely pubescent, to 
nearly 20 cm. long and half as broad, on petioles 2-5 cm. long; 
peduncles axillary, rarely geminate or ternate; calyx villous within; 
utricle to about 1.5 cm. long, the tube 2 cm. long, broad and little 
ampliate, the lip 3.5 cm. long; capsule oblong, obtusely hexagonous. 
Leaves all only 5-nerved in 3 sheets comprising the type at Madrid, 
the 2 lateral basal nerves much fainter, so the leaves at base are 
prominently only 3-nerved. The measurements are from Ruiz's 
illustration, loc. cit. The illustration in Lambert, Cinchona 173, 
shows the tip of the lip with numerous small warts, placing the 
species (Schmidt) in the Papillatae. The flowers are said to be 
brown-red. Neg. 4914. 

Huanuco: Posuso, Ruiz. "Bejuco de estrella," "contrayerba." 

Aristolochia Guentheri 0. C. Schmidt, Repert. Sp. Nov. 27: 
292. 1930. 

Stems, especially the older ones, erose; stipules none; petioles 
3-4.5 cm. long; leaves 10-15 cm. long, 5.5-9 cm. wide, gradually 
acuminate, 5-7-nerved; flowers 4-5, glabrous, the peduncles 3-6 
cm. long; utricle ellipsoid, 2 cm. long, 8 mm. thick, arachnoid within, 
the tube about 1.5 cm. long, the limb peltiform, spotted with deep 
purple, 2.5-3 cm. wide. Allied with A. Pilgeriana. The flowers of 
the Peruvian specimen are larger, the limb about 4 cm. long and 
more than 3 cm. wide (Schmidt). Neg. 4917. 

Junin : Chanchamayo Valley, 1,200 meters, Schunke 392. Bolivia. 

Aristolochia iquitensis 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 10: 196. 1927. 

Stipules none; leaves sublanceolate, 15-17 cm. long, 5.5-8 cm. 
wide, the sinus of the cordate base broad; bracts narrowly oblong- 
cordate, 1-1.5 mm. long; pedicels 2.5-3.5 cm. long; flower base 
obovoid, 7-9 mm. long, glabrous without, the obliquely truncate tip 
transitional to the erect, cylindric tube 7-8 mm. long, the upper 
lip of the dilated limb consisting of 2 lobes 1.2-1.6 cm. long and 
6-8 mm. wide, these rounded and dilated at the apex; capsule 
narrowly elliptic, 7 cm. long; seeds narrowly oblong-cordate, 5 mm. 
long. Similar to A. lingulata Ule, but the flowers different, these 
bright yellow and red-brown (Klug), basally green, purplish red 
above. 



438 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Iquitos, Tessmann 5120 (type). Mishuyacu, 100 meters, 
in forest, Kittip & Smith 29905; King 66, 1388. 

Aristolochia Killipiana O. C. Schmidt, Repert. Sp. Nov. 38: 
111. 1935. 

Stems slender, more or less sparsely pilose; petioles pilose, to 
2.5 cm. long; leaf blades to 9.4 cm. long, 5 cm. wide, the sinus to 
12 mm. deep and 15 mm. broad, thin, short-pubescent above, pilose 
beneath, especially on the 5-7 prominent nerves; flowers solitary, 
pilose, the utricle obovate-elliptic, the nearly erect tube apically 
dilated and almost bilabiately expanded, the subovate limb acumi- 
nate. Allied to A. triangularis Cham., with glabrous, scarcely 
cordate leaves, the perianth lip not abruptly expanded. Only young 
flowers are known, for which the author gives the measurements: 
peduncle 3 cm. long; utricle 1 cm. long; tube 5 mm. long; lip 13 mm. 
long, 12 mm. broad. 

Cuzco: Machupicchu, about 2,100 meters (Cook & Gilbert 841, 
type). 

Aristolochia lingulata Ule, Verh. Bot. Ver. Brandenb. 47: 
123. 1905. 

Herbaceous, glabrous, with slender, round branchlets; leaves 
cordate-orbicular, acute, glaucous beneath, thick, 8-10 cm. long, 
7.5-9 cm. broad, on petioles 5-6 cm. long; flowers solitary, axillary, 
12-14 cm. long, the pedicel about as long, straw-colored with dark 
purple markings; tube base ventricose, the tube 2 cm. long; limb 
2-lobed, the upper lobe lanceolate, acute, 4.5 cm. long, the lower 
8-9 cm. long, lanceolate-lingulate, constricted toward the tip and 
there expanded; capsule elliptic, 8 cm. long. Neg. 4921. 

San Martin: Tarapoto, Ule 6581 (type). Chazuta, 260 meters, 
Klug 3988. Loreto: Iquitos, Tessmann 5157. Santa Rosa, 135 
meters, in forest, Kittip & Smith 28923. 

Aristolochia Macbrideana Standl., sp. nov. 

Scandens herbacea, caulibus crassiusculis subteretibus pilis 
minutis brunnescentibus adpressis vel valde adscendentibus pilo- 
sulis; folia mediocria membranacea, petiolo gracili 3-3.5 cm. longo 
minute adpresso-pilosulo; lamina late triangulari-ovata 7-12.5 cm. 
longa 4.5-8 cm. lata acute acuminata, basi profunde (ad 2.5 cm.) 
cordata, sinu lato aperto, lobis basalibus late rotundatis, in sicco 
fusca, supra subsparse pilis brevissimis subadpressis brunnescentibus 
conspersa, subtus concolor, undique sed sparse minutissime adpresso- 



FLORA OF PERU 439 

pilosula, basi 5-nervia, nervis elevatis, venulis prominentibus laxe 
reticulatis; flores laxe racemosi, racemis paucifloris, rhachi plus 
minusve elongata, subsessiles, ovario clavato-lineari pilis minutis 
brevissimis patentibus vel subreflexis induto; utriculus obovoideus 
circa 1.8 cm. longus sordide minute adpresso-puberulus, abrupte in 
tubum 18 mm. longum medio 3 mm. crassum constrictus, labio 
(perfecto non viso) ut videtur ovato-oblongo apice obtuso vel 
rotundato extus minute adpresso-puberulo intus sparsissime villoso. 
Flowers violet, green, and yellow. 

San Martin: Chazuta, Rio Huallaga, altitude 260 meters, April, 
1935, G. King 4079; type in Herb. Field Mus. 

Aristolochia maranonensis 0. C. Schmidt, Repert. Sp. Nov. 
23: 296. 1927. 

Slender stems and petioles glabrous, the latter 3-7 cm. long; 
leaf blades glabrate above, densely arachnoid-pilose beneath and 
reticulate, 9.5-18 cm. long, 10-18 cm. wide, the sinus to 18 mm. 
deep; flowers 6 or fewer, greenish yellow, 8-10 cm. long; utricle 
obovoid, 3-3.5 cm. long, puberulent within and without as also the 
(1.5 cm. long) tube; limb broadly ovate, short-acuminate, 8.5-10.5 
cm. long, with 2 brown-purple spots; gynostemium 5 mm. long, the 
lobes lanceolate, the anthers linear; capsule narrowly elliptic, 8 cm. 
long; seeds short-acuminate, 5 mm. long. Neg. 4922. 

Loreto: Puerto Mele"ndez, Tessmann 4865 (type). 

Aristolochia Mathewsii Duchartre in DC. Prodr. 15, pt. 1: 
497. 1864. A. reticulata Seem. Bot. Voy. Herald 193. 1852, not Nutt. 

Branches pubescent; leaves oblong, cordate at the base, 3-5- 
nerved, obtuse or short-cuspidate, 5-12 cm. long, 2.5-3.5 cm. wide, 
glabrous above, pubescent and reticulate beneath; flowers solitary; 
perianth about 8 cm. long, the cordate limb villous without; capsule 
3.5 cm. long. By error Seemann's name has been written "utricu- 
lata" by Duchartre and later students. A. Buchtienii 0. C. Schmidt, 
Repert. Sp. Nov. 27: 292. 1830, from northwestern Bolivia, has 
leaves 12-20 cm. long, 2-4 cm. wide, narrowed to the tip, densely 
short-pilose beneath, and flowers about 6.5 cm. long. 

San Martin: Tarapoto (Mathews). 

Aristolochia mishuyacensis 0. C. Schmidt, Repert. Sp. Nov. 
32: 96. 1933. A. Williamsii 0. C. Schmidt, op. cit. 30: 70. 1932, 
not Rusby, 1910. 



440 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Stems slender, glabrous; petioles 5.5-7 cm. long; leaves broadly 
hastate, 12-18.5 cm. long, 8-10 cm. wide, usually acute, reniform- 
cordate at the base, the broadly rounded lobes 1.5-3 cm. long, 
glabrous, thin; stipules ovate, cordate at the base, acuminate, 25 
mm. long, 18 mm. wide; flowers solitary, axillary, glabrous without, 
the peduncle with the ovary about 9 cm. long; utricle 5 cm. long, 
2.2 cm. thick, glabrous within; upper (outer) part of the limb curved- 
cymbiform, the base truncate, to 5 cm. long, the apex subabruptly 
cordate, the appendage about 14 cm. long; ovary attenuate at the 
base; gynostemium 6.5 mm. long, the stipe 1.5 mm. long. Flowers 
green and red-brown or greenish yellow, and dark violet (Klug). 

Loreto: Fortaleza, Rio Huallaga, 200 meters, Williams 4314 
(type). Mishuyacu, 100 meters, in forest, Klug 1177, 1302. 

Aristolochia pandurata Jacq. Hort. Schoenbr. 4: 49. pi. 497. 
1804; 475. 

Stems sulcately angled; lower leaves mostly pandurate-hastate, 
the upper hastate, acuminate, the deeply cordate base with oblong- 
ovate, obtuse, divergent lobes, 7-nerved, 8-16 cm. long, 3.5-6 
cm. wide; peduncles longer than the petioles, these 3-4.5 cm. long; 
flowers within lineate or reticulate with white or yellow; utricle 22 
mm. long, the tube 10-15 mm. long, the limb 7 cm. long; capsule 
oblong, beaked, 7.5 cm. long. Flowers yellow and whitish with 
black-purple markings (Ule). 

Rio Acre: Ule 9339. Bolivia to Central America and Venezuela. 

Aristolochia peruviana 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 9: 136. 1924. 

Stems below corky-ridged, to 1 cm. thick; petioles slender, 
tortuous, glabrous, 4-5 cm. long; leaves chartaceous, finely silvery- 
pubescent beneath, glabrous above, 5-nerved at the base, cordate- 
ovate, 8-17 cm. long, 6-11.5 cm. wide, the sinus rarely as much 
as 2 cm. deep; flowers glabrous, developing fasciculately from the 
lower stem, 6-10 together, the inflorescence to 6 cm. long; peduncles 
to 2.5 cm. long; limb peltate, 2.5-3.2 cm. long, 1.8-2.5 cm. broad, the 
tube slender, 7 mm. long, 4 mm. thick; utricle oblong-cylindric, 
rounded at the base, 12-15 cm. long, 4-7 mm. wide; perianth within 
deep red with black and white spots, the throat of the tube white, 
without spotted with pale lilac-red, the throat orange; gynostemium 
5 mm. long, with 6 spreading lobes; anthers stipitate, 2 mm. long; 
ovary glabrous. Neg. 4925. 

Loreto: Yarina-cocha, Tessmann 3459, type. 



FLORA OF PERU 441 

Aristolochia physodes Ule, Verb. Bot. Ver. Brandenb. 47: 
119. 1905. 

Similar to A. deltoidea, but the leaves often constricted medially 
and therefore subtrilobate, and somewhat silvery-glaucescent 
beneath; utricle 18 mm. long; tube strongly inequilateral, 6-7 mm. 
long; throat 3-3.5 cm. long, about 2 cm. broad; limb peltate, ovate, 
long-mucronate, the base bilobate-emarginate. Ground color green- 
ish white but the lip within reddish with dark purple marks and 
orange at the throat, the tube itself greenish white. Here would 
key A. pseudotriangularis 0. C. Schmidt, Repert. Sp. Nov. 38: 110. 
1935, with glabrous leaves. Neg. 4927. 

San Martin: Tarapoto, Ule 6582 (type). 

Aristolochia Pilgeriana 0. C. Schmidt, Repert. Sp. Nov. 23: 
297. 1927. 

Similar to A. peruviana but the leaves 7-nerved and with no 
marked sinus; leaf nerves beneath densely and finely pilose; utricle 
pink with purple nerves, purple-pubescent within, usually 2.5 cm. 
long, obovoid; tube to 14 mm. long, 5-6 mm. thick, glabrous; limb 
ovate-suborbicular, cordate at the base, obtuse, mucronate, to 3.5 
cm. long; gynostemium obconic, to nearly 4 mm. long, the linear 
anthers 1.8 mm. long; ovary glabrous. By slip of the pen A. Pilgeri 
0. C. Schmidt, op. cit. 27: 293. 1930. Neg. 4926. 

Loreto: Mouth of Rio Santiago, Tessmann 4374 (type). 

Aristolochia pilosa HBK. Nov. Gen. & Sp. 2: 146. pi 113. 
1817; 454. 

Stems slender, spreading-hirsute with brown hairs; petioles 2.5 
cm. long; leaf blades ovate, cordate or subhastate-cordate, the deep 
sinus with nearly parallel, rounded lobes, glabrous above, 7 cm. 
long, 4-4.5 cm. wide at the base; peduncles 1.5 cm. long; flowers 
sparsely pilose; utricle 22 mm. long, the tube 3 cm. long, the lip 18 
mm. long. 

Peru: (Surely). Bolivia to Central America. 

Aristolochia putumayensis 0. C. Schmidt, Repert. Sp. Nov. 
38: 112. 1935. 

Slender-stemmed and glabrous except the leaves beneath and the 
flowers within; petioles to 5 cm. long; leaves oblong-ovate or deltoid- 
ovate, to 14.5 cm. long, truncate, angulately rounded, short-acumi- 
nate, chartaceous, shortly grayish-silvery-pilose beneath ; flowers few, 



442 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

reddish brown and orange, the peduncles about 6 cm. long; utricle 
narrowly obovoid, arachnoid within, to nearly 3 cm. long and 2 
cm. wide, the suberect tube to over 4 cm. long, its oblique limb over 
2 cm. long; ovary pilose; capsule narrowly elliptic, about 6.5 cm. 
long, very thin. Related to A. Pilgeriana, according to the author. 
A. KlugiiO.C. Schmidt, op. cit. 30: 66, is smaller, the tube only 3 
cm. long but the limb to 7 cm. long and flaring, as in A. iquitensis. 

Loreto: Florida, Rio Putumayo, 200 meters (King 2032). 

Aristolochia Rimbachii 0. C. Schmidt, Repert. Sp. Nov. 23: 
287. 1927. 

Stems slender; petioles 3.5-4.5 cm. long; leaves narrowly cordate, 
the sinus nearly 2 cm. deep, 4 cm. wide, short-cuspidate, chartaceous, 
5(-7)-nerved, more or less densely but very shortly pilose beneath; 
flowers whitish without, glabrescent; utricle obovoid, 10-12 mm. 
long; tube about 12 mm. long, 2 mm. thick, the opening yellow; limb 
narrowly peltiform, 5-7.5 cm. long, 3 cm. broad, glabrous, reddish 
brown with pale reddish nerves; capsule oblong-ovoid, acuminate, 
6 cm. long, 2.5 cm. thick. Near A. odoratissima L., widely dis- 
tributed in South America, and to be expected; its utricle is about 
twice as long, the flowers violet and purple, like those of A. pandurata, 
the capsule 12 mm. thick. 

Peru: (Probably). Ecuador; Bolivia. 

Aristolochia Ruiziana Duchartre in DC. Prodr. 15, pt. 1: 476. 
1864. A. Duchartrei Andre", Le Mouvem. Hortic. 61. 1867; Fl. 
Serres 18: 35. 1869. 

Stem smooth and lustrous; petioles stout, to 12 cm. long; leaf 
blades coriaceous, green and lustrous above, very glaucous, shortly 
crisped-puberulent, and conspicuously reticulate-veined beneath, 
15 cm. long or longer and nearly as wide, short-acuminate, the base 
angled-truncate, scarcely cordate; flowers large, the connate limb 
12 cm. long or longer, the obovoid utricle 7.5 cm. long and 3 cm. 
thick, the tube 3.5 cm. long, the throat 12-18 cm. long. The meas- 
urements (as in other descriptions usually) are by Schmidt, in this 
instance from Colombian material, the type being imperfect. The 
flowers without are yellowish with brown veinlets, the throat within 
thickly flecked with black-purple on a whitish ground, the mouth of 
the tube white. Klotzsch assigned the name in herbarium under 
Howardia. Illustrated, Fl. Serres 18: 35. Neg. 4930. 

Huanuco: Chicoplaya, Ruiz & Pavdn. Colombia. 



FLORA OF PERU 443 

Aristolochia truncata Field. & Gardn. Sert. PL 1: pi 44. 1844. 
Howardia truncata Klotzsch, Monatsb. Akad. Berlin 1859: 610. 1859. 
A. tarapotina Ule, Verh. Bot. Ver. Brandenb. 47: 123. 1905. 

Closely allied to A. amazonica, but the leaves often 10-14 cm. 
long and 7-8 cm. wide; flowers yellowish green, the oval-ovate lip 
purple-splotched within and sparsely papillose with fleshy processes. 
Negs. 4937, 27818. 

San Martin: Tarapoto, 750 meters, Ule 6501; Williams 5454, 
5560, 5570. Brazil. "Oreja de perro." 

Aristolochia Weberbaueri 0. C. Schmidt, Repert. Sp. Nov. 
23:294. 1927. 

A vine with somewhat angled branches; petioles 1.5-4 cm. long; 
leaf blades deltoid-cordate, 4.5-10.5 cm. long, 3-6 cm. broad, the 
sinus 8-12 mm. deep, 7-nerved, glabrescent above, pilose on the 
nerves beneath; stipules ovate, 6-12 mm. long; flowers axillary, 
solitary, glabrous without, 4.5 to nearly 6 cm. long, reddish brown; 
utricle broadly ovate, 12-18 mm. long, sparsely arachnoid within; 
tube 1.5-2.5 cm. long; limb lanceolate, 2.5-3.5 cm. long, about 1.5 
cm. wide, mucronate, lightly pilose at the base; gynostemium 5.5 mm. 
long, the narrowly linear anthers 3 mm. long; ovary puberulent. 
Distinguished by the author from A. fragrantissima by the absence 
of papillae on the lip (but cf. descr.). 

Huancavelica: In grasslands above Colcabamba, 2,200 meters, 
Weberbauer 6444 (type). 

57. RAFFLESIACEAE. Rafflesia Family 
Reference: Harms: Pflanzenfam. ed. 2. 16b: 243-281. 1935. 
Among the most singular plants in the world, these parasites, 
practically without vegetative development other than the flower 
which is produced directly from the roots or stems of the host plant, 
are represented in Peru only by a small-flowered plant on the branches 
of Casearia, or of other shrubs, possibly. It is surprising to learn 
that these tiny flowers, like elongate knobs projecting horizontally 
from branches of the host, are closely related to the strangely formed 
and colored Rafflesias that may measure 1 meter across (see Pflan- 
zenfam. 262 for a photograph of one). 

Besides the following genus, the scarcely distinct Pilostyles Guill. 
may be expected, especially on Leguminosae inhabiting the grass 
steppes. It is separated by having the perianth lobes broadened 
at the base and the placentae indefinitely or poorly developed. 



444 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1. APODANTHES Poit. 

Little plants that practically are only small flowers borne directly 
and divaricately on the trunks and branches of Casearia. Perianth 
divisions free, narrowed at the base, subtended by 2 opposite, tiny, 
scale-like leaves. Placentae 4, broad. 

Apodanthes Caseariae Poit. Ann. Sci. Nat. 3: 422. pi. 26. 1824. 

Flowers about 5 mm. long, waxy white or reddish, often many 
close together. Curiously enough, only the pistillate plants have 
ever been observed. Illustrated, Mart. Fl. Bras. 4, pt. 2: pi. 27. 

Loreto: Sierra de Ponasa, Ule 6937b. Guianas and Brazil. 

58. POLYGONACEAE. Buckwheat Family 

By Paul C. Standley 
Reference: Meisner in DC. Prodr. 14: 1-186. 1857. 

Herbs, shrubs, or trees, sometimes scandent; leaves mostly alter- 
nate and penninerved, entire or essentially so; stipules represented 
by tubular, membranaceous to coriaceous structures (ocreae), these 
persistent or deciduous, sometimes wanting; flowers usually small, 
sometimes large and showy, perfect or unisexual, most often race- 
mose, the racemes simple or paniculate, the flowers clustered in 
distinct nodules; bractlets, if present, more or less connate to form 
ocreolae; pedicels articulate; perianth segments 5-6, free or connate, 
equal or unequal, often accrescent in fruit; stamens 5-10, the fila- 
ments free or connate at the base, the anthers longitudinally dehis- 
cent; ovary free, 1-celled, 1-ovulate; styles 1-3, free or partially 
connate; fruit a lenticular or 3-4-angulate achene, usually enclosed 
in the perianth. 

Plants with tendrils, these terminating the inflorescence. 

1. Antigonon. 
Plants without tendrils. 

Plants herbaceous or merely suffrutescent. 

Perianth segments 6, in 2 series, the inner ones accrescent in 
fruit and enveloping the achene, sometimes with granules 
on the outer surface 2. Rumex. 

Perianth segments 3-5, equal or nearly so, never with granules. 
Flowers perfect; perianth not fleshy 3. Polygonum. 

Flowers dioecious or polygamo-dioecious; perianth fleshy in 
fruit . . . 4. Muehlenbeckia. 



FLORA OF PERU 445 

Plants woody, trees or shrubs, rarely scandent. 

Flowers perfect; perianth segments 5 5. Coccoloba. 

Flowers dioecious; perianth segments 6. 

Inner perianth segments enlarged in fruit, not red; stamens 

numerous 6. Symmeria. 

Outer perianth segments enlarged in fruit, red; stamens 9. 

Achenes obtusely trigonous 7. Ruprechtia. 

Achenes very acutely trigonous 8. Triplaris. 

1. ANTIGONON Endl. 

Plants scandent, herbaceous or suffrutescent, the branches 
angulate; leaves petiolate, cordate, entire or nearly so; ocreae oppo- 
site the leaves, obsolete and scale-like; flowers racemose, opposite 
the leaves, solitary, simple, terminating in a tendril, the flowers 
fasciculate in the raceme; sepals 5, colored, unequal, the 3 outer ones 
cordate, the 2 inner ones narrower; stamens 8, equal; achene included 
in the calyx, 3-angulate. The genus is easy of recognition because 
of the tendril-bearing racemes. 

Antigonon leptopus Hook. & Arn. Bot. Beechey Voy. 308. pi. 
69. 1839-40. 

A large vine, somewhat pubescent; leaves deltoid or broadly 
ovate, deeply cordate at the base, acute to obtuse; flowers at first 
small and inconspicuous but enlarging and finally 1 cm. long or more, 
bright rose-pink. 

Loreto: Iquitos, Williams 3549, 1529, 1354- Yurimaguas, 
Williams 4046. Caballo-cocha, Williams 2286. La Victoria, 
Williams 2818. San Martin: Tarapoto, Williams 5949. Native 
of Mexico and Central America, cultivated commonly for ornament 
in South America, as in most tropical regions, and often becoming 
naturalized. "Lazo de amor," "amor enredado." 

A very handsome vine because of its great abundance of large, 
handsomely colored flowers, which persist for a long time, and are as 
showy in fruit as during anthesis. 

2. RUMEX L. 

By K. H. Rechinger f. 

Reference: K. H. Rechinger f., Die slid- und zentralameri- 
kanischen Arten der Gattung Rumex, Ark. Bot. 26A, No. 3. 1933. 

Flowers dioecious, polygamous, or androgynous; perianth 6- 
parted, herbaceous, the 3 inner segments often larger, enlarging 



446 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

after flowering and enclosing the nutlet, sometimes developing a 
grain on the midrib near the base; anthers 6, inserted in pairs on the 
base of the 3 outer perianth lobes; stigmas 3, penicillate; achenes 
3-angled. 

Flowers dioecious or polygamous; leaves hastate; valves (inner 
perianth segments) not enlarged in fruit R. Acetosella. 

Flowers usually androgynous; leaves not hastate; valves enlarged in 
fruit. 

Valves entire. 

Valves without grains; plants very tall R. peruanus. 

Valves with grains; plants of medium size. 

Leaves short, obovate, thick; plants ascending, low. 

R. cuneifolius. 
Leaves oblong or lanceolate, thin; plants erect, of medium size. 

Lower leaves cordate at the base; flower verticels remote, 
nearly all with a leaf; pedicels not longer than the fruit. 

R. conglomerate. 

Leaves narrowed at both ends; flower verticels not or not 
all remote, only the lowest sometimes with a leaf; 
pedicels about twice as long as the fruit . . . . R. crispus. 

Valves toothed. 

Lower leaves large, deeply cordate at the base, thin; pedicels 
about 2.5 times longer than the fruit, jointed near the base. 

R. obtusifolius. 
Lower leaves small, slightly cordate at the base, thickish; 

pedicels as long as the fruit, jointed at the middle. 

R. pulcher. 
Rumex Acetosella L. Sp. PI. 338. 1753; 47. 

A low, slender perennial with linear or lanceolate, hastate leaves; 
valves entire, not enlarged in fruit, not larger than the nutlet, grain- 
less. A weed of European origin, naturalized nearly throughout the 
world. The subspecies angiocarpus Murbeck, Beitr. Fl. Sudbosn. 
46. 1891, is remarkable in the union of the valves and the nutlets 
into a single body; it is known from Chile, Prov. Cautin, Werdermann 
1254, and from Falkland Islands, Port Stanley, Birger. Illustrated, 
Reichenb. Icon. Fl. Germ. 24: pi. 192. 

Junin: La Quinua, 3,600 meters, clay bank along trail, 2010. 
Colombia; Brazil; Argentina; Chile. 



FLORA OF PERU 447 

Rumex conglomerates Murr. Prodr. Fl. Getting. 52. 1770; 45. 

Lower leaves cordate at the base, plane; branches of the panicle 
divergent; flower verticels nearly all with a leaf, remote; pedicels 
usually not longer than the fruit; valves small, about 2.5 mm. long, 
entire, all bearing large grains. A weed of European origin, natural- 
ized in extratropic parts of the New World, but not so common as 
R. crispus. Illustrated, Reichenb. Icon. Fl. Germ. 24: pi. 166. 

Huanuco: Mito, abandoned river pasture, 2,700 meters, 1557. 
Huanuco, 2,100 meters, ditch banks, 3508. Junin: Tarma, 3,100 
meters, shaded stream banks, Killip & Smith 21864- Lima: Aman- 
caes, Savatier 1602. San Lorenzo Island, Andersson. Venezuela; 
Bolivia; Uruguay; Argentina; Chile. "Acelga." 

Rumex crispus L. Sp. PI. 335. 1753; 44. 

Stem strict, erect; leaves narrowed at both ends, lanceolate, 
acute, crisped; panicle elongate, narrow; pedicels longer than the 
fruit; valves round-ovate, somewhat cordate at the base, entire, 
4-5 mm. long, usually all grain-bearing, but the grains of the same 
perianth often unlike in size or even 2 of them absent. A common 
weed of European origin, introduced to most other parts of the world 
and naturalized in extratropical regions. A polymorphic species. 

Lima: Lima, W. Nation. San Martin: San Roque, 1,400 meters, 
Williams 7777. Huanuco: Mito, weed in pasture, 2,700 meters, 
1683. Bolivia; Argentina; Chile. 

Rumex cuneifolius Campd. Mon. Rum. 95. 1819; 20. 

Widely creeping, with somewhat fleshy, obovate leaves and small 
fruiting panicles; leaves somewhat crisped marginally, the rather 
short petioles and leaf nerves beneath scabrous; branches of the 
panicle few, short; flower verticels usually approximate, without 
leaves; pedicels thick, shorter than the fruit, usually jointed at the 
middle; valves firm, triangular-ovate, entire, 4-5 mm. long, all with 
a prominent grain; ripe achenes dark brown, broadest at the middle, 
2.5 mm. long. This species of peculiar habit can not be confused 
with any other Peruvian one. It is widely spread through a large 
part of southern South America and introduced to some ports of 
North America and Europe. The height of stem, thickness of leaves, 
and size of valves are variable. A hybrid, R. mirabilis Rech. f. 
(R. crispus X cuneifolius} op. cit. 48, is known from Bolivia: La Paz, 
Buchtien 4488. Illustrated, Rech. f. op. cit. pi. 5. 

Arequipa: Arequipa, Holway 766. Viktortal, "La Chorunga," 
1,050 meters, Weberbauer 1447. Cuzco: San Cristobal, 3,450 meters, 



448 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Herrera 2175. Below Cuzco, Rose. Puno: Lake Titicaca, Lechler. 
Junin: Oroya, 3,600 meters, margin of brook, 985. Tacna: Alto de 
Tacora, Isern 2032. Bolivia; Uruguay; Argentina; Chile. "Llague." 

Rumex obtusifolius L. Sp. PL 335. 1753; 46. 

Lower leaves broad, deeply cordate at the base, flat, the upper 
rounded at the base, narrower, lanceolate; branches of the panicle 
divergent; only the lower flower verticels with leaves and remote; 
pedicels slender, to 2.5 times as long as the fruit, jointed near the 
base; valves 5-6 mm. long, usually one of them bearing a grain, with 
2 or 3 often very pronounced teeth on each side. A weed of Euro- 
pean origin, very polymorphic in Europe, introduced to South 
America only as subsp. agrestis (Fries) Danser, Nederl. Kruidk. 
Arch. 1925: 424. 1926 (R. obtusifolius ft agrestis Fries, Novit. Fl. 
Suec. ed. 2. 99. 1828), to which the following specimens belong. For 
a discussion of this variable species see Rechinger f., Vorarbeiten zu 
einer Monographic der Gattung Rumex I, Beih. Bot. Centralbl. 49, 
Abt. 2: 41. 1932. Illustrated, Reichenb. Icon. Fl. Germ. 24: pi. 181. 

Cuzco: Cuzco, Herrera. Colinas de Sacsahuaman, 3,450 meters, 
Herrera 2347. Huanuco: Mito, 2,700 meters, river shore, 1708. 
Lima: Rio Blanco, 3,600 meters, grassy river bank, 820. Brazil; 
Argentina. "Paico." 

Rumex peruanus Rech. f. Ark. Bot. 26A, No. 3: 6. 1933. 

Plant completely smooth, vigorous, the simple, strict, trunk- 
like stem to 2 meters tall or taller, the internodes elongate, with 
short, caducous ocreae; leaves rigid but thin (dried), the radical 
rounded or truncate at the base, elongate-elliptic, plane, broadest 
at the middle, about 2.5 times as long as wide; lateral nerves 
numerous, forming an angle of 60-70 with the midrib; petioles of 
basal leaves one-third to one-fourth shorter than the blade, with 
very large, pale brown basal ocreae; pedicels thin, 2.5-4 times as 
long as the fruit, obscurely jointed near the base; valves truncate at 
the base, roundish, entire, finely reticulate, the midnerve somewhat 
thicker but never grain-bearing; achene 2.5-3 mm. long, dark brown 
when ripe, narrowed at both ends. This species was first described 
from an incomplete specimen in the Berlin Herbarium, consisting 
of three basal leaves, a fragment of stem, and some ripe fruits. 
The upper leaves, inflorescence, and flowers are thus unknown. In 
the form, consistency, and nervation of the basal leaves and the 
grain-bearing valves it is similar to R. tolimensis Wedd. Ann. Sci. 
Nat. III. 13: 262. 1849, from Colombia, which differs especially by 



FLORA OF PERU 449 

its short internodes, stouter stems, and leaves covered by a dense, 
yellowish or brownish pubescence beneath. Illustrated, Rechinger 
f. op. cit. /. 1. 

Peru: Without locality, Weberbauer 5473 (type). Ancash: Pomo- 
pampa, 4,200 meters, in patches in bog holes of dryish flats, 2492. 
Cuzco: Valle de Paucartambo, Hacienda Churu, Herrera 2313. 

Rumex pulcher L. Sp. PI. 336. 1753; 46. 

Lower leaves small, somewhat crisped marginally, cordate at the 
base, often pubescent; branches of the panicle very divergent, often 
intricate in fruit; flower verticels partly with leaves, all remote; 
pedicels thick, not longer than the fruit, jointed at the middle; 
valves toothed, 4.5-6 mm. long, 2.5-4.5 mm. wide, usually all 
bearing a grain, but the grains often unequal in size; achenes 3-4 
mm. long, broadest a little below the middle. A weed originally 
from the Mediterranean Basin, naturalized in regions of the New 
World climatically suitable. A variable species, especially as regards 
shape of the valves; the subsp. eu-pulcher Rech. f. Vorbarbeiten 
Monogr. Rumex I, Beih. Bot. Centralbl. 49, Abt. 2: 25. 1932, with 
usually fiddle-shaped and relatively narrow but long, toothed valves, 
and the subsp. divaricatus (L.) Murb. Beitr. Fl. Siidbosn. 45. 1891 
(R. divaricatus L. Sp. PI. ed. 2. 478. 1762), with leaves nearly ovate 
and relatively broad but short, toothed valves, are known from 
South America and almost surely will be found in Peru. Illustrated, 
Reichenb. Icon. Fl. Germ. 24: pi. 183. 

Peru: Probably. Brazil; Uruguay; Argentina; Chile. 

3. POLYGONUM L. 

Plants herbaceous, erect or prostrate, sometimes scandent; 
ocreae membranaceous or with a green, herbaceous border, some- 
times scarious; flowers small, perfect, green or colored, axillary and 
solitary or aggregate or in racemes or spikes, the pedicels articulate; 
calyx normally 5-parted, the lobes subequal, often gland-dotted; 
stamens usually 8, the filaments subulate; achene included in the 
calyx, lenticular or 4-angulate. 

Flowers axillary, solitary or fasciculate. 

Ocreae very large and conspicuous, deeply lacerate; upper leaves 
linear P. lacerum. 

Ocreae small, inconspicuous, not deeply lacerate; upper leaves 
oblong P. aviculare. 



450 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers in long-pedunculate racemes or spikes. 

Leaf blades shallowly cordate at the base P. Meisnerianum. 

Leaf blades acute to long-attenuate at the base. 
Stems hirsute or hispid. 

Ocreae with a large, green, spreading, herbaceous border. 

P. hispidum. 

Ocreae membranaceous throughout, not green. 

P. peruvianum. 
Stems glabrous. 

Ocreae glabrous throughout, not with marginal bristles. 

P. portoricense. 

Ocreae often strigose, with long marginal bristles. 
Perianth with numerous small, dark glands . . P. punctatum. 
Perianth not gland-dotted P. hydropiperoides. 

Polygonum aviculare L. Sp. PI. 362. 1753. 

Plants glabrous, annual, prostrate and much branched, often 
forming dense mats; leaves short-petiolate, oblong or narrowly 
oblong, mostly 2-3 cm. long, obtuse or acutish, pale green; flowers 
green or whitish, sometimes tinged with pink. 

Lima: Matucana, 2,400 meters, steep, rocky slope, 91. Native 
of the Old World, naturalized almost throughout the cooler regions 
of America. 

The single Peruvian collection is referable here at least in the 
broad concept of the species. It seems better referable to P. avicu- 
lare than to any of the several South American species that are 
closely related but presumed to be indigenous. 

Polygonum hispidum HBK. Nov. Gen. & Sp. 2: 178. 1817. 

A large, coarse perennial, often a meter high, with stout, hispid 
stems; ocreae 1-3 cm. long, densely hispid, with large, spreading, 
green borders; leaves long-petiolate, lanceolate to narrowly ovate, 
8-20 cm. long, long-acuminate, sparsely or densely hispid; racemes 
thick and dense, often paniculate, 2-10 cm. long or larger, the flowers 
pink or deep red; achene lenticular, 4 mm. long, black and shining. 

Reported from Peru (Meisn. in DC. Prodr. 14: 122) upon the 
basis of a Ruiz collection, the locality not known. To be expected 
in the eastern lowlands. Widely distributed in South America, 
ranging northward to Guatemala; growing usually in swampy places. 



FLORA OF PERU 451 

Polygonum hydropiperoides Michx. Fl. Bor. Amer. 1: 
239. 1803. 

Plants erect or ascending, annual or perennial, the slender stems 
glabrous, less than a meter high; ocreae 1-3 cm. long, appressed, 
strigose, with long bristles on the margin; leaves almost sessile, 
lanceolate or linear-lanceolate, 5-15 cm. long, long-attenuate, 
strigose at least on the margins and costa; racemes erect, dense, 
10 cm. long or less, the flowers usually deep pink; achene more 
or less trigonous, sometimes almost lenticular. Illustrated, Mem. 
Dept. Bot. Columb. Coll. 1: pi. 27. 

Cajamarca: Celendin, 2,625 meters, Woytkowski 10; a common 
weed, filling ditches and marshy spots on plain. Cuzco: Valle del 
Paucartambo, 3,000 meters, Herrera 3861 ; at 3,400 meters, Hacienda 
Churu, Herrera 2330. Huanuco: Mito, 2,700 meters, between rocks 
along river, 1545. Lima: Chosica, 900 meters, edge of ditch, 2861. 
Widely distributed in the cooler regions of America, also in the tropics. 

Polygonum lacerum HBK. Nov. Gen. & Sp. 2: 179. 1817. 

Plants apparently erect and perennial, inclined to be suffrutescent 
at the base, glabrous, pale, the stems branched, striate; ocreae often 
equaling the upper internodes, very conspicuous, whitish, scarious; 
upper leaves linear, fleshy-coriaceous, the lower ones linear-oblong, 
obtuse or acutish, short-petiolate, mostly 2-3 cm. long; flowers 
axillary, inconspicuous. Neg. 4977. 

Cajamarca: Type collected at hot springs near Cajamarca, 
Humboldt. Bolivia and Argentina. 

Polygonum Meisnerianum Cham. & Schlecht. Linnaea 3: 
40. 1828. 

Plants very slender, perennial, subscandent, the stems sparsely 
glandular-hispidulous and with larger, recurved, prickle-like hairs 
at the nodes; leaves sessile or nearly so, linear or lance-linear, 5-15 
cm. long, 5-15 mm. wide, attenuate, aculeolate beneath along the 
costa, elsewhere glabrous or nearly so; ocreae naked at the margin; 
inflorescences dichotomous, the racemes few, few-flowered, the 
peduncles glandular; perianth pink; achenes 3-angulate, lustrous. 

San Martin: Zepelacio, near Moyobamba, 1,200-1,600 meters, 
Klug 3580; flowers cream-colored. Argentina to Mexico and south- 
eastern United States. 

Polygonum peruvianum Meisn. in DC. Prodr. 14: 122. 1857. 

Plants erect, slender, the stems glandular-hispid below the nodes, 

finally glabrate; ocreae 2.5 cm. long, strigose; leaves oblong-lanceo- 



452 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

late, acuminate, subsessile, appressed-pilose, 7-10 cm. long, acute 
to rounded at the base; racemes geminate, oblong, 2.5-3.5 cm. long, 
the flowers large; achene lenticular. 

Type collected at some unspecified locality in Peru, Mathews 
3120. Amazonas: Chachapoyas, Weberbauer 4436 (fide Macbride). 

Polygonum portoricense Bert, ex Small, Mem. Bot. Columb. 
Coll. 1: 46. 1895. P. glabrum Cham. & Schlecht. Linnaea 3: 46. 
1828, non Willd. 1799. 

A stout, glabrous perennial, sometimes 1.5 meters high; ocreae 
very large, often as long as the nodes; leaves slender-petiolate, 
the blades lanceolate, 15-30 cm. long, long-acuminate, with con- 
spicuous lateral nerves; racemes slender, 5-13 cm. long, dense, 
erect, the flowers white or pink; achenes lenticular or 3-angulate, 
black and shining. 

Loreto: Pebas, Williams 1994- Iquitos, 120 meters, Williams 
8086. Caballo-cocha, aquatic, Williams 2455. Rio Masana, 
Williams 3. Extending to Argentina, the West Indies, and southern 
United States. "Tabaco de lagarto." 

Polygonum punctatum Ell. Bot. S. C. & Ga. 1: 455. 1817. 
P. acre HBK. Nov. Gen. & Sp. 2: 179. 1817, non Lam. 1778. 

Plants slender, annual or perennial, glabrous or nearly so, erect 
or ascending, usually less than 1 meter high; ocreae appressed, 
fringed with long bristles; leaves short-petiolate, linear-lanceolate, 
mostly 5-10 cm. long, acuminate; racemes very slender, 2-8 cm. 
long, the nodes remote, the flowers greenish white; achene lenticular 
or 3-angulate, black and lustrous. Illustrated, Mem. Bot. Columb. 
Coll. I: pi. 31. 

Junin: Puerto Bermudez, 375 meters, in thickets, Killip & Smith 
26682. Loreto: Yurimaguas, 200 meters, edge of small stream, 
Williams 3842. Mishuyacu, 100 meters, in clearing, Klug 338. 
Iquitos, 120 meters, Williams 1410, 7984. San Martin: San Roque, 
in pasture, 1,400 meters, Williams 7691. Tarapoto, Williams 5657. 
Widely distributed in tropical America, often a common weed, 
extending northward to the United States. "Yacu shutiri." 

4. MUEHLENBECKIA Meisn. 

Shrubs or suffrutescent plants, the stems usually scandent or 
prostrate, angulate or sulcate, the ocreae membranaceous, obliquely 
truncate, cylindric, usually soon deciduous; leaves petiolate, often 
cordate or sagittate; flowers small and inconspicuous, dioecious or 



FLORA OF PERU 453 

polygamo-dioecious, whitish or greenish, solitary or fasciculate in 
the leaf axils or racemose or paniculate, the pedicels articulate; 
calyx herbaceous, 5-parted, the lobes subequal or the 2 inner ones 
smaller, in fruit more or less accrescent, sometimes thick and suc- 
culent; stamens 8 in the staminate flower, about equaling the sepals; 
styles 3 and short or none; achene 3-angled, enclosed in the fleshy 
perianth or more or less exserted. The oldest name for the genus 
is Calacinum Raf., and the name Muehlenbeckia is not conserved. 
Since the latter has been in general use for the group, it seems 
preferable to continue its use, in the confident expectation that it 
will be conserved. 

Flowers in sessile glomerules; leaves mostly acute or attenuate at 
the base, never cordate or hastate-lobate. 

Plants usually prostrate and matted; leaves small, mostly less 
than 15 mm. long and acute or acutish, sometimes obtuse 
or rounded M. volcanica. 

Plants erect or scandent; leaves mostly larger and 2 cm. long 
or more, but sometimes small, broadly rounded to very 
obtuse at the apex. 

Plants erect; leaf blades mostly conspicuously longer than broad. 

M. fruticulosa. 

Plants scandent; leaf blades mostly as broad as long. 

M. Nummularia. 

Flowers in simple or paniculate racemes; leaves cordate at the base 
or more or less evidently hastate-lobate. 

Leaves not cordate at the base, more or less hastate-lobate; 
racemes simple, often greatly reduced M. hastulata. 

Leaves cordate at the base; racemes simple or paniculate. 
Racemes simple; leaves often puberulent beneath . .M. peruviana. 
Racemes paniculate; leaves glabrous or puberulent beneath. 

Leaves densely puberulent or tomentulose beneath; achenes 
often semi-exserted M. tiliifolia. 

Leaves glabrous; achenes covered by the sepals . . M. tamnifolia. 

Muehlenbeckia fruticulosa (Walp.) Standl., comb. nov. Poly- 
gonum fruticulosum Walp. Nov. Act. Acad. Leop. 19: Suppl. 1: 
407. 1843. M. rupestris Wedd. Ann. Sci. Nat. III. 13: 256. 1850. 
Sarcogonum fruticulosum Rusby, Mem. Torrey Club 4: 251. 1895. 
Calacinum fruticulosum Macbr. Field Mus. Bot. 4: 115. 1927. 



454 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A glabrous, rather densely branched, erect shrub about a meter 
high; ocreae small, often rather long-persistent; leaves petiolate, 
the blades thick and fleshy or coriaceous, elliptic-oblong to broadly 
oval, the larger ones 1.5-2.5 cm. long, rounded or very obtuse at 
the apex, cuneately narrowed at the base; flowers short-pedicellate, 
few or numerous at each node; achene included in the perianth. 
Neg. 4981 (M. rupestris). 

Arequipa: Yura near Arequipa, 2,600 meters, Weberbauer 6840. 
Moquehua: Carumas, 3,100 meters, Weberbauer 7483. Puno: 
Puno, 4,000 meters, Soukup 372. Lake Titicaca, Meyen. Sandia(?) : 
Pascomayo to Moyobamba (Stuebel 37a). Bolivia. 

It is doubtful whether this is more than an ecological form 
of M. vokanica, and it should probably be reduced to varietal rank 
under that species. 

M uehlenbeckia hastulata (Smith) Standl., comb. nov. Rumex 
hastulata Smith in Rees, Cycl. 29. 1802-20. M. chilensis Meisn. 
in DC. Prodr. 14: 148. 1856. M. chilensis var. fascicularis Meisn. 
loc. cit. Sarcogonum chilense Rusby, Bull. Torrey Club 27: 128. 
1900. Calacinum chilense and C. hastulatum Macbr. Field Mus. 
Bot. 4: 116. 1927. 

Plants woody, scandent, pale green when dried, glabrous; ocreae 
large and brown, conspicuous, rather long-persistent; leaves fleshy- 
coriaceous, rather long-petiolate, the blades broadly hastate-ovate 
to linear-hastate, truncate to attenuate at the base, obtuse to 
acuminate; flowers short-pedicellate, in axillary and terminal, short 
or somewhat elongate racemes, the pedicels short; achene included 
in the perianth or somewhat exserted. Negs. 7431, 27770. 

Arequipa: Above Arequipa, open, rocky slopes, 2,800 meters, 
Pennell 13246. Near Arequipa, Lechler 2746.- Cuzco: Pachar, bushy 
bank, 2,900 meters, Pennell 13691. Urubamba, Weberbauer 4914- 
Junin: Yanahuanca, 3,000 meters, 1221. Bolivia and Chile. 
"Huano negro." 

The plant is extremely variable in leaf form, and some of the 
forms are perhaps worthy of varietal rank. 

Muehlenbeckia Nummularia H. Gross, Bot. Jahrb. 49: 346. 
1913. 

A much branched, scandent, glabrous shrub; ocreae rather long- 
persistent; leaves petiolate, the blades coriaceous, broadly oval to 
orbicular, 8-20 mm. long, 8-17 mm. wide, broadly rounded and 



FLORA OF PERU 455 

sometimes apiculate at the apex, abruptly decurrent into the petiole; 
flowers densely fasciculate in the leaf axils; achene enclosed in the 
perianth. 

Ancash: Below Hacienda Cajabamba, between Samanco and 
Caraz, 3,000-3,500 meters, Weberbauer 3172. 

This is probably only a form or variety of M. fruticulosa or M. 
volcanica. There is known only the type collection, of which I have 
seen fragmentary material. 

Muehlenbeckia peruviana Meisn. in DC. Prodr. 14: 150. 1856. 
Calacinum peruvianum Macbr. Field Mus. Bot. 4: 117. 1927. 

A scandent shrub, the branches glabrous or somewhat tomentu- 
lose; ocreae conspicuous and long-persistent; leaves slender-petio- 
late, the blades subcoriaceous, ovate or oval-oblong, mostly 3.5-5 
cm. long, obtuse or subacute, puberulent beneath or almost glabrous, 
shallowly cordate at the base; racemes short and dense, usually 
shorter than the leaves, sometimes greatly reduced; achenes more 
or less exserted. Neg. 4980. 

Junin: Huasa-huasi, Dombey210 (type); Ruiz&Pavdn. Cuzco: 
Paso de Tres Cruces, 3,500-3,800 meters, in thickets, Pennell 13894. 
Yanamanche, 3,500 meters (Weberbauer 4951). 

Muehlenbeckia peruviana var. cuspidata Standl., var. nov. 
M. cuspidata H. Gross in herb. 

A forma typica foliis solemniter acuminatis vel longiacuminatis 
differt. Apparently differentiated from the typical form only by 
the conspicuously acuminate leaves. 

Ancash: Pichin, Weberbauer 2929 (type, a fragment of this num- 
ber in Herb. Field Mus., received from Herb. Berlin). Huanuco: 
Monzon, Weberbauer 3373. Without locality: Ruiz & Pawn 13-95. 
Also in Ecuador (Loja-Zamora, Andre 4521). 

Muehlenbeckia tamnifolia (HBK.) Meisn. Comm. 2: 227. 
1840. Polygonum tamnifolium HBK. Nov. Gen. & Sp. 2: 180. 1817. 
M. tamnifolia var. laxiflora Meisn. in DC. Prodr. 14: 149. 1856. 
M. leptobotrys Meisn. loc. cit. Sarcogonum tamnifolium Rusby, 
Mem. Torrey Club 6: 111. 1896. Calacinum tamnifolium and C. 
leptobotrys Macbr. Field Mus. Bot. 4: 116. 1927. 

A large, scandent shrub, glabrous throughout or nearly so; 
ocreae large, scarious, deciduous; leaves on short or elongate petioles, 
the blades firm-membranaceous, broadly ovate or oval, mostly 
5-9 cm. long, acuminate to rounded and abruptly cuspidate-acumi- 



456 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nate, usually deeply cordate at the base; racemes mostly elongate 
and arranged in lax panicles, densely or sparsely flowered; achene 
included in the perianth. Negs. 4983, 4979. 

Arequipa: Quequena, Isern 2036. Cajamarca(?) : Palco, Ruiz & 
Pavdn. Above San Pablo (Weberbauer 3806). Huanuco: Ambo, 
2,100 meters, 3166. Huanuco, Ruiz. Without locality: Ruiz & 
Pavdn 13-97. Junin: Huacapistana, Weberbauer 2175 (determined 
by H. Gross as a new species). Sandia: (Weberbauer 648, 835). 
Ancash: Chiquian (Weberbauer 2826). Ancash (Weberbauer 3164). 
Northwestern Argentina and Bolivia to Colombia and Venezuela; 
Mexico and Central America. 

M. leptobotrys (type collected near Huanuco by Ruiz; Coccoloba 
carinata Ruiz in herb, ex Meisn. in DC. Prodr. 14: 150. 1856) is 
merely a form with lax inflorescences. It may be called M. tamnifolia 
var. laxiflora Meisn., although it scarcely seems to deserve special 
designation. 

Muehlenbeckia tiliifolia Wedd. Ann. Sci. Nat. III. 13: 255. 
1850. Calacinum tiliifolium Macbr. Field Mus. Bot. 4: 117. 1927. 

A large, more or less woody vine; ocreae very large and con- 
spicuous, often long-persistent; leaves thin, slender-petiolate, the 
blades ovate or ovate-oval, mostly 5-13 cm. long, rounded and 
mucronate to abruptly cuspidate-acuminate at the apex, deeply 
cordate at the base, green and glabrate on the upper surface, densely 
puberulent or tomentulose beneath and often grayish; racemes 
elongate, usually very dense, in large or small panicles; achene often 
exserted from the perianth. 

Ancash: Ocros, Weberbauer 2733. Pampa Ramos (Weberbauer 
3187). Cuzco: Marcapata, 3,100 meters, Weberbauer 7777, flowers 
white; 7778.' Huanuco: Huacachi, 1,950 meters, in clearing, 4165, 
4189; fruits red. Lima: Viso, 2,700 meters, trailing over rocks and 
shrubs, 754' Near Viscas, 2,100 meters, thickets along river, fruit 
purple-black, Pennell 14447. Bolivia. 

Muehlenbeckia volcanica (Benth.) Endl. Gen. Suppl. 4, pt. 2: 
51. 1847. Polygonum volcanicum Benth. PI. Hartw. 81. 1841. Sarco- 
gonum volcanicum Rusby, Mem. Torrey Club 4: 252. 1895. Cala- 
cinum volcanicum Macbr. Field Mus. Bot. 4: 115. 1927. 

Plants depressed and forming small, dense mats, or sometimes 
ascending, the branches usually 30 cm. long or less, woody, glabrous 
throughout; ocreae small and soon deciduous; leaves subsessile, 



FLORA OF PERU 457 

fleshy-coriaceous, mostly rhombic-elliptic, acute or obtuse, cuneate- 
attenuate at the base, mostly 7-14 mm. long; flowers small, green, 
fasciculate in the upper leaf axils, short-pedicellate; achene enclosed 
in the perianth. Illustrated, Wedd. Chlor. And. 2 : pi. 89. Neg. 7432. 
Ayacucho: Pampalca, 3,200 meters, open hillside, Killip & Smith 
23253. Ancash : Tallenga, 3,600 meters (Weberbauer 2870) . Huaraz 
(Weberbauer 3228). Cuzco: Colinas del Sacsahuaman, 3,500 meters, 
Herrera 2350. Paso de Tres Cruces, 3,900 meters, rocky banks, 
edge of paramo, Pennell 13837; fruit black. Huanuco: Six miles 
south of Mito, 3,000 meters, stony, open slope, 1834. Tambo de 
Vaca, 3,900 meters, mossy, rocky upland, 4406. Junin: Carpapata, 
2,400 meters, open hillside, Killip & Smith 24344', stems as much 
as 60 cm. long, prostrate. La Quinua, 3,600 meters, 2022. Lima: 
Rio Blanco, 4,500 meters, 3002. Canta, 2,700-3,200 meters, Pennell 
14339. Huaros, 3,400 meters, rock slide, Pennell 14714. Rio 
Blanco, 3,000-3,500 meters, creeping over rocks, Killip & Smith 
21546. Matucana, 2,400 meters, 245. Puno: Araranca, 4,200 
meters, ledges of siliceous rock, Pennell 13453. Asangaro, Lechler 
1748. Sandia: Cuyocuyo (Weberbauer 849). Without locality: 
Ruiz & Pavon. Bolivia to Ecuador. "Mullaca," "pasamullaca," 
"zoczocma." 

5. COCCOLOBAL. 

Reference: Lindau, Bot. Jahrb. 13: 106-229. 1890. 

Trees or shrubs, usually glabrous or nearly so; ocreae coriaceous- 
membranaceous, cylindric, eciliate, truncate, deciduous; leaves often 
deciduous, usually coriaceous; flowers perfect, in spike-like, axillary 
or terminal, simple or rarely branched racemes, the bracts ocreiform, 
subtending several flowers, the pedicels short or elongate, articulate 
at the apex; calyx green or whitish, small, the 5 subequal segments 
united at the base, the tube or the lobes accrescent and enclosing 
the fruit, usually becoming much thickened and succulent; stamens 
8, equal; achene subtrigonous-globose, small or large. The genus 
is in need of critical revision in the light of recently accumulated 
material. Many of the species seem to be based upon vague or 
variable characters. The fleshy calyces at maturity are often very 
juicy and edible. 

Flowers in panicled racemes C. mollis. 

Flowers in simple racemes. 

Calyx tube accrescent and enclosing the achene, the calyx lobes 
very small. 



458 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Leaves conspicuously short-pilose beneath, at least when 
young, short-acuminate, more or less cordate at the base. 

C. excelsa. 
Leaves glabrous beneath, or barbate along the costa in the axils 

of the nerves. 
Bracts and ocreolae 0.5 mm. long. Leaves large, obovate, 

cordate at the base C. nutans. 

Bracts and ocreolae more than 1 mm. long. 
Veinlets conspicuously elevated and closely reticulate on 

the upper leaf surface C. sphaerococca. 

Veinlets inconspicuous on the upper leaf surface. 

C. Barbeyana. 

Calyx lobes accrescent and enclosing the achene. 
Leaves glabrous beneath, rounded at the apex, 4-7 cm. long. 

C. Ruiziana. 

Leaves more or less pubescent beneath, at least on the costa, 
or barbate in the axils of the nerves. 

Leaves large, mostly 7-12 cm. wide C. Williamsii. 

Leaves smaller, chiefly 2-5 cm. wide. 
Leaves narrowly lance-oblong, long-acuminate. 

C. acuminata. 
Leaves elliptic to obovate, obtuse or acute. 

Rachis of the inflorescence glabrous or nearly so; leaves 

mostly acute C. peruviana. 

Rachis of the inflorescence short-pilose; leaves obtuse. 

C. gracilis. 

Coccoloba acuminata HBK. Nov. Gen. & Sp. 2: 176. 1817. 

A slender shrub or tree 2-6 meters high, reported to attain a 
height of 10 meters, the elongate branchlets glabrous or ferruginous- 
pubescent; leaves short-petiolate, the blades narrowly lance-oblong, 
long-acuminate, 12-20 cm. long, acute or acutish at the base, more 
or less puberulent beneath in the axils of the nerves, otherwise 
glabrous; racemes slender, spike-like, longer than the leaves, the 
rachis puberulent, the pedicels very short; fruits 6 mm. long, pink 
or bright red. Neg. 4939. 

Loreto: Yurimaguas, 200 meters, in forest, Williams 4651, 4606. 
Portal, 135 meters, Kittip & Smith 29263. Puerto Arturo, 135 
meters, Killip & Smith 27879; edge of forest, Williams 5271. 



FLORA OF PERU 459 

Santa Rosa, 200 meters, in forest, Williams 4804- Contamana, 150 
meters, dense forest, Killip & Smith 26879. Extending northward 
to Central America. 

A well marked species because of the narrow, relatively thin, 
long-acuminate leaves. A handsome shrub when in fruit because of 
the great profusion of bright red racemes. 

Coccoloba Barbeyana Lindau, Bot. Jahrb. 13: 185. 1890. 

A shrub or tree, sometimes 8 meters high, glabrous throughout, 
with stout branches; leaves short-petiolate, coriaceous, the blades 
obovate-oblong to elliptic-oblong or obovate, mostly 9-27 cm. long, 
obtuse to acuminate or rounded and abruptly short-acuminate at 
the apex, obtuse or rounded at the base and sometimes shallowly 
emarginate, the lateral nerves very oblique, the veinlets promi- 
nulous and closely reticulate beneath; racemes rather slender, 
shorter or longer than the leaves, the pedicels slightly longer than 
the ocreolae, or sometimes more elongate; fruit subglobose, almost 
1 cm. long. Flowers described as cream-colored, white, or green. 

Type collected in Peru by Ruiz and Pavon, the locality unknown. 
Loreto: Puerto Arturo, 200 meters, in forest, Williams 5138. 
Florida, 180 meters, in forest, King 2260. Mishuyacu, 100 meters, 
in forest, Klug 1077. Yurimaguas, 200 meters, Williams 4528. 
Iquitos, 100 meters, Killip & Smith 27494- Rancho Indiana, 
overflowed creek bank, 110 meters, Mexia 6426. Rio Huallaga, 
135 meters, Killip & Smith 29006. San Martin: Chazuta, 260 
meters, in forest, Klug 4127. "Nemono-o" (Huitoto name). 

Goccoloba excelsa Benth. in Hook. Lond. Journ. Bot. 4: 
624. 1845. 

A scandent shrub, the young branches ferruginous-puberulent 
or glabrous; leaves short-petiolate, the blades coriaceous, broadly 
ovate to oval or rounded-obovate, 15-22 cm. long, obtuse to rounded 
at the apex and usually abruptly acuminate, slightly narrowed to 
the shallowly cordate base, glabrous above or nearly so, short-pilose 
beneath, at least when young, the veinlets prominent beneath and 
closely reticulate; racemes slender, shorter than the leaves, the 
rachis usually puberulent, the nodes 1-3-flowered, the pedicels 
equaling or longer than the ocreolae; fruit globose, 9 mm. long. 
Neg. 4958. 

Loreto: Yurimaguas, edge of forest, 200 meters, Williams 4538. 
Amazonian Brazil, Surinam, and British Guiana. 



460 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Coccoloba gracilis HBK. Nov. Gen. & Sp. 2: 176. 1817. C. 
peruviana Willd. ex Lindau, Bot. Jahrb. 13: 214. 1890, in syn. 

A shrub or tree of 3.5-6 meters, the branchlets stout, glabrous; 
leaves short-petiolate, coriaceous, the blades oblong-elliptic to ellip- 
tic, 5-7 cm. long, 2-4 cm. wide, obtuse, rounded or obtuse at the 
base, the veinlets scarcely prominent beneath, closely reticulate; 
racemes lax or dense, slightly longer than the leaves, the rachis 
short-pilose or puberulent, the nodes mostly 1-2-flowered, the pedi- 
cels 2 mm. long. Neg. 4945. 

Loreto: Rio Cachiyaco, Humboldt, type. Without locality: 
Weberbauer 6982. 

Coccoloba mollis Casar. Nov. Stirp. Bras. Dec. 8: 72. 1842-45. 
C. polystachya Wedd. Ann. Sci. Nat. III. 13: 261. 1850. 

A small or medium-sized tree, the branches glabrous; leaves 
short-petiolate, subcoriaceous, broadly ovate to oblong-ovate, 12-25 
cm. long, acuminate or long-acuminate, cordate to truncate at the 
base, minutely puberulent or glabrate; racemes slender, paniculate, 
the panicles often very large, the rachis tomentulose, the nodes 
1-flowered; flowers white; fruit ovoid, 1 cm. long. Neg. 21412. 

Loreto: Florida, 200 meters, in forest, Klug 1991. Brazil, 
Ecuador, and the Guianas. "Tangarana" (Klug; signifying ant tree, 
and presumably indicating that the tree is inhabited by biting ants). 

On young, sterile plants, as indicated by Ecuador material, the 
leaves are often extremely large, as much as 60 cm. long and 50 
cm. wide. 

Coccoloba nutans HBK. Nov. Gen. & Sp. 2: 175. 1817. 

A tree, the branchlets glabrous; leaves petiolate, subcoriaceous, 
the blades obovate, 20 cm. long and 13 cm. wide, short-acuminate, 
cordate at the base, glabrous, the veinlets prominulous and closely 
reticulate; racemes dense, nutant, the nodes 1-flowered, the flowers 
in bud almost sessile. Known to the writer only from the 
descriptions. 

Libertad: Trujillo, Bonpland (type). 

Coccoloba peruviana Lindau, Bot. Jahrb. 13: 213. 1890. 

A rather slender shrub or small tree, the branchlets somewhat 
puberulent; leaves short-petiolate, chartaceous, the blades oblong- 
obovate, 5.5-12 cm. long, acute or acuminate, narrowed to the acute 
base, tomentose or barbate beneath along the nerves, the venation 
closely reticulate but not conspicuous; racemes dense, shorter than 



FLORA OF PERU 461 

the leaves, the rachis glabrous or nearly so, the nodes 1-flowered, 
the pedicels 1.5 mm. long. Neg. 4960. 

Loreto: Caballo-cocha, in forest, Williams 2482. San Martin: 
Juan Guerra, 720 meters, in forest, Williams 6847, 6852. Without 
locality: Ruiz & Pavon 229 (type), 13-89. "Cunchu-caspi." 

Coccoloba Ruiziana Lindau, Bot. Jahrb. 13: 215. 1890. 

A shrub or tree, as much as 5 meters high, the branchlets obscurely 
puberulent or glabrate; leaves short-petiolate, chartaceous, the 
blades subovate to oblong or more often rounded-obovate, 3-7 cm. 
long, 2-5 cm. wide, broadly rounded to obtuse at the apex, shallowly 
cordate at the base, glabrous, the venation prominulous, laxly retic- 
ulate; racemes usually much longer than the leaves, dense, the 
rachis puberulent, the nodes 1-5-flowered, the pedicels 1.5-2 mm. 
long; fruits globose, scarcely 5 mm. long. Neg. 27791. 

Lambayeque: Supo, 1,450 meters, Townsend A1S5; a form with 
very small leaves. Piura: Between Frias and Chulucanas, 300-400 
meters, Weberbauer 6435. Negritos, Haught F15. Tumbez : Between 
Haciendas Casitas and Ricaplaya, 100 meters, dry river bed, Weber- 
bauer 7738. Without locality, Ruiz & Pavdn (type) . Also in Ecuador. 
"Liquanco" (Ruiz & Pavon), "analque." 

Coccoloba sphaerococca Lindau, Bot. Jahrb. 13: 185. 1890. 

Reported as a shrub, a tree, or a woody vine, the stout branchlets 
glabrous; leaves on short, thick petioles, coriaceous, the blades oblong- 
ovate to broadly elliptic or rounded-obovate, 7-20 cm. long, 4-11 
cm. wide, rounded or very obtuse at the apex, sometimes obtuse- 
acuminate, rounded or obtuse at the base, often very lustrous, 
glabrous, the venation closely reticulate and prominent; racemes 
dense, usually longer than the leaves, the rachis glabrous, the 
pedicels slightly longer than the ocreolae; fruit globose, 7 mm. in 
diameter. 

Loreto: Yurimaguas, in pasture, 200 meters, Williams 3805; 
in forest, Killip & Smith 29027, 27958. San Martin: Tarapoto, 
Spruce (type). 

Coccoloba Williamsii Standl. Field Mus. Bot. 11: 148. 1936. 

A shrub or tree, said to be sometimes scandent, the branchlets 
stout, glabrous; ocreae as much as 3 cm. long; leaves petiolate, 
coriaceous, the blades oval, oblong-oval, or oblong-elliptic, 12-20 
cm. long, 7-12 cm. wide, obtuse or rounded at the apex and some- 
times short-acuminate, more or less oblique at the rounded base, 



462 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sometimes shallowly emarginate, puberulent beneath along the 
costa and nerves, elsewhere glabrous; racemes 20-25 cm. long, 
the rachis densely puberulent or pilose, the nodes 1-3-flowered, the 
pedicels in anthesis shorter than the ocreolae; fruit purple, ovoid- 
globose, 5-6 mm. in diameter. 

Loreto: Lower Rio Huallaga, 200 meters, Williams 4803 (type). 
Yurimaguas, in forest, Williams 4597; Killip & Smith 28304- Lower 
Rio Nanay, Williams 673. Caballo-cocha, in forest, Williams 2423. 
Florida, 180 meters, in forest, Klug 2240. "Palo meta-caspi," 
"tangarana mashau," "eseri-ey" (the last a Huitoto name). 

6. SYMMERIA Benth. 

Shrubs or small trees; leaves alternate, coriaceous, the petiole 
subvaginate, no true ocreae present; flowers small, dioecious, panic- 
ulate, the staminate small, subsessile and glomerate, the pistillate 
larger, pedicellate; staminate calyx 6-parted, spreading, the seg- 
ments orbicular, the 3 outer ones smaller; stamens numerous, the 
filaments very short; pistillate calyx 6-parted, the outer segments 
small, oblong, not accrescent, the inner ones cordate-ovate, erect- 
connivent, accrescent and closely investing the 3-angulate achene. 
The genus consists of a single species. 

Symmeria paniculata Benth. in Hook. Lond. Journ. Bot. 4: 
630. 1845. 

A shrub or small tree, sometimes 8 meters high, the young 
branches and inflorescence somewhat ferruginous-tomentulose, other- 
wise glabrous or nearly so; leaves short-petiolate, coriaceous, the 
blades oval to oblong, mostly 10-20 cm. long and 2.5-10 cm. wide, 
obtuse or acutish, rounded or cordate at the base; staminate panicles 
large, open, with slender branches, the pistillate panicles smaller; 
fruiting bracts 12-18 mm. long. Flowers described as yellowish, 
brown-yellow, and golden yellow. Illustrated, Mart. Fl. Bras. 5, 
pt. 1: pi. 23. 

Loreto: Rio Masana, Williams 8, 101, 8150, 8145. Mishuyacu, 
100 meters, in forest, Klug 727, 457, 1181. Iquitos, 120 meters, 
Williams 7993. Amazonian Brazil, Guianas, and Colombia. "Tan- 
garana" (ant tree; presumably inhabited by ants, like Triplaris). 
Sometimes called "manguirana" in Amazonian Brazil. 

7. RUPRECHTIA C. A. Mey. 

Shrubs or trees; ocreae deciduous; flowers dioecious, fasciculate 
within small bracts, arranged in simple or paniculate racemes; 



FLORA OF PERU 463 

perianth 6-parted ; stamens 9; outer segments of the pistillate perianth 
erect, narrow, greatly enlarged in fruit and bright-colored, the 3 
inner ones small, linear, sometimes minute or obsolete; achene 
obtusely 3-angulate, pyramidal, 3- or 6-sulcate. 

Leaves acute or acuminate; inner segments of the pistillate perianth 
obsolete R. apeiala. 

Leaves obtuse or rounded at the apex; inner segments of the pistillate 
perianth developed R. Jamesonii. 

Ruprechtia apetala Wedd. Ann. Sci. Nat. III. 13: 268. 1849. 

A shrub; leaves short-petiolate, subcoriaceous, the blades ovate 
to oblong or obovate, 2.5-6.5 cm. long, acute or acuminate, obtuse or 
subacute at the base, entire or undulate, soft-pilose beneath or 
glabrate; staminate racemes slender, lax, longer than the leaves, the 
flowers short-pedicellate; fruiting perianth 2.5 cm. long, pubescent, 
the segments oblong-lanceolate, obtuse, 3-nerved. Neg. 7414. 

Cajamarca: Between Ja6n and Bellavista, 600 meters, Weber- 
bauer 6206. Without locality: Ruiz & Pavdn 33-98. Bolivia; a 
variety reported from Amazonian Brazil. 

The Weberbauer collection is staminate. It is not certain that the 
Ruiz and Pavon collection is really conspecific with Weddell's type. 

Ruprechtia Jamesonii Meisn. in DC. Prodr. 14: 179. 1857. 

A shrub 1.5-4 meters high; leaves short-petiolate, coriaceous, 
the blades oblong to oval or broadly obovate, mostly 3-7.5 cm. 
long and 1.5-5 cm. wide, obtuse or rounded at the apex, obtuse at 
the base, glabrate above, densely velutinous-pilose beneath with 
short hairs or finally glabrate, the venation very prominent beneath 
and closely reticulate; racemes very dense, usually longer than the 
leaves; fruiting perianth 2.5-3 cm. long, the outer segments glabrate, 
bright red or purplish red, linear-oblong or oblong-spatulate, obtuse 
or rounded at the apex, the 3 inner segments very short, subulate. 

Piura: Cerro Viento, 30 miles east of Talara, H aught F 29, 13 la. 
Cana Dulce, about 30 miles east of Cabo Blanco, Haught F29. 
Tumbez: North of Tumbez, Oleson; (Haught 131). Also in Ecuador. 

8. TRIPLARIS Loefl. 

Trees, the branches usually hollow and septate; ocreae deciduous; 
leaves large, short-petiolate, often with 3-6 longitudinal, distant 
lines on each side of the costa, these indicating folds of the blade in 
bud; flowers dioecious, racemose, the racemes paniculate or fas- 



464 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

ciculate, dense, the bracts small, ovate, acute, the ocreolae larger, 
long-acuminate, deeply slit on the anterior side; staminate perianth 
segments 6, subequal; stamens 9; segments of the pistillate perianth 
6, the 3 outer ones connate into a short or elongate tube, in fruit 
greatly enlarged and colored with red, the 3 inner segments free 
or partially adnate to the tube, small and narrow, little if at all 
exceeding the tube, usually shorter; achene trigonous, the angles 
usually acute. The genus is a difficult one because of the apparent 
inconstancy of most of the characters usually depended upon for 
separating species. The best characters for separation of species 
probably are those of the fruiting calyx, but since the majority of 
the Peruvian specimens are staminate, fruit characters alone are 
unsatisfactory for separating the forms. The genus is greatly in 
need of critical and careful revision. Several Peruvian specimens 
probably represent species additional to those recognized here, but 
material of them is insufficient for description. The branches of 
Triplaris trees practically always are inhabited by small ants that 
bite severely and painfully. 

Branchlets hirsute or hispid, at least about the nodes. 
Leaves rather densely and evenly hirsute on both surfaces. 

T. fulva. 
Leaves glabrous or nearly so except for a few very long and stiff, 

appressed hairs along the costa T. Poeppigiana. 

Branches glabrous or nearly so, the hairs, if any, appressed or 

strongly ascending. 

Bracts of the inflorescence sparsely short-hispid near the apex, 
otherwise glabrous. Fruiting calyx 5-6 cm. long, glabrous; 
leaves oblong, acute or attenuate at each end, conspicuously 

black-punctate beneath T. punctata. 

Bracts densely sericeous or pilose; fruiting calyx shorter, usually 

abundantly pilose. 

Inner perianth lobes in fruit longer than the tube; pubescence 

of the inflorescence very short, mostly appressed, of silky 

or very stiff hairs; leaves usually oblong. 

Outer lobes of the fruiting perianth little longer than the 

tube, with a large, rounded or emarginate lobe in each 

sinus; inner lobes spatulate-linear, not auriculate at 

the base T. peruviana. 

Outer lobes of the perianth much longer than the tube, with 
only minute and inconspicuous lobes in the sinuses; inner 
lobes linear, auriculate at the base T. auriculata. 



FLORA OF PERU 465 

Inner perianth lobes in fruit shorter than the tube; pubescence 
of the inflorescence of long, spreading, silky hairs; leaves 
mostly elliptic to oval, conspicuously broadest at or near 
the middle. 

Sinuses of the perianth tube naked T. Pavonii. 

Sinuses bearing small lobes T. Bonplandiana. 

Triplaris auriculata Meisn. in DC. Prodr. 14: 174. 1857. 

Leaves oblong or ovate-oblong, 15-20 cm. long, short-acuminate, 
rounded or acute at the base, sparsely appressed-pilose or almost 
wholly glabrous; inflorescence densely grayish-sericeous; fruiting 
perianth (immature) as much as 3.5 cm. long, densely sericeous on 
both surfaces, the outer lobes lanceolate, 4-6 mm. wide, obtuse, the 
inner lobes linear, adnate to the tube for more than half their length. 

The original locality is given as "In Nova-Granata? et Mexico 
(Pavon! in herb. Shuttlew.). T. Americana Pavon! mss." The 
specimens are unlike those of the only species known from Mexico, 
and are probably of South American origin. There is at hand a 
specimen from the Madrid Herbarium (Ruiz & Pavon 33-99), 
labeled as from Peru (or Chile). I suspect that the plant is the 
same as T. guayaquilensis Wedd. of Ecuador, and therefore to be 
excluded from the Peruvian flora, but it is included here because 
of the possibility of its Peruvian origin. 

Triplaris Bonplandiana Wedd. Ann. Sci. Nat. III. 13: 
262. 1849. 

Branchlets glabrous; leaves unknown; tube of the fruiting 
perianth subglobose, subappressed-pilose on both surfaces, the 
sinuses dentate, 18 mm. long, the outer lobes oblong, obtuse, the 
inner segments linear, adnate at the base to the tube, much shorter 
than the achene. 

The type was collected by Bonpland in "prov. Marannon" 
(Maranon), presumably somewhere in eastern Peru. Meisner (in 
DC. Prodr. 14: 174) associates with it "T. americana Bonpl. mss. 
n. 3599! in herb. Kunth." The type was without leaves, but the 
leaves of this specimen are described by Meisner as follows: Leaves 
oblong, 10-12.5 cm. long, 4-7.5 cm. wide, attenuate-acuminate at 
each end, glabrous above, puberulent beneath, the lateral lines 
obsolete. I have seen nothing to represent the species, and have 
been unable to associate any of the available material with it. 



466 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Triplaris fulva Huber, Bol. Mus. Goeldi 4: 560. 1906. 

Petioles only 5 mm. long, the blades oblong, acutely acuminate, 
unequal at the base and rounded or subcordate, membranaceous, 
hirsute on both sides with fulvous hairs; petioles and branchlets 
densely fulvous-hirsute. 

Loreto: Paca, Rio Ucayali, Huber 1565, type. 

The species, strangely enough, was based upon a sterile branch, 
apparently taken from a young plant. I have seen a portion of a 
leaf of the original material, which shows that the leaf pubescence 
is distinctive for the specimen, but it may not be characteristic 
for mature material. It is probable that the proper treatment of 
the name will remain obscure until the type locality is revisited. 
As a matter of fact, it is not altogether certain that the plant belongs 
to the genus Triplaris. 

Triplaris Pavonii Meisn. in DC. Prodr. 14: 172. 1857. T. 
boliviana Britton, Bull. Torrey Club 27: 130. 1900. 

A small or medium-sized tree, 5-16 .meters high, the stout 
branchlets glabrous or nearly so, the much branched, ample in- 
florescence densely sericeous-villous with soft, spreading, long, pale 
hairs; leaves large, short-petiolate, usually subcoriaceous, glabrous 
or nearly so in age, mostly elliptic and acute at each end or abruptly 
short-acuminate at the apex, the lateral nerves numerous and close 
together; fruiting perianth about 3.5 cm. long, densely pilose on 
both surfaces, the tube ovoid, the outer lobes obtuse, narrowly 
oblanceolate, obtuse or rounded at the apex, red or purple-red, 
the inner lobes lance-linear, shorter than the tube. 

Cuzco: Cuquipata, Diehl 2436 (determination very doubtful; 
perhaps undescribed, but material unsatisfactory; vernacular name 
Palo santo). Junin: La Merced, 600 meters, on river cliff, 5448. 
Loreto: Lower Rio Nanay, Williams 424- Yurimaguas, 200 meters, 
Williams 4176. Cachipuerto, 250 meters, in forest, Klug 3134 
(very doubtful, perhaps new; staminate, the large, thin leaves 
densely and softly pubescent beneath). Mishuyacu, 100 meters, 
Klug 947; staminate flowers cream-colored. Pongo de Manseriche, 
Killip & Smith 29124- Florida, 200 meters, riverside forest, Klug 
2162, 2087. San Martin: Zepelacio, 1,100 meters, Klug 3656 
(leaves rounded or subcordate at the base; perhaps a different 
species). Without locality: Ruiz & Pavon 34-1 (probably type 
material; fruiting perianths only), 33-94, 33-98. Bolivia and 
Amazonian Brazil. "Tangarana," "maicharo-ey" (Huitoto name). 



FLORA OF PERU 467 

Like other species of the genus, this is a handsome and ex- 
ceedingly showy tree when covered with the mature pistillate 
inflorescences, which are brightly colored in various shades of red, 
and remain upon the tree for a long time. 

Triplaris peruviana Fisch. & Mey. Mm. Acad. St. Pe'tersb. 
VI. 6: 149. 1845. 

A tree 3-16 meters high, the stout branchlets glabrous; leaves 
coriaceous, on short, stout petioles, the blades oblong, acuminate, 
acute to obtuse at the base, glabrous or nearly so, the lateral nerves 
rather few and distant; fruiting perianth about 3.5 cm. long, sparsely 
appressed-pilose or almost glabrous, the tube tubular-campanulate, 
the outer segments little longer than the tube, narrowly oblong, 
obtuse or rounded at the apex, each sinus with a large, ovate lobe. 
Neg. 8490. 

Loreto: Timbuchi, Williams 1008. Florida, 200 meters, in forest, 
Klug 2057 (uncertain, perhaps somewhat abnormal; if not referable 
here, the species must be undescribed). Rio Huallaga, Stuebel 1. 
Mishuyacu, 100 meters, in forest, Klug 1451; staminate flowers 
white. San Martin: Chazuta, in forest, 260 meters, Klug 4153. 
Without locality: Mathews 1620 (type). Amazonian Brazil. "Tan- 
garana," "tangarana blanca." 

Triplaris Poeppigiana Wedd. Ann. Sci. Nat. III. 13: 265. 
1849. Blochmannia peruviana Poepp. ex Meisn. in DC. Prodr. 14: 
173. 1857, in syn. T. hispida Britton, Mem. Torrey Club 6: 111. 
1896. T. longifolia Huber, Bol. Mus. Goeldi 4: 559. 1900. 

A shrub or tree, sometimes 20 meters high, the stout branchlets 
sparsely hispid with very long, spreading, stiff, brownish hairs; 
leaves large, oblong, acuminate or cuspidate-acuminate, usually 
rounded or obtuse at the base, sometimes cordate, the lateral nerves 
numerous and rather close together, sparsely hispid or appressed : 
hispid beneath along the costa, otherwise glabrous or nearly so, or 
sometimes sparsely hispid over almost the whole lower surface; 
branches of the broad inflorescence more or less hispid with long, 
stiff, fulvous hairs; fruiting perianth about 5 cm. long, the tube 
sparsely or densely hirsute, the outer segments linear-oblong, twice 
as long as the tube, obtuse, the inner lobes lanceolate, short, adnate 
at the base to the tube. Negs. 27773, 4972. 

Junin: La Merced, 600 meters, in forest, 5401. Puerto Yessup, 
400 meters, in forest, Kittip & Smith 26392, 26342. Rio Pichis, 
350 meters, Kittip & Smith 26700. Rio Paucartambo Valley, 700 



468 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

meters, Killip & Smith 25244- Loreto: Cerro de Canchahuaya, 
Huber 1458, type material of T. longifolia. San Martin: Tocache, 
Poeppig 1957 (type). Without locality: Ruiz & Pavdn 33-97. Also 
in Bolivia and doubtless in Amazonian Brazil. 

T. longifolia was based upon sterile material taken, apparently, 
from a young shoot. I have seen a leaf of the original material, 
and this shows along the costa a few of the long, stiff hairs that are 
so characteristic for this species. 

Triplaris punctata Standl., sp. nov. 

Arbor 10-18-metralis, ramulis crassis glabris brunneo-puncticu- 
latis; folia breviter petiolata crasse membranacea in sicco olivaceo- 
viridia, petiolo crasso 1-1.5 cm. longo; lamina anguste lanceolato- 
oblonga 18-30 cm. longa 4.5-8 cm. lata anguste attenuato-acumi- 
nata, basi plus minusve inaequali acuta, glabra, subtus ubique dense 
puncticulis nigrescentibus notata, costa gracili elevata, nervis 
lateralibus numerosis approximatis angulo acuto adscendentibus, 
striis obsoletis; racemi masculi fasciculati densissimi 7-17 cm. 
longi, rhachi glabra vel obscure puberula, bracteis latis tantum 
prope apicem hispidulis, aliter glabris, sepalis strigosis, staminibus 
bene exsertis; racemi fructiferi breves densi, rhachi glabra; peri- 
gonium fructiferum in sicco pallidum omnino glabrum circa 5.5 cm. 
longum, tubo late campanulato vix ultra 1 cm. longo, sinu lobulo 
ovato brevi acuminate onusto, alis oblongis tubo triple longioribus 
obtusis 10-12 mm. latis arete reticulato-venosis, lobis interioribus 
lineari-subulatis tubo brevioribus media longi tudine tubo adnatis; 
achaenium 13 mm. longum ovoideum profunde 3-sulcatum, angulis 
valde compressis alariformibus. 

Loreto: Balsapuerto, 220 meters, in forest, Klug 3009; staminate 
flowers cream-colored. Brazil: Near mouth of Rio Macauhan, a 
tributary of Rio Yaco, Territory of Acre, on terra firma, Krukoff 
5277 (Herb. Field Mus., type; in fruit), 5333 (staminate). 

The species is well marked because of its long, narrow leaves, 
glabrate inflorescence, and large, glabrous fruiting perianths. The 
conspicuous dark dots of the lower leaf surface are not confined to 
this species, but they are more noticeable than in any other I have 
observed. The leaves are strikingly like those of T. longifolia Huber, 
but in that there are no black dots, and this plant does not have the 
distinctive, long hairs found on the lower leaf surface in T. longifolia, 
i.e., T. Poeppigiana. 



FLORA OF PERU 469 

59. CHENOPODIACEAE. Goosefoot Family 
By Paul C. Standley 

Herbs or shrubs, the pubescence often of minute, inflated hairs, 
the stems sometimes jointed; leaves opposite or alternate, without 
stipules, never regularly serrate; flowers perfect or unisexual, usually 
small and greenish, solitary or glomerate, often arranged in spikes 
or cymes; perianth simple, herbaceous or membranaceous, normally 
of 2-5 segments, these united below, often enlarged in fruit; stamens 
usually 2-5, the filaments slender, the anthers dorsifixed, 4-celled; 
ovary superior, 1-celled; stigma capitate, or the styles 2-3, or the 
stigmas 2-5 and sessile; ovule solitary; fruit a utricle, containing a 
single small seed. 

Besides the species and genera listed below, the beet (remolacha; 
Beta vulgaris L.) and spinach (espinaca; Spinacia oleracea L.) some- 
times are cultivated as edible vegetables in Peru. 

Stems jointed, fleshy; leaves reduced to minute scales; flowers 
sunken in groups of 3-5 in the joints of the stem . . 1. Salicornia. 

Stems not jointed; leaves usually well developed; flowers not sunken 
in the stem. 

Embryo spirally coiled; leaves terete or semiterete, very fleshy; 
flowers all axillary 2. Suaeda. 

Embryo not spirally coiled; leaves flat; flowers often spicate or 
cymose. 

Flowers perfect, not bracteolate 3. Chenopodium. 

Flowers unisexual, the pistillate ones subtended by 2 bracteoles, 
these enlarging with age and enclosing the fruit. 

4. Atriplex. 
1. SALICORNIA L. 

Annuals or perennials, sometimes suffrutescent, fleshy, glabrous, 
with jointed branches, the joints dilated at the apex into a short 
sheath; flowers perfect or polygamous, immersed in groups of 3-7 
on opposite sides of the joints, the flowering joints forming cylindric, 
terminal spikes; perianth obpyramidal, fleshy, 3-4-dentate; stamens 
1-2; fruit a minute utricle, included in the perianth; seed erect, 
compressed, minutely hairy. 

Salicornia fruticosa L. Sp. PI. ed. 2. 5. 1762. S. pemviana 
HBK. Nov. Gen. & Sp. 2: 193. 1817. S. Gaudichaudiana Moq. 
Chenop. Enum. 115. 1840. S. biloba Kunze ex Fenzl in Mart. Fl. 



470 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Bras. 5, pt. 1: 158. 1864, in syn. S. Neei Lag. Mem. PI. Barrill. 51. 
1817. S. equisetifolia Willd. ex Moq. loc. cit. as syn. 

An erect or prostrate perennial, suffrutescent at the base, much 
branched, the secondary branches ascending or erect, usually 
branched, the joints mostly 1-2 cm. long; sheaths rounded or with 
acutish lobes; flower spikes about 2 cm. long and 3 mm. thick, the 
flowers in groups of 3; seed yellowish brown, covered with short, 
conic hairs. Neg. 7354 (S.Gaudichaudiana). 

Arequipa: Mollendo, R. S. Williams 2543. Lima: Lurin, in salt 
marsh, 5931. Callao, Wilkes Expl. Exped. Type of S. peruviana 
collected by Humboldt and Bonpland near Huarmei (Dept. Lima). 
Widely distributed on seashores of both hemispheres. 

2. SUAEDA Forsk. 

Annuals or perennials, erect or prostrate, glabrous or pubescent, 
herbaceous or suffrutescent; leaves alternate, terete or semiterete, 
rarely flat, entire, fleshy; flowers minute, chiefly perfect, solitary or 
glomerate in the leaf axils; perianth fleshy, 5-lobed; stamens 5; fruit 
a compressed or depressed utricle, enclosed in the perianth; seed 
horizontal or erect, smooth or roughened. 

Suaeda foliosa Moq. in DC. Prodr. 13, pt. 2: 156. 1849. 

Apparently perennial, glabrous or nearly so, much branched, 
the stout branches roughened by the persistent leaf bases of fallen 
leaves; leaves mostly 5-8 mm. long, glaucous, very thick and fleshy, 
obtuse or acutish, semiterete, glabrous; flowers minute, green, 
solitary or in clusters of 3. Neg. 7356. 

Lima: Lima and Callao, Wilkes Expl. Exped. Lima, Rose 18576. 
Type said to have been collected somewhere in Peru by Gaudichaud. 
Also in Chile. 

Suaeda foliosa var. tenuifolia (Phil.) Standl., comb. nov. S. 
tenuifolia Phil. Anal. Univ. Chile 91: 432. 1895. S. divaricata Moq. 
var. tenuifolia Reiche, Fl. Chil. 6: 175. 1911. 

Leaves longer and proportionately narrower, as much as 18 mm. 
long; flowers in clusters of 3-5. 

Arequipa: Tingo, 2,200 meters, open, rocky places, Pennell 
13104- Lima: Chorrillos, near Lima, 150 meters, loose, stony, 
seaside slopes, 5867. Also in Chile. 

3. CHENOPODIUM L. 

Annual or perennial herbs, often strong-scented, usually either 
glandular or covered with a mealy pubescence of small, white, 



FLORA OF PERU 471 

inflated hairs; leaves alternate; flowers perfect or rarely unisexual, 
chiefly 5-parted; perianth calyx-like, herbaceous, with usually 5 
lobes, these often keeled dorsally, persistent; stamens 5 or fewer; 
fruit a utricle, erect or depressed, free from or adherent to the seed; 
seed horizontal or vertical, the embryo completely or incompletely 
annular. 

Seed vertical. Plants white- villous when young; leaves deeply 

pinnatifid C. multifidum. 

Seed horizontal. 

Embryo not completely encircling the endosperm; leaves and 
inflorescence glandular. 

Pericarp gland-dotted; flowers in spikes C. ambrosioides. 

Pericarp not gland-dotted; inflorescence dichotomous, some of 

the flowers pediceled C. incisum. 

Embryo completely encircling the endosperm; plants without 
glands. 

Leaves lustrous on the upper surface, rhombic. Inflorescences 
chiefly axillary C. murale. 

Leaves dull on the upper surface. 

Seeds white or nearly so C. Quinoa. 

Seeds black or blackish. 

Inflorescence naked or almost so, usually with many slender, 

spreading branches. Plants tall and much branched, 

slender; seed about 1 mm. broad.... C. petiolare. 

Inflorescence usually very leafy, dense, the branches stout, 

chiefly erect. 

Plants low and spreading; leaves small, 1-2 cm. long and 
wide, fleshy; seed about 1.2 mm. broad. 

C. pallidicaule. 

Plants tall, erect, stout; leaves much larger, thin; seed 
about 1.8 mm. broad C. hircinum. 

Chenopodium ambrosioides L. Sp. PI. 219. 1753. C. anthel- 
minticum L. Sp. PI. 220. 1753. 

Plants evil-scented, erect or ascending, 1 meter high or usually less, 
annual or perennial, glabrous or puberulent, usually glandular- 
villous about the inflorescence; lower leaves petiolate, the blades 
oblong to ovate or lanceolate, 3-12 cm. long, coarsely sinuate- 
dentate or sinuate-pinnatifid, conspicuously gland-dotted; flowers 



472 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

glomerate, forming short or elongate, leafy or naked spikes; upper- 
most leaves narrow, frequently spatulate or linear, often entire; 
seed 0.6-0.8 mm. broad, black. 

Cuzco: San Sebastian, 3,300 meters, Herrera 557 (reported by 
Aellen). Cuzco, Herrera 142 (ex Aellen). Huanuco: Mito, 2,700 
meters, a weed in corral, 1703. Lima: Lima, 150 meters, roadsides, 
70. Loreto: Iquitos, Williams 3556, 7914- Lower Rio Nanay, 
Williams 339. Yurimaguas, 200 meters, Williams 4048. Mishu- 
yacu, in clearing, Klug 117. Fortaleza, Yurimaguas, in garden, 
Williams 4499. Aripari, 150 meters, Killip & Smith 29178. Santa 
Rosa, 135 meters, in clearing, Killip & Smith 28868. San Martin : 
Tarapoto, Williams 6247, 6189. Generally distributed in tropical 
America, and naturalized in the United States and many regions of 
the Old World. 

"Paico" (Amazon Valley), "camatai," "cashiva" (Mito). The 
plant has an exceedingly offensive odor, which is retained in dried 
specimens. Although widely dispersed in tropical America, this 
species seldom is abundant, and is confined chiefly to the vicinity of 
dooryards and barnyards. In Peru it sometimes is cultivated as a 
medicinal plant, being employed locally as a remedy for intestinal 
parasites and for affections of the stomach. The seeds are the worm- 
seed or Mexican wormseed of the pharmacopoeia, being considered 
a highly efficient vermifuge. 

Chenopodium ambrosioides L. subsp. chilense (Schrad.) 
Aellen, Repert. Sp. Nov. 26: 36. 1929. C. chilense Schrad. Ind. Sem. 
Hort. Goett. 1832: 2. 1832. C. vagans Standl. N. Amer. Fl. 21: 
26. 1916. 

Similar to the species, frequently suffrutescent; stems often more 
or less white- villous; leaves chiefly smaller than in the typical form 
and more deeply pinnatifid, the uppermost leaves commonly coarsely 
dentate or deeply pinnatifid. 

Ancash: Recuai, 2,700 meters, rock cliffs, forming clumps, 
2518. Puno: Azangaro, 4,000 meters, Weberbauer (ex Aellen). 
Puno, 4,000 meters, Soukup 222. Ranging to Chile, Argentina, and 
Brazil, and adventive in California. 

Macbride reports the vernacular name as "amush," and states 
that the plant is employed as a remedy for insect bites. 

Chenopodium hircinum Schrad. Ind. Sem. Hort. Goett. 1833: 
2. 1833. C. hircinum subsp. eu-hircinum Aellen, var. andinum 
Aellen, Repert. Sp. Nov. 26: 122. 1929. 



FLORA OF PERU 473 

Plants tall and coarse, erect, the pale stems more or less striate; 
leaves long-petiolate, broadly rhombic, coarsely sinuate-dentate or 
often somewhat 3-lobate, green but sparsely and minutely mealy; 
inflorescence dense, narrow, leafy, loosely mealy. 

Junin: Tarma, 2,100 meters, along trail, 1024 (type of var. 
andinum). Huancayo, 3,400 meters, Weberbauer 6597. The species 
in its various subspecies, varieties, and forms has been reported from 
most of the South American countries. In general appearance the 
plant is similar to the common C. album of the United States and 
Europe. Called "sacha-quinua" in Ecuador. 

Chenopodium incisum Poir. in Lam. Encycl. Suppl. 1: 392. 
1811. C. incisum var. Bangii Murr, f. rotundifolium Aellen, Repert. 
Sp. Nov. 26:40. 1929. 

An erect annual, 60 cm. high or less, strong-scented, often much 
branched, sparsely pubescent or glabrate, glandular; leaves petiolate, 
the blades deltoid to oblong or rounded, sinuate-pinnatifid or laciniate- 
pinnatifid, bright green, bearing many yellow glands on the lower 
surface; inflorescence of numerous loosely few-flowered, axillary 
cymes; flowers sessile in the forks of the cymes and solitary at the 
ends of the slender lateral branches, the pedicellate flowers chiefly 
abortive, their pedicels becoming spinose; seed 0.5-0.8 mm. broad, 
dark brown. "Arcapaico" (Cook). 

Arequipa: Tingo, 2,200 meters, open, rocky slope, Pennell 13128. 
Arequipa, 2,500 meters, open, gravelly soil, Pennell 13166. Cuzco: 
Ollantaitambo, 3,000 meters, Cook & Gilbert 807. Temple of Vira- 
cocha, near Tinta, 3,500 meters, among volcanic rocks, Cook & Gil- 
bert 208 (type of f. rotundifolium). In its various forms the species 
ranges to Argentina and northward to southwestern United States. 

One of the Peruvian collections has been referred to C. graveolens 
Lag. & Rodr. That is an older name than C. incisum, and may 
pertain to the same plant, but its identity is doubtful. 

Chenopodium multifidum L. Sp. PI. 220. 1753. Roubieva 
multifida Moq. Ann. Sci. Nat. II. 1: 293. 1834. 

A prostrate perennial, strong-scented, much branched, the 
branches usually short and more or less villous; leaves oblong, 
1-4.5 cm. long, sparsely villous and glandular, deeply pinnatifid or 
the lowest merely coarsely lobed ; flowers green, solitary or clustered 
n the leaf axils, sessile; seed 1 mm. in diameter, black and shining. 



474 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Reported by Aellen (Repert. Sp. Nov. 26: 41. 1929) from Peru, 
Dombey 217. The record is doubtful. The species occurs in Chile, 
Argentina, and Brazil, and is adventive occasionally in other parts 
of the world. 

Chenopodium murale L. Sp. PI. 219. 1753. 

An erect or ascending annual, rarely more than 50 cm. high, 
bright green but somewhat mealy, usually much branched from the 
base; leaves slender-petiolate, the blades chiefly ovate-rhombic, 
3-8 cm. long, irregularly sinuate-dentate; flowers mealy, sessile, the 
small glomerules arranged in lax or dense, chiefly leafless cymes or 
panicles; seed 1.2-1.5 mm. broad, finely puncticulate. 

Arequipa: Tingo, 2,200 meters, open, rocky slopes, Pennell 
13145- Arequipa, Rose 19007. Junin: Tarma, 2,100 meters, mud 
wall of old house, 1019; moist ditch, Killip & Smith 21880. Lima: 
Chorrillos, near Lima, 150 meters, along trail on seaside hill, 5876. 
Lima, 150 meters, roadside, 67. A native of the Old World, but 
naturalized as a weed in many parts of America. " Yerba de gallinazo' ' 
(Cook). 

Chenopodium pallidicaule Aellen, Repert. Sp. Nov. 26: 
126. 1929. 

Plants low and much branched, 50 cm. high or less, sparsely and 
coarsely mealy, pale; leaves long-petiolate, the blades 2 cm. long or 
less, thick and fleshy, shallowly trilobate, attenuate at the base to 
the petiole, the uppermost leaves sagittate or entire; flower spikes 
shorter than the leaves, axillary; seed minutely and irregularly 
punctate. 

Puno: Santa Rosa, south side of La Raya Pass, 400 meters, Cook 
& Gilbert 161 (type). Between Santa Rosa and Araranca, Cook & 
Gilbert 170. Junin: Oroya, Kalenborn 129 (determination uncer- 
tain). Also in Bolivia. 

Aellen names also a forma purpureum (based on part of the type 
number), in which the plant is more or less tinged with red. 

Chenopodium petiolare HBK. Nov. Gen. & Sp. 2: 191. 1817. 
C. paniculatum Hook. Bot. Misc. 2: 237. 1831. C. paniculatum var. 
reniforme Murr, Bull. Herb. Boiss. II. 4: 994. 1904. C. paniculatum 
var. incanum Murr, Allg. Bot. Zeitschr. 12: 54. 1906. C. petiolare 
f. incanum Aellen, Repert. Sp. Nov. 26: 150. 1929. C. petiolare f. 
Hookeri Aellen, loc. cit. C. petiolare f. hastatum (Phil.) Aellen, loc. 
cit. C. petiolare f. scutatum Aellen, Repert. Sp. Nov. 26: 151. 1929. 



FLORA OF PERU 475 

C. petiolare f . trilobum Aellen, loc. cit. C. petiolare f . reniforme Aellen, 
loc. cit. 

Plants erect or spreading, slender, much branched, pale, the 
branches rather densely mealy; leaves slender-petiolate, the blades 
thin, densely farinose or sometimes green and glabrate, very variable 
in outline, small, usually more or less deltoid and distinctly hastate- 
lobate at the base, otherwise entire or remotely sinuate-dentate; 
inflorescence large and open, much branched, the spikes slender and 
much interrupted, naked or with few reduced leaves. 

Arequipa: Arequipa, Rose 19006; gravel ridges along stream bed, 
2,800 meters, Pennell 13252. Posco, 550 meters, Cook & Gilbert 41a. 
Mollendo, on benches and sheltered places, Johnston 3561; sandy 
desert hills, Hitchcock 22363. Ayacucho: Coracora, 2,900 meters, 
Weberbauer 5797. Cuzco: Ollantaitambo, 3,000 meters, a weed in 
cultivated ground, Cook & Gilbert 432. Junin: Tarma, 3,600 meters, 
along trail, 1072. Lima: Matucana, 2,400 meters, gravelly river 
canyon slope, 242. Chorrillos, near Lima, 150 meters, in rocks on 
upper slopes of seaside hills, 5873. San Geronimo, Lima, 150 meters, 
rocky slope, 5901. Rio Chillon, near Viscas, 1,900 meters, bare, 
open, loam slopes, Pennell 14453. Lima, Rose 18590. Moquehua: 
Torata, 3,200 meters, Weberbauer 7465. The species ranges from 
Ecuador to Bolivia and Chile. "Lipcha" (Cook). 

The forms named by Aellen are based upon variations in leaf 
form, which are of slight importance. 

Chenopodium Quinoa Willd. Sp. PL 1: 1301. 1797. C. pur- 
purascens Jacq. var. punctulatum Moq. in DC. Prodr. 13, pt. 2: 67. 
1849. C. Nuttalliae Safford, Journ. Wash. Acad. Sci. 8: 523. 1918. 
C. Canihua Cook, in herb. C. Quinoa f. purpureum Aellen, Repert. 
Sp. Nov. 26: 124. 1929. 

A coarse, erect annual, frequently 1 meter high or more, sparsely 
and finely mealy, green or somewhat purplish (f . purpureum) ; leaves 
slender-petiolate, the blades large, broadly rhombic, sinuate-dentate, 
sometimes obscurely lobate at the base; inflorescences erect, leafy, 
very dense and compact; seeds whitish, about 1.5 mm. broad. 

Cuzco: Tinta, Cook & Gilbert 232. Santa Rosa, 4,000 meters, 
Cook & Gilbert 231 . Cuzco, 3,400 meters, in market, Cook & Gilbert 
84- Junin: Tarma, 3,100 meters, Killip & Smith 21907. Puno: 
Pomata, 3,820 meters, Weberbauer 7964- Without locality: Ruiz & 
Pavon. Also in Ecuador, Bolivia, and Argentina, and in Mexico. 
"Quinoa," "quinua," "canihua." 



476 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

The plant has long been an important food staple in the Andes, 
where it is planted extensively for its nutritious seeds. Most com- 
monly the fresh flower spikes are cooked and the whole is eaten, or 
else the seeds are stripped from the spikes with the teeth. 

4. ATRIPLEX L. 

Shrubs or herbs, more or less furfuraceous, farinose, or canes- 
cent with inflated hairs; leaves alternate or opposite; flowers monoe- 
cious or dioecious, usually glomerate, the glomerules axillary or dis- 
posed in solitary or panicled spikes; staminate perianth 3-5-parted; 
stamens 3-5; pistillate flowers bibracteolate, the bractlets accres- 
cent, free or united, enclosing the fruit, the perianth usually none; 
seed erect or inverted, rarely horizontal. The genus is most 
abundantly represented in arid regions, and there are numerous 
species in Chile and Argentina. 

Plants prostrate, herbaceous; leaves small, mostly 4-8 mm. long. 

A. serpyllifolia. 
Plants erect or ascending, often shrubby; leaves large, chiefly 1.5-6 

cm. long. 
Bracts foliaceous, dentate, usually 6-9 mm. long or longer; leaves 

mostly dentate or crispate A. rotundifolia. 

Bracts not foliaceous, entire or nearly so, usually about 5 mm. long. 

A. peruviana. 

Atriplex peruviana Moq. in DC. Prodr. 13, pt. 2: 102. 1849. 
Chenopodium cinereum Moq. Chenop. Enum. 27. 1840. 

A shrub 1-2 meters high, much branched, grayish or whitish and 
densely farinose or furfuraceous throughout, the branches slender, 
pale; leaves slender-petiolate, the blades rhombic or broadly ovate to 
rounded, mostly 1.5-3 cm. long, entire or nearly so but often cris- 
pate; flowers monoecious, the staminate glomerules forming large 
and rather open, terminal panicles, the pistillate solitary or clustered 
in the leaf axils; fruiting bracts rhombic or broadly triangular, obtuse 
or acutish, densely furfuraceous. Neg. 7355. 

Arequipa: Rio de Lomas and Rio Yanca, 1,900 meters, Weber- 
bauer 5757. Rio de Lomas, 700 meters, Weberbauer 5747. Rio de 
Lomas, Acari, 900 meters, Weberbauer 5735. Tiabaya, 2,100 meters, 
rocky river bank, Pennell 13088. Tacna: Candarave, 2,900 meters, 
Weberbauer 7366, 7367. The type is Gaudichaud 68 from some 
unknown locality of Peru, and Moquin reports also a collection by 
Mathews. The species is reported by the same author from Chile. 



FLORA OF PERU 477 

It is uncertain whether the specimens cited really are referable 
to A. peruviana, since they do not agree quite satisfactorily with 
Moquin's descriptions. It is questionable, also, whether the speci- 
mens cited are specifically distinct from A. rotundifolia. 

Atriplex rotundifolia (Moq.) Dombey ex Moq. Chenop. Enum. 
70. 1840, in syn. Obione rotundifolia Moq. loc. cit. A. Espostoi 
Speg. Physis 2: 241. /. 1916. 

Plants herbaceous or suffrutescent, coarse and stout, 50 cm. high 
or more, minutely farinose or more often green and glabrate, sparsely 
branched, the branches angulate; leaves slender-petiolate, the blades 
rather thin but sometimes thick and crispate, rounded-deltoid to 
broadly rhombic or ovate, commonly 4-6 cm. long, very obtuse or 
rounded at the apex, often coarsely dentate; flowers monoecious, the 
staminate heads in small but lax panicles, often blackish when dried, 
the pistillate solitary or clustered in the upper leaf axils; fruiting 
bracts variable but most of them large and greenish, finely farinose, 
deltoid or broadly rhombic, commonly tuberculate near the base. 

Lima: San Lorenzo Island, 400 meters, Weberbauer 5923; Wilkes 
Expl. Exped. Near Lima, Rose 18588. Without locality: Weber- 
bauer 5710. Type collected in Peru by Dombey. Moquin reports 
also a specimen collected by Mathews. Type of A. Espostoi collected 
on the coast of Peru by Nicolas E. Esposto. 

If all the collections cited above really belong to this species 
some of them are incomplete it is a remarkably variable one. 

Atriplex Herzogii Standl. Field Mus. Bot. 11: 125. 1931. A. 
serpyllifolia Herzog, Med. Rijks Herb. 27: 11. 1915, not Bunge, 1877. 

Perennial from a woody root, much branched, the branches 30 
cm. long or less, pale, prostrate, forming dense mats; leaves subsessile, 
oblong to obovate, minute, whitish, obtuse or rounded at the apex 
and usually mucronate, entire, narrowed to the base; flowers monoe- 
cious, the staminate in few terminal heads scarcely exceeding the 
leaves, the pistillate axillary; fruiting bracts broadly rhombic or 
rounded, about 2 mm. long, finely dentate and usually tuberculate 
dorsally. 

Junin: Tarma, 2,100 meters, dry wash on hillside, 1000. 
Arequipa: Tingo, 2,200 meters, Pennell 13106. Also in Bolivia. 

This probably is the species reported by Moquin (in DC. Prodr. 
13, pt. 2: 110. 1849) from Peru, collected by Pavon, provided that the 
specimen so reported was Peruvian. Another Pavon collection 
reported by Moquin (p. 113) likewise is probably A. Herzogii, if 



478 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

really Peruvian. The Pennell collection was determined by Ulbrich 
as A. prostrata Phil., a Chilean species. The specimen is unsatis- 
factory, but may well be referable to A. Herzogii. 

60. AMARANTHACEAE. Amaranth Family 
By Paul C. Standley 

Reference: Schinz, Nat. Pflanzenfam. ed. 2. 16c: 7-85. 1934. 

Herbs or shrubs; leaves simple, opposite or alternate, without 
stipules; flowers small, usually inconspicuous and green, sometimes 
white or colored, perfect, monoecious, polygamous, or dioecious, 
usually spicate or capitate, the bracts and bractlets scarious; petals 
none; sepals 0-5, distinct or slightly united at the base, equal or the 
inner ones smaller; stamens 1-5, opposite the sepals, the filaments 
free or connate or united with the intervening pseudostaminodia; 
anthers 1-2-celled; ovary 1-celled, containing 1 or few ovules, the 
style short or elongate, the stigmas 1-5; fruit usually a utricle, 
sometimes baccate or capsular, circumscissile, irregularly dehiscent, 
or indehiscent. 

There are few families in which characters for separating genera 
and species are so nearly confined to the flowers, foliage characters 
being usually of slight or no importance. The genera and in some 
cases the species can be determined only after careful and often 
difficult dissection of the flowers. 

Ovules 2 or more. Anthers 4-celled ; plants herbaceous or suffrutes- 

cent, never scandent; leaves alternate. 
Fruit somewhat baccate; perianth segments more or less spreading 

in age 1. Pleuropetalum. 

Fruit dry; perianth segments erect 2. Celosia. 

Ovule solitary. 

Leaves alternate. Anthers 4-celled. 

Seeds arillate; plants often scandent and frequently more or 

less woody; filaments connate at the base. . .3. Chamissoa. 

Seeds not arillate; plants never scandent, herbaceous; filaments 

distinct 4. Amaranthus. 

Leaves opposite. 
Anthers 4-celled; segments of the sterile flowers with uncinate, 

spine-like tips 5. Cyathula. 

Anthers 2-celled; segments of the flowers never with uncinate 
tips. 



FLORA OF PERU 479 

Perianth segments united into a tube, this indurate in fruit 
and bearing spine-like appendages or dentate crests. 

6. Froelichia. 
Perianth segments usually distinct, never with spines or crests 

in age. 

Stamens perigynous; flowers glomerate in the leaf axils, 
sessile. Plants perennial, the prostrate, much branched 
stems forming dense mats; plants densely lanate. 

7. Guilleminea. 
Stamens hypogynous; flowers in heads or spikes, these sessile 

or more often pedunculate. 

Stigma capitate or sometimes shallowly bilobate. 

Stamen tube 5-lobate, without pseudostaminodia, the 

lobes 3-lobate, dentate, or laciniate. . .8. Pfaffia. 

Stamen tube 4-10-lobate, the antheriferous lobes 

alternating with pseudostaminodia, or the tube 

5-lobate but with entire lobes. . .9. Alternanthera. 

Stigma 2-3-lobate, the lobes subulate or filiform. 

Stamen tube with broad lobes, these usually trilobate, 

dentate, or laciniate; flowers in broad, dense heads 

or spikes, these often subtended at the base by 

leaves; flowers perfect 10. Gomphrena. 

Stamen tube with entire, subulate lobes; flowers in 
slender spikes, these naked at the base; flowers 
often dioecious 11. Iresine. 

1. PLEUROPETALUM Hook. f. 

Glabrous shrubs; leaves alternate, petiolate; flowers perfect, 
pedicellate, racemose or paniculate, greenish; perianth segments sub- 
equal, striate-nerved, obtuse, spreading in fruit; stamens 5-8, the 
subulate filaments connate at the base into a short cup; stigmas 2-4, 
short, subulate; fruit baccate, rupturing irregularly, containing few 
or numerous seeds. 

Pleuropetalum Sprucei (Hook, f.) Standl. N. Amer. Fl. 21: 
96. 1917. Melanocarpum Sprucei Hook. f. in B. & H. Gen. PI. 3: 
24. 1880. 

A slender shrub with elongate, sometimes subscandent branches; 
leaves blackish when dried, short-petiolate, the blades oblong- 
lanceolate to ovate-elliptic, 10-18 cm. long, long-acuminate, acumi- 



480 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

nate to subobtuse at the base; flowers paniculate, the panicles 3-6 
cm. long, dense; sepals greenish, or becoming orange-red in fruit, 
2.5-4 mm. long; fruit red or finally black, globose, 5 mm. in diameter. 
Illustrated, Curtis's Bot. Mag. pi. 6674. 

Ayacucho: Near Kimpitiriki, 400 meters, dense forest, Killip 
& Smith 23002. Ranging to southern Mexico. 

2. CELOSIA L. 

Annual or perennial herbs or shrubs; leaves alternate, usually 
petiolate; flowers perfect, in dense, terminal or axillary spikes, 
or fascicled along the branches; perianth 5-parted, the segments 
scarious; stamens 5, the filaments connate at the base into a short 
cup; style elongate, short, or none; stigmas 2-3, subulate or capitate; 
ovules 2 or more; utricle included or exserted, usually circumscissile, 
containing 2 to many seeds. 

Sepals 3-4 mm. long, brownish when dried; leaves narrowly oblong- 
lanceolate C. Persicaria. 

Sepals 5-9 mm. long; leaves mostly ovate. 

Flower spikes about 7 mm. thick, brownish when dried; sepals 
5-6 mm. long C. virgata. 

Flower spikes 1.5-2 cm. thick, white, red, purple, or yellow; sepals 
6-9 mm. long C. argentea. 

Celosia argentea L. Sp. PI. 205. 1753. C. cristata L. loc. cit. 

A coarse, erect annual; leaves long-petiolate, the blades linear to 
rounded-ovate, acute or acuminate, often tinged with purple; flowers 
forming dense, thick spikes terminating the branches, variable in 
color from white to red, purple, or yellow; sepals 6-9 mm. long; seeds 
3-8. 

Loreto: Caballo-cocha, Williams 2396. Near Iquitos, Williams 
1349, 3535. Pebas, Wittiams 1832. Puerto Arturo, in chacara, 
Williams 5133. San Martin : Rumizapa, Williams 6806. San Roque, 
Williams 7300. Tropics of both hemispheres. 

The form or variety cristata of this species, a form with fasciate 
inflorescence, is the common cockscomb ("cresta de gallo") of 
gardens, a favorite ornamental plant of tropical American gardens. 
It is probably a native of tropical America, but is not known in a 
wild state, although it sometimes persists or escapes from gardens. 
Cockscomb is cultivated for ornament in most tropical and temperate 
regions of the earth. 



FLORA OF PERU 481 

Celosia Persicaria Schinz, Bull. Herb. Boiss. II. 3: 4. 1903. 

Plants herbaceous, 50 cm. high; leaves glabrous, petiolate, the 
blades narrowly oblong-lanceolate, about 13 cm. long and 2.5 cm. 
wide, acute or acuminate, long-attenuate to the base; flower spikes 
simple or branched, the flowers densely glomerate, the glomerules 
sessile, remote along the slender rachis; sepals elongate-oval, obtuse. 
Neg. 27888. 

San Martin: Near Tarapoto, Spruce 4929 (type). 

Celosia virgata Jacq. Coll. Bot. 2: 279. 1788. C. virgata var. 
paniculata Suesseng. Repert. Sp. Nov. 39: 5. 1935. 

Plants erect, a meter high or less, glabrous, herbaceous or suf- 
frutescent; leaf blades broadly ovate to lanceolate or elliptic, 4-15 
cm. long, acuminate or cuspidate-acuminate, at the base abruptly 
contracted and long-decurrent upon the petiole; spikes sessile or 
pedunculate, short or elongate, in terminal or axillary panicles. 

Cajamarca: Ja6n, Raimondi 637, type of var. paniculata. Palmal, 
Prov. Santa Rosa, Raimondi 1131. Ranging northward to the West 
Indies and southern Mexico. 

3. CHAMISSOA HBK. 

Erect or scandent herbs or shrubs; leaves alternate, petiolate, 
with broad blades; flowers perfect or monoecious, each subtended by 
usually 3 bracts, spicate, the spikes axillary or terminal, simple or 
paniculate; sepals 5; stamens 5, connate at the base; staminodia 
none; ovule 1; style short or elongate, the 2 stigmas short or elongate; 
utricle thin-walled, circumscissile, surrounded by the persistent 
calyx; seed involved in an aril, or the aril minute. 

Aril minute; style elongate. Fruit subtruncate at the apex. 

1. C. Maximiliani. 
Aril well developed, involving the seed; style short. 

Fruit rounded at the apex 2. C. macrocarpa. 

Fruit truncate and emarginate at the apex, or conspicuously 
areolate 3. C. altissima. 

Chamissoa altissima (Jacq.) HBK. Nov. Gen. & Sp. 2: 197. 
1817. Celosia paniculata L. Sp. PI. ed. 2. 298. 1762, not L. 1753. 
Achyranthes altissima Jacq. Enum. PI. Carib. 17. 1760. A. baccata 
Pavon ex Moq. in DC. Prodr. 13, pt. 2: 250. 1849, in syn. Kokera 
paniculata Kuntze, Rev. Gen. 542. 1891. 



482 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plants herbaceous or suffrutescent, stout, often much elongate 
and scandent, sometimes lower and suberect, glabrous or nearly so; 
leaves slender-petiolate, the blades oval to lanceolate, 6-18 cm. 
long, acute or acuminate, often rather thick; panicles naked or leafy, 
the spikes usually dense and thick; flowers greenish white or green, 
the sepals 3-4 mm. long, ovate or broadly ovate, acute or acuminate; 
utricle equaling or slightly exceeding the sepals; seed black, 2-2.5 
mm. in diameter. 

Loreto: Lower Rio Nanay, Williams 432, 489, 521, 520, 510, 504, 
498. Pebas, in forest, Williams 1852. La Victoria, edge of forest, 
in pasture, or along stream, Williams 2670, 2666, 2650, 2523. 
Caballo-cocha, in forest or cleared land, Williams 2417, 2295. 
Iquitos, Williams 8196, 8028. San Martin: Lamas, 840 meters, 
Williams 6486. Pongo de Cainarachi, 230 meters, Klug 2648. 
Without locality, Weberbauer 6964> Generally distributed in the 
lowlands of tropical America. 

The plant is abundant in many regions, growing usually in clear- 
ings or thickets. In some plants all or most of the flowers are sterile, 
the inflorescence then being much more lax and slender than in 
fertile plants. 

Chamissoa macrocarpa HBK. Nov. Gen. & Sp. 2: 197. 1817. 
Celosia tomentosa Willd. ex R. & S. Syst. Veg. 5: 531. 1819. 

Plants low, herbaceous or suffrutescent, often scandent; leaves 
usually bright green when dried, slender-petiolate, the blades oblong- 
ovate or ovate-elliptic, 15 cm. long or less, acute or acuminate, 
glabrous or nearly so; panicles rather lax and open, the flowers 4 mm. 
long, green; sepals ovate-oblong; utricle usually surpassing the 
calyx, circumscissile below the middle. 

Loreto: Balsapuerto, 220 meters, Klug 2997 (determination 
uncertain). San Martin: Morales, near Tarapoto, in forest, Williams 
5666. Tarapoto, Williams 6577. Juanjui, 400 meters, in forest, 
Klug 3859. Ranging to Brazil and Colombia, and perhaps also to 
Mexico. 

Chamissoa Maximiliani Mart, ex Moq. in DC. Prodr. 13, pt. 
2: 251. 1849. 

Plants rather slender, herbaceous or suffrutescent, erect or sub- 
scandent, usually 1 meter long or less; leaves slender-petiolate, 
commonly drying bright green, the blades ovate or ovate-oblong, 
4-8 cm. long, acuminate, acute to rounded at the base, puberulent 
on the veins or almost glabrous; inflorescence usually rather lax and 



FLORA OF PERU 483 

open; sepals 3 mm. long, acuminate; utricle shorter than the calyx, 
areolate at the apex; seed black, 1.5 mm. in diameter. Neg. 7358. 

Junin: Colonia Perene", 600 meters, in forest, Killip & Smith 
25163. San Martin: Chazuta, 260 meters, in forest, King 4077. 
Juanjui, 400 meters, in forest, Klug 3782. Paraguay to Surinam 
and Costa Rica. 

All the Peruvian species of Chamissoa are almost identical in 
general appearance, and can be distinguished only by fruit char- 
acters. Specimens of C. Maximiliani usually are bright green when 
dried, and the inflorescence is more lax and open than is usual in C. 
altissima. 

4. AMARANTHUS L. 

Annuals; leaves alternate, petiolate; flowers usually green, 
monoecious, dioecious, or polygamous, bracteate and bibracteolate, 
glomerate, the glomerules axillary or disposed in spikes, the spikes 
often paniculate; sepals usually 5, sometimes 1-3; stamens normally 
5, the subulate or filiform filaments distinct; ovary 1-ovulate; style 
short or wanting, the style branches 2-3; utricle usually included in 
the perianth, indehiscent, cirsumscissile, or bursting irregularly, 
2-3-dentate at the apex. There is considerable doubt regarding the 
validity of some of the forms listed here as species. The forms re- 
lated to A. hybridus are variable, and authors are not in agreement 
as to their status. 

Plants armed with axillary spines A. spinosus. 

Plants unarmed. 

Flowers all in axillary clusters A. peruvianus. 

Flowers chiefly in terminal spikes. 
Utricle indehiscent. 

Utricle rugose A. gracilis. 

Utricle smooth A. deflexus. 

Utricle circumscissile. 

Sepals of the pistillate flowers spatulate, contracted into a 
claw, urceolate in age, pinnate-nerved .... A. Haughtii. 
Sepals of the pistillate flower oblong to obovate, not con- 
tracted into a claw or, if somewhat spatulate, not 
urceolate. 

Seeds white or whitish A. caudatus. 

Seeds black. 



484 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Bracts conspicuously longer than the sepals. 
Bracts green, usually twice as long as the sepals; 
terminal spike commonly short and thick, erect. 

A. hybridus. 

Bracts deep red or purple, less than twice as long as the 
sepals; terminal spike slender, much elongate, 

recurved A. cruentus. 

Bracts not exceeding the sepals, often much shorter. 
Flower spikes short, very dense, usually 1 cm. thick 

or more A. celosioides. 

Flower spikes much elongate, slender and often inter- 
rupted, commonly 5-7 mm. thick. . . .A. dubius. 

Amaranthus caudatus L. Sp. PI. 990. 1753. 

Plants coarse and erect, often 2 meters high or more, at least in 
cultivation, sparsely branched, glabrous, or sparsely villous about the 
inflorescence; leaves slender-petiolate, the blades lanceolate to 
rhombic-ovate, 20 cm. long or less, acute at base and apex; panicles 
usually very large and composed of numerous long, slender, dense 
spikes, these often drooping; bracts twice as long as the sepals or 
shorter; sepals of the pistillate flower 1.5-2 mm. long, usually spatu- 
late, often red or purple; utricle circumscissile at the middle; seeds 
whitish in cultivated forms, dull, not shining and black as in most 
species. 

Without locality: Weberbauer 5426. Probably native of the 
Old World tropics. 

Called "Inca pachaqui" in Bolivia. In some parts of the Andes 
the plant is grown extensively for its seeds, which are boiled and 
eaten, or ground into meal. The plant is grown for food in central 
Mexico, as well as in Abyssinia and Tibet and other regions of the 
Old World. 

Amaranthus celosioides HBK. Nov. Gen. & Sp. 2: 194. 1817. 

Plants tall and stout or more often low, often much branched; 
leaves on long, slender petioles, the blades orbicular to broadly 
ovate or ovate-rhombic, 6 cm. long or less, broadly rounded to acutish 
at the apex; flower spikes green or sometimes sparsely tinged with 
red, short, thick, very dense, erect; bracts mostly shorter than the 
sepals; sepals 5; utricle circumscissile. 

Huanuco: Mito, 2,700 meters, a dooryard weed, 1684- Junin: 
Tarma, 3,100 meters, open hillside, Killip & Smith 21926. Lima: 



FLORA OF PERU 485 

Lima, 150 meters, roadsides, 68. Piura: Parinas Valley, Haught 
F179. Without locality, Weberbauer 5332. Chile to Venezuela. 
"Ratago." Called "bledo" in Ecuador. 

Amaranthus cruentus L. Syst. Veg. ed. 10. 1269. 1759. A. 
paniculatus L. Sp. PI. ed. 2. 1406. 1763. 

Plants stout, erect, 2 meters high or less, simple or much branched, 
the stems usually pubescent, villous about the inflorescence; leaves 
long-petiolate, the blades rhombic-ovate to ovate-lanceolate, 30 
cm. long or less, acute or acuminate, acute to attenuate at the base; 
panicles usually large, composed of numerous spikes, the terminal 
spike usually much elongate, slender, and recurved, deep red or 
purple; bracts usually half longer than the sepals; sepals of the pis- 
tillate flower 1.5 mm. long, often obtuse or rounded at the apex; 
utricle exceeding the sepals, circumscissile at the middle. 

Loreto: Lower Rio Nanay, Williams 348. Cultivated for orna- 
ment and escaped in tropical and temperate regions of both hemi- 
spheres; perhaps native of tropical America. 

Known in Ecuador by the names "bledo Colorado," "sangorache," 
and "ataco." 

Amaranthus deflexus L. Mant. PI. 2: 295. 1771. Euxolus 
deflexus Raf. Fl. Tell. 3: 42. 1837. 

Plants much branched, glabrous or nearly so, ascending or 
decumbent, 60 cm. high or less; leaves slender-petiolate, the blades 
rhombic-ovate to lanceolate, small, narrowed to the obtuse apex, 
rounded to cuneate at the base; flowers green or purplish, chiefly 
in short, dense, stout, terminal spikes 2-8 cm. long; bracts shorter 
than the flowers; utricle oblong, smooth, longer than the sepals. 

Arequipa: Posco, 550 meters, Cook & Gilbert 1*3. Probably native 
of the Old World, but adventive in the American tropics; specimens 
seen from Chile, Argentina, and Uruguay. 

Amaranthus dubius Mart. PI. Hort. Erlang. 197. 1814. A. 
tristis Willd. Hist. Amaranth. 21. 1790, saltern ex parte, not L. 1753. 

Plants slender, erect, 1 meter high or less, often much branched, 
glabrous throughout or nearly so; leaves slender-petiolate, the 
blades ovate or rhombic-ovate, 3-12 cm. long, narrowed to the obtuse 
or rounded and often emarginate apex, rounded to acutish at the 
base, thin; spikes green or whitish, 5-25 cm. long, most of the flowers 
sterile; bracts scarious, mostly shorter than the sepals; utricle thin- 
walled, often exceeding the sepals. 



486 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ayacucho: Aina, 750-1,000 meters, open hillside, Killip & Smith 
23110. Junin: La Merced, 700 meters, Killip & Smith 24057 
Loreto: La Victoria, edge of forest, Williams 2893. Recreo, in 
chacara, Williams 4155. Generally distributed in tropical America. 

Amaranthus gracilis Desf. Tabl. Bot. 43. 1804. Chenopodium 
caudatum Jacq. Coll. Bot. 2: 325. 1788, not Amaranthus caudatus L. 
1753. Euxolus caudatus Moq. in DC. Prodr. 13, pt. 2: 274. 1849. 

Plants rather slender, usually less than 1 meter high, much 
branched, glabrous; leaves long-petiolate, thin, the blades ovate or 
ovate-rhombic, 8 cm. long or less, rounded or narrowed at the apex, 
with emarginate tip, rounded to broadly cuneate at the base; spikes 
panicled, 4-12 cm. long, 4-8 mm. thick; bracts ovate, scarious, 
much shorter than the flowers; sepals 3, oblong or linear-oblong, 
acute or obtuse, cuspidate; utricle strongly rugose, equaling or longer 
than the sepals. 

Libertad: Salaverry, sandy slope of Morro Hill, Johnston 3520. 
Loreto: Mishuyacu, in clearing, Klug297. La Victoria, in pasture, 
Williams 2731. Iquitos, in pasture, Williams 8188. Piura: Parinas 
Valley, Haught 265. Tropics of both hemispheres. "Ataco." 

Amaranthus Haughtii Standl. Field Mus. Bot. 11: 149. 1936. 

Plants nearly glabrous, 20-60 cm. high; leaves on long, slender 
petioles, the blades lance-linear to ovate-elliptic, narrowed to the 
obtuse or rounded apex, long-attenuate to the base; flowers dioecious, 
pale green, arranged in axillary clusters and in short, dense, terminal 
spikes; sepals 5, the pistillate broadly obovate-spatulate, 2-2.5 
mm. long, broadly rounded or subtruncate at the apex, with green 
nerves; utricle much shorter than the sepals, tridentate at the apex, 
circumscissile near the middle. 

Piura: Parinas Valley, Haught F147 (type), F148, 221, 227. 
Talara, along sandy draws and on cliff talus, Johnston 3509. 
Without locality, Weberbauer 5960. 

Easily recognized by the dioecious flowers and spatulate pistillate 
sepals. 

Amaranthus hybridus L. Sp. PI. 990. 1753. A. hypocondriacus 
L. Sp. PL 991. 1753. A. chlorostachys Willd. Hist. Amaranth. 34. 1790. 

Plants low or tall and coarse and as much as 2.5 meters high, 
usually much branched, erect, glabrous, rough-puberulent, or some- 
what villous; leaves slender-petiolate, the blades lanceolate to 
rhombic-ovate, 3-15 cm. long, acute to rounded at the apex, some- 



FLORA OF PERU 487 

what pubescent or glabrous; flower spikes axillary and terminal, 
stout and dense, the terminal ones erect; bracts lanceolate or ovate, 
with spinose tips; sepals of the pistillate flower 1.5-2 mm. long, 
1-nerved, usually mucronate; utricle equaling or shorter than the 
sepals, circumscissile at the middle, often rugose. 

Cuzco: Ollantaitambo, 3,000 meters, Cook & Gilbert 715. Ha- 
cienda Capana, Paucartambo Valley, 3,450 meters, Hen era 1053. 
Vilcanota Valley, 2,900 meters, H err era 1073. Valle Lares, Diehl 
2443- Lima: Matucana, 2,400 meters, grassy stream margin, 320. 
San Geronimo, along trail, 150 meters, 5915. Tropical and temperate 
regions of almost the whole earth; usually a weed in cultivated or 
waste ground. "Tataco," "jataco." 

The species is a somewhat variable one* 

Amaranthus peruvianus (Schauer) Standl., comb. nov. Mengea 
peruviana Schauer, Nov. Act. Nat. Cur. 19: Suppl. 1: 406. 1843. 

Plants small and prostrate, glabrous, the branches 15 cm. long 
or less, very leafy; leaves on rather long and somewhat margined 
petioles, the blades orbicular or nearly so, mostly 4-7 mm. long, 
broadly rounded and often emarginate at the apex, contracted or 
narrowed at the base, with rather conspicuous, pale margins and few 
nerves; flowers few in the axils, the bracts white and scarious; sepal 
only 1; stamen 1; utricle membranaceous, greenish, slightly longer 
than the sepals. 

Puno: Type from Lake Titicaca, at 3,870 meters, Meyen (photo, 
seen, ex hb. Berol.). Also in Bolivia. 

Easily recognized by the prostrate habit, very small leaves, and 
few flowers. 

Amaranthus spinosus L. Sp. PI. 991. 1753. A. caracasanus 
HBK. Nov. Gen. & Sp. 2: 195. 1817. 

Plants nearly glabrous, erect or ascending, 1 meter high or less, 
much branched, armed with stout, fleshy spines in the leaf axils; 
leaves slender-petiolate, the blades rhombic-ovate to lanceolate, 12 
cm. long or less, narrowed to the obtuse apex; pistillate flowers in 
dense, globose, axillary clusters, the staminate in long, slender, 
terminal, pale spikes; bracts usually shorter than the sepals, some- 
times longer; sepals 5 in the pistillate flower, 1.5 mm. long, obtuse 
or acute; utricle about equaling the sepals, irregularly and imper- 
fectly circumscissile. 

Cuzco: Rio Alto Urubamba, Diehl 2491. Junin: Chanchamayo, 
Isern 2365. Piura: Near Saucecito, Haught 125. San Martin: 



488 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Tarapoto, in pasture, Williams 5624- San Roque, Williams 7246. 
Pongo de Cainarachi, 230 meters, Klug 2733. Tropical and tem- 
perate regions of both hemispheres, probably native of tropical 
America. "Ataco," "ataco casha." 

The plant is a common weed of the lowland tropics of America, 
and extends far northward into the United States. In some regions 
the leaves and young shoots are cooked and eaten like spinach. 

5. CYATHULA Lour. 

Annual or perennial herbs; leaves opposite, petiolate; flowers 
fasciculate, each fascicle consisting of 1-2 perfect flowers and few or 
many sterile ones, the fascicles spicate or capitate, reflexed in age; 
segments of the sterile flowers finally produced into elongate bristles, 
these uncinate at the apex; stamens 5, the filaments united at the 
base, the staminodia linear or quadrate and lacerate; stigma capitate; 
utricle included in the perianth, indehiscent. 

Cyathula achyranthoides (HBK.) Moq. in DC. Prodr. 
13, pt. 2: 326. 1849. Desmochaeta achyranthoides HBK. Nov. Gen. 
&Sp. 2:210. 1818. 

A branched annual or perennial, 1 meter high or less, often genic- 
ulate and rooting at the base; leaves short-petiolate, the blades 
oval to rhombic-elliptic, thin, 5-15 cm. long, acuminate, cuneate at 
the base, strigose or glabrate; spikes 4-20 cm. long, 6-7 mm. thick, 
the flowers green; perfect flowers 2. Neg. 7390. 

Huanuco: Pampayacu, 1,050 meters, forest clearing, 5071. 
Junin: Chanchamayo Valley, 1,200 meters, Schunke 138, 599, 600. 
La Merced, 700 meters, Kittip & Smith 23703. Loreto: Mishu- 
yacu, 100 meters, Klug 148. Puerto Arturo, 135 meters, dense 
forest, Killip & Smith 27907. Fortaleza, Yurimaguas, Williams 
4269. Paraiso, Williams 3351. Leticia, borders of forest and river 
banks, common, Williams 3071. La Victoria, Williams 2561. Ca- 
ballo-cocha, Williams 2032. Iquitos, Williams 1347. Lower Rio 
Nanay, Williams 671, 543. San Martin: San Roque, Williams 6955. 
Tarapoto, Williams 5712. Bolivia and Brazil to Mexico. 

The hooked bristles of the flowers adhere tenaciously to clothing 
and other articles, and penetrate the flesh easily and painfully. 

6. FROELICHIA Moench 

Annual or perennial herbs, erect or procumbent, with simple or 
branched stems; leaves opposite, sessile or petiolate; flowers perfect, 



FLORA OF PERU 489 

spicate, the spikes sessile or pedunculate; perianth 5-lobate, the lobes 
glabrous, the tube lanate, indurate in age and longitudinally winged 
or cristate, or bearing longitudinal rows of spines; stamens 5, the 
filaments united into an elongate tube, this 5-lobate at the apex; 
anthers sessile in the sinuses between the lobes; style elongate, with a 
capitate stigma, or the stigma sessile and penicillate; utricle inde- 
hiscent; seed 1. 

Froelichia interrupta (L.) Moq. in DC. Prodr. 13, pt. 2: 421. 
1849. Gomphrena interrupta L. Sp. PI. 224. 1753. F. lanata Moench, 
Meth. 50. 1794. Oplotheca tomentosa Mart. Nov. Gen. & Sp. 2: 48. 
1826. F. tomentosa Moq. in DC. Prodr. 13, pt. 2: 421. 1849. 

Plants perennial from a thick root, or sometimes flowering the 
first year from seed, the stems ascending or decumbent, usually 
much less than 1 meter high, white-lanate or sericeous; leaves petio- 
late or the upper subsessile, the blades ovate-orbicular to oblong, 
3-10 cm. long, obtuse or acute, sericeous or floccose-tomentose 
beneath; inflorescence lax and interrupted; fruiting calyx deltoid, 
broadly winged laterally, the thin wings entire or crenulate, the 
sides of the calyx tube with or without spine-like tubercles. Negs. 
3199, 3196. 

Arequipa: Mollendo, Weberbauer 1491. Piura: Between Piura 
and Romala, 100-250 meters, Weberbauer 5951. Tumbez: South- 
east of Hacienda La Choza, 100-200 meters, Weberbauer 7722. 
Chile to Paraguay and northward to Texas and the West Indies. 

7. GUILLEMINEA HBK. 

Prostrate, much branched, densely leafy, lanate, perennial herbs, 
forming small mats; leaves opposite, those of a pair unequal, petio- 
late, the petioles connate at the base; flowers small, perfect, glomerate 
in the leaf axils; perianth lanate, with an obconic tube; stamens 5, 
perigynous, the filaments short-connate; style short, the stigma 
emarginate; ovule 1; utricle membranaceous, indehiscent. 

Guilleminea densa (Willd.) Moq. in DC. Prodr. 13, pt. 2: 338. 
1849. Illecebrum densum Willd. ex R. & S. Syst. Veg. 5: 517. 1819. 
G. illecebroides HBK. Nov. Gen. & Sp. 6: 42. 1823. Achyranthes 
conferta Pavon ex Moq. loc. cit. in syn. Brayulinea densa Small, 
Fl. Southeast. U. S. 394. 1903. 

Stems numerous from a thick, vertical root, 25 cm. long or less, 
densely lanate; leaves on short, winged petioles, the blades elliptic 
to broadly oval, 3-15 mm. long, obtuse or acute, abruptly contracted 



490 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

at the base, densely villous or lanate beneath; flowers white, the 
glomerules much shorter than the leaves; bracts ovate, acute, scari- 
ous; calyx 2-2.5 mm. long. Negs. 3233, 3201. 

Cajamarca: Cascas, Raimondi 7692. Cuzco: Valle Lares, Diehl 
2475, 2429a. Hacienda Chancamayo, Diehl 2478. Valle del Apuri- 
mac, Herrera in 1929. Huanuco: Huanuco, 2,100 meters, stony 
slopes, 3242; Ruiz & Pavdn 5-69. Huanuco and Cajatambo, Ruiz. 
Junin: Tarma, 3,100 meters, open hillside, Killip & Smith 21830. 
Huancayo, 3,400 meters, rocky slope, Ledig 42. Lima: Matucana, 
2,400 meters, gravelly river trail, 185, 2924- Argentina and Brazil 
to the southwestern United States. 

Ruiz records the vernacular name as "yerba de la sangre." 

8. PFAFFIAMart. 

Herbs or shrubs, erect or scandeht; leaves opposite, sessile or 
short-petiolate; flowers mostly perfect, bracteate and bibracteolate, 
capitate or spicate, the spikes or heads chiefly pedunculate, often 
numerous and paniculate; perianth sessile, terete, the 5 segments 
free, pilose or lanate; filaments united in a 5-lobed tube, the lobes 
fimbriate, dentate, or 3-lobate; staminodia none; style very short or 
wanting, the stigma capitate or bilobate; utricle indehiscent. 

Flower spikes 1 cm. in diameter; leaves sericeous beneath with very 

long, lax, silvery hairs P. holosericea. 

Flower spikes about 6 mm. in diameter; leaves never silvery-sericeous. 
Spikes short, very dense; leaves linear or lanceolate. 

P. stenophylla. 

Spikes elongate, slender, much interrupted; leaves broad. 
Panicle branches appressed-pilose with simple, straight hairs. 

P. paniculata. 

Panicle branches puberulent with fine, stellately branched or 
hispidulous, spreading hairs P. grandiflora. 

Pfaffia grandiflora (Hook.) Fries, Arkiv Bot. 16, No. 12: 10. 
1920. Iresine grandiflora Hook. Icon. PI. 2: pi. 102. 1837. Hebanthe 
decipiens Hook. f. in B. & H. Gen. PI. 3: 41. 1880. Gossypianthus 
decipiens Kuntze, Rev. Gen. 543. 1891. 

A large, scandent shrub, similar to P. paniculata; leaves glabrate 
in age, when young copiously puberulent beneath with short, conic 
hairs, these hispidulous near the base, erect; inflorescence and flow- 
ers like those of P. paniculata; lateral lobes of the filaments minute 
or absent, the filaments in P. paniculata being deeply trifid. 



FLORA OF PERU 491 

Cuzco: Valle Lares, Diehl 2407. Quebrada Versalles, Diehl 
245 la. Junin: La Merced, 600 meters, edge of forest, 5379, 5796. 
Loreto: Yurimaguas, 135 meters, Killip & Smith 29074- Puno: 
Sangaban (Lechler in 1854). Bolivia; Brazil. 

The specimens from Cuzco could be referred equally well to P. 
Bangii Fries, a Bolivian species, distinguished by having the lower 
leaf surface covered with elongate, soft, articulate, more or less 
hispidulous hairs, but it is doubtful whether that is more than a 
casual variant of P. grandiflora. 

P. grandiflora, if I have identified it correctly, is readily dis- 
tinguishable from the widespread P. paniculata by its pubescence. 
There are supposed to be stamen differences also, but there is doubt 
regarding their importance or value. 

Pfaffia holosericea (Moq.) Standl., comb. nov. Gomphrena 
holosericea Moq. in DC. Prodr. 13, pt. 2: 386. 1849. Hebanthe 
holosericea Mart, ex Moq. loc. cit. in syn. G. holosericea Moq. op. cit. 
463. 1849. 

Plants large and much branched, commonly trailing or more or 
less scandent, suffrutescent, the slender stems pilose with long, 
appressed or somewhat spreading hairs; leaves short-petiolate, large 
or small, lanceolate or linear-lanceolate, long-attenuate, obtuse or 
acute at the base, green and thinly pilose on the upper surface, cov- 
ered beneath, at least when young, with very long, silvery hairs, in 
age sometimes almost glabrous; heads whitish, subglobose, short- 
pedunculate, in lax, open panicles; bracts short and broad, lanate; 
sepals linear-lanceolate, acute, 3-nerved, 4-5 mm. long, pilose at the 
base with long, straight, white hairs as long as the sepals. 

Huanuco: La Merced, 600 meters, openly wooded slope, 5359. 
Loreto: Balsapuerto, 220 meters, in forest, Klug 3011 (determination 
doubtful). Bolivia and Brazil. 

Pfaffia paniculata (Spreng.) Kuntze, Rev. Gen. 543. 1891. 
Iresine paniculata Spreng. Syst. Veg. Cur. Post. 103. 1827. Hebanthe 
paniculata Mart. Nov. Gen. & Sp. 2: 43. pi. 140, 142. 1826. Gom- 
phrena paniculata Moq. in DC. Prodr. 13, pt. 2: 385. 1849. 

A large, scandent shrub, the branches appressed-pilose with 
simple hairs or glabrate; leaves short-petiolate, the blades firm, 
elliptic to ovate-oblong, acute or acuminate or sometimes obtuse, 
rounded to acute at the base, glabrate above, beneath rather sparsely 
appressed-pilose or soon glabrate; spikelets usually much interrupted 
and many-flowered, in large, open panicles; sepals ovate or oblong, 



492 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

obtuse, 3-4 mm. long, 3-nerved, pubescent, pilose at the base with 
long, straight, whitish hairs. 

Huanuco: Pampayacu, 1,050 meters, open, brushy hills, 5122. 
Loreto: Rio Itaya, Williams 52. Lower Rio Nanay, in thicket, 
Williams 503. San Martin: Tarapoto, edge of pasture, Williams 
5481, 6749, 6741, 6309, 5513. San Roque, Williams 7367. Zepe- 
lacio, 1,100 meters, in forest, Williams 3715. Bolivia and Brazil 
to French Guiana and Ecuador. 

Pfaffia stenophylla (Spreng.) Stuchl. Repert. Sp. Nov. 12: 357. 
1913. Gomphrena stenophylla Spreng. Syst. Veg. 1: 823. 1825. P. 
glauca Spreng. Cur. Post. 107. 1827. Serturnera glauca Mart. Nov. 
Gen. & Sp. 2: 37. pi. 136. 1826. G. glauca Moq. in DC. Prodr. 13, 
pt. 2: 384. 1849. 

Plants chiefly herbaceous, branched, slender, the older branches 
glabrate, the young ones sparsely appressed-pilose; leaves short- 
petiolate, green when dried, densely appressed-pilose when young 
but often almost glabrous in age; flower heads whitish, globose or 
in age oblong; bracts very short; sepals glabrous, obtuse, 2 mm. long, 
conspicuously 3-nerved. Neg. 7377. 

Loreto: Iquitos, Williams 7942, 3553, 1507, 8132. Mishuyacu, 
in clearing, Klug 1237. Argentina and Bolivia to the Guianas. 
"Trompetero." 

Some of Williams' specimens are taken from garden plants, 
although there is no apparent reason why so inconspicuous a plant 
should be in cultivation. 

9. ALTERNANTHERA Forsk. 

Herbs or shrubs, often scandent; leaves opposite; flowers perfect, 
bracteate and bibracteolate, capitate or spicate, usually compressed, 
the heads axillary or terminal; sepals distinct, unequal; filaments 
united to form a short or elongate tube, this with 3-5 antheriferous 
lobes and as many short or elongate staminodia; ovary 1-ovulate; 
style short or elongate; stigma capitate. 

Flower heads sessile, all or chiefly axillary, never paniculate. 

Staminodia shorter than the filaments; pubescence of simple or 
merely scaberulous hairs; sepals with greatly elongate, spine- 
like tips; flower heads mostly 7-10 mm. thick. 

Sepals 3-5 mm. long, densely villous; leaf blades longer than 
broad A. repens. 



FLORA OF PERU 493 

Sepals 6-7 mm. long, sparsely villous or glabrate above; leaf 

blades usually as broad as long A. pungens. 

Staminodia equaling or much longer than the filaments; pubes- 
cence of stellate or hispidulous hairs; sepals with short, spinose 
tips or merely acute; flower heads mostly smaller. 
Outer bracts laciniate-lobed ; leaves often colored with red or 

purple A. Bettzickiana. 

Outer bracts entire, like the others. 

Leaves glabrate, usually acute A. ficoidea. 

Leaves densely stellate-tomentose, usually rounded at the 
apex. 

Flower heads globose A. halimifolia. 

Flower heads oblong or often much elongate. 

A. peruviana. 

Flower heads pedunculate, or sometimes sessile but then terminal, 

often in large panicles. 
Flowers conspicuously stipitate within the bractlets, the stipe 

5-sulcate. 
Bractlets equaling or longer than the sepals, broadly cristate 

toward the apex A. dentata. 

Bractlets much shorter than the sepals, narrowly cristate or 

ecristate. 

Stems densely and minutely stellate-tomentose . . A. flavida. 
Stems pilose, villous, or glabrate. 
Stems pilose with very long, tawny, spreading hairs. 

A. villosa. 

Stems pilose with short and spreading or appressed hairs. 

Lower leaves mostly rounded or very obtuse at the apex; 

stems densely villous-tomentose with spreading 

hairs A. tomentosa. 

Lower leaves chiefly long-acuminate; stems usually gla- 
brate or appressed-pilose A. brasiliana. 

Flowers sessile within the bractlets. 
Peduncles simple, bearing a single head. 

Plants dwarf and matted, the stems rarely more than 5 cm. 

long A. lupulina. 

Plants large, with much elongate stems. 

Sepals pilose A. Lehmannii. 



494 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Sepals glabrous. 

Flower heads pink to purple or blackish; bractlets usually 
pubescent over nearly the whole surface. 

A. elongala. 
Flower heads usually white; bractlets glabrous or pilose 

only on the costa. 

Heads 11-14 mm. broad; leaves glabrous or nearly so. 

A. philoxeroides. 
Heads usually smaller; leaves copiously pilose. 

Bracts glabrous A. lanceolata. 

Bracts pilose on the costa A. microcephala. 

Peduncles trifid, or bearing a cluster of 2-3 heads, some of the 
peduncles on the plant, especially the lower ones, frequently 
with a single head. 
Sepals pilose. 

Sepals 2.5-3 mm. long A. porrigens. 

Sepals 5-8 mm. long A. pubiflora. 

Sepals glabrous, or rarely inconspicuously pilose at the base. 
Stems and leaves densely stellate-tomentose . .A. cakicola. 
Stems and leaves not stellate-tomentose. 
Clusters of heads naked at the base. Sepals 3-4 mm. 

long A. paniculata. 

Clusters of heads, some or all of them, subtended at the 

base by leaves. 
Heads 1 cm. broad or larger; plants large and sub- 

scandent A. piurensis. 

Heads much less than 1 cm. broad; plants small. 
Bracts at the base of the head with a conspicuous 

red blotch; heads all sessile A. Macbridei. 

Bracts green; heads partly pedunculate. 

A. Dominii. 

Alternanthera Bettzickiana (Regel) Standl. Field Mus. Bot. 
3: 254. 1930. Telanthera Bettzickiana Regel, Gartenflora 11: 178. 
1862. Achyranthes picta Pass. Giardini 9: 515. 1863. Alternanthera 
spathulata Lem. 111. Hort. 12: pi. 445. 1865. Telanthera picta C. 
Koch, Wochenschr. Gartn. 9: 15. 1866. 

Plants erect or ascending, annual or perennial, 40 cm. high or less, 
branched, the stems villous when young but soon glabrate; petioles 



FLORA OF PERU 495 

slender, equaling or shorter than the blades; leaf blades rhombic or 
rhombic-obovate, sometimes spatulate, 1-3.5 cm. long or somewhat 
larger, acuminate or abruptly acute and apiculate, abruptly long- 
attenuate to the base, undulate or crispate, sparsely appressed- 
pilose when young but soon glabrate, green or often colored with 
purple or yellowish, often variegated; heads axillary, sessile, usually 
solitary, ovoid or oblong, whitish; bracts and bractlets broadly ovate, 
aristate-acuminate, at least the lowest ones laciniately lobed, gla- 
brous; sepals lance-oblong, acute or acuminate, 3-nerved, sparsely 
pilose; staminodia equaling the filaments, ligulate, laciniateat the apex. 

Loreto: Pro, on the Amazon, in forest, Williams 1975. Lower 
Rio Nanay, in forest, Wittiams 666. Pebas, in forest, Williams 1922. 
Yurimaguas, Williams 4056. Iquitos, Williams 3574- Brazil to 
the Guianas; often escaped from cultivation elsewhere in tropical 
America. "Paloma chaqui." 

The plant often is grown for ornament in northern gardens. 
Probably it is a cultivated derivative of Alternanthera ficoidea, and 
it is rather doubtful whether it is known anywhere in a truly 
wild state. 

Alternanthera brasiliana (L.) Kuntze, Rev. Gen. 537. 1891. 
Gomphrena brasiliana L. Cent. PI. 2: 13. 1756. Gomphrena brasili- 
ensis Lam. Encyl. 1: 119. 1783. Achyranthes capituliflora Bert, in 
Colla, Hort. Ripul. 4. pi. 18. 1824. Mogiphanes straminea Mart. 
Nov. Gen. & Sp. 2: 35. 1826. M. ramosissima Mart. op. cit. 36. 
1826. Telanthera capituliflora Moq. in DC. Prodr. 13, pt. 2: 381. 
1849. Achyranthes geniculata Pavon ex Moq. loc. cit. in syn. T. 
ramosissima Moq. loc. cit. T. brasiliana Moq. op. cit. 382. 1849. 

Plants usually large and much branched, herbaceous or suffru- 
tescent, often scandent, the branches slender or stout, pilose with 
rather long and appressed or sometimes short and somewhat spread- 
ing hairs, often glabrate or almost glabrous from the first; leaves 
slender-petiolate, the blades oblong or lanceolate to ovate, 4-10 cm. 
long or even larger, acute or acuminate, rounded to acute at the base, 
appressed-pilose or sericeous, often nearly glabrous; flower spikes 
globose or short-oblong, 1-2.5 cm. long, 8-12 mm. thick, stramine- 
ous or whitish; peduncles simple or trifid; bracts and bractlets half 
as long as the sepals or shorter, glabrous or nearly so, sometimes 
narrowly and obscurely cristate near the apex, the crest denticulate; 
sepals ovate-lanceolate or lance-oblong, 3-5 mm. long, rigid, acute, 
short-pilose; staminodia longer than the filaments, ligulate, laciniate 
at the apex. 



496 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cuzco: Quebrada Versalles, Diehl 24-50. Machupicchu, Soukup 
188. Huanuco: Cueva Grande, 1,050 meters, dense thicket, 4796. 
Muna, 2,100 meters, edge of thicket, 3960. Huanuco: Pillao, Ruiz 
& Pavdn 5-61. Junin: La Merced, 600 meters, stream bed, 5238; 
in thicket, Killip & Smith 23381 . Colonia Perene", 680 meters, open, 
rocky bank, Killip & Smith 24985. San Martin: Rio Mayo, Wil- 
liams 6194. Tarapoto, edge of forest and along river, Williams 5691, 
5445. Brazil to southern Mexico and the West Indies. 

The species is a somewhat variable one, but the Peruvian material 
is fairly uniform. Moquin lists Poeppig 152 from Peru as Telan- 
thera Moquini Webb, and I presume that the plant is Alternanthera 
brasiliana. 

Alternanthera calcicola Standl. Field Mus. Bot. 11: 151. 1936. 

Perennial, erect or procumbent, the branches densely stellate- 
tomentose; leaves subsessile or short-petiolate, lance-oblong to 
rounded-oval, 8-22 mm. long, obtuse or rounded at the apex, densely 
stellate- tomentose; flower heads few, triglomerate, terminal or a few 
axillary, chiefly in a small, dense, almost head-like inflorescence, 
stramineous or whitish; bracts almost equaling the calyx, glabrous; 
sepals 2.5 mm. long, glabrous, acutish or obtuse. 

Junin: La Oroya, 3,600 meters, limestone cliff ledges, 944* Hua- 
riaca, 2,850 meters, shrubby canyon side, 3110. 

Well marked by the dense, whitish, stellate tomentum. 

Alternanthera dentata (Moench) Stuchl. ex Fries, Arkiv Bot. 
16, No. 13: 11. 1921. Gomphrena brasiliensis Jacq. Coll. Bot. 2: 
278. 1788, non L. 1756. G. dentata Moench, Meth. Suppl. 273. 1802. 
Mogiphanes brasiliensis Mart. Nov. Gen. & Sp. 2: 34. 1826. M. 
Jacquini Schrad. Ind. Sem. Goetting. 1834: 4. 1834. Telanthera 
dentata Moq. in DC. Prodr. 13, pt. 2: 378. 1849, ex parte. Alter- 
nanthera brasiliana var. Jacquini Kuntze, Rev. Gen. 2: 538. 1891. 
Achyranthes Jacquini Standl. Journ. Wash. Acad. Sci. 5: 74. 1915. 
?A. Moquini var. grandiceps Suesseng. Repert. Sp. Nov. 35: 302. 1934. 

Plants perennial, erect or ascending, sometimes subscandent, 
herbaceous or suffrutescent, usually much branched, the stems 
appressed-pilose or glabrate; leaves slender-petiolate, the blades 
oval or ovate to oblong, mostly 4-10 cm. long, acute or acuminate, 
abruptly acute at the base, thin, sparsely or densely appressed- 
pilose or sericeous, sometimes glabrate; peduncles simple or trifid, 
elongate; flower heads globose or short-cylindric, 1-2.5 cm. long, 
about 1 cm. in diameter; bracts short, white, long-acuminate, gla- 



FLORA OF PERU 497 

brous; bractlets equaling or longer than the sepals, oblong, acute, 
sparsely villous, cristate dorsally, the crest serrulate; sepals lance- 
oblong, rigid, 3-nerved, acute, appressed-pilose, 3-3.5 mm. long; 
staminodia longer than the filaments, ligulate, lacerate at the apex. 

Loreto: Yurimaguas, in chacara, Williams 3916; Killip & Smith 
27584. Mishuyacu, 100 meters, in clearing, Killip & Smith 29918; 
Klug 1511. Piura: Cafia Dulce, 20 miles inland from Negritos, 
along watercourse, Haught FSl.Tumbez: Plain southeast of 
Hacienda La Choza, 100-200 meters, Weberbauer 7696. Brazil to 
the West Indies. 

The leaves are sometimes pink or deep purple. Killip and Smith 
report that they are used for coloring pottery. The type of A. 
Moquini var. grandiceps is Weberbauer 6366 from Piura. 

Alternanthera Dominii Schinz, Pflanzenfam. ed. 2. 16c: 76. 
1934. Telanthera nana Moq. in DC. Prodr. 13, pt. 2: 374. 1849, not 
A. nana R. Br. 1810. 

A slender annual, erect or decumbent, simple or branched, 30 
cm. high or less, the stems glabrous or nearly so; leaves on rather 
slender petioles, the blades thin, broadly rhombic-ovate to ovate or 
obovate, 2 cm. long or less, obtuse, contracted and decurrent at the 
base, sparsely pilose above, usually glabrous beneath; heads white, 
globose, 4-5 mm. in diameter, chiefly terminal, often in twos, one 
sessile, the other pedunculate and composed of 2-3 sessile heads, the 
clusters subtended by 2 slightly reduced leaves; bracts ovate, mu- 
cronate, glabrous, half as long as the calyx; sepals 1.5 mm. long, 
ovate-oblong, obscurely serrulate near the apex, obscurely 3-nerved, 
glabrous or pubescent at the base; staminodia equaling the filaments, 
laciniate at the apex. 

Lima: Matucana, 2,400 meters, among rocks on moist, grassy 
slope, 106. Type of T. nana from Peru, without locality, Mathews 
779. 

It is not at all certain that the collection cited is the same as the 
plant described as Telanthera nana. 

Alternanthera elongata (Willd.) Schinz, Nat. Pflanzenfam. 
ed. 2. 16c: 75. 1934. Gomphrena elongata Willd. ex R. & S. Syst. Veg. 
5: 542. 1819. Brandesia elongata Mart. Beitr. Amarant. 105. 1826. 
Telanthera elongata Moq. in DC. Prodr. 13, pt. 2: 372. 1849. 

Plants large and much branched, usually trailing or scandent, 
slender, pilose with appressed or spreading hairs; leaves mostly 



498 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

short-petiolate, the blades ovate to lance-oblong, acute or acuminate, 
pilose with appressed or spreading hairs, often densely sericeous- 
pilose beneath; peduncles axillary and terminal, short or elongate, 
simple or trifid; heads subglobose or oblong, about 7 mm. broad, 
rarely more than 1 cm. long, pink or purple, sometimes fuscous when 
dried; bractlets ovate, acuminate and short-mucronate, a third as 
long as the calyx, usually densely pubescent; sepals 2.5-3 mm. long, 
oblong, acutish or obtuse, glabrous, obscurely 3-nerved; staminodia 
longer than the filaments, laciniate at the apex. Negs. 3231, 7387. 

Amazonas: Chachapoyas, 2,700 meters, Williams 7535. 
Huanuco: San Rafael, 2,550 meters, river canyon slopes, 3140. 
Northward to Colombia. 

Alternanthera elongata var. nigriceps (Hook.) Suesseng. 
Repert. Sp. Nov. 35: 299. 1934. A. nigriceps Hook. Bot. Misc. 2: 
237. 1831. A. nigripes Steud. Norn. Bot. ed. 2. 1: 65. 1840. Telan- 
them elongata var. nigriceps Moq. in DC. Prodr. 13, pt. 2: 372. 1849. 
"Alternanthera nigrescens Hook." ex Moq. loc. cit. in syn. Achy- 
ranthes atra Pavon ex Moq. loc. cit. in syn. 

Like the species, except that the flowers in age become black or 
purple-black; heads sometimes 8-9 mm. in diameter. Neg. 7386. 

Huanuco: Llata, 2,100 meters, grassy ledges of river cliffs, 
2,100 meters, 2231. Junin: Yanahuanca, 3,000 meters, rocky slopes, 
1247. Tarma, 3,100 meters, open hillside, Kittip & Smith 21936. 
Between Viques and Ingahuasi, 3,150 meters, open hillside, Killip 
& Smith 22169. Lima: Type from Obrajillo. San Buenaventura, 
2,700 meters, open, rocky slopes, Pennell 14511. Matucana, 2,400 
meters, stony slope, 117. Canta, 2,800 meters, open, rocky slope, 
Pennell 14348. Canta and Obrajillo, Ruiz & Pavon. 

The variety, a well marked color form, but probably only a slight 
variation from the type, is known only from Peru. The heads often 
are somewhat larger than in the typical form of A. elongata, but not 
always so. Suessenguth has described (Repert. Sp. Nov. 35: 299. 
1934) two forms of this variety which have but scant systematic 
interest: f. major (Pennell 14511 from San Buenaventura), and /. 
minor (Raimondi 1528 from Dept. Ancash, and Weberbauer 3037, 
collected between Samanco and Caraz). 

Alternanthera ficoidea (L.) R. Br. Prodr. 417. 1810. Gom- 
phrena ficoidea L. Sp. PI. 255. 1753. Achyranthes ficoidea Lam. 
Encycl. 1: 548. 1785. Bucholzia ficoidea Mart. Nov. Gen. & Sp. 2: 51. 
1826. Telanthera ficoidea Moq. in DC. Prodr. 13, pt. 2: 363. 1849. 



FLORA OF PERU 499 

Perennial, prostrate or ascending, herbaceous, the branches at 
first pubescent with short, appressed, branched or hispidulous, white 
hairs; leaves petiolate, the blades elliptic to oblong, 2.5-5.5 cm. long, 
acute or rarely obtuse, when young finely pubescent with short, 
hispidulous hairs; heads axillary, solitary or glomerate, sessile, about 
as broad as long, white or yellowish; bracts and bractlets half as long 
as the sepals, broadly ovate, acuminate to a rigid, spinose tip, usu- 
ally sparsely pilose; sepals lance-oblong to ovate, 3-3.5 mm. long, 
acuminate, mucronate, 3-nerved, short-pilose; staminodia ligulate, 
equaling or shorter than the filaments, laciniate at the apex; utricle 
suborbicular. 

Lima: Callao, Wilkes Exped. Loreto: Iquitos, in pasture, 
Williams 3593; waste places, Killip & Smith 27498. Mishuyacu, 
in clearing, Klug 1209, 1049. Piura: Between Piura and Nomala, 
100-250 meters, Weberbauer 5957. General through tropical America. 

Alternanthera flavida Suesseng. Repert. Sp. Nov. 35: 300. 
1934. A. asterophora Standl. Field Mus. Bot. 11: 150. 1936. 

A shrub 2 meters high, the branches densely stellate- tomentose; 
leaves on rather long petioles, the blades 3-6.5 cm. long, rounded- 
ovate to broadly elliptic, rounded to acutish at the apex, acute to 
truncate at the base, tomentose on both surfaces, more densely so 
beneath, with closely appressed, yellowish, stellate or hispidulous 
hairs; heads stramineous, 12-15 mm. wide, globose or short-oblong, 
paniculate, the peduncles trifid; bracts broadly ovate, short, obtuse 
or acute, pilose; sepals pilose, at least below, 5-6 mm. long, obtuse. 

Huancavelica: Valley of Rio Mantaro below Colcabamba, 1,700 
meters, Weberbauer 6464, type (also type of A. asterophora}. 

Alternanthera halimifolia (Lam.) Standl. ex Pittier, PI. 
Usual. Venez. 145. 1926. Achyranthes halimifolia Lam. Encycl. 1: 
547. 1785. Illecebrum frutescens L'He>. Stirp. Nov. 75. 1788. /. 
alsinaefolium Scop. Delic. Insub. 3: 27. 1788. /. limense Dum. 
Cours. Bot. Cult. 1: 646. 1802. Alternanthera truxillensis HBK. 
Nov. Gen. & Sp. 2: 206. 1817. Telanthera frutescens Moq. in DC. 
Prodr. 13, pt. 2: 365. 1849. Celosia peruviana Van Spand. ex Moq. 
loc. cit. in syn. Achyranthes incana Moq. loc. cit. in syn. T. frutes- 
cens var. acutifolia Moq. op. cit. 366. 1849. T. frutescens var. 
manillensis Moq. loc. cit. T. densiflora Moq. op. cit. 366. 1849. T. 
truxillensis Moq. op. cit. 367. 1849. Alternanthera ficoidea var. 
halimifolia Kuntze, Rev. Gen. 2: 539. 1891. T. halimifolia A. Stewart, 
Proc. Calif. Acad. IV. 1: 58. 1911. 



500 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A much branched perennial, prostrate or procumbent, the stems 
stellate-puberulent and with spreading, hispidulous hairs, in age 
sometimes glabrate; leaves rather thin, short-petiolate, the blades 
oblong to oval or obovate-oblong, 1.5-6 cm. long or larger, rounded 
to acutish at the apex, when young densely stellate-pubescent and 
often pilose with spreading, hispidulous hairs, in age sometimes 
glabrate; heads chiefly axillary, whitish, sessile, globose, solitary or 
glomerate, the flowers stramineous or whitish; bracts and bractlets 
half as long as the sepals, ovate, acuminate, mucronate, appressed- 
pilose, the tips not appressed; sepals 3-4 mm. long, ovate-oblong, 
acute, 3-5-nerved, densely pubescent with short, hispidulous hairs; 
staminodia ligulate, longer than the filaments, ligulate at the apex. 
Neg. 3230. 

Huanuco: Huanuco, 2,100 meters, in hedge, 2319, 3540; Ruiz 
& Pavon 5-66. Libertad: Trujillo, Humboldt & Bonpland, type 
of A. truxillensis (photo, ex hb. Berol.). Salaverry, sandy slopes, 
Johnston 3521. Lima: Type of Achyranthes halimifolia from Lima. 
Rio Chillon, near Viscas, 1,900 meters, open, rocky slopes, Pennell 
14456. Lima, Huanuco, and Chancai, Ruiz & Pavon 5-66. Lurin, 
moist, open soil, Pennell 12210. Lima, Cook & Gilbert 2070. Chosica, 
900 meters, dry gulch, 2864- San Lorenzo Island, Andersson. Callao, 
slopes of seaside gulches, 5882; Wilkes Exped. Obrajillo, Wilkes 
Exped. Santa Clara, Rose 18567, 18775. Piura: Serran, 250 meters, 
Weberbauer 5977. Talara, Haught 54. Ten miles east of Talara, 
abundant on low hills near the sea, Haught F99. Paita, Rose 18512; 
shale cliff above the sea, Pennell 14817. Chile, Ecuador, Jamaica, 
and probably elsewhere. "Yerba blanca," "sanguinaria." 

The plants are somewhat variable, and the species is too closely 
related to A.ficoidea, there being no sharp differences between the 
two. In some plants the leaves are very densely and finely silvery- 
pubescent, even in age, while in others the leaves are soon green 
and glabrate. 

Alternanthera lanceolata (Benth.) Schinz, Nat. Pflanzenfam. 
ed. 2. 16c: 75. 1934. Brandesia lanceolata Benth. PI. Hartweg. 247. 
1839. Telanthera lanceolata Moq. in DC. Prodr. 13, pt. 2: 371. 1849. 
Mogiphanes soratensis Rusby, Bull. N. Y. Bot. Gard. 6: 503. 1910. 
A. mexicana var. gracilis Suesseng. Repert. Sp. Nov. 35: 301. 1934. 

Plants annual or perennial, slender, erect or more often pro- 
cumbent and rooting at the nodes, or elongate and straggling, 
much branched, the stems pilose with rather long, pale, appressed 
or spreading hairs, often glabrate in age; leaves slender-petiolate, the 



FLORA OF PERU 501 

petioles short or elongate, the blades usually thin, narrowly lanceo- 
late to lance-ovate, oblong-elliptic, or even somewhat obovate, acute 
or often long-acuminate, usually attenuate at the base, pilose with 
long, appressed or spreading, weak hairs, rarely glabrate; peduncles 
slender, densely pilose, axillary or terminal, usually simple, rarely 
trifid; heads short-oblong, whitish or greenish white, 6-8 mm. wide, 
rarely more than 1 cm. long; bractlets glabrous, much shorter than 
the flowers (or in bud sometimes projecting beyond them), abruptly 
contracted into a long, stiff, spine-like mucro, this often recurved; 
sepals glabrous, rigid, 2.5-3 mm. long, acute, obscurely 3-nerved; 
staminodia longer than the filaments, 4-6-fid. 

Ayacucho: Carrapa, 1,500 meters, dense forest, Killip & Smith 
22351. Cuzco: Quillabamba, Soukup 187. Rio Yanamayo, 2,300 
meters, bank in forest near stream, Pennell 14037. Huanuco: 
Muna, 2,100 meters, 3996 (type of A. mexicana var. gracilis), 4013. 
Cani, 2,550 meters, in clearing, 3441- Libertad: Cushi, 1,500 meters, 
4820. Bolivia to Colombia and Venezuela. 

Alternanthera Lehmannii Hieron. Bot. Jahrb. 20: Beibl. 49: 
8. 1895. Achyranthes Lehmannii Standl. Journ. Wash. Acad. Sci. 5: 
74. 1915. 

Plants slender, much branched, probably supported on other 
plants, weak, the stems pilose with rather long, chiefly spreading 
hairs; petioles slender, short or elongate; leaf blades lance-oblong 
to oblong-elliptic, thin, abruptly acuminate or rarely obtuse, acute 
or attenuate at the base, copiously pilose with usually widely spread- 
ing hairs; peduncles chiefly axillary, long and slender, simple; heads 
white or stramineous, globose or short-oblong, 1 cm. long or less; 
bractlets whitish, ovate, very long-aristate, shorter than the sepals, 
glabrous; sepals 2.5-3 mm. long, oblong, acute, obscurely 3-nerved, 
copiously short-pilose; staminodia longer than the filaments, lacin- 
iate-dentate at the apex. Neg. 3219. 

San Martin: San Roque, in forest, Williams 7403. Ranging to 
Colombia. "Picurullum-sisa." 

The determination of the single Peruvian collection is somewhat 
doubtful. A. Lehmannii probably is only a variety of A. lanceolata. 
By Suessenguth it is reduced, apparently, to synonymy under A. 
mexicana (Moq.) Hieron. 

Alternanthera lupulina HBK. Nov. Gen. & Sp. 2: 206. 1817. 
Bucholzia lupulina Mart. Beitr. Amarant. 108. 1825. Illecebrum 



502 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

lupulinum Spreng. Syst. Veg. 1: 820. 1825. Telanthera andicola 
Moq. in DC. Prodr. 13, pt. 2: 373. 1849. Achyranthes parviflora 
Pavon ex Moq. loc. cit. in syn. T. lupulina Moq. op. cit. 374. 1849, 
excluding description. Achyranthes lupulina Willd. ex Moq. loc. cit. 
in syn. 

Plants perennial, densely branched, the stems glabrate, suf- 
frutescent, forming dense mats 10 cm. broad, densely leafy; leaves 
petiolate, spatulate, obtuse, glabrous, 5 mm. long or less; heads 
small, white, naked, on short, slender, glabrate, terminal peduncles 
slightly surpassing the leaves, subglobose; bracts about equaling the 
calyx, ovate, acutish, glabrous; sepals 2.5 mm. long, lance-oblong, 
obtuse, glabrous, scarious, obscurely 3-nerved; staminodia equaling 
the filaments, slightly dilated at the apex and laciniate-dentate. 
Negs. 7385, 3221. 

Ancash: Mountains near Conchucos, 4,500 meters, Weberbauer 
7246. Cajamarca: Near Cajamarca, Humboldt & Bonpland, type 
(photo, seen ex hb. Berol.). Junin: Morococha, 4,500 meters, steep, 
short grass slope, 880. La Oroya, 3,300-3,900 meters, Kalenborn 
14. Without definite locality: Andes, Ruiz & Pavdn. High Andes 
of Peru, Wilkes Exped. Moquin reports specimens collected by 
Pavon, MacLean, and Mathews. 

The species is a high alpine plant, growing at a higher elevation 
than any other Andean species. It is different from other Peruvian 
species in its small size and dense, cespitose habit. The plant 
Moquin described as Telanthera lupulina is Alternanthera Macbridei. 

Alternanthera Macbridei Standl. Field Mus. Bot. 11: 151. 
1936. Telanthera lupulina Moq. in DC. Prodr. 13, pt. 2: 374. 1849, 
as to description and as to synonymy in part, not A. lupulina HBK. 
Achyranthes lactea Moq. loc. cit. in syn. 

Plants perennial from a thick root, the stems 60 cm. long or less, 
simple or much branched from the base, erect or decumbent, sparsely 
pilose or glabrate; leaves short-petiolate, the blades small, oblong- 
spatulate to elliptic or obovate, acute or obtuse, sparsely pilose or 
glabrate; flower heads white, globose or short-oblong, very obtuse, 
sessile in clusters of 3 at the ends of the branches, each cluster sub- 
tended by 2 leaves, these with a conspicuous central red blotch; 
bracts and bractlets shorter than the sepals, acute and aristate- 
mucronate, glabrous; sepals linear-oblong, acute or obtuse, 3 mm. 
long, glabrous; staminodia longer than the filaments, ligulate, 
slightly dilated and subtrifid at the apex. Neg. 3218. 



FLORA OF PERU 503 

Huanuco: Chinchao, Ruiz & Pavdn (photo, ex hb. Berol). 
Cheuchin, Ruiz & Pavdn. Junin: La Oroya, 3,600 meters, base of 
limestone cliff, 967, type; Rose 18704; Kalenborn 121. La Quinua, 
3,600 meters, steep, grassy slope, 2027. Near Huancayo, 3,400 
meters, open, rocky hillside, Killip & Smith 23355. Lima: Rio 
Blanco, 3,000-3,500 meters, open hillside, Killip & Smith 21708.- 
Cuzco: Cuzco, Soukup 200. 

A well marked species, easily recognized by the red-blotched 
leaves subtending the cluster of flower heads. 

Alternanthera microcephala (Moq.) Schinz, Nat. Pflanzen- 
fam. ed. 2. 16c: 75. 1936. Brandesia mexicana Schlecht. Linnaea 7: 
392. 1832. Telanthera microcephala Moq. in DC. Prodr. 13, pt. 2: 
371. 1849. T. mexicana Moq. op. cit. 372. 1849. A. mexicana 
Hieron. Bot. Jahrb. 20: Beibl. 49: 8. 1895, non Moq. 1849. 

Plants herbaceous, erect or ascending, often reclining, much 
branched, weak, slender, the stems pilose with long, white, chiefly 
spreading or sometimes retrorse hairs; petioles short or elongate, 
the blades thin, lance-oblong to oval-ovate, 3-10 cm. long, rather 
abruptly long-acuminate, obtuse to long-attenuate at the base, 
pilose on both surfaces; peduncles chiefly axillary and simple, 
elongate, very slender; spikes subglobose or short-cylindric, 5-7 
mm. thick, about 1 cm. long, white; bractlets broadly ovate, half as 
long as the calyx, long-aristate, villous on the costa; sepals narrowly 
oblong, 2.5-3.5 mm. long, acute or acutish, 3-nerved, glabrous; stam- 
inodia longer than the anthers, ligulate, laciniate at the apex. 

Ayacucho: Aina, 750-1,000 meters, in thicket, Killip & Smith 
22792. Cuzco: San Miguel, Urubamba Valley, 1,800 meters, Cook 
& Gilbert 904. Huanuco: Mito, 2,700 meters, shady thicket, 3274 
Junin: Enenas, 1,600-1,900 meters, dense forest, Killip & Smith 
25741. Carpapata, 2,700-3,200 meters, edge of forest, Killip & 
Smith 24440. Bolivia to Colombia and Mexico. 

It is doubtful whether this is more than a form of Alternanthera 
lanceolata. 

Alternanthera paniculata HBK. Nov. Gen. & Sp. 2: 208. 
1817. Illecebrum paniculatum Spreng. Syst. Veg. 1: 819. 1825. Tel- 
anthera paniculata Moq. in DC. Prodr. 13, pt. 2: 377. 1849. T. 
luzuloides Moq. op. cit. 378. 1849. T. Meyeriana Regel & Koern. 
Ind. Sem. Hort. Petrop. 24. 1856. T. Bangii Rusby, Mem. Torrey 
Club 6: 110. 1896. Mogiphanes paniculata Rusby, Bull. N. Y. Bot. 



504 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Gard. 6: 503. 1910. Achyranthes Bangii Standl. Journ. Wash. Acad. 
Sci. 5: 74. 1915. 

Plants large and much branched, suberect or more often scandent, 
sometimes 3 meters long or more, the slender branches usually softly 
pilose with short or long, spreading hairs, sometimes appressed- 
pilose; leaves slender-petiolate, green, rather thin, ovate to elliptic 
or lance-oblong, acute or acuminate, acute or obtuse at the base, 
usually densely appressed-pilose, especially beneath, rarely some- 
what glabrate; flower heads white or pink to purple, naked at the 
base, forming large, open panicles, usually sessile in clusters of 3, 
sometimes solitary and long-pedunculate, the clusters long-pedun- 
culate, the heads globose or short-oblong; bracts and bractlets much 
shorter than the calyx, lanceolate or ovate, glabrous or pubescent, 
mucronate; sepals 3-4 mm. long, glabrous, linear-oblong, acute, 
3-nerved; staminodia longer than the filaments, laciniate at the 
apex. Negs. 3222, 3224. 

Huanuco: Yanano, 1,800 meters, in thicket, 3733, 4934. Mito, 
2,700 meters, sprawling on stone wall, 1536. Lima: Matucana, 2,400 
meters, among rocks on steep canyon slope, 260. Rio Chillon above 
Obrajillo, 3,300 meters, on rocks about cascade, Pennell 14700. 
Rio Blanco, 3,000-3,500 meters, open hillside, Kittip & Smith 21658. 
Bolivia to Colombia. 

The Peruvian material referred here varies conspicuously in 
the size of the flowers, and it is altogether possible that it includes 
two species. The flowers usually are white, but occasionally pink 
or purplish. 

Alternanthera peruviana (Moq.) Suesseng. Repert. Sp. Nov. 
35: 302. 1934. Telanthera peruviana Moq. op. cit. 366. 1849. A. 
peruviana f. globifera Suesseng. op. cit. 303. 

Plants perennial, much branched, procumbent or prostrate, the 
branches often elongate, densely silvery-strigose; leaves sessile or 
short-petiolate, thick, oblong to obovate, mostly 1.5-4.5 cm. long, 
rounded to acutish at the apex, narrowed to the base, when young 
densely strigose or silvery-sericeous on both surfaces, in age glabrate 
or the pubescence more often persistent; flower heads axillary, sessile, 
solitary or glomerate, short-oblong or sometimes as much as 2.5 
cm. long, whitish or stramineous; bracts ovate, acutish, short- 
villous, the tips appressed or even incurved ; sepals somewhat indurate, 
ovate to oblong, acute, the outer sometimes glabrous, the inner 
short- villous; staminodia equaling the filaments, ligulate, laciniate 
at the apex. 



FLORA OF PERU 505 

Type of Telanthera peruviana collected somewhere in Peru, 
Cuming 971. Piura: Talara, common on sandhills and sandy ground, 
forming tangled masses of branches, H aught 32; open, arid cliff, 
Pennell 12203. Tumbez: Between Zorritos and Cancas, sandy 
seashore plain, Weberbauer 7749. Also in Ecuador. 

The type of f . globifera was collected at Paita by Ball. 

Alternanthera philoxeroides (Mart.) Griseb. Abh. Ges. Wiss. 
Goett. 24: 36. 1879. Bucholzia philoxeroides Mart. Nov. Act. Acad. 
Leop. Carol. 13, pt. 1: 315. 1826. Telanthera philoxeroides Moq. in 
DC. Prodr. 13, pt. 2: 362. 1849. Achyranthes philoxeroides Standl. 
Journ. Wash. Acad. Sci. 5: 74. 1915. 

A herbaceous perennial, the stout stems often fistulous, ascend- 
ing or decumbent, simple or branched, 1 meter long or less, villous 
in the leaf axils, elsewhere glabrous or sparsely villous; leaves short- 
petiolate or subsessile, elliptic-linear to obovate, 4-12 cm. long, 
acute or obtuse, rather thick, glabrous; peduncles simple, axillary 
or terminal, 1-5 cm. long; flower spikes globose, 14-17 mm. thick, 
white; bracts one-fourth as long as the sepals, acute or acumi- 
nate, glabrous; sepals ovate-oblong, 6 mm. long, acute or acutish, 
obscurely nerved ; staminodia ligulate, exceeding the anthers, lacerate 
at the apex. 

Loreto: Iquitos, wet soil, Williams 1466, 3592. Rio Itaya, 
Killip & Smith 29403. Argentina to the Guianas and Colombia; 
Mexico and Central America. 

The plant grows commonly in very wet soil, and often in water. 

Alternanthera piurensis Standl. Field Mus. Bot. 11: 152. 1936. 

Plants much branched, probably subscandent, the stems glabrous 
or when young sparsely pilose, ferruginous; leaves slender-petiolate, 
the blades ovate, acuminate or cuspidate-acuminate, 1.5-4 cm. long, 
obtuse or rounded at the base and decurrent, sparsely pilose with 
lax, spreading hairs; flower heads terminal and axillary, usually in 
clusters of 3, all sessile or nearly so, the compound heads appearing 
like simple heads, usually bracted at the base with reduced leaves; 
bracts glabrous or pilose on the costa; sepals glabrous, linear-oblong, 
5 mm. long, acute or acuminate, the nerves very obscure. 

Piura: Below Ayavaca, 2,600 meters, Weberbauer 6358, type. 

Alternanthera porrigens (Jacq.) Kuntze, Rev. Gen. 538. 1891. 
Achyranthes porrigens Jacq. Hort. Schoenbr. 3 : pi. 350. 1798. Celosia 
elongata Spreng. in Schrad. Journ. 2: 196. 1800. C. peruviana Zucc. 



506 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

in Roem. Coll. Bot. 133. 1809. Alternanthera sericea HBK. Nov. 
Gen. & Sp. 2: 207. 1817. Alternanthera gomphrenoides HBK. op. 
cit. 2: 208. 1817. Illecebrum gomphrenoides Willd. ex R. & S. Syst. 
Veg. 5: 515. 1819. Brandesia porrigens Mart. Beitr. Amarant. 106. 
1825. I. sericeum Spreng. Syst. Veg. 1 : 819. 1825. Telanthera por- 
rigens Moq. in DC. Prodr. 13, pt. 2: 377. 1849. Achyranthes pur- 
purea Pavon ex Moq. loc. cit. in syn. T. gomphrenoides Moq. loc. cit. 
Gomphrena sericea Moq. op. cit. 385. 1849, not Spreng. T. Riveti 
Dang. & Cherm. Bull. Mus. Hist. Nat. Paris 28: 438. 1922. 

Plants suffrutesceht, large, much branched, rather stout, erect 
or often scandent, the stems densely pilose with appressed, white 
hairs, rarely glabrate in age; leaves short-petiolate, the blades ovate 
to elliptic or lance-oblong, acute or acuminate, commonly densely 
appressed-pilose on both surfaces, more conspicuously so beneath; 
flower heads small, numerous, forming a large panicle, purple or 
pink, or rarely almost white, mostly sessile in clusters of 3 at the 
tips of the branches, sometimes solitary on long, axillary peduncles, 
the clusters not bracted; bracts half as long as the calyx, ovate, 
mucronate, usually pubescent over the whole surface; sepals densely 
pilose, or sometimes glabrate above, usually long-pilose at the base, 
2.5-3 mm. long, lance-oblong, acute, 1-nerved or obscurely 3-nerved; 
staminodia equaling the filaments, laciniate at the apex. Negs. 
3225, 3216, 3204. 

Huanuco: Huanuco, 2,100 meters, river cliffs, 3496; Ruiz, type 
of Achyranthes purpurea Ruiz (photo, ex hb. Berol.); Kanehira 266; 
Ruiz & Pavdn 5-63. Taruca, Sawada 51. Pampayacu, Sawada 
P54- Uspachaca, 2,550 meters, dry slope, 1315. Junin: Tarma, 
2,100 meters, heavy, stony soil of steep slope, 1004; Killip & Smith 
21794- Lima: Lima to Obrajillo, Wilkes Exped. Without locality, 
Weberbauer 5529. Moquin reports Mathews 508. Chile to Colombia. 

The material referred here shows less variation than that placed 
under other species. The type of Achyranthes porrigens is Peruvian. 

Alternanthera pubiflora (Benth.) Kuntze, Rev. Gen. 538. 
1891. Brandesia pubiflora Benth. Bot. Voy. Sulph. 157. 1844. 
Telanthera pubiflora Moq. in DC. Prodr. 13, pt. 2: 375. 1849. T. 
pubiflora var. glomerata Moq. op. cit. 376. 1849. T. glomerata Moq. 
loc. cit. in syn. A. albo-squarrosa Suesseng. Repert. Sp. Nov. 39: 
1. 1935. 

Plants perennial, slender or stout, sometimes suberect but more 
often scandent, much branched, the stems sparsely or often very 



FLORA OF PERU 507 

densely pilose with white, chiefly appressed hairs; leaves short- 
petiolate, the blades ovate or ovate-elliptic to lance-oblong, sparsely 
or very densely pilose with weak, subappressed, white hairs; flower 
heads white, globose or short-oblong, laxly paniculate, usually in 
clusters of 3 sessile heads at the ends of the branches, the lateral 
heads pedunculate, naked or the clusters often leafy-bracted at 
the base; bractlets much shorter than the calyx, acute or mucro- 
nate, pilose; sepals 5-8 mm. long, linear-lanceolate, densely pilose with 
spreading hairs, acuminate, 3-nerved, the tips often somewhat 
recurved; staminodia longer than the filaments, laciniate at the 
apex. Negs. 3229, 3217, 3226, 27880. 

Arequipa: Tingo, 2,200 meters, open, rocky slopes, Pennell 
13131; Rose 18800. Arequipa, Isern 2506. Mollendo, sandy hills, 
Hitchcock 22372. Ayacucho: Plateau between Rio de Lomas and 
Rio Yauca, 1,900 meters, Weberbauer 5759, type of A. albo-squarrosa. 
Junin: Below Surco, 1,900 meters, Weberbauer 5205. Lima: 
Near Lima, Cuming 1003 ex Moq. Chosica, Rose 18544- South of 
Santa Clara, Rose 18622. Posco, Rose 18803. Piura: Between 
Piura and Nomala, 100-250 meters, Weberbauer 5930. Somate, 90 
meters, Townsend 821 . Bolivia and Chile to Colombia. 

The Peruvian specimens are rather more uniform in pubescence 
than material from other parts of South America, but they exhibit 
some variation in size of flowers. A. albo-squarrosa may be distinct, 
having larger flowers than some of the other specimens, but there 
appear to be intergrading forms. 

Alternanthera pulverulenta (Mart.) Moq. in DC. Prodr. 13, 
pt. 2: 351. 1849. Trommsdorffia pulverulenta Mart. Nov. Gen. & Sp. 
2: 42. 1826. Illecebrum pulverulentum Spreng. Syst. Veg. Cur. Post. 
103. 1827. Iresine latifolia D. Dietr. Syn. PL 1: 870. 1839. 

Type collected in Peru by Haenke. Moquin places the plant in 
a group now usually referred to the genus Iresine. No plant of that 
particular alliance is known from Peru. I suspect that A. pulveru- 
lenta is really a Pfaffia, a genus that can easily be confused with 
Iresine argentata and its relatives. The description does not agree, 
however, with the Pfaffia species represented by recent Peruvian 
material. 

Alternanthera pungens HBK. Nov. Gen. & Sp. 2: 206. 1817. 
Celosia echinata Humb. & Bonpl. ex R. & S. Syst. Veg. 5: 531. 
1819. Alternanthera echinata Smith in Rees, Cycl. Suppl. No. 
10. 1802-1820. Telanthera pungens Moq. in DC. Prodr. 13, pt. 2: 



508 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

371. 1849. Alternanthera Achyrantha var. leiantha Seub. in Mart. 
Fl. Bras. 5, pt. 1: 183. pi 55. 1875. 

Plants annual or perennial, the stems prostrate, much branched, 
villous with obscurely scaberulous, white hairs; leaves on short, 
marginate petioles, green when dried, rather thin, the blades orbic- 
ular or rhombic, 1.5-5 cm. long, rounded and apiculate at the apex, 
appressed-pilose when young, soon glabrate; heads stramineous, 
globose or oblong, axillary, sessile, usually solitary; bracts and 
bractlets lance-oblong, equaling the perianth, attenuate to an aristate 
apex, glabrous; sepals oblong, 5-6 mm. long, acuminate to a long, 
rigid tip, 3-nerved, sparsely villous near the base; staminodia tri- 
angular, remotely dentate; utricle truncate. 

Huanuco: Huanuco, Ruiz & Pavdn 5-70. Ambo, 2,100 meters, 
prostrate on stony river flat, 3195. Bolivia and Argentina to Mexico 
and the West Indies. 

Called "yerba de polio" in Argentina and Uruguay. 

Alternanthera repens (L.) Kuntze, Rev. Gen. 536. 1891. 
Achyranthes repens L. Sp. PI. 205. 1753. Illecebrum Achyrantha L. 
Sp. PI. ed. 2. 299. 1762. Alternanthera Achyrantha R. Br. Prodr. 
417. 1810. Alternanthera caracasana HBK. Nov. Gen. & Sp. 2: 205. 
1817. Celosia humifusa Willd. ex R. & S. Syst. Veg. 5: 531. 1819. 
Alternanthera Achyrantha var. parvifolia Moq. in DC. Prodr. 13, pt. 
2: 359. 1848. Telanthera caracasana Moq. op. cit. 370. 1848. Alter- 
nanthera parvifolia Fawc. & Rendle, Fl. Jam. 3: 139. 1914. 

Perennial from a thick, woody, vertical root, the prostrate or 
procumbent, branched stems forming dense mats, white-villous or 
glabrate; leaves petiolate, the blades rhombic-ovate to obovate, 
1-2.5 cm. long, obtuse, sparsely villous when young, soon glabrate; 
heads ovoid or short-cylindric, 5-8 mm. thick, sessile and usually 
glomerate in the leaf axils, pale yellowish; bracts and bractlets 
shorter than the sepals, ovate, mucronate-pungent, glabrous or 
pilose; sepals very unequal, the outer oval or broadly ovate, 3-5 
mm. long, acutish, 3-nerved, villous along the nerves with articulate, 
scabrous hairs; staminodia usually shorter than the filaments, 
triangular or subulate, entire or rarely denticulate. Neg. 3232. 

Cuzco: Below Cuzco, Rose 19071. Vilcanota Valley, 2,900 
meters, Herrera 1071 a. Lucumayo Valley, Cook & Gilbert 1349. 
Junin: Tarma, 2,100 meters, heavy, stony soil, 1003; open hillside, 
Killip & Smith 21860. Huancayo, 3,400 meters, in ravine, Ledig 41- 
Lima: Matucana, 2,400 meters, shale slope along trail, 203. 



FLORA OF PERU 509 

Generally distributed in tropics of both hemispheres. "Yerba 
del moro." 

The plant's favorite habitat in most regions of tropical America 
seems to be among cobblestones of village streets. 

Alternanthera tomentosa (Moq.) Schinz, Nat. Pflanzenfam. 
ed. 2. 16c: 76. 1934. Telanthera tomentosa Moq. in DC. Prodr. 13, 
pt. 2:380. 1849. 

Plants suberect, suffrutescent, stout, 1 meter high or less, the 
branches densely villous-tomentose with fulvous or reddish hairs; 
leaves very shortly petiolate, the blades ovate or broadly elliptic to 
rounded-ovate, mostly 4-6 cm. long, acute to rounded at the apex, 
mucronate, rounded to truncate or subcordate at the base, rather 
thick, densely pilose above, densely villous-tomentose beneath with 
tawny hairs, the nerves conspicuous beneath; peduncles simple or 
trifid, short or elongate; flower heads globose, stramineous or whitish, 
about 12 mm. broad; bractlets lance-oblong, slightly shorter than the 
sepals, acute and apiculate, sparsely pilose or glabrate; sepals pilose, 
6 mm. long, thick, 3-nerved, acute; staminodia longer than the fila- 
ments, laciniate-dentate at the apex. Neg. 27881. 

Amazonas: Type from Chachapoyas, Mathews 3130. Cajamarca: 
Montana de Nancho, 2,310 meters, Raimondi 6172; at 2,460 meters, 
Raimondi3164, 5247, 4593. Between Chota and Ninabamba, Raimondi 
9987 (Raimondi collections det. by Suessenguth). Huanuco: Ambo, 
river canyon slopes, 2,100 meters, 3155. Also in Bolivia. 

I have seen no authentic material of this species. The collection 
from Huanuco agrees fairly well with the original description, and 
it is the only collection examined that could be referred to A. 
tomentosa, as described by Moquin. 

Alternanthera villosa HBK. Nov. Gen. & Sp. 2: 208. 1817. 
Illecebrum villosum Willd. ex R. & S. Syst. Veg. 5: 516. 1819. Telan- 
thera villosa Moq. in DC. Prodr. 13, pt. 2: 379. 1849. 

Plants herbaceous or suffrutescent, more or less scandent, densely 
tawny-hirsute throughout with long, spreading hairs; leaves short- 
petiolate, the blades ovate or broadly ovate, 2.5-6.5 cm. long, short- 
acuminate, rounded to acutish at the base; peduncles simple, 
terminal and axillary, elongate; heads globose, stramineous, 1.5 cm. 
broad; bracts unequal, somewhat shorter than the sepals, more or 
less pilose, obscurely cristulate and dentate near the apex; sepals 
6-7 mm. long, attenuate to an acute apex, thinly pilose; staminodia 
ligulate, slightly longer than the filaments, laciniate at the apex. 



510 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Piura: Type collected near Ayavaca, Humboldt & Bonpland. 
Ayavaca, 2,700 meters, Weberbauer 6366. Also in Ecuador. 

Easily recognized by the abundant pubescence of very long, 
tawny hairs. 

10. GOMPHRENA L. 

Annual or perennial herbs, rarely suffrutescent; leaves opposite; 
flowers perfect, bracteate and bibracteolate, spicate or capitate, 
the heads solitary or glomerate, terminal or axillary, naked or sub- 
tended by leaves, white, yellow, or red; bractlets carinate, often 
winged or cristate dorsally; perianth sessile, terete or compressed, 
5-lobate or 5-parted, usually lanate at the base; stamens 5, the fila- 
ments united to form a tube, this 5-lobate at the apex, the lobes 
bifid or emarginate, the anthers sessile or stipitate in the sinuses; 
stigmas 2 or rarely 3, subulate or filiform, on a short or elongate 
style; ovary 1-ovulate. 
Stems scapose, naked; a cespitose perennial from a thick root, the 

leaves and scapes arising from its summit G. Meyeniana. 

Stems leafy and usually branched. 

Bracts and bractlets very obtuse or rounded at the apex; plants 
small, the stems less than 8 cm. long. 

Plants perennial; stems densely pilose G. oroyana. 

Plants annual; stems glabrous G. umbellata. 

Bracts and bractlets acute or acuminate; plants normally much 
larger. 

Flower heads not subtended by leaves, naked G. elegans. 

Flower heads subtended by large or small leaves. 
Sepals not exceeding the bractlets. 

Heads 2-2.5 cm. broad G. globosa. 

Heads 1 cm. broad G. decumbens. 

Sepals longer than the bractlets G. pulchella. 

Gomphrena decumbens Jacq. Hort. Schoenbr. 4: 41. pi. 482. 
1804. ?G. bicolor Mart. Beitr. Amarant. 92. 1826. Xeraea decum- 
bens Kuntze, Rev. Gen. 2: 545. 1891. G. ixiamensis Rusby, Bull. 
N. Y. Bot. Card. 6: 502. 1910. 

A prostrate or decumbent annual, rarely erect, usually much 
branched, the stems appressed-pilose; leaves short-petiolate, the 
blades obovate to oblong or oval, rounded or obtuse at the apex, 
acute to attenuate at the base, appressed-pilose; heads subglo- 
bose, white or pinkish, 1 cm. broad, usually solitary and pedunculate, 



FLORA OF PERU 511 

subtended normally by 2 leaves, these shorter or longer than the 
heads; bractlets long-acuminate, narrowly cristate at the apex; 
perianth shorter than the bractlets, densely lanate, the sepals oblong- 
linear, long-attenuate. 

The species is widely distributed in tropical America, from Bolivia 
to Uruguay and northward to Mexico. It is included for Peru only 
because G. bicolor, whose type was collected in Peru by Haenke, 
appears, from description, to be a probable synonym. Gomphrena 
decumbens has been collected in Bolivia, and may well be expected 
in Peru. 

Gomphrena elegans Mart. Nov. Gen. 2: 17. pi. 119. 1826. G. 
perennis L. f. villosa (Mart.) Stuchl. subf. boliviana Stuchl. Repert. 
Sp. Nov. 11: 154. 1913. G. elegans var. brunnea and var. Mando- 
nioides Suesseng. Repert. Sp. Nov. 35: 310. 1934. 

A perennial herb 60 cm. high or less, sometimes taller, branched, 
the branches densely pilose at first, glabrate; leaves slender-petiolate, 
the blades ovate-oblong to rounded-ovate, 3-7 cm. long, acuminate 
to acutish, at the base obtuse or acutish, sparsely or densely ap- 
pressed-pilose above with long, slender hairs, beneath usually whitish 
and densely pilose as well as more or less tomentose, in age rarely 
glabrate; heads white or whitish, subglobose, 1-1.5 cm. broad, 
chiefly terminal and long-pedunculate, naked at the base; bracts 
and bractlets rather densely pilose, acute or acuminate; sepals 
linear-lanceolate, 5 mm. long, acute, lanate below, hirtellous above; 
staminodia linear- triangular, acutish. 

Ayacucho: Ayacucho, 3,100 meters, Weberbauer 5529, type of 
var. brunnea. Cuzco: Ollantaitambo, 3,000 meters, Cook & Gilbert 
289; open, rocky slope, Pennell 13667. Without locality, Herrera 
2587. Huasao, 3,200 meters, Herrera 3035. Gucay, Urubamba 
Valley, 2,900 meters, Herrera 1133. Urubamba Valley, 2,760 
meters, Herrera 2109. Cuzco, 3,000-3,600 meters, Herrera in 1923. 
Huancavelica: Below Colcabamba, 2,200 meters, Weberbauer 644$, 
type of var. Mandonioides. Junin: Huancayo, 3,400 meters, rocky 
cliff, Ledig 38. Huacapistana, 1,800-2,400 meters, open hillside, 
Killip & Smith 24193. Huancayo, 3,400 meters, rocky cliff, Ledig 
38; Killip & Smith 22017. Loreto: Leticia, edge of forest, Williams 
3056. Rio Nanay, in forest, Williams 547. Bolivia. "Pieccacissacj." 

The Loreto specimens come from far outside the expected range 
of the species, and their determination is decidedly doubtful, but 
in most characters the plants seem to agree well with those of higher 



512 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

elevations. Some of the Peruvian material is probably referable to 
G. Mandonii Fries, a species perhaps not distinct from G. elegans. 

Gomphrena globosa L. Sp. PI. 224. 1753. 

A stout annual 1 meter high or less, much branched, the stems 
sparsely or densely appressed-pilose; leaves short-petiolate, the 
blades oblong to oval, ovate, or spatulate, acute, appressed-pilose; 
peduncles simple, elongate; heads subtended by usually 2 short 
leaves, globose or short-cylindric, white, red, or yellow; bractlets 
broadly cristate, the crest serrulate; sepals densely lanate, lance- 
subulate, 1-nerved. 

Loreto: Fortaleza, Yurimaguas, in garden, Williams 4489. San 
Salvador, Williams 1 555. Mishuyacu, in clearing, Klug 1 79. Lower 
Rio Nanay, Williams 347. San Martin: Tarapoto, Williams 6750, 
6515. Cultivated generally in tropical and temperate countries. 
"Siempreviva," "manto de Cristo." 

This is the globe amaranth, bachelor's button, or immortelle of 
northern gardens, where it is a favorite ornamental plant. Probably 
it is a native of tropical America, although perhaps unknown in the 
wild state. The plant sometimes escapes from tropical gardens to 
waste ground and roadsides, but seldom persists long. In many 
regions it is employed commonly for making funeral wreaths. 

Gomphrena Meyeniana Walp. Nov. Act. Nat. Cur. 19: Suppl. 
1: 404. 1843. G. acaulis R<my, Ann. Sci. Nat. III. 6: 350. 1846. G. 
Conwayi Rusby, Bull. N. Y. Bot. Card. 8: 89. 1912. 

Plants perennial from a long, thick, vertical root, the scapes and 
leaves clustered at its apex and surrounded by dense, brownish wool ; 
leaves long-petiolate, the blades obovate to oblanceolate or even 
ovate, 1-3 cm. long, obtuse, pilose with long, soft hairs or often 
glabrate; heads white, subtended by several small leaves, globose, 
1-1.5 cm. broad or sometimes slightly larger; bracts and bractlets 
very obtuse, thin and soft; sepals cuneate-spatulate, scarious, cren- 
ulate at the apex. Neg. 3252. 

Arequipa: San Ignacio, near Cailloma, 4,400 meters, Weberbauer 
6886. Puno: Pisaloma (Pisacoma?), 4,500 meters, near Lake Titi- 
caca, Meyen (photo, of type ex hb. Berol.). Tuapata (Puno?), 
3,750 meters, Watkins in 1916. Puno, 4,000 meters, Soukup 99. 
Chuquibambilla, 4,000 meters, rocky, clay slope on puna, Pennell 
13352. Collacache, Stordy in 1920. Chile, Bolivia, and Argentina. 
"Pimpinela." 

An alpine plant, distinguished by dwarf habit and scapose stems. 



FLORA OF PERU 513 

Gomphrena oroyana Standl. Field Mus. Bot. 11: 150. 1936. 

A perennial from a thick, vertical root, the numerous stems 
probably ascending, leafy, 5 cm. long or less, long-pilose; basal leaves 
long-petiolate, obovate, with blades 12 mm. long; cauline leaves on 
short, broad petioles, the blades elliptic-oblong to rounded-ovate, 
acutish or obtuse, 3-6 mm. long, sparsely and laxly white-pilose; 
heads dense, white, naked at the base, 8 mm. broad; bracts and 
bractlets hyaline, glabrous; sepals 3 mm. long, obtuse, densely 
lanate near the base. 

Junin : La Oroya, Kalenborn 89 (type) ; Stevens 6. 

Gomphrena pulchella Mart. Beitr. Amarant. 94. 1826. G. 
Haenkeana Mart. Nov. Act. Nat. Cur. 13: 299. 1826. G. pulchella var. 
rosea Stuchl. Repert. Sp. Nov. 12: 523. 1913. Xeraea Haenkeana 
Kuntze, Rev. Gen. 545. 1891. G. pulchella var. Haenkeana Suesseng. 
Repert. Sp. Nov. 35: 311. 1934. 

Plants annual, erect or ascending, almost simple or much 
branched, usually 30-60 cm. high, the stems pilose with long, chiefly 
appressed, white hairs; leaves short-petiolate, rather thick, oblong 
or lance-oblong, acute at each end, densely appressed-pilose; pe- 
duncles simple, elongate; heads subglobose, 2-3 cm. broad, usually 
pink, subtended by 2-4 leaves, these commonly shorter than the 
flowers; sepals elongate-linear, glabrous above, villous below, erose- 
denticulate at the apex; bractlets long-acuminate, narrowly cristate 
at the apex. Neg. 3247. 

Type of G. Haenkeana collected in Peru by Haenke. Bolivia to 
Argentina and Brazil. 

I have seen a photograph of the type of G. Haenkeana, from the 
Berlin herbarium. Although the specimen is a fragmentary one, it 
seems definitely referable to G. pulchella, in spite of the fact that 
Moquin (in DC. Prodr.) placed it near G. globosa, and compared it 
with that species. If G. pulchella really occurs in Peru, it is rather 
strange that it has not been found there by recent collectors. It 
may well be that the type of G. Haenkeana was obtained in some 
other country. 

Gomphrena umbellata Re"my, Ann. Sci. Nat. III. 6: 349. 1846. 
Xeraea umbellata Kuntze, Rev. Gen. 545. 1891. 

A diminutive annual, glabrous almost throughout, umbellately 
branched, 7.5 cm. high or less; leaves somewhat fleshy, spatulate, 
sometimes as much as 1 cm. long, obtuse, clasping at the base, the 
radical leaves linear-spatulate and petiolate; heads globose, dense, 



514 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

rather few-flowered, pedunculate or sometimes almost sessile; outer 
bracts rounded-obovate, greenish, broadly rounded or subtruncate 
at the apex, ciliate with long, tangled hairs; bractlets white, oblong- 
obovate, thin, scarious, glabrous; sepals 1.5 mm. long, linear or sub- 
spatulate, obtuse or acutish, diaphanous. 

Arequipa: Near Sumbay, 3,900 meters, Weberbauer 6908. Bolivia 
and Argentina. 

11. IRESINE P. Br. 

Plants herbaceous or woody, sometimes scandent, rarely small 
trees; leaves opposite, petiolate; flowers perfect, polygamous, or 
dioecious, bracteate and bibracteolate, capitate or spicate, the spikes 
numerous and paniculate; perianth terete, sessile, the 5 segments 
distinct, usually lanate or pilose; stamens united at the base into a 
short tube, the 5 filaments subulate, entire, the pseudostaminodia 
usually short or wanting; style short or none, the stigmas 2-3, 
subulate or filiform; ovule 1; utricle compressed, membranaceous, 
indehiscent. 
Plants herbaceous. 
Leaves bilobate at the apex, usually colored with purple-red, pink, 

and yellow I. Herbstii. 

Leaves acute or acuminate. 

Leaves densely pilose beneath with very long, lax, weak, spread- 
ing hairs /. Macbridei. 

Leaves with variable pubescence but never as above, more 

often glabrous or nearly so. 

Pubescence of the lower leaf surface and inflorescence in part 
of closely appressed, lustrous, yellow hairs. 

/. spiculigera. 
Pubescence never of appressed, lustrous, yellow hairs. 

/. Celosia. 

Plants fruticose or at least suffrutescent. 
Leaves white or whitish beneath, covered with a very dense and 

close tomentum of branched hairs /. Weberbaueri. 

Leaves green beneath, glabrate, the sparse pubescence of straight, 
simple hairs. 

Pseudostaminodia well developed /. Hassleriana. 

Pseudostaminodia obsolete /. tennis. 

Iresine angustifolia Euphr. (7. elatior Rich.) with lanceolate or 
linear-lanceolate, glabrous leaves and perfect flowers, occurs in both 
Bolivia and Ecuador and is to be expected along the coast of Peru. 



FLORA OF PERU 515 

Iresine Celosia L. Syst. Nat. ed. 10. 1291. 1759. Celosia panicu- 
lata L. Sp. PI. 206. 1753. I. celosioides L. Sp. PL ed. 2. 1456. 1763. 
7. diffusa Humb. & Bonpl. ex Willd. Sp. PI. 4: 765. 1805. 7. poly- 
morpha Mart. Nov. Gen. & Sp. 2: 56. 1826. I . paniculata Kuntze, 
Rev. Gen. 2: 542. 1891, not Poir. 1813. 

Plants essentially annual, but often persisting, sometimes low 
and erect, but frequently more or less scandent, nearly glabrous, the 
stems often villous at the nodes; leaves slender-petiolate, the blades 
chiefly ovate or deltoid-ovate, 5-14 cm. long, acute or acuminate, 
truncate to cuneate at the base, glabrous or sometimes sparsely 
short- villous beneath, very rarely finely pubescent; panicles usually 
broad and lax, sometimes narrow, the spikelets alternate, opposite, 
or verticillate, 5-25 mm. long; flowers usually white, the pistillate 
with copious, long wool at the base; sepals 1-1.5 mm. long, obtuse. 
Negs. 3271, 3272. 

Ayacucho: Estrella, 500 meters, dense woods, Killip & Smith 
23062. Cuzco: Hacienda Chiraura, Prov. Quispicanchi, 3,250 meters, 
Herrera 2610. Huanuco: Yanano, 1,80Q meters, in thicket by 
river, 3S05. Posuso, 600 meters, in thicket or forest, 4631, 4642, 
4686. Vilcabamba, 1,800 meters, open slopes, 4980. Mufia, 2,100 
meters, 4015. Junin: La Merced, 600 meters, edge of forest, 5378, 
5345, 5576; Killip & Smith 23790. Dos de Mayo, 1,800 meters, 
Killip & Smith 25832. Chanchamayo Valley, 1,500 meters, Schunke 
15, 482. Above San Ramon, Schunke A94; Killip & Smith 24732. 
Puerto Bermudez, 375 meters, in thickets, Killip & Smith 26631. 
Loreto: Yurimaguas, in pasture, Williams 4726. Mishuyacu, in 
clearing, Klug 548. Florida, 180 meters, Klug 2351. Alto Rio 
Itaya, Williams 3337. Iquitos, Williams 8251. Piura: Talara, 
Haught 73. Cerro Prieto, 720 meters, HaughtFllO. San Martin: 
Pongo de Cainarachi, Klug 2614- Zepelacio, 1,500 meters, Klug 
3510. Tarapoto, Williams 6696, 5794. Without locality, Weber- 
bauer 6666; Ruiz & Pavdn 1642. General in tropical America. 

The plant is one of the most abundant weedy species of tropical 
America. Usually it is most plentiful in lowlands, but it grows also 
in mountains, often at rather high elevations. In the Peruvian 
material several reasonably distinct forms may be recognized, but 
when all specimens of the species from other regions are considered, 
it appears impossible to separate the Peruvian forms. 

Iresine Hassleriana Chod. Bull. Herb. Boiss. II. 3: 390. 1903. 
7. Hassleriana var. guaranoides Suesseng. Repert. Sp. Nov. 35: 
321. 1934. 



516 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A large, woody vine, the branches appressed-pilose or glabrate; 
leaves oblong to ovate or oblong-elliptic, short-acuminate, rounded 
to subacute at the base, sparsely appressed-pilose beneath, in age 
nearly glabrous; panicles very large and lax, the branches densely 
short-pilose with mostly appressed, ochraceous hairs. Neg. 27889. 

Loreto: Iquitos, Tessmann 3663. Yarina-cocha, 155 meters, Tess- 
mann 5393. Mouth of Rio Morona, Tessmann 3837. Bolivia, Brazil, 
and Paraguay. 

The determinations of the Tessmann collections, which I have 
not seen, are by Suessenguth. 

Iresine Herbstii Hook. Card. Chron. 1864: 654. 1864. Achy- 
ranthes Verschaffeltii Lem. 111. Hort. 11: pi. 409. 1864. /. Ver- 
schaffeltii Lem. 111. Hort. 11: sub pi. 418. 1864. 

An erect annual, short-villosulous at the nodes; leaves long- 
petiolate, the blades rounded, 3-6.5 cm. long, deeply retuse at the 
apex, rounded to truncate at the base and decurrent, mottled or 
colored with purple-red, green, pink, and yellow, often with lustrous, 
appressed hairs on the lower surface; panicles usually ample and 
rather lax, 10-20 cm. long; flowers yellowish, the sepals 1 mm. long. 

Loreto: Pebas, in forest, Williams 1924- Mishuyacu, in clearing, 
100 meters, Klugl21. San Martin: Moyobamba (Mathews). Brazil. 

The plant is much cultivated for ornament in tropical America, 
because of its brightly colored leaves of peculiar form, and it is seen 
sometimes in northern parks or hothouses. It is probably only a 
horticultural form developed from I. spiculigera. 

Iresine Macbridei Standl. Field Mus. Bot. 11: 152. 1936. 

Plants suffrutescent, erect or subscandent, the young stems 
densely and laxly tomentose; leaves on very short petioles, the 
blades ovate or ovate-oblong, 6-8 cm. long, acuminate, obtuse at 
the base, conspicuously pilose on both sides with very long, weak, 
spreading hairs, or in age glabrate; flowers dioecious, in large, yel- 
lowish panicles; sepals 2 mm. long. 

Junin: Huacapistana, 1,800-2,400 meters, open hillside, Killip 
& Smith 24207, type; Macbride 5823. 

Closely related to /. Celosia, but apparently distinct in its 
pubescence. 

Iresine spiculigera Seub. in Mart. Fl. Bras. 5, pt. 1: 228. pi. 
70. 1875. I. acicularis Standl. Contr. U. S. Nat. Herb. 18: 93. 
1916. /. spiculigera f. pauciglandulosa Herzog, Med. Rijks Herb. 46: 
9. 1922. 



FLORA OF PERU 517 

Plants erect or often subscandent, large, the nodes of the branches 
pilose; leaves usually larger than in /. Celosia, thin, lance-ovate to 
broadly ovate, 10-25 cm. long, acuminate, rounded to attenuate at 
the base, more or less villous beneath, sometimes densely so, and 
furnished with few or numerous, closely appressed, glistening, golden 
hairs, similar hairs present also in the inflorescence; panicles usually 
large and open, yellowish; sepals 1.5 mm. long. 

Cuzco: Valle del Apurimac, Herrera. Cuzco, Soukup 260, 96. 
Huanuco: Huacachi, 1,950 meters, 4139. Junin: La Merced, 600 
meters, in forest, 5426; Killip & Smith 24067. Rio Pinedo, 800 
meters, Killip & Smith 23665. Loreto: Mishuyacu, 100 meters, in 
clearing, Klug 1380. San Martin: San Roque, in forest, Williams 
7200, 7629. Without locality: Ruiz & Pawn 1640. Argentina and 
Brazil to Central America. 

The plant is doubtfully distinct from I. Celosia. It grows usually 
at rather high elevations, or at least seldom in the lowlands. /. 
spiculigera var. picta Suesseng. (Repert. Sp. Nov. 35: 323. 1934) 
is a form in which the leaves are conspicuously striped with pink or 
red. The type is Raimondi 8397 from Chanchamayo, Junin, and 
the variety occurs also in the Putumayo region of Colombia ( Klug 
1756, 1893). 

Iresine tenuis Suesseng. Repert. Sp. Nov. 35: 32. 1934. /. 
tenuis var. discolor Suesseng. loc. cit. 

Branches slender, appressed-pilose or glabrous, fruticose; leaves 
short-petiolate, the blades ovate to oblong, short-acuminate, rounded 
to acute at the base, appressed-pilose beneath or glabrous; branches 
of the inflorescence very slender, sparsely pilose with short, ascending 
hairs, the flowers mostly inserted remotely on the ultimate branch- 
lets, up to 5 mm. long. 

San Martin: Near Tarapoto, Spruce. San Roque, Williams 7466 
(type of var. discolor). Ecuador. 

The Peruvian collections are determined by Suessenguth. 

Iresine Weberbaueri Suesseng. Repert. Sp. Nov. 35: 323. 1934. 

An erect (?) shrub of 3 meters, the branches and lower leaf surface 
covered with an exceedingly dense and fine, whitish tomentum of 
branched hairs; leaves linear-lanceolate or linear-oblong, mostly 
6-10 cm. long, short-petiolate, coriaceous, acute or obtuse and 
apiculate, attenuate at the base, glabrous on the upper surface; 
inflorescence very large, dense, and many-flowered, the flowers 
tomentose, 1 mm. long. 



518 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Libertad: Valley of the Maranon above Uchos, 1,800 meters, 
Weberbauer 7022, type. Amazonas: Above Balsas, 1,000 meters, 
Weberbauer 4267 (fide Suessenguth). 

A very distinct species because of the abundant, pale tomentum 
and very narrow leaves. 

61. NYCTAGINACEAE. Four-o'clock Family 
By Paul C. Standley 

References: Heimerl, Nat. Pflanzenfam. 16c: 86. 1934; Standley, 
The Nyctaginaceae of northwestern South America, Field Mus. 
Bot. 11: 73. 1931. 

Herbs, shrubs, or trees, sometimes scandent, occasionally armed 
with spines; leaves simple, opposite, alternate, or verticillate, 
estipulate, usually entire, often marked with conspicuous raphids; 
flowers perfect or unisexual, sometimes dioecious, usually bracteate 
or involucrate, the involucre, when present, calyx-like; perianth 
inferior, commonly corolla-like, in fruit usually enclosing closely the 
pericarp, the tube short or elongate, the limb truncate or with 3-5 
teeth or lobes; stamens 1-many, hypogynous; filaments usually 
united at the base, unequal, filiform, the anthers didymous, dorsi- 
fixed near the base, dehiscent by lateral slits; ovary included in the 
perianth tube, sessile or stipitate, 1-celled; style 1, the stigma simple; 
fruit an anthocarp, composed of the persistent, indurate base of the 
perianth tube enclosing the indehiscent utricle and adherent to it, 
costate, sulcate, or winged; seed 1, erect, the radicle inferior. A 
specimen of Pisonia macranthocarpa Donn. Smith in the herbarium 
of Field Museum was collected by Ruiz and Pavon (No. 33-78), 
possibly in Peru. It is more probable, however, that it came from 
Ecuador, where this species is known to occur. The family Nycta- 
ginaceae is almost wholly American. 

Embryo straight; plants trees or shrubs, sometimes armed with 
spines; fruit juicy; flowers dioecious; leaves chiefly opposite or 
verticillate. (Pisonieae.) 

Stamens included 1. Neea. 

Stamens exserted 2. Torrubia. 

Embryo curved; plants chiefly herbaceous, sometimes woody vines; 

fruit dry; flowers perfect; leaves opposite or alternate. 
Leaves alternate. 

Flowers not bracteate; plants chiefly herbaceous. (Salpi- 
antheae.) 3. Cryptocarpus. 



FLORA OF PERU 519 

Flowers bracteate, usually in clusters of 3, each flower adnate 
to a large, colored bract; plants woody, often armed with 

spines. (Bougainvilleae.) 4. Bougainvillea. 

Leaves opposite. 

Perianth lobes valvate; leaves commonly verticillate; plants 
often scandent; flowers minute, in umbels, not bracteate. 

(Colignonieae.) 5. Colignonia. 

Perianth lobes induplicate- valvate; leaves chiefly opposite; 
plants not scandent; flowers small or large, bracteate or 
involucrate. (Mirabileae.) 

Anthocarp lenticular, the broad, thin margins dentate. 
Flowers in 3's, surrounded by a 3-parted involucre. 

6. Allionia. 
Anthocarp not lenticular. 

Flowers subtended by a gamophyllous, calyx-like involucre. 

9. Mirabilis. 
Flowers not involucrate. 

Anthocarp with 5 or fewer angles 7. Boerhaavia. 

Anthocarp terete, 10-costate 8. Commicarpus. 

1. NEEA R. & P. 

Shrubs or trees; leaves opposite or verticillate, rarely alternate, 
petiolate, entire, coriaceous to membranaceous; flowers unisexual 
and normally dioecious, small, sessile or pedicellate, tribracteolate 
at the base, arranged in lateral, axillary, or terminal cymes; stami- 
nate perianth urceolate, globose, or tubular, shortly 4-5-dentate; 
stamens 5-10, included, the filaments unequal; pistillate perianth 
urceolate or tubular, constricted above the ovary, often contracted 
at the mouth; stigma penicillate; anthocarp narrowly or broadly 
ellipsoid, crowned by the persistent free portion of the perianth. 

The Amazon region of Peru and Brazil is the area in which this 
genus is most richly represented. The species are poorly marked, 
although when each is represented by a single specimen the difficulties 
in separating them are not so great! When there has been accumu- 
lated a large series of specimens, like that in the herbarium of Field 
Museum, the case is different. Almost every specimen seems to 
differ slightly from every other one, and it becomes necessary to dis- 
tinguish species by petty differences or to consider the material as 
representing a few variable species, which even then are not definitely 
separable from one another. With the ample material at hand, it 



520 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

seems impossible to prepare a satisfactory account of the Peruvian 
species, but it is suspected that additional material will complicate 
rather than simplify the matter. In the Berlin herbarium Dr. 
Heimerl, foremost student of the group, has indicated as new a large 
number of Amazonian species still unpublished, that are represented 
in Field Museum by photographs and fragments. I doubt that it 
will be possible to prepare a practical key to the forms he has dis- 
tinguished, and some of them, indeed, appear to be identical. The 
key here presented will not be found very useful, except with ample 
named material for comparison. 
Leaves sparsely or densely pilose or hirsute beneath, sometimes in 

age only along the costa. 

Leaves bright green when dried, those of a pair often very unequal, 

the larger several times as long as the smaller. Inflorescence 

pendent on a very long and slender peduncle . . N. Williamsii, 

Leaves fuscous when dried, those of a pair mostly subequal. 

Bractlets much elongate and almost setiform, about equaling 

the perianth N. hirsuta. 

Bractlets small and inconspicuous, much shorter than the 

perianth. 

Bractlets 2-2.5 mm. long, almost half as long as the perianth. 

N. oppositifolia. 

Bractlets minute, mostly 1 mm. long or less. 
Nerves of the leaves chiefly straight, not arcuate. 

A 7 , subpubescens. 

Nerves strongly arcuate and upcurved A 7 , parvi flora. 

Leaves glabrous, at least in age. 

Leaves all or chiefly alternate, green when dried N. virens. 

Leaves all or mostly opposite or verticillate. 

Inflorescences mostly lateral on naked branches or on the trunk. 

N. floribunda. 

Inflorescences terminal or in the axils of leaves. 
Leaves bright green when dried; inflorescence lax, many- 
flowered, pendent on a long and very slender peduncle. 

N. laxa. 
Leaves fuscous when dried. 

Leaf blades small, mostly 2-5 cm. wide; inflorescence lax 
and open, glabrate or sparsely puberulent or hirtous. 

N. Spruceana. 



FLORA OF PERU 521 

Leaf blades usually 6-12 cm. wide or more; inflorescence 

sometimes densely tomentulose or rufous-hirtous. 
Petioles mostly 4-5 cm. long; leaf blades oblong-lanceo- 
late, very large, about equally attenuate to each end. 

N. verticillata. 

Petioles less than 2.5 cm. long; leaf blades not oblong- 
lanceolate, sometimes oblanceolate. 
Inflorescence long-pedunculate, lax and many-flowered, 
glabrate or appressed-tomentulose. 

N. macrophylla. 

Inflorescence short-pedunculate, dense, usually densely 
rufous-hirtous or coarsely tomentulose. 

N. divaricata. 

Neea divaricata Poepp. & Endl. Nov. Gen. & Sp. 2: 45. 
pi. 161. 1838. 

A shrub or small tree, the branchlets glabrous or when young 
puberulent or tomentulose; leaves opposite or verticillate, fuscous 
when dried, subcoriaceous or thick-membranaceous, glabrous or 
practically so, the petioles stout, mostly 1.5 cm. long or less; leaf 
blades oblong to elliptic, mostly 10-23 cm. long and 5-10 cm. wide, 
abruptly acute to long-acuminate, acute or obtuse at the base and 
frequently oblique; inflorescences terminal or axillary, usually erect, 
short-pedunculate, commonly small and rather dense, densely rufous- 
hirtous or tomentulose except sometimes in fruit; perianth tubular- 
urceolate, ferruginous-puberulent or glabrate, 4-6 mm. long; antho- 
carp oblong, glabrous or nearly so, 1 cm. long. Neg. 3156. 

Amazonas: Rio Maranon, Tessmann 3995 (photo, and fragm. ex 
hb. Berol.; indicated as a new species by Heimerl). Loreto: Iquitos, 
Williams 8062. Rio Ucayali, Tessmann 3029 (photo, and fragm. ex 
hb. Berol.; indicated by Heimerl as a new species). La Victoria, in 
forest, Williams 2678. Florida, 180 meters, in forest, King 2334. 
San Martin: Rumizapa, near Tarapoto, Williams 6830. Juan 
Guerra, near Tarapoto, Williams 6906. Tarapoto, Williams 6507. 
Huanuco: Cochero, Poeppig 1493 (photo, of type ex hb. Berol.). 
Colombia and Amazonian Brazil. 

Neea floribunda Poepp. & Endl. Nov. Gen. & Sp. 2: 46. 1838. 
N. cauliflora Heimerl, Pflanzenfam. ed. 2. 16c: 129. 1934. 

A shrub or tree, sometimes 6 meters high, glabrous or practically 
so except sometimes in the inflorescence, there sparsely and minutely 
puberulent; bark whitish; leaves opposite, fuscous when dried, sub- 



522 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

coriaceous, the stout petioles mostly 1-2 cm. long; leaf blades obo- 
vate-elliptie to oblanceolate-oblong, 8-21 cm. long, 3-9 cm. wide, 
abruptly acuminate or rarely cuspidate-acuminate, the tip obtuse 
or acute, at the^base acute or obtuse, or sometimes long-attenuate; 
inflorescences produced on the older branches below the leaves, or 
also on the trunk, small or large, mostly many-flowered and lax, 
the flowers sessile or nearly so, the bractlets minute; perianth 
glabrous, pink, 4 mm. long, tubular-urceolate; stamens 6; anthocarp 
ellipsoid-oblong, 9-13 mm. long, glabrous. Negs. 3148, 3145. 

. Loreto: Pebas, in forest, Williams 1673. Balsapuerto, 350-550 
meters, dense forest, Killip & Smith 28518. Puerto Arturo, Williams 
5023. Yurimaguas, 150 meters, Williams 4708, 4535; Killip & Smith 
27980, 27553. Rio Acre, Ule 9365b. San Martin: Mainas, Poeppig 
(photo, of type ex hb. Berol.). Amazonian Brazil. 

It is not at all certain that all the collections cited are conspecific, 
but the material is not sufficiently extensive to indicate how it may 
be divided into forms or species. The vernacular name is given 
as "mullo caspi." 

Ule 9365b from the Rio Acre, indicated by Heimerl as a new 
species, probably is not separable from N. floribunda. 

Neea hirsuta Poepp. & Endl. Nov. Gen. & Sp.2: 45. pi. 161. 1838. 

A tall shrub, the branchlets hirsute; leaves ternate or quarternate, 
short-petiolate, the blades elliptic-oblong, short-acuminate, cuneate 
at the base, hirtous on both surfaces, membranaceous, about 12.5 
cm. long and 5 cm. wide; peduncles axillary, pendent, long and 
slender, hirsute, the inflorescence lax, corymbiform, 3.5-5 cm. long, 
the flowers short-pedicellate; bractlets densely ciliate, 4-6 mm. 
long, equaling the perianth; perianth of the staminate flower tubular, 
red, glabrous; stamens 8-10. 

San Martin: In forests of the Province of Mainas, Poeppig, type. 

I have seen no material of this species, which, according to the 
original plate, must be a strikingly distinct one. 

Neea laxa Poepp. & Endl. Nov. Gen. & Sp. 2: 45. pi. 162. 1838. 

A shrub or small tree, sometimes 9 meters high, glabrous except 
in the inflorescence, there usually sparsely rufous-puberulent or 
hirtulous; leaves opposite or ternate, those of a pair often very 
unequal, the smaller ones mostly rounded and only 2-3 cm. long, 
the petioles commonly very short; leaf blades oblong to oblong- 
elliptic, occasionally oblong-ovate, commonly 10-17 cm. long and 



FLORA OF PERU 523 

3-7.5 cm. wide, abruptly acute or acuminate, sometimes long- 
acuminate, the tip usually obtuse, bright green when dried, mem- 
branaceous or subcoriaceous, acute to rounded at the base; inflores- 
cences axillary, usually on very long and slender, pendent peduncles, 
the panicles broadly pyramidal, laxly many-flowered, the flowers 
sessile or nearly so; bractlets lance-subulate, minute; branches of the 
inflorescence slender, sometimes colored pink; perianth tubular or 
narrowly urceolate, 6-10 mm. long; stamens 8; anthocarp dark red 
or purple, oblong or narrowly ellipsoid, 12-15 mm. long, glabrous. 

Junin: Rio Paucartambo Valley, 700 meters, in forest, Killip 
& Smith 25306. Loreto: Type from Yurimaguas, Poeppig. Puerto 
Arturo, in forest, Williams 5370, 5089; Killip & Smith 27767, 
27926, 27729. San Ramon, in forest, Williams 4554. Yurimaguas, 
135 meters, Williams 4691; Killip & Smith 27700, 30688. Santa 
Rosa, 135 meters, dense forest, Killip & Smith 28727. Lower Rio 
Nanay, in forest, Williams 583. Caballo-cocha, in forest, Williams 
2025. San Martin: Tarapoto, 750 meters, Williams 5876, 6108. 
Juanjui, 400 meters, in forest, Klug 3905; flowers cream-colored. 

Vernacular name, "puca huayo." Weberbauer 6949, determined 
by Heimerl as a new species, is closely related but possibly distinct. 

Neea macrophylla Poepp. & Endl. Nov. Gen. & Sp. 2: 46, 1838. 

A shrub or tree 3-12 meters high, glabrous or nearly so except 
in the inflorescence, there sparsely or rather densely puberulent 
or tomentulose; leaves opposite or verticillate, subcoriaceous, fus- 
cous when dried, large, short-petiolate, the petioles mostly 1-2.5 
cm. long, stout; leaf blades variable, mostly elliptic-oblong to 
elliptic, sometimes obovate or oblanceolate-oblong, 14-35 cm. 
long, 6.5-16 cm. wide, abruptly or gradually acuminate or long- 
acuminate, at the base acute to rounded, often oblique; inflorescences 
axillary, long-pedunculate, probably pendent, commonly large and 
many-flowered, the flowers sessile or short-pedicellate; staminate 
perianth tubular, about 12 mm. long, white or green; branches of the 
inflorescence pink or red; anthocarp oblong, glabrous, 15 mm. long, 
white. Neg. 3151. 

Amazonas: Upper Maranon, Tessmann 4808 (photo, and fragm. 
from hb. Berol.; indicated as a new species by Heimerl; probably 
referable here). Junin: Puerto Bermudez, 375 meters, dense forest, 
Killip & Smith 26638. Loreto: Mainas, Poeppig (photo, of type ex 
hb. Berol.). Puerto Arturo, in forest, 200 meters, Williams 5079. 
Yurimaguas, 200 meters, in forest, Williams 4665, 3857, 3874; 



524 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Killip & Smith 27699. Mishuyacu, near Iquitos, 100 meters, dense 
forest, Killip & Smith 29886; King 546, 14, 16, 799. Soledad, on 
Rio Itaya, 110 meters, dense forest, Killip & Smith 29635. Iquitos, 
100 meters, in woods, Killip & Smith 27286, 29845. San Martin: 
Zepelacio, 1,100 meters, Klug 3731. Pongo de Cainarachi, 230 
meters, Klug 2649. Chazuta, 260 meters, Klug 4089. 

Here, as in the case of some of the other species, it is possible 
that the specimens cited represent more than a single species. 

Neea oppositifolia R. & P. Syst. 91. 1798; Fl. Peruv. 4: pi. 329. 

A shrub or small tree about 7 meters high; leaves opposite, 
oblong-elliptic to elliptic-obovate, abruptly short-acuminate or merely 
acute, short-petiolate, at the base acute or acutish, beneath sparsely 
pilose; staminate inflorescences arising from the upper leaf axils, 
apparently pendent on rather long and slender peduncles, the cymes 
small and rather few-flowered ; stamens 8. 

Junin: Forests of Pillao near Chacahuasi, Ruiz & Pavdn. 

Neea parviflora Poepp. & Endl. Nov. Gen. & Sp. 2: 46. 1838. 

A shrub or small tree, 3-7.5 meters high, the young branchlets 
rufous-hirtellous; leaves mostly opposite, short-petiolate, thick- 
membranaceous, fuscous when dried, elliptic-oblong or narrowly 
elliptic, 7-20 cm. long, 2.5-7 cm. wide, abruptly long-acuminate, 
acute at the base and often very oblique, glabrous above or nearly 
so, beneath sparsely or densely pilose with short, spreading, rufous 
hairs; inflorescences axillary, small and few-flowered, erect, stout, 
short-pedunculate, usually umbelliform, densely rufous-hirtous or 
rufous-tomentose, the flowers glomerate, sessile; bractlets minute, 
scarcely 1 mm. long; pistillate perianth densely rufous-tomentulose, 
4-5 mm. long; anthocarp oblong, 1 cm. long, pilose. Neg. 7397. 

Junin: Pichis Trail, Santa Rosa, 625-900 meters, dense forest, 
Killip & Smith 26191. Puerto Bermudez, 375 meters, dense 
forest, Killip & Smith 26514- Loreto: Iquitos, 100 meters, in woods, 
Killip & Smith 27423; Williams 8120. La Victoria, in forest, 
Williams 2720. Between Yurimaguas and Balsapuerto, 135-150 
meters, dense forest, Killip & Smith 28229. Santa Rosa, Williams 
4915. Yurimaguas, in forest, Williams 4108. San Martin: Type 
from forests of the Province of Mainas, Poeppig. Cerro de Ponasa, 
Ule 6824 (fragm. ex hb. Berol.). Also in Colombia. 

Called "yana muco" in the Putumayo region of Colombia. The 
leaves are chewed by the Indians, making their teeth black but 
preserving them. 



FLORA OF PERU 525 

Neea Spruceana Heimerl, Notizbl. Bot. Gart. Berlin 6: 131. 
1914. N. Weberbaueri Heimerl, Bot. Jahrb. Engl. 54: Beibl. 117: 
38. 1916. 

A shrub 1-4 meters high, the young branchlets glabrous, sparsely 
rufous-puberulent, or sparsely hirtulous; leaves opposite, ternate, or 
quaternate, fuscous when dried, membranaceous or subcoriaceous, 
the petioles mostly 4-15 mm. long, puberulent or hirtulous at first; 
leaf blades elliptic to oblong, mostly 7-16 cm. long and 2-5 cm. 
wide, sometimes somewhat broader, gradually or abruptly acumi- 
nate or long-acuminate, at the base acute to attenuate, often oblique, 
glabrous or practically so except sometimes when very young; 
inflorescences mostly erect, on a short, slender peduncle, sparsely 
puberulent or hirtulous, small, lax, few-flowered or many-flowered; 
perianth yellowish or reddish, sparsely rufous-puberulent or almost 
glabrous, 5.5-7.5 mm. long; bractlets 1.5 mm. long or less; stamens 
5-9. Negs. 21777, 3167. 

Amazonas: Pongo de Manseriche, Tessmann 4768 (photo, and 
fragm. ex hb. Berol.; indicated by Heimerl as type of a new species, 
and perhaps distinct). Upper Maranon, Tessmann 4285 (photo, and 
fragm. ex hb. Berol.; indicated by Heimerl as a new species, and 
possibly distinct). Huanuco: Monson in forest, 1,200 meters, 
Weberbauer 3495 (photo, and fragm. of type of N. Weberbaueri, ex 
hb. Berol.). Junin: La Merced, 600 meters, 5421. Loreto: Yarina- 
cocha, Rio Nanay, in forest, Williams 623. Soledad on Rio Itaya, 
110 meters, dense forest, Killip & Smith 30693. Pumayacu, Klug 
3179. Florida, Klug 2026. Iquitos, edge of forest, Williams 8187. 
Rio Ucayali, Tessmann 3146 (indicated by Heimerl as a new species; 
photo, ex hb. Berol.). San Martin: Tarapoto, Spruce 4858 (type), 
4883; Ule 6498 (photo, and fragm. ex hb. Berol.). Rio Acre, UU 
9367 (photo, ex hb. Berol.; indicated as a new species by Heimerl). 
Tarapoto, Williams 5885. San Roque, in forest, Williams 7654. 
Juan Guerra near Tarapoto, 720 meters, Williams 6912. Juanjui, 
Klug 3849, 3864. Zepelacio, Klug 3670. Pongo de Cainarachi, 
230 meters, Klug 2609. Colombia. 

Vernacular name, "topamaka blanca." The material referred 
here is variable, but it is doubtful whether it can be divided con- 
servatively into two or more distinct species. 

Neea subpubescens Heimerl, Beitr. Syst. Nyctag. 36. 1897. 
A small or medium-sized tree, the young branchlets rufous- 
hirtellous; leaves short-petiolate, membranaceous, mostly opposite, 



526 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the petioles 5-10 mm. long; leaf blades lance-oblong to obovate- 
oblong, 7-10.5 cm. long, 2.5-4 cm. wide, acuminate or abruptly 
acuminate, acute to attenuate and more or less oblique at the base, 
glabrous or nearly so above, beneath densely or sparsely pilose with 
ferruginous, spreading hairs; inflorescences small, short-pedunculate, 
lax or dense, rather few-flowered, erect, broadly pyramidal, densely 
hirtellous, the flowers subsessile; pistillate perianth oblong-urceolate, 

5 mm. long, ferruginous-puberulent near the base, glabrate above. 
Neg. 3169. 

San Martin : Rio Mayo near Tarapoto, 360-900 meters, in forest, 
Williams 6267. Rumizapa near Tarapoto, Wittiams 6815, 6802. 
Also in the upper Amazon Valley of Brazil. "Yntutu caspi." 

It is rather probable that there should be referred to this species 
Spruce 4196 from Tarapoto (not seen by the present writer), which is 
referred doubtfully by Heimerl (Beitr. Syst. Nyctag. 38. 1897) to 
N. Selloiana Heimerl. 

Neea verticillata R. & P. Syst. 90. 1798; Fl. Peruv. 4: pi. 328. 

A small tree, about 4.5 meters high, glabrous except in the inflo- 
rescence, there sparsely and inconspicuously puberulent; leaves 
ternate or quaternate, rather long-petiolate, the blades fuscous 
when dried, subcoriaceous, very large, narrowly oblong-lanceolate 
or elliptic-oblong, 25-40 cm. long, 7-10.5 cm. wide, very long- 
acuminate; inflorescence pendent, long-pedunculate, many-flowered, 
large and much branched, the branches stout, deep red; anthocarp 
oblong, glabrous, about 13 mm. long, greenish white. Neg. 27760. 

Huanuco: Type from Posuso, Ruiz & Pavdn. Vitoc, Ruiz 

6 Pavon. Junin : La Merced, 600 meters, in forest, 55.47. Without 
locality: Ruiz & Pavdn 11-95. 

The collection from La Merced agrees perfectly with the Ruiz 
and Pavon illustration. The collectors of the type state that the 
Indians, in order to protect themselves against the bites of mosquitoes 
or gnats, were accustomed to rub their faces, hands, and feet with 
the juice of the fruit. The species is easy to recognize because of 
the exceptionally large and narrow leaves. 

Neea virens Poepp. ex Heimerl, Beitr. Syst. Nyctag. 38. 1897. 

A shrub or small tree, 1-4.5 meters high, glabrous throughout or 
nearly so; leaves mostly alternate, membranaceous, bright green 
when dried, the petioles 1-2.5 cm. long; leaf blades narrowly to 
broadly oblong-elliptic, sometimes oblong-obovate, mostly 10-21 
cm. long and 3-7 cm. wide, gradually or abruptly acuminate or long- 



FLORA OF PERU 527 

acuminate, acute or cuneate at the base, the nerves 9-12 pairs; 
inflorescence axillary, small and few-flowered, usually scarcely 
longer than the petiole, sometimes as much as 6 cm. long, the 
branches very thick, pink, glabrous or very sparsely puberulent, the 
peduncles very short or rarely as much as 2.5 cm. long; staminate 
perianth 11-12 mm. long, oblong, glabrous; stamens 10; anthocarp 
dark purple, 1 cm. long. Neg. 3175. 

Ayacucho: Rio Apurimac Valley near Kimpitiriki, 400 meters, 
dense forest, Killip & Smith 22891, 22877. Junin: Cochero, Poeppig 
1330 (photo, of type ex hb. Berol.). Loreto: La Victoria, in forest, 
Williams 2838; edge of forest, Williams 2661, 2682. Iquitos, 100 
meters, in woods, Killip & Smith 27449. Balsapuerto, 300 meters, 
dense forest, Killip & Smith 28577. Lower Rio Nanay, Williams 592. 
Also in Brazil. 

The flowers are described as either pink or white, and some- 
times as greenish. 

Neea Williamsii Standl. Field Mus. Bot. 8: 309. 1931. 

A shrub, the very slender branches pilose; leaves short-petiolate, 
thick-membranaceous, bright green when dried, those of a pair very 
unequal, the larger blades broadly elliptic-oblong, about 17 cm. 
long and 7.5-8 cm. wide, abruptly short-acuminate, oblique at the 
base, lustrous above, with prominulous, reticulate venation, beneath 
rather densely spreading-pilose; blades of the smaller leaves rounded 
or broadly elliptic, 1.5-2.5 cm. long; pistillate inflorescence lax 
and open, broadly pyramidal, 5.5-9 cm. broad, the lower branches 
opposite or verticillate, the flowers glomerate, sessile or nearly so; 
peduncle very slender, 13-16 cm. long; bractlets linear-subulate, 
1.5-2 mm. long; immature perianth 3 mm. long, sparsely rufous- 
puberulent. 

San Martin: Tarapoto, 360-900 meters, Williams 6567 (type). 

2. TORRUBIA Veil. 

Unarmed shrubs or trees, glabrous or pubescent; leaves chiefly 
opposite and petiolate, entire, membranaceous to coriaceous; flow- 
ers small, dioecious, reddish, greenish, or yellowish, exinvolucrate, 
2-3-bracteolate, sessile or pedicellate, arranged in lateral or terminal, 
pedunculate cymes; staminate perianth obconic-campanulate, the 
limb 5-dentate; stamens 6-10, exserted, the filaments unequal; 
pistillate perianth tubular; anthocarp drupaceous, red to black, 
oblong, the exocarp fleshy and juicy. 



528 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

The genus is represented in South America by a large number 
of species, but only a few are known from the western part of the 
continent, and only one has been discovered thus far in Peru. 

Torrubia myrtiflora Standl. Field Mus. Bot. 8: 307. 1931. 

A shrub or small tree, nearly glabrous except in the inflorescence; 
leaves opposite or ternate, fuscous when dried, the slender petioles 
12-20 mm. long; leaf blades chartaceous, lance-oblong or narrowly 
elliptic-oblong, 9-13 cm. long, 3.5-5 cm. wide, short-acuminate, at 
the base acute or short-acuminate; staminate inflorescence long- 
pedunculate, many-flowered, 7-10 cm. broad, rather densely rufous- 
tomentulose; perianth 4-4.5 mm. long, densely rufous-tomentulose. 

Loreto: Puerto Arturo, Yurimaguas, 200 meters, edge of forest, 
Williams 5162 (type). "Clavo-caspi." 

3. GRYPTOCARPUS HBK. 

Large herbs or small shrubs, pubescent and usually viscid; 
leaves alternate, petiolate, the blades entire; flowers small, perfect, 
neither involucrate nor bracteate, glomerate or racemose, forming 
terminal and axillary, panicled cymes; perianth pyriform, green 
or reddish, persistent, not accrescent, the limb with 4-5 teeth or 
lobes; stamens 4-5; anthocarp subglobose, coriaceous, minute. 

Cryptocarpus pyriformis HBK. Nov. Gen. & Sp. 2: 188. 
pi 124. 1817. C. cordifolius Moric. PI. AmeY. 75. pi. 50.^1830. 
C. cordiformis cordifolius Moq. in DC. Prodr. 13, pt. 2: 88. 1849. 
Salpianthus pyriformis Standl. Field Mus. Bot. 11: 95. 1931. 

Plants herbaceous or suffrutescent, scandent or trailing over 
other plants, the stems 2-5 meters long, densely viscid-puberulent; 
leaves on stout petioles, the blades rhombic-ovate to broadly ovate- 
deltoid or rounded-ovate, mostly 2.5-5 cm. long, usually cordate 
or deeply cordate at the base, rounded or very obtuse at the apex, 
densely viscid-puberulent or tomentulose; inflorescence raceme-like, 
very leafy or naked, densely viscid-puberulent, the flowers subsessile; 
perianth less than 2 mm. long, green, pyriform in age, with spreading, 
white lobes; stamens 4; anthocarp globose, blackish, 1 mm. in 
diameter. 

Cajamarca: Type collected by Humboldt and Bonpland near 
Contumasa and Cascas. Piura: Region of Negritos, Haught F63. 
Tumbez: Coastal plain between Zorritos and Tumbez, Weberbauer 
7747. Also in Ecuador and the Galapagos Islands. Vernacular name, 
"nacupillo." 



FLORA OF PERU 529 

Haught supplies the following notes regarding the occurrence 
of the plant about Negritos: "A characteristic and very common, 
scrambling shrub of watercourses. Grows about 5 meters high when 
supported by other plants, elsewhere only 1-3 meters high. Trailing 
shoots as much as 10 meters long. Found in this region practically 
everywhere that there is any soil." 

4. BOUGAINVILLEA Commers. 

Shrubs or small trees, often scandent, usually armed with stout, 
simple or branched spines; leaves alternate, petiolate, entire; flowers 
perfect, either solitary and subtended by 3 bracts or usually in a 
3-flowered, axillary inflorescence consisting of 3 large, persistent, 
often brightly colored bracts, a flower borne on the inner surface 
of each bract, its pedicel confluent with the costa of the bract; 
perianth tubular, the limb usually shallowly 5-lobed, the tube 
terete or 5-angled; stamens 5-10, somewhat unequal, connate at 
the base into a short cup; anthocarp fusiform, coriaceous, 5-costate. 
Spines forked at the apex; leaves linear-spatulate or oblong-spatu- 
late; inflorescences 1-flowered, surrounded by 3 bracts. Sub- 
genus Tricycla B. spinosa. 

Spines simple, or often wanting; leaves broader; inflorescences 
normally 3-flowered. Subgenus Eubougainvillea. 

Perianth glabrous B. peruviana. 

Perianth variously pubescent. 

Perianth tube hirsute or villous B. spectabilis. 

Perianth tube merely puberulent. 

Leaves thin, glabrous or nearly so, at least in age; bracts 
when fully developed mostly 3-4 cm. long, usually 

acute or acuminate B. glabra. 

Leaves thick, densely puberulent except when old; bracts 
usually less than 3 cm. long, very obtuse or rounded at 
the apex B. pachyphylla. 

Bougainvillea glabra Choisy in DC. Prodr. 13, pt. 2: 437. 1849. 
B. spectabilis var. glabra Hook, in Curtis's Bot. Mag. pi. 4810. 1854. 

A large, spiny, woody vine, the branchlets puberulent or glabrate; 
leaf blades broadly ovate to ovate-lanceolate, 4-10 cm. long, grad- 
ually or abruptly acute or acuminate, puberulent when young but 
quickly glabrate; bracts usually purplish red, broadly ovate or 
oval, mostly 2.5-4.5 cm. long, mostly acute or acuminate, sparsely 
puberulent or glabrous; perianth 15-20 mm. long; anthocarp 7-13 
mm. long, sparsely puberulent or glabrate. Neg. 7405. 



530 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Near Iquitos, Williams 1477, 3525, 8110. Yurimaguas, 
Williams 4052. Fortaleza, Yurimaguas, Williams 4324- La Victoria, 
Williams 2513. Native of Brazil. 

In Peru the plant is known only in cultivation, unless it may 
have escaped from cultivation in the Amazon region, or persists 
about the sites of former dwellings. It is grown commonly as an 
ornamental vine in tropical regions of the whole globe, and in the 
United States it often is seen in hothouses. 

Bougainvillea pachyphylla Heimerl ex Standl. Field Mus. 
Bot. 8: 308. 1931. 

Usually a shrub or small tree, but sometimes scandent, armed 
with numerous spines, the young branchlets finely puberulent; leaf 
blades ovate to broadly elliptic, thick and somewhat leathery, 
3-6.5 cm. long, acute or acuminate, or gradually narrowed to an 
obtuse apex, broadly rounded at the base, finely and usually densely 
puberulent; bracts rose-colored, broadly elliptic-ovate to almost 
orbicular, glabrate; perianth densely puberulent, 9-11 mm. long. 
Neg. 3094. 

Piura: Without definite locality, Raimondi 8703 (photo, of type 
from hb. Berol.). Frias, Weberbauer 6432 (fragm. from hb. Berol.). 
Amotape Hills, Haught F24. "Papelillo." 

Bougainvillea peruviana H. & B. PL Aequin. 1: 174. pi. 49. 
1808. Tricycla peruviana Poir. Encycl. Suppl. 5: 359. 1817. 

A shrub 3-7 meters high, erect or scandent, armed with numerous 
slender spines 1-2.5 cm. long, the branches sparsely puberulent or 
glabrate; leaves thin, slender-petiolate, the blades broadly ovate to 
suborbicular, 5-7 cm. long, subobtuse or abruptly acute, glabrous 
or nearly so except when very young; bracts bright rose, 1.5-3.5 
cm. long, obtuse or rounded at the apex, glabrous except along the 
puberulent costa; perianth 16-20 mm. long, white or whitish; antho- 
carp 10 mm. long, glabrous. Neg. 27752. 

Tumbez: Mountains southeast of Hacienda La Choza, Prov. 
Tumbez, 200-300 meters, Weberbauer 7725. Piura: Type from Rio 
Huancabamba, Humboldt & Bonpland. Reported from Jae*n and 
Corazi. Occurring also in Ecuador and Colombia. 

Bougainvillea spectabilis Willd. Sp. PI. 2: 348. 1789. 

A large, woody vine, armed with stout spines, the branches 
fulvous-villous; leaf blades broadly ovate to suborbicular, 5-10 cm. 
long, rounded to acutish at the base and often short-decurrent, 



FLORA OF PERU 531 

abruptly acute or acuminate at the apex, usually densely villous 
beneath; bracts purplish red, ovate-oval or broadly ovate, 2-4.5 cm. 
long, abruptly acute or acuminate or rarely obtuse, sparsely puberu- 
lent or short-villous; perianth 15-30 mm. long, the limb yellowish; 
anthocarp 11-14 mm. long, densely short-villous. 

Lima: Purchased in the Lima market, Cook & Gilbert 2075. 
Native of Brazil. 

This Bougainvillea is grown widely for ornament in tropical 
regions, but in cultivation it is much less common than B. glabra. 
The vernacular name at Lima is given as "papelillo." 

Bougainvillea spinosa (Cav.) Heimerl in Engl. & Prantl, Nat. 
Pflanzenfam. 3, Ib: 27. 1889. Tricycla spinosa Cav. Anal. Cienc. 
Nat. 5: 6B.pl. 40. 1802. 

A shrub 2-4 meters high, the stout branches sparsely puberulent 
on the younger parts, armed with stout, rigid spines 2 cm. long or 
less, these furcate at the apex; leaves linear-spatulate or oblong- 
spatulate, 9-15 mm. long, 2-4 mm. wide, thick and fleshy, obtuse 
or rounded at the apex, gradually narrowed to the base, very shortly 
petiolate, glabrous or nearly so; flower borne on a slender peduncle 
3-6 mm. long, solitary; bracts green, shorter than the flower, broadly 
cordate-ovate to orbicular, about 12 mm. long, sparsely rufous- 
puberulent; perianth 8-13 mm. long, glabrous or sparsely puberulent; 
anthocarp 6-7 mm. long. 

Tacna: Torata, Prov. Moquehua, 2,300 meters, Weberbauer 7414. 
Also in southern Argentina. 

The occurrence of this species so far from its range in Argentina 
furnishes an interesting and rather remarkable case of interrupted 
distribution. 

5. COLIGNONIA Engl. 

Plants herbaceous or suffrutescent, often clambering or sub- 
scandent; leaves entire, opposite or verticillate, long-petioled, those 
of the inflorescence often white; flowers perfect, without bracts or 
involucre, arranged in small and usually very numerous umbels; 
perianth commonly white or greenish, very small, campanulate or 
funnelform, 3-5-parted, the tube 3-5-angled; stamens 5-6, slightly 
unequal, about equaling the perianth; anthocarp ellipsoid to fusiform 
or orbicular, angled or winged. 
Perianth normally 5-parted; anthocarp obpyramidal, angled but 

not winged; leaves with numerous pale raphids on the lower 

surface. (Apterae.) 



532 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Upper leaves acute or acuminate, narrow C. acutifolia. 

Upper and lower leaves broad, usually rounded or very obtuse 
at the apex. 

Leaves glabrous or nearly so beneath C. Weberbaueri. 

Leaves laxly tomentose beneath C. pubigera. 

Perianth 3-parted; anthocarp 3-winged; leaves without conspicuous 
raphids. 

Leaves densely rufous- tomentose beneath C. rufopilosa. 

Leaves glabrous beneath or nearly so. 
Leaf blades small, mostly 1-2 cm. long, about as broad as 

long C. microphylla. 

Leaf blades larger, most of them 3-5 cm. long. 

Stamens about equaling the perianth C. scandens. 

Stamens about twice as long as the perianth . .C. biumbellata. 

Colignonia acutifolia Heimerl, Denkschr. Akad. Wiss. Wien 
70: 136. 1900. C. glomerata var. acutifolia Heimerl, loc. cit., in syn. 

Plants herbaceous, about 1 meter long, the stems glabrate; leaves 
of a pair strongly unequal, the upper, bract-like ones whitish, lanceo- 
late, the lower ones ovate-lanceolate, 5-10.5 cm. long, attenuate or 
acuminate, paler beneath, sparsely hirtulous beneath along the veins, 
elsewhere glabrous, the petioles equaling or much shorter than the 
blades; umbels densely many-flowered, the pedicels scarcely 1 mm. 
long, sparsely hirtulous; perianth 3 mm. long, sparsely hirtulous at 
the base, the lobes oblong, obtuse; anthocarp almost pyriform, 3-3.5 
mm. long, attenuate to the base. 

Cajamarca: Type collected at Cutervo, Jelski. Heimerl reports 
also Mathews 3122, without locality. 

Colignonia biumbellata Ball, Journ. Linn. Soc. 22: 54. 1885. 

An erect shrub, the branches nearly glabrous; leaves of a pair 
subequal, the petioles equaling or shorter than the blades; leaf 
blades broadly deltoid-ovate, about 4 cm. long, subobtuse at the 
apex, subtruncate or broadly rounded at the base, paler beneath, 
glabrous except for sparse hairs along the costa; umbels 10-20- 
flowered, the capillary pedicels 3-5 mm. long, glabrous; perianth 
more than 3 mm. long; stamens about twice as long as the perianth. 

Lima: Type collected by Ball in the upper valley of the Rimac, 
near Tamboraque, at 2,700 meters. 

I have seen no material of this species, but from the description 
probably it is scarcely distinguishable from C. scandens, and it may 



FLORA OF PERU 533 

well be synonymous with it. It may be remarked here that most 
species of Colignonia are closely related, and the characters by which 
some of them are separated are of apparently slight consequence. 

Colignonia microphylla Heimerl, Bot. Jahrb. 54: Beibl. 117: 
37. 1916. 

A scandent shrub, the branches brown, puberulent at the nodes; 
leaves more or less unequal, small, the uppermost white, the slender 
petioles equaling or shorter than the blades, tomentulose or glabrate; 
leaf blades rounded-ovate or orbicular, mostly 1-2 cm. long, usually 
rounded at base and apex, appearing glabrous but really furnished 
beneath with an exceedingly minute, microscopic, pale tomentum; 
umbels few in the upper axils, few-flowered, the slender pedicels 
glabrous, 2 mm. long; perianth white, glabrous, 3 mm. long; 
immature anthocarp 2.5 mm. long, 3-winged, elliptic. Neg. 3133. 

Cuzco: Yanamanche, between Cuzco and Santa Ana, among 
shrubs, 3,400-3,500 meters, Weberbauer 4957 (type). Lucumayo 
Valley, 1,800-3,600 meters, Cook & Gilbert 1295. 

The vernacular name is given by Cook as "sachaparaquay." 

Colignonia pubigera Heimerl, Bot. Jahrb. 42: 79. 1908. 

Plants herbaceous or suffrutescent, trailing over herbs and 
shrubs, sometimes 2 meters long, the stems sparsely puberulent at 
the nodes; leaves opposite or verticillate, unequal, the slender 
petioles equaling or shorter than the blades, puberulent or glabrate; 
leaf blades mostly ovate-deltoid and 4-7 cm. long, broadly rounded 
or subtruncate at the base, narrowed to the obtuse or acutish apex, 
glabrate above, beneath rufous-tomentulose, especially along the 
veins; umbels about 15-20-flowered, the glabrous, slender pedicels 
3-4 mm. long, or in fruit as much as 6 mm. ; perianth white, 3.5 mm. 
long, sparsely hirtulous below; stamens equaling the perianth lobes; 
anthocarp clavate, 5-angled, 4.5 mm. long. Negs. 3132, 6988. 

Ancash: Below Cajabamba, between Samanco and Caraz, in 
thickets along stream, 3,000-3,100 meters, Weberbauer 3132 (type). 
Huanuco: Eighteen miles northeast of Huanuco, 3,000 meters, 
2150. Locality uncertain: Weberbauer 6988. 

Weberbauer 6988 has been indicated by Heimerl as a new species, 
but it does not appear to differ appreciably from C. pubigera. 

Colignonia rufopilosa Kuntze, Rev. Gen. 3, pt. 2: 264. 1898. 

Plants herbaceous or suffrutescent, 1-3 meters long, usually 

clambering, the stout branches densely ferruginous-tomentose; leaves 



534 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

chiefly opposite, those of a pair subequal, the reduced ones of the 
inflorescence often white, the slender petioles shorter than the 
blades; leaf blades ovate or ovate-elliptic, 3-8 cm. long, narrowed 
to an obtuse apex, or the uppermost leaves often acute, green and 
sparsely tomentulose above, paler beneath and densely rufous- 
tomentose; umbels numerous, 10-25-flowered, the capillary pedicels 
glabrous or nearly so, 2-4 mm. long, or in fruit 5 mm.; perianth 
white, 2.5-3 mm. long, glabrous; anthocarp glabrous, suborbicular, 
5 mm. long and broad, rounded at the base, 3-winged. Neg. 3138. 

Cuzco: Lucumayo Valley, 1,800-3,600 meters, Cook & Gilbert 
1353.Huanuco: Vilcabamba, 1,800 meters, 5148. Cushi, 1,500 
meters, 454%- Junin: Carpapata, 2,700-3,200 meters, Killip & 
Smith 24416. Also in Ecuador and Bolivia. 

This is the most easily recognized of the Peruvian species, by 
its abundant, rusty pubescence. 

Colignonia scandens Benth. PL Hartw. 148. 1844. 

Plants herbaceous or suffrutescent, erect or subscandent, the 
stems 1-2 meters long, glabrous or nearly so; leaves often subverti- 
cillate, the small ones of the inflorescence frequently white, the 
petioles mostly shorter than the blades but sometimes equaling them; 
leaf blades elliptic to rhombic-ovate or oblong-ovate, 2-6 cm. long, 
rounded or very obtuse at the apex, sometimes acute at the base, 
sparsely ferruginous-puberulent beneath along the veins when 
young, in age glabrous or nearly so; umbels commonly 8-12-flowered, 
the filiform pedicels glabrous, 3-4 mm. long, or in age 6 mm. ; perianth 
green or greenish, 2.5-3 mm. long, glabrous; anthocarp suborbicular, 
4-5 mm. long and broad, glabrous, rounded at the base, 3-winged. 

Piura: Above Frias, 2,600-2,700 meters, Weberbauer 6407. Also 
in southern Ecuador. 

Colignonia Weberbaueri Heimerl, Bot. Jahrb. 42: 78. 1908. 

Plants herbaceous or suffrutescent, erect or clambering, some- 
times 2 meters long, the pale, hollow stems glabrous or nearly so; 
leaves chiefly verticillate and unequal, the slender petioles mostly 
shorter than the blades; leaf blades ovate or deltoid-ovate, 3-7 cm. 
long, narrowed to the obtuse or rounded or occasionally acute apex, 
acute to subcordate at the base, thin, slightly paler beneath, glabrous 
or nearly so; umbels 12-20-flowered, the slender, glabrous pedicels 
2-3 mm. long; perianth white, glabrous, 3-3.5 mm. long; stamens 
equaling or slightly exceeding the perianth lobes; anthocarp turbi- 



FLORA OF PERU 535 

nate-obpyramidal, 4 mm. long, glabrous, 5-angled, tapering to the 
base. Neg. 3139. 

Amazonas: Mito, 2,700 meters, 1564- Ancash: Below Ocros at 
Alnos, 2,400-2,900 meters, Weberbauer 2712 (fragm. of type from 
hb. Berol.). Between Chiquian and Tallenga, grassy thickets, 
3,000-3,600 meters, Weberbauer 2853 (according to Heimerl). 
Apurimac: Rio Pinco and Apurimac, 3,300-3,400 meters, Weberbauer 
5906. Cuzco: Hacienda Churo, Paucartambo Valley, 3,500 meters, 
Herrera 1400. Valley of the Apurimac, Herrera. Torontoy, Uru- 
bamba Valley, 2,400 meters, Bingham 731, 555. Lucumayo Valley, 
1,800-3,000 meters, Cook & Gilbert 1371. Marcapata, 3,200 meters, 
Weberbauer 7776. Paucartambo to Tres Cruces, 3,200-3,400 meters, 
Pennell 14141- Lima: Matucana, 411. Obrajillo, Wilkes Exped. 
Rio Blanco, 3,000-3,500 meters, open hillside, Kittip & Smith 21683. 
San Buenaventura, 2,700-2,800 meters, rocky canyon, Pennell 
14525. Rio Blanco, 450 meters, 714- Tacna: Carumas, Prov. 
Moquehua, Weberbauer 7285. Department uncertain: San Mateo, 
Isern 2246. 

Weberbauer states that the bracts usually are white on the upper 
surface but green beneath. The vernacular names are reported as 
"lluplunsha" (Mito) and "sachaparaquay" (Lucumayo Valley). 
This species is closely related to C. glomerata Griseb., of Bolivia and 
Argentina, differing chiefly, if at all, in the form of the fruit. It 
may be that some of the Peruvian specimens, with flowers only, 
really are referable to C. glomerata. 

6. ALLIONIA L. 

Annual or perennial herbs; leaves opposite, petiolate, those 
of a pair unequal, entire or sinuate; flowers perfect, in axillary 
clusters of 3, each subtended by a broad, green bract, the bracts 
cucullate, enclosing the fruit; perianth corolla-like, the limb oblique, 
4-5-lobed; stamens 4-7, exserted; anthocarp coriaceous, obovoid 
or oval, strongly compressed, 3-costate or cristate on the inner 
surface, the outer surface bearing 2 parallel, longitudinal rows of 
stipitate glands, the thin margins dentate or entire, inflexed. 

Allionia incarnata L. Syst. Nat. ed. 10. 890. 1759. Wedelia 
incarnata Kuntze, Rev. Gen. 2: 533. 1891. Wedeliella incarnata 
Cockerell, Torreya 9: 167. 1909. 

Perennial from a slender or thick, woody root, the stems short 
or elongate, glandular-puberulent or viscid-villous; petioles 5-20 mm. 
long; leaf blades mostly oval to oblong, 1-6 cm. long, unequal at 



536 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

the base and subcordate or rounded, obtuse or rounded at the apex, 
paler beneath and glandular-puberulent or viscid- villous; involucres 
on slender peduncles 5 mm. long or less, the bracts obovate-orbicular, 
5-8 mm. long, rounded or obtuse at the apex; perianth 7-15 mm. 
long, purple-red; anthocarp 3-4.5 mm. long, the inner side 3-costate, 
the margins with 3-5 low, broad teeth, or the teeth slender and more 
numerous. Negs. 3088, 3087, 3089. 

Arequipa: Tingo, open, rocky slopes, 2,200 meters, Pennell 
13133. lea (?): Mountains between valley of Rio de las Trancas 
and "Llaxwa," 1,200-1,800 meters, Hrdlicka. Junin: Huanuco, 
stony slopes, 2,100 meters, 3249. Lima: Chosica, rocky, sandy 
slope, 900 meters, 2869. Below Cocachacra, 1,300 meters, Weber- 
bauer 5267. Mountains near Chosica, 1,100 meters, Weberbauer 
5331. Piura: Tablazo, north of Parinas, mostly in calcareous soil, 
Haught F132, 226. Tacna: Torata, 2,200 meters, Weberbauer 
7417; at 1,600 meters, Weberbauer 7417a. Without definite locality: 
Weberbauer 6506. Cheuchin, Ruiz & Pavon 12-66. Also found 
in Chile and Argentina, and ranging northward to southwestern 
United States. 

The Peruvian specimens are variable as to amount and character 
of pubescence, and toothing of the margins of the anthocarp. 

7. BOERHAAVIA L. 

Annual or perennial herbs, sometimes somewhat suffrutescent 
and subscandent, the stems often with viscous areas in the internodes; 
leaves opposite, those of a pair often unequal, petioled, the blades 
entire or sinuate; flowers perfect, small, variously arranged, bracteate, 
the bracts distinct and usually very small; perianth corolla-like, 
campanulate, nearly rotate, or funnelform, the limb shallowly 
5-lobate; stamens 1-5, exserted or included, unequal; anthocarp 
cylindric to obovoid or obpyramidal, terete or 3-10-angulate, some- 
times 3-5-winged, glabrous or pubescent, often furnished with 
stipitate glands. 

Flowers in racemes; anthocarp winged B. verbenacea. 

Flowers not in racemes; anthocarp not winged. 
Flowers in cymes; anthocarp truncate at the apex, glabrous; 

perianth white or pink; plants annual, erect B. erecta. 

Flowers in heads or glomerules; anthocarp not truncate, usually 
pubescent; perianth dark red; plants perennial, usually pro- 
cumbent or prostrate. 



FLORA OF PERU 537 

Branches of the inflorescence glabrous; flowers mostly in clusters 
of 2-4 B. coccinea. 

Branches of the inflorescence puberulent or glandular-puberu- 
lent; flowers chiefly in many-flowered heads . . .B. caribaea. 

Boerhaavia caribaea Jacq. Obs. Bot. 4: 5. 1771. B. polymorpha 
Rich. Act. Soc. Hist. Nat. Par. 1: 185. 1792. B. hirsuta Willd. 
Phytogr. 1. 1794. B. viscosa Lag. & Rodr. Anal. Cienc. Nat. 4: 256. 
1801. B. patula Domb. ex Vahl, Enum. 1: 287. 1804. 

Plants perennial from a thick, woody root, the stems decumbent 
or prostrate, below viscid-puberulent and often hirsute or villous, 
above densely glandular-puberulent; leaves petiolate, thick, the 
blades suborbicular to oval or oblong, 1.5-5.5 cm. long, truncate to 
rounded at the base, rounded to obtuse at the apex, glabrous or 
often puberulent or densely villous or hirsute; flower heads axillary 
or forming open cymes; perianth 2 mm. broad, puberulent or glan- 
dular-puberulent; stamens 1-3; anthocarp narrowly obovoid, 2.5-3 
mm. long, 5-sulcate, densely glandular-puberulent. Neg. 27751. 

Ancash: Chosica, on sandy gravel, 900 meters, 5^1. Cuzco: 
Santa Ana, 900 meters, Cook & Gilbert 1471. Huanuco: Huanuco, 
Ruiz & Pavdn B28. Lima: Santa Clara, Rose 18627. Piura: 
Mainly along watercourses, between the Amotape Hills and the 
sea, Haught F115, 269. Tacna: Hills southeast of Moquehua, 
only one plant seen, 1,600 meters, Weberbauer 7460. Without 
locality, Weberbauer 6461. 

The name "raiz de China" is reported from Santa Ana. 

Boerhaavia coccinea Mill. Gard. Diet. ed. 8. Boerhaavia No. 4. 
1768. B. paniculata Rich. Act. Soc. Hist. Nat. Par. 1: 105. 1792. 
B. adscendens Willd. Sp. PI. 1: 19. 1797. B. decumbens Vahl, Enum. 
PI. 1: 284. 1804. 

Plants perennial from an often fusiform and fleshy root, the 
stems ascending or procumbent, 1 meter long or less, minutely 
puberulent below or often villous, especially at the nodes, glabrous 
above; leaves somewhat fleshy, the blades rhombic-orbicular to 
rhombic-ovate or oval, 2-5.5 cm. long, subcordate to broadly rounded 
at the base, rounded or obtuse at the apex, pale beneath and not 
punctate, glabrous or sparsely puberulent, sometimes villous along 
the veins; peduncles 3-10 mm. long; perianth 2 mm. broad, minutely 
glandular-puberulent; stamens 2; anthocarp narrowly obovoid, 3-4 
mm. long, 5-sulcate, densely glandular-puberulent or glandular- 
pilose. 



538 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cuzco: Santa Ana, 900 meters, Cook & Gilbert 1471. Hacienda 
Chancamayo, Diehl 2453. Huanuco: Huanuco, dry gulch slopes 
above river valley, 2,100 meters, 2329. Loreto: Paranapura, lower 
Rio Huallaga, 200 meters, Williams 4591. Yurimaguas, Williams 
4086. San Martin: Juan Guerra, 720 meters, Williams 6867. 
Widely distributed as a weed in tropical America. 

Boerhaavia coccinea Mill. var. leiocarpa (Heimerl) Standl. 
Field Mus. Bot. 11: 108. 1931. B. paniculata f. leiocarpa Heimerl, 
Oesterr. Bot. Zeitschr. 56: 252. 1906. 

Anthocarp glabrous; otherwise as in the typical form of the 
species. 

Huanuco: Ambo, steep, clay slopes, 2,100 meters, 3167. Extend- 
ing to Colombia and Argentina. 

Boerhaavia erecta L. Sp. PI. 3. 1753. 

Plants annual, usually erect, 1 meter high or less, much branched 
from the base, the slender branches finely puberulent below, the 
middle internodes often with brown, viscous bands, the upper ones 
glabrous or minutely puberulent; leaves slender-petiolate, the blades 
broadly ovate-rhombic or deltoid-ovate to oval or oblong, 2-6 cm. 
long, truncate to rounded at the base, rounded or obtuse or rarely 
acute at the apex, pale beneath and usually brown-punctate, glabrous 
or sparsely puberulent; flowers on pedicels 1-5 mm. long; perianth 
1-1.5 mm. long, glabrous; stamens 2-3; anthocarp narrowly ob- 
pyramidal, 3-3.5 mm. long, glabrous, 5-angled. 

Piura: Parinas Valley, Haught 222, F131. Between Piura and 
Nomala, 100-250 meters, Weberbauer 5954- Widely distributed as a 
weed in tropical America. 

Boerhaavia verbenacea Killip, Journ. Wash. Acad. Sci. 16: 
567. 1926. 

An erect or ascending annual, 1 meter high or less, the stems 
puberulent and more or less viscid below, glabrous above; leaves 
long-petiolate, rhombic-ovate to lance-oblong, mostly 2-4.5 cm. 
long, broadly rounded to acutish at the apex, commonly rounded 
or truncate at the base, pale beneath, often sparsely black-punctate, 
puberulent or glabrate; racemes 3-7 cm. long, paniculate, the flowers 
very shortly pedicellate or subsessile, the bracts large, persistent; 
perianth 1.5-2 mm. long, white or pale pink; anthocarp glabrous, 
3 mm. long, broadly 5-winged. Neg. 3083. 



FLORA OF PERU 539 

Piura: Talara, Prov. Paita, Haught 8 (type). Parinas Valley, 
very abundant after rains, Haught F133, 229. Between Piura and 
Nomala, 100-250 meters, Weberbauer 5933. 

The species is noteworthy because it is the only one with racemose 
inflorescence that is known from South America. 

8. COMMICARPUS Standl. 

Perennial herbs or shrubs, usually reclining or more or less 
scandent; leaves opposite, petiolate, the broad blades fleshy, entire 
or undulate; flowers perfect, umbellate or verticillate, pedicellate; 
perianth funnelform or campanulate, corolla-like, usually with a 
distinct tube, the limb shallowly 5-lobate; stamens 2-5, exserted; 
anthocarp cylindric-fusiform, finely costate vertically, bearing num- 
erous large glands. 

Gommicarpus tuberosus (Lam.) Standl. Contr. U. S. Nat. 
Herb. 18: 101. 1916. Boerhaavia tuberosa Lam. 111. Gen. 1: 10. 
1791. B. excelsa Willd. Phytogr. 1. 1794. B. litoralis HBK. Nov. 
Gen. & Sp. 2: 216. 1817. 

Plants much branched, suberect or scandent, the stems sometimes 
2 meters long, pale, glabrous except at the nodes; leaves thick and 
fleshy, long-petiolate, the blades broadly ovate or deltoid-ovate, 
3.5-5.5 cm. long, acutish to short-acuminate, often subcordate at 
the base, glabrous or nearly so; flowers umbellate, the numerous 
umbels 4-7-flowered, the pedicels long and slender; perianth purple 
or purplish, 8-10 mm. long, glabrous or nearly so; anthocarp linear- 
clavate, 6.5 mm. long, bearing a few stalked glands near the apex. 

Lima: Chosica, river valley among large rocks, 900 meters, 499. 
Near Corcona, Rose 18676. Piura: Talara, Prov. Paita, Haught 100. 
Also in the Galapagos Islands. 

This species usually has been confused with Boerhaavia scandens 
L., which in South America is known only from northern Colombia 
and Venezuela. B. tuberosa was described from Peru, and B. litoralis 
from Patibilca, Peru. 

9. MIRABILIS L. 

Perennial herbs, erect or procumbent, often viscid-pubescent, 
the branches commonly swollen at the nodes; leaves opposite, petio- 
late or sessile, the blades entire or undulate; flowers perfect, involu- 
crate, the involucre 1-several-flowered, 5-lobate, in fruit often accres- 
cent and becoming almost rotate; perianth corolla-like, tubular to 
campanulate, often oblique, the limb 5-lobed, the perianth soon 



540 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

withering and deciduous; stamens 3-5, unequal, usually exserted; 
anthocarp coriaceous, smooth or 5-angled or 5-sulcate, often con- 
stricted at the base, mucilaginous when wet. 

Perianth large, 3-5.5 cm. long M. Jalapa. 

Perianth smaller, less than 2 cm. long. 

Anthocarp constricted at the base, conspicuously 5-sulcate; invo- 
lucre strongly accrescent in age, becoming almost rotate and 
entire, membranaceous, and reticulate- veined. 
Anthocarp glabrous; leaf blades deeply cordate at the base. 

M. viscosa. 

Anthocarp pubescent; leaf blades obtuse or rounded at the base. 

M. ovata. 

Anthocarp not constricted at the base, smooth or nearly so; 

involucre not at all or scarcely accrescent, unchanged in age. 

Perianth tubular, about 15 mm. long, with only very short, 

erect lobes M. Weberbaueri. 

Perianth campanulate or short-funnelform, with a broad limb, 

in most species less than 15 mm. long. 

Leaves thin, mostly truncate or subcordate at the base, com- 
monly acuminate or long-acuminate, with long and 
slender petioles. Stamens 3 ; anthocarp shorter than the 

involucre M. prostrata. 

Leaves relatively thick, mostly rounded to acute at the base, 

sometimes truncate, broadly rounded to obtuse at the 

apex, or the uppermost leaves acute, the petioles usually 

stout and often very short. 

Stamens 5; anthocarp subglobose, much exceeding the 

involucre M. intercedens. 

Stamens normally 3; anthocarp shorter than the involucre. 
Involucre 6.5-10 mm. long; stems thick and somewhat 
fistulose, glabrous below the inflorescence. 

M. elegans. 
Involucre 3.5-5 mm. long; stems not fistulose, glabrous 

or pubescent. 
Involucres very shortly pedunculate, in dense clusters; 

stamens usually included M. expansa. 

Involucres on slender peduncles 4-7 mm. long, forming 
lax, open cymes; stamens exserted. 

M. campanulata. 



FLORA OF PERU 541 

Mirabilis campanulata Heimerl, Bot. Jahrb. 42: 75. 1908. 
Allionia campanulata Standl. Contr. U. S. Nat. Herb. 18: 101. 1916. 

Plants procumbent, the branches rather slender, glabrous below 
or hirtulous at the nodes, in the inflorescence glandular-pilosulous; 
leaves rather thick, the petioles stout and mostly very short, the 
blades ovate to oblong-ovate, 7 cm. long or less, rounded to acute at 
the base, narrowed to the obtuse or acutish apex, glabrous or nearly 
so; inflorescence broad and open, dichotomous; involucres slender- 
pedunculate, narrowly campanulate, densely glandular-puberulent 
or viscid-puberulent, 3.5-4 mm. long, the lobes lance-triangular, 
subobtuse or acutish, equaling the tube; perianth red-purple, about 
1.5 cm. broad; anthocarp 4 mm. long, dark brown, glabrous, nearly 
smooth. Neg. 3076. 

Ancash: Below Pampa Romas, between Samanco and Caraz, 
open thickets, 2,000-2,200 meters, Weberbauer 3198 (photo, and 
fragm. of type ex hb. Berol.). Lima: Matucana, in decomposed 
granite, 2,400 meters, 297; dry, rocky slope, 661. 

Mirabilis elegans (Choisy) Heimerl in Engl. & Prantl, Nat. 
Pflanzenfam. 3: Abt. Ib: 24. 1889. Oxybaphus elegans Choisy in 
DC. Prodr. 13, pt. 2: 431. 1849. Allionia elegans Kuntze, Rev. 
Gen. 2: 533. 1891. M. arenaria Heimerl, Bot. Jahrb. 42: 74. 1908. 
A. arenaria Standl. Contr. U. S. Nat. Herb. 18: 101. 1916. 

Plants apparently erect and 1 meter high or less, the branches 
stout and fistulose, glabrous below the inflorescence, there glandular- 
pubescent; leaves on very stout and usually short petioles, the upper- 
most often sessile or nearly so, the blades thick, rounded-oval to 
ovate-rounded, deltoid-ovate, or rhombic-ovate, 4-9 cm. long, 
truncate or rounded at the base and more or less oblique, rounded to 
acutish at the apex, glabrous or nearly so; inflorescence large and 
open, dichotomous, the branches separating usually at a wide angle, 
the leaves of the inflorescence much reduced; involucres narrow, 
6.5-10 mm. long, densely viscid-villosulous, 1-flowered, the lobes 
lance-triangular, about equaling the tube; perianth purple-red, 15 
mm. long or less; stamens exserted; anthocarp ellipsoid, 5 mm. long, 
brown, obscurely costate. Neg. 7409. 

Arequipa: Mollendo, sandy soil, 200 meters, Weberbauer 1521 
(photo, of type of M. arenaria ex hb. Berol.). Tiabaya, open, rocky 
hills, 2,100 meters, Pennell 13080. Also in Chile. 

Mirabilis expansa (R. & P.) Standl. Field Mus. Bot. 11: 113. 
1931. Calyxhymenia expansa R. & P. Fl. 1: 45. pi. 75, f. a. 1798. 



542 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Oxybaphus expansus Vahl, Enum. PI. 2: 41. 1806. Allionia expansa 
Kuntze, Rev. Gen. 2: 533. 1891. 

A coarse herb, suberect or even subscandent, the stems some- 
times 2.5 meters long, very sparsely puberulent or villosulous or 
almost glabrous; leaves rather thick, on stout and usually short 
petioles, the blades ovate-rounded to rhombic-ovate, 2.5-5 cm. long, 
truncate to acute at the base and often abruptly decurrent, obtuse 
to rounded at the apex, or the uppermost leaves acute, sparsely 
puberulent or pilosulous or glabrate; inflorescence cymose-paniculate, 
rather small, the involucres mostly in small, dense clusters; involucre 
1-flowered, 4-5 mm. long, viscid-villosulous, the lobes ovate-triangu- 
lar, much shorter than the tube; perianth red-purple, 6 mm. long; 
stamens 3, usually not exserted; anthocarp ellipsoid, 3 mm. long, 
dark olivaceous, glabrous, almost smooth. 

Cuzco: San Sebastian, rocky slope of canyon, 3,300 meters, 
Pennell 13624. Junin: Tarma, 3,000-3,200 meters, Killip & Smith 
21854. Lima: Described from "collibus aridis Chancay et Limae 
ad Amancaes," Ruiz & Pavdn. Without locality, Ruiz & Pavdn 
12-37, probably type material. Rio Blanco, loose soil, 3,600 meters, 
715. Lima, Rose 18592. Rio Blanco, open hillside, 3,000-3,500 
meters, Killip & Smith 21664- Ranging to Venezuela and Chile. 

Mirabilis intercedens Heimerl, Bot. Jahrb. 54: Beibl. 117: 36. 
1916. 

An ascending herb 1 meter high or less, the branches rather stout, 
minutely puberulent below or often glabrate; leaves short-petiolate, 
the upper sometimes sessile, broadly rhombic-ovate to deltoid-ovate 
or the upper oblong-ovate, mostly 4-6 cm. long, rounded or truncate 
at the base and usually somewhat oblique, sometimes abruptly 
decurrent or even acute at the base, obtuse at the apex, rather fleshy, 
glabrous or nearly so; panicles dichotomous, usually densely leafy; 
involucres pedunculate, villosulous or puberulent and viscid, 4-5 
mm. long, 1-flowered, the lobes ovate-lanceolate, acutish, equaling 
the tube; perianth purple-red, 12-16 mm. long, glabrous, the limb 
15-22 mm. wide; stamens 5, exserted; anthocarp globose, black, 
glabrous, as much as 13 mm. in diameter. Neg. 5270. 

Lima: San Bartolome", between Lima and Oroya, open thickets, 
1,400-1,600 meters, Weberbauer 5270, type collection. Matucana, 
steep, rocky, northern slope, 2,400 meters, 145; loose soil, southeast 
slope, 2927. Tacna: Mountains between Moquehua and Torata, 
stony places, 2,000 meters, Weberbauer 7435. Carumas, 2,900 
meters, Weberbauer 7268. 



FLORA OF PERU 543 

This species may be recognized readily by the fruit, altogether 
unlike that of any other species. The perianth does not increase 
in size after anthesis, and it finally is scarcely noticeable at the base 
of the fruit. In dried specimens all the fruits are pressed flat, indicat- 
ing that they must be very soft in the fresh state, and it may well 
be that the mature and hardened ones are still larger than is indicated 
in the description. 

Mirabilis Jalapa L. Sp. PI. 177. 1753. M. odorata L. Cent. 
PI. 1: 7. 1755. M. dichotoma L. Sp. PL ed. 2. 252. 1762. 

A stout, bushy-branched perennial 1 meter high or less, the 
branches glabrous, puberulent, or rarely short-villous; leaves slender- 
petiolate, the blades ovate-deltoid to lance-oblong, 5-14 cm. long, 
subcordate to rounded at the base, acute to acuminate at the apex, 
glabrous or rarely puberulent, usually ciliate; flowers in dense 
clusters at the ends of the branches; involucre short-pedunculate 
or subsessile, 7-15 mm. long, glabrous, puberulent, or short-villous, 
the lobes linear-lanceolate to lance-ovate, longer than the tube, 
acute or acuminate; perianth 3-5.5 cm. long, purplish red or white, 
yellow, or variegated; stamens 5; anthocarp 7-9 mm. long, 5-angled, 
verrucose or rugose, dark brown or black, glabrous or puberulent. 

Cuzco: Alturas del Valle de Lares, Diehl 2474- Junin: Posuso, 
600 meters, a common weed, 4659. Rio Pinedo, north of La Merced, 
700-900 meters, Kittip & Smith 23599. La Merced, in thickets, 
700 meters, Killip & Smith 23818. Loreto: Iquitos, Williams 8198, 
3599. La Victoria, Williams 3108. Puerto Arturo, Yurimaguas, 
Williams 5010, 5229. Mishuyacu, near Iquitos, 100 meters, in clear- 
ing, Klug 67. Fortaleza, Yurimaguas, Williams 4495. San Martin: 
Tarapoto, Williams 6501. 

This plant, the four-o'clock or marvel of Peru, is known in 
cultivation in most tropical regions as well as in temperate countries, 
and it is one of the most common garden flowers of the United States. 
So far as known, it does not grow wild anywhere, although often it 
escapes from cultivation into waste ground, and may persist there 
if not too hardly crowded by native plants. The early botanical 
writers assumed the plant to be a native of Peru, but it is more 
probable that it is of Mexican origin. The following vernacular 
names are recorded from Peru: "Buenas tardes," "clavanilla" 
(clavellina?). The kernels of the seeds are said to be used some- 
times in Peru as a substitute for soap. The sweet-scented flowers 
open in the evening and close early the following morning. 



544 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Mirabilis ovata (R. & P.) Meigen, Bot. Jahrb. Engl. 18: 
436. 1894. Calyxhymenia ovata R. & P. Fl. 1: 45. pi. 75, /. 6. 
1798. Oxybaphus ovatus Vahl, Enum. PI. 2: 41. 1806. 0. chilensis 
Steud. Norn. Bot. ed. 2. 2: 213. 1841. 0. campestris Griseb. PL 
Lorentz. 39. 1874. Allionia campestris Kuntze, Rev. Gen. 2: 533. 
1891. A. ovata Standl. Contr. U. S. Nat. Herb. 13: 409. 1911. 

Plants coarse and stout, erect, 1 meter high or less, the branches 
densely viscid- villous; leaves on stout petioles, the blades thick and 
fleshy, ovate or deltoid-ovate, 2-3 cm. long and larger, rounded or 
subtruncate at the base, obtuse, rather densely viscid-pilose; in- 
florescence large, lax and open, the involucres short-pedunculate, 
densely viscid-villous, in fruit 2 cm. broad or more, the lobes short, 
obtuse or rounded; perianth purple-red, about 1 cm. long, sparsely 
villosulous; stamens 3, short-exserted; anthocarp 5 mm. long, con- 
spicuously 5-angled, minutely pubescent. Neg. 18240. 

Lima: Described from the hills of Huaura and the mountains 
of Lachay, Pasamayo, and Jequan, Ruiz & Pavon. Occurring also 
in Argentina and Chile. 

No Peruvian specimens have been seen by the writer, but the 
original description and figure agree well with material from Chile 
and Argentina. 

Mirabilis prostrata (R. & P.) Heimerl, Beitr. Syst. Nyctag. 
21. 1897. Calyxhymenia prostrata R. & P. Fl. 1: 46. pi. 75, /. c. 
1798. Oxybaphus prostratus Vahl, Enum. PL 2: 40. 1806. 0. 
micranthus Choisy in DC. Prodr. 13, pt. 2: 432. 1849. 

Plants much branched, erect or procumbent, said to be sometimes 
2 meters long, the slender branches puberulent, viscid-villosulous, 
or glabrate; leaves slender-petioled, thin, the uppermost short- 
petioled, the blades ovate or ovate-deltoid, often elongate-deltoid, 
mostly 3-7 cm. long, truncate or subcordate at the base, acute to 
long-acuminate, rarely obtuse, sparsely villosulous or puberulent 
or more often glabrate; inflorescence cymose-paniculate, usually 
large and open, often leaf y-bracted ; involucres 4-5 mm. long, 
narrow, densely viscid-villosulous, the lobes narrowly triangular, 
mostly acute, equaling or shorter than the tube; perianth purplish 
red or pink, 7-10 mm. long; stamens 3, included or exserted; antho- 
carp 3 mm. long, glabrous, fuscous, nearly smooth. Negs. 27754, 
3079. 

Cuzco: Ollantaitambo, 3,000 meters, Cook & Gilbert 415, 325, 357. 
Cuzco, 3,350 meters, Herrera 255. Junin: Huacachi, 1,950 meters, 



FLORA OF PERU 545 

sprawling at base of shrubby cliff, 4160. Lima: Viso, trailing on 
rocks along river, 2,700 meters, 769. Lima, Weberbauer 1624- 
San Geronimo, 150 meters, in rocks, 5892. Paita: Cerro Viento, 
Haught 96. Junin: Tarma, 3,000-3,200 meters, open hillside, Killip 
& Smith 21778, 21835. Tumbez: Mountains east of Hacienda 
Chicama, in deciduous bush wood, 600 meters, Weberbauer 7661. 
Without locality: Ruiz & Pavon 12-59. Ecuador, Bolivia, and 
Chile. 

Ruiz and Pavon report the vernacular name as "pegapega." 

Mirabilis viscosa Cav. Icon. 1: 13. pi. 19. 1791. Nyctago parvi- 
flora Salisb. Prodr. 57. 1796. Calyxhymenia viscosa R. & P. Fl. 
1: 46. 1798. Calymenia viscosa Pers. Syn. PL 1: 36. 1805. Vitmania 
viscosa Turra ex Steud. Nom. Bot. 140, in syn. 1821. Oxybaphus 
viscosus L'He"r. ex Choisy in DC. Prodr. 13, pt. 2: 430. 1849. Allionia 
viscosa Kuntze, Rev. Gen. 2: 533. 1891. 

A coarse herb 0.5-1.5 meters high or even larger, the stems some- 
times as much as 5 cm. thick, the branches densely viscid-pilose; 
leaves rather thick, long-petiolate, the blades broadly cordate-ovate 
or ovate-deltoid, 3-10 cm. long, usually cordate and abruptly short- 
decurrent at the base and acute to acuminate at the apex, short- 
villous or puberulent, especially beneath; inflorescence large, openly 
paniculate, the branches chiefly opposite, densely viscid-villous; 
involucres at anthesis 3-5 mm. long, in fruit 15-25 mm. broad, 
densely viscid-pilose; flowers solitary or rarely 2-3 in the involucre, 
the perianth 8-20 mm. long, purplish red, pink, or white, the limb 
15-25 mm. broad; stamens 3; anthocarp 5 mm. long, glabrous, 
densely covered with large, coarse tubercles. 

lea: Above Pisco, between Pampano and Huaytara, 1,300 meters, 
Weberbauer 5389. Huanuco: Huanuco, Ruiz & Pavdn 12-60. Type 
from Huanuco. Huanuco, 2,100 meters, in gulches on steep, dry 
slopes, 2036; on stony slopes in shale and gravel, 3208. Chaulki, 
Huanuco, Sawada P73. Lima: Open, rocky slopes, along Rio 
Chillon, near Viscas, 1,800-2,000 meters, Pennell 14459. Above 
San Bartolome", Prov. Huarochiri, 1,500-1,800 meters, Weberbauer 
5261. Colombia, Ecuador, and Mexico. 

Mirabilis Weberbaueri Heimerl, Bot. Jahrb. 42: 73. 1908. 

Plants apparently procumbent, the branches densely viscid- 
villosulous; leaves slender-petiolate, the blades broadly ovate, 6.5 cm. 
long or smaller, rounded to subcordate at the base, obtuse or rounded 
at the apex, viscid-villosulous; inflorescence cymose-paniculate, lax 



546 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and open, the involucres slender-pedunculate, densely glandular- 
puberulent, broadly campanulate, 6-7 mm. long, 1-flowered, the 
lobes triangular, acutish, slightly longer than the tube; perianth 
purplish, tubular, 15 mm. long, hirtulous along the nerves, very 
shortly 5-lobate; stamens 3, exserted; anthocarp 4-5 mm. long, 
brownish, glabrous. Neg. 3082. 

Ancash: Below Pampa Romas, between Samanco and Caraz, 
2,100-2,400 meters, Weberbauer 3185 (photo, and fragm. of type 
ex hb. Berol.). 

62. BATIDACEAE. Batis Family 

This family consists of a single species, widely distributed on 
tropical and subtropical shores of the New World, but the plant 
seems to be rare on the coast of Peru. Compare Weberbauer 
(p. 18, Part I of this work), whose remark implies rather that it is 
rarely collected. 

1. BATIS L. 

A smooth, sprawling, oppositely branched shrub with fleshy, 
narrow, nearly terete, opposite leaves and dioecious flowers in stro- 
biles solitary in the leaf axils; staminate strobiles sessile, the pistillate 
pedunculate. 

Batis maritima L. Syst. ed. 10. 1289. 1759. 

The plant is known from Hawaii, and on American shores it 
extends north to those of Florida and California. Illustrated, Mart. 
Fl. Bras. 3, pt. 3: pi 126. 

Lambayeque: Raimondi (det. Ulbrich). Tumbez: Coastal plain 
between Tumbez and Zarumilla, a small shrub in salt soil, Weberbauer 
7621. Piura: Negritos, just behind the beach, 25 miles north of 
Cabo Blanco, growing about 1 meter high, seen also near Negritos 
and 6 miles south of Paita, Haught 204- Zorritos, Haught 204- 

63. PHYTOLACCACEAE. Pokeweed Family 
Reference: H. Walter, Pflanzenreich IV. 83. 1909. 

In Peru, as elsewhere, this family is known for the high repute 
in which some of its members are held as medicinal plants, for which, 
however, there appears to be no scientific basis. Included here for 
convenience is the genus Achatocarpus Triana, the type genus of 
the family Achatocarpaceae Heimerl, a group which is academically, 
that is to say logically, separable; its near affinity is with the Aizoa- 
ceae (see Heimerl, Pflanzenfam. ed. 2. 16c: 174-178. 1934). 



FLORA OF PERU 547 

In addition to the following, Ercilla spicata (Bert.) Moq. (E. 
volubilis Juss.) has been repeatedly accredited to Peru on the basis 
of a Dombey specimen, in this instance, as suggested by Harms, 
Pflanzenfam., op. cit. 154, certainly mislabeled, as are so many Dom- 
bey plants. 

Inflorescence a simple raceme or spike (except Phytolacca thyrsiflora). 

Racemes 6-10-flowered, crowded, partly in clusters of 2 or 3; 

shrubs 1. Achatocarpus. 

Racemes mostly many-flowered, solitary; plants herbaceous or 

green, at least above. 

Flowers minute (about 1 mm. long) ; fruits rough ... 2. Microtea. 
Flowers often small but longer than 1 mm.; fruits smooth. 
Flowers remote, closely appressed to the rachis . . 3. Petiveria. 
Flowers often crowded, never appressed to the rachis. 
Fruits of solitary carpels. 
Sepals broad, widely spreading to reflexed. 

4. Trichostigma. 

Sepals often narrow, erect or suborbicular. 
Sepals 3-connate; pedicels short (2, rarely 3.5 mm. 

long) 6. Hilleria. 

Sepals all free; pedicels 3-6 mm. long. 

Fruit baccate; stamens 4 5. Rivina. 

Fruit dry; stamens 12-15 7. Schindleria. 

Fruit a ring of carpels 9. Phytolacca. 

Inflorescence paniculate (in Peruvian species). 

Fruit baccate; sepals pale, paper-like 8. Flueckigera. 

Fruit a samara; sepals herbaceous. 

Sepals 4, erect 10. Gallesia. 

Sepals 5, becoming reflexed 11. Seguieria. 

1. ACHATOGARPUS Triana 

A stout shrub with short, gray-barked branchlets, short-clustered 
racemes, and broadly oval leaves. Flowers unisexual. Stamens 
12-15, irregularly inserted. Fruit baccate. Regarding the position 
of this genus see remarks at the beginning of the family. 

Achatocarpus pubescens C. H. Wright, Kew Bull. 1906: 6. 
1906. A. mottis H. Walt. Pflanzenr. IV. 83: 136. 1909. 



548 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

A shrub 3-4 meters high, the young branches and leaves beneath 
densely pilose; petioles 5 mm. long; leaf blades usually rounded or 
obtuse at the apex, 10 cm. long and 4.5 cm. broad; racemes 6-10- 
flowered, pilose, the pedicels 2 mm. long. Illustrated, Pflanzenr. 
IV. 83: 135. 

Prov. Tumbez at 200 meters, Weberbauer 7693. Ecuador. 

Achatocarpus praecox Griseb. Goett. Abh. 24: 32. 1879. 
Similar to A. pubescens, but the branchlets shortly spinescent and 
the more lanceolate leaves only about half as large. Neg. 27741. 
Peru: Without locality (Pavori). Argentina. 

2. MICROTEA Sw. 

The Peruvian species slender annuals with short or long racemes 
of tiny flowers. Stamens 3-9, in one series. Carpels 3, minute. 

Microtea debilis Sw. Prodr. 53. 1788; 128. 

Weak and prostrate or ascending; leaves thin, usually somewhat 
obovate; racemes dense, in fruit to 3 cm. long; pedicels scarcely 1 
mm. long; sepals and stamens 5, about 0.5 mm. long; fruit tuber- 
culate or echinate. 

San Martin: Tarapoto, Raimondi (det. Heimerl); at 750 meters, 
Williams 5573. Loreto: Maucallacta, 200 meters, in clearing, 
Klug 3959. Yurimaguas, 200 meters, Williams 4081; in waste 
places, Killip & Smith 28215. Rio Putumayo, Klug 1647. Yuri- 
maguas, Poeppig 2303. Rio Acre: Vie 9363. Puno: Sangaban, 
Lechler 2443. Widely distributed in tropical America. 

Microtea maypurensis (HBK.) G. Don in Loud. Hort. Brit, 
ed. 2. 98. 1839; 132. Ancistrocarpus maypurensis HBK. Nov. Gen. 
& Sp. 2: 186. 1817. 

Similar to M. debilis but often erect, the leaves narrower, to 
linear-lanceolate, and the racemes lax, the slender pedicels 1.5 mm. 
long or longer; stamens 6-8, nearly 1 mm. long. Illustrated, 
Pflanzenreich IV. 83: 133. 

Tumbez: Prov. Tumbez, 200 meters, Weberbauer 7729. Puno: 
Sangaban (Lechler}. South America. 

3. PETIVERIA L. 

Well characterized by the long, slender, spike-like racemes of 
remote flowers that are ascending or closely appressed to the rachis. 
Stamens 4-8. Sepals free. Fruit elongate, with 4-6 setae at the 



FLORA OF PERU 549 

apex. Standley, Contr. U. S. Nat. Herb. 23: 265, has compiled an 
amazingly long list of reputed medicinal uses; its reputation as a 
medicinal plant apparently results from the garlic odor it emits 
when crushed. 

Petiveria alliacea L. Sp. PL 342. 1753; 118. 

Slender, usually somewhat woody below; petioles to 6 mm. long; 
leaves oblong-elliptic, nearly glabrous, about 10 cm. long and half 
as broad; racemes erect, axillary and terminal, exceeding the leaves; 
pedicels rarely 1 mm. long; fruit to 8 mm. long, with 4 or rarely 5 
setae. P. tetrandra Gomez (P. alliacea var. tetandra Hauman- 
Merck, Anal. Mus. Nac. Buenos Aires 24: 501, 513, 1913) of Brazil 
is a form only that has fruit with 6 setae. According to Raimondi, 
the plant has diuretic properties. Illustrated, Pflanzenreich IV. 
83: 119. 

Ayacucho: Prov. La Mar, Raimondi. Aina, 750-1,000 meters, in 
clearing, Killip & Smith 22828. Cajamarca: Jae"n, 1,000 meters, 
Weberbauer 6197. Cuzco: Prov. Convencion, Raimondi. Quebrada 
Versalla, Diehl 2499a.Loreto: Iquitos, Williams 3534, 7913. 
Puerto Arturo, 200 meters, Williams 5221, 5183. Pinto-cocha, 
Williams 823. Mishuyacu, 100 meters, in clearing, Klug 439, 946. 
Yurimaguas, 200 meters, Williams 4652. San Martin: Tarapoto, 
800 meters, Williams 6901, 5762. Juanjui, 400 meters, Klug 3833. 
Tumbez: Raimondi. South America to southern United States. 
"Chanviro," "mucura." 

4. TRICHOSTIGMA A. Rich. 

Villamilla R. & P. Fl. pi. 402, ex Moq. in DC. Prodr. 13, pt. 2: 
10. 1849. 

This often clambering shrub is similar to Rivina, but the stamens 
are 8 to many and the stigma is sessile and penicillate. 
Petioles stout, 0.5-3 cm. long, more or less pubescent . . T. peruvianum. 
Petioles slender, 2-4 cm. long, glabrous T. octandrum. 

Trichostigma octandrum (L.) H. Walt. Pflanzenr. IV. 83: 
109. 1909. Rivina octandra L. Cent. PI. 2: 9. 1756. Villamilla 
octandra Hook. f. in Benth. & Hook. f. Gen. 3: 81. 1880. 

Scandent, the elongate branches pale verruculose-lenticellate; 
petioles to 2 cm. long; leaf blades chartaceous, glabrous, about 10 
cm. long and 4.5 cm. wide; racemes of white or purple flowers spread- 
ing, often to 10 cm. long and 1 cm. thick; pedicels to 6 mm. long; 



550 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

sepals green, ovate, 4 mm. long, little enlarged in fruit; fruit baccate, 
black. Illustrated, Pflanzenreich IV. 83: 109. 

Loreto: Mainas (Poeppig 2404)- San Seidro, Maranon, Tess- 
mann 4992. Middle Rio Santiago, Tessmann 4563. Yarina-cocha, 
Tessmann 3395 (det. Schmidt). Iquitos, in forest, 120 meters, Wil- 
liams 7910, 8045. Fortaleza, 200 meters, edge of forest, Williams 
4275. Rio Nanay, Williams 540. San Martin: Juan Guerra, 720 
meters, in forest, Williams 6854- Argentina, Bolivia, and the 
Amazon region to Mexico and the West Indies. 

Trichostigma peruvianum (Moq.) H. Walt. Pflanzenr. IV. 
83: 111. 1909. Rivina peruviana Moq. in DC. Prodr. 13, pt. 2: 10. 
1849. Villamilla peruviana Hook. f. in Benth. & Hook. f. Gen. 3: 81. 
1880. V. tinctoria R. & P. Fl. pi. 402, ex Moq. loc. cit. 

Scarcely distinguishable from the preceding except by the 
pubescent young branchlets, often subcordate leaves, and larger 
inflorescence of dark brown flowers, this sometimes over 30 cm. 
long and 15 cm. wide. The fruit is unknown. Neg. 29487. 

San Martin: Tarapoto, Williams 6953 (det. Schmidt). Ama- 
zonas: Moyobamba, 1,100 meters, Weberbauer 4555 (type); (Ma- 
thews 1455). Junin: Chanchamayo, Raimondi. Loreto: Balsa- 
puerto, Klug 2976; Killip & Smith 28690. Vochiza, Ruiz & Pawn. 
Between Yurimaguas and Balsapuerto, 110 meters, Killip & Smith 
28344; a shrub of 1.5-2.5 meters, the flowers white. Soledad, 110 
meters, in forest, Killip & Smith 29696; a tree of 3-4.5 meters; 
calyx pink. 

5. RIVINA L. 

In aspect like Hilleria, but the sepals free and the fruit baccate. 
Stamens 4. Style short, the stigma capitate. 

Rivina humilis L. Sp. PI. 121. 1753; 102. 

A somewhat woody plant with slender, dichotomous branchlets; 
leaves ovate to elliptic, membranaceous, often lightly pubescent on 
the nerves, 7.5 cm. long and 4 cm. broad or smaller or larger; inflo- 
rescence racemose, little or not exceeding the leaves; sepals 2 mm. 
long, spreading and enlarged in fruit; fruit red or orange, to 4 mm. 
thick. The glabrous form is var. glabra L. The plant has spread 
to Madagascar, Australia, and elsewhere as a weed, and when eaten 
by cows is said to impart to the milk a disagreeable odor. The 
English gardener Philip Miller recorded the placing of the berries 
in water for cut flowers, in order to color the blossoms. The plant 



FLORA OF PERU 551 

was already in cultivation in Europe in 1699. Illustrated, Pflan- 
zenreich IV. 83: 103. 

Cajamarca: Jae"n, Raimondi. Junin: Chanchamayo, Raimondi. 
Tropical and subtropical America. 

6. HILLERIA Veil. 

Herbaceous or somewhat suffrutescent, and in aspect not unlike 
Phytolacca, but often distinctive in the herbarium by the nearly 
black dried inflorescences. Leaves usually slightly pubescent. 
Flowers perfect. Carpels solitary. Perianth oblique, of 3 sepals. 

Stamens always 4; style shorter than the ovary or none. 

Stigma sessile H. latifolia. 

Stigma evident H . secunda. 

Stamens usually 5-9; style equaling the ovary. 

Stamens 5-7; style elongate H. Meziana. 

Stamens 8-13; stigma sessile H. longifolia. 

Hilleria latifolia (Lam.) H. Walt. Pflanzenr. IV. 83: 81. 1909. 
Rivina latifolia Lam. Illustr. 1: 324. 1791. 

Erect, suffrutescent, often 1.5 meters high, with slender branch- 
lets, the younger green and pilose; petioles to 4 cm. long; leaf blades 
elliptic-ovate, thin, finally glabrate or pilose on the nerves beneath, 
to nearly 9 cm. long and half as broad; racemes dense, erect, scarcely 
or little exceeding the leaves, to 14 cm. long and 9 mm. thick; pedicels 
2 mm. long; sepals 1.8 mm. wide, in fruit enlarged, papery, reticu- 
late-veined; stigma sessile, slightly penicillate. Illustrated, Pflan- 
zenreich IV. 83: 81. 

San Martin: Zepelacio, 1,200-1,600 meters, King 3438. San 
Roque, 1,400 meters, Williams 7611. Tarapoto, Williams 6884, 
5757. Near Moyobamba, King 3438.Loreto: Fortaleza, 200 
meters, edge of forest, Williams 4274, 4432. Santa Rosa, 135 
meters, Killip & Smith 28904. Yurimaguas, Poeppig 2163b. Yarina- 
cocha, Tessmann 3408. Rio Acre: Ule 9362 (det. Pilger). Cuzco: 
Alto Rio Urubamba, Diehl 2423a. South America; Africa. 

Hilleria longifolia (H. Walt.) Heimerl, Pflanzenfam. ed. 2. 16c: 
151. 1934. H. latifolia (Lam.) H. Walt. var. longifolia H. Walt. 
Pflanzenr. IV. 83: 82. 1909. 

Leaves subequal; inflorescence about 20 cm. long; stamens 
numerous. Otherwise like H . latifolia. 

Huanuco: Rio Chinchao, Poeppig 1541. 



552 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Hilleria Meziana H. Walt. Pflanzenr. IV. 83: 83. 1909. Mohla- 
na Meziana H. Walt. Bot. Jahrb. 37: Beibl. 85: 25. 1906. 

In general similar to other species, but the mature leaves charta- 
ceous, somewhat pubescent on the margins, and rather prominently 
reticulate- veined beneath; stamens 4-7, 4 alternate with the sepals; 
style cylindric-filiform, slightly curved, equaling the ovary, the 
stigma penicillate. Neg. 5723. 

San Martin: Near Tarapoto, Ule 6500 (type). Morales, in forest, 
Williams 5668. 

Hilleria secunda (R. & P.) H. Walt. Pflanzenr. IV. 83: 82. 
1909. Rivina secunda R. & P. Fl. 1: 65. 1794. R. inaequalis Hook. 
Icon. pi. 130. 1837. 

Very similar to H . latifolia, but the costa only obscurely mucronu- 
late and the fleshy style about 0.5 mm. long with a scarcely penicillate 
stigma. Neg. 27746. 

Huanuco: Rio Huallaga Canyon, 1,200 meters, 1+221. Posuso, 
Vitoc, Ruiz 211 (type). San Martin: Rumizapa, Williams 6753. 
San Roque, 1,400 meters, Williams 7489. Tarapoto, Spruce 4306. 
Alto Rio Huallaga, Williams 6709. Cajamarca: Jae*n, 1,000 meters, 
Weberbauer 6194. Tumbez: Raimondi (det. Heimerl). Cuzco: Prov. 
Convention, Raimondi (det. Heimerl). Junin: La Merced, 600 
meters, 5310. Chanchamayo, Isern 2369, 2066. South America. 

7. SCHINDLERIA H. Walt. 

Shrubs, or at least woody below, resembling Rivina but with 
12-15 irregularly disposed stamens and a dry fruit. 

Plant drying black S. glabra. 

Plant drying yellowish green S. Weberbaueri. 

Schindleria glabra H. Walt. Bot. Jahrb. 36: Beibl. 85: 24. 1906. 

Erect, glabrous throughout; leaves nearly ovate-elliptic, narrowly 
acuminate, about 15 cm. long, 6 cm. wide, the petioles to 5.5 cm. 
long; racemes dense, suberect, to 15 cm. long and 1.5 cm. thick, 
the pedicels to 6 mm. long; sepals 4, 3 mm. long, red and enlarged 
in fruit; stamens about 16. The related Bolivian S. racemosa 
(Britton) H. Walt. Bot. Jahrb. 37: Beibl. 85: 24. 1906, and S. mollis 
H. Walt. Pflanzenr. IV. 83: 116. 1909, have leaves pubescent beneath, 
the first with short, the second with elongate pedicels. Illustrated, 
op. cit. 115. 

Puno: Chunchosmayo, 900 meters, Weberbauer 1168. 



FLORA OF PERU 553 

Schindleria Weberbaueri 0. C. Schmidt, Notizbl. Bot. Gart. 
Berlin 8: 313. 1923. 

Similar to S. glabra, but the leaves obtusely acuminate, on petioles 
to 12 mm. long; sepals scarcely longer than 2.5 mm., little enlarged 
in fruit, greenish. S. rivinioides (Rusby) H. Walt. Bot. Jahrb. 37: 
Beibl. 85: 24. 1906, has flowers in loose racemes; S. rosea (Rusby) 
H. Walt. Bot. Jahrb. 37: Beibl. 85: 24. 1906, has rose-colored flowers 
in dense racemes. A shrub of 3 meters; flowers white. 

Huanuco: Mairo Valley, 400 meters, Weberbauer 6758 (type). 
Cuzco: Lares Valley, 1,900 meters, border of forest, Weberbauer 
7924, 7924a (det. Schmidt). 

8. FLUECKIGERA Kuntze 

Ledenbergia Klotzsch. This genus is well marked by the long, 
pendulous inflorescence, composed of several to many slender racemes 
that are particularly conspicuous in fruit, because of the papery, 
stellately spreading sepals. The 12 stamens are regularly disposed, 
8 alternate with the sepals. The original name of Klotzsch com- 
memorates, though falsely written, that of a professional politician; 
the same individual had been honored by Ladenbergia Klotzsch in 
1846, for which reason Kuntze changed the later name to Flueckigera. 
I follow Kuntze, believing that to accept two scientific names of the 
same personal origin which differ by an accidental (not deliberate) 
change in spelling as slight as this is condonable in neither the name 
of science nor of common sense, all arbitrary rules and regulations 
to the contrary notwithstanding. 

Flueckigera peruviana (0. C. Schmidt) Macbr., comb. nov. 
Ledenbergia peruviana O. C. Schmidt, Notizbl. Bot. Gart. Berlin 8: 
313. 1923. 

A shrub 4 meters high, the slender branchlets minutely lineate- 
puberulent; petioles about 3.5 cm. long; leaf blades elliptic, acumin- 
ate at both ends, glabrous, to 10 cm. long and half as wide; panicle 
to 25 cm. long, 18 cm. wide; pedicels 3 mm. long; sepals narrowly 
oblong, rounded at the apex, white, 4 mm. long, becoming 2 mm. 
longer in fruit, the fruit 2.5 cm. thick. 

Piura: Ayavaca, Weberbauer 6413 (type). 

9. PHYTOLACCA L. 

Nearly smooth, green-branched shrubs or trees with ovate-elliptic, 
pointed leaves and racemose or spicate flowers. Inflorescences 
terminal but finally pseudo-lateral. Carpels 5-16, forming a de- 



554 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

pressed-globose, baccate fruit. The young shoots and leaves of 
several species are cooked as "greens," and the roots or other parts, 
which contain saponin, rubbed in water, form suds that serve in 
place of soap or as an aid to it. 
Racemes spiciform, the pedicels very short, rarely 3.5 mm. long 

and then the lower not longer than the upper. 
Inflorescence shorter than or little exceeding the leaves. 

Flowers scarcely touching; carpels free at the tip . . .P. dioica. 

Flowers very crowded, touching; carpels connate. .P. bogotensis. 
Inflorescence much longer than the leaves , . P. icosandra. 

Racemes open, the flowers sometimes contiguous but at least some 
of the pedicels 5 mm. long or longer. 

Inflorescence simply racemose. 

Stamens little or not at all exceeding the sepals ... P. rivinoides. 

Stamens much exceeding the sepals, conspicuous. P. Weberbaueri. 
Inflorescence paniculate at the base P. thyrsiflora. 

Phytolacca bogotensis HBK. Nov. Gen. & Sp. 2: 183. 1817; 
60. P. australis Phil. Anal. Univ. Chile 43: 536. 1873. 

A smooth, green, tree-like shrub, the stout branches scarcely 
or little angled; petioles to 4 cm. long, grooved and angled; leaf 
blades oblong-elliptic or broadly lanceolate, acute at both ends or 
acuminate, the base decurrent, usually chartaceous and white- 
punctate, to about 10 cm. long and 4 cm. broad; racemes suberect, 
many-flowered, to only 4.5 cm. long and 13 mm. thick, the peduncles 
to 1 cm. long; flowers perfect, the pedicels 3 mm. long; stamens 
(8-12) and sepals subequal; ovary 8-10-carpellate, the carpels com- 
pletely joined; fruit baccate. In Colombia called "guaba." Herrera 
records from Cuzco, and Ball from Matucana, probably as an intro- 
duction or escape, the similar P. octandra L., a species ranging from 
Mexico to Colombia and widely naturalized elsewhere; its racemes, 
to 50 cm. long, about equal the leaves. Neg. 5713. 

Cajamarca: Nancho, Raimondi. Amazonas: Chachapoyas to 
Moyobamba, Raimondi. Huanuco: Twelve miles south of Panao, 
2214- Mito, 2,700 meters, 1549. Ayacucho: Huanto, Weberbauer 
5592 (det. Pilger). Huancayo: Chavez (coll.?; det. Schmidt). 
Cuzco: Machupicchu, Soukup 193. Prov. Calca, Herrera 354. 
Prov. Urubamba, 3,200 meters, Herrera 510, 668 (det. Mansfeld), 
688. Valle de Paucartambo, Herrera 3373. Without locality 
(Dombey). Junin: Yanahuanca, 1215. Huasa-huasi, Ruiz & Pavon 



FLORA OF PERU 555 

(det. Schmidt). Huanuco: Vitoc, Isern 2558, 2224- Colombia to 
Chile. "Chocllo-chocllo," "ailambo." 

Phytolacca dioica L. Sp. PL ed. 2. 632. 1763; 47. 

Except for the pedicels, a glabrous, evergreen shrub, pale- 
verruculose at the stout and angled nodes; petioles slender, to 4 cm. 
long; leaf blades ovate, acute or acuminate, rounded but slightly 
decurrent at the base, thick, lightly reticulate-veined beneath, 
sometimes over 10 cm. long; racemes more or less nodding, to 12 
cm. long, on peduncles to 1.5 cm. long; flowers dioecious, the ovary 
rudimentary in the staminate, the pistillate rarely with staminodia; 
pedicels in the former rarely 4 mm. long, in the latter 3 mm. long; 
sepals elliptic, white-punctate, 3.5 mm. long, much exceeded by the 
20-30 stamens; ovary globose, 7-10-carpellate, the carpels connate 
at the base. This abundantly flowered, gigantic-based plant, often 
planted for shade, is known in the southern part of its range as 
"ombu" or "umbu" or elsewhere as "bella sombra" or "belombra." 
See Record & Mell, Timbers Trop. Amer. 157. 1924, for a description 
of the wood. Illustrated, Pflanzenreich IV. 83: 48. 

Peru: Without locality: Raimondi; (Haenke). Extending to 
Argentina. 

Phytolacca icosandra L. Sp. PL 631. 1753; 60. 

A shrub allied with and in general similar to P. bogotensis, but 
generally recognizable by the spicate or racemose inflorescence, which 
is usually terminal and much longer than the leaves; flowers sub- 
sessile or rarely on pedicels 2.5 mm. long; stamens 16-22; carpels 
8-10, all connate. Illustrated, Bot. Mag. pi. 4967. 

Cajamarca: Huancabamba, Chota, Raimondi. West Indies and 
Mexico to Peru. "Huailampo." 

Phytolacca rivinoides Kunth & Bouch^, Ind. Sem. Hort. 
Berol. 15. 1848; 55. 

A scandent or straggling herb, to 5 meters high, the stout branches 
herbaceous; leaves ovate-acuminate, membranous, often over 10 
cm. long and 6 cm. wide, minutely white-punctate; racemes lax, 
suberect, much exceeding the leaves, to 40 cm. long and 2.5 cm. 
thick, the peduncles to 5 cm. long; flowers perfect, the slender pedicels 
to 1 cm. long; sepals broadly elliptic, 2 mm. long; stamens 10-20; 
carpels 10-16, connate. The leaves and young shoots, cooked, often 
serve for "greens." Illustrated, Field Mus. Bot. 10: pi. 35. Neg. 
5717. 



556 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cuzco: Marcapata (Hen era 1175). Huanuco: Hacienda Pam- 
payacu, 1,050 meters, in clearing, 5073. Junin: Chanchamayo, 
Raimondi; Isern 2368. Chanchamayo Valley, 1,200-1,500 meters, 
Schunke 1473, 1791, 8. Above San Ramon, 1,500 meters, in forest, 
Kittip & Smith 24530. Rio Pinedo, 800 meters, in forest, Kittip & 
Smith 23634. Puerto Yessup, 400 meters, Kittip & Smith 26287; 
petals and anthers white; fruit black. Yapas, 1,500 meters, Kittip 
& Smith 25477; fruit purple. Loreto: Leticia, Ule 6190. Pinto- 
cocha, Williams 809. La Victoria, in forest, Wittiams 2556. Puerto 
Arturo, edge of water, Williams 5235. Pongo de Manseriche, 200 
meters, on landslides in sun, Mexia 6231. Caballo-cocha, Wittiams 
2495. Lower Rio Nanay, Williams 436. Near Iquitos, Wittiams 
1378, 3710, 1323; Kittip & Smith 26991. Yurimaguas, 200 meters, 
Wittiams 3849. Puno: Sangaban, Raimondi. San Martin: Zepela- 
cio, 1,200-1,600 meters, Klug 3494- South America to Mexico and 
the West Indies. "Apacas," "airambo," "jaboncillo," "jaboncillo 
airambo." 

Phytolacca thyrsiflora Fenzl ex J. A. Schmidt in Mart. Fl. 
Bras. 2: 343. pi. 80. 1872; 50. 

Allied to P. rivinoides but more robust; inflorescence thyrsiform, 
to 15 cm. long and at the base 2.5 cm. wide, slightly exceeding the 
leaves; peduncles to 6 cm. long, the pedicels finally 7 mm. long; 
sepals nearly 3 mm. long; carpels 7-8, connate. 

Peru: (Mathews 3118). Paraguay; Brazil; Dominican Republic. 

Phytolacca Weberbaueri H. Walt. Pflanzenr. IV. 83: 49. 1909. 

Similar to P. dioica, but so far as known only a shrub, sometimes 
6 meters high, with an erect inflorescence much exceeding the leaves 
or to nearly 30 cm. long; pistillate flowers and fruit unknown. Neg. 
5722. 

Cajamarca: East of Chepe"n, 700 meters, Weberbauer 4817 (type). 
Lambayeque: Raimondi. "Santo Torne," "yumbi." 

10. GALLESIA Casar. 

A much branched, often tall tree with ovate-elliptic, coriaceous 
leaves and ample, axillary and terminal inflorescences of panicled 
racemes. Flowers perfect, 4-parted, the free sepals erect in fruit. 
Fruit samara-like. Kuntze wrote the name Gallesioa, the genus 
being named for Gallesio, an Italian botanist. 

Gallesia integrifolia (Spreng.) Harms, Pflanzenfam. ed. 2. 16c: 
144. 1934; 84. Thouinia integrifolia Spreng. Neue Entdeck. 2: 155. 



FLORA OF PERU 557 

1821; Syst. 2: 221. 1825. G. Gorarema (Veil.) Moq. in DC. Prodr. 
13, pt. 2: 8. 1849. Crataeva Gorarema Veil. Fl. Flum. Text 1: 200. 
1825; Icon. 5: pi. 4.1827. 

Leaves very smooth, mostly over 10 cm. long and nearly 5-6 cm. 
wide, borne on slender petioles; inflorescence 25 cm. long and nearly 
as broad; sepals enlarged in fruit to 6 mm. long and half as wide; 
fruit wing about 2 cm. long and 9 mm. broad. A tree, readily known 
in flower by its onion-like odor and in fruit by its maple-like seeds; 
a Brazilian name is "pao d'alho." The leaves are used as soap. 
Record & Mell, Timbers Trop. Amer. 158, have described the wood, 
rich in potash (under the name G. scorododendron Casar.). Illus- 
trated, Pflanzenreich IV. 83: 84. 

San Martin: Tarapoto, Spruce 4156. Rio Acre: Krukoff 5405, 
5216. Brazil. 

Gallesia ovata O. C. Schmidt, Repert. Sp. Nov. 32: 97. 1933. 

Branchlets and inflorescence more or less tomentose; petioles 
3.5-5.5 cm. long; leaf blades ovate, 10-17.5 cm. long, 6-10 cm. wide, 
rounded at the base, narrowed to the rounded but mucronulate 
apex, thin, glabrous above, barbate in the 8-9 nerve axils beneath; 
racemes 12-15 cm. long; flowers 5 mm. long, subsessile, the sepals in 
fruit to 7 mm. long; fruit wing abruptly dilated from a nearly linear 
base, to 1.5 cm. long. 

Cuzco: Valle de Yanabili, Raimondi (type). 

11. SEGUIERIA Loefl. 

Similar to Gallesia, but often shrubby or scandent, and some- 
times with tubercle-like stipules. Sepals 5, reflexed in fruit. 

Seguieria foliosa Benth. Trans. Linn. Soc. 18: 235. 1839; 93. 

A glabrous shrub or tree with slender, chestnut-colored branches; 
petioles 3 mm. long, with 2 spine-like stipules at the base; leaf 
blades ovate, gradually acuminate, coriaceous, 2.6 cm. long and 
nearly half as wide, minutely pellucid-punctate; inflorescence broadly 
spicate, to 20 mm. long, the branchlets to 4 cm. long; pedicels about 
5 mm. long; sepals 4.5 mm. long and 3 mm. broad; fruit wing to 2.5 
cm. long and 1.5 cm. broad, the seed-bearing portion with 3 or 4 
small wings. Neg. 27745. 

Rio Acre: Ule 9487 (det. Schmidt), 9486 (det. Pilger). 

Seguieria macrophylla Benth. Trans. Linn. Soc. 18: 235. 
1839; 195. 



558 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Distinguishable from the preceding by its twice longer petioles, 
its larger (to 30 cm. long) inflorescence, and especially by its smooth 
fruits with somewhat narrower wings. 

Loreto: Yurimaguas, Poeppig D2176. Bolivia. 

64. AIZOACEAE. Carpet-weed Family 
Well known as is the leafy vegetable New Zealand spinach 
(Tetragonia expansa), established in Peru, which belongs to this 
family, the group is equally interesting as containing the genus 
Mesembryanthemum sens, lat., which shares with cacti the great 
popularity among cultivated succulents. In Peru the family is 
represented only by the following herbs, mostly of sandy places. 
Leaves opposite or whorled, at least apparently; calyx free. 

Plants finely pubescent with branched hairs 1. Glinus. 

Plants glabrous or not as above. 

Leaves whorled 2. Mollugo. 

Leaves opposite, fleshy. 

Stipules present; ovary 1-2-celled 5. Trianthema. 

Stipules none; ovary 3-5-celled 3. Sesuvium. 

Leaves alternate: calyx tube adnate to the ovary. . . .4. Tetragonia. 

1. GLINUS L. 

Annual or perennial herbs with opposite or whorled leaves and 
pediceled flowers clustered in the leaf axils. Petals none. Stamens 
3-20. Capsule loculicidal. Funicle filiform. The related Glischro- 
thamnus Ulei Pilger, Bot. Jahrb. 40: 396. 1908, of Brazil has closely 
clustered, subsessile flowers and leaves densely punctate with dark 
glands. 

Glinus radiatus (R. & P.) Rohrb. in Mart. Fl. Bras. 14, pt. 2: 
238. 1872. Mollugo radiata R. & P. Fl. 1: 48. 1798. 

A prostrate or ascending, stellate-pubescent annual; leaves 
obovate, usually 1-2 cm. long, scarcely acute; flowers densely 
clustered; sepals 3-4 mm. long, enclosing the capsule; seeds 
many, lustrous, brown. Illustrated, Fawcett & Rendle, Fl. Jam. 3: 
pi 59. 

Peru: (Probably). Chile (type locality) to Texas and West Indies. 

2. MOLLUGO L. Carpet-weed 

Branched annuals with linear leaves in uneven whorls, or the 
leaves all basal. Sepals 5, the stamens usually of the same number 
or fewer, rarely more numerous. Capsule loculicidal. 



FLORA OF PERU 559 

Mollugo verticillata L. Sp. PI. 89. 1753. 

Prostrate, much branched, with usually 5-6 linear-spatulate 
leaves in each whorl, the longest leaves 2.5 cm. long or sometimes 
longer; flowers axillary, pediceled, apetalous, 2 mm. long, the sepals 
slightly exceeded by the capsule; seeds ridged, brown. Common in, 
temperate and tropical America, often in cultivated places. To 
be expected is M. nudicaulis Lam., the leaves all in a basal tuft, 
narrowed to a long petiole and about 4 cm. long, the flowers in a 
long-peduncled cyme. 

Arequipa: Near Mejia (Gunther & Buchtien 3). Piura: Eleven 
miles east of Cabo Blanco, Haught Fl 73. Lima: Quive, 1,000 
meters, open, rocky slope, Pennell 14303. Nearly cosmopolitan. 

3. SESUVIUM L. 

Perennial strand plants with sessile or shortly pediceled, appar- 
ently axillary flowers. Styles and capsule cells usually 3-5, with 
numerous seeds in each cell. 

Sesuvium Portulacastrum L. Syst. ed. 10. 1058. 1759. 

Glabrous, the often long stems rooting at the nodes; leaves 
fleshy, nearly oblong, sessile, 1.5-4 cm. long; flowers 6-10 mm. long, 
with numerous rose-purple stamens; petals none. Urban has shown 
that the flowers are really terminal. Used in lea in the manufacture 
of glass and elsewhere in soap making, in place of soda (Ruiz & 
Pavon). Illustrated, Pflanzenfam. ed. 2. 16c: 229. 

Piura: Paita, Gaudichaud. lea: (Ruiz & Pavon). Lima: Lurin, 
Ruiz & Pavdn. Arequipa: Mollendo (Weberbauer, 148). Mejia 
(Gunther & Buchtien 315, 316). Tacna: Arica, Brenning 141. On 
nearly all shores. "Litho." 

4. TETRAGONIA L. 

Somewhat fleshy herbs or woody-based plants with solitary or 
few axillary flowers. Petals none. Fruit nut-like, indehiscent, some- 
times armed or winged. Foliage and stems usually more or less 
whitened by spiculate, glistening scales or pubescence. 

A shrub or half-shrub with narrow leaves T. maritima. 

Annual or perennial herbs, little or not at all woody below. 
Leaves oblong, oblong-linear, or narrowly obovate. 

Flowers distinctly pediceled T. pedunculata. 

Flowers sessile T. macrocarpa. 

Leaves ovate to suborbicular. 



560 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plants perennial, the greenish leaves often 5 cm. wide. 

T. expansa. 
Plants annual, the often whitish leaves smaller. 

Fruit quadrangulately lobed, beaked, 4-celled . . T. crystalline,. 

Fruit globose, somewhat quadrate in drying T. ovata. 

Fruit obscurely 4-angled, 5-7-celled T. vestita. 

Tetragonia crystallina L'HeY. Nov. Hist. 81. pi 39. 1784. 

Openly branched, 10-30 cm. high; leaves sessile, ovate, acute, 
crystalline-papillose, narrowed to the apex, 4-6 cm. long, 2-3 cm. 
wide; flowers solitary or rarely binate, subsessile, 8 mm. wide, in 
fruit pedicellate, the ovate-acute divisions spreading, crystalline- 
papillose without; filaments 16, yellow; styles 4; fruit 4-angulately 
lobed, beaked. 

Lima: Chancai (Dombey). Lima, Abadia. Chorrillos, loose, 
stony, upper slopes of seaside hills, 150 meters, 5855. 

Tetragonia expansa Murr. Comment. Gott. 6: 13. 1785. 
Demidovia tetragonoides Pall. Hort. Demid. 150. pi. 1. 1781. T. 
tetragonoides Kuntze, Rev. Gen. 1: 264. 1891. 

A nearly smooth, coarse, subprostrate perennial with rhombic- 
ovate, petioled leaves 2.5-8 cm. long; flowers solitary, axillary, 
5-6 mm. wide, the 4-knobbed, subsessile fruit about 1 cm. thick. 
The earliest name if validly published (publication not seen) is 
T. tetragonoides (Pall.) Kuntze, but in any case this binomial, from 
an intelligent if not from an academic standpoint, merely repeats 
the generic name and therefore its use is contrary to the International 
Rules. The leaves, cooked, are used as "greens." Illustrated, 
Pflanzenfam. ed. 2. 16c: 233. 

Arequipa: Mejia (Gunther & Buchtien 317; det. Bruns). Widely 
distributed and cultivated. "New Zealand spinach." 

Tetragonia macrocarpa Phil. Fl. Atac. 19. 1860; Viaje Des. 
Atac. 12, 193. 1860. 

More or less spiculate-villous, to 10 cm. high, usually lower, 
branching, mostly horizontally, from the base; leaves oblong-obovate, 
obtuse, 1-2 cm. long, narrowed to the sessile base; flowers solitary, 
sessile; ovary 4-6-celled; fruit sessile or short-pediceled, angled with 
4-6 ridges, these sometimes nearly wing-like, to 1 cm. long. 

Tacna: At 650 meters, Werdermann 721. Chile. "Pasto 
aguanoso." 



FLORA OF PERU 561 

Tetragonia maritima Barnh. in Gay, Fl. Chile 2: 469. 1846. 

A dioecious, stoutly branched, spiculate-villose shrub; leaves 
oblong-lanceolate, obtuse, attenuate basally to a short petiole; 
flowers solitary, short-pediceled, the staminate with a rudimentary 
ovary; stamens 15; styles papillose; fruit 3-celled, longitudinally 
3-winged. According to Johnston, a dense bush 30-120 cm. high 
with yellowish green, succulent foliage. Allied to T. angustifolia 
Barnh., with oblong-linear leaves, 15-20 stamens, the 3-4-celled 
fruit 4- winged. 

Tacna: Arica, Ball. Arequipa: Mejia, Gunther & Buchtien 313, 
314, 318 (det. Bruns). Chile. 

Tetragonia ovata Phil. Anal. Univ. Chile 85: 168. 1893. 

A small annual with crowded, elongate, ovate leaves about 3 cm. 
long, gray-green with papillose-spiculate pubescence; flowers yellow, 
short-pediceled; fruit prismatic, 4-ridged or angled. Johnston 
describes the plant as "usually with widely spreading branches, the 
fruit subglobose, red, juicy, only somewhat 4-angled in drying." 

Tacna: At 800 meters, Werdermann 725 (det. Werdermann as 
aff.). Cachendo, Gunther & Buchtien 319 (det. Bruns). Moquehua: 
Mountains between Moquehua and Torata, 2,000 meters, Weber- 
bauer 7422. Chile. 

Tetragonia pedunculata Phil. Anal. Univ. Chile 717. 1872. 

Low, densely long-spiculate-villose; leaves oblong-linear; flowers 
distinctly pediceled, the pedicels becoming 1-1.5 cm. long; fruit 
5-6-celled, pyramidal-prismatic, the 4-6 ridges united at the base. 

Tacna: Woitschach in 1891. Lima: San Agustin, Weberbauer 4- 
Chile. "Lechuga." 

Tetragonia vestita I. M. Johnston, Contr. Gray Herb. 81: 

88. 1928. 

An annual herb 5-15 cm. high, the few branches ascending in 
part; leaves ovate or oblong-ovate, 2.5 cm. long, 1.5-3 cm. broad, 
crystalline-papillose and more or less villous; calyx 4-5 mm. long, 
the petal-like lobes obtuse, yellowish; stamens 15-20; stigmas 5-7, 
the ovary with as many cells and seeds, the fruit usually about 6 
mm. long; ovary very densely villous with subulate or conic, 1-celled 
trichomes. 

Arequipa: Just below the fertile belt on hill back of Mollendo, 
Johnston 3533; (Weberbauer 144?). 



562 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

5. TRIANTHEMA L. 

The genus is represented in America by a single species. 

Trianthema Portulacastrum L. Sp. PI. 223. 1753. T. monan- 
thogyna L. Mant. 1: 69. 1767. 

An erect or ascending, glabrous annual, the leaves rounded- 
obovate, fleshy, bright green; calyx 5-parted, colored within; stamens 
6-10; style 1. 

Piura: La Brea, 150 meters, Weberbauer 7764,' flowers rose- 
colored. Parinas Valley, Haught 268; flowers rose. Negritos, 
Haught F 84- Eight miles northeast of Cape Parinas, Haught 173. 
Widely distributed in the warmer parts of America. 

65. PORTULACACEAE. Purslane Family 

Apart from the curiously long-hairy Portulacas, the Calandrinias 
are the most interesting Peruvian purslanes because of their variety 
and their often pretty, flax-like blossoms. 

Flowers not glomerulate and scarious-bracted. 
Capsule circumscissile, adnate below to the calyx tube; plants 

usually pilose, especially in the leaf axils 1. Portulaca. 

Capsule valvate, (like the ovary) entirely free; plants not charac- 
teristically pilose. 

Petals distinct; seeds usually many. 

Sepals usually caducous; seeds smooth; inflorescence ample. 

2. Talinum. 
Sepals persistent; seeds usually rough; inflorescence often 

small 3. Calandrinia. 

Petals united below; seeds few; a small annual of wet places. 

4. Montia. 

Flowers glomerulate in the axils of colored, scarious bracts. 

5. Phillippiamra. 

1. PORTULACA L. 

Somewhat fleshy, annual or perennial herbs, decumbent, ascend- 
ing, or erect, commonly with the flowers and fruits involved in long, 
white or brownish hairs. Uppermost leaves forming an involucre 
around the sessile flowers. Calyx deciduous. Ovary partially 
inferior. Capsule circumscissile. P. grandiflora Hook., with very 
large and showy flowers of various colors, native of Brazil and 
Argentina, often is grown for ornament. The local name is "flor 



FLORA OF PERU 563 

de las once." Since the preparation of this account, Karl von Poell- 
nitz has published a revision of the genus, Repert. Sp. Nov. 37: 
240-320. 1934. Unfortunately, he treats only those species accepted 
as valid, and with specimens of the Peruvian plants no longer avail- 
able to me it has been impossible to place them in his treatment or to 
give them their correct nomenclature. 

Plants inordinately soft-pilose, the minute, crowded leaves hidden. 

P. pilosissima. 

Plants more or less pilose or glabrous, the leaves not minute. 
Plants distinctly perennial, with thick caudex and stems. 
Pubescence short, soft; seeds black, radiately stellate- tuber- 

culate P. peruviana. 

Pubescence long or stiff; seeds tardily red-black, tuberculate. 

P. lanuginosa. 

Plants evidently annual or the stems not thick or much branched. 
Leaves terete or narrowly linear. 

Seeds not or scarcely metallic-lustrous, more or less stellate- 

tuberculate; plants usually branching. 
Flowers typically red-purple; seeds black, stellate-tuber- 

culate P. pilosa. 

Flowers yellow; seeds red-brown, tuberculate, faintly 

stellate P. Haughtii. 

Seeds strongly metallic-lustrous, finely tuberculate; plants 

simple or nearly so P. amboensis. 

Leaves flat, oblong-lanceolate. 

Plants glabrous P. oleracea. 

Plants pilose. 

Involucral leaves few; seeds metallic-lustrous, stellate- 

tuberculate P. tingoensis. 

Involucral leaves many; seeds black, tuberculate. 

P. rubricaulis. 

Portulaca amboensis Macbr. Candollea 5: 350. 1934. 

Similar to P. tingoensis but the leaves terete, as many flowers 
axillary as terminal or more, and the stems not enlarged above; 
seeds distinctly purple-opalescent, 0.5 mm. broad. The 3-4 linear- 
terete leaves subtending the terminal flower cluster are no larger 
(8-15 mm. long) than the promptly deciduous cauline leaves. Stems 
simple or rarely with one or two branches. Growing on stony 
river flats. 



564 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Ambo, 2,100 meters, Macbride 3202 (type). Huanuco, 
Ruiz & Pavdn. 

Portulaca Haughtii Macbr. Field Mus. Bot. 11: 21. 1931. 

A slender annual with few to many loosely spreading branches; 
stems moderately white-pilose in the axils, the hairs about 5 mm. 
long; leaves alternate, perhaps subterete, narrowly linear, 1-1.5 cm. 
long, those of the stem scarcely crowded or caducous; hairs about 4 
mm. long, only partially concealing the flowers and capsules; stamens 
about 10; style 4-parted; capsule circumscissile about the middle; 
seeds acutely and finely tuberculate, the tubercles scarcely mani- 
festly stellate basally, reddish brown. Named for Mr. Oscar Haught, 
who made a nearly complete collection of the plants of Piura. 

Piura: High plain near Parinas Valley, Haught F 180, type. 

Portulaca lanuginosa HBK. Nov. Gen. & Sp. 6: 74. 1823. 

Stems procumbent, branching, with short to very long but 
always firm hairs in the axils; leaves 5-7 mm. long, approximate, 
fleshy, terete, blunt; stamens 16-25; styles 3-7; capsule circum- 
scissile below the middle; seeds tuberculate, tardily red-black and 
somewhat opalescent. A coarse perennial with typically yellow 
flowers, but in my material (possibly referable to P. peruviana) 
these red-purple. Poellnitz, op. cit. 262, thinks the HBK. plant 
belongs to the P. pilosa complex, following Rohrbach in Mart. Fl. 
Bras. 14, pt. 2: 304. 1872, who treats it as a race with short, obtuse 
leaves; supposedly the same plant has received the name P. pilosa 
var. eriocarpa (Casar.) Hauman. Probably, too, the HBK. name 
is not tenable; P. lanuginosa Crantz, Inst. 2: 428. 1778, was not 
cited by HBK. but it has been referred here. 

On the Marafion (Humboldt, type). Amazonas: Balsas, Weber- 
bauer 4266; 156. Huanuco: Stony river flat, Ambo, 2,100 meters, 
3201; flowers red-purple. Junin: Tarma, Weberbauer 2367. 

Portulaca oleracea L. Sp. PI. 445. 1753. 

Plants annual, very fleshy, prostrate, glabrous; leaves cuneate- 
obovate; flowers small, yellow, sessile in the leaf axils; capsule 
circumscissile. 

Cuzco : Valle del Urubamba, Machupicchu, 2,200 meters, Herrera 
3235. Huanuco: Huanuco, along road, 2,100 meters, 3537. Loreto : 
Rio Putumayo, in clearing, Klug 1620. Puerto Arturo, 200 meters, 
in pasture, Williams 5033. Fortaleza, 200 meters, Williams 4365. 
Pisco, Williams 1283. Rio Masana, Williams 244- A species of 
cosmopolitan distribution. "Verdolaga." 



FLORA OF PERU 565 

Portulaca peruviana I. M. Johnston, Contr. Gray Herb. 81: 
88. 1928. 

A perennial, fleshy, much branched at the base, with often 
numerous stems a few to many cm. high; axil pubescence short and 
dense or even lacking; leaves alternate, 5-10 mm. long, linear- 
spatulate, many of them persistent, the shorter involucral ones 
scarcely exceeding the purple flowers or the capsules; capsule dehis- 
cent near or toward the base; seeds black, little lustrous, radiately 
stellate-tuberculate. Open, rocky slopes. Perhaps not distinct 
from P. lanuginosa, unless in the shorter pubescence. 

Arequipa: Tingo, 2,200 meters, Pennell 13140 (type); Stubel 81. 
Mt. Chachani, 3,000 meters, sandy pampa, Hinkley 1. Moquehua: 
Carumas, 2,700 meters, Weberbauer 7278. Lima: Prostrate on 
granitic slopes, Matucana, 2,400 meters, 278; Weberbauer 90, 90a, 
163, 166. San Bartolome", Weberbauer 5281. Ancash: Near Huaraz, 
Raimondi. "Verdolaga." 

Portulaca Philippii I. M. Johnston Contr. Gray Herb. 85: 39. 
1929. 

A fleshy perennial, often ligneous below, the branches 5-15 cm. 
long, procumbent or ascending, usually branched; leaves alternate, 
numerous, suberect, 5-8 mm. long, 1.5-2.5 mm. broad, persistent; 
axillary hairs moderately long; flowers solitary or 2-3, the triangular, 
ovate, acute involucral leaves about 4 mm. long; sepals ovate, 
8 mm. long; stamens 25-35; capsule stipitate, circumscissile below 
the middle; seeds opaque, 0.5 mm. wide, blackish, minutely tubercu- 
late-stellate. 

Puno: Lake Titicaca (Pentland). Bolivia; Chile. 

Portulaca pilosa L. Sp. PI. 445. 1753. 

An ascending-erect annual, more or less conspicuously pubescent 
in the leaf axils; leaves narrowly linear, acute, to 1.5 cm. long, 
alternate; flowers typically red or purple, to a few mm. long, the 
subtending leaves 2-3 mm. long, involved in long hairs; stamens 15 
or more; styles 3-6; capsule circumscissile about the middle; seeds 
0.5 mm. wide, minutely tuberculate, each obtuse (or acute?) cone 
star-like at the base. Urban's interpretation of this species, Symb. 
Antill. 5: 343. /. A-B. 1907, is accepted here. Possibly the following 
Peruvian material (not studied by me) is referable to other species, 
very probably at least in part. Tessmann found the species growing 
near houses, the leaves rubbed on the hair to "smooth" it and, 
"das Kinder gut gehen," on their knees. 



566 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Loreto: Yarina-cocha, Tessmann 5484. Pinto-cocha, Williams 
815, 1278. Iquitos, Williams 7939. Leticia, Williams 3028,Cuzco: 
Valle del Vilcanota, 3,550 meters (Hickeri). Ollantaitambo, 2,800 
meters, Herrera 627. Arequipa: Mollendo (Gunther & Buchtien 
1679, fide Bruns). South America to the United States. "Flor 
de seda," "flor de las once," "bustirao chama." 

Portulaca pilosissima Hook. Bot. Misc. 2: 220. 1831. 

Inordinately and softly white-pilose, the small, terete leaves and 
flowers concealed in the long, rusty pubescence; stems decumbent, 
branched. Characteristically a much branched, matted perennial. 
Poellnitz, op. cit. 261, treats it as a form of P. pilosa. 

Lima: Yazo, Valley of Canta (Cruckshank). Tacna: Woitschach. 
Arequipa: Mollendo, Weberbauer 1483; 144. Mejia (Gunther & 
Buchtien 303, 304). 

Portulaca rubricaulis HBK. Nov. Gen. & Sp. 6: 73. 1823. 

A well marked species, at least in Peru, by its flat leaves about 
1 cm. long and 3-4 mm. wide, its glabrous stems (but the soft leaf 
axil pubescence sometimes so long that it reaches from leaf node to 
leaf node!), and its 6-9 involucral leaves; flowers purple, usually 
several and terminally congested; capsule about medially circum- 
scissile; seeds black, lustrous, tuberculate. The stems are often 
branched but mostly only above the base. Our material has some- 
what shorter leaves than those of the type. Poellnitz refers the HBK. 
plant to P. pilosa with a query; very possibly the Peruvian collec- 
tions were misdetermined. The type is from Venezuela. 

San Martin: Tarapoto, Wittiams 5686, 5787. 

Portulaca tingoensis Macbr. Field Mus. Bot. 11: 21. 1931. 

A few cm. high or rarely higher, simple or branching near the 
base, the stems or branches strongly enlarged toward the tips; 
leaves alternate, caducous, apparently flat, oblong-spatulate, about 
1 cm. long, the involucral few; inflorescence densely involved in 
hairs 5-7 mm. long; capsules sometimes axillary, solitary, circum- 
scissile below the middle; seeds iridescent, bluntly tuberculate, the 
scales radiate at the base, nearly 0.6 mm. broad. Flowers unknown. 
This may prove to be referable to P. simpliciuscula Mart., with 
shorter pubescence and eradiate scales. 

Arequipa: Open, rocky and sandy slopes in early April, Tingo, 
2,200 meters, Pennell 13111 (type). 



FLORA OF PERU 567 

2. TALINUM Adans. 

Glabrous, often slender herbs with flat leaves and generally 
an amply racemose or paniculate inflorescence. Sepals promptly 
or tardily deciduous. Ovary free. Capsule 3-valved. 

Flowers 5 mm. long or shorter, paniculate T. paniculatum. 

Flowers 6 mm. long or longer, usually racemose T. triangulare. 

Talinum paniculatum (Jacq.) Gaertn. Fruct. 2: 219. 1791. 
Portulaca paniculata Jacq. Enum. 22. 1760. P. patens Jacq. Hort. 
Vind. 2:71. 1772. T. patens Willd. Sp. PI. 2: 863. 1800. T.reflexum 
Cav. Icon. 1: 1. pi. 1. 1791. T. dichotomum R. & P. Syst. 118. 1798. 

Often fleshy-suffrutescent at the base and several dm. high; 
leaves more or less obovate and somewhat petiolate, usually 5-10 
cm. long; pedicels terete, filiform; flowers rose, flesh-colored, or 
yellow; stamens 15-20. Illustrated, Fa we. & Rendle, Fl. Jamaica 
3, pt. 1:171. 1914. 

Junin: La Merced, 700 meters, Killip & Smith 23721. Loreto: 
Fortaleza, 200 meters, in pasture, Williams 4409. Yurimaguas, 200 
meters, in pasture, Williams 3955. Puerto Arturo, 200 meters, 
Williams 51 05. Huanuco : Posuso, Esposto (det. Ulbrich) . Huanuco, 
Ruiz & Pawn. Huanuco, 2,100 meters, loose soil on cliffs, 3257. 
Lima: Chancai (Ruiz & Pawn). San Bartolome", Weberbauer 5307 
(det. Ulbrich). San Martin: San Roque, 1,400 meters, Williams 
7364. Tarapoto, Ule 6338; Williams 5593. Tumbez: East of 
Hacienda Chicama, 1,000 meters, Weberbauer 7652. Without 
locality: Weberbauer 6463. South America to the southern United 
States. "Cuchi-yuya," "sacha-culantro," "cuchi-yuyu." 

Talinum triangulare (Jacq.) Willd. Sp. PI. 2: 862. 1800. 
Portulaca triangularis Jacq. Enum. PI. Carib. 22. 1760. P. racemosa 
L. Sp. PI. ed. 2. 640. 1762. T. crassifolium Willd. Sp. PI. 2: 862. 
1800(?). 

Similar to T. paniculatum but the pedicels 3-angled and the 
stamens often more numerous. Flowers sometimes white. The 
plant is sometimes cooked and eaten as a vegetable. 

Piura: Chanso, Weberbauer 5998 (det. Ulbrich). Tumbez: 
Hacienda Ci4naga, 250 meters, Weberbauer 7629.- San Martin: 
Tarapoto, Ule 6334; Williams 5728. Widely distributed in tropical 
America. 



568 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3. CALANDRINIA HBK. 

Cosmia Domb. ex Juss. Gen. 312. 1789; Baitaria R. & P. Prodr. 
63. pi. 36. 1794. 

Low herbs, rarely acaulescent, with axillary flowers, or more 
frequently the flowers in racemes or panicles terminating the often 
leafless stems. Leaves mostly crowded toward the base of the plant. 
Stamens few to many, variable in number in the same species. 
Sepals more or less enclosing the mature capsule. Seeds usually 
puncticulate. Kuntze, Rev. Gen. 1 : 14. 1893, referred the genus to 
Claytonia Gronov. The generic name of HBK. has been conserved. 
The fleshy leaves are used as an emollient. 

Plants caulescent, the stems sometimes short, particularly in C. ciliata. 

Flowers several to many, terminal; stamens usually more than 10. 

Leaves, at least the lower, subrotund-obovate, often several 

cm. wide. 
Petals entire. 

Capsule little exserted; seeds sparsely hispid. . . .C. Ruizii. 
Capsule well exserted; seeds densely hispidulous. 

C. paniculata. 

Petals crenate C. crenata. 

Leaves linear to narrowly obovate, few if any of them more 

than 1 cm. wide. 
Leaves oblong-obovate, all or most of them 6-10 mm. wide. 

C. pachypoda. 

Leaves linear-oblanceolate, 2-4 mm. wide or rarely wider. 
Plants annual or biennial; leaves often 3-5 cm. long. 
Capsule little if at all exceeding the sepals. 
Flowers usually umbellate-fasciculate; capsule a little 

exserted C. Weberbaueri. 

Flowers racemose or paniculate; capsule and sepals 

subequal C. lingulata. 

Capsule twice as long as the sepals C. quivensis. 

Plants perennial; leaves 2-2.5 cm. long C. linomimeta. 

Flowers solitary, axillary; stamens often fewer than 10. 
Seeds scarcely lustrous, distinctly puncticulate; leaves obovate. 

C. alba. 

Seeds very lustrous, not strongly punctate; leaves nearly linear. 

C. ciliata. 
Plants acaulescent, perennial, with a fusiform root C. acaulis. 



FLORA OF PERU 569 

Calandrinia acaulis HBK. Nov. Gen. & Sp. 6: 78. 1823. 

A tufted, little perennial with solitary, white, or later roseate 
flowers more or less hidden in the numerous elongate, linear leaves; 
pedicels a few mm. to 2 cm. long, recurving in fruit; leaves 2-4 
mm. wide, 1 to several cm. long, glabrous; flowers 1 cm. long or 
smaller or, in var. magna Macbr. (Candollea 5: 350. 1934), over 2 
cm. long; stamens few; capsule elliptic, included; seeds roundish- 
ovate, very minutely puncticulate, the puncticulations scarcely dis- 
cernible under magnification less than 10. Sandy or stony soils. 
"Usually on the puna to 3,200 meters" (Weberbauer 88, 196, 211). 
The Lechler specimen, distributed under a new name by Barneoud, 
seems to be referable here. 

Arequipa: Open, sandy places along streams, Nevado de Cha- 
chani, 3,800 meters, Pennell 1327 b. Puno: Agapata, Lechler 2672. 
Sandia region, Weberbauer 956; 219. Lima: Canta, Ruiz & Pavon. 
Viso, 2,700 meters, 622, 606 (var. magna). Above Lima, Weberbauer 
169Junm: Near Yauli, Weberbauer 380; 221. Rio Blanco, 4,500 
meters, 2981. Near Oroya, Weberbauer 232; 223. Ancash: Ocros, 
Weberbauer 2671. Huanuco: Mito, 2,700 meters, 1946. Cuzco: 
Valle del Paucartambo, Hacienda Lauramarca, 2,600 meters, 
Herrera 2339. "Thurpa." 

Calandrinia alba (R. & P.) DC. Prodr. 3: 359. 1828. Talinum 
album R. & P. Syst. 1: 116. 1798. Claytonia alba Kuntze, Rev. Gen. 
1:14.1891. 

Glabrous, with slender, often elongate stems and rather remote, 
lance-spatulate leaves to 3.5 cm. long and 8 mm. broad, strongly 
narrowed to a petiole-like base about 1 cm. long, rounded to an 
acutish apex; pedicels in fruit 8 mm. long; capsule exserted about 
1 mm. ; seeds nearly round, very turgid, scarcely lustrous, distinctly 
puncticulate, nearly 1 mm. wide. The flowers of the original 
collection seen by me are undeveloped; in the Weberbauer material 
(det. Berlin) they are purple, in mine white. Stony places. 
Neg. 8434. 

Lima: Chancai, Ruiz & Pavon. San Geronimo, 150 meters, 
5.908. Arequipa: Mollendo, Weberbauer 1509. Mejia, 200 meters 
(Gunther & Buchtien 306, fide Bruns); also at 1,000 meters, at 
Cachendo. 

Calandrinia ciliata (R. & P.) DC. Prodr. 3: 359. 1828. Talinum 
ciliatum R. & P. Syst. 1: 116. 1798. C. caulescens HBK. Nov. Gen. 
& Sp. 6: 78. pi. 526. 1823. Talinum caulescens Spreng. Syst. 2: 453. 



570 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

1825. Phacosperma peruviana Haw. Phil. Mag. 1: 124. 1827(?). 
C. Phacosperma DC. loc. cit. 

A little annual, at first cespitose-tufted, the stems in fruit develop- 
ing to several cm. long; leaves linear-spatulate, to about 3 cm. long, 
often glabrous; flowers purple, or white fading to purple, subsessile 
or pediceled, solitary, axillary; capsule scarcely exserted; seeds tur- 
gid-lenticular, with a thin margin, lustrous, scarcely prominently 
puncticulate. The leaves of the type at Madrid are essentially 
glabrous, the few cilia being obscure, or lacking on some leaves. 
The original description reads: Herb to about 30 cm. high: leaves 
linear-oblong, ciliate; flowers solitary. Neg. 8433. 

Arequipa: Open gravel on rocky slopes, 4,000 meters, Nevado de 
Chachani, Pennell 13277. Mollendo, Johnston 3546 (det. Johnston 
as C. caulescens). Moquehua: Torata, 2,700 meters, Weberbauer 
7415. Lima: Banos and Obrajillo (Wilkes Exped.}. Viso, 2,700 
meters, 598. Junin: Huasa-huasi, Ruiz & Pavon; Dombey. Near 
Oroya, BalL Huanuco: Mito, 2,700 meters, 1719. Cajamarca: 
Cutervo, Jelski 245. 

Calandrinia crenata (R. & P.) Macbr. Field Mus. Bot. 11: 20. 
1931. Talinum crenatum R. & P. Syst. 115. 1798. 

An herb 30 cm. high with subrotund-obovate, acuminate leaves 
and racemose flowers with crenulate petals. Apparently unknown, 
there being no specimens at Madrid, and perhaps not distinct from 
C. Ruizii. Its characters may be proved or disproved by exploration. 

Arequipa: On the hills about Pongo, Atiquipa, April-September 
(Ruiz & Pavon}. "Lengua de vaca." 

Calandrinia lingulata (R. & P.) DC. Prodr. 3: 358. 1828. 
Talinum lingulatum R. & P. Syst. 115. 1798. 

An herb about 20 cm. high, branching from the base, with gla- 
brous, lingulate leaves 2-3 (-4) mm. wide; racemes crowded; bracts 
2 mm. long; stamens 10-15; pedicels ascending, 5-10 mm. long; 
sepals 3-4 mm. long; capsule finally about 5 mm. long. There is 
only one capsule on the Madrid specimen, but this seems to be well 
exserted. One sheet of T. ciliatum, as labeled in Herb. Boiss., is evi- 
dently the same as the type at Madrid of T. lingulatum. Neg. 8435. 

Lima: In sands, Chancai at Jequan, Ruiz & Pavon; Dombey. 

Calandrinia linomimeta Diels, Bot. Jahrb. 37: 400. 1906. 
Suffrutescent below, with strict, herbaceous branches 10 cm. tall 
or taller that are leafless above and often forked at the racemose 



FLORA OF PERU 571 

inflorescence of showy, roseate flowers; leaves crowded, oblanceolate, 
mucronulate; pedicels in fruit 1-1.5 cm. long, recurved; petals 1.5 
cm. long; stamens many; stigmas 3-parted; capsules almost bright 
yellow, 6-7 mm. long, 5 mm. thick; seeds estrophiolate, lustrous, 
smooth. As the author notes, suggestive of Linum. Neg. 27676. 

Amazonas: Prov. Luya, stony slopes, 900 meters (Weberbauer 
4788; 155). 

Calandrinia pachypoda Diels, Bot. Jahrb. 37: 399. 1906. 

A fleshy perennial with very thick, branching, procumbent stems 
and densely leafy, ascending flowering branches about 10 cm. high; 
leaves 1.5-3 cm. long, 6-10 mm. wide, narrowed to a petiole-like 
base, scarcely acute but often apiculate; flowers 3 or 4, white, about 
1 cm. long; sepals acuminate, half as long; stamens many; seeds dis- 
tinctly muriculate, minutely strophiolate. Suggests Talinum crassi- 
folium Willd. (type poor), but the leaves are narrower. Stony slopes. 
Neg. 27678. 

Cajamarca: East of Chepe"n, 800 meters, Weberbauer 4816 (type); 
153. Junin: Weberbauer 5280(1}. Lima: Chosica, 900 meters, 
493, 2875. 

Calandrinia paniculata (R. & P.) DC. Prodr. 3: 358. 1828. 
Talinum paniculatum R. & P. Syst. 114. 1798. C. adenosperma I. M. 
Johnston, Contr. Gray Herb. 81 : 89. 1928. 

A stout, glabrous, glaucous perennial, sometimes over 40 cm. 
high; leaves obovate-oblong, acuminate, fleshy, the largest 10 cm. 
long and 6 cm. wide; flowers racemose, the pedicels 3-7 times longer 
than the conspicuous bracts; sepals suborbicular, 8-10 mm. long; 
petals red, to about 1 cm. long, capsule longer than the calyx; 
seeds black, opaque, densely hispidulous or glandular-echinate. Very 
near C. grandiflora Lindl., typically, at least, with larger flowers, 
smaller bracts, and eglandular seeds; but Esposto 16 had yellow 
flowers 2.5 cm. wide. Neg. 29565. 

Arequipa: Pongo, Atiquipa (Ruiz & Pavori). Chachani, 2,700 
meters, Hinkley 35, type of C. adenosperma. Cerros de Caracoles, 
Esposto 16. Arequipa: Hopp 6. Mt. Chacani, 2,745 meters, Hinkley 
35. Lima: Lomas at Chancai, Ruiz & Pavon (type); Dombey. 
"Oreja de perro," "castanuela," "lloto del cerro." 

Calandrinia quivensis Macbr. Field Mus. Bot. 11: 21. 1931. 
An annual or biennial, strictly erect and more or less branched 
above the base, several dm. high; leaves crowded, linear, to about 



572 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

4 cm. long, 2-3 mm. wide; racemes often branching; pedicels spread- 
ing in fruit, 5-7 mm. long; petals purple, 5 mm. long; sepals orbicular, 
2.5 mm. long, the capsule nearly twice as long; seeds opaque, muric- 
ulate, minutely strophiolate. Very possibly this is well developed, 
mature material of C. lingulata. 

Lima: Open, rocky slope, 900 meters, Quive, Pennell 14-299 
(type). Chorrillos, 150 meters, 5850. 

Calandrinia Ruizii Macbr. Field Mus. Bot. 11: 20. 1931. Tali- 
num polyandrum R. & P. Syst. 115. 1798, not C. polyandra Benth. 

Stems ascending-decumbent below and there clothed with 
broadly obovate or subrotund leaves, originally described as "acumi- 
nate" but, so far as seen, merely apiculate; flowers pink, racemose; 
pedicels finally becoming several cm. long; stamens many; petals 
obcordate, entire, less than 1 cm. long; capsule a little exserted; seeds 
black, minutely and sparsely white-hispid. Very fleshy plants of 
disintegrating rocky slopes, sometimes 30 cm. high. Type not seen; 
Weberbauer specimens determined at Berlin. 

Arequipa: Lomas near Pongo (Ruiz & Pavori). Mollendo, Weber- 
bauer 1484,' 144. Tacna: Woitschach. Lima: Barranco, Weberbauer 
1598; 144. Chorrillos, 150 meters, 5849. Matucana, 2,400 meters, 432. 

Calandrinia Weberbaueri Diels, Bot. Jahrb. 37: 399. 1906. 

Annual or biennial, much branched at the base, the ascending or 
erect stems a few cm. to several dm. high; flowering branches leafless; 
pedicels slender, 5-8 mm. long; sepals 3 mm. long; petals purple; 
capsule 4 mm. long. Nearly C. lingulata, except for the crowded, 
almost umbellate flowers. 

Arequipa: In sand, Mollendo, Weberbauer 1507 (type). Moque- 
hua: Between Moquehua and Torata, 2,000 meters, Weberbauer 
7432. Lima: Below Cocachachua, Weberbauer 5269. Near Chosica, 
1,000 meters, Weberbauer 5221 (leaves broader). 

4. MONTIA L. 

Reference: Briquet, Prodr. Fl. Cors. 1: 474-476. 1910. 

Our species a small, ascending or procumbent, sometimes tufted, 
branching annual (or rarely more enduring) with opposite leaves and 
axillary or terminal, few-flowered racemes of tiny flowers. Petals 
white, unequal, connate at the base but the tube open on one side. 
Seeds few. The rather similar Monocosmia monandra (R. & P.) 
Pax is Chilean, not Peruvian, as has been sometimes indicated. 
Urban observed the remarkable ejection of the seeds: the three 
capsule segments upon opening at once curl up from beneath, thus 



FLORA OF PERU 573 

pressing together and raising the seeds which are ordinarily thrown 
50-80 cm., or even as far as 2 meters and as high as 60 cm. 

Montia fontana L. Sp. PI. 87. 1753. 

Leaves obovate to narrowly spatulate, from minute to 12 mm. 
long, with a petiole-like base; calyx scarcely 2 mm. long, the petals 
little longer, the globose pod included. There are two or three 
forms of which only one, var. repens Pers. Syn. 1: 111. 1805 (M. 
rivularis Gmel. Fl. Bad. 1: 301. 1805), has apparently been found in 
Peru. It has somewhat lustrous seeds with tubercles developed in 
varying manner and degree. Also to be expected is var. erecta Pers. 
(M. minor Gmel.) with dull, lineately muricate seeds. In water or 
very wet places in cool regions of the world. 

Puno: Sachapata, Lechler 2688. Huanuco: Six miles south of 
Mito, 3,150 meters, in bog, 1831. Tambo de Vaca, 3,900 meters, 
edge of forest, 4402. Cuzco: Paso de Tres Cruces, 3,700 meters, 
streamlet in paramo valley, Pennell 13908; an aquatic herb; petals 
white. La Raya, 4,500 meters, in Distichia cushions, Pennell 13499 
(var. repens}. 

5. PHILIPPIAMRA Kuntze 

Silvaea Phil. Fl. Atac. 21. pi. 1J.C. 1860, not H. & A. 1836. 

A widely branching herb with small flowers crowded in apical 
inflorescences. Involucral leaves persistent. Sepals 3-4. Stamen 
1. Style 2-toothed. Fruit not splitting explosively nor irregularly 
dehiscent, with 1 seed. 

Philippiamra pachyphylla (Phil.) Kuntze, Rev. Gen. 1: 58. 
1891. Silvaea pachyphylla Phil. loc. cit.; Viaje Des. Atac. 38, 
195. 1860. 

Well marked by the nearly bushy habit and the conspicuous, 
scarious bracts, often tinted, about the glomerulate flowers. There 
are several closely related species in northern Chile to be expected, 
which, however, Johnston (Contr. Gray Herb. 85: 39. 1929) thinks 
are probably all forms of one species. The leaves are sometimes 
nearly amplexicaul, sometimes petioled. 

Tacna: Werdermann 231 (det. Werdermann). Chile. 

66. BASELLACEAE. Basella Family 
Reference: Ulbrich, Pflanzenfam. ed. 2. 16c: 262-271. 1934. 
These low, glabrous, somewhat twining plants with rather fleshy, 
obovate leaves and axillary spikes or racemes of tiny flowers, are 



574 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

important in Peru as furnishing the tubers called "ullucos," com- 
monly cultivated and eaten like potatoes. The remains of them 
have been found, apparently, in old Peruvian graves (Harms). 
Flowers sessile; filaments straight; petals not caudate. . .1. Basella. 
Flowers pedicellate (Peruvian forms) ; filaments curved or the petals 

caudate. 
Filaments in bud straight; petals tailed; bractlets prominent. 

2. Ullucus. 
Filaments in bud curved; petals blunt or not tailed. 

Floral bractlets not dorsally winged in fruit . . 3. Boussingaultia. 
Floral bractlets with a wide, fin-like wing in fruit . . 4. Anredera. 

1. BASELLA L. 

Succulent, branching, twining herbs with spikes of tiny, tubular, 
white, reddish, or purplish flowers. Bracts minute, caducous. Calyx 
lobes 2. Anthers extrorse. 

Basella alba L. Sp. PI. 272. 1753. B. rubra L. loc. cit. 

Often 1 meter long; leaves sometimes nearly round, several 
cm. to 15 cm. long, brittle when fresh; corolla lobes 2 mm. long, 
obtuse; fruit red, black, or white. B. rubra, according to Ulbrich, is 
only the more'or less reddish-colored race of this variable plant, which 
has been in cultivation since 1688 and is used as a vegetable or salad. 
Illustrated, Pflanzenfam. ed. 2. 16b: 267. 

Lima: Matucana, Weberbauer 5274 (det. Ulbrich); at 2,400 
meters, trailing over rocks, 166. Viso, Weberbauer 156 (det. Ulbrich). 
Chorrillos, 150 meters, stony slopes, seaside hills, 5851. Junin: 
Tarma, 3,100 meters, open hillside, Killip & Smith 21815. Arequipa: 
Hopp (det. Ulbrich). Arequipa, 2,800 meters, open, rocky slopes, 
Pennell 13248. Moquehua: Raimondi. Native of Asia or Africa, 
widely established after cultivation. 

2. ULLUCUS Lozano 

An herb with potato-like tubers, at first erect but soon sprawling 
and somewhat twining, succulent. Perianth parts spreading, 
narrowed abruptly from an oval base to an elongate, nearly linear, 
appendage-like tip. Anthers dehiscent apically. Important as a 
food plant. In the cultivated form almost bushy in habit. The 
tubers attain the size of small potatoes, but are usually more elon- 
gate, and generally pale yellow with violet "eyes." When cooked 
they are rather sweet and palatable but "wet." The Peruvian 



FLORA OF PERU 575 

peasants freeze them after soaking them in water, then they are 
sun-dried or pressed and dried, for preservation for an indefinite 
period, in the same manner in which they treat potatoes. After this 
treatment they resemble "chufio" and like it emit an odor in cooking 
that is very disagreeable to one not accustomed to it, but after some 
experience, as in the case of the "chuiio" or potato so preserved, they 
may be eaten with some relish, at least upon necessity. Herrera 
records the name "lingli" for the dried product; Pflanz, "tunta." 
Ruiz and Pavon found them a common food but eaten also as an aid 
in child-birth. 

Ullucus tuberosus Lozano in Caldas, Semanario de Nueva 
Granada 185. 1809. Basella tuberosa HBK. Nov. Gen. & Sp. 2: 
189. 1817. Melloca pemviana Moq. in DC. Prodr. 13, pt. 2: 225. 
1849. U. Kunthii Moq. Bibl. Univ. Geneve 11: 80. 1849. 

Leaves long-petioled, cuneate to subcordate at the base, rounded 
or obtuse at the apex, often mucronulate, sometimes 20 cm. broad; 
racemes of greenish yellow flowers shorter than the leaves, bracteo- 
late, the lower bractlets longer than the pedicels. According to 
Ulbrich, the tubers of the wild form are the size of pigeon eggs, rose- 
violet in color, the plants developing little tubers in the leaf axils and 
attaining a length of 50 cm. The species is variable, and Herrera, 
Contr. Fl. Cuzco ed. 2. 70-72. 1921, has recorded the native names 
of cultivated and wild races as they occur in that department 
as follows: 

Tubers white: Mestiza chchuccha, yurac-lisas, or yurac-ckolla, 
the last with tubers produced in abundance but very small. 

Tubers yellow: Ckello-chuccha, ckello-lisas, or ckello-ckolla. 

Tubers yellowish red: Bela-api-chchuccha. 

Tubers rose-colored: Muru-chchuccha, muru-lisas, or muru- 
ckolla, the last large. 

Tubers reddish : Puca-lisas. 

Herrera notes further, without indication of color: Chucchan- 
lisas, long and of superior quality: Ckolla-lisas, considered inferior; 
Kitalisas (wild "papas lisas"), extremely bitter, not edible; and Atoc- 
lisas or kipa ullucu (kipa ul juku), (wild smooth potatoes), esteemed 
fresh. The former has been found in the provinces of Paucartambo, 
Urubamba, and Huaipo, the latter in Anta, Quispicanchi, and 
Urubamba. The species is said to ascend to over 3,800 meters in a 
wild state in Bolivia, and in Peru it is found in the central sierra 



576 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and interandean regions (Weberbauer), where especially frequent in 
cultivation. Illustrated, Pflanzenfam. ed. 1. 16c: 267. 

Lima: Near Tambo de Viso, Weberbauer 147; 169. Rocky places 
above Chicla (Ball). Matucana, Weberbauer 170. Rio Blanco, 
4,500 meters, 3044. Without locality (MacLean, type of M. peruvi- 
ana). Colombia; Bolivia. "Papa lisa," "lisas," "ullucu," "ul juku." 

3. BOUSSINGAULTIA HBK. 

Tandonia Moq. in DC. Prodr. 13, pt. 2: 226. 1849. 

Reference: Hauman, Anal. Mus. Nac. Buenos Aires 33: 347-359. 
1923-1925. 

Glabrous, more or less twining, rather sparsely leaved herbs, 
usually drying almost black, including the axillary racemes of small 
flowers. Sepals blunt, shell-shaped. Style 1, often basally 3-parted. 
Fruit included in the sometimes keeled sepals or bractlets. United 
by Baillon, Hist. PI. 9: 198. 1888, with Anredera Juss., with which 
it naturally belongs, but unfortunately, from a practical standpoint, 
the union now is inconvenient, since all the known species except 
one have been described under the later name of HBK. 

Leaves definitely cordate; pedicels filiform B. filiformis. 

Leaves little or not at all cordate; pedicels thickish, short. 
Leaves not oblongish; racemes usually simple, the flowers often 
tinted. 

Leaves broadly elliptic .B. diffusa. 

Leaves obovate-rhombic B. minor. 

Leaves oblong-lanceolate; racemes often branched; flowers white. 

B. Weberbaueri. 

Boussingaultia diffusa (R. & P.) Volkens, Pflanzenfam. 3, la: 
128. 1893. Basella diffusa R. & P. in lit. in syn. Tandonia diffusa 
Moq. in DC. Prodr. 13, pt. 2: 226. 1849. 

Leaves broadly elliptic, acuminate, about 3 cm. wide; racemes 
spike-like, several cm. to 10 cm. long; sepals narrowed to the base; 
bractlets longer than wide; flowers tiny, subsessile, whitish, yellow- 
ish, or purplish. B. Volkensii Ulbrich in Engl. & Prantl, Nat. 
Pflanzenfam. ed. 2. 16c: 269. 1934 (B. cordifolia Volkens), B. obovata 
(HBK.) Haum. Anal. Mus. Nac. Buenos Aires 33: 352. 1925, and B. 
marginata (HBK.) Britton, all of Ecuador, may be expected; the 
first has sessile flowers and keeled sepals; the others broad-based 
sepals, B. obovata with obovate, obtuse leaves, B. marginata with 



FLORA OF PERU 577 

leaves oval and acute. B. boliviensis (Hauman) Macbr., comb. nov. 
(B. ramosa Hemsl. var. boliviensis Hauman, op. cit. 353), to be 
expected, has keeled sepals like B. Volkensii but pediceled flowers; 
it differs from B. ramosa of Mexico in having bractlets about twice 
as long as the pedicels, lighter-colored, larger leaves, etc. To B. 
diffusa apparently belongs all Peruvian material formerly referred to 
B. baselloides HBK., which has pedicellate flowers fully 2 mm. long 
with oblong sepals similar to the petals. Often cultivated, it may 
well occur, as also B. cordifolia Tenore (B. gracilis Miers, f. pseudo- 
baselloides Hauman, Anal. Mus. Nac. Buenos Aires 33: 356. 1925, 
probably the same according to Harms) confused with it in literature 
but readily distinct by its 3-parted style. The fleshy rhizomes are 
a common food (Ruiz & Pavon). The plant ranges from the coast 
to over 2,000 meters in the sierras. 

Lima: Huara, Lima, and Pillao, Ruiz & Pawn. Chosica, Weber- 
bauer 5316 (det. Ulbrich). San Agustin, Weberbauer 5229 (det. 
Ulbrich). Matucana, Weberbauer 161. Amazonas: Between Cha- 
chapoyas and Moyobamba, Raimondi. Ayacucho: Chavina, Weber- 
bauer 5787. Ancash: Ocros, Weberbauer 2730. Cuzco: Raimondi. 
Ollantaitambo, 2,800 meters, Herrera 3410. Arequipa: Above Posco 
(Gunther & Buchtien 302; det. Bruns as B. baselloides). Southern 
slopes of Chachani Mountain, 3,660 meters, Hinkley 78. Huanuco: 
San Rafael, 2,550 meters, 3144- Cabello, 2,400 meters, 1337. 
"Lloto del cerro." 

Boussingaultia filiformis (Moq.) Hauman, Anal. Mus. Nac. 
Buenos Aires 33: 352. 1925. Tandonia filiformis Moq. in DC. 
Prodr. 13, pt. 2: 227. 1849. 

Well marked by its slender pedicels as long as the flowers, 1.5 
mm. long, its triangular, obtuse sepals, and especially by its cordate 
leaves, acute at the apex. 

Peru: Without locality, Ruiz & Pavon. 

Boussingaultia minor Diels, Bot. Jahrb. 37: 400. 1905. 

Slender, scarcely twining, with subsessile, oblong-ovate or rhombic- 
ovate leaves, long-cuneate at the base, 2-3 cm. long, 6-15 mm. wide; 
spikes shorter or little longer than the leaves; flowers sessile, the 
unequal, concave segments 2 mm. long, white within, reddish 
without; bracts lanceolate, acute; ovary nearly 4 mm. long, the 
single style about twice as long. 

Junin: Tarma, 3,000 meters, Weberbauer 2351. 



578 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Boussingaultia Weberbaueri Ulbrich, Pflanzenfam. ed. 2. 
16c: 270. 1934. 

A twining half shrub; leaves lanceolate, acute; involucral leaves 
plane; flowers white, 2-2.5 mm. long, abundant in peduncled, often 
much branched racemes; style 1; stigmas 3, strongly papillose. 
Neg. 29513. 

Cajamarca: Huambos, 2,700-3,000 meters, Weberbauer 4185, 
type. Huanuco: San Rafael, 2,550 meters, 3143. Lima: San 
Buenaventura, 2,800 meters, open, rocky slopes, Pennell 14508. 

4. ANREDERA Juss. 

Similar to Boussingaultia HBK., but restricted to the one original 
species, characterized by densely flowered racemes about 8 mm. 
thick, the conspicuously pediceled flowers yellowish and somewhat 
translucent as are also the straw-colored, dorsally winged fruiting 
sepals. Unlike species of Boussingaultia, this plant dries light green. 

Anredera scandens (L.) Moq. in DC. Prodr. 13, pt. 2: 230. 
1849. Polygonum scandens L. Sp. PL 364. 1753. 

Leaves narrowed at each end, acute; style completely 3-parted. 
The tuberous roots are used for food. Cultivated, especially in 
Spain. 

Peru: (Weberbauer, 307). North to Texas and the West Indies. 

67. CARYOPHYLLACEAE. Pink Family 

Reference: Pax & Hoffmann, Pflanzenfam. ed. 2. 16c: 275-364. 
1934. 

Herrera, in one of his several useful "Contributions to the Flora 
of the Department of Cuzco," has recorded Dianthus Caryophyllus 
L., the carnation or "clavel" as cultivated in abundance in gardens 
for its beauty and for making perfume, and far to the north in the 
Department of Ancash, near the coast, I saw it grown in rows many 
rods in extent for the same purposes. Most of the genera, as 
Pax and Hoffmann state for Paronychia, "bedarf dringend einer 
systematischen Bearbeitung," so that the following synopsis of the 
Peruvian genera is largely based on scattered references and on the 
partial revisions of Rohrbach and Williams. Mattfeld in 1922, 
Repert. Sp. Nov. 18: 171, announced a report on the Muschler 
species based on Weberbauer plants, but none has been forthcoming 
except for one genus (Pycnophyllum). Accordingly, except for a 
few of the Muschler species seen, they have not been included in the 



FLORA OF PERU 579 

keys or new combinations made, for his descriptions are unreliable 
and his names, as Mattfeld suggests, op. cit., may be considered 
nomina nuda. Nevertheless, since they often represent new species, 
it seems best to give the apparently salient characters from his 
descriptions, sometimes correct, in order to have a record of the 
collections to which they refer. 

The genera of this family are for the most part intangible con- 
cepts, often not separable or tenable on the basis of logic which 
is one reason why Plettkea Mattfeld is questionably a valid or 
expedient segregate because the species themselves are plastic 
(see note under Stellaria media}. All generic keys, therefore, as the 
following, are only suggestive, and one should be warned that there 
are exceptions in most characters for most genera. In addition to 
the genera listed here, Corrigiola L. may occur; it is rather similar 
to Paronychia, but the stipules are deciduous below or minute, and 
the styles 3 instead of 2. Heliospermum (Heliosperma Reichenb.), 
accredited to Peru by Weberbauer, 175, 354, must be a reference to 
some species of Silene sens. lat. (Melandrium or Lychnis). The hairy 
Agrostemma Githago L., of Europe, with linear calyx lobes that well 
exceed the rather showy petals, may be found adventive in cultivated 
fields, especially in stands of maize or other grain. A number of 
species, especially those proposed by Muschler, are listed alpha- 
betically in the genera to which they belong without, however, 
making the transfer. 

It is fitting to acknowledge here the kindness of Dr. Charles 
Baehni in making herbarium comparisons and careful dissections 
for me. 
Sepals slightly if at all united. 

Plants without stipules (cf. Drymarid). 

Fruit capsular; plants scarcely if at all cushion-forming, some- 
times densely matted. 

Plants mostly low-matted, or with congested inflorescences; 
capsule teeth of the same number as the styles, or in 
Arenaria twice as many. 

Petals entire or merely lobed 5. Arenaria. 

Petals, if present, minute 1. Sagina. 

Petals none 2. Colobanthus. 

Plants mostly open-growing and open-flowering; capsules 
with twice as many teeth as styles. 



580 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petals 2-lobed; pods dehiscent by twice as many teeth as 

styles 3. Cerastium. 

Petals 2-parted or 2-lobed; pods splitting into as many 

valves as styles 4. Stellaria. 

Fruit not capsular; a cushion-like plant. . . .6. Pycnophyllum. 
Plants usually with stipules. 
Fruit not capsular; dense herbs or half-shrubs with conspicuous, 

silvery or white stipules. 
Sepals abruptly spinulose from a mostly cucullate tip. 

7. Paronychia. 

Sepals gradually spine-tipped 8. Cardionema. 

Fruit capsular; plants more or less openly branched, or simple 

herbs, the stipules relatively inconspicuous. 
Styles more or less united; leaves often broader than linear. 
Stipules minute, often caducous; inflorescence usually 
openly cymose, sometimes head-like .... 9. Drymaria. 
Stipules rather obvious; flowers densely cymose. 

Sepals keeled 10. Polycarpon. 

Sepals not keeled 11. Polycarpaea. 

Styles free; leaves linear 12. Spergularia. 

Sepals definitely united, forming a tube 13. Silene. 

1. SAGINA L. 

Small or diminutive herbs bearing several long-stalked, whitish 
flowers. Stamens in one or two whorls. Petals sometimes obsolete. 

Sagina apetala Ard. Animadv. Bot. 2: 22. pi 8. 1763. 

A tiny annual, rarely more than a few cm. tall, with filiform 
stems and long, capillary pedicels, these straight or slightly curved 
at maturity; leaves setaceous, 3-7 mm. long, sometimes puberulent 
or with a few cilia at the base; sepals 4 and 1.5 mm. long, all or 
some of them obtuse; petals evanescent, minute, sometimes appar- 
ently lacking; capsule 2 mm. long. The following specimen was 
recorded by Weberbauer, 171, as S. ciliata Fries, of Europe, more 
properly var. ciliata (Fries) Mert. & Koch., with curved pedicels 
and appressed sepals. I did not find the specimen at Dahlem. The 
cosmopolitan S. procumbens L., with 4 or 5 obtuse sepals (sometimes 
with some flowers with 4, others with 5 on the same plant!), is dis- 
tinguished otherwise by its creeping habit, the stems rooting and 



FLORA OF PERU 581 

tending to become perennial. S. saginoides (L.) Dalla Torre (S. 
Linnaei Presl), with curved pedicels and floral parts always in 5's, 
may occur; its sepals, according to Fernald, are typically 2-3 mm. 
long, its capsules 3.4 mm. long. S. humifusa (Camb.) Fenzl, of 
Brazil and perhaps eastern Peru, also pentamerous, has petals and 
calyx 2 mm. long. 

Ayacucho: Pampalca, 3,200 meters, Killip & Smith 23281. 
Cajamarca: Meadow moor, 3,500 meters, Ocros (Weberbauer, 171?; 
see note above). Ecuador; Canary Islands; Australia; South 
America (Pax & Hoffmann). 

2. COLOBANTHUS Bartl. 

Thick-rooted plants bearing a solitary flower, the short stem or 
peduncle thickened above; stamens 4-5, alternate with the same 
number of sepals. Illustrated, Presl, Rel. Haenk. 2: pi. 4-9. 

Colobanthus crassifolius (D'Urv.) Hook. f. Fl. Ant. 248. 
1844-47. Sagina crassifolia D'Urv. Me"m. Soc. Linn. Par. 4: 617. 
1826. C. quitensis Bartl. in Presl, Rel. Haenk. 2: 13. 1831. S. 
quitensis HBK. Nov. Gen. & Sp. 6: 19. 1823. 

A little, tufted plant with grass-like, linear, rigid or soft leaves 
1-1.5 cm. long or sometimes longer, and slender scapes as long or a 
little longer, bearing the single flower; sepals acutish, ovate-oblong; 
capsule exserted. The following collection, determined by Gray as 
C. crassifolius Hook, f., apparently lacked the bracts sometimes 
borne on the scape or peduncle; the plant is variable in size. 

Junin: Casa Cancha (WilkesExped.). East of Canta, open slope, 
Pennell 14671. Colombia to Patagonia. 

3. CERASTIUM L. 

Reference; Rohrbach, Linnaea 37: 283-308. 1872. 

Plants usually pubescent, often densely so, the Peruvian forms 
frequently perennial. Flower parts regularly 5 or infrequently 
reduced to 3 or 4, or the stamens from 10 to 5 or 4. Capsule most 
often curved, especially when exserted. Descriptions of the older 
forms are mostly taken directly from Rohrbach. F. N. Williams 
published a partial list with notes in Journ. Bot. in 1898, 1899, 
1921, 1922 (not only in the first three years as cited in Pflanzenfami- 
lien), which has been referred to, as likewise a partial key in Bull. 
Herb. Boiss. I. 6: 893. 1898. I am indebted also to Fernald & 
Wiegand, Rhodora 22: 171, 177, 178. 1920, for the key characters 



582 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

and descriptions of C. arvense and C. vulgatum. The key otherwise 
is based in large part on descriptions. The types of Muschler species, 
not seen by me, are especially dubious. Herrera has listed for 
Cuzco (Herrera 2114, Valle de Santa Ana) a "C. biflorum Poepp.," a 
name I have not found. 
A. Perennials. 

Bracts broadly scarious-margined. 
Basal branches herbaceous, with few axillary tufts; petals and 

sepals subequal C. vulgatum. 

Basal branches finally marcescent, the axillary tufts conspicuous; 

petals 2-3 times as long as the sepals C. arvense. 

Bracts herbaceous, or the uppermost very narrowly scarious- 
margined. 

Plants low or diminutive, pulvinate or densely tufted, the parts 

above ground only 1-4 cm. high, or the stems, if longer, 

creeping-assurgent, densely clothed with dead leaves; 

leaves soft, thin (cf. C. vulgatum var. andinum Gray). 

Leaves more or less pubescent or glabrous, never white- 

tomentose. 

Capsule well exserted; pedicels enlarged toward the base. 

C. crassipes. 

Capsule often scarcely if at all exserted; pedicels slender. 
Plants densely pulvinate. 

Plants more or less pubescent, often robust. 

Calyx 2-4 mm. long C. Behmianum. 

Calyx 5-8 mm. long C. soratense. 

Plants essentially glabrous, diminutive. . . .C. nanum. 
Plants rather loosely tufted, the stems often assurgent. 

Sepals narrowly oblong C. imbricatum. 

Sepals elliptic or elliptic-oblong C. peruvianum. 

Leaves closely white-tomentose, glabrate only in age. 

C. candicans. 

Plants often cespitose but open in habit, the principal, erect or 
suberect stems to about 10 cm. high or higher; leaves rather 
firm or somewhat rigid (see also C. trichocalyx, not included 
in the key). 

Plants densely white-lanate or floccose-pubescent throughout. 
Inflorescence soon open; stem leaves scarcely imbricate. 

C. mollissimum. 



FLORA OF PERU 583 

Inflorescence crowded; stem leaves well imbricate. 

C. floccosum. 

Plants green but often more or less villous. 
Flowers crowded, pseudo-umbellate, terminal. 
Sepals and often the leaves glandular-pilose. 

Capsule well exserted; pedicels more or less nodding or 

refracted. 

Leaves densely short-pubescent, more or less revolute. 

C. Danguyi. 
Leaves sparsely and loosely pilose, plane. 

C. Trianae. 

Capsule included or slightly exserted; pedicels erect. 

C. mucronatum. 
Sepals eglandular, the leaves glabrous above. 

C. tucumanense. 

Flowers axillary, solitary, few C. subspicatum. 

A. Annuals (C. subspicatum as a biennial might be sought here). 

Petals ciliate below C. viscosum. 

Petals glabrous. 
Capsule teeth erect; plants more or less viscid-pubescent. 

C. nutans. 
Capsule teeth recurved-revolute; plants glabrate. 

C. humifusum. 

Cerastium arvense L. Sp. PI. 1: 438. 1753. 
A matted or tufted perennial, with depressed or trailing, tough 
basal branches bearing marcescent leaves and abundant axillary 
fascicles or leafy tufts; flowering branches ascending, simple to freely 
branched, 2-60 cm. high, glabrous to densely villous, glandless to 
densely glandular; leaves linear-subulate to narrowly ovate, flaccid 
to rigid, acute to obtuse, glabrous to velutinous, glandless or glandu- 
lar, 1-6 cm. long, 0.5-13 mm. broad, mostly confined to the lower 
two-thirds of the branch; inflorescence few- to many-flowered, its 
bracts scarious-margined; sepals 4.5-8.5 mm. long, glabrous, pilose, 
or glandular; petals 2-3 times as long as the sepals, the broad lobes 
spreading in an thesis, the claw glabrous; capsule cylindric, equaling 
to much exceeding the calyx; seeds reddish, 0.3-0.7 mm. in diameter, 
the testa close and tuberculate. Description by Fernald & Wiegand, 
Rhodora 22: 178. 1920. Forma oblongifolium (Torr.) Pennell, with 
elongate capsule, near Lima according to Weberbauer, 169, is scarcely 



584 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

distinct, as also C. arvense var. arvensiforme (Wedd.) Rohrb. (C. 
arvensiforme Wedd. Ann. Sci. Nat. V. 1: 296. 1864), accepted by 
F. N. Williams, which is, apparently, a large-leaved, large-flowered 
form, the sepals 6-8 mm. long; var. nervosum (Naud.) Reiche, Fl. 
Chile 1: 184. 1896, glandular with elliptic-oblong leaves 4-8 mm. 
long, seems better marked. Neg. 27711 (var. nervosum). 

Puno: Titicaca highlands (Weberbauer, 219 as C. nervosum). 
Lima: At about 3,000 meters (Weberbauer, 169, as C. oblongifolium). 
North and South America; Eurasia. 

Cerastium Behmianum Muschl. Bot. Jahrb. 45: 446. 1911. 
C. imbricatum HBK. var. Mandonianum Rohrb. Linnaea 37: 293. 
1872. 

Stems ascending, 3-5 cm. long, sparsely if at all branched, becom- 
ing glabrous; leaves at first rather pubescent, somewhat ciliate, 
finally glabrate, oblong or ovate-oblong, acutish or rarely obtusish, 
0.5-2 cm. long, 1-3 mm. wide; flowers few, solitary, on pubescent 
pedicels, in fruit semi-erect and somewhat longer than the calyx; 
sepals oblong, obtuse or acutish, scarious-margined, apically gla- 
brous, equaled by the petals; capsule scarcely exserted, the teeth 
revolute; seeds tuberculate, with inflated epidermis. Description 
drawn in part from an apparent cotype in Herb. Delessert. Negs. 
29833, 29834. 

Junin: Yauli, 4,400 meters, Weberbauer 558. Bolivia. 

Cerastium candicans Wedd. Ann. Sci. Nat. V. 1: 295. 1864; 302. 

Stems ascending or suberect, many, some of them sterile, crisply 
short-pubescent, 3-10 cm. high; leaves narrowly lanceolate, acute, 
white-tomentose but becoming smooth above, 1.5-2 cm. long, 3 mm. 
wide; flowers few, loosely umbellate, the tomentose fruiting pedicels 
longer than the lanceolate, acute sepals, these only 5 mm. long, 
dorsally tomentose, apically glabrous, scarious-margined; petals as 
long as the sepals, emarginate; stamens 10; capsule cylindric, to 
nearly twice as long as the calyx; seeds obtusely granulate. 
Neg. 27707. 

Puno: Titicaca highlands, 4,400 meters (Weberbauer, 219). 
Ecuador. 

Cerastium crassipes Bartl. in Presl, Rel. Haenk. 2: 18. 1831; 
296. C. orophilum Wedd. Ann. Sci. Nat. V. 1: 295. 1864 (fide 
Rohrbach). 



FLORA OF PERU 585 

Stems crowded, cespitose, diffuse, crisply white-strigose or pilose, 
as also the leaves, peduncles, and sepals; lower rosulate leaves linear- 
oblanceolate, 15-25 mm. long, 2-4 mm. wide, the upper shorter and 
broader, acuminate; inflorescence glomerulate and finally umbellate; 
pedicels 2-3 times longer than the calyx, strongly enlarged below the 
middle, incurved and glabrescent in fruit; sepals oblong, obtuse, 
scarious-margined, 4-5 mm. long; stamens 5 or 10; petals slightly 
longer, bifid one- third their length, scarcely clawed; capsule half 
again as long as the calyx; seeds large, densely and coarsely tuber- 
culate. Sometimes a tiny plant, as the Weddell specimen, again 
robust with fistulose stems 15 cm. long and leaves 3 cm. long (a 
Chilean form). The Weddell species is maintained by Pax and 
Hoffmann in Pflanzenfamilien. Neg. 27709. 

Huanuco: Montana (Haenke, type). Puno (?): Macusani 
(Lechler). Junin: Casapalca, 5,000 meters, 840. Bolivia. 

Cerastium Danguyi Macbr., nom. nov. C. Trianae Danguy & 
Cherm. Bull. Mus. Hist. Nat. Paris 28: 432. 1922, not Briq., 1911. 

Stems ascending, 25-40 cm. high, with the leaves shortly but 
densely pubescent, more or less glandular toward the tip; leaves 
numerous, the lower somewhat crowded, the upper about equaling 
the internodes, linear-lanceolate, 1.5-2 cm. long, 3^4 mm. broad at 
the base, acute, the margin manifestly revolute; inflorescences 
rather contracted, few-flowered; pedicels at first nodding, 0.5-1 cm. 
long, glandular-pilose like the lanceolate, obtusish sepals, these 5-6 
mm. long; petals 7-8 mm. long; capsule 10-12 mm. long; seeds 
tuberculate. Apparently, with C. Trianae Briq., well distinct from 
C. Kunthii Briq. (C. glutinosum HBK. Nov. Gen. & Sp. 6: 29. 1823) 
of Colombia, which is very glutinous, with erect pedicels 15-25 mm. 
long and flat leaves (fide Danguy). Neg. 29835. 

Huanuco: Tambo de Vaca, 4,000 meters, wet, mossy uplands, 
4360. Ecuador; Colombia. "Puscala." 

Cerastium floccosum Benth. PL Hartw. 162. 1839; 300. 

In general similar to C. mollissimum, but the pubescence tomen- 
tose-floccose, composed of branched and simple hairs; calyx early 
campanulate, 10-12 mm. long, the petals and capsule about as long. 
C. candicans Wedd. has smaller leaves becoming glabrous above, 
the calyx lobes only 5 mm. long. Neg. 27710. 

Peru: Without locality, Ruiz & Pavon (det. Mattfeld). Junin: 
Mount Juntai, 4,700 meters, Killip & Smith 22059. Hacienda 
Runatullu, 4,400 meters, Weberbauer 6625. Ecuador; Colombia. 



586 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Cerastium humifusum Camb. in St. Hil. Fl. Bras. 2: 120. 1829. 

A nearly glabrous annual with spreading, creeping stems, branch- 
ing, to 20 cm. long or longer; leaves oblong, narrowed to the base, 
obtusish, 1-3 cm. long, exceeded by the filiform, glandular-puberu- 
lent pedicels; sepals ovate-oblong, obtusish, 4 mm. long, very sparsely 
if at all pubescent, twice exceeded by the oblong, bifid petals and the 
ovate capsule, the teeth of the latter recurved-revolute; seeds obtusely 
and coarsely granulate. Widely distributed in South America, 
according to Pax and Hoffmann, so probably frequent in Peru. 
Illustrated, Mart. Fl. Bras. 14, pt. 2: pi. 64. 

Lima: At 3,000 meters, above Lima (Weberbauer, 169). South 
America. 

Cerastium imbricatum HBK. Nov. Gen. & Sp. 6: 28. 1823. 

Much branched, the creeping stems covered below with the bases 
of dead leaves; leaves sessile, more or less densely imbricate, spatu- 
late-oblong, soft-villous, 8-15 mm. long, 2-5 mm. broad; flowers 
terminal, the puberulent pedicels 2 mm. long, or longer in fruit; 
calyx short-pilose, campanulate, the sepals oblong, obtusish, the 
interior scarious-margined, 2-4 mm. long, twice exceeded by the 
oblong, cuneate-based petals; capsule curved, equaling or slightly 
longer than the calyx; seeds brown, puncticulate-tuberculate. 

Cajamarca: Puccha Valley, 4,400 meters (Weberbauer, 224). 
Ancash: Huaraz (Weberbauer, 225). Bolivia; Ecuador; Colombia. 

Cerastium mollissimum Poir. Diet. Suppl. 2: 164. 1811; 298. 
C. Willdenvwii HBK. Nov. Gen. & Sp. 6: 29. 1823. 

Perennial, the usually suberect branches to 50 cm. tall, these 
with the leaves and, to greater or less extent, the calyx densely 
white-lanate with stellately branched trichomes; leaves lanceolate, 
long-acuminate, 3-5 cm. long, 5-10 mm. broad; flowers few to many, 
often laxly umbellate, the filiform pedicels 3 cm. long or longer; 
sepals lanceolate, acute, green, narrowly margined, 7-10 mm. long; 
petals to even twice as long, shallowly bilobate, obovate-cuneate; 
capsule teeth straight, the capsule long-exserted or in var. diffusum 
Fenzl less so; seeds brown, acutely tuberculate. C. andinum Benth. 
PI. Hartw. 162. 1839, is a "lusus," fide Rohrbach, with calyx lobes 
white-lanate; C. Willdenowii is the less pubescent, typical form; both 
are maintained by Pax and Hoffmann as species in Pflanzenfamilien. 

Peru: (Jussieu, type). Without locality, Ruiz & Pavon (det. 
Mattfeld). Huanuco: Huallaga, 3,800 meters, Weberbauer 6726. 



FLORA OF PERU 587 

Tambo de Vaca, 4,500 meters, 4426. Southern South America 
and in the Andes north to Colombia. 

Cerastium mucronatum Wedd. Ann. Sci. Nat. V. 1: 294. 
1864; 301. 

A softly pubescent and more or less glutinous perennial, 5-15 
cm. tall, with a number of densely leafy, fascicled and creeping 
sterile stems; leaves linear or spatulate-lanceolate, narrowed at the 
base, mucronulate, densely hirsute-scabrous, 3-5 cm. long, 3-8 mm. 
wide; flowers few, subtended by leafy, acuminate, mucronate bracts; 
pedicels and truncate-campanulate calyx subequal; sepals densely 
glandular, 7-8 mm. long, shorter than the ovate-oblong, emarginate 
petals; capsule probably little exserted. Allied by Rohrbach to 
C. floccosum Benth., but certainly very different in pubescence. 
Neg. 27712. 

Puno(?): (Weddell). Lima: Viso, wet, rocky crevices, 617. 
Cuzco(?) : Marcapata, 3,300 meters, in shrubwood, Weberbauer 7794. 
Bolivia. 

Cerastium nanum Muschl. Bot. Jahrb. 45: 447. 1911. 

Stems crowded, simple, 1-2 cm. long, glabrate; leaves at first 
minutely or obscurely ciliate, narrowly ovate-oblong, acutish, 
scarcely 1 mm. long, to 0.5 mm. broad; flowers apparently solitary; 
pedicels concealed by the leaves; calyx lobes oblong-acutish, green, 
subscarious-margined, glabrous, 1.5-2 mm. long; petals slightly 
longer; capsule twice as long, nearly straight(?); seeds small, 
obtusely granulate (according to Muschler). Neg. 29721. 

Cajamarca: Ocros, 4,400 meters, Weberbauer 2781 (type). 
Moquehua: Carumas, 4,400 meters, Weberbauer 7323. 

Cerastium nutans Raf. Pre"c. Decouv. 36. 1814; 288. C. 
longipedunculatum Muhl., a nomen nudum (fide Gray, Man.) but 
in general use. 

Stems ascending, erect, more or less branching, to 40 cm. high, 
viscid-pubescent; leaves lanceolate, acutish, 2-4 cm. long, 3-6 mm. 
broad; flowers many, often subumbellate; pedicels somewhat glan- 
dular-puberulent, straight except at the tip, or deflexed in fruit, 
usually 3-5 times longer than the calyx; bracts herbaceous; sepals 
lanceolate, obtusish, scarious-margined, puberulent, 4-5 mm. long, 
the petals usually half again as long; capsule curved, to 3 times 
longer than the calyx; seeds lightly and obtusely granulate. 
See also C. subspicatum Wedd. 



588 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Peru (?). Widely distributed in North and South America 
(Pax & Hoffmann). 

Cerastium peruvianum Muschl. Bot. Jahrb. 45: 445. 1911. 

Caespitosum depressum ramosissimum repens undique molliter 
villosum; ramis brevibus confertis densissime foliosis basibus foliis 
marcidis obtectis ad 3-4 cm. longis, caulibus floriferis 3-4 cm. altis 
plerumque 3-floris; foliis dense imbricatis oblongo-ellipticis obtusis 
circa 7 mm. longis 4 mm. latis superioribus paullo reductis sub- 
ovatis subacutis; pedicellis 3-5 mm. longis plerumque suberectis; 
sepalis extus villoso-glandulosis et nigro-punctatis et lineatis intus 
glabris fere oblongis obtusis 6 mm. longis quam petala emarginata 
paullo brevioribus; capsulis cylindrico-oblongis 8 mm. longis leviter 
curvatis dentibus erectis margine vix vel haud revolutis; seminibus 
brunneis acute denseque tuberculatis. A description is offered for 
the Cuzco plant, the type of Muschler not having been seen and 
anyway probably poorly described. 

Cuzco: Stony places west of Marcapata, 4,600 meters, Weber- 
bauer 7775. Without locality, Weberbauer 7078. Junin: La Oroya 
to Tarma, 4,300 meters (Weberbauer 2598, part of type). Ancash: 
Ocros, 4,400 meters (Weberbauer 2798, part of type). Ecuador. 

Cerastium soratense Rohrb. Linnaea 37: 291. 1872. 

Stems ligneous below, ascending-prostrate, cespitose, little 
branching, like the pedicels and calyx puberulent, 5-8 cm. long; 
leaves linear or spatulate-lanceolate, obtusish, both sides and margins 
hirsutulous, 15-20 mm. long, 2-4 mm. wide; inflorescence terminal, 
few-flowered, the subrecurving pedicels and calyx subequal; sepals 
oblong-lanceolate, acuminate or obtusish, green, scarcely scarious- 
margined, 5-8 mm. long; petals white, a third longer than the 
calyx, ovate-oblong, lobed one-third their length, with 2 ovate, 
obtuse lobes, barely clawed; capsule slightly curved, not at all or 
barely exserted; seeds brown, minutely and acutely tuberculate. 
C. imbricatum HBK. has softer, longer pubescence and subequal 
sepals and petals. Neg. 27713. 

Puno: Lake Titicaca (Meyeri). Bolivia. 

Cerastium subspicatum Wedd. Ann. Sci. Nat. V. 1: 295. 
1864; 287. 

Biennial or scarcely more enduring, soft-hirsutulous and more 
or less glandular or viscid throughout; leaves linear-oblong or lance- 
olate; inflorescences soon loose and elongate; bracts herbaceous; 



FLORA OF PERU 589 

fruiting pedicels equaling or longer than the calyx and more or less 
divaricate or refracted; sepals acute, hirsutulous, exceeded by the 
curved capsule and by the petals, or these lacking, then the stamens 
mostly only 5 (var. apetalum Wedd.). Plants with the habit of 
C. vulgatum, to which Rohrbach reduced it, but the bracts not at 
all scarious-margined. Except that it is more enduring and more 
cespitose or tufted at the base, it seems scarcely distinguishable from 
C. nutans. C. obscurum Triana & Planch, belongs here. Neg. 29836. 
Junin: Morococha, Isern 221+7. Huaron, bunchgrass slopes, 
4,000 meters, 1134- Cerro de Pasco, grassy slopes, 4,000 meters, 
3080. Mount La Juntai, Killip & Smith 22088. Without locality: 
(Weddell, type). Huanuco: Near Mito, bunchgrass slope, 1885. 
Puno: Araranca, rocky knoll, 4,100 meters, Pennell 13461. Lima: 
Rocky canyon wall east of Canta, Pennell 14644- Bolivia; Colombia. 

Cerastium Trianae Briq. Ann. Cons. Jard. Geneve 13-14: 
282. 1911. C. caespitosum Triana & Planch. Ann. Sci. Nat. V. 17: 
152. 1862, non Gilib., 1781. 

Stems many, decumbent-ascending, densely leafy, 10-15 cm. 
long; leaves lance-linear, gradually narrowed to the acute or obtusish 
apex, plane, 2.5 cm. long, 4-6 mm. wide, loosely and sparsely glan- 
dular-pilose; flowers few; pedicels thick, densely pubescent, after 
anthesis elongating and refracted; sepals ovate-lanceolate, obtuse, 
densely glandular-pilose, 6 mm. long; petals half again as long as 
the sepals, as also the capsule; seeds minutely punctulate. 

Huanuco: Chasqui, wet, grassy slope, 3291; like the type except 
that the capsule is more curved. Ecuador; Colombia. 

Cerastium trichocalyx Muschl. Bot. Jahrb. 45: 445. 1911. 

Apparently very similar, if not referable, to C. soratense or C. 
imbricatum; stems to about 20 cm. long; leaves finally subglabrous, 
1-3 cm. long, 5-10 mm. broad; bracts herbaceous; sepals oblong, 
acutish, scarious-margined, 7 mm. long; petals slightly longer than 
the calyx, obcordate, bifid, clawed; capsule shorter than the calyx. 

Ancash: Near Huaraz, 4,400-4,600 meters (Weberbauer 2983, 
type). Lima: Viso, 3,000 meters, rock crevices and shallow soil, 605; 
perhaps new, the leaves linear-oblong, glabrate. 

Cerastium tucumanense Pax, Bot. Jahrb. 18: 25. 1893. 

Perennial, with many erect or ascending, nearly simple, more or 
less densely villous, eglandular stems, to 25 cm. high, with elongate 
internodes; leaves lanceolate or linear-lanceolate, acute or the apex 



590 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

obtusish, glabrous except for the densely ciliate margin, 3-4 cm. 
long, 4 mm. broad; flowers many, pseudo-umbellate; pedicels shorter 
than the sepals or sometimes as long, recurved; bracts herbaceous; 
calyx campanulate, the sepals oblong-lanceolate, acute, apically 
scarious or the inner scarious-margined, all villous, 7-8 mm. long; 
petals 1 cm. long, 4 mm. wide; calyx bilobate, obovate-spatulate; 
stamens 5; capsule cylindric, 1 cm. long; seeds brown, acutely tuber- 
culate. Differs from C. mucronatum and C. soratense in the merely 
ciliate leaves, scarious sepals, 5 stamens, and cylindric capsule 
(Pax). 

Puno: Titicaca region, 4,000 meters (Weberbauer, 186). Tacna: 
Candarave, 4,000 meters, tola heath, Weberbauer 7374; perhaps new, 
the flowers smaller and the seeds bluntly tuberculate. Argentina. 

Cerastium viscosum L. Sp. PI. 437. 1753; 283. C. glomeratum 
Thuill. Fl. Paris 225. 1799. 

Pubescent and viscid-hairy like C. vulgatum, but annual and 
lacking the sterile leafy shoots, nearly erect, often more than 10 cm. 
tall; leaves ovate or more elongate and more or less obovate; bracts 
herbaceous; flowers in close clusters, the pedicels, even in fruit, 
shorter than the sharply pointed sepals, these exceeding the petals. 
Var. consanguineum (Wedd.) Rohrb. (C. consanguineum Wedd. Ann. 
Sci. Nat. V. 1: 296. 1864) is a short-pubescent form with narrowly 
linear-lanceolate leaves; calyx lobes apically glabrous; petals want- 
ing. It is retained as a species by Pax and Hoffmann in Pflanzen- 
familien. Neg. 27708. 

Junin: La Quinua, 2031. Morococha, 891. Tarma, bunchgrass 
slopes, 1056 (?). Huanuco: Mito, 1738. Lima: San Lorenzo Island, 
500 meters, Weberbauer 5925. Chancai, Ruiz & Pavon (det. Matt- 
feld). Rio Blanco, 692. Lurin, 5948. Without locality: (Dombey). 
Rio Moho, central Peru (Weddell, type of the var.). Puno: Chu- 
quibambilla, 3,900 meters, rocky puna slope, Pennell 13415. 
Cuzco: Huancaro, Vargas 3166. Nearly cosmopolitan. 

Cerastium vulgatum L. Fl. Suec. ed. 2. 158. 1755; 286. C. 
caespitosum Gilib. Fl. Lith. 159. 1781. C. triviale Link, Enum. 
Hort. Berol. 1:433. 1821. 

Flowering stems rather slender, 10-65 cm. high, simple or slightly 
branching, hirsute or rarely glandular; internodes elongate, the 
median becoming 2-12 cm. long; leaves of the season 3-7 pairs, 
oblong to narrowly oval, hirsute on both surfaces, the median 0.5-4 
cm. long, 1.5-15 mm. broad; bracts similar but smaller, broadly 



FLORA OF PERU 591 

scarious at the margin and apex; inflorescence 3-60-flowered, forming 
a terminal, finally very dichotomous cyme, at first rather congested, 
in fruit with the lower pedicels divergent or reflexed and 2-4 times as 
long as the calyx; sepals 4-7 mm. long, ovate-lanceolate, acute, 
hirsute, the margins scarious; petals 4-8 mm. long, equaling or very 
slightly exceeding the sepals, cleft to the middle, with a ciliate claw; 
capsule 7-11 mm. long; seeds 0.5-0.7 mm. in diameter, reddish, 
tuberculate. Description by Fernald & Wiegand, Rhodora 22: 177. 
1920. See Briq. Prodr. Fl. Corse 1: 505-506. 1910, also Hauman & 
Irigoyen, Anal. Mus. Hist. Nac. Buenos Aires 32: 172. 1923-25, for 
arguments for the elimination of this name and also that of C. vis- 
cosum L., which does not seem to be justified, the elimination itself 
being bound to result in confusion; cf. Fernald & Wiegand, op. cit. 
Var. hirsutum Fries is glandless; var. hirsutum f. glandulosum (Boenn.) 
Druce is glandular. 

Ancash: Samanco, 3,000-3,500 meters (Weberbauer, 170). 
Hualgayoc, 4,000 meters (Weberbauer, 271). Junin: Yauli, 4,400 
meters, between Calamagrostis tufts (Weberbauer, 222). Almost 
cosmopolitan. 

Cerastium vulgatum L. var. peruvianum Gray, Bot. U. S. 
Expl. Exped. 120. 1854. 

Later flowers apetalous; stems above and pedicels glandular- 
pubescent; leaves sessile, linear, 12-20 mm. long, 2 mm. wide; petals, 
if present, longer than the glandular calyx. Another form is var. 
andinum Gray, loc. cit., scarcely glandular, condensed, the leaves 
oblong-linear, the petals a little shorter than the calyx. With the 
above broad interpretation of C. vulgatum, these varieties appear to 
have no significance. See also C. subspicatum. 

Lima: Banos (Wilkes Exped., type); also Obrajillo, Alpamarca 
(var. andinum). Cuzco: Paucartambo, 3,400 meters ( Herrera 2318). 
Ollantaitambo, 2,800 meters, Herrera 3401. Puno: Chuquibambilla, 
rocky, limestone knoll, 3,900 meters, Pennell 13385. Near Puno, 
4,000 meters, Soukup 71. Ecuador. 

4. STELLARIA L. 

Generally lax perennials with narrow, often green leaves and 
2-parted or 2-lobed petals, or these rarely reduced or lacking. 
Leaves broader than linear. 

Petals shorter than the calyx or none; pubescence restricted to a 
line on stems and petioles S. media. 



592 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Petals longer than the calyx or lacking; pubescence not charac- 
teristically in lines. 

Leaves all shorter than 1 cm S. serpyllifolia. 

Leaves all or nearly all distinctly longer. 

Sepals acute; petioles all short S. ovata. 

Sepals obtuse; lower petioles as long as or longer than the 

leaves S. cuspidata. 

Leaves linear or awl-shaped. 

Leaves linear-oblong, narrowed at each end S. xanthospora. 

Leaves narrowly ovate at the base, awl-shaped . S. aphananthoidea. 

S tell aria aphananthoidea Muschl. Bot. Jahrb. 45: 444. 1911. 

Branches crowded, cespitose, to 30 cm. long, sometimes slightly 
branched, at first minutely puberulent, as also the leaves, these 
linear, more or less spreading, 1-3 cm. long, 1-2 mm. broad, acute; 
flowers solitary, the mostly erect pedicels 2-3 cm. long, sometimes 
glandular-puberulent; sepals densely glandular-pubescent, ovate- 
lanceolate, 1 cm. long, obtusish, 3-nerved, green; petals scarcely 
as long, ovate. Neg. 29837. 

Lima: Matucana, 2,500 meters, Weberbauer 133 (type). 

Stellaria cuspidata Willd. ex Schlecht. Ges. Naturf. Berl. 
Mag. 7: 196. 1816; HBK. Nov. Gen. & Sp. 6: 27. 1823. S. ciliata 
Vahl ex Pers. Syn. 1: 503. 1805, not Gilib., 1781. S. prostrata Bald- 
win, as to Weberbauer, 170 (probably). S. leptopetala Benth. PI. 
Hartw. 163. 1839 (S. leptosepala, Weberbauer, 183). 

Prostrate-ascending, sometimes rooting at the nodes, the weak 
branches usually more or less pubescent or puberulent; lower petioles 
often 10-15 mm. long, the upper becoming obsolete; leaf blades 
ovate-oblong, somewhat acuminate, more or less cordate at the 
base, typically glabrous, thin and to 4 cm. long or sometimes more 
or less coriaceous and as short as 1 cm.; peduncles elongate; sepals 
oblong-lanceolate, obtuse, ciliate toward the base, villous or less 
frequently glabrous, about 4 mm. long, the petals slightly longer 
to twice as long. Apparently very variable and scarcely, as re- 
marked by Gray, distinct from S. nemorum L. of Europe, with 
longer-petioled leaves, to which the following Weberbauer specimens 
with viscid pedicels and calyces have been referred in herb. Berlin. 
S. micrantha Spruce ex Rohrb. op. cit. 280 has broadly ovate, cordate- 
based leaves to 4 cm. long and half as broad, and acuminate sepals 
to 3 mm. long, half as long as the nearly 2-parted petals. 



FLORA OF PERU 593 

Lima: Matucana, 110. Obrajillo (Wilkes Exped., det. Gray). 
Montana (Jussieu, type of S. ciliata Vahl, a small, glabrous var.). 
Mountains of Mongomarca, 300-400 meters, Weberbauer 5686. Rio 
Blanco, 3,000-3,500 meters, open hillside, Kittip & Smith 21571. 
Canta, 3,100 meters, moist bank, Pennell 14606. Atocongo, 250- 
500 meters, open, calcareous hills, Pennell 14775. Cuzco: Paso de 
Tres Cruces, 3,800 meters, rocky banks and knolls on paramo, 
Pennell 13830. San Martin: Zepelacio, 1,200-1,600 meters, in forest, 
Klug 3530. Puno: Cuyocuyo, Sandia, 3,800 meters, Weberbauer 
925 (det. Muschler). Loreto(?): San Antonio, Rio Cumbaso, Ule 
6700 (det. Muschler). Ancash: Lomas, 300 meters, Weberbauer 
5686. Cajamarca: Ocros, Weberbauer 2677. Ayacucho: Osno, 
3,300 meters, Weberbauer 5569. Huanuco: Cani, 3393. Chile to 
Mexico. 

Stellaria media (L.) Vill. Hist. PI. Dauph. 3: 615. 1789. Alsine 
media L. Sp. PI. 272. 1753. 

A bright green, small annual, the weak, diffuse or suberect 
stems pubescent in a line, as the petioles; leaves ovate, the upper 
sessile; petals shorter than the calyx and bifid, or none; stamens 
2-10; styles 3-4; capsule narrowly ovoid, subcylindric; seeds more 
or less tuberculate. Variable, especially in floral parts. For an 
interesting account of variation as illustrated in this plant see 
Matzke, Amer. Journ. Bot. 19: 477. 1932. Although Burnat as 
early as 1892 pointed out in Fl. Alp. Mar. 1: 257, that Cirillo, 
Essent. PI. Char. Comm. 36. 1784, merely remarked that Alsine 
media L. is a species of Stellaria and failed to make the transfer, 
many authors have continued to attribute the combinations to him. 
Briquet, Fl v Cors. 1: 498. 1910, again called attention to the common 
error and, with his usual precision, corrected Burnat's spelling of 
the name of the Naples professor of botany from "Cyrillo," an 
error common also in America. Ball, Proc. Journ. Linn. Soc. 22: 31, 
records it as frequent about Chicla and thinks it is probably native. 

Lima: Matucana, 2,370 meters (Weberbauer 146). Chicla (Ball). 
Lurin, rock slide, 5961. Huanuco: Mito, in clearing, 1713; sunny 
stream flat, 1909. Cosmopolitan in cool regions. 

Stellaria ovata Willd. ex Schlecht. Ges. Naturf. Berl. Mag. 
7: 196. 1816; HBK. Nov. Gen. & Sp. 6: 26. 1823. 

Similar to S. cuspidata; leaves roundish or subrhombic-ovate, 
minutely apiculate, scarcely acute, about 1.5 cm. long and 1 cm. 
wide, glabrous but ciliate, like the (3 mm. long) petioles; pedicels 



594 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

2.5 cm. long, divergent or reflexed in fruit; sepals elliptic-oblong, 
acute, pilose at the base, the petals nearly twice as long; seeds 
lenticular, laterally rugulose, tuberculate, brown. 

Lima: Rio Blanco, 694- Matucana, 477. Obrajillo (Wilkes 
Exped., det. Gray). San Martin: Tarapoto, Spruce 4454- San 
Roque, Williams 7274, 7380. Near Moyobamba, King 3530 (det. 
as S. nemorum; see remark under S. cuspidata). To Mexico and 
Venezuela. 

Stellaria serpyllifolia Willd. ex Schlecht. Ges. Naturf. Berl. 
Mag. 7: 196. 1816; HBK. Nov. Gen. & Sp. 6: 26. 1823. 

Much branched, procumbent; leaves all short-petioled, ovate, 
acute, submucronate, rotund at the base, densely ciliate on the 
margins and midnerve beneath, the largest cauline leaves 8 mm. 
long and 6 mm. wide; sepals pubescent, oblong, obtuse; stamens 10. 

Peru: (Probably). Ecuador. 

Stellaria xanthospora Chod. & Wilcz. Bull. Herb. Boiss. II. 
2: 293. 1902. 

Stems weak, to 15 cm. long; leaves linear-oblong, acute at each 
end, glabrous, to 15 mm. long, 2.5 mm. wide; flowers axillary, on 
pedicels longer than the leaves, recurving below the tip; capsule 
slightly more than half as long as the calyx; seeds brown, inflated, 
lenticular. "Affinis S. lanuginosae Rohrb. Monogr. in Linnaea 38: 
260," according to Chodat and Wilczek. That plant, however, 
is Arenaria lanuginosa (Michx.) Rohrb. Linnaea 37: 259 (not "38: 
260"), or S. lanuginosa (Michx.) Torr. & Gray, if preferred. Ac- 
cording to Pax and Hoffmann, Pflanzenfamilien, the species of Chodat 
and Wilczek is a valid Stellaria, known from Chile and Colombia; 
the type locality, however, is Argentina! The species has not been 
seen by me. 

Peru: (Probably, according to the range given by Pax and 
Hoffmann). Argentina; Chile; Colombia. 

5. ARENARIA L. 

Minuartia L. Sp. PI. 89: 1753; Alsine Gaertn. Fruct. 2: 223. 1791. 

Reference: F. N. Williams, Journ. Linn. Soc. 33: 326-437. 1898. 

Plants often cespitose, usually perennial, with most often sub- 
ulate, rigid leaves. Petals typically entire, sometimes wanting. 
Stamens 10, rarely 5. Disk more or less developed, often prominent. 
Styles normally 2 or 3. Following Fernald, Minuartia is here 
included. 



FLORA OF PERU 595 

Sepals more or less inflexed and connivent; styles usually 2; cushion 

plants suggesting Pycnophyllum. 
Petals longer than the calyx, or at least about as long. 

Petals spatulate; plants densely matted A. dicranoides. 

Petals oblong; plants open, the stems rooting at the nodes. 

A. aphanantha. 

Petals distinctly shorter than the calyx or lacking. 
Stamens 10; plants often robust or forming ample cushions. 

Leaves deeply bisulcate dorsally A. bisulca. 

Leaves not sulcate or not markedly so. 
Sepals broadly membranous-margined. 

Sepals ovate, obtuse A. pycnophylla. 

Sepals oblong, acuminate A. pycnophylloides. 

Sepals narrowly scarious, ovate-oblong, acutish. 

A. boliviano,. 
Stamens 5; diminutive plants 1-5 cm. high. 

Leaves cuspidate; capsule included A. alpamarcae. 

Leaves acute; capsule exserted A. pedunculosa. 

Sepals not inflexed; styles usually 3-4; plants sometimes densely 
matted but not infrequently lax and open in habit. 

Staminal disk conspicuous, lobed; plants glabrate, diffuse; leaves 
flat; seeds large, lustrous, obsoletely puncticulate. 

A. tetragyna. 
Staminal disk ring-like, often obscure; plants frequently tufted 

or pubescent; seeds smooth or tuberculate. 
Fertile stamens 5; seeds roughened (except A. Orbignyana). 
Roots not at all fusiform or tuber-like. 

Sepals indurate at the base A. nitida. 

Sepals not indurate. 

Petals none; seeds granulate A. andina. 

Petals present; seeds smooth A. Orbignyana. 

Roots fusiform, tuber-like A. crassipes. 

Fertile stamens 10; seeds smooth, lustrous; sepals not indurate. 
Plants more or less densely pulvinate-cespitose or, if diffuse, 

low and with creeping-assurgent stems or branches. 
Flowers 3-5 together, involucrate by the uppermost leaves. 

A. Mandoniana. 



596 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Flowers solitary but often only in the upper axils. 

Leaves awl-like, more or less pungent and rigid; flowers 

subsessile in the upper axils. 
Petals present. 

Petals and sepals subequal; leaves pungent-aristate. 

A. aphanantha. 
Petals much shorter than the sepals, leaves acute. 

A. Standleyi. 
Petals none A. Jamesoniana. 

Leaves not awl-like, soft; flowers often scattered and 
conspicuously pediceled. 

Calyx lobes acute A. parvifolia. 

Calyx lobes obtuse A. digyna. 

Plants typically lax and open, with strongly ascending or 

erect stems and branches. 
Flowers few to several, borne terminally in a loose or 

cymose inflorescence. 

Sepals oblong, the petals slightly longer . .A. Poeppigiana. 
Sepals ovate, distinctly exceeded by the petals. 

A. Stuebelii. 
Flowers solitary, axillary. 

Sepals 4.5-5.5 mm. long A. soratensis. 

Sepals 2.5-4 mm. long. 

Sepals much exceeded by the petals. . . . A. megalantha. 

Sepals and petals subequal A. lanuginosa. 

Arenaria alpamarcae Gray, Bot. U. S. Expl. Exped. 116. 1854. 

Glabrous and cespitose, forming pulvinate tufts a few cm. high; 
leaves crowded or imbricate but somewhat spreading, ovate-lanceo- 
late, 4-6 mm. long, rigid, carinate-cuspidate, the margins thickened 
and smooth or very obscurely ciliate near the connate base; peduncle 
2-4 mm. or in fruit 6-10 mm. long, 1-flowered; sepals usually 5, 
resembling the leaves, 3-4 mm. long; petals apparently none; capsule 
rather shorter than the calyx, usually 6-valved to the base; ovules 
about 20, the seeds 5 or 6, globular, smooth. Mattfeld (in herb.) 
reduces this to A. nitida. 

Junin: Alpamarca (Wilkes Exped., type). Also above Banos and 
Obrajillo (Dept. Lima). 



FLORA OF PERU 597 

Arenaria andina Rohrb. Linnaea 37: 255. 1872; 372. 

Similar to A. Orbignyana; leaves slightly aristate, thin at the base, 
glabrous, 5 mm. long, 1-1.5 mm. broad; pedicels glabrous, 1.5 mm. 
long; calyx lobes ovate-oblong or oblong, obtusish, yellow, 2.5-3 mm. 
long; capsule nearly exserted; seeds elegantly striate-granulate. 
Neg. 27727. 

Peru: (Probably). Bolivia. 

Arenaria aphanantha Wedd. Ann. Sci. Nat. V. 1: 293. 1864. 

Cespitose, rather openly branched, the diffuse stems often rooting 
at the nodes; leaves laxly imbricate, subcoriaceous, narrowly lanceo- 
late, aristate, glabrous or toward the base ciliate, 3-4.5 mm. long; 
flowers solitary, subsessile among the upper leaves; calyx glabrous, 
the lanceolate, very acute lobes 3 mm. long equaling or slightly 
shorter than the oblong, obtuse, entire petals; stamens 10; capsule 
ovate-oblong; seeds minute, acutely grooved dorsally. Omitted by 
Williams; by Rohrbach placed between A. dicranoides and A. bolivi- 
ano,, with the remark, "in habit and character strongly accedes to 
Stellaria laevis (Bartl.) Rohrb. but is distinguishable in the narrower, 
less imbricate leaves, very acute sepals, and form of the petals." 

Andes of Peru: (Gay 1776, type). 

Arenaria bisulca (Bartl.) Rohrb. Linnaea 37: 248. 1872. Cher- 
leria bisulca Bartl. in Presl, Rel. Haenk. 2: 12. 1831. 

Diffuse, much branched, finely puberulent, the stems to 10 cm. 
long, the ascending flowering portion sparsely pubescent; leaves 
lanceolate, acutely mucronate, lustrous, marked by 2 deep, narrow 
grooves, the thick margin shortly setulose-ciliate, nearly amplexicaul; 
calyx coriaceous, lustrous, the ovate-acuminate, lanceolate sepals 
glabrous; petals linear-oblong, emarginate, twice as long as the calyx: 
styles 2-3; capsule included, with 6-10 seeds. 

Huanuco: Montana (Haenke). Argentina(?). 

Arenaria boliviana Williams, Proc. Journ. Linn. Soc. 33: 425. 
1898. A. conferta Wedd. Ann. Sci. Nat. V. 1: 293. 1864, not Boiss. 

Stems often densely pulvinate, cespitose, much branched, the 
short branches glabrous, clothed below with old leaves; leaves minute, 
oblong-lanceolate, subacute, narrowly scarious-margined, ciliate, 
sometimes hispid throughout; flowers solitary, terminal, sometimes 
few in the upper axils; pedicels hispid, twice longer than the calyx, 
6 mm. long; sepals ovate-oblong, obtusish, green or scarcely scarious- 



598 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

margined; petals none; capsule sometimes slightly exserted; seeds 
brown, lustrous, smooth. Neg. 27726. 

Junin: Cerro de Pasco, 4,400 meters, Weddell. Bolivia. 

Arenaria caespitosa Muschl. Bot. Jahrb. 45: 449. 1911. 

Diffuse, with we&k, procumbent, branching stems 3-6 cm. long, 
these more or less angled and pubescent above; leaves linear- or 
rarely ovate-lanceolate, thin or fleshy, mostly much longer than the 
internodes, acuminate or submucronulate, narrowed to an almost 
petiolar base, 5-10 mm. long, 2-4 mm. wide, somewhat puberulent 
at the base and on the nerves; flowers axillary on filiform, spreading, 
densely puberulent or rarely glabrous pedicels 1.5-2 cm. long, in 
fruit curving to the apex; sepals ovate-oblong, green-carinate, 
acuminate, puberulent-scabrous, sometimes ciliate, scarious-mar- 
gined, 3-4 mm. long, slightly exceeded by the oblong-oval, obtuse 
petals, or these wanting; anthers pale yellow; styles 3; capsule 
usually a little longer than the calyx, 6-valved, 8-seeded, the seeds 
smooth, lustrous. 

Cajamarca: Ocros (Weberbauer 2796, type). 

Pycnophyllum carinatum Muschl. Bot. Jahrb. 45: 457. 1911. 

Densely cespitose, densely leafy, the stems 4-5 cm. long, with 
slender branchlets, glabrescent; leaves ovate to lanceolate, very 
acute, thickly keeled, glabrous and lustrous but minutely ciliate- 
margined, deeply concave; pedicels slender, glabrate or scaly- 
pulverulent, 3-4 mm. long; sepals broadly ovate or roundish, scarious, 
entire, nerved; petals apparently lacking; fertile stamens to 3 mm. 
long, slightly exceeded by the elongate style; stigmas 3. A species 
of Arenaria, section Dicranella, fide Mattfeld. Cf. A. boliviana 
Williams. 

Arenaria crassipes Baehni & Macbr., sp. nov. 

Radix crassa fusiformis 4 cm. longa ad apicem 4-6 mm. crassa; 
caulibus 7-12 prostratis simplicibus 3-5 cm. longis glabris vel 
glabratis laxe foliatis, internodiis 3-4 mm. longis; foliis suboblongis 
breviter acuminato-apiculatis 5-7 mm. longis 2-2.5 mm. latis sub- 
coriaceis vel demum membranaceis laxis haud sulcatis marginem 
versus pilosis vel toto margine piloso-ciliatis, demum glabratis; 
floribus terminalibus circa 5-7 in apice ramorum fere umbellato- 
congestis, pedicellis circa 4 mm. longis leviter curvatis; sepalis 
anguste lanceolato-acuminatis circa 3 mm. longis glabris haud vel 
angustissime marginatis planis ad basin obscure induratis; stamini- 
bus 5; petalis nullis; seminibus puncticulatis. With much the 



FLORA OF PERU 599 

habit of A. nitida but remarkable in its tuber-like root that resembles 
in shape a small carrot. It differs also in the complete absence 
of petals. 

Lima: Prostrate on upland slopes, 3,000 meters, Rio Blanco, 
Macbride 812, type in Herb. Field Museum. 

Arenaria dicranoides HBK. Nov. Gen. & Sp. 6: 34. 1823. 
Pycnophyllum dicranoides Muschl. Bot. Jahrb. 45: 454. 1911. 

Cespitose, the crowded, densely leafy stems a few cm. high, 
glabrous except the ciliate, ovate-lanceolate, acute or obtusish 
leaves, these imbricate in 4 rows, 3-4 mm. long; flowers sessile at 
the branchlet tips; sepals oblong, acute, subequal, exceeded by the 
spatulate petals; styles 2; ovules 4. The leaves of the older stems 
are mostly triangular-lanceolate and more or less acute, those of the 
younger branches rather ovate and obtuse. The specimen from 
Alpamarca has smaller and blunter leaves and might be referred 
to A. bryoides Willd., but I can not draw any line between the 
different forms. A cushion-like or tufted plant, it simulates Distichia 
except for the shorter, awl-like leaves. 

Puno: (Meyen; det. Walpers as A. bryoides). Junin: Near Yauli, 
4,500 meters Weberbauer 5120, 221 (det. Muschler). Cerro de Pasco 
(Mathews 688). Near Casa Cancha, Culnai, and Alpamarca (Wilkes 
Exped.). Ecuador; Bolivia. 

Arenaria digyna Schlecht. Ges. Naturf. Berl. Mag. 7: 201. 
1813. A. scopulorum HBK. Nov. Gen. & Sp. 6:31. 1823. A. serpens 
HBK. op. cit. 32. A. serpylloides Naud. in Gay, Fl. Chile 1: 271. 
1845. 

Procumbent to suberect, diffuse, much branched, the branchlets 
glabrous or rarely hirsutulous; leaves 1-nerved, thin or somewhat 
fleshy, glabrous or puberulent or marginally and dorsally ciliate, 
scarcely over 6 mm. long, 1 mm. wide; flowers solitary, erect, the 
glabrous or pubescent pedicels usually only 2-3 mm. long; calyx 
often as long, glabrous or rarely sparsely hirsute, the oblong-obtuse 
lobes a little shorter than the ovate-oblong, obtuse, entire petals; 
seeds 2-3, lustrous, smooth; capsule elliptic, scarcely 2 mm. long, 
slightly exserted. A. parvifolia Benth. has leaves 4-8 mm. long, 
1-3 mm. wide, petals none or shorter than the often puberulent 
calyx. The Humboldt specimen in Herb. Berlin is pubescent, the 
leaves acute. 

Cajamarca: Micuipampa, Humboldt, type of A. scopulorum. 
Lima: Casa Cancha, a little distance below snow (Wilkes Exped.; 



600 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

det. Gray as A. scopulorum}. Banos (Wilkes Exped.; det. Gray). 
Junin: Cerro de Pasco, Mathews. Puno: Near Poto, 4,600 meters 
Weberbauer (as A. serpylloides) . Lake Titicaca region, 3,919 meters, 
Mandon 963, det. Rohrb. Cuzco: Prov. Canchis, Sicuani, 3,550 
meters (Hickeri). Chile to Bolivia and Mexico. 

Arenaria Engleriana Muschl. Bot. Jahrb. 45: 449. 1911. 

A pulvinate perennial with weak, procumbent, diffuse, but 
never rooting stems 3-6 cm. long, the branches more or less angled 
and pubescent toward the tip; leaves usually ovate-lanceolate, acute 
or mucronulate, broadly truncate at the sessile base, where puncticu- 
late and puberulent-scabrous on the nerves, lightly ciliate, usually 
thick, 5-10 mm. long, 2-5 mm. broad; pedicels filiform, spreading, 
densely puberulent, in fruit curved, 1 cm. long; sepals ovate-oblong, 
green-keeled, acuminate, slightly scabrous, the broadly scarious 
margin somewhat ciliate, 3-4 mm. long; petals lanceolate, obtuse, 
often slightly longer than the calyx; anthers pale yellow; styles 3; 
capsule often slightly exserted, with 8 smooth, lustrous seeds. 

Junin: Between Tarma and Oroya (Weberbauer 251+2, type). 

Pycnophyllum horizon tale Muschl. Bot. Jahrb. 45: 454. 1911. 

Stems densely cespitose, tortuous, about 7 cm. long, glabrate, 
the many branches often rooting; leaves linear- or rarely ovate- 
lanceolate, acute, scarious, lustrous, glabrous except the ciliate 
margins, densely and quadrately imbricate but horizontally spread- 
ing; flowers apical, sessile; sepals lanceolate or ovate, scarious, acute, 
about equaled or slightly exceeded by the oblong-linear petals; 
anthers elliptic, red; style trifid apically; ovules 3-4, affixed to a 
central placenta; capsule 3-valved. This is a species of Arenaria, 
section Dicranella, according to Mattfeld. Cf. A. aphanantha Wedd. 

Lima: Andes above Lima, 4,500 meters (Weberbauer, type). 

Arenaria lanuginosa (Michx.) Rohrb. in Mart. Fl. Bras. 14, 
pt. 2: 274. pi. 63. 1872. Spergulastrum lanuginosum Michx. Fl. 
Bor. Amer. 1: 275. 1803. Stellaria lanuginosa Torr. & Gray, Fl. 
N. Amer. 1: 187. 1838. A. nemorosa HBK. Nov. Gen. & Sp. 6: 
35. 1823. A. diffusa Ell. Sketch. Bot. Car. & Georgia 1: 519. 1821. 
Stellaria laxa Muschl. Bot. Jahrb. 45: 443. 1911. 

More or less densely hirsutulous to glabrescent, the diffuse, 
weak stems sulcate or angled above; leaves linear to lanceolate, 
usually acutish, 5-30 mm. long, 2-5 mm. broad, puncticulate or 
slightly scabrous; flowers solitary, axillary on filiform, spreading, 



FLORA OF PERU 601 

densely puberulent pedicels 1.5-3 cm. long; sepals ovate-oblong, 
carinate, acuminate, the green keel scabrous, the scarious margin 
sometimes ciliate, 2.5-4 mm. long; petals typically lacking but 
usually shorter than the calyx; stamens 10; capsule slightly longer 
than the calyx, 8-seeded; seeds smooth, lustrous, spheroid-lenticular. 
Very variable in leaf form and degree of pubescence (Rohrbach). 
Used as an astringent for hemorrhages of the uterus (Herrera). 

Lima: Viso, in limestone slide rock, 766. Matucana, Weberbauer 
153; 167 (type of Stellaria laxa). Banos, Obrajillo, and Culnai 
(Wilkes Exped. ; det. Gray, using the name A. diffusa). Huanuco: 
Cochero, Poeppig 1402b. Mito, 1744, 1844- Panao, 3619. Aya- 
cucho: Prov. Huanta, 2,600 meters, Weberbauer 5603. Carrapa, 
1,200-2,500 meters, open or wooded hillsides, Killip & Smith 
22295, 22421. Amazonas: Chachapoyas, Mathews 2098. Cuzco, 
Sacsahuaman, 3,600 meters, Herrera 2354, 2177. Pillahuata, 2,500 
meters, in thickets, Pennell 13995. Machupicchu, 2,200 meters, 
Herrera 3250. Huarocondo, 3,200 meters, Herrera 3627a. Cuzco, 
Soukup 78. San Sebastian, Pennell 13614- Puno: Tabina, Lechler 
1889. Junin: Huacapistana, 1,800-2,400 meters, thickets and open 
woods, Killip & Smith 24122. Chanchamayo Valley, 1,500 meters, 
Schunke 350, 383, 486. San Martin: San Roque, 1,400 meters, 
Williams 7453. Without locality: Ruiz & Pavon (det. Mattfeld). 
Bolivia, north in the Andes to Mexico and southeastern United 
States. "Celedonia," "tauchchalli." 

Arenaria Mattfeldii Baehni, sp. nov. 

Perennis, dense caespitosa, 3-5 cm. alta; folia 1.5-2 mm. longa, 
spiraliter disposita, 4 series parallelos efformantia, imbricata, cras- 
siuscula, subtriangulata, apice obtusa, margine ciliata et paululum 
involuta; flores brevissime pedicellati, ex foliis inferioribus exserti, 
apetali; sepala 2 mm. longa, subtriangularia, crassiuscula, ad basin 
gibbosa, apice recurvata; discus conspicuus, carnosus; stamina 10 
sepala aequantia; ovarium 0.75-1 mm. longum, glabrum, seminibus 
8; styli 2 usque ad basin distincti. This little plant has the appear- 
ance of a Pycnophyllum. 

Junin: Yauli, above the Hacienda Arapa, near the Lima-Oroya 
railroad, Weberbauer 353 (type, in Herb. Deless.). 

Arenaria nitida (Bartl.) Rohrb. Linnaea 37: 249. 1872; 375. 
Cherleria nitida Bartl. in Presl, Rel. Haenk. 2: 12. 1831. 

Stems weak, procumbent at the base; leaves ovate, mucronate, 
basally connate, plane, coriaceous, glabrous, lustrous, remotely 



602 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

bisulcate beneath, 4-7 mm. long; flowers subterminal; calyx glabrous, 
coriaceous, turbinate; sepals lanceolate, acute, not sulcate; stamens 5, 
with 5 staminodia; petals half as long as the calyx, linear-oblong, 
subacute or subemarginate, attenuate to the base; capsule narrow; 
seeds nearly globose, seriate-tuberculate. 

Junin: Yauli, 4,200 meters, 902 (det. Mattfeld). Mount La Juntai, 
4,700 meters, Kittip & Smith 22107 (det. Mattfeld). Near Tarma, 
4,500 meters, Killip & Smith 21953. Lima: Obrajillo (Haenke, 
type). 

Arenaria Orbignyana Wedd. Ann. Sci. Nat. V. 1: 293. 1864; 
384. 

Diminutive, the diffuse or creeping, pulverulent or glabrous 
stems only 1-5 cm. long; leaves thickish, somewhat imbricate, 
nearly spatulate-lanceolate, acute, serrulate-ciliate especially toward 
the base, 3-4 mm. long, 1-1.5 mm. broad; flowers solitary, terminal, 
shortly pediceled; calyx lobes ovate-lanceolate, obtusish, scarious- 
margined, glabrous, 4 mm. long, scarcely equaled by the obovate- 
lanceolate, obtuse petals; capsule ovate-oblong, included, dehiscent 
by 3 biparted valves; seeds globose, reniform, lustrous. Description 
by Rohrbach, from Orbigny 1511, as represented in Herb. DC. 
Weddell describes the plant as glabrous, flowers sessile, and petals 
longer than the calyx. The calyces are notably large for the size 
of the plant. Neg. 27730. 

Cuzco: Chuquibambilla, Pennell 13392. Sacsahuaman, 3,600 
meters, Herrera 2384- Paucartambo Valley, Herrera 2325. Bolivia. 

Arenaria pallens Muschl. Bot. Jahrb. 45: 450. 1911. 

A densely pulvinate annual with diffuse, procumbent stems 2-4 
cm. long, the more or less sulcate or angled stems and branches 
glabrous; leaves usually linear-lanceolate, thick or rigid, very acute 
or mucronate, 5-8 mm. long, 2-3 mm. broad, petiolate, on the nerves 
and at the base puberulent, ciliate; pedicels filiform, spreading, 1 cm. 
long, becoming glabrate, curved in fruit; sepals oblong-ovate, green- 
carinate, puberulent, the broadly scarious margins mostly ciliate, 
3-4 mm. long, the mostly ovate petals usually a little longer; other- 
wise like A. Engleriana. 

Junin: Near Yauli, 4,400 meters (Weberbauer 267). 

Arenaria parvifolia Benth. PI. Hartw. 163. 1839; 382. 
Greatly resembling A. digyna, but the leaves often proportion- 
ately broader, ovate or lanceolate-spatulate, more or less pubescent 



FLORA OF PERU 603 

or glabrous except the midnerve beneath, with the petiole 4-8 mm. 
long, 1-3 mm. broad. 

Peru: Without locality, Ruiz & Pavon. North to Mexico. 

Pycnophyllum peruvianum Muschl. Bot. Jahrb. 45: 457. 1911. 

Stems 3-4 cm. long, densely cespitose, densely leafy, glabrous; 
leaves lanceolate or narrowly so, very acute, fleshy or rigid, densely 
ciliate-margined, otherwise glabrous, lustrous; pedicels 0.5 mm. long, 
glabrous; sepals linear- to ovate-lanceolate, scarious, acutish, nerved; 
petals narrowly ovate; fertile stamens 2-3 mm. long, the style 
slightly longer; seeds triangular, compressed, lustrous. Belongs to 
Arenaria, section Dicranella, according to Mattfeld. 

Junin: Near La Oroya, 430 meters (Weberbauer 2597, type). 

Scleranthus peruvianus Muschl. Bot. Jahrb. 45: 460. 1911. 

A densely cespitose perennial, with many ascending, slightly 
branched, glabrous stems 6-10 cm. long, clothed below with dead 
leaf bases; leaves dilated at the base, linear to lanceolate, plano- 
convex, densely ciliate-margined, glabrous, 6-7 mm. long, 1-2 mm. 
broad; flowers subsessile, in loose cymes; sepals oblong to ovate, 
the midnerve thick, mostly glabrate; petals 5, subulate, about as 
long as the sepals; stamens 5, the filaments strongly dilated at the 
base. This plant is a species of Arenaria, according to Mattfeld in 
Herb. Berlin. 

Cajamarca: Above Hualgayoc, 4,000 meters (Weberbauer 3985, 
type). 

Arenaria Poeppigiana Rohrb. Linnaea 37: 265. 1872. 

Closely allied to A. lanuginosa, but the quadrangular stems with 
many short, sterile, very leafy branchlets, a few elongate, ascending- 
erect, floriferous at the apex, 5-20 cm. high; pubescence of simple 
and stellate trichomes intermixed; sepals acutish, green, narrowly 
scarious-margined, the outer puberulent, the inner glabrescent except 
the hirsutulous nerve. A. soratensis Rohrb. op. cit. 266, has ovate- 
lanceolate leaves 3-4 mm. broad and long-acuminate sepals, scarious- 
ciliate, green, the nerve scabrous, shorter than the petals. A. Jame- 
soniana Rohrb. op. cit. 267, with leaves scarcely 1 mm. broad, calyx 
lobes acuminate, puberulent, petals none, capsule half as long as the 
calyx, the plant glabrescent, may occur. Neg. 27731. 

Huanuco: Casapi, Poeppig 1402a (type). Huacachi, rock pockets, 
3867. Huallaga, 1,600 meters, Weberbauer 6814. 



604 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Arenaria pycnophylloides Pax, Bot. Jahrb. 18: 29. 1893. 

Glabrous or more or less ciliate; leaves lanceolate, acuminate, 
lustrous, subcoriaceous, densely ciliate to the apex and on the promi- 
nent midnerve beneath, above more or less concave; sepals oblong, 
acuminate, ciliate, broadly thin-margined; petals obtuse, minute 
or wanting; ovary depressed-globose; styles 3. Stems greatly 
abbreviated, densely imbricate-leafy, forming low cushions (var. 
compacta Pax) ; var. Moritziana Pax ex Williams has longer branches, 
leafless below, and eciliate leaves and sepals. 

Junin: Cerro de Pasco, Ruiz & Pavon (det. Mattfeld as "aff."). 
Colombia; Argentina. 

Alsine rupestris Muschl. Bot. Jahrb. 45: 448. 1911. 

Densely pulvinate, with many ascending or rarely decumbent, 
usually simple, glabrous stems 3-10 cm. long; leaves lanceolate or 
often ovate, with an amplexicaul base, acute, densely scaly-puberu- 
lent or glabrate, ciliate, many-nerved, scarcely 8 mm. long, 2-3 mm. 
wide; pedicels 5 mm. long, erect, more or less curved beneath the 
calyx, rather densely spreading-puberulent or glabrate, as also the 
green, oblong sepals, these 3.5-5 mm. long; petals ovate-cuneate, 
4.5 mm. long, equaling the capsule; seeds minutely tuberculate. 

Puno: Above Ananco, 5,100 meters (Weberbauer 1042, type). 

Arenaria soratensis Rohrb. Linnaea 37: 266. 1872. 

Similar in habit to A. lanuginosa, but with stouter stems and 
firmer foliage, the former ashy-puberulent, the latter less so, drying 
yellowish green; leaves ovate-lanceolate, to 1 cm. long, 3-4 mm. 
broad, the lower imbricate; pedicels spreading or curved below the 
calyx, 10-20 mm. long; sepals ovate-lanceolate, long-acuminate, the 
narrow apex recurving; petals ovate-oblong, obtuse, slightly exserted; 
capsule included; seeds lenticular, smooth, lustrous. Neg. 27732. 

Peru: (Probably). Bolivia. 

Arenaria Standleyi Baehni & Macbr., sp. nov. 

Caespitosa ramosissima; caulibus e radice repente numerosis 
brevibus dense fastigiatis 1-2 cm. altis; foliis imbricatis ovato-lan- 
ceolatis subcoriaceis acutis obscure pungentibus vix 3 mm. longis 
circa 1 mm. latis plus minusve involutis ad marginem inferne minu- 
tissime ciliatis; floribus solitariis inter folia suprema ramorum; 
sepalis 2.5-3 mm. longis acutis paullo inflexis; petalis obovato- 
ellipticis 1 mm. longis calyce multo brevioribus; staminibus 10, disco 
obscuro. Apparently nearly A. aphanantha Wedd., which, however, 



FLORA OF PERU 605 

has subequal petals and sepals and aristate-pungent leaves. The 
styles are either 2 or 3. 

Ayacucho: Forming small, flat cushions in rocks, Mt. Razuhuillca, 
Prov. Huanta, 4,200 meters, Weberbauer 7489, type, Herb. Field 
Museum. 

Arenaria Stuebelii Hieron. Bot. Jahrb. 21: 307. 1895. 

Stems numerous, ascending-erect, little branched unless at the 
base, to 10 cm. high, more or less minutely hirsutulous-puberulent; 
leaves glaucous, linear-lanceolate, acute or subacuminate, minutely 
puberulent and marginally scabrous, about 8 mm. long, 1 mm. broad; 
pedicels erect, to 1.5 cm. long; sepals ovate, scarious-margined, the 
outer acuminate, ciliate at the base, scabrous, 3.5-4 mm. long, the 
interior obtuse, mucronate; petals ovate-oblong, obtuse, 6.5 mm. 
long, 3 mm. wide; ovary ovate. 

Peru: (Probably). Bolivia. 

Arenaria tetragyna Willd. ex Schlecht. Ges. Naturf. Berl. Mag. 
7: 201. 1813. 

Much branched, creeping or ascending, diffuse, glabrous or 
sparsely pilose; leaves more or less approximate, pale green, plane, 
oblong-lanceolate, acutish or obtuse, membranous, scarcely narrowed 
to the base, very shortly ciliate, 6-10 mm. long, 1.5-2 mm. broad; 
flowers solitary at the branchlet tips, often long-peduncled; sepals 
obtuse, narrowly scarious-margined, glabrous, 3.5-4 mm. long; 
petals ovate-elliptic, short-clawed, shorter than the calyx; capsule 
tardily dehiscent, equaling the calyx; seeds large, obsoletely punctu- 
late. Some at least of the material in herbaria as A. serpens HBK. 
matches our plant exactly, and is rather referable to A. tetragyna 
because of the disk glands. 

Peru: (Dombey). Lima: Rio Blanco, 5,000 meters, 3019. Cuzco: 
Raya, 4,350 meters, Pennell 13528. Chile; Ecuador; Mexico. 

Alsine Weberbaueri Muschl. Bot. Jahrb. 45: 448. 1911. 

Densely pulvinate, with an indurate caudex, the soon glabrous, 
closely leafy stems 2-4 cm. long; leaves linear or linear-lanceolate, 
narrowed to a petiole, acute, the younger pubescent on both sides 
and often ciliate, many-nerved, thick, 10-15 mm. long, 0.5-2 mm. 
wide; pedicels filiform, 1 cm. long, more or less curved below the 
calyx, somewhat puberulent or sometimes glabrous; sepals oblong, 
3-5 mm. long, the ovate, cuneate petals about as long as the capsule. 

Lima: At 4,500 meters above Lima (Weberbauer 5160, type). 



606 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

6. PYCNOPHYLLUM R<my 

Plettkea Mattf . ; Pycnophyllopsis Skottsb. 

Reference: Mattfeld, Repert. Sp. Nov. 18: 167-179. 1922. 

Characteristically cushion plants, sometimes small and then 
distinguished in flower from a few species of Arenaria only by the 
presence of 1-4 ovules instead of several in the 1-celled ovary, the 
fruit a utricle. Leaves small or minute, bract-like, densely imbricate 
in rows or spiraled. Flowers small or nearly minute, terminal or 
apparently lateral, often concealed by the bracts. Styles 2-3, entirely 
free (Plettkea, Pycnophyllopsis), free nearly to the base (P. Lech- 
lerianum), or more or less united (remaining species). Receptacle 
infundibuliform (Plettkea) or campanulate, with a truncate or some- 
what sunken base (Pycnophyllopsis), the ovary in the former case 
more or less sunken in the tube. The species with completely 
divided styles approach in this single character some genera of the 
Alsineae and are more closely allied than indicated by Pax and 
Hoffmann to Habrosia and Scleranthus, which, however, are totally 
different in aspect. Therefore, logical though Mattfeld's segregation 
of Pycnophyllum may be (see Pflanzenfam. ed 2. 16c: 367. 1934), 
it seems to serve no useful purpose in floristic or phytogeographic 
work, and accordingly is not accepted here, where the convenience 
of a single group name for these often conspicuous cushion plants of 
the high Andes is a first consideration. In monographic work, where 
a prime intent is to show relationship in development, Plettkea could 
and perhaps should be maintained as either a generic or sectional 
entity. To avoid duplication of a specific name, one transfer is made. 
Besides the following, Pycnophyllopsis keraiopetala Mattf. op. cit. 22, 
may be expected; it is distinguished from P. glomeratum Mattf. by 
its oblong-lanceolate, ciliate leaves and 3 completely divided styles. 

Leaves scarious, orbicular, apiculate, appressed; branches mostly 

only 8-15 mm. long, 2 mm. thick P. Aschersonianum. 

Leaves not as above, at least not in all respects. 
Leaves more or less spreading, at least the tips of the upper aristate, 

scarious. 

Leaves all spreading, the tips filiform, curling. . . .P. aristatum. 
Leaves aristate, but abruptly, the points straight. 

P. Holleanum. 

Leaves all closely appressed or not scarious and not aristate. 
Leaves scarious (thin) or orange-colored and scarious-margined, 
often conspicuously calloused; petals present. 



FLORA OF PERU 607 

Flowers 4-5.5 mm. long; leaves not orange-colored in age. 
Leaves constricted below the middle, minute, orbicular, 
neither apiculate nor notably calloused; branchlets 
club-shaped; petals well exserted . . . P. macropetalum. 
Leaves not constricted below the middle. 
Leaves lightly calloused; branchlets not or scarcely club- 
shaped; sepals broadly obovate P. molle. 

Leaves conspicuously calloused; branchlets filiform; sepals 

obovate P. Markgrafianum. 

Flowers 3-4 mm. long; leaves orange-colored in age. 

P. bryoides. 

Leaves opaque, firm, not at all scarious or not orange-colored. 

Branches not quadrate, slender, all but the ultimate about 

2 cm. long; leaves loose; styles nearly 3-parted; petals 

none P. Lechlerianum. 

Branches quadrate, the lower often only about 1 cm. long; 

styles completely separated, 2-3; petals present. 
Leaves straw-colored in age, 5-7 mm. long, clearly spiraled. 
Sepals 5; styles 3; ovary almost completely immersed in 

the calyx base P. macrophyllum. 

Sepals 4; styles 2; ovary partly immersed in the calyx 

base Plettkea cryptantha. 

Leaves about 2.5 mm. long, not clearly spiraled. 

Flowers 3.5-4 mm. long; leaf cilia 1-celled . . P. Mattfeldii. 

Flowers 2.5 mm. long; leaf cilia 2-celled. .P. Weberbaueri. 

Branches short, quadrate; leaves not clearly spiraled; styles 

united; petals none. 
Branches 1.5-2 mm. thick. 

Branchlets 5-18 mm. long, few P. tetrastichum. 

Branchlets 3-4 mm. long, many P. glomeratum. 

Branches very slender, 1 mm. thick. 

Flowers 2.5 mm. long; branches 1 mm. thick. P.filiforme. 
Flowers 4-4.5 mm. long; branches nearly 1.5 mm. thick. 

P. leptothamnum. 

Pycnophyllum aristatum Mattf. Repert.Sp.Nov.18: 178. 1922. 

Closely allied to P. Holleanum, but the awns of the ovate leaves 
3-5 mm. long, the bracts sheathed and lightly calloused, the petals 
3 mm. long. 



608 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Ancash: Mt. Ururupay, Prov. Pallasca, in nearly sterile stone 
slides, 4,800 meters, Weberbauer 7250. Cabrococha, 4,600 meters, 
Weberbauer. 

Pycnophyllum Aschersonianum Muschl. Bot. Jahrb. 45: 
455. 1911. 

Densely cespitose; branches apically enlarged, very densely leafy; 
leaves opposite, concave, 3-4-seriate, orbicular-obovate, rarely 1 
mm. broad, 2-3 mm. long. According to Mattfeld, op. cit. 179, 
this species, so far as Muschler's description of the flowers is con- 
cerned, is based on his well known imagination, the type material 
being sterile. However, the species appears to be closely related 
to P. molle and P. spathulatum. 

Ancash: Vicinity of Huaraz, 4,500 meters, Weberbauer 3293 (type). 

Pycnophyllum bryoides (Phil.) Rohrb. Linnaea 36: 662. 1870. 
Stichophyllum bryoides Phil. Fl. Atac. 19. pi 1. 1860. 

Stems numerous, much branched, to 15 cm. high, the branchlets 
scarcely enlarging apically; leaves ovate-lanceolate, acuminate, 
broadly scarious-margined, minutely serrulate, 2-2.5 mm. long; 
flowers pedicellate; calyx 3 mm. long, the lanceolate, obtuse lobes 
scarious, longitudinally nerved; petals 5, oblong-linear, bifid nearly 
to the base, the lanceolate divisions very acute, nearly 2 mm. long, 
about equaled by the fertile stamens; style simple, the stigmas 3. 
Peruvian specimens have flowers 4-4.5 mm. long (Mattfeld). 

Arequipa: Nearly sterile, sandy slopes, eastern side of Mt. 
Misti, 4,400 meters, Weberbauer 1420; 218 as P. argentinum Pax. 
Puno: Picutani, 4,400 meters, Weberbauer 501b. Chile. 

Plettkea cryptantha Mattf. Schrift. Ver. Nat. Unterweser 
N. F. 7: 15. 1934. 

Pulvinate, the woody branches flabellately branching, the older 
naked, 3 mm. thick, the younger, with the spirally imbricate leaves, 
5 mm. in diameter, the ultimate in pairs with a solitary flower 
concealed in the tip; leaves ovate-lanceolate, amplexicaul, acute, 
mucronulate, coriaceous, the margins ciliolate above, 5-6 mm. long, 
about 3 mm. broad at the base; flowers sessile, perfect, tetramerous, 
5.5 mm. long; receptacle infundibuliform, to 1.5 mm. high; sepals 
4 mm. long, the outer oblong, ciliolate, the inner ovate or elliptic, 
glabrous; petals scarcely 2 mm. long, parted nearly half their length, 
broadly obovate; styles 2, not at all joined outside the ovary wall, 
the ovary itself half sunken in the receptacle; ovules 4. 

Junin: Morococha (Raimondi; type of the genus Plettkea). 



FLORA OF PERU 609 

Pycnophyllum filiforme Mattf. Repert. Sp. Nov. 18: 172. 1922. 

Forming large cushions, the much branched branches filiform, 
1-1.5 mm. thick; leaves rarely spiraled, the white, membranous 
sheaths 0.5-0.8 mm. broad, toward the base scarious-margined and 
minutely papillose, calloused, mucronulate, 2 mm. long; flowers 
dioecious, the staminate 2.5 mm. long, exceeded by the ovate bracts; 
sepals ovate-elliptic, with oblong callus, 3-nerved. 

Arequipa: Lake Villafro, 4,700 meters, Weberbauer 6885. 

Pycnophyllum glomeratum Mattf. Repert. Sp. Nov. 18: 
171. 1922. 

Branches and branchlets erect, short, about 5 cm. long, the 
ultimate pulvinate or glomerulate; leaves densely imbricate, oppo- 
site, decussate or often spiraled; sheaths glabrous, infundibuli- 
form, to 1.2 mm. long; leaf blade broadly ovate, about as long, 
membranous-margined, mucronulate, cymbiform-concave; bractlets 
3 mm. long; flowers dioecious, the staminate cylindric, not at all 
gibbous, 4 mm. long; sepals 3-nerved from the base; pistillate 
flowers similar, with sterile stamens; style 1 mm. long, with branches 
0.2 mm. long; seeds 1-2, minutely reticulate. To be expected 
is P. Stuebelii Mattf. of Bolivia with the leaves 3-3.5 mm. long, 
including the half shorter sheath, this densely short-papillose; 
branchlets about 12 cm. long. 

Puno: Picutani, 4,400 meters, Weberbauer 501 a (type). Sandia, 
near Poto, 4,600 meters, Weberbauer 952; 219, as P. convenum 
Griseb. Cuzco: Below glaciers of Auzangate, Weberbauer 7774 
(det. Mattf eld). 

Pycnophyllum Hoi lean um Mattf. Repert. Sp. Nov. 18: 
177. 1922. 

Stems ligneous, the erect, flabelliform branches to 20 cm. long, 
2 mm. thick; leaves densely spiraled, obovate, 2.5 mm. long, abruptly 
awned, the awn 0.5-1 mm. long; upper bracts ecallose; petals retuse, 
1.8 mm. long, the broadly obovate sepals 5 mm. long. 

Junin: Prov. Huancayo, above Hacienda Acocarpa, 4,900 meters, 
Weberbauer 6522. 

Stellaria laevis (Bartl.) Rohrb. Linnaea 37: 275. 1872. Cher- 
leria laevis Bartl. in Presl, Rel. Haenk. 2: 12. 1831. 

A nearly glabrous perennial, much branched from the base, 
the many diffuse stems covered below with dead leaves; leaves 4-6 
mm. long, coriaceous, lustrous, ovate, connate at the base, acutely 



610 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

acuminate, mucronulate, ciliolate; flowers solitary at the tips of the 
branches, sessile; sepals scarcely 2 mm. long, oblong-lanceolate, 
obtusish, faintly 3-nerved; stamens 5; petals mostly bifid, inserted 
with the 5 stamens in a ring; ovary deep in the calyx; stigmas 3. 
Mattfeld, Schrift. Ver. Naturk. Unterweser N. F. 7: 21. 1934, 
thinks that this belongs to his genus Plettkea. There is in Herb. 
Delessert a scrap of a Pycnophyllum-like plant, apparently sterile, 
and purporting to be by Pa von, which is labeled by Fenzl "S. 
lycopodioides Fenzl," a manuscript name mentioned by Rohrbach. 
op. cit., as referable to the Presl plant. 
Lima: Obrajillo. 

Pycnophyllum Lechlerianum Rohrb. Linnaea 36: 664. 
1870; 174. 

Laxly cespitose, the branches not club-shaped; leaves coriaceous, 
with a small, medial callus, broadly ovate-lanceolate, attenuate 
apically, narrowed basally, 6 mm. long; calyx broadly obovoid, 
strongly enlarged, 3 mm. long; sepals nearly scarious, lanceolate, 
obtuse, mucronulate; petals none; ovary hexagonous; stigmas 3, 
parted nearly to the base. Leaves, according to Mattfeld, mem- 
branous. Neg. 29832. 

Puno: Azangaro, 3,820 meters, Lechler 1742 (type). 

Pycnophyllum leptothamnum Mattf. Notizbl. Bot. Gart. 
Berlin 10: 1051. 1930. 

With the habit and long, slender stems of P. filiforme but in 
other characters resembling most P. glomeratum; leaves slightly if 
at all spiraled, the sheath 0.8 mm. long, the blade 2 mm. long, 
softly muticous at the narrowed tip, opaque, straw-colored, ob- 
scurely calloused, narrowly scarious; floral bracts 4 mm. long; 
sepals mucronulate; 1 petal rarely present; style 1.5 mm. long with 
3 or sometimes 2 short branches. 

Cuzco: In puna near glaciers of Auzangate, 4,600 meters, Weber- 
bauer 7772 (type). 

Pycnophyllum macropetalum Mattf. Repert. Sp. Nov. 18: 
176. 1922. 

Stems to 15 cm. long, the thicker branches to 3.5 mm. thick 
toward the tip; leaves densely spiraled, shortly sheathed, the blades 
obovate, conspicuously constricted below the middle, 2-2.5 mm. long, 
rounded at the apex, minutely mucronulate, pale straw-colored, 
obscurely calloused; bracts spatulate, longer than 4 mm., not 



FLORA OF PERU 611 

calloused; flowers dioecious, the pistillate obovoid, 6 mm. long, 
the nearly orbicular sepals papillose below the apex, with a green 
callus 4 mm. long; petals obovate, deeply bifid, 5 mm. long; styles 
1.5 mm. long, the branches 0.5 mm. long. P. spathulatum Mattf., to 
be expected, has obovate, obtuse or denticulate petals to 1.5 mm. long. 
Puno: Cerro Tacora, 4,500 meters (Stuebel 110, type). 

Pycnophyllum macrophyllum Muschl. Bot. Jahrb. 45: 458. 
1911 ; 179. Plettkea macrophylla Mattf. Schrift. Ver. Nat. Unterweser 
N. F. 7:18. 1934. 

With the habit of Plettkea cryptantha, and recalling Lycopodium 
Selago; leaves lanceolate or ovate-lanceolate, acuminate, densely 
ciliate, 5 mm. long; flowers sessile, terminal, 4.5 mm. long, the 
infundibuliform receptacle 2 mm. deep; sepals ovate, coriaceous, 
2-5 mm. long, the slightly shorter petals obovate; ovules about 3. 
Description from Mattfeld, op. cit. 18. 

Ancash: Huaraz, 4,500 meters, Weberbauer 2975 (type). 

Pycnophyllum Markgrafianum Mattf. Repert. Sp. Nov. 18: 
175. 1922. 

Similar to P. molle, but the sepals obovate or obovate-elliptic, 
narrowed to the base, and the leaf callus light brown. 

Ancash: Prov. Cajatambo, 4,400 meters, Weberbauer 2783 (type); 
224, as P. molle. Lima: At 4,500 meters, at the silver mine Alpa- 
mena, Weberbauer 5121. 

Pycnophyllum Mattfeldii Macbr., nom. nov. Plettkea tetra- 
sticha Mattf. Schrift. Ver. Nat. Unterweser N. F. 7: 21. 1934, not 
Pycnophyllum tetrastichum Re"my. 

Very similar to P. Weberbaueri, but more robust; leaves and 
flowers slightly longer, the sepals 2.5 mm. long; cilia of the leaves 
all 1-celled. 

Cajamarca: Above Ocros, 4,800 meters, Weberbauer 2804a (type). 

Pycnophyllum molle Re"my, Ann. Sci. Nat. III. 6: 355. pi. 20. 
1846; 174. P. aculeatum Muschl. Bot. Jahrb. 45: 456. pi. 1. 1911. 

Stems many, to 20 cm. long, scarcely enlarged toward the tip; 
leaves spiraled, orbicular-obovate, very blunt, lustrous, scarious- 
margined, scarcely 2 mm. long; leaf callus green or pale straw- 
colored; bracts obovate; sepals broadly ovate; petals oblong-linear, 
not dilated, obtuse, entire or emarginate, bifid or 3-dentate; ovary 
trigonous; seeds pyriform, smooth. Flowers 4.5-5.5 mm. long; 



612 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

leaves of the Peruvian plant less densely imbricate (Mattfeld). 
Illustrated, Weberbauer 205. 

Puno: Sandia, stony ground near Poto, 4,400 meters, Weberbauer 
946, 950; 219. Tacora, 4,500 meters, Stuebel 109. Arequipa: 
Vincocaya, 4,100 meters, Weberbauer 1374; 205 (type of P. aculea- 
tum). Nevado de Chachani, 4,400 meters, Pennell 13315. Caja- 
marca: Ocros (Weberbauer 224)- Ancash: Huarapasca, 4,900 meters, 
2488. Moquehua: Carumas, 4,500 meters, Weberbauer 7315, 7356 
(det. Mattfeld). Bolivia. 

Pycnophyllum tetrastichum Re"my, Ann. Sci. Nat. III. 6: 
356. pi. 20. 1846; 172. 

Branches erect, forming very dense cushions, the several slender 
branchlets scarcely if at all enlarged apically; leaves narrowed to 
the apex, gradually convex-curved, the callus not definitely defined, 
broadly ovate, 4-5 mm. long; calyx 4.5 mm. long, the ovate-oblong 
sepals acutish, scarious, 3-nerved; petals none; style short-trifid at 
the apex; ovary trigonous. Leaves, according to Mattfeld, as 
short as 2 mm. 

Puno: Pisalomo, 4,450 meters, Mcyen. Lake Titicaca, Hill 53. 
Bolivia. 

Pycnophyllum Weberbaueri Muschl. Bot. Jahrb. 45: 455. 
1911; 179. Plettkea Weberbaueri Mattf. Schrift. Ver. Naturk. 
Unterweser N. F. 7: 19. 1934. 

Stems much branched, forming cushions, to 4 cm. long; leaves 
ovate or ovate-lanceolate, 1 mm. long, clasping at the base, blunt, 
mucronulate, subcoriaceous, marginally ciliate, the lower hairs 
2-celled, the upper 1-celled; sepals oval, 1.8 mm. long, papillose- 
ciliolate, about twice longer than the rotund-ovate petals; styles 
3-parted to the base; ovules about 3. Description from Mattfeld, 
loc. cit. 

Arequipa: Vincocaya, 4,100 meters, Weberbauer 1373, type. 

7. PARONYCHIA L. 

Annual herbs or even half shrubby, with sometimes falsely 
whorled leaves. Flowers few, usually axillary, often concealed by 
the large stipules. Sepals usually 5, the petals (staminodia) some- 
times lacking. Fruit fragile. Embryo curved. 

Leaves more or less concealed by the imbricate stipules, or also 
imbricate, tiny; flowers mostly solitary. 



FLORA OF PERU 613 

Stipules extraordinarily imbricate, concealing all or nearly all 

the leaves P. membranacea. 

Stipules, except toward the branchlet tips, not densely imbricate, 

the leaves more or less evident. 
Calyx 2.5-3 mm. long, the lobes densely soft-pubescent even 

to the mucro P. andina. 

Calyx scarcely 2 mm. long, the lobes more or less puberulent or 

glabrescent P. Mandoniana. 

Leaves markedly longer than the slightly or not at all imbricate 

stipules; flowers usually glomerulate. 
Calyx glabrous, the mucronate apex of the sepals about one-third 

as long as the herbaceous base P. brasiliana. 

Calyx more or less pubescent, the lobes short-mucronulate. 

Leaves exceeding the internodes P. chilensis. 

Leaves shorter than the internodes P. microphylla. 

Paronychia andina Gray, U. S. Expl. Exped. 128. 1854. 

Depressed, tufted, the caudex branches much crowded, the 
stems 5-15 cm. long, nearly glabrous; stipules imbricate, oblong- 
lanceolate; leaves coriaceous, nearly imbricate, ovate-oblong, awn- 
pointed, puberulent or glabrate, 2-3 mm. long, scarcely exceeding 
the stipules; flowers subsessile, terminally congested with the 
silvery-scarious bracts; calyx softly puberulent, the sepals oblong, 
scarious, the midrib shortly awn-produced; stamens 5, setaceous, 
shorter than the sepals. Growing from 3,000 meters to the limit of 
vegetation (Gray). 

Lima: Viso, sterile, stony summit, 624- Banos (Wilkes Exped., 
type). Rio Blanco, 4,500 meters, stony slope, 3048. Casa Cancha 
to Culnai and Alpamarca (Wilkes Exped.) . Same region (McLean). 
Junin: Cerro de Pasco (Mathews). Cuzco: Cerro de Colquipata, 
4,100 meters, open, grassy puna, Pennell 13740. Hills of Sacsa- 
huaman (Herrera 21140); at 3,600 meters, Herrera 2375. Bolivia. 

Paronychia bonariensis DC. Prodr. 3: 370. 1828. P. bra- 
siliana DC. in Lam. Encycl. 5: 23. 1804. 

Cespitose, the numerous stems prostrate; leaves elliptic, acute 
at the base, acuminate, lightly pubescent on both sides, 5-10 mm. 
long, 1-2 mm. broad; stipules lanceolate, long-acuminate, often 
lacerate; flowers glomerate in the leaf axils; sepals shorter than 
2 mm., glabrous, tipped with a slender, straight awn; petals seti- 
form. Illustrated, Mart. Fl. Bras. 14, pt. 2: pi. 57. 



614 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Arequipa: Meyen. Lima: Banos (Wilkes Exped.; det. Gray). 
Without locality (Haenke). Chile; Brazil. 

Paronychia chilensis DC. Prodr. 3: 570. 1828. 

Diffuse, prostrate or ascending, glabrescent or more or less 
pubescent; leaves linear or oblong-lanceolate, attenuate to each 
end, 8-15 mm. long, 1-2 mm. broad; stipules lanceolate, often 
lacerate; flowers axillary, solitary or few, subsessile, the scarious, 
long-acuminate bracts about equaling the turbinate calyx, this to 
2.5 mm. long, deeply 5-parted, the ovate-oblong lobes rigidly mucron- 
ulate. P. Hartwegiana Rohrb. op. cit. 204, of Ecuador, has the 
calyx pubescent only at the base and ovate leaves shorter than the 
internodes; in this last character it resembles P. bogotensis Triana & 
Planch, of Colombia and Ecuador, scarcely distinct from P. chilensis 
except that the sepals are not at all scarious. Illustrated, Mart. 
Fl. Bras. 14, pt. 2: pi. 57. Neg. 7952. 

Puno: At 4,200-5,400 meters (Meyen). Chuquibambilla, 3,900 
meters, crevices of limestone rock, Pennell 13406. Arequipa: 
Tingo, 2,200 meters, open, rocky slopes, Pennell 13136. Huanuco: 
Huanuco, 2,100 meters, stony slopes, 3243. Yanashallas, 4,800 
meters, loose, stony slopes, 2480. Huanuco, rocky cliff 2326 (?). 
Cuzco: Between Pisac and Paucartambo, 4,100 meters, Weber- 
bauer 691 9. Lima: Viso, 758 (?). Chile to Mexico. 

Paronychia Mandoniana Rohrb. Linnaea 37: 208. 1872. 

Cespitose, suffrutescent, densely short-puberulent; leaves crowded, 
spreading, narrowly to broadly lanceolate, cuneate at the base, 
pungently mucronate, densely scabrous, 6-9 mm. long, 1-2 mm. 
broad; stipules ovate-lanceolate, long-acuminate, rarely bifid, 
shorter than the leaves; flowers solitary in the upper leaf axils, 
subsessile; calyx urceolate-turbinate, pale greenish brown, 2 mm. 
long, somewhat puberulent, deeply 5-parted, the narrowly lanceo- 
late, 3-nerved lobes broadly scarious-margined; petals subulate, 
half as long as the calyx lobes; ovary punctate, scabrous, the style 
twice as long, bifid to the middle. Very much like Cardionema in 
appearance. Neg. 30177. 

Lima: Casapalca, 5,000 meters, 852 (or P. andina sens, lat.). 
Puno: Azangaro, Lechler 1760. Bolivia. 

Paronychia membranacea Muschl. Bot. Jahrb. 45: 460. 1911. 

Densely cespitose, the few-branched stems becoming naked 

below, 4-5 cm. long; leaves ovate, mucronulate, minutely puberulent, 



FLORA OF PERU 615 

scarcely 2 mm. long; stipules appressed, broadly oval or suborbic- 
ular, longer than the leaves which they completely conceal; flowers 
solitary in the upper leaf axils, subsessile; calyx oblong-turbinate, 
2.5-3 mm. long, densely puberulent, the 3-nerved, mucronulate, 
oval-oblong lobes whitish except the brownish base; petals yellowish, 
subulate, half as long as the calyx; ovary smooth, the bifid style 
short. The description of the flowers is by Muschler; a cotype in 
Herb. Delessert appears to be sterile. Neg. 30179. 

Cajamarca: Near Ocros, 4,400 meters, Weberbauer 2784. 
Junin: Near Oroya, in limestone rocks, 4,000 meters (Weberbauer 
2536}. 

Paronychia microphylla Phil. Anal. Mus. Nac. Chile Bot. 
1891: 26. 1891. 

Woody, with the form of a small shrub, ascending from the 
base, about 15 cm. high, somewhat villous; leaves elliptic, mucron- 
ulate, hirsute, only 5 mm. long, exceeded by the stipules; petals very 
hirsute without, obtuse, mucronulate. 

Arequipa: At 2,300 meters (Weberbauer, 129). Arequipa, 
2,600 meters, gravelly stream bed, Pennell 13159; rocky ledges, 
Pennell 13203. Southern slopes of Mount Chachani, 3,355 meters, 
Hinkley 75. Tiabaya, 2,100 meters, open, rocky slope, Pennell 
13078. Junin: Yanahuanca, 3,000 meters, shrubby slope, 1200. 
Chile. "Gateadora." 

Paronychia polygonoides Muschl. Bot. Jahrb. 45: 459. 1911. 

Suffrutescent, densely cespitose, prostrate or ascending, soon 
glabrous; leaves crowded, spreading, ovate-lanceolate or narrower, 
gradually narrowed to the base, pungently mucronate, densely 
scabrous, 6-10 mm. long, 3-4 mm. wide; stipules broadly ovate, 
long-acuminate, rarely bifid, glabrous, usually much shorter than 
the leaves; flowers mostly solitary in the upper leaf axils, subsessile; 
calyx urceolate, turbinate, 3 mm. long, puberulent or above glabres- 
cent, the lanceolate or ovate-lanceolate lobes narrowly scarious- 
margined, prominently 3-nerved ; petals subulate, half as long as the 
calyx; stamens 5; style twice as long as the ovary; seeds lenticular, 
lustrous, microscopically reticulate. Neg. 30178. 

Cajamarca: Ocros, 3,500-3,700 meters (Weberbauer 2702, type). 

Paronychia rigida Muschl. Bot. Jahrb. 45: 458. 1911. 
Cespitose, procumbent, densely matted, the much branched 
stems covered below with dead leaf bases, 10-20 cm. long; leaves 



616 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

narrowly ovate, subtruncate or nearly amplexicaul at the base, 
gradually narrowed to the conspicuously aristate apex, minutely 
puberulent, especially toward the apex, becoming glabrate, 3-5 
(-8?) mm. long, 1-2 (-3?) mm. wide; stipules lanceolate, long- 
acuminate; flowers short-pediceled, in few-flowered glomerules; 
bracts scarious, lanceolate, acute, about equaling the ovate-oblong 
calyx, this 2 mm. long, glabrous, its divisions narrowly membranous- 
margined, apically cucullate-mucronate, the spinulose tips nearly 
one-third as long as the calyx lobes; petals setiform, scabrous; style 
minute; fruit tuberculate, included, the seeds brown, smooth, 
narrowly sulcate beneath. All the Weberbauer specimens are cited 
by Muschler, without designation of the type. Negs. 29281 (R. & 
P. specimen), 30180. 

Junin: Near Yauli, 4,400 meters (Weberbauer 291). Oroya, 
3,700-4,300 meters (Weberbauer 2626, 2627). Tarma, 3,500 meters 
(Weberbauer 2410); Ruiz & Pavon (det. Mattfeld). Puno: Poto, 
4,500 meters (Weberbauer 984)- 

8. CARDIONEMA DC. 

Pentacaena Bartl. 

Depressed herbs with needle-like leaves and conspicuous, white 
stipules. Flowers tiny, appressed. Sepals 5, slightly concave, enclos- 
ing the thin-walled fruit that opens irregularly. Stamens 3-5. Petals 
minute, entire or 2-lobed. Embryo straight. 

Cardionema ramosissima (Weinm.) Nels. & Macbr. Bot. 
Gaz. 56: 473. 1913. Loeflingia ramosissima Weinm. Bot. Zeit. 3: 
608. 1820. Pentacaena polycnemoides Bartl. in Presl, Rel. Haenk. 
2: 5. pi. 49. 1831. 

Densely matted, the much branched stems more or less lanate- 
pubescent above and closely clothed with scarious, appressed 
stipules and glabrate, subulate, mucronate leaves, these 4-8 mm. 
long; flowers sessile in the leaf axils; calyx lobes glabrate or pubes- 
cent, often very unequal, the 3 longer usually 4-6 mm. long, long- 
mucronate, the interior shortly so; fertile stamens 3-5; seeds obovoid, 
smooth. Several forms have been recognized, perhaps not spe- 
cifically distinct, including C. camphorosmoides (Camb.) Nels. & 
Macbr. of Brazil, more diffuse, with puberulent, carinate leaves and 
subequal calyx lobes; C. andina (Phil.) Nels. & Macbr. of Chile, a 
tiny plant with solitary flowers; C. congesta (Benth.) Nels. & Macbr., 
of Ecuador (Neg. 30175), apparently better marked by reason of its 



FLORA OF PERU 617 

shorter leaves that are spinulose, the cartilaginous tip fully half as 
long as the herbaceous part of the leaf. Illustrated, DC. Me"m. 
Paronychia pi. 4; Mart. Fl. Bras. 14, pt. 2: pi. 56, apparently a 
diffuse form or species. 

Huanuco: Chavanillo, 2,400 meters, dry bank, 1976. Lima: 
Rio Blanco, 4,500 meters, dry, stony slope, 809. Puno: Chuqui- 
bambilla, 4,000 meters, rocky, clay soil on puna, Pennell 13354. 
Western North America; Chile, Brazil, Argentina. 

9. DRYMARIA Willd. 

Slender, rather openly branched herbs, rarely simple, with tiny 
flowers in axillary or terminal cymes. Sepals and petals (stami- 
nodia) 5, the latter with 2 or 4 parts. Stamens 5, sometimes by 
abortion fewer. Besides the following, D. arenarioides Willd., D.. 
ramosissima Schlecht., and D. sperguloides Gray were reported by 
Weberbauer, the first from the Titicaca region and Huaraz, the 
second from above Supe in the Department of Lima, and the third 
from above Samanco, in Ancash. As the first two are otherwise 
known only from Mexico and the last from Texas, it is more than 
probable that the Weberbauer references apply to one or the other 
of the following species, which, however, may not all be distinct 
from northern species. D. molluginea (Lag.) Didr., as to Peru 
(Weberbauer, 139, 144, and Bruns, Mitt. Inst. Allg. Bot. Hamburg 
8: 48. 1929), is Spergularia collina I. M. Johnston. 

Petals none; plants coarse. 

Calyx 4-5 mm. long D. apetala. 

Calyx 8-10 mm. long D. macrantha. 

Petals present, sometimes shorter than the sepals. 
Plants shrubby or perennial and more or less enduring at the base. 

Shrub with awl-like, appressed leaves D. frutescens. 

Perennials, more or less woody below, the leaves not awl-like. 
Leaves oblong-lanceolate, glabrate or not ovate. 

Leaves lance-acuminate D. auriculipetala. 

Leaves oblong-ovate, obtuse, apiculate D. squarrosa. 

Leaves ovate, puberulent D. stereophylla. 

Plants annual or biennial or at least herbaceous to the base. 

Flowers large, about 10 mm. long D. grandiflora. 

Flowers much smaller. 



618 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Pubescence, at least on pedicels, definitely viscid-glandular; 
flowers rarely densely fasciculate. 

Leaves petioled or subpetioled, the radical evanescent. 
Fruiting pedicels divaricate or deflexed, the lower 

elongate D. divaricata. 

Fruiting pedicels erect or suberect, 2-10 mm. long. 
Stems glabrous; seeds bluntly tuberculate. 

D. viscidula. 
Stems glandular; seeds acutely tuberculate. 

D. agapatensis. 

Leaves of the stem sessile, the radical persisting . D. praecox. 
Pubescence eglandular or the glandulosity very obscure. 
Leaves mostly well petioled, the lower petioles often 6 mm. 

long. 
Leaves, at least some of them, conspicuously if sparsely 

long-hirsute D. hirsuta. 

Leaves not at all hirsute with long hairs. 
Leaves puberulent, 2.5-3.5 cm. long; peduncles villous. 

D. ovata. 
Leaves glabrous or glabrate, shorter; peduncles not 

villous. 
Leaves well petioled, even the upper. 

Plants many-flowered, the flowers about 3.5 mm. 

long D. divaricata. 

Plants few-flowered, the flowers 1-3 or few, about 

2.5 mm. long D. pauciflora. 

Leaves subsessile or the petioles very short. 

D. cordata. 

Leaves sessile or subsessile, even the lower, the basal some- 
times petioled. 

Sepals more or less rigidly cuspidate-acuminate. 
Cymes head-like, the flowers not clearly separated. 

D. fasciculata. 
Cymes dense but dichotomous, each flower bracted 

and separated D. devia. 

Sepals obtuse or acute but not aculeate-acuminate. 
Sepals keeled by the strong midnerve . . D. Engleriana. 



FLORA OF PERU 619 

Sepals not keeled, the midnerve obsolete or incon- 
spicuous. 
Petals and sepals about equal in length or the former 

shorter. 

Inflorescence an open cyme. 
Sepals oblongish; petals gradually clawed. 
Radical leaf rosettes marcescent; leaves round- 
ish, obtuse or acute D. cordata. 

Radical leaf rosettes persisting; leaves ovate, 

apiculate D. praecox. 

Sepals roundish; petals abruptly clawed. 

D. sphagnophila. 
Inflorescence not an open cyme. 

Flowers cymose-fasciculate D. nitida. 

Flowers in short, dense racemes . . D. glaberrima. 
Petals distinctly exceeding the sepals. 

Flowers showy; hill plant D. rotundifolia. 

Flowers rather inconspicuous, the petals slightly 
longer than the sepals; loma plant. 

D. Weberbaueri. 

Drymaria agapatensis Baehni & Macbr., sp. nov. 

Herba annua suberecta ad 20 cm. alta; ramis paucis fere strictis 
teretibus, internodiis elongatis e basi fere ad apicem minute glandu- 
losis; stipulis setaceis; foliis late rotundato-reniformibus obtusis vel 
subacutis valde ad basin angustatis breviter petiolatis vel sub- 
sessilibus membranaceis glabris, interdum circa 1.5 cm. longis latis- 
que, nervis tenuibus vix notatis; floribus subglobosis laxe dichotomo- 
cymosis; pedicellis plerumque erectis vel suberectis 5-10 mm. longis 
minute glandulosis; sepalis subelliptico-ovatis anguste scarioso- 
marginatis nervo medio inferne incrassato glanduloso 4.5 mm. longis, 
stylo ad medium vel ultra 3-partito; seminibus acute tuberculatis. 
This was distributed as D. divaricata, but it seems clearly distinct 
in its short petioles, glandular stems, and suberect pedicels. 

Puno: Agapata, Lechler 1947, type in Herb. Delessert. 

Drymaria apetala Bartl. in Presl, Rel. Haenk. 2: 7. 1831. 

Erect, the erect branchlets more or less compressed at the nodes; 
leaves ovate or ovate-lanceolate or the lowest subcordate, acute, 
subsessile, 12-16 mm. long, 8-10 mm. broad, the smaller upper ones 



620 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

acuminate; flowers many, in terminal, fastigiate corymbs; sepals 
4-5 mm. long, oblong, acutish, slightly and minutely puberulent like 
the always erect, somewhat longer pedicels; capsule included, oblong; 
seeds about 20, blackish. 

Ancash: Ocros, 2,700 meters, Weberbauer 5814- Ayacucho, 3,300 
meters, Weberbauer 5827. Lima: Canta, Ruiz & Pavon (det. Matt- 
feld). Obrajillo, cliff ledges, Pennell 14321 (det. Mattfeld). Chile. 

Drymaria auriculipetala Mattfeld, sp. nov. in herb. 

Suifrutex vel ramis juvenilibus herbaceis, ramis simplicibus 
gracilibus numerosis ad 60 cm. longis sat angulatis valde nodulosis 
demum teretibus, glabris vel superne minutissime sparseque pul- 
verulentis; stipulis rigido-setaceis 1.5 mm. longis tarde caducis; 
foliis coriaceis glabris vel glabratis anguste ovato-lanceolatis acumina- 
tis sessilibus ad 10 mm. longis, 2 mm. latis, subtrinerviis plus minusve 
cartilagineo-marginatis, nervis supra vix notatis subtus prominenti- 
bus praesertim costa media; floribus solitariis vel in cymis paucifloris, 
pedunculis interdum dichotomis; pedicellis 1-2 mm. longis; sepalis 
oblongo-lanceolatis cuspidato-acutis, basi conspicue incrassatis, 
8 mm. longis; petalis basi auriculatis apice bifidis, 7 mm. longis; 
seminibus complanatis fere orbiculatis tuberculatis. 

Huanuco: Pendent from cliff ledges, 2,500 meters, Llata, Mac- 
bride 2264, type in Herb. Field Mus. 

Drymaria cordata (L.) Willd. ex R. & S. Syst. 5: 406. 1819. 
Holosteum cordatum L. Sp. PI. 88. 1753. 

Plants tiny and simple or nearly so or larger and loosely much 
branched, glabrous or slightly pubescent above; leaves nearly orbicu- 
lar or broader than long, usually exceeded by the often forked 
peduncles; pedicels and calyces subequal, the petals shorter than the 
acute sepals, these 1.5-3 mm. long. Sometimes rather weed-like in 
moist, disturbed ground and, distributed nearly everywhere in 
tropical America, no doubt in Peru, where probably masquerading 
under one or more of the specific names here included ; see Rohrbach 
in Mart. Fl. Bras. 14, pt. 2: 259, where illustrated, pi. 60. 

Lima: Matucana, 189. Huanuco: Mito, in clearing, 1714- 
Mufia, 3918. Tropical America. 

Drymaria devia Baehni & Macbr., sp. nov. 

Planta annua minima plus minusve diffusa fere glabra, caulibus 
1-3 (-6) cm. longis simplicibus vel paullo ramosis; foliis sessilibus 
coriaceo-rigidis ovatis acutis vel apiculatis 3-5 mm. longis, 2-3.5 mm. 



FLORA OF PERU 621 

latis, nervis obscuris; floribus subsessilibus binato-scarioso-bracteatis 
plerumque 3 in fasciculam plus minusve congestis; bracteis 2.5 mm. 
longis acuminatis; sepalis oblongo-lanceolatis aculeato-acuminatis 
margine scariosis dorso glanduloso-puberulis, 4 mm. longis; petalis 
ad medium bifidis, lobis lineari-oblongis, 3 mm. longis; seminibus 
irregulariter tuberculatis. Differs from D. fasciculata in size and 
habit and placement of the bibracteate flowers, these, though fasci- 
cled, being borne one below the other. We associate doubtfully here 
the plant from Puno. Our species is probably D. leptoclados Hemsl. 
var. peruviana Ball, Journ. Linn. Soc. 22: 32. 1885, but as we have not 
seen Ball's specimen, we do not take up his name. The Peruvian 
species differs markedly from Hemsley's, of Mexico, in habit, size, 
fewer flowers, petals less divided, and capsule much shorter than calyx. 
Lima: Open, rocky slopes near Canta, 4,000 meters, Pennell 
14655, type in Herb. Field Museum. Puno: Rocky knoll in siliceous 
slope, 4,000 meters, Araranca, Pennell 13462(1). 

Drymaria divaricata HBK. Nov. Gen. & Sp. 6: 24. 1823. 

Stems glabrous, weak, filiform, branched, 20 cm. long or longer; 
leaves glabrous, broadly rounded-ovate, rounded at the base but 
decurrent to the petiole, this 6-8 mm. long, acute, 14-18 mm. long; 
peduncles irregularly branched, divaricate, many-flowered, filiform, 
glabrate, with lanceolate, persistent, opposite bracts; pedicels filiform, 
viscid-pubescent, in fruit 16-18 mm. long, divergent and deflexed; 
petals not clawed, deeply bifid, a little longer than the glabrous 
sepals, these oblong, acute, 3-nerved; capsule ovate, shorter than 
the calyx, about 5-seeded, the seeds brown-tuberculate. 

Lima: (Humboldt). Amancaes, Ruiz & Pavon (det. Mattfeld, 
ex descr.). 

Drymaria Engleriana (Muschler) Baehni & Macbr., comb. nov. 
Polycar pon Englerianum Muschler, Bot. Jahrb. 45: 452. 1911. 

Planta annua prostrata basi ramosissima, ramis subsimplicibus 
gracilibus paullo flexuosis 5-10 cm. longis; stipulis setaceis; foliis 
sessilibus late ovatis acutis membranaceis leviter pilosis, nervis subtus 
vix notatis, plerumque 4 mm. longis 2.5-3 mm. latis, superioribus 
gradatim reductis; floribus subsessilibus 2-4 in fasciculam dispositis; 
bracteis scariosis circa 3 mm. longis; sepalis oblongo-ovatis acutis 
valde carinatis, costa media prominente, 3 mm. longis; petalis pro- 
funde, usque ad medium, bifidis inferne abrupte constrictis 2-2.5 mm. 
longis; seminibus puncticulatis. 



622 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Lima: Prostrate on stony slopes, Viso, Macbride 500. Ancash: 
Huaraz, 4,300 meters, Weberbauer 310, type. 

Drymaria fasciculata Gray, U. S. Expl. Exped. 125. 1854. 
D. leptoclados Hemsl. var. peruviana Ball, Journ. Proc. Linn. Soc. 
22: 32. 1885(?). 

A nearly glabrous, erect, simple or little branched annual ; leaves 
only 3 or 4 pairs, sessile, ovate-oblong, acute, 1-2 cm. long, obscurely 
3-nerved; peduncle erect and strict, to 10 cm. long, minutely glandu- 
lar toward the apex; flowers subsessile, crowded in a dense fascicle, 
subtended by oblong-ovate, cuspidate, scarious bracts little shorter 
than the calyx; sepals oblong-lanceolate, cuspidate-acuminate, 6 mm. 
long, scarious-margined and conspicuously 3-nerved, subequaling 
the 5 oblong-linear, 2-lobed petals; capsule shorter than the calyx; 
seeds scabrous. D. stellarioides Willd. ex. R. & S. Syst. 5: 406. 1819, 
of Ecuador, has oblong-lanceolate leaves only 2 mm. wide but, 
apparently, would key here. 

Lima: Matucana, moist, rocky slope, 187 (det. Mattfeld). Rio 
Blanco, open, stony slope, 693 (det. Mattfeld). Obrajillo (Wilkes 
Exped.). Chicla (Ball, probably). 

Drymaria frutescens Mattfeld, sp. nov. in herb. 

Fruticosa ubique glabra ut videtur suberecta vel plus minusve 
diffusa circa 30 cm. alta, ramis flexuosis adscendentibus subtetragonis 
pauce ramosis fere nudatis griseo-brunneis, ramulis terminalibus 
congestis 6-10 cm. longis dense foliatis; foliis adpresso-imbricatis 
sessilibus rigidis conspicue trinerviis subulato-acuminatis vix 4 mm. 
longis; floribus subsessilibus solitariis fere 6 mm. longis; sepalis 
oblongo-lanceolatis apiculato-acuminatis 7 mm. longis; petalis bifidis 
basi cuneatis 4-5 mm. longis; seminibus ut videtur papillosis. 

Peru: Without locality, 1909-1914, Weberbauer 7203, type in 
Herb. Field Museum. 

Drymaria glaberrima Bartl. in Presl, Rel. Haenk. 2: 7. 1831. 

Glabrous, much branched from the base, 20-25 cm. high; leaves 
subsessile, narrowed to the base, acutish, cuspidate or the upper 
acuminate, more or less 3-nerved, broadly ovate; stipules minute, 
promptly caducous; flowers in short, few-flowered racemes, the 
common axillary peduncle greatly elongate; pedicels all shorter 
than the calyx, always erect, 3-4 in each corymb; bracts lanceolate, 
scarious; calyx 4 mm. long, the oblong, acute divisions white-trans- 
lucent, slightly longer than the semi-bifid petals; stamens 5, much 
shorter than the calyx, this nearly twice exceeding the capsule. 



FLORA OF PERU 623 

Huanuco: Huanuco, stony slopes, 3506. Montana (Haenke). 
Lima: Moist places, Huamantanga (Mathews, det. Planchon). 

Drymaria grandiflora Bartl. in Presl, Rel. Haenk. 2: 7. 1831. 

Basal branches sometimes nearly 30 cm. high, remotely branched, 
finely pubescent toward the tip; leaves very remote, broadly ovate, 
subsessile, rounded or subcordate at the base, acuminate, mucronate, 
5-nerved, sparsely pubescent or glabrate, to nearly 2 cm. broad, 
the upper much smaller; stipules scarious, nearly lanceolate, acumi- 
nate, caducous; flowers in axillary and terminal, dichotomous corymbs, 
5-7, the terminal corymbs long-peduncled ; pedicels always erect, 
many times longer than the calyx, this 6 mm. long, minutely pubes- 
cent, the oblong, acutish sepals scarious-margined; petals 5, 2-parted 
to the base, the divisions linear, slightly exceeding the calyx, this 
a little longer than the capsule. 

Huanuco: Montana (Haenke, type). Lima: Huaros, bushy, 
rocky slopes, Pennell 14724 (det. Mattfeld). 

Drymaria hirsuta Bartl. in Presl, Rel. Haenk. 2: 8. 1831. 

Many-stemmed, spreading, hirsutulous nearly throughout, the 
diffuse, procumbent branches to 15 cm. long; leaves reniform, 
rounded at the apex or rarely minutely cuspidate, broadly cordate 
at the base, the lowest shortly narrowed to the (6-8 mm. long) 
petiole, obsoletely nerved, 10-12 mm. broad, the much smaller upper 
ones subsessile; stipules obsolete, hair-like; flowers few, the filiform, 
glabrous common peduncle equaling the internodes; pedicels finally 
twice as long as the calyx, deflexed in fruit; calyx 2 mm. long, the 
ovate-oblong sepals obtusish; petals deeply bifid, very narrow; 
stamens 3-4; capsule a little longer than the calyx. Neg. 27717. 

Huanuco: Mito, 3266, 1543. Montana (Haenke). Puno: 
Sandia, Weberbauer 581. Ayacucho: Carrapa, Killip & Smith 
23212. Bolivia. 

Drymaria macrantha Gray, U. S. Expl. Exped. 126. 1854. 

Many-stemmed, laxly branched, slightly puberulent, especially 
above; leaves sessile, cordate-ovate, acute, 3-5-nerved, 10-14 mm. 
long; peduncle 2.5 cm. long or less; cymes rather close but even the 
lateral pedicels 6-12 mm. long; bracts narrowly subulate, greenish, 
with narrow, scarious margins below, 4-6 mm. long; sepals ovate- 
oblong, 8-10 mm. long, slightly 3-nerved, minutely glandular except 
the broad, hyaline margins; petals none; stamens 5; capsule shorter 



624 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

than the calyx; seeds scabrous. This is referred by Mattfeld (in 
herb.) to D. apetala. 

Lima: Obrajillo (Wilkes Exped., type). 

Drymaria nitida Ball, Journ. Proc. Linn. Soc. 22: 31. 1887. 

A small, glabrous annual, branching from the base; branches 
suberect, 8-20 cm. high; leaves broadly elliptic-ovate, acute or 
cuspidate, sessile; stipules minute, setaceous, fugacious; cymes 
terminal, dense, the flowers short-pediceled; bracts scarious, ovate; 
sepals broadly ovate, obtusish, purplish on the margin; petals 
narrowed to a claw, deeply parted; capsule subglobose, 8-seeded. 
Near D. divaricata HBK., but that has larger, laxer inflorescences, 
acute sepals, and less deeply parted petals (Ball). 

Lima: Chicla (Ball, type). Matucana, 188. Junin: Tarma, 
Kittip & Smith 21810. 

Drymaria ovata Willd. ex R. & S. Syst. 5: 406. 1819. 

Stems weak, procumbent, the branches puberulent or villous 
toward the tips; petioles 6 mm. long, villous; leaf blades ovate, 
acute, puberulent on both sides, 2.5-3.5 cm. long, 2-2.5 cm. wide; 
stipules persistent; peduncles bifid, few-flowered, the pubescent 
pedicels reflexed in fruit, 6-8 mm. long; calyx glabrous, the sepals 
oblong, acute, equaled by the deeply bifid petals. The determination 
of the following material from southern Peru is probably open to 
question, in view of the fact that the species appears to be unknown 
otherwise except from northern Peru and Ecuador. 

Arequipa: Pasco; Mejia (Gunther & Buchtien 295, 296, 297, det. 
Bruns). Lima: Above Supe, 3,000 meters (Weberbauer 169}. 
Without locality: Weberbauer 5327a, 6543. Ecuador to Venezuela. 

Drymaria pauciflora Bartl. in Presl, Rel. Haenk. 2: 8. 1831. 

Slender, sparsely branched, procumbent or diffuse, less than 
30 cm. high; leaves remote, cordate-suborbicular, sometimes nearly 
truncate at the base, often subsinuately margined, sparsely pubes- 
cent, lightly 3-5-nerved, 6-12 mm. broad; petiole slender, usually 
shorter than the leaf blade; stipules hyaline; flowers few, less than 
2 mm. long; peduncles binate or ternate, 1-2-flowered, glabrous, 
reflexed in fruit, with 2 tiny bracts at the middle; sepals glabrous, 
scarcely 2 mm. long, obtusish, nearly twice exceeded by the ovate 
capsule; petals 5, deeply bifid; stamens 2-3. 

Huanuco: Montana (Haenke, type). Huanuco: Cushi, rocky, 
partly sunny slopes, 4835 (det. Mattfeld). San Martin: San Roque, 
Williams 7621. 



FLORA OF PERU 625 

Drymaria praecox Baehni & Macbr., sp. nov. 

Herba annua stricta 3-5 cm. alta, caulibus solitariis vel interdum 
3-4 glabris; foliis radicalibus persistentibus dense rosulatis brevissime 
petiolatis obovato-spathulatis minute parceque hirsutulis 3-6 mm. 
lohgis; stipulis subulato-setaceis; foliis superioribus plerumque solum 
2 sessilibus ovatis apiculato-acutis glabris 4 mm. longis, 3 mm. latis; 
floribus circa 5 in cymulas sat laxas terminales dispositis; pedicellis 
minutissime glanduloso-puberulis erectis 2-4 mm. longis; sepalis 
oblongo-lanceolatis 3 mm. longis 3-nerviis extus obscure puberulis; 
petalis profunde, ultra medium, partitis, 3 mm. longis; stylo ad 
medium 3-partito. Scarcely a diminutive form of D. cordata, in 
view of the persistent rosettes of root leaves and the uniformly 
strict habit; probably a loma or desert plant. 

Peru: Without locality, Weberbauer 6916, type in Herb. Field 
Museum. 

Drymaria rotundifolia Gray, U. S. Expl. Exped. 123. 1854. 

Glabrous, erect and slender, 10 cm. or sometimes several dm. 
tall; leaves roundish, sessile, the larger about 6 mm. long; cymes 
corymbose, many-flowered, on peduncles 1-3 cm. long; pedicels 
slender, erect even in fruit, sometimes minutely glandular, the outer 
2-4 mm. long; bracts minute, entirely scarious; calyx 3 mm. long; 
sepals oval, very obtuse, faintly 3-nerved, narrowly scarious-mar- 
gined; petals about a third longer than the calyx, narrowly cuneiform, 
cleft to the middle; capsule ovoid, a little longer than the calyx; 
seeds muricate-tuberculate. 

Lima: Open, rocky slopes, San Buenaventura, Pennell 14516. 
Above Obrajillo, Pennell 14390. Obrajillo (Wilkes Exped., type). 
Open hillside, Killip & Smith 21761. Matucana, disintegrated 
granite, 261; wet, rocky slope, 183. 

Drymaria sphagnophila Baehni & Macbr., sp. nov. 

Planta flaccida diffusa plus minusve pilosa, caulibus ut videtur 
subprostratis remote ramosis sat foliosis glabris 5-10 cm. longis; 
stipulis capillaceo-setaceis minutis; foliis sessilibus subtus plus 
minusve pilosis, pilis caducis, fere rotundatis 4-6 mm. longis latisque 
haud vel vix ad apicem angustatis floribus circa 4 racemoso-cymosis; 
pedunculis glabris filiformibus circa 1 cm. longis; pedicellis circa 
2 mm. longis; floribus fere globosis; sepalis rotundato-ellipticis 
obtusis 2 mm. longis; petalis ultra medium bifidis; lobis acutis, 
2-2.5 mm. longis, basi abrupte constrictis longe unguiculatis; semini- 
bus tuberculatis lenticularibus. 



626 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Huanuco: Sunny sphagnum bog, 3,000 meters, Mito, Macbride 
1542, type in Herb. Field Museum. 

Drymaria stereophylla Mattfeld, sp. nov. in herb. 

D. auriculipetalae similis; planta interdum plus minusve caespitosa 
et ubique plus minusve minute pulverulenta vel granulato-glan- 
dulosa; stipulis interdum obsoletis; foliis ovatis apiculatis 6-8 mm. 
longis, 3-5 mm. latis, nervis costa media excepta utrinque vix 
notatis vel subtus paullo prominentibus; sepalis valde scarioso- 
marginatis oblongo-ellipticis cuspidatis, 6-7 mm. longis; petalis 
basi subauriculatis, apice bifidis, 8 mm. longis; seminibus fere 
laevibus. 

Junin: In crevices of limestone cliffs, 4,000 meters, La Oroya, 
Macbride 962, type in Herb. Field Mus. Rocky hillside, Huan- 
cayo, 3,300 meters, Kittip & Smith 22134, 22132; Weberbauer 6600. 
Huanuco: In mats on shaded canyon ledges, Chasqui, 1761. 
Lima: In clumps, loose, steep canyon sides, Viso, 563. 

Drymaria viscidula Gray, U. S. Expl. Exped. 124. 1854. 

Stems diffuse, to 30 cm. long, glabrous except at the tip; branch- 
lets and inflorescence glandular-pubescent; leaves subreniform-ovate, 
6-8 mm. long, 8-10 mm. broad, abruptly contracted to a somewhat 
hairy petiole; stipules capillary-setaceous; peduncles to 2.5 cm. long, 
glandular-glutinous; cyme 5-15-flowered, the bracts setaceous-sub- 
ulate; lateral pedicels 2-4 mm. long, erect even in fruit; calyx 3 mm. 
long, the oblong, acute sepals about equaling the narrowly cuneate, 
2-cleft petals and capsule; seeds obscurely tuberculate. 

Lima: Obrajillo (Wilkes Exped., type). Open, rocky slopes, San 
Buenaventura, Pennell 14518. 

Drymaria Weberbaueri Muschl. Bot. Jahrb. 45: 451. 1911. 

Branching at the base and sparsely above, about 15 cm. high, 
glabrous or most obscurely pruinose-glandular; leaves ovate-orbi- 
cular, slightly cordate or truncate at the sessile base; peduncles 1- 
several cm. long, bearing 1-3 head-like fascicles of several more or 
less spreading or nodding flowers, the pedicels 2-3 mm. long; sepals 
acutish, 4 mm. long, the somewhat longer petals more or less con- 
spicuous. All the determinations are by Mattfeld. 

Lima: Barranca in the loma (Weberbauer 1662, 1657, types). 
Loose, stony, upper slopes of seaside hills, Chorrillos, 5863. Rocky 
hillside, San Geronimo, 5904. Sandy lomas along the sea, Lurin, 
5963. Open, sandy slopes, Atocongo, Pennell 14783. 



FLORA OF PERU 627 

10. POLYCARPON L. 

Small herbs with scarious stipules and bracts, the tiny flowers 
in crowded, terminal or partly axillary cymes. Petals minute. 
Stamens 3-5. Styles short. Loefling wrote the name Polycarpa. 

Polycarpon apurense HBK. Nov. Gen. & Sp. 6: 33 (40). 1823. 

Stems many, more or less matted and branched or diffuse; leaves 
verticillate, the upper opposite, linear-spatulate, obtusish, narrowed 
to the base, ciliate, 12 mm. long, half as long as the internodes; 
stipules ovate, acuminate, pubescent; cymes dichotomous, axillary 
and terminal; flowers pedicellate, the bractlets similar to the stipules; 
calyx glabrous, persistent, the lanceolate, obtuse lobes longer than 
the petals; styles 3; capsule oblong, acute; seeds many, oblong, 
brown, scabrous. Description from HBK. Included by Rohrbach 
in P. depressum (L.) Rohrb., an untenable name, as pointed out by 
Harms, Pflanzenfam. ed. 2. 16c: 308, because of P. depressum Nutt. 
of California (and the Andes?), a glabrous plant; the synonomy is 
involved. The nearly cosmopolitan P. tetraphyllum L. is quite 
glabrous, with a leafless inflorescence, has obovate leaves only 5-10 
mm. long and oblong, carinate sepals 2 mm. long. 

Peru: (Probably). South America (Pax & Hoffmann). 

Polycarpon Englerianum Muschl. Bot. Jahrb. 45: 452. 1911. 

A densely cespitose annual or biennial with numerous spreading- 
ascending, much branched stems 2-4 cm. long; leaves obovate or 
obovate-lanceolate, obtuse or acutish, gradually narrowed to a short 
petiole, more or less densely crisp-pubescent, 5-6 mm. long, 3-3.5 
mm. broad; stipules scarious, triangular-lanceolate, 1 mm. long, 
minutely and sparsely puberulent; flowers in dense clusters, mostly 
subsessile; bracts scarious, ovate, acuminate; calyx lobes oblong, 
obtusish, prominently carinate, membranous-margined, more or 
less densely puberulent, the outer 2 mm., the inner 3 mm. long; 
petals 5 or fewer or none, lanceolate, sometimes emarginate, about 
half as long as the calyx; stamens 3 or rarely 5; capsule about as long 
as the calyx, the minute seeds reddish. Probably only a densely 
cespitose form of P. apurense. Neg. 27705. 

Ancash: Huaraz, 4,300 meters (Weberbauer 3101, type). 

Polycarpon Urbanianum Muschl. Bot. Jahrb. 45: 452. 1911. 

A densely cespitose, much branched annual, the stems 2-4 cm. 
long; leaves ovate or ovate-lanceolate, usually acute, narrowed to a 
rather long petiole, more or less densely puberulent, 6-8 mm. long, 



628 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

4-5 mm. broad; stipules 1 mm. long; pedicels, if present, short and 
densely puberulent; outer calyx lobes 1 mm., the inner nearly 2 mm. 
long; otherwise like P. Englerianum, but much less branched. 
Ancash: Huaraz, 4,300 meters (Weberbauer 2985, type). 

11. POLYCARPAEA Lam. 

Erect herbs with mostly rosulate leaves and more or less crowded, 
silvery, bracted inflorescences of tiny flowers. Stamens 5, united in 
a ring with the petals or staminodia. Styles elongate. Microphyes 
Phil, may occur on the coast of southern Peru; it is a tiny, branched 
herb, distinguished by styles not longer than the ovary. 

Polycarpaea corymbosa (L.) Lam. 111. 2: 129. 1791. Achy- 
ranthes corymbosa L. Sp. PI. 205. 1753. P. filifolia Muschl. Bot. 
Jahrb. 45:453. 1911. 

A slender annual with mostly rosulate, filiform leaves, the stem 
closely branched at the summit and bearing a lustrous, scarious- 
bracted cyme; calyx lobes scarious, white, 2.5-3.5 mm. long, the 
petals yellowish in age, the capsule scarcely half as long; seeds 5-6, 
lightly grooved on the back. 

San Martin: Moyobamba, 1,000 meters, Weberbauer 4587, type of 
P. filifolia. South America. 

12. SPERGULARIA (Pers.) J. & C. Presl 
Reference: Rohrbach, Linnaea 37: 219-244. 1872. 
Annual, perennial, or sometimes even suffrutescent plants with 
narrow, opposite leaves and more or less prominent stipules, the 
flowers in cymes or short racemes. Sepals characteristically thin 
at the edges. Styles commonly 3. Spergula arvensis L. may occur 
as an introduction from Europe; it usually has 5 styles, flowers borne 
in terminal, cymose panicles, and crowded, apparently whorled 
leaves. S. squarrosa Muschl. is scarcely a Spergularia and is omitted 
from the key; see note after the description. Persoon established 
the name as a section of Arenaria; a few authors have united the 
group with Spergula L., and there is no reason except the con- 
venience of tradition against their action. The name has been con- 
served in place of Tissa Adans. 

Seeds winged or rarely a few wingless; stamens mostly 10. 
Seeds tuberculate; stipules connate for one- third their length. 

S. villosa. 



FLORA OF PERU 629 

Seeds smooth. 

Stipules connate only at the base S. grandis. 

Stipules connate about half their length S. media. 

Seeds mostly wingless; stamens 5 or fewer. 

Plants annual or short-lived perennials S. salina. 

Plants perennial S. macrotheca. 

Seeds all wingless. 

Plants diminutive, the stems only 2-5 cm. long; perennial. 

S. andina. 
Plants usually about 10 cm. tall. 

Leaves imbricate; stipules finely lacerate S. congestifolia. 

Leaves, at least the lower, not imbricate; stipules if lacerate, 
only at the tip, or rarely fimbriate half their length. 

Stems more or less glandular-villous or glandular-puberulent. 
Glandular-puberulent perennials or biennials. 
Calyx 6-7 mm. long; branches ascending. 

Lowest internodes much exceeding the leaves; stipules 
lacerate above S. Stuebelii. 

Lowest, like the upper, internodes slightly shorter 
than the leaves; stipules all fimbriate one-half 
their length S. laciniata. 

Calyx 3-4.5 mm. long; branches usually prostrate. 

S. campestris. 

Glandular-villous annual, erect with ascending branches. 

S. collina. 
Stems glabrate, with sessile or short-stipitate glands. 

S. stenocarpa. 

Spergularia andina Rohrb. Linnaea 37: 234. 1872. 

Stems prostrate or ascending, 2-5 cm. long, densely crisp-pubes- 
cent; leaves short-mucronulate, glabrescent, 10-15 mm. long, 0.5 
mm. broad; stipules ovate-lanceolate, long-acuminate, 3-4 times 
longer than broad, 5-6.5 mm. long, connate below; flowers few, 
terminal and solitary; pedicels densely white-hirsute; sepals ovate- 
lanceolate, obtusish, broadly scarious-margined, often reddish, more 
or less hirsute, 4-5 mm. long, scarcely exceeding the broadly ovate, 
obtuse, suberose petals; capsule scarcely exserted. Neg. 27713. 

Puno: Azangaro, Lechler 1772. Bolivia. 



630 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Spergularia campestris (L.) Aschers. Fl. Mark. Brandenb. 
194. 1864. Arenaria rubra var. campestris L. Sp. PI. 423. 1753. 

Biennial or sometimes a short-lived perennial, the stems nearly 
always prostrate, to 15 cm. long, glabrescent below, glandular- 
puberulent above; leaves linear-filiform, mucronate, glabrate, 1-2 
cm. long; stipules ovate-lanceolate, attenuate, acuminate, 2 or 3 
times longer than broad, short-connate; inflorescences scorpioid in 
fruit, racemose, the pedicels mostly shorter than the calyces or some- 
times apparently much exceeding them; sepals 3.5-4.5 mm. long, 
nearly equaled by the petals and capsule; seeds brown, tuberculate. 
The correctness of the name is not certain. 

Peru: (Probably). Cosmopolitan. 

Spergularia collina I. M. Johnston, Contr. Gray Herb. 81: 
89. 1928. 

A loosely and dichotomously branched annual, 5-10 cm. high, 
abundantly but shortly glandular- villous; leaves spreading, linear, 
1-2 cm. long; stipules ovate, acuminate, often more or less lacerate, 
connate basally, 1 mm. long; flowers axillary, on slender, spreading 
or reflexed pedicels as long as or longer than the leaves; sepals nearly 
oblong, 4-5 mm. long, the white petals slightly shorter; stamens 10; 
capsule ovoid, included; seeds black, opaque, all wingless, com- 
pressed, tuberculate or glandular-mucronulate. Illustrated, Weber- 
bauer, 144 as Drymaria molluginea. 

Arequipa: Sandy, open places behind Mollendo, Johnston 3568 
(type). 

Spergularia congestifolia I. M. Johnston, Contr. Gray Herb. 
81:90. 1928. 

Perennial, with many branching, prostrate stems 2-8 cm. long; 
peduncles ascending, glandular-villous, 3-8 cm. long; leaves strongly 
congested, linear-setaceous, mucronulate, falcate, hispidulous, some- 
what glandular, 3-9 mm. long, longer than the internodes; stipules 
black, about equaling the leaves, deeply and finely laciniate, basally 
connate; flowers 3-12, laxly cymose; pedicels 5-12 mm. long; sepals 
lance-oblong, shortly glandular-villous, 4-6 mm. long, scarious- 
margined; petals white, slightly longer than the sepals; stamens 10; 
capsule about 5 mm. long, somewhat exceeding the calyx. 

Arequipa: Prostrate on open places in the fertile belt behind 
Mollendo, Johnston 3567 (type). Mollendo, Weberbauer 1503. 



FLORA OF PERU 631 

Spergularia grandis (Pers.) Camb. in St. Hil. Fl. Bras. 2: 177. 
1829; 236. Spergula grandis Pers. Syn. 1: 522. 1805. Spergularia 
macrocarpa Presl, Rel. Haenk. 2: 9. 1831. 

Stems many from a thick, woody rhizome, erect, simple or 
sparsely branched, to several dm. high, glandular-puberulent above; 
leaves narrowly linear, acute, 2.5-4.5 cm. long; stipules long-acumin- 
ate, scarcely connate, 4-5 mm. long; pedicels slender, many times 
longer than the calyx, reflexed in fruit; sepals shortly glandular- 
pubescent, 5-6 mm. long, half as broad, about equaled by the pur- 
plish-tipped petals; styles 3-5; capsule ovate-oblong, sometimes 
well exserted; seeds reddish brown, broadly winged. 

Huanuco: Montana (Haerike). Without locality, Ruiz & Pavon 
(det. Mattfeld). Southern South America. "Romerillo." 

Spergularia laciniata Baehni & Macbr., sp. nov. 

Ut videtur perennis, caulibus prostratis vel plus minusve adscen- 
dentibus mediocriter ramosis plerumque 10-15 cm. longis e basi ad 
apicem satis dense cum pilis minutis patentibus pubescentibus haud 
vel solum superne paullo glandulosis; foliis undique quam internodiis 
plerumque paullo longioribus patentibus glabris linearibus acuto- 
apiculatis 10-15 mm. longis, interdum 1.5 mm. latis; stipulis 6-10 
mm. longis ad medium connatis superne multo-fimbriatis; floribus 
circa 6 in cymam laxam dispositis; pedicellis 6-10 mm. longis minute 
glanduloso-pilosis; sepalis oblongis scarioso-marginatis glandulosis 
circa 6 (-7.5) mm. longis; staminibus 10; petalis albis circa 7.5 mm. 
longis integris; seminibus ignotis. Differs from S. Stuebelii as 
indicated in the key, also in the exserted petals; from S. congestifolia, 
its not at all imbricate leaves and less deeply laciniate stipules 
distinguish it readily. 

lea: Loma near the sea, 500 meters, Bahia de la Independencia, 
Weberbauer 7958, type in Herb. Field Museum. 

Spergularia macrotheca (Hornem.) Heynh. Nom. Bot. 689. 
1840. Arenaria macrotheca Hornem. ex Cham. & Schlecht. Linnaea 
1: 53. 1826. 

A sturdy, half erect perennial, sometimes apparently flowering 
as a biennial, viscid-pubescent throughout; leaves 1.5-3.5 cm. long; 
stipules to 6 mm. long; flowers becoming racemose, white or pink, 
on pedicels 5-15 mm. long; sepals 5-8 mm. long; stamens 5; seeds 
wingless except those at the base of the capsule, this sometimes 
slightly exserted. 

Peru: (Probably). Widely distributed; Chile. 



632 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Spergularia media (L.) Griseb. Spic. Fl. Rumel. 1: 213. 1843. 
Arenaria media L. Sp. PI. ed. 2. 606. 1762. 

Perennial, more or less glandular-pubescent, especially above; 
stems prostrate or ascending, rather strongly nodose, sometimes 
several dm. long; leaves linear-filiform or subulate, rarely mucronu- 
late, 1-3 mm. long; flowers few to many, the slender pedicels often 
elongate and spreading or reflexed ; calyx 4-4.5 or exceptionally 6 mm. 
long, equaled by the white, purple-tipped petals; capsule included or 
distinctly exserted; seeds smooth. This plant is called S. marginata 
(DC.) Kittel in Taschenb. Fl. Deutschl. ed. 2. 1003. 1844 by Pax and 
Hoffmann, perhaps rightly, the name media being abandoned as "a 
source of confusion" by some authors, since it is Arenaria media L. 
Sp. PI. ed. 2. 606. 1762 only in part. 

Ancash: Ocros, 3,000 meters, Weberbauer 2751(1}. Nearly 
cosmopolitan. 

Spergularia salina J. & C. Presl, Fl. Cech. 95. 1819. 

In general similar to S. macrotheca, but an annual or a short-lived 
perennial; leaves 1-5 cm. long; pedicels often little longer than the 
calyx, this 3-4 mm. long; petals reddish, about as long as the sepals, 
the capsule slightly to well exserted. S. marina (L.) Griseb. Spic. 
Fl. Rumel. 1: 213. 1843, is apparently the same, based on Arenaria 
rubra var. marina L. Sp. PI. 423. 1753; S. marina Pallas, Reise 3: 
603. 1776. 

Peru: (Probably). Widely distributed. 

Spergularia stenocarpa (Phil.) I. M. Johnston, Contr. Gray 
Herb. 85: 41. 1929. Arenaria stenocarpa Phil. Fl. Atac. 10. 1860; 
Viaje Des. Atac. 19, 184. 1860. 

Flowering as an annual but sometimes more enduring; leaves 
narrowly linear, mucronulate, to 16 mm. long; stipules long-acumi- 
nate, 5 mm. long; flowers numerous, in dichotomous cymes, the 
oblong-lanceolate sepals exceeding the white petals; capsule cylindric, 
5 mm. long, exserted; seeds black, lustrous. Johnston reports it as 
usually an annual but sometimes fruticulose, the yellowish green 
stems glabrate except for small, sessile or short-stipitate glands. 
The Peruvian collections were determined by Bruns. 

Arequipa: Cachendo, 1,000 meters (Gunther & Buchtien 291, 
293a). Mollendo, 50 meters, Gunther & Buchtien 291b, 292. Mejia, 
40 meters (Qunther & Buchtien 293). Chile. 

Spergularia squarrosa Muschl. Bot. Jahrb. 45: 461. 1911. 



FLORA OF PERU 633 

Stems many, much branched, especially at the base, erect, 
30-50 cm. high, strongly enlarged at the reddish nodes, becoming 
glabrous; leaves usually much shorter than the internodes, ovate- 
lanceolate or the upper linear, mucronulate-aristate, glabrescent 
or rarely densely puberulent, 1.5-2 cm. long, 5-10 mm. wide or the 
uppermost half as large; stipules minute, lanceolate, connate one- 
third their length, long-acuminate and often multilaciniate, 5-6 mm. 
long; flowers many, terminal, nearly capitate, on pubescent pedicels; 
sepals lance-oblong, obtusish, broadly scarious-margined, densely 
hirsutulous, 3.5-4 mm. long; petals ovate-oblong, obtusish, a little 
shorter than the calyx; capsule short-stipitate, sometimes slightly 
exserted; seeds brown, rotund-obovate, tuberculate. This is prob- 
ably a species of Drymaria if, indeed, it is not D. glaberimma BartL; 
specimen seen, sterile. Neg. 30174. 

Lima: Near Matucana, in rocks, 2,570 meters (Weberbauer 57, 
type). 

Spergularia Stuebelii (Hieron.) I. M. Johnston, Contr. Gray 
Herb. 81 : 90. 1918. Tissa Stuebelii Hieron. Bot. Jahrb. 21 : 308. 1895. 

Stems numerous, erect, probably to 20 cm. high, below glabrate, 
above shortly glandular-puberulent throughout; leaves to 1 cm. long, 
scarcely 0.5 mm. broad, mucronate; lower internodes much longer 
than the leaves; stipules ovate, basally connate, acuminate, about 
5 mm. long, finally mostly lacerate at the tip; flowers few, pedicellate; 
calyx lobes narrowly lanceolate, broadly hyaline-margined, glan- 
dular, 6-7 mm. long, about 1.5 mm. wide; petals white, ovate, 
acute, 5 mm. long, the capsule slightly shorter; seeds minutely 
papillose-puberulent. 

Arequipa: Base of Mt. Misti (Stuebel 79). Arequipa, 2,500-2,900 
meters, gravelly stream beds and open, rocky slopes, Pennell 13247, 
13156. Southern slopes of Mt. Chachani, 3,050 meters, Hinkley 10. 
"Estrellita del cerro." 

Spergularia villosa (Pers.) Camb. in St. Hil. Fl. Bras. 2: 178. 
1829; 237. Spergula villosa Pers. Syn. 1: 522. 1805. 

Similar to S. grandis, but often more glandular-pubescent; 
stipules connate one-fourth to one-third their length, usually entire, 
5-8 mm. long; pedicels erect or spreading; capsule slightly exserted; 
seeds yellow, the wing sometimes lacerate, tuberculate. S. laevis 
Camb., allied but the petals 3-4 times shorter than the glabrous 
calyx, may be expected. Illustrated, Mart. Fl. Bras. 14, pt. 2: pi. 61. 

Peru: (Probably). Southern South America. 



634 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

13. SILENE L. 

Melandrium Roehl. 

Reference: Rohrb. Linnaea 36: 196-258. 1869. 

Annuals or perennials, the latter often with somewhat creeping- 
ascending rootstocks bearing rosettes of spatulate-lanceolate or 
somewhat ovate leaves from which arises one sparsely leafy, brac- 
teate or naked stem (or this sometimes apparently lacking) with 
solitary or few (generally) erect or often nodding flowers, or the 
inflorescence well developed, but not in the Peruvian species. Calyx 
10-many-nerved, often inflated. Styles 3-5. Ovary if not 1-celled 
more or less 3-5-celled toward the base, or the capsule not clearly 
more than 1-celled. Capsule valves usually twice as many as the 
styles. The inclusion of Melandrium (see Harms, Pflanzenfam. ed. 
2. 16c: 341 for a discussion of the spelling) in Silene as a section 
does not preclude, for those who wish, the grouping together of the 
species with partially more than 1-celled ovary without the incon- 
venience of two group names for plants that otherwise, both as 
regards aspect and technical characters, are identical. The present 
nomenclatorial confusion (cf. S. thysanodes, for example) is thus 
avoided, and one definite group name is available for those whose 
interest in the plants extends no further. Logically Lychnis L. 
could be united, for reasons of tradition, together with the certain 
aspect many species have, but it may conveniently be kept distinct 
even as Stellaria, Cerastium, and Arenaria are arbitrarily maintained. 
Pax and Hoffmann, op. cit. 345, quote Fries' remark made even in 
1843 as to the difficulty of separating Melandrium, Viscaria, and 
Heliosperma; there is obviously one natural genus made difficult 
to recognize by adopting sectional characters as generic. Besides 
the following species, the Index Kewensis records an S. Aucheriana 
Boiss. for "Peru," evidently a misprint for Persia. 

Plants annual. 

Flowers paniculate; stems usually glandular-banded at the 
internodes S. antirrhina. 

Flowers racemose; stems not glandular-banded S. gallica. 

Plants perennial. 

Cespitose plants, the flowers sessile in the rosettes of leaves. 

Melandrium Mandonii. 

Stems well developed, even in flower, sometimes scapose. 
Pubescence even of the calyx obscurely if at all glandular. 



FLORA OF PERU 635 

Seeds with a narrow, inflated margin; petals obtusely auricled, 

exserted S. andicola. 

Seeds emarginate; petals usually acutely auricled. 

Melandrium chilense. 
Pubescence definitely glandular, especially below the calyx. 

S. thysanodes. 

Silene andicola Gill, in Hook. Bot. Misc. 3: 147. 1833. Melan- 
drium andicolum Rohrb. Linnaea 36: 221. 1869. Lychnis andicola 
Britton, Bull. Torrey Club 16: 61. 1889. 

Plants with a creeping, woody caudex, producing leafy rosettes 
and sterile, erect shoots; lower leaves rosulate, linear-lanceolate, 
acuminate, glabrous except the ciliate margins, the cauline leaves 
nearly linear; flowers mostly erect, solitary or few, short-pedicellate; 
calyx ovate-turbinate, inflated, densely pubescent, the nerves bi- 
parted at the base; calyx teeth one- third to one-half as long as the 
tube, lanceolate, acutish, densely lanuginose-ciliate; petals white, 
obtusely auricled, the blade sometimes bifid, with 2 minute appen- 
dages; capsule ovate-oblong; seeds narrowly margined. Neg. 30181. 

Junin: Huaron, rocky slopes, 1118; wet rock ledges, 1146. Casa- 
palca, loose soil of alpine basin slopes, 5,000 meters, 834- La Qiiinua, 
Poeppig. Bolivia; Chile. 

Silene antirrhina L. Sp. PI. 419. 1753. 

Similar to S. gallica, but usually simple and generally well 
marked by a dark, glutinous band in each internode. 
Peru: (Probably). Cosmopolitan. 

Melandrium chilense (C. Gay) Rohrb. Linnaea 36: 222. 1869. 
Lychnis chilensis C. Gay, Fl. Chil. 1: 256. 1845. 

Stems simple, erect, somewhat hirsute, from a thick, ascending 
caudex; leaves lanceolate, linear, acute or obtusish, attenuate at the 
base, puberulent and densely ciliate, the few cauline leaves ovate- 
acuminate; flowers erect or somewhat nodding, solitary or with 1 or 
2 lateral ones; calyx little inflated, ovoid, densely pubescent; petals 
white or violet, the blade bifid; seeds tuberculate. Var. cucubaloides 
(Fenzl) Hoss. Trab. Inst. Bot. Farm. Buenos Aires 33. 1915 (M. 
cucubaloides Fenzl ex Rohrb. op. cit. 223), is distinguished principally 
by the obtuse auricles. 

Ancash: Hacienda Cajabamba, 3,000 meters, Weberbauer 3169 
(det. Muschler as M. cucubaloides}. Samanco, 3,700 meters (Weber- 
bauer 171}. Junin: Tarma, 4,000 meters, Weberbauer. Chichla, 



636 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

3,720 meters, Weberbauer 243 (det. Muschler as M. cucubaloides). 
Near Huancayo, 4,700 meters, Killip & Smith 22082. Huanuco: 
Chasqui, moist cliff edges, 3290. Lima: Rio Blanco, steep, grassy 
slopes, 666, 2961 ; Killip & Smith 21610. Chile. 

Silene gallica L. Sp. PI. 417. 1753. 

An erect, villous-viscid annual, generally much branched, 10-20 
cm. tall; leaves spatulate, obtuse, mucronate, 1-3 cm. long; flowers 
in one-sided racemes; calyx ovoid, 7-9 mm. long, slightly exceeded 
by the white or pinkish petals; capsule included. All determinations 
by Wolff. 

Lima: Matucana, 3,200 meters, 316, 330; Weberbauer 5728; 91. 
Tambo de Viso, Weberbauer 136. Rio Blanco, 699. Barranco, 300 
meters, Weberbauer 1651. Lima, Ruiz & Pawn (det. Mattfeld). 
Amazonas: Chachapoyas 2,700 meters, Williams 7544- Huanuco: 
Huanuco, 2093. Cajamarca: At 2,200 meters, Weberbauer 3861. 
Introduced from Europe. 

Melandrium macrocalyx Muschl. Bot. Jahrb. 45: 441. 1911. 

Perennial, with a solitary, nearly or quite erect, densely long- 
pubescent stem about 30 cm. high, branching above; basal rosulate 
leaves lanceolate, narrowed to a petiolar base, acute, 3.5 cm. long, 
0.5-1 cm. broad, green, on the margin and midnerve densely pubes- 
cent; flowers cymose, on pedicels 3^4.5 cm. long, these more or less 
curved below the calyx, densely brown-pubescent; calyx broadly 
tubular-campanulate, the evident nerves pubescent like the stems, 
the teeth triangular; petals 2-parted, obliquely truncate, the broad 
lobes approximate, purplish; capsule ovoid, included in the ventri- 
cose calyx; seeds globose-reniform, black-tuberculate. 

Cajamarca: Prov. Hualgayoc, 3,100 meters (Weberbauer 4015, 
type). 

Melandrium Mandonii Rohrb. Linnaea 36: 222. 1869. 

Densely cespitose from a woody caudex, the flowers sessile in the 
rosulate leaves, these linear-lanceolate, obtusish, membranous! y 
connate at the base, densely scabrous-ciliate, otherwise glabrous; 
calyx short-campanulate, glabrous, the teeth one-third to one-half 
as long as the tube, lanceolate, recurving, densely ciliate; petals 
white, obtusely auricled, the appendages minute; capsule ovoid- 
campanulate; seeds smooth, broadly margined. M. Rimbachii 
Mattf. Notizbl. Bot. Gart. Berlin 10: 115. 1927, has a densely tomen- 
tose calyx, the short stems lengthening in fruit to even 10 cm. tall. 



FLORA OF PERU 637 

Doubtfully more than the alpine, reduced state of S. andicola. 
Neg. 27700. 

Lima: Stony, grassy hillsides, 5,000 meters, 2980. Bolivia. 

Melandrium rhizophorum Muschl. Bot. Jahrb. 45: 452. 1911. 

Perennial from very tortuous rhizomes to 6 cm. long and 5 mm. 
thick, bearing a caudex clothed with the remains of the rosulate 
basal leaves from which arises an erect or suberect, short-pubescent 
stem; leaves ovate-lanceolate, gradually narrowed to a petiole 5-10 
mm. long, somewhat acute, glabrous beneath, more or less densely 
scaly-puberulent above, the margins densely ciliate, yellowish, some- 
what fleshy, 2-3 cm. long, 1-2 cm. broad; pedicels 2-2.5 cm. long; 
petals yellowish purple; seeds globose; otherwise like M. macrocalyx 
Muschl. Differs from M. thysanodes in that the pedicels and calyx 
are not glandular (Hieronymus in Herb. Berlin). 

Lima: At 4,500 meters, above Lima (Weberbauer 5098). 

Silene thysanodes Fenzl in Endl. Stirp. Nov. Dec. 4: 31. 1839. 
Melandrium thysanodes Gray, U. S. Expl. Exped. 114. 1854. Lychnis 
thysanodes Hook. f. Fl. Ant. 246. 1844-47. 

A cespitose perennial with ascending, 1-few-flowered stems some- 
times 15 cm. high, glandular-pubescent, especially above; leaves 
lanceolate or the lower spatulate, ciliate-tomentose and glandular as 
also the peduncle and calyx, the last broadly campanulate, 8 mm. 
long, 5-parted to the middle, the ovate, obtuse lobes very tomentose- 
ciliate; petals about a third longer than the calyx, purplish, with 2 
minute appendages or these obsolete; stamens 10, woolly at the very 
base, inserted in a fleshy disk, free from the 1-celled ovary; styles 
5; capsule about equaling the calyx, 5-valved, the valves 2-cleft; 
seeds smooth, with a wide and thickish or inflated wing. The 
description is by Gray, based on the Wilkes material; the type from 
Ecuador is described as having 3 styles and partly 3-celled ovary, 
thus it must be a true Silene and probably the Peruvian plant of Gray 
represents another species, perhaps one of those proposed by Muschler. 

Junin: Above Bafios and Alpamarca (Wilkes Exped.). Edge of 
grassy bank, 2007 (apparently the species as interpreted by Gray). 
Chinche, 4,000 meters, bunchgrass slope, 1259 (like No. 2007). 
Colombia. 

Melandrium Weberbaueri Muschl. Bot. Jahrb. 45: 442. 1911. 

Perennial from a stout taproot; caudex much branched, clothed 

with leaf remains, glabrous, with internodes 2-20 mm. long; leaves 



638 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

densely rosulate, linear or rarely lanceolate, with the rather long 
petiole 3-5 cm. long, 5-7 mm. broad; bractlets minute; pedicels 1 cm. 
long, mostly glabrous; petals yellowish white; otherwise like M. 
macrocalyx. 

Junin: Near Yauli, 4,400 meters (Weberbauer S37, type). 

68. NYMPHAEACEAE. Water-lily Family 
By Paul C. Standley 

Reference: Caspary in Mart. Fl. Bras. 4, pt. 2: 130-184. 1878. 

Perennial, aquatic plants, acaulescent or with elongate stems, 
the leaves floating or submerged; flowers perfect, terminating 
elongate scapes or solitary in the leaf axils; sepals 3-6; petals 3 to 
many, often gradually passing into stamens or staminodia; stamens 
few or numerous; ovary of few or numerous free or united carpels, 
the stigmas sessile or united to form a disk with radiating stigmatic 
lines; fruit indehiscent, of distinct or united carpels. Caspary, op. 
cit., gives a questionable report of the genus Victoria from the 
Rio Ucayali; see also Rev. Hort. 405. 1866. 

Plants acaulescent; leaves floating, deeply cordate at the base, entire 
or nearly so I. Nymphaea. 

Plants with elongate stems; leaves mostly submerged and dissected 
into capillary segments 2. Cabomba. 

1. NYMPHAEA L. Water-lily 

Reference: Conard, The Waterlilies: A monograph of the genus 
Nymphaea. Carnegie Inst. Wash. Publ. 4. 1905. 

Coarse, perennial herbs with thick, fleshy, horizontal rootstocks; 
leaves floating, large, deeply cordate at the base, entire or somewhat 
undulate or coarsely dentate; flowers large and showy. It is probable 
that other species of the genus may be found in eastern Peru. 
Leaves thick, red-purple beneath, usually undulate or somewhat 

dentate; flowers open during the day N. ampla. 

Leaves thin, green beneath, entire; flowers open only at night. 

A T . blanda. 

Nymphaea ampla (Salisb.) DC. Syst. 2: 54. 1821. Castalia 
ampla Salisb. Parad. Lond. 1: pi. 14- 1805. 

Leaves 15-46 cm. in diameter, coarsely sinuate-dentate or almost 
entire; flowers white, raised above the water, 8-16 cm. broad, very 
showy; sepals oblong-lanceolate, obtuse or acutish, the outer surface 
green with purple-black lines; petals 12-21. 



FLORA OF PERU 639 

Loreto: Balsapuerto, open swamp, 150-350 meters, Killip & 
Smith 28682. Widely dispersed in tropical America, extending 
northward to Texas and the West Indies. Probably common in 
eastern Peru. 

Nymphaea blanda Mey. Prim. Fl. Esseq. 201. 1818. 

Plants smaller and more slender than those of the preceding 
species; leaves mostly 10-15 cm. long, broadly rounded at the 
apex, thin, green on both sides; flowers creamy white, 8-9.5 cm. 
broad, the sepals 3.5-4.5 cm. long, green; petals 16. Illustrated, 
Mart. Fl. Bras. 4, pt. 2: pi. 36, f. 9-16. 

Loreto: Iquitos, in pool, 100 meters, Killip & Smith 27173. 
Ranging northward to Central America. 

2. CABOMBA Aubl. 

Slender, mostly submerged plants; leaves of two kinds, the sub- 
merged opposite or verticillate, palmately dissected into numerous 
capillary segments, the floating leaves (usually absent) few, alternate, 
centrally peltate; sepals and petals each 3; stamens 3-6; fruit of 
2-4 indehiscent carpels. 

Cabomba aquatica Aubl. PI. Guian. 1: 321. pi. 124. 1775. 

Stems often much elongate; submerged leaves opposite or 
verticillate, usually long-petiolate, the floating leaves elliptic; 
flowers axillary, pedunculate, 1-1.5 cm. broad, yellow. Illustrated, 
Mart. Fl. Bras. 4, pt. 2: pi. 37. 

Loreto: Rio Itaya above Iquitos, floating in river, 110 meters, 
Killip & Smith 29282. Widely distributed in tropical America. 

69. RANUNCULACEAE. Buttercup Family 

The visitor to Peru from a northern land may enjoy the thrill 
of seeing some of the largest buttercups that grow, the remark- 
able Laccopetalum giganteum and Krapfia Raimondii. Their hand- 
some leaves 40-70 cm. long and huge flowers, that measure to 10 
cm. across and even larger, place them among the notable plants of 
their group. The buttercup family in Peru includes a number of 
genera that even the most casual student may recognize, as Caltha 
and Clematis, Anemone, and Thalictrum, not to mention many sorts 
of yellow buttercups that, unless one is a professional botanist, 
appear like those at home, if that is in a northern clime. In the 
preparation of the following compilation I acknowledge especially 
my indebtedness to the work of Weddell and of Ulbrich. 



640 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Carpels 1-ovuled, the fruit a head of achenes. 
Plants not scandent, herbaceous. 
Sepals unappendaged. 
Sepals 5-many, petaloid; petals none. 
Low, tufted, alpine perennials, the leaves not fern-like. 

Leaves divided 1. Anemone. 

Leaves entire 2. Capethia. 

Tall perennials of middle altitudes with compound leaves. 

3. Thalictrum. 

Sepals 5, herbaceous or petaloid, but petals present, usually 
5-many. 

Receptacle rather disk-like, not clavate, the gynophore 
woody or spongy, the perianth caducous. 

5. Ranunculus. 

Receptacle fleshy, somewhat clavate, the perianth per- 
sistent. 

Petals thin, with one nectary .^. . . . 6. Krapfia. 

Petals fleshy, foveolate with many nectaries. 

7. Laccopetalum. 
Sepals spurred; fruiting receptacle spike-like ... 8. Myosurus. 

Plants scandent, fruticose 4. Clematis. 

Carpels many-ovuled, the follicular fruits dehiscent; sepals 5, petaloid ; 
petals none; leaves appendaged at the base 9. Caltha. 

1. ANEMONE L. 

Petals none. Flowers often cymose. Low perennials with erect 
scapes. Growing on rock outcrops, at 2,200-3,600 meters. 
Achenes glabrous, in globose heads; divisions of the leaves broad. 

A. helleborifolia. 

Achenes densely tomentose or sericeous, the heads elongate-oblong; 
divisions of the leaves narrowly linear A. triternata. 

Anemone helleborifolia DC. Syst. 1: 211. 1818. A. aequi- 
noctialis Poepp. Fragm. Syn. PI. Chile 28. 1833. A. perumana Britton, 
Ann. N. Y. Acad. Sci. 4: 225. 1892(7). 

Radical leaves subcoriaceous, numerous, arising from a fleshy, 
densely fibrillose caudex; petiole 7-10 cm. long, pilose, the leaf 
blade pedatisect, the glabrous, thickish, cuneately petiolulate 
divisions trifid, their acute lobes serrate, sometimes nearly trifid; 



FLORA OF PERU 641 

stems erect, fistulose, 30 cm. high or more; umbel compound, usually 
3-4-radiate, the elongate peduncle glabrous; involucres all 3-foliate, 
subsessile; flowers white; sepals 5, oval; achenes glabrous, nearly 
oblong, turgid, distinctly stipitate, gibbous, with a short, strongly 
revolute or hooked style, forming a globular head, the receptacle 
villous. Description of achenes by Gray, who records that the 
leaves may be thin or more or less coriaceous. Ulbrich, Bot. Jahrb. 
37: 237. 1905, remarks that from description A. peruviana seems to 
be "only a form." It was based on Mathews 537 and McLean. 

Huanuco: Piedra Grande, 1,500 meters, 3689. Huanuco (Dom- 
bey; Ruiz). Cajabamba, Weberbauer 3150 (det. Ulbrich). Junin: 
Palca, 2,700 meters, Weberbauer 1788; 247 (det. Ulbrich). Huasa- 
huasi (Dombey). Tarma (Ruiz & Pavon). Lima: Huaros, 3,500 
meters, Pennell 14729. Matucana, 479; Weberbauer 196 (det. 
Ulbrich). Obrajillo, Banos (Wilkes Exped.). Cajamarca: San 
Pablo, Weberbauer 3862 (det. Ulbrich). San Pablo, 2,200 meters 
(Weberbauer, 257). Ancash: Above Samanco, 3,000 meters (Weber- 
bauer, 169). Cuzco: Paucartambo, 3,500 meters (Herrera 1395, 
as A. decapetala L.). Ollantaitambo, 2,800 meters, Herrera 3404- 
Without locality, Weberbauer 6490. Puno: Sandia, 3,000 meters 
(Weberbauer, 237). "Polizonis," "Soliman," "arracacha cimarrona." 

Anemone triternata Vahl, Symb. 3: 74. pi. 65. 1794. 

Radical leaves long-petiolate, much divided into numerous very 
narrow, slender segments, almost glabrous; stems about 30 cm. high, 
slender, 1-flowered, the much-divided involucre borne above the 
middle; sepals 10-12, white, linear-oblong; fruit heads very densely 
sericeous. 

Cuzco: Colinas del Sacsahuaman, 3,600 meters, Herrera 2362. 
Extending to Brazil, Argentina, and Chile. 

2. CAPETHIA Britton 

A small, depressed plant with entire leaves and solitary flowers. 
Fruit indehiscent, ligneous. Flowers with both involucre and calyx. 
Otherwise similar to Anemone. 

Capethia integrifolia (DC.) Britton, Ann. N. Y. Acad. Sci. 6: 
234. 1891. C. Weddellii Britton, op. cit. 235(?). Anemone integrifolia 
HBK. ex Wedd. Chlor. And. 2: 298. 1857. Hepatica integrifolia DC. 
Syst. 1:217. 1818. 

Petioles and scapes densely villous with long, white hairs; radical 
leaves many, coriaceous, ovate, subacute, entire, somewhat pubes- 



642 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

cent; involucral leaves none or merged with the many sepals, these 
lanceolate or linear; carpels villous, ovoid, 3 mm. long. Petals and 
scapes very short, the flowers large, or the scape sometimes 2-3 cm. 
high, the flowers then smaller (Weddell). Flowers white or yellow! 
(Ulbrich, Bot. Jahrb. 37: 401. 1905). Illustrated (as C. Weddellii), 
Wedd. Chlor. And. 2: pi. 83. 

Cajamarca: Between Micuipampa and Hualgayoc, 3,600 meters 
(Humboldt & Bonpland, type). Hualgayoc, 4,000 meters (Weber- 
bauer, 271). Cuzco: (Gay). Huanuco: Chavanillo, dry, grassy 
slope, 2,400 meters, 1965. Puno : Sachapata (Lechler 2706). Bolivia; 
Ecuador. 

3. THALICTRUM L. 

Reference: Lecoyer, Bull. Soc. Bot. Belg. 24: 78-324. 1885. 

This well-known genus is represented in Peru by only 4 closely 
related species, all characterized by 2-4-pinnate leaves, the leaflets 
small and in greater or less degree paler beneath. The oblong-oval 
perianth segments are, with the stamens, greenish yellow or more 
or less lined or tinged with purple. 

Styles 4-5 mm. long or longer; leaflets mostly not regularly 3-lobed 

or 3-toothed. 
Leaflets stipellate. 
Achenes slender-veined; leaf sheaths ovate-lanceolate. 

T. longistylum. 
Achenes coarsely nerved, the nerves thick or nodose; leaf 

sheaths ample, broadly ovate T. vesciculosum. 

Leaflets estipellate T. rutidocarpum. 

Styles about 3 mm. long; most of the leaflets regularly 3-lobed or 
3-toothed T. podocarpum. 

Thalictrum longistylum DC. Syst. 1: 171. 1818; 126. 

Leaflets often somewhat pubescent, subentire or usually with 
1-several teeth of unequal size, stipellate; anthers finally 4-6 mm. 
long; achenes stipitate or sometimes subsessile, reticulate-costate, 
enodose. The type by Dombey ("Am. merid.") was probably from 
Peru. Illustrated, Deless. Icon. pi. 7; Lecoyer, op. cit. pi. 2 (fruit). 

Lima: Cheuchin: Ruiz & Pawn. Purruchuca (Mathews). Viso, 
2,600 meters, Weberbauer 144; 169. Matucana, shrubby thicket, 
2,400 meters, 409. Canta, open, rocky slope, 3,000 meters, Pennell 
14338. Rio Blanco, 3,000-3,500 meters, open hillside, Killip & 
Smith 21572. Bolivia to Mexico. 



FLORA OF PERU 643 

Thalictrum podocarpum HBK. Nov. Gen. & Sp. 5: 38. 
1821; 129. 

Glabrous; leaflets mostly very evenly 3-lobed across the tip, 
stipellate; anthers finally 4-6 mm. long; achenes sessile or subsessile, 
reticulate-nerved, compressed. Illustrated, Lecoyer, op. cit. pi. 2. 

Cajamarca: Ocros, 3,000 meters, Weberbauer 2739. Lima: 
Matucana, 2,400 meters, 269. Near Viso (Weberbauer, 169). 
Amazonas: Region of Moyobamba (Weberbauer, 266; 267). 
Huanuco(?): Rio Chiquie'n (Weberbauer, 178). Huanuco: Tambo 
de Vaca, 3,600 meters, 4466. Junin: Near Huancayo, open, rocky 
hillside, 3,400 meters, Killip & Smith 22125. Cuzco: Colinas del 
Sacsahuaman, 3,200-3,600 meters, Herrera 829. Urubamba, Soukup 
40. Quiquijana, 3,220 meters, grassy wayside, Pennell 13540. 
Hacienda Churu, Prov. Paucartambo, 3,500 meters, Herrera 1047 a. 
Bolivia to Venezuela. 

Thalictrum rutidocarpum DC. Syst. 1: 172. 1818; 127. 

Allied to T. longistylum, but the small leaflets estipellate and 
pubescent and the achenes nodose. The type from "Am. merid." 
by Dombey may well have come from Peru, to which country 
Lecoyer accredits it. 

Peru: Probably. Ecuador. 

Thalictrum vesiculosum Lecoyer, Bull. Soc. Bot. Belg. 24: 
130. 1885. 

Glabrous; leaflets sometimes 3- toothed but not usually nor as 
regularly as in T. podocarpum, stipellate; anthers 3 mm. long or 
finally often much longer; achenes stipitate. The salient and 
perhaps single constant character of this species is found in the 
achene nervation, which is strongly marked. Illustrated, Lecoyer, 
op. cit. pi. 2. 

Amazonas: Chachapoyas, Mathews. Puno: Sandia Valley, 
Weberbauer 553; 237. Bolivia to Colombia. 

4. CLEMATIS L. 

Reference: Eichler in Mart. Fl. Bras. 13, pt. 1: 143-150. 1864. 

These familiar vines, characterized by opposite, pinnately com- 
pound leaves, the leaflets usually more or less toothed, petaloid 
sepals but no petals, and achenes conspicuous by the persistent, 
plumose or silky, elongate style, are obscurely separable in Peru 
into several species that more probably represent races of fewer 



644 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

forms. The following key, only general in character, is at best 
merely suggestive. The questionable C. Seemanni Kuntze, not seen, 
is omitted. 

Leaves finely dissected, at least many of the leaflets about 1 cm. wide 
or smaller or even filiform or, when wider, acuminate. 

Leaflets or their divisions linear-filiform C. millifoliolata. 

Leaflets at least 5-15 mm. wide. 

Leaflets glabrous or sparsely pubescent beneath; petaloid fila- 
ments lacking; leaflets often only 3-5 mm. wide. 

C. parvifrons. 
Leaflets more or less distinctly pubescent beneath, often larger; 

petaloid filaments often present C. peruviana. 

Leaves with ample leaflets mostly 2-3 cm. wide, coarsely lobed or 

entire. 

Leaves evenly and usually densely pubescent beneath . . C. sericea. 
Leaves glabrous or sparsely and unevenly pubescent beneath. 
Flowers reddish white, the segments barely 10 mm. long; most 

of the leaflets irregularly roundish-lobed C. alborosea. 

Flowers greenish or yellowish white, often longer; leaflets entire 

or more or less equally 3-lobed. 
Leaflets more or less lobed or toothed. 

Leaflets rather regularly 3-lobed C. Hilarii. 

Leaflets irregularly and remotely lobed or toothed. 

C. campestris. 

Leaflets entire or mostly so C. dioica. 

Clematis alborosea Ulbrich, Notizbl. Bot. Gart. Berlin 8: 
325. 1923. 

Leaves once or twice trifoliolate, 10 cm. long or longer, the ovate 
or obliquely ovate, irregularly and deeply crenate terminal leaflets 
to nearly 6 cm. long and broad, the lateral ones smaller; inflorescence 
ample, glabrous or the flowers sparsely pilose; sepals 5-6, to 12 mm. 
long and 4-5 mm. wide; filaments glabrous, 7-8 mm. long; pistillate 
flowers unknown. 

Junin: Near Comas, below Hacienda Runatullu, 3,300 meters, 
Weberbauer 6609 (type). Cajamarca: Nancho, Raimondi(t}. 

Clematis campestris St. Hil. Fl. Bras. 1: 4. 1825; 145. 
Stems 6-angled, 1-striate between the angles, tomentose, becoming 
glabrescent; petioles divaricate, flexuous or cirrose; leaves pinnate, 



FLORA OF PERU 645 

15 cm. long or longer, with 2-5 pairs of leaflets, these ternate, cor- 
date-lanceolate or linear, attenuate, acute, entire; panicles shorter 
than the leaves, 3-5 or rarely 7-flowered; upper bracts oblong-linear; 
bractlets remote, lanceolate, flowers perfect, 12-18 mm. broad; 
petals lanceolate, grayish-tomentose within or without. By Kuntze 
regarded as C. dioica, sens. lat. Illustrated, Mart. Fl. Bras. 13, pt. 1 : 
pi. 33, 35. 

Moquehua: Moquehua, climbing over stone walls, 1,500 meters, 
Weberbauer 7452. Southern South America and Brazil. 

Clematis dioica L. Amoen. Acad. 5: 398. 1759; Sp. PI. ed. 2. 
765. 1762; 147. C. thalictroides Steud. Flora 39: 407. 1856. C. 
Haenkiana Presl, Rel. Haenk. 2: 69. 1831(7), fide Kuntze. 

Stems 10-12-angled, glabrous as the entire plant or more or less 
tomentose; leaves simple to pinnate with 3 pinnae, the leaflets ternate, 
entire or dentate, ovate or ovate-cordate, acute or acuminate; panicles 
as long as or often longer than the leaves. Very variable; description 
by Eichler. The Isern specimen seems to approach C. sericea. 

Amazonas: Chachapoyas (Mathews). Lima: Matucana, 2,400 
meters, 398. Quive, 800-1,200 meters, in thickets, Pennell 14295. 
Near Viscas, 2,100 meters, in thicket, Pennell 14448. Banos (Wilkes 
Exped.). Matucana, Isern 2086. Cajamarca: Above Supi, 1,600 
meters (Weberbauer, 162). Huaraz, 2,200 meters (Weberbauer, 172); 
at 2,400 meters (Weberbauer, 173). Ayacucho: Carrapa, wooded 
hillside, 1,200 meters, Kittip & Smith 23225, 22316. Cuzco: Ha- 
cienda Chancamayo, Valle Lares, Diehl 2462. Machupicchu, 2,400 
meters, Herrera 2005. Below Pillahuata, 2,200 meters, in forest, 
Pennell 1 4061 . Huanuco : Muna, 2, 100 meters, 3986, 3901 . Yanano, 
1,800 meters, 3727. Llata, 2,100 meters, on shrubs along stream, 
2260. Cani, 2,550 meters, on shrubs in sun, 3474- Junin: Hua- 
capistana, 1,800-2,400 meters, Kittip & Smith 24329. Colonia 
Perene 1 , 680 meters, in thicket, Killip & Smith 25050. Loreto: 
Balsapuerto, 220 meters, King 2922. San Martin: Zepelacio, 1,200- 
1,600 meters, King 3526, 3465. San Roque, 1,400 meters, Williams 
7770. Without locality: Weberbauer 5377. Widely distributed in 
tropical America. 

Clematis Hilarii Spreng. Syst. Veg. 5: 177. 1828; 146. 

Similar to C. campestris, but the leaves more or less ternate, the 
leaflets mostly 3-lobed, the lobes very acute. Treated as a sub- 
species of C. dioica by Kuntze. 

Peru: Probably. Mexico to Uruguay. 



646 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Clematis millifoliolata Eichl. in Mart. Fl. Bras. 13, pt. 1: 
150. 1864. 

Gray-tomentose, or the 6-angled, striate stem finally glabrescent; 
leaves 10-12 cm. long and broad, the ultimate segments only 2-4 mm. 
long, linear-lanceolate; panicles scarcely equaling the leaves, 3-7- 
flowered; bracts filiform, the bractlets broadly ovate-cordate, 3- 
lobed, densely villous-tomentose like the sepals, these 4, elliptic- 
cordate, pubescent within below the apex, about equaled by the 
stamens. 

Peru: (Besser, type). Moquehua: Carumas, 3,200 meters, climb- 
ing in thicket of moist ravine, Weberbauer 7288. Arequipa: Nevado 
de Chachani, open, rocky ravine, 3,600 meters, Pennell 13263. 

Clematis peruviana DC. Syst. 1: 141. 1818. 

Stems terete, somewhat purplish, pubescent; petioles terete, 
pubescent; leaves bipinnatifid, the segments sometimes ternate, 
sometimes 3-parted, ovate at the base, acutely incised, dentate 
above, glabrous or pubescent on the upper surface, villous beneath; 
peduncles axillary, as long as the leaves, sparsely branched, few- 
flowered, the opposite branchlets 1-flowered; bracts oval, subacute, 
entire; flowers subpaniculate, polygamous, some staminate, others 
perfect; sepals 4, widely spreading, ovate-oblong, acute, nearly twice 
longer than the stamens, these sometimes very many, sometimes few, 
the glabrous filaments finally reflexed; anthers oval; ovaries about 40. 

Peru: Without locality (Dombey, type). Lima: Bafios (Wilkes 
Exped.). Obrajillo, Cullnai (Cruckshanks). Lima, 3,500 meters 
(Weberbauer, 170; 169). Ayacucho: Between Chavina and Cora- 
cora, 3,100 meters, Weberbauer 5786. Cuzco: (Herrera, det. Ul- 
brich). Huanuco: Rio Maranon region, 3,300 meters (Weberbauer, 
178). Rio Blanco, 3,000-3,500 meters, open hillside, Kittip & Smith 
21563. Huaros, 3,400 meters, open, rocky banks, Pennell 14706. 
Canta, 3,300 meters, rocky thickets, Pennell 14617. Viso, 2,700 
meters, trailing over shrubs, 575. 

Clematis parvifrons Ulbrich, Notizbl. Bot. Gart. Berlin 8: 
326. 1923. 

Glabrous except the young parts and the lightly pilose, rotund- 
elliptic perianth segments, these 8 mm. long; leaves bipinnate or 
tripinnate, the segments acutely toothed, ovate-lanceolate or sub- 
orbicular; stamens scarcely 4 mm. long; fruit glabrous, ovoid, 4-5 mm. 
long, the styles about 4 cm. long, rather laxly pilose. The native 
name means "winding here and there" (Weberbauer). 



FLORA OF PERU 647 

Ayacucho: Hacienda Totorabamba, 3,500 meters, Weberbauer 
5464 (type). Huanta, 3,500 meters, Weberbauer 7488, 7488a; Killip & 
Smith 23325. Prov. Cangallo, Raimondi. Cuzco: Huasao, 3,300 
meters, Herrera. Sacsahuaman, 3,500 meters, Herrera; Soukup 80. 
Arequipa : Near Ancara and Calf ara, Raimondi. ' 'Arhuiarhui. ' ' 

Clematis Seemanni Kuntze, Verb. Bot. Ver. Brandenb. 26: 
146. 1885. 

Branches short, with many crowded, few-flowered branchlets; 
leaflets lobed or 2-parted, sometimes biternately bipinnate, the ulti- 
mate divisions small, ovate, not at all dentate, glabrous; peduncles 
longer than the leaves; sepals 5-6, oblong, obtuse, 1-1.5 cm. long; 
bracts none or narrow. Probably a race of C. dioica, sens, lat., or of 
C. peruviana. 

Common about the village of Andahuageas (Pearce, type). 
Lima: Obrajillo, San Mateo (Mathews). Bolivia (?). 

Clematis sericea DC. Syst. 1 : 144. 1818. 

Similar to C. dioica, but the many cauline leaves pinnate, usually 
densely pubescent and evenly so beneath, the leaflets entire, dentate, 
or more or less divided. C. brasiliana DC. Syst. 1 : 143. 1818, regarded 
as a subspecies of C. dioica by Kuntze, seems to be the form with 
entire or few-dentate leaflets ranging from Mexico to Uruguay 
through all tropical America (Kuntze). Kuntze treats this as a sub- 
species of C. dioica, and credits it to Peru. 

Puno: Sandia Valley, 2,000-3,000 meters (Weberbauer, 237); 
6589. Cuzco: Valle del Apurimac, Herrera in 1929. Paucartambo 
to Tres Cruces, 3,300 meters, in thicket, Pennell 14150. Huanuco: 
Huacachi, 1,950 meters, 4098. Mito, 2,700 meters, on shrubs in 
sunny places, 1553. North to Mexico and south to Argentina. 

5. RANUNCULUS L. Buttercup 
Reference: Weddell, Chlor. And. 2: 297-307. 1857-62. 
This almost universally distributed group requires no charac- 
terization here. It is interpreted to include Oxygraphis, which is at 
least a convenient disposition for practical purposes. Several of the 
following species are doubtfully distinct, but a revision of all the 
Andean forms is needed before they can be properly disposed. 

Plants aquatic. 

Floating leaves more or less lobed and cordate. . . R. trichophyllus . 
Floating leaves entire, not cordate R. Mandonianus. 



648 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

Plants terrestrial (R. flagelliformis sometimes subaquatic). 
Leaves all alike, the plants tufted or creeping, glabrous or rarely 

slightly pilose. 

Leaves entire or barely crenate, cordate-rotund or truncate at 
the base, often 1.5-2.5 cm. wide; flowers small; plants 

creeping R. flagelliformis, R. sarmentoso-repens. 

Leaves rarely entire, the plants then tufted and the flowers 
large; leaves usually at least crenate, sometimes 3-5-parted. 
Leaves mostly more or less crenate or lobed; flowers small. 
Plants glabrous. 
Flowers yellow or yellowish; leaves rarely lobed. 

R. Cytnbalaria. 

Flowers white; leaves 3-5-lobed R. hemignostus. 

Plants slightly hairy. 

Leaves crenate R. nubigenus. 

Leaves 3-5-parted R. breviscapus. 

Leaves entire; flowers nearly 10 mm. wide. . .R. arequipensis. 

Leaves often dissimilar; stems or cauline leaves or both developed 
and the latter often differing from the basal leaves, or at 
least the plants pubescent. 
Leaves, at least, glabrous or nearly so, the basal ones not parted 

or deeply lobed. 
Basal leaves cordate-rotund, crenate; achenes not muricate. 

Sepals densely villous R. peruvianus. 

Sepals glabrous or lightly pilose R. bonariensis. 

Basal leaves flabellate, coarsely toothed; achenes muricate. 

R. muricatus. 
Leaves, as the stems, usually pubescent, deeply 3-lobed, trifid, 

or pinnate. 
Leaves trifid or deeply 3-5-lobed or, if subpinnate, the plants 

low and tufted (R. sibbaldioides). 
Stems well developed. 

Leaves mostly or usually only deeply 3-5-lobed, or, in 

any case, the plants erect or suberect. 
Leaf lobes, at least the middle one, deeply 3-lobed 

or twice cleft. 
Basal leaves mostly or all ternate . . . .R. geranioides. 



FLORA OF PERU 649 

Basal leaves palmately 3-5-parted nearly to the base. 

R. acris. 

Leaf lobes merely toothed R. chilensis. 

Leaves all or some of them trifid, the plants spreading- 

repent R. repens. 

Stems not obvious, the peduncle 3-4 cm. high, little if at 
all exceeding the leaves. 

Calyx spreading R. sibbaldioides. 

Calyx appressed R. breviscapus. 

Leaves more or less pinnately lobed or divided; plants tall. 
Plants usually markedly pubescent, with coarsely divided, 

dissimilar leaves. 
Uppermost reduced leaves usually 2 or more; plants 

often tall, stout, and stiff. 
Petals obovate; style straight or nearly so. 

Petals 5-6; style straight R. pilosus. 

Petals 10; style a little inclined. .R. argemonifolius. 
Petals oblong-cuneate; style often recurved. 

R. chiclensis. 

Uppermost reduced leaf often bract-like; plants often 
relatively lax and low. 

R. praemorsus, R. setoso-pilosus. 

Plants sparsely appressed-pilose, with finely divided leaves, 
the ultimate segments often only 1-2 mm. wide. 

R. palimbifolius, R. Sodiroi. 

Ranunculus acris L. Sp. PI. 554. 1753. 

More or less strigose-hirsute, several dm. tall; basal leaves 
3-several cm. wide, the broadly rhombic divisions twice parted into 
lanceolate lobes; petals 1 cm. long or longer, at least twice longer 
than the sepals; achene beak slightly curved, about a fourth as long 
as the achene. 

Peru: Probably. Nearly cosmopolitan. 

Ranunculus arequipensis Ulbrich, Notizbl. Bot. Gart. Berlin 
8: 326. 1923. 

A small, glabrous, stoloniferous herb, the flowering stems tufted; 
petioles often 2-5 cm. long; leaves 3-8 mm. long, 3-6 mm. broad, 
almost nerveless; flowers axillary, solitary, the peduncle 2.5-4 cm. 
long; sepals 3 and 5-6 mm. long; petals mostly 6, oblong-linear, 



650 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

4-5 mm. long, 1-1.5 mm. broad; carpels very many, nearly 1 mm. 
long, obliquely obovoid, attenuately stipitate, apiculate. Related 
to R. boliviensis Phil., with smaller flowers, 5 sepals, and few (12-15) 
carpels (Ulbrich). 

Arequipa: Near Cailloma, 4,500 meters, Weberbauer 6887 (type). 
Lake Titicaca, Meyen(1}. 

Ranunculus argemonifolius Griseb. Goett. Abh. 19: 70. 1874. 

Perennial, the pilose stems about 60 cm. high, divided above 
into few-flowered branches; blades of the lower leaves 7 cm. long; 
leaves distant, pinnatisect, pilose, oblong, the divisions with few 
pinnae, these broadly cuneate, unequally 3-pinnatifid, the lobes 
ovate or oblong, callous- tipped, roundish-acute; pedicels terminal 
and axillary; flowers yellow, the 10 spatulate-oblong, subretuse 
petals 12-14 mm. long, more than twice as long as the 5 glabrous, 
reflexed sepals; achenes ovate, compressed, reticulate, the elongate, 
subulate style a little inclined. Allied by the author to R. palimbi- 
folius. 

Junin: Palca, Isern 2107. Huacapistana, Weberbauer 2199. 
Puno: Near Puno, 4,000 meters, Soukup 15. Ayacucho: Yana- 
monte, 2,700 meters, Weberbauer 5653. Cuzco: Pillahuata, 3,300 
meters, open, grassy paramo, Pennell 14140. Huanuco: Mito, 2,700 
meters, 1441- Huacachi, moist grasslands, 1,950 meters, 3887. 
Southeast of Huanuco, 3,150 meters, barley field, 2096. Junin: 
Chinche, 3,450 meters, grassy slope, 1264- La Quinua, 3,600 meters, 
wet meadow, 2014- Lima: Rio Blanco, 3,600 meters, along irrigation 
ditch, 727. Extending to Argentina. 

Ranunculus bonariensis Poir. Diet. 6: 102. 1804. 

Glabrous or nearly so, the creeping to erect stems often branched ; 
leaves cordate, orbicular, entire or crenate, dentate, on greatly 
elongate petioles, the blades to 5 cm. wide or larger; petals usually 
only 2 or 3, shorter than the finally reflexed, glabrous or slightly 
pilose sepals; achenes numerous, tiny, not at all compressed, ob- 
scurely margined, forming a cylindric head. Cf. R. arequipensis. 
Illustrated, Deless. Icon. Sel. 1: pi. 29. 

Peru: Probably. South America. 

Ranunculus breviscapus DC. Syst. 1: 253. 1818. Oxygraphis 
breviscapa Prantl, Bot. Jahrb. 9: 263. 1888. 

Tufted, with creeping offshoots; basal leaves many, glabrous or 
pilose, orbicular-cordate, 3-5-parted, the lobes incised-dentate, the 



FLORA OF PERU 651 

teeth acute; petioles 7 cm. long, the 1-flowered peduncles half as 
long; flowers small, the petals obtuse; calyx glabrous, appressed. 
The type, by Dombey without locality, may be rather from Chile. 
Peru: (Dombey}. 

Ranunculus chiclensis Ball, Journ. Proc. Linn. Soc. 22: 
28. 1885. 

Plants 30 cm. high or taller, the entire plant hirsute with sub- 
appressed hairs; radical and lower leaves pinnatifid, the alternate 
segments 3-parted or pinnate, the upper leaves oblong-cuneate, 
variously incised; flowers long-peduncled, rather large; petals 5-10; 
nearly oblong-cuneate, truncate or lightly emarginate, obscurely 
lineate; achenes compressed, gradually narrowed to the beak, this 
often but not always recurved. A very distinct species, especially 
remarkable by the form of the petals (Ball). 

Lima: Chicla (Ball). 

Ranunculus chilensis DC. Syst. 1: 286. 1818. 

Procumbent stems and petioles hispid, the shorter flowering 
stems densely so, the longer sterile ones sparsely; leaves subvillous, 
the petioles of the numerous radical leaves hispid, 5 cm. long, the 
blades orbicular, deeply cordate, 3-5-parted to below the middle, the 
lobes coarsely dentate apically; petioles of cauline leaves 2.5 cm. long, 
dilated at the base into a glabrous, membranous sheath; peduncles 
opposite the leaves, densely pilose, 1-flowered, the flowers small; 
calyx densely villous. Flowers 1.5 cm. broad according to Reiche. 
Neg. 27645. 

Huanuco: Tambo de Vaca, 3,900 meters, mossy, rocky uplands, 
4386. Chile. 

Ranunculus Cymbalaria Pursh, Fl. Amer. Sept. 392. 1814; 
300. R. tridentatus HBK. ex DC. Syst. 1: 252. 1818. Oxygraphis 
Cymbalaria Prantl, Bot. Jahrb. 9: 263. 1888. 

A diminutive, glabrous annual, usually less than 3 cm. high, ces- 
pitose or sometimes the stems elongate and creeping; leaves 2-6 mm. 
long, ovate or rotund, 3-lobed at the apex or coarsely and obtusely 
dentate, or merely crenate, rather thick; peduncle usually 1-flowered; 
sepals and petals 5, the former elliptic, the latter obovate-spatulate, 
unequal, obtuse; fruit heads ovate, 4-6 mm. long, the achenes very 
many, minute, compressed, ovate, acute, veiny, acutish-margined. 
Compare the doubtfully distinct R. nubigenus. Includes, following 
Weddell and Fries, R. minutus Gay, Fl. Chil. 1: 38. 1845. The 



652 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

diminutive R. psychrophilus Wedd. op. cit. 300 is comparable. The 
Peruvian specimen determined by Ulbrich, "compare tridentatus." 
Leaves vary from 5-20 mm. long (Fries). In wet, probably always 
saline places. 

Puno: Raimondi. North America, south in the Andes to 
Patagonia. 

Ranunculus flagelliformis Smith in Rees, Cycl. 29: No. 13. 
1802-20. 

Glabrous, the weak stems often creeping and rooting at the nodes; 
leaves long-petioled, orbicular-cordate, entire, obscurely crenate, the 
petioles to 5 cm. long; peduncles opposite the leaves; flowers white, 
2-4 mm. broad; petals 2-3, slightly shorter than the 3-4 finally 
reflexed sepals; achenes few, elliptic or oblong, bluntly pointed, 
slightly margined, foveolate under a lens, forming a subglobose head. 
Description from Eichler. Illustrated, Mart. Fl. Bras. 13, pt. 1: 
pi. 34. 

Lima: Cheuchin, Dombey. Huanuco: Tomaiquichua, 2,550 
meters, floating in shallow pool, 2435. Mito, 2,700 meters, on wet 
mud, 1537. Without locality: Weberbauer 6931. San Martin: 
Tarapoto, Spruce 4470. Southern South America to Colombia. 

Ranunculus geranioides HBK. ex DC. Syst. 1: 286. 1818. 

Stems ascending, simple, about 10 cm. high, silky-hirsute like 
the ternate radical leaves, these especially so beneath, 3 cm. broad; 
leaflets cuneately petiolate, incised-serrate, the rhombic lateral ones 
2-3-parted; petiole to 5 cm. long, pubescent like the leaves and with a 
membranous sheath; cauline leaves more or less 3-parted; flowers 
generally 2, the pubescent peduncle about 2 cm. long to twice as long 
and divaricate in fruit; sepals hirsute, reflexed, slightly shorter than 
the petals; achenes obliquely obovate, compressed, the beak some- 
what curved, forming a small, subglobose head. Illustrated, HBK. 
Nov. Gen. & Sp. 5: pi. 427. 

Huanuco: Mufia, 2,400 meters, open, wet, grassy places, 4295. 
North to Colombia. 

Ranunculus hemignostus Steud. Flora 39: 404. 1856. 

Glabrous throughout; stems scarcely 2.5 cm. long; petioles elon- 
gate, nerved and dilated basally; leaves fleshy, oblong, mostly 3- 
lobed or rarely 5-many-lobed, the lobes obtuse; flowers solitary, the 
petals white; achenes numerous, glabrous, marginate, apiculate, 
forming an ovoid, subrotund head. 

Puno: Azangaro (Lechler 1793). 



FLORA OF PERU 653 

Ranunculus Mandonianus Wedd. Chlor. And. 2: 299. 1857. 

A glabrous aquatic with the habit of Alisma natans; stems elon- 
gate, rooting at the nodes; leaves floating, long-petioled, the blades 
1.5-2 cm. long, elliptic, obtuse at each end, obsoletely 3-nerved; 
flowers small, long-pediceled; sepals orbicular; petals 3-4, white, 
obovate-spatulate, scarcely equaling the calyx, the nectary at the 
base of the blade; stamens fewer than 5; fruit globose, 4 mm. thick, 
the achenes oblong-ovate, apiculate, smooth. Neg. 27652. 

Ancash: Huaraz, 4,400 meters, Weberbauer 3108; 227. Huanuco: 
Punco, 4,050 meters, 2474. Mito, 2,700 meters, 3355. Bolivia. 

Ranunculus muricatus L. Sp. PL 78. 1753. 

A nearly glabrous, erect or diffuse annual, the radical or lower 
leaves suborbicular or reniform, 3-lobed and coarsely crenate, the 
upper 3-parted, cuneate at the base; petals slightly longer than the 
sepals; achenes compressed, many, sharply tuberculate, prominently 
beaked, with a well-defined, unarmed margin. 

Lima: Environs of Lima, Raimondi. Nearly cosmopolitan. 

Ranunculus nubigenus HBK. ex DC. Syst. 1: 253. 1818; 300. 
Oxygraphis nubigenus Prantl, Bot. Jahrb. 9: 263. 1888. 

A diminutive annual with long-petioled radical leaves, these with 
suborbicular, rather fleshy, glabrous, 3-7-crenate blades 3-5 mm. 
broad, the petioles pilose; stems ascending, filiform, nearly leafless, 
about 2-flowered, pubescent at the tip; bracts or cauline leaves linear, 
somewhat 3-toothed; flowers minute; calyx appressed, the corolla 
reduced. In Peru, according to Prantl; cf. R. Cymbalaria Pursh, f. 
hebecaulis Fernald, Rhodora 16: 162. 1914. 

Peru: Probably. Ecuador. 

Ranunculus palimbifolius Wedd. Chlor. And. 2: 303. 1857. 

Plants glabrate, 20-40 cm. high, few-leaved, 2-3-flowered; radical 
leaves long-petioled, the blades 4-8 cm. long, oblong, twice or thrice 
pinnatifid, the linear, pilose or glabrate divisions callous- tipped ; 
upper cauline leaves sometimes entire; petals 5, yellow, obovate, 
obtuse, striate without, half again as long as the reflexed, villous 
sepals; anthers oblong; achenes numerous, compressed, the style 
nearly 2 mm. long. Flowers the size of those of R. acris. Neg. 27656. 

Cajamarca: At 3,300 meters (Raimondi}. Ecuador. 

Ranunculus peruvianus Pers. Syn. 2: 103. 1807; 301. 
Perennial from long, fibrous roots, with a fibrillose caudex; stems 
slender, few-flowered, sparsely pilose; basal leaves long-petioled, 



654 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

somewhat pilose, the few cauline leaves sessile, parted into entire, 
linear lobes; flowers conspicuously pediceled; calyx spreading, very 
villous; petals 5, yellow, streaked with red without, suborbicular, 
half as long again as the calyx; achenes ovate, compressed, smooth, 
the fruit head globose. Illustrated, Deless. Icon. pi. 37. 

Cajamarca: Between Cajamarca and Chachapoyas, 3,900 meters 
(Raimondi). Hualgayoc, grass steppes, 3,900 meters (Weberbauer, 
271). Without locality (Jussieu, type). Ecuador. 

Ranunculus pilosus HBK. ex DC. Syst. 1: 287. 1818. 

Stems ascending, dichotomously branched above, hirsute; radical 
leaves petiolate, ternate, appressed-pilose on both sides, 2.5 cm. long 
and 2 cm. broad or larger; leaflets incised-serrate, the lateral ones 
ovate-rhombic, entire or 2-3-lobed, the intermediate ones petiolu- 
late, subrotund, 3-lobed or trifid like the terminal lobes; petioles 
pilose, 5 cm. long; cauline leaves deeply divided, the upper gradually 
more nearly simple; peduncle silky-pilose like the reflexed sepals, 
these 2 or 3 times shorter than the 5 or 6 petals; achenes beaked by 
the straight, persistent style, glabrous, compressed, obliquely sub- 
rotund-obovate, forming a globose head. R. erodiifolius Gay, at 
least as to Herrera, apparently. Plant poisonous, with an acrid 
taste, killing rabbits that eat it (Herrera). Also in the Department 
of Apurimac, according to Herrera. 

Amazonas: Chachapoyas, 3,000 meters, Williams 7393.Cuzco: 
Valle de Urubamba ( Herrera 638}. Sacsahuaman, 3,500 meters, 
Herrera 830. Valle de Paucartambo, 3,500 meters, Herrera 1047, 
3368, 1406, 286. Prov. Canchis, Sicuani, 3,550 meters (Hicken). 
Huasao, 3,200 meters, Herrera 3021. North to Colombia. "Sicuta," 
"chchapo-chchapo," "huarenccaiso," "cicuta." 

Ranunculus praemorsus HBK. in DC. Syst. 1: 292. 1818; 302. 

Perennial, nearly throughout with silky, appressed hairs, the 
caudex thickened with the remains of dead leaves; stems 1-2-flowered ; 
leaves trisected, the segments obtusely incised-dentate, the medial 
ones petiolate; cauline leaves 1-2, 3-parted; flowers rather large; 
calyx reflexed, pubescent, exceeded by the 10-15 cuneate petals; 
achenes ovate, compressed, the fruit globose. Var. Bonplandianus 
(HBK.) Wedd. is taller, slender, the leaves glabrate or glabrous 
above, the cauline pubescence subappressed. Plant used as a caustic 
(Raimondi). The Wilkes Expedition specimen was determined by 
Gray as a variety of R. Bonplandianus HBK., without assigning a 
name, but with the description "upper leaves pubescent, pilose 



FLORA OF PERU 655 

also above; stems decumbent or reclining and flowers as large as 
those of R. acris." Low forms like Isern 2220 seem to approach 
R. sibbaldioides. 

Lima: Bafios, Wilkes Exped. Cuzco: (Gay); (Herrera 106). 
Puno: Sandia, grass steppes, 2,500 meters, Weberbauer 597. Cara- 
baya, Raimondi. Junin: Palca, Isern 2220. Cajamarca: Nancho, 
Cutervo, Raimondi. Ancash: Cajatambo, Raimondi. Bolivia to 
Colombia. "Yerba centilla," "Soliman," "huarancayza," "cienta." 

Ranunculus repens L. Sp. PL 554. 1753. 

Creeping, nearly glabrous or more or less pubescent; leaves 
palmately 3-parted, the roundish lobes deeply 3-lobed and serrate, 
frequently variegated or spotted with white; flowers 2 cm. broad or 
broader, the sepals spreading only in fruit; achenes mostly beaked 
by the short, recurved, usually persistent style. Introduced from 
Europe. 

Junin: Tarma, 3,100 meters, shaded stream bank, Killip & 
Smith 21904- Widely distributed. 

Ranunculus sarmentoso-repens Steud. Flora 39: 403. 1856. 

Glabrous throughout, the sarmentose, creeping stems with diffuse, 
weak branches; leaves subfasciculate, small, coriaceous, rotund or 
rarely ovate, more or less cordate at the base, entire, 1.5 cm. broad; 
flowers small; achenes few, ovate, glabrous, scarcely mucronate. 
Seems to be related to R. humilis Colla (Steudel). 

Puno: Azangaro (Lechler 1778). 

Ranunculus setoso-pilosus Steud. Flora 39: 404. 1856. 

Stems low, branching, the well-petioled radical leaves pinnately 
dissected with obtuse, short, simple or denticulate lobes, setulose- 
pilose throughout; peduncle equaling or longer than the petiole, 
1-flowered; flowers 12-15 mm. wide; sepals setulose-pilose, some- 
what exceeded by the obovate, striate petals; achenes ovate, reflexed, 
acuminate, the style uncinate. Stems weak, 4-6 cm. long. Prob- 
ably the same as R. praemorsus. 

Puno: Sachapata (Lechler 2709). 

Ranunculus sibbaldioides HBK. ex DC. Syst. 1: 293. 
1818; 303. 

A low perennial, the solitary flowers seemingly sessile in the 
congested radical leaves, these petioled, pinnatifid, pilose beneath, 
the segments often 2 on each side, ovate or obovate, obtuse, incised- 



656 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

dentate, subtrifid at the tip; flowers small; calyx spreading, the 
sepals pilose without, shorter than the oblong-obtuse petals; achenes 
ovate, subcompressed, smooth. R. Sprucei Briq. of Ecuador has 
spreading pubescence, the petals 5-6 mm. long. 

Junin: Yauli, 4,400 meters, Weberbauer 277; 221. Huanuco: 
Punco, 4,050 meters, alpine grassland, 2473. Ecuador. 

Ranunculus Sodiroi Franchet, Bull. Acad. Intern. Geogr. Bot. 
10: 209. 1901. 

Roots fibrous, the ascending stem often rooting at the lower 
nodes; leaves tripinnate, glabrous, mostly radical, the very long 
petiole more or less villous; petals yellow, entire, longer than the 
reflexed sepals; achenes long-beaked, greenish brown and velutinous. 
Suggests R. acris in appearance, but easily recognized by the leaves, 
composed of seven 3-lobed segments spaced regularly, all glabrous, 
and with divisions of equal size (Franchet). 

Cajamarca: Raimondi 3200. Ecuador. 

Ranunculus trichophyllus Chaix in Villars, Hist. PI. Dauph. 
1: 335. 1786. R. aquatilis L. vars. Wedd. Chlor. And. 2: 299. 1857. 

Aquatic, usually somewhat pubescent, often with both more or 
less dilated floating and dissected immersed leavee, the formsr 
variously lobed, their stipular sheaths more pubescent, broader, and 
less completely adnate than those of the dissected leaves, the latter 
usually collapsing out of water; flowers generally 1-1.5 cm. broad; 
petals rarely contiguous, 3-9-veined, about twice longer than the 
calyx; stamens usually 10-12; carpels ordinarily 16-24; achenes to 
1.5 mm., rarely to 2 mm. long, pubescent or glabrous; style tip 
usually deciduous, the remaining short beak subterminal or sub- 
lateral. Very variable. Description based on Drew, Rhodora 
38: 17. 1936. 

Loreto: Pebas, Isern 2267. Puno: Lake Titicaca, Weberbauer 
1365. Chuquibambilla, 3,900 meters, in rivulet on puna, Pennell 
13400. Temperate South and North America; Eurasia. 

6. KRAPFIA DC. 

Aspidophyllum Ulbrich; Rhopalopodium Ulbrich. 

Caulescent or acaulescent, perennial herbs with fleshy leaves. 
Flowers medium-sized to very large, greenish or purplish to orange, 
the colored sepals 5-9. Petals thin, with 1 nectary. Stamen column 
fleshy and clavate. Plants of moist, often limestone grasslands, 



FLORA OF PERU 657 

with one exception. With Laccopetalum they could be retained more 
conveniently in Ranunculus as sections of that polymorphic genus. 

Leaves entire or toothed but neither lobed nor divided. 

Petioles slender, not at all winged K. clypeata. 

Petioles thick, winged. 

Flowers purplish or roseate green or yellow; leaves, at least 
above, densely long-pubescent, conspicuously ligulate by 
the extended petiole wing. 

Leaves glabrous beneath K. Raimondii. 

Leaves more or less pubescent beneath, at least at first. 

K. cochlearifolia. 
Flowers greenish yellow; leaves above short-pubescent; petiole 

wing not or not conspicuously extended K . Herrerae. 

Leaves deeply toothed, lobed, or dissected, at least in part. 
Lobes never pinnately dissected; some leaves sometimes merely 

toothed. 
Petals with a stipe-like claw 1-2 cm. long, roseate; leaves 

glabrescent or beneath loosely pilose K . Weberbaueri. 

Petals not or scarcely clawed, orange or yellow; leaves, especially 

beneath, more or less densely appressed-villous. 
Leaves deeply 3-5-lobed, the lobes deeply toothed. 

Sepals red; petals orange K. Gusmanni. 

Sepals roseate; petals yellow K. flava. 

Leaves more or less 3-lobed, but the lobes merely coarsely 

toothed K. ranunculina. 

Lobes more or less pinnately dissected K. Lechleri. 

Krapfia clypeata (Ulbrich) Standl. & Macbr., comb. nov. 
Aspidophyllum clypeatum Ulbrich, Notizbl. Bot. Gart. Berlin 8: 
268. 1922. 

Lower leaves 10 cm. long or longer, the blades 6-12 cm. long, 
4-8 cm. broad, acutely serrulate, sometimes incised, silky-villous 
except in age; stems erect, villous, taller than the basal leaves, 
branching above and bearing terminal, solitary flowers 2-2.5 cm. 
broad; sepals 5, about 2 cm. long and broad, yellowish green, silky- 
villous without; petals 5, broadly clawed, to 15 mm. long and broad, 
glabrous; nectary indistinctly limited; stamen column about 8 mm. 
high. Related technically to K. Raimondii. Ulbrich at one time 
in Herb. Berlin referred this plant there, but since, in his later 



658 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

judgment, it constitutes a separate genus on the basis of its ligneous 
roots, greenish sepals, and ligneous pericarp, I have not felt free to 
take up his herbarium name. 

Huanuco: Above Mufia, 3,800 meters, Weberbauer 6728. 

Krapfia cochlearifolia (R. & P.) Standl. & Macbr., comb. nov. 
Ranunculus cochlearifolius R. & P. ex DC. in Deless. Icon. Select. 
1 : 10. 1820. R. macropetalus DC. Prodr. 1 : 29. 1824. Rhopalopodium 
cochlearifolium Ulbrich, Notizbl. Bot. Gart. Berlin 8: 263. 1922. 

Very similar to K. Raimondii and probably not specifically 
distinct; leaves, petiole wings, and ligule one- third to one-half 
smaller. Illustrated, Hook. Icon. pi. 1821. Neg. 27651. 

Huanuco: Above Muna, 3,800 meters, Weberbauer 6729; (Pearce 
525}. Junin: Huancayo, 4,200 meters, Weberbauer 6684- Without 
locality, Ruiz & Pawn (type). 

Krapfia flava (Ulbrich) Standl. & Macbr., comb. nov. Rhopa- 
lopodium flavum Ulbrich, Notizbl. Bot. Gart. Berlin 8: 260. 1922. 

Similar to K. Gusmanni, but the rosulate basal leaves often more 
deeply 3-5-lobed, the upper strongly reduced; stems erect, the 
solitary flowers 2.5-3 cm. broad, the outer segments dark rose, the 
inner pale yellow; nectary with small lateral plicae. Mossy, boggy 
places. 

Huanuco: Above Muna, 3,500 meters, Weberbauer 6790 (type). 
Tambo de Vaca, 3,900 meters, 4349 (det. Ulbrich). 

Krapfia Gusmanni (HBK.) Standl. & Macbr., comb. nov. 
Ranunculus Gusmanni HBK. Nov. Gen. & Sp. 5: 43. 1821. Rhopa- 
lopodium Gusmanni Ulbrich, Notizbl. Bot. Gart. Berlin 8: 258. 1922. 

Basal leaves 3-parted or lobed, the lobes more or less toothed 
but never pinnately dissected; flowers typically 3-6 cm. broad, 
often more than one; sepals and petals mostly 5, the latter scarcely 
clawed; fruit and stamen column subequal; nectary plane. Some 
of the plants are var. Weberbaueri (Ulbrich) Standl. & Macbr., 
comb. nov. Rhopalopodium Gusmanni var. Weberbaueri Ulbrich, 
loc. cit., with leaves glabrate above and flowers 2-3 cm. broad. 
Illustrated, Wedd. Chlor. And. 2: pi. 82b (typical); Deless. Icon. 1: 
pi. 35. 1820 ("to var. Weberbaueri" Ulbrich). 

Cuzco: Valle del Paucartambo, 3,700-4,700 meters, Herrera 2336, 
2329. San Martin: Moyobamba, Stuebel 50a. Amazonas: Above 
Balsas, 3,500 meters, Weberbauer 4293 (type of var.). Junin: 
Near Yurac-yaco, Raimondi (det. Ulbrich). Huanuco: Southeast 



FLORA OF PERU 659 

of Huanuco, 3,300 meters, 2161. Puno: Agapata to San Jose", 
Raimondi (det. Ulbrich). Without locality, Weberbauer 6790. 

Krapfia Lechleri (Schlecht.) Standl. & Macbr., comb. nov. 
Ranunculus Lechleri Schlecht. ex Steud. Flora 39: 406. 1856. R. 
Gusmanni var. Lechleri Wedd. Chlor. And. 2: 304. 1857. R. haeman- 
thus Ulbrich, Bot. Jahrb. 37. 404. 1906. Rhopalopodium haeman- 
thum Ulbrich, Notizbl. Bot. Gart. Berlin 8: 257. 1922. 

A low, stemless or nearly stemless plant with basally rosulate, 
3-5-parted leaves; flowers subglobose, about 2 cm. broad, the reddish 
sepals 6-9, the shortly clawed petals 5-6; fruit head globose. This 
small or diminutive plant is easily recognized by the shaggy pubes- 
cence and finely divided leaves. The stem is more or less developed, 
rarely to 14 cm. high. Illustrated, Weberbauer, 206. Neg. 27647. 

Junin: Yauli, 4,600 meters, Weberbauer 286. Alpamina, 4,500 
meters, Weberbauer 5101. Morococha, Hauthal 372. Cerro de 
Pasco, Sawada 87. Puno: Sangaban (Lechler 2216}. Bolivia. 

Krapfia Raimondii (Wedd.) Standl. & Macbr., comb. nov. 
Ranunculus Raimondii Wedd. Chlor. And. 2: 305. 1857. Rhopa- 
lopodium Raimondii Ulbrich, Notizbl. Bot. Gart. Berlin 8: 262. 1922. 

Basal leaves to 60 cm. long or longer, the broadly oval or orbic- 
ular blades about 20 cm. long, the petiole wings 4-7 cm. broad, 
sometimes elongate-ligulate (to 7 cm. long and 6 cm. wide), per- 
manently silky-pubescent above; flowers 6-8 cm. broad, the sessile, 
suboblong sepals 5 cm. long, the broadly obovate-cuneate petals 
somewhat longer and narrowed to a claw 1 cm. long; sepals and 
petals purplish green. A magnificent plant. Raimondi and Weber- 
bauer have explained the common names as referring to the folk 
custom of tossing the flowers to the mouths of children who were 
slow to talk. 

Huancavelica: Asapara, 4,200 meters, Weberbauer 6684- Prov. 
Pataz, 4,000 meters (Raimondi). Huanuco: Tambo de Vaca, 
Raimondi; 3,900 meters, wet, rocky uplands, 4902. "Rima-rima," 
"lima-lima." 

Krapfia ranunculina DC. Syst. 1: 228. 1818. Ranunculus 
Gusmanni var. Krapfia Wedd. Chlor. And. 2: 304. 1857. Rhopalo- 
podium ranunculinum Ulbrich, Notizbl. Bot. Gart. Berlin 8: 
260. 1922. 

Similar to K. flavum, but the basal leaves not parted, the pubes- 
cence sparser, the leaves of the 1-2-flowered, slender stems bract- 



660 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII 

like, and the flowers yellow, the outer parts finally brown. Illus- 
trated, Deless. Icon. 1 : pi. 35. 

Junin: Huacapistana, 3,400 meters, Weberbauer 2248. Huasa- 
huasi (according to Delessert). 

Krapfia Weberbaueri (Ulbrich) Standl. & Macbr., comb. nov. 
Rkopalopodium Weberbaueri Ulbrich, Notizbl. Bot. Gart. Berlin 8: 
261. 1922. 

Basal leaves 3-5-parted above the middle, with the strongly 
sheathed base 10-30 cm. long or even longer, the thick blades gla- 
brous above, reniform or suborbicular; stems erect, their 1-3 leaves 
greatly reduced; flowers terminal, solitary, to 5 cm. broad, the 5-6 
outer segments 4-4.5 cm. long, the inner 1.5-2 cm. long, to 2-3 mm. 
broad; nectary elongate-sulcate, with sharply differentiated, circular 
border. Related to K. Gusmanni and allies. The native name, 
according to the collector, means "speak, speak." 

Huanuco: Near Monzon, 3,400 meters, Weberbauer 3339 (type). 
Junin: Andamarca, Raimondi (det. Ulbrich, form with leaves in 
part merely toothed). Ancash: Above Huaraz, 4,200 meters, Weber- 
bauer 3745. "Rima-rima," "lima-lima." 

7. LACCOPET ALUM Ulbrich 

Leaves sinuate-serrate. Flowers solitary, the leathery, nearly 
orbicular sepals much longer than the deeply pitted, fleshy petals. 
Fruit heads globose. Technically separated from Ranunculus and 
Krapfia by the many and deep nectar pores in the petals. See 
Ulbrich, Notizbl. Bot. Gart. Berlin 8: 271-272. 1921. A decoc- 
tion of this beautiful plant is said to be used in Peru as a remedy for 
throat and pulmonary affections, and, according to Weddell, the 
inhabitants feed the plant to breeding cattle as an excitant and 
to assure fecundity. Since this author confused this species and 
Krapfia Raimondii (cf. Ulbrich), these uses may be shared with 
the latter and similar forms. 

Laccopetalum giganteum (Wedd.) Ulbrich, Bot. Jahrb. 37: 
404. 1906. Ranunculus giganteus Wedd. Chlor. And. 2: 30. 1857-62. 

Basal leaves to 70 cm. high; cauline leaves 1-3 and 6-8 cm. broad; 
flowers greenish, 10-15 cm. broad; sepals 5 (-6), 8-10 cm. long, 6-8 
cm. broad, the pores 2-4 mm. broad; carpels glabrous, scarcely 1 mm. 
long. Var. insignis Ulbrich has flowers to 10 cm. broad, petals less 
densely fimbriate, basal leaves shorter (to 45 cm. long) and wider 



FLORA OF PERU 661 

(to 17 cm. wide), with few coarser, fimbriate teeth. Illustrated, 
Weberbauer, 269, and Ulbrich, op. cit. 405, 406. 

Cajamarca: Between Cajamarca and Hualgayoc, 4,100 meters, 
Weberbauer 4224- Prov. Pataz, Raimondi. Ancash: Cordillera 
Pelagatos, 4,200 meters, Weberbauer 7020a. "Huamanripa," "pac- 
ra," "pagra," "pacra-pacra." 

8. MYOSURUSL. 

A small annual of very wet places, marked by its linear leaves 
and spike-like receptacle. Besides the following, M. minimus L., 
widely dispersed, may be expected as an introduction and may be 
known by its nearly filiform leaves and generally spurred petals. 

Myosurus apetalus Gay, Fl. Chile 1: 130. 1845. 

Leaves linear; petals none; sepals shortly spurred. Treated by 
Huth as a variety of M. aristatus Benth., a later published name. 
Illustrated, Gay, Atlas pi. 1 . 

Peru: Probably. Chile. 

9. CALTHAL. 

Reference: Hill, Ann. Bot. 32: 421-435. 1918. 

A small, glabrous, tufted, rather fleshy plant, well marked by 
its leaves that bear at the base prominent appendages. Sepals petal- 
like. Follicles dehiscent along the ventral suture. 

Caltha alata Hill, Ann. Bot. 32: 428. 1918. 

About 2 cm. high, from an elongate, fleshy root; petioles 1 cm. 
long, amply vaginate below; blades shortly cordate-sagittate, 6-8 
mm. long and broad, obtuse or retuse, more or less sinuate-margined ; 
appendages erect, wing-like, oblong, obtuse, arising from the leaf 
base near the midrib; peduncle about 1.5 cm. long; perianth segments 
5, ovate or elliptic, obtuse, yellow; carpels many, curved, erect, 
acutely beaked, mostly 6 mm. long; seeds about 6. This is C. 
sagittata of authors in part, not Cav., a distinct species, as shown by 
Sir Arthur W. Hill, confined to the Magellanic region. It has been 
confused also with C. andicola (Gay) Walp., definitely Chilean. 
C. involuta Hill, op. cit. 427, of Ecuador, is to be expected; its broadly 
ovate- or cordate-sagittate leaves are 12-15 mm. long and the appen- 
dages are infolded from the leaf base, erect, and parallel to the 
midnerve. Illustrated, Wedd. Chlor. And. pi. 83. 

Puno: Carabaya (Weddell}. Agapata, Lechler 1953. Poto, be- 
tween Distichia cushions (Weberbauer, 219). Bolivia.